Wikipedia - Readings On Life Evolutionary History V1

Evolutionary history of living beings
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Khalid Chraibi
01 February 2011
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Evolutionary history of life

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"History of evolution" redirects here. It is not to be confused with History of evolutionary thought.
"History of life" redirects here. For other uses, see Life history.
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History
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The evolutionary history of life on Earth traces the processes by which living and fossil organisms
evolved. It stretches from the origin of life on Earth, thought to be over 3,500 million years ago, to the present
day. The similarities between all present day organisms indicate the presence of a common ancestor from
which all known species have diverged through the process of evolution.[1]
Microbial mats of coexisting bacteria and archaea were the dominant form of life in the early Archean
and many of the major steps in early evolution are thought to have taken place within them.[2] The evolution
of oxygenic photosynthesis, around 3,500 million years ago, eventually led to the oxygenation of the
atmosphere, beginning around 2,400 million years ago.[3] The earliest evidence of eukaryotes (complex cells
with organelles), dates from 1,850 million years ago,[4][5] and while they may have been present earlier, their
diversification accelerated when they started using oxygen in their metabolism. Later, around 1,700 million
years ago, multicellular organisms began to appear, with differentiated cells performing specialised functions.
[6]
The earliest land plants date back to around 450 million years ago,[7] though evidence suggests that
algal scum formed on the land as early as 1,200 million years ago. Land plants were so successful that they
are thought to have contributed to the late Devonian extinction event.[8] Invertebrate animals appear during
the Vendian period,[9] while vertebrates originated about 525 million years ago during the Cambrian
explosion.[10]
During the Permian period, synapsids, including the ancestors of mammals, dominated the land,[11]
but the Permian–Triassic extinction event 251 million years ago came close to wiping out all complex life.[12]
During the recovery from this catastrophe, archosaurs became the most abundant land vertebrates,
displacing therapsids in the mid-Triassic.[13] One archosaur group, the dinosaurs, dominated the Jurassic
and Cretaceous periods,[14] with the ancestors of mammals surviving only as small insectivores.[15] After
the Cretaceous–Tertiary extinction event 65 million years ago killed off the non-avian dinosaurs[16] mammals
increased rapidly in size and diversity.[17] Such mass extinctions may have accelerated evolution by
providing opportunities for new groups of organisms to diversify.[18]
Fossil evidence indicates that flowering plants appeared and rapidly diversified in the Early
Cretaceous, between 130 million years ago and 90 million years ago, probably helped by coevolution with
pollinating insects. Flowering plants and marine phytoplankton are still the dominant producers of organic
matter. Social insects appeared around the same time as flowering plants. Although they occupy only small
parts of the insect "family tree", they now form over half the total mass of insects. Humans evolved from a
lineage of upright-walking apes whose earliest fossils date from over 6 million years ago. Although early
members of this lineage had chimpanzee-sized brains, there are signs of a steady increase in brain size after
about 3 million years ago.
Contents
[hide]
• 1 Earliest history of Earth
• 2 Earliest evidence for life on Earth
• 3 Origins of life on Earth
• 3.1 Life "seeded" from elsewhere
• 3.2 Independent emergence on Earth
• 3.2.1 Replication first: RNA world
• 3.2.2 Metabolism first: Iron-sulfur world
• 3.2.3 Membranes first: Lipid world
• 3.2.4 The clay theory
• 4 Environmental and evolutionary impact of microbial mats
• 5 Diversification of eukaryotes
• 6 Multicellular organisms and sexual reproduction
• 6.1 Multicellularity
• 6.2 Evolution of sexual reproduction
• 6.3 Fossil evidence for multicellularity and sexual
reproduction
• 7 Emergence of animals
• 8 Colonization of land
• 8.1 Evolution of soil
• 8.2 Plants and the Late Devonian wood crisis
[edit] Earliest history of Earth
History of Earth and its life
-4500 —
–
-4000 —
–
-3500 —
–
-3000 —
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-2500 —
–
-2000 —
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-1500 —
–
-1000 —
–
-500 —
–
0—
Hadean
Archean
Protero
-zoic
Phanero
-zoic
Eo
Paleo
Meso
Neo
Paleo
Meso
Neo
Paleo
Meso
Ceno
←
Solar system formed
←
Impact formed Moon
←
? Cool surface, oceans, atmosphere
←
Late Heavy Bombardment
←
? Earliest evidence of life
←
Oxygenation of atmosphere
←
Earliest multicellular organism[19]
←
Earliest known fungi
←
Earliest known cnidarians
←
? Cambrian explosion
←
Earliest land invertebrates and plants
←
Earliest land vertebrates
←
Earliest known dinosaur
←
Extinction of non-avian dinosaurs
Scale:
Millions of years
Main article: History of the Earth
The oldest meteorite fragments found on Earth are about 4,540 million years old, this, coupled
primarily with the dating of ancient lead deposits, has put the estimated age of Earth at around that time.[20]
The Moon has the same composition as Earth's crust but does not contain an iron-rich core like the Earth's.
Many scientists think that about 40 million years later a planetoid struck the Earth, throwing into orbit crust
material that formed the Moon. Another hypothesis is that the Earth and Moon started to coalesce at the
same time but the Earth, having much stronger gravity, attracted almost all the iron particles in the area.[21]
Until recently the oldest rocks found on Earth were about 3,800 million years old,[20] leading
scientists to believe for decades that Earth's surface had been molten until then. Accordingly, they named
this part of Earth's history the Hadean eon, whose name means "hellish".[22] However analysis of zircons
formed 4,400 to 4,000 million years ago indicates that Earth's crust solidified about 100 million years after the
planet's formation and that the planet quickly acquired oceans and an atmosphere, which may have been
capable of supporting life.[23]
Evidence from the Moon indicates that from 4,000 to 3,800 million years ago it suffered a Late Heavy
Bombardment by debris that was left over from the formation of the Solar system, and the Earth should have
experienced an even heavier bombardment due to its stronger gravity.[22][24] While there is no direct
evidence of conditions on Earth 4,000 to 3,800 million years ago, there is no reason to think that the Earth
was not also affected by this late heavy bombardment.[25] This event may well have stripped away any
previous atmosphere and oceans; in this case gases and water from comet impacts may have contributed to
their replacement, although volcanic outgassing on Earth would have contributed at least half.[26]
[edit] Earliest evidence for life on Earth

The earliest identified organisms were minute and relatively featureless, and their fossils look like
small rods, which are very difficult to tell apart from structures that arise through abiotic physical processes.
The oldest undisputed evidence of life on Earth, interpreted as fossilized bacteria, dates to 3,000 million
years ago.[27] Other finds in rocks dated to about 3,500 million years ago have been interpreted as bacteria,
[28] with geochemical evidence also seeming to show the presence of life 3,800 million years ago.[29]
However these analyses were closely scrutinized, and non-biological processes were found which could
produce all of the "signatures of life" that had been reported.[30][31] While this does not prove that the
structures found had a non-biological origin, they cannot be taken as clear evidence for the presence of life.
Geochemical signatures from rocks deposited 3,400 million years ago have been interpreted as evidence for
life,[27][32] although these statements have not been thoroughly examined by critics.
[edit] Origins of life on Earth

Evolutionary tree showing the divergence of modern species from their common ancestor in the
center.[33] The three domains are colored, with bacteria blue, archaea green, and eukaryotes red.
Further information: Evidence of common descent, Common descent, and Homology (biology)
Biologists reason that all living organisms on Earth must share a single last universal ancestor,
because it would be virtually impossible that two or more separate lineages could have independently
developed the many complex biochemical mechanisms common to all living organisms.[34][35] As previously
mentioned the earliest organisms for which fossil evidence is available are bacteria, cells far too complex to
have arisen directly from non-living materials.[36] The lack of fossil or geochemical evidence for earlier
organisms has left plenty of scope for hypotheses, which fall into two main groups: 1) that life arose
spontaneously on Earth or 2) that it was "seeded" from elsewhere in the universe.
[edit] Life "seeded" from elsewhere

Main articles: Panspermia, Life on Mars, Fermi paradox, and Rare Earth hypothesis
The idea that life on Earth was "seeded" from elsewhere in the universe dates back at least to the
fifth century BCE.[37] In the twentieth century it was proposed by the physical chemist Svante Arrhenius,[38]
by the astronomers Fred Hoyle and Chandra Wickramasinghe,[39] and by molecular biologist Francis Crick
and chemist Leslie Orgel.[40] There are three main versions of the "seeded from elsewhere" hypothesis:
from elsewhere in our Solar system via fragments knocked into space by a large meteor impact, in which
case the only credible source is Mars;[41] by alien visitors, possibly as a result of accidental contamination by
micro-organisms that they brought with them;[40] and from outside the Solar system but by natural means.
[38][41] Experiments suggest that some micro-organisms can survive the shock of being catapulted into
space and some can survive exposure to radiation for several days, but there is no proof that they can
survive in space for much longer periods.[41] Scientists are divided over the likelihood of life arising
independently on Mars,[42] or on other planets in our galaxy.[41]
[edit] Independent emergence on Earth

Main article: Abiogenesis
Life on Earth is based on carbon and water. Carbon provides stable frameworks for complex
chemicals and can be easily extracted from the environment, especially from carbon dioxide. The only other
element with similar chemical properties, silicon, forms much less stable structures and, because most of its
compounds are solids, would be more difficult for organisms to extract. Water is an excellent solvent and has
two other useful properties: the fact that ice floats enables aquatic organisms to survive beneath it in winter;
and its molecules have electrically negative and positive ends, which enables it to form a wider range of
compounds than other solvents can. Other good solvents, such as ammonia, are liquid only at such low
temperatures that chemical reactions may be too slow to sustain life, and lack water's other advantages.[43]
Organisms based on alternative biochemistry may however be possible on other planets.[44]
Research on how life might have emerged unaided from non-living chemicals focuses on three
possible starting points: self-replication, an organism's ability to produce offspring that are very similar to
itself; metabolism, its ability to feed and repair itself; and external cell membranes, which allow food to enter
and waste products to leave, but exclude unwanted substances.[45] Research on abiogenesis still has a long
way to go, since theoretical and empirical approaches are only beginning to make contact with each other.
[46][47]
[edit] Replication first: RNA world

Main articles: Last universal ancestor and RNA world
Even the simplest members of the three modern domains of life use DNA
to record their "recipes" and a complex array of RNA and protein molecules to
"read" these instructions and use them for growth, maintenance and self-
replication. This system is far too complex to have emerged directly from non-
living materials.[36] The discovery that some RNA molecules can catalyze both
their own replication and the construction of proteins led to the hypothesis of
earlier life-forms based entirely on RNA.[48] These ribozymes could have formed
an RNA world in which there were individuals but no species, as mutations and
horizontal gene transfers would have meant that the offspring in each generation
were quite likely to have different genomes from those that their parents started
with.[49] RNA would later have been replaced by DNA, which is more stable and
therefore can build longer genomes, expanding the range of capabilities a single
organism can have.[49][50][51] Ribozymes remain as the main components of
ribosomes, modern cells' "protein factories".[52]
Although short self-replicating RNA molecules have been artificially
produced in laboratories,[53] doubts have been raised about where natural non-
biological synthesis of RNA is possible.[54] The earliest "ribozymes" may have
been formed of simpler nucleic acids such as PNA, TNA or GNA, which would
have been replaced later by RNA.[55][56]
In 2003 it was proposed that porous metal sulfide precipitates would
assist RNA synthesis at about 100 °C (212 °F) and ocean-bottom pressures near
hydrothermal vents. In this hypothesis lipid membranes would be the last major The replicator in
cell components to appear and until then the proto-cells would be confined to the virtually all known life is
pores.[57] deoxyribonucleic acid.
DNA's structure and
replication systems are
far more complex than
those of the original
replicator.[36]
[edit] Metabolism first: Iron-sulfur world
Main article: Iron-sulfur world theory
A series of experiments starting in 1997 showed that early stages in the formation of proteins from
inorganic materials including carbon monoxide and hydrogen sulfide could be achieved by using iron sulfide
and nickel sulfide as catalysts. Most of the steps required temperatures of about 100 °C (212 °F) and
moderate pressures, although one stage required 250 °C (482 °F) and a pressure equivalent to that found
under 7 kilometres (4.3 mi) of rock. Hence it was suggested that self-sustaining synthesis of proteins could
have occurred near hydrothermal vents.[58]
[edit] Membranes first: Lipid world

It has been suggested that double-walled "bubbles" of
lipids like those that form the external membranes of cells may
have been an essential first step.[59] Experiments that = water-attracting heads of lipid
simulated the conditions of the early Earth have reported the
formation of lipids, and these can spontaneously form
liposomes, double-walled "bubbles", and then reproduce = water-repellent tails
themselves. Although they are not intrinsically information-
carriers as nucleic acids are, they would be subject to natural
selection for longevity and reproduction. Nucleic acids such as
RNA might then have formed more easily within the liposomes
than they would have outside.[60]
Cross-section through a liposome.

[edit] The clay theory
Main articles: Graham Cairns-Smith#Clay Theory and RNA world
RNA is complex and there are doubts about whether it can be produced non-biologically in the wild.
[54] Some clays, notably montmorillonite, have properties that make them plausible accelerators for the
emergence of an RNA world: they grow by self-replication of their crystalline pattern; they are subject to an
analog of natural selection, as the clay "species" that grows fastest in a particular environment rapidly
becomes dominant; and they can catalyze the formation of RNA molecules.[61] Although this idea has not
become the scientific consensus, it still has active supporters.[62]
Research in 2003 reported that montmorillonite could also accelerate the conversion of fatty acids
into "bubbles", and that the "bubbles" could encapsulate RNA attached to the clay. These "bubbles" can then
grow by absorbing additional lipids and then divide. The formation of the earliest cells may have been aided
by similar processes.[63]
A similar hypothesis presents self-replicating iron-rich clays as the progenitors of nucleotides, lipids
and amino acids.[64]
[edit] Environmental and evolutionary impact of microbial mats

Main articles: Microbial mat and Oxygen catastrophe
Modern stromatolites in Shark Bay, Western Australia.
Microbial mats are multi-layered, multi-species colonies of bacteria and other organisms that are
generally only a few millimeters thick, but still contain a wide range of chemical environments, each of which
favors a different set of micro-organisms.[65] To some extent each mat forms its own food chain, as the by-
products of each group of micro-organisms generally serve as "food" for adjacent groups.[66]
Stromatolites are stubby pillars built as microbes in mats slowly migrate upwards to avoid being
smothered by sediment deposited on them by water.[65] There has been vigorous debate about the validity
of alleged fossils from before 3,000 million years ago,[67] with critics arguing that so-called stromatolites
could have been formed by non-biological processes.[30] In 2006 another find of stromatolites was reported
from the same part of Australia as previous ones, in rocks dated to 3,500 million years ago.[68]
In modern underwater mats the top layer often consists of photosynthesizing cyanobacteria which
create an oxygen-rich environment, while the bottom layer is oxygen-free and often dominated by hydrogen
sulfide emitted by the organisms living there.[66] It is estimated that the appearance of oxygenic
photosynthesis by bacteria in mats increased biological productivity by a factor of between 100 and 1,000.
The reducing agent used by oxygenic photosynthesis is water, which is much more plentiful than the
geologically-produced reducing agents required by the earlier non-oxygenic photosynthesis.[69] From this
point onwards life itself produced significantly more of the resources it needed than did geochemical
processes.[70] Oxygen is toxic to organisms that are not adapted to it, but greatly increases the metabolic
efficiency of oxygen-adapted organisms.[71][72] Oxygen became a significant component of Earth's
atmosphere about 2,400 million years ago.[73] Although eukaryotes may have been present much earlier,
[74][75] the oxygenation of the atmosphere was a prerequisite for the evolution of the most complex
eukaryotic cells, from which all multicellular organisms are built.[76] The boundary between oxygen-rich and
oxygen-free layers in microbial mats would have moved upwards when photosynthesis shut down overnight,
and then downwards as it resumed on the next day. This would have created selection pressure for
organisms in this intermediate zone to acquire the ability to tolerate and then to use oxygen, possibly via
endosymbiosis, where one organism lives inside another and both of them benefit from their association.[2]
Cyanobacteria have the most complete biochemical "toolkits" of all the mat-forming organisms.
Hence they are the most self-sufficient of the mat organisms and were well-adapted to strike out on their own
both as floating mats and as the first of the phytoplankton, providing the basis of most marine food chains.[2]
Eukaryotes
Bikonta
Apusozoa
[edit] Diversification of eukaryotes Archaeplastida (Land plants

Main article: Eukaryote green algae, red algae, and
glaucophytes)
Eukaryotes may have been present long before the
oxygenation of the atmosphere,[74] but most modern eukaryotes
require oxygen, which their mitochondria use to fuel the Chromalveolata
production of ATP, the internal energy supply of all known cells.
[76] In the 1970s it was proposed and, after much debate, widely
accepted that eukaryotes emerged as a result of a sequence of Rhizaria
endosymbioses between "procaryotes". For example: a
predatory micro-organism invaded a large procaryote, probably
an archaean, but the attack was neutralized, and the attacker Excavata
took up residence and evolved into the first of the mitochondria;
one of these chimeras later tried to swallow a photosynthesizing
cyanobacterium, but the victim survived inside the attacker and
the new combination became the ancestor of plants; and so on.
After each endosymbiosis began, the partners would have Unikonta
eliminated unproductive duplication of genetic functions by re-
arranging their genomes, a process which sometimes involved Amoebozoa
transfer of genes between them.[79][80][81] Another hypothesis
proposes that mitochondria were originally sulfur- or hydrogen- Opisthokonta
metabolising endosymbionts, and became oxygen-consumers
later.[82] On the other hand mitochondria might have been part Metazoa
of eukaryotes' original equipment.[83] (Animals)
Choanozoa
Eumycota
(Fungi)
There is a debate about when eukaryotes first appeared: the presence of steranes in Australian
shales may indicate that eukaryotes were present 2,700 million years ago;[75] however an analysis in 2008
concluded that these chemicals infiltrated the rocks less than 2,200 million years ago and prove nothing
about the origins of eukaryotes.[84] Fossils of the alga Grypania have been reported in 1,850 million-year-old
rocks (originally dated to 2,100 million years ago but later revised[5]), and indicates that eukaryotes with
organelles had already evolved.[85] A diverse collection of fossil algae were found in rocks dated between
1,500 million years ago and 1,400 million years ago.[86] The earliest known fossils of fungi date from 1,430
million years ago.[87]
[edit] Multicellular organisms and sexual reproduction

[edit] Multicellularity
Main articles: Multicellular organism , Evolution of multicellularity , and Sexual reproduction
The simplest definitions of "multicellular", for example "having multiple cells", could include colonial
cyanobacteria like Nostoc. Even a professional biologist's definition such as "having the same genome but
different types of cell" would still include some genera of the green alga Volvox, which have cells that
specialize in reproduction.[88] Multicellularity evolved independently in organisms as diverse as sponges and
other animals, fungi, plants, brown algae, cyanobacteria, slime moulds and myxobacteria.[5][89] For the sake
of brevity this article focuses on the organisms that show the greatest specialization of cells and variety of
cell types, although this approach to the evolution of complexity could be regarded as "rather
anthropocentric".[90]
A slime mold solves a maze. The mold (yellow) explored and filled the maze (left). When the
researchers placed sugar (red) at two separate points, the mold concentrated most of its mass there and left
only the most efficient connection between the two points (right).[91]
The initial advantages of multicellularity may have included: increased resistance to predators, many
of which attacked by engulfing; the ability to resist currents by attaching to a firm surface; the ability to reach
upwards to filter-feed or to obtain sunlight for photosynthesis;[92] the ability to create an internal environment
that gives protection against the external one;[90] and even the opportunity for a group of cells to behave
"intelligently" by sharing information.[91] These features would also have provided opportunities for other
organisms to diversify, by creating more varied environments than flat microbial mats could.[92]
Multicellularity with differentiated cells is beneficial to the organism as a whole but disadvantageous
from the point of view of individual cells, most of which lose the opportunity to reproduce themselves. In an
asexual multicellular organism, rogue cells which retain the ability to reproduce may take over and reduce the
organism to a mass of undifferentiated cells. Sexual reproduction eliminates such rogue cells from the next
generation and therefore appears to be a prerequisite for complex multicellularity.[92]
The available evidence indicates that eukaryotes evolved much earlier but remained inconspicuous
until a rapid diversification around 1,000 million years ago. The only respect in which eukaryotes clearly
surpass bacteria and archaea is their capacity for variety of forms, and sexual reproduction enabled
eukaryotes to exploit that advantage by producing organisms with multiple cells that differed in form and
function.[92]
[edit] Evolution of sexual reproduction

Main article: Evolution of sexual reproduction
The defining characteristic of sexual reproduction is recombination, in which each of the offspring
receives 50% of its genetic inheritance from each of the parents.[93] Bacteria also exchange DNA by
bacterial conjugation, the benefits of which include resistance to antibiotics and other toxins, and the ability to
utilize new metabolites.[94] However conjugation is not a means of reproduction, and is not limited to
members of the same species – there are cases where bacteria transfer DNA to plants and animals.[95]
The disadvantages of sexual reproduction are well-known: the genetic reshuffle of recombination
may break up favorable combinations of genes; and since males do not directly increase the number of
offspring in the next generation, an asexual population can out-breed and displace in as little as 50
generations a sexual population that is equal in every other respect.[93] Nevertheless the great majority of
animals, plants, fungi and protists reproduce sexually. There is strong evidence that sexual reproduction
arose early in the history of eukaryotes and that the genes controlling it have changed very little since then.
[96] How sexual reproduction evolved and survived is an unsolved puzzle.[97]
The Red Queen Hypothesis suggests that sexual reproduction provides protection against parasites,
because it is easier for parasites to evolve means of overcoming the defenses of genetically identical clones
than those of sexual species that present moving targets, and there is some experimental evidence for this.
However there is still doubt about whether it would explain the survival of sexual species if multiple similar
clone species were present, as one of the clones may survive the attacks of parasites for long enough to out-
breed the sexual species.[93]
The Mutation Deterministic Hypothesis assumes that each organism has more than one harmful
mutation and the combined effects of these mutations are more harmful than the sum of the harm done by
each individual mutation. If so, sexual recombination of genes will reduce the harm that bad mutations do to
offspring and at the same time eliminate some bad mutations from the gene pool by isolating them in
individuals that perish quickly because they have an above-average number of bad mutations. However the
evidence suggests that the MDH's assumptions are shaky, because many species have on average less
than one harmful mutation per individual and no species that has been investigated shows evidence of
synergy between harmful mutations.[93]
Horodyskia apparently re-arranged itself into fewer but larger main masses as the sediment grew
deeper round its base.[5]
The random nature of recombination causes the relative abundance of alternative traits to vary from
one generation to another. This genetic drift is insufficient on its own to make sexual reproduction
advantageous, but a combination of genetic drift and natural selection may be sufficient. When chance
produces combinations of good traits, natural selection gives a large advantage to lineages in which these
traits become genetically linked. On the other hand the benefits of good traits are neutralized if they appear
along with bad traits. Sexual recombination gives good traits the opportunities to become linked with other
good traits, and mathematical models suggest this may be more than enough to offset the disadvantages of
sexual reproduction.[97] Other combinations of hypotheses that are inadequate on their own are also being
examined.[93]
[edit] Fossil evidence for multicellularity and sexual reproduction

The Francevillian Group Fossil, dated to 2,100 million years ago, is the
earliest known fossil organism that is clearly multicellular.[19] This may have
had differentiated cells.[99] Another early multicellular fossil, Qingshania,[note
1] dated to 1,700 million years ago, appears to consist of virtually identical
cells. The red alga called Bangiomorpha, dated at 1,200 million years ago, is
the earliest known organism which certainly has differentiated, specialized
cells, and is also the oldest known sexually-reproducing organism.[92] The
1,430 million-year-old fossils interpreted as fungi appear to have been
multicellular with differentiated cells.[87] The "string of beads" organism
Horodyskia, found in rocks dated from 1,500 million years ago to 900 million
years ago, may have been an early metazoan;[5] however it has also been
interpreted as a colonial foraminiferan.[98]
Horodyskia may
have been an early
metazoan,[5] or a colonial
foraminiferan[98]
[edit] Emergence of animals Bilaterians
Main articles: Animal, Ediacara biota, Cambrian
Explosion, Burgess shale type fauna, and Stem group Deuterostomes
Animals are multicellular eukaryotes,[note 2] and are (chordates
distinguished from plants, algae, and fungi by lacking cell walls. hemichordates
[101] All animals are motile,[102] if only at certain life stages. All echinoderms
animals except sponges have bodies differentiated into separate Protostomes
tissues, including muscles, which move parts of the animal by
Ecdysozoa
contracting, and nerve tissue, which transmits and processes
(arthropods,
signals.[103]
nematodes, tardigrades,
The earliest widely-accepted animal fossils are rather etc.)
modern-looking cnidarians (the group that includes jellyfish, sea
anemones and hydras), possibly from around 580 million years Lophotrochozoa
ago, although fossils from the Doushantuo Formation can only (molluscs, annelids,
be dated approximately. Their presence implies that the brachiopods, etc.)
cnidarian and bilaterian lineages had already diverged.[104]
The Ediacara biota, which flourished for the last
40 million years before the start of the Cambrian,[105] were the
first animals more than a very few centimeters long. Many were
flat and had a "quilted" appearance, and seemed so strange that
there was a proposal to classify them as a separate kingdom,
Vendozoa.[106] Others, however, been interpreted as early
molluscs (Kimberella[107][108]), echinoderms (Arkarua[109]),
and arthropods (Spriggina,[110] Parvancorina[111]). There is still debate about the classification of these
specimens, mainly because the diagnostic features which allow taxonomists to classify more recent
organisms, such as similarities to living organisms, are generally absent in the Ediacarans. However there
seems little doubt that Kimberella was at least a triploblastic bilaterian animal, in other words significantly
more complex than cnidarians.[112]
The small shelly fauna are a very mixed collection of fossils found between the Late Ediacaran and
Mid Cambrian periods. The earliest, Cloudina, shows signs of successful defense against predation and may
indicate the start of an evolutionary arms race. Some tiny Early Cambrian shells almost certainly belonged to
molluscs, while the owners of some "armor plates", Halkieria and Microdictyon, were eventually identified
when more complete specimens were found in Cambrian lagerstätten that preserved soft-bodied animals.
[113]
Opabinia made the largest single contribution to modern interest in the Cambrian explosion.[114]
In the 1970s there was already a debate about whether the emergence of the modern phyla was
"explosive" or gradual but hidden by the shortage of Pre-Cambrian animal fossils.[113] A re-analysis of
fossils from the Burgess Shale lagerstätte increased interest in the issue when it revealed animals, such as
Opabinia, which did not fit into any known phylum. At the time these were interpreted as evidence that the
modern phyla had evolved very rapidly in the "Cambrian explosion" and that the Burgess Shale's "weird
wonders" showed that the Early Cambrian was a uniquely experimental period of animal evolution.[115] Later
discoveries of similar animals and the development of new theoretical approaches led to the conclusion that
many of the "weird wonders" were evolutionary "aunts" or "cousins" of modern groups[116] – for example that
Opabinia was a member of the lobopods, a group which includes the ancestors of the arthropods, and that it
may have been closely related to the modern tardigrades.[117] Nevertheless there is still much debate about
whether the Cambrian explosion was really explosive and, if so, how and why it happened and why it
appears unique in the history of animals.[118]
Acanthodians were among the earliest vertebrates with jaws.[119]

Most of the animals at the heart of the Cambrian explosion debate are protostomes, one of the two
main groups of complex animals. One deuterostome group, the echinoderms, many of which have hard
calcite "shells", are fairly common from the Early Cambrian small shelly fauna onwards.[113] Other
deuterostome groups are soft-bodied, and most of the significant Cambrian deuterostome fossils come from
the Chengjiang fauna, a lagerstätte in China.[120] The Chengjiang fossils Haikouichthys and Myllokunmingia
appear to be true vertebrates,[121] and Haikouichthys had distinct vertebrae, which may have been slightly
mineralized.[122] Vertebrates with jaws, such as the Acanthodians, first appeared in the Late Ordovician.
[123]
[edit] Colonization of land
Adaptation to life on land is a major challenge: all land organisms need to avoid drying-out and all
those above microscopic size have to resist gravity; respiration and gas exchange systems have to change;
reproductive systems cannot depend on water to carry eggs and sperm towards each other.[124][125]
Although the earliest good evidence of land plants and animals dates back to the Ordovician period
(488 to 444 million years ago), modern land ecosystems only appeared in the late Devonian, about
385 to 359 million years ago.[126]
[edit] Evolution of soil

Before the colonization of land, soil, a combination of mineral particles and decomposed organic
matter, did not exist. Land surfaces would have been either bare rock or unstable sand produced by
weathering. Water and any nutrients in it would have drained away very quickly.[126]
Lichens growing on concrete

Films of cyanobacteria, which are not plants but use the same photosynthesis mechanisms, have
been found in modern deserts, and only in areas that are unsuitable for vascular plants. This suggests that
microbial mats may have been the first organisms to colonize dry land, possibly in the Precambrian. Mat-
forming cyanobacteria could have gradually evolved resistance to desiccation as they spread from the seas
to tidal zones and then to land.[126] Lichens, which are symbiotic combinations of a fungus (almost always
an ascomycete) and one or more photosynthesizers (green algae or cyanobacteria),[127] are also important
colonizers of lifeless environments,[126] and their ability to break down rocks contributes to soil formation in
situations where plants cannot survive.[127] The earliest known ascomycete fossils date from 423 to 419
million years ago in the Silurian.[126]
Soil formation would have been very slow until the appearance of burrowing animals, which mix the
mineral and organic components of soil and whose feces are a major source of the organic components.
[126] Burrows have been found in Ordovician sediments, and are attributed to annelids ("worms") or
arthropods.[126][128]
[edit] Plants and the Late Devonian wood crisis

Main article: Evolutionary history of plants
Reconstruction of Cooksonia, a vascular plant from the Silurian
Fossilized trees from the Mid-Devonian Gilboa fossil forest
In aquatic algae, almost all cells are capable of photosynthesies and are nearly independent. Life on
land required plants to become internally more complex and specialized: photosynthesis was most efficient
at the top; roots were required in order to extract water from the ground; the parts in between became
supports and transport systems for water and nutrients.[124][129]
Spores of land plants, possibly rather like liverworts, have been found in Mid Ordovician rocks dated
to about 476 million years ago. In Mid Silurian rocks 430 million years ago there are fossils of actual plants
including clubmosses such as Baragwanathia; most were under 10 centimetres (3.9 in) high, and some
appear closely related to vascular plants, the group that includes trees.[129]
By the Late Devonian 370 million years ago, trees such as Archaeopteris were so abundant that they
changed river systems from mostly braided to mostly meandering, because their roots bound the soil firmly.
[130] In fact they caused a "Late Devonian wood crisis",[131] because:
• They removed more carbon dioxide from the atmosphere, reducing the greenhouse effect
and thus causing an ice age in the Carboniferous period.[132] In later ecosystems the carbon dioxide
"locked up" in wood is returned to the atmosphere by decomposition of dead wood. However the
earliest fossil evidence of fungi that can decompose wood also comes from the Late Devonian.[133]
• The increasing depth of plants' roots led to more washing of nutrients into rivers and seas by
rain. This caused algal blooms whose high consumption of oxygen caused anoxic events in deeper
waters, increasing the extinction rate among deep-water animals.[132]
[edit] Land invertebrates
Animals had to change their feeding and excretory systems, and most land animals developed
internal fertilization of their eggs. The difference in refractive index between water and air required changes
in their eyes. On the other hand in some ways movement and breathing became easier, and the better
transmission of high-frequency sounds in air encouraged the development of hearing.[125]
Some trace fossils from the Cambrian-Ordovician boundary about 490 million years ago are
interpreted as the tracks of large amphibious arthropods on coastal sand dunes, and may have been made
by euthycarcinoids,[134] which are thought to be evolutionary "aunts" of myriapods.[135] Other trace fossils
from the Late Ordovician a little over 445 million years ago probably represent land invertebrates, and there
is clear evidence of numerous arthropods on coasts and alluvial plains shortly before the Silurian-Devonian
boundary, about 415 million years ago, including signs that some arthropods ate plants.[136] Arthropods
were well pre-adapted to colonise land, because their existing jointed exoskeletons provided protection
against desiccation, support against gravity and a means of locomotion that was not dependent on water.
[137]
The fossil record of other major invertebrate groups on land is poor: none at all for non-parasitic
flatworms, nematodes or nemerteans; some parasitic nematodes have been fossilized in amber; annelid
worm fossils are known from the Carboniferous, but they may still have been aquatic animals; the earliest
fossils of gastropods on land date from the Late Carboniferous, and this group may have had to wait until leaf
litter became abundant enough to provide the moist conditions they need.[125]
The earliest confirmed fossils of flying insects date from the Late Carboniferous, but it is thought that
insects developed the ability to fly in the Early Carboniferous or even Late Devonian. This gave them a wider
range of ecological niches for feeding and breeding, and a means of escape from predators and from
unfavorable changes in the environment.[138] About 99% of modern insect species fly or are descendants of
flying species.[139]
[edit] Land vertebrates

Main article: Tetrapod
Acanthostega changed views about the early evolution of tetrapods.[140]

"Fish"
Acanthostega
Ichthyostega
Tulerpeton
Early
amphibians
Anthracosauria
Amniotes
Family tree of tetrapods[141]
Tetrapods, vertebrates with four limbs, evolved from other rhipidistian fish over a relatively short
timespan during the Late Devonian, between 370 million years ago and 360 million years ago.[142] From the
1950s to the early 1980s it was thought that tetrapods evolved from fish that had already acquired the ability
to crawl on land, possibly in order to go from a pool that was drying out to one that was deeper. However in
1987 nearly-complete fossils of Acanthostega from about 363 million years ago showed that this Late
Devonian transitional animal had legs and both lungs and gills, but could never have survived on land: its
limbs and its wrist and ankle joints were too weak to bear its weight; its ribs were too short to prevent its
lungs from being squeezed flat by its weight; its fish-like tail fin would have been damaged by dragging on
the ground. The current hypothesis is that Acanthostega, which was about 1 metre (3.3 ft) long, was a wholly
aquatic predator that hunted in shallow water. Its skeleton differed from that of most fish, in ways that
enabled it to raise its head to breathe air while its body remained submerged, including: its jaws show
modifications that would have enabled it to gulp air; the bones at the back of its skull are locked together,
providing strong attachment points for muscles that raised its head; the head is not joined to the shoulder
girdle and it has a distinct neck.[140]
The Devonian proliferation of land plants may help to explain why air-breathing would have been an
advantage: leaves falling into streams and rivers would have encouraged the growth of aquatic vegetation;
this would have attracted grazing invertebrates and small fish that preyed on them; they would have been
attractive prey but the environment was unsuitable for the big marine predatory fish; air-breathing would have
been necessary because these waters would have been short of oxygen, since warm water holds less
dissolved oxygen than cooler marine water and since the decomposition of vegetation would have used
some of the oxygen.[140]
Later discoveries revealed earlier transitional forms between Acanthostega and completely fish-like
animals.[143] Unfortunately there is then a gap of about 30 million years between the fossils of ancestral
tetrapods and Mid Carboniferous fossils of vertebrates that look well-adapted for life on land. Some of these
look like early relatives of modern amphibians, most of which need to keep their skins moist and to lay their
eggs in water, while others are accepted as early relatives of the amniotes, whose water-proof skins and
eggs enable them to live and breed far from water.[141]
[edit] Dinosaurs, birds and mammals

Main articles: Dinosaur evolution, Origin of Birds, and Evolution of mammals
Amniotes
Synapsids
Early synapsids
(extinct)
Pelycosaurs
Extinct pelycosaurs
Therapsids
Extinct therapsids
Mammaliformes
Extinct mammaliformes
Mammals
Sauropsids
Diapsids
Squ
(lizards and
snakes)
Archosaurs
Extinct
archosaurs
Crocodilians
Pterosaurs
(extinct)
Dinosaurs
Theropods
Extinct
theropods
Birds
Sauropods
(extinct)
Ornithischians
(extinct)
Possible family tree of dinosaurs, birds and mammals[145][146]
Amniotes, whose eggs can survive in dry environments, probably evolved in the Late Carboniferous
period, between 330 million years ago and 314 million years ago. The earliest fossils of the two surviving
amniote groups, synapsids and sauropsids, date from around 313 million years ago.[145][146] The synapsid
pelycosaurs and their descendants the therapsids are the most common land vertebrates in the best-known
Permian fossil beds, between 229 million years ago and 251 million years ago. However at the time these
were all in temperate zones at middle latitudes, and there is evidence that hotter, drier environments nearer
the Equator were dominated by sauropsids and amphibians.[147]
The Permian-Triassic extinction wiped out almost all land vertebrates,[148] as well as the great
majority of other life.[149] During the slow recovery from this catastrophe, estimated to be 30M years,[150] a
previously obscure sauropsid group became the most abundant and diverse terrestrial vertebrates: a few
fossils of archosauriformes ("shaped like archosaurs") have been found in Late Permian rocks,[151] but by
the Mid Triassic archosaurs were the dominant land vertebrates. Dinosaurs distinguished themselves from
other archosaurs in the Late Triassic, and became the dominant land vertebrates of the Jurassic and
Cretaceous periods, between 199 million years ago and 65 million years ago.[152]
During the Late Jurassic, birds evolved from small, predatory theropod dinosaurs.[153] The first birds
inherited teeth and long, bony tails from their dinosaur ancestors,[153] but some developed horny, toothless
beaks by the very Late Jurassic[154] and short pygostyle tails by the Early Cretaceous.[155]
While the archosaurs and dinosaurs were becoming more dominant in the Triassic, the mammaliform
successors of the therapsids could only survive as small, mainly nocturnal insectivores. This apparent set-
back may actually have promoted the evolution of mammals, for example nocturnal life may have
accelerated the development of endothermy ("warm-bloodedness") and hair or fur.[156] By 195 million years
ago in the Early Jurassic there were animals that were very nearly mammals.[157] Unfortunately there is a
gap in the fossil record throughout the Mid Jurassic.[158] However fossil teeth discovered in Madagascar
indicate that true mammals existed at least 167 million years ago.[159] After dominating land vertebrate
niches for about 150 million years, the dinosaurs perished 65 million years ago in the Cretaceous–Tertiary
extinction along with many other groups of organisms.[160] Mammals throughout the time of the dinosaurs
had been restricted to a narrow range of taxa, sizes and shapes, but increased rapidly in size and diversity
after the extinction,[161][162] with bats taking to the air within 13 million years,[163] and cetaceans to the sea
within 15 million years.[164]
[edit] Flowering plants
Main articles: Flowering plant and Gymnosperm
Gymnosperms Gymnosp
Gnetales
(gymnosperm)
Welwitschia Cycads
(gymnosperm) (gymnosperm)
Bennettitales
Ephedra
(gymnosperm)
Gingk
Bennettitales
Angiosperms Gnetales
(flowering plants) (gymnosperm)
Conifers
(gymnosperm)
One possible family tree of flowering plants.[165]
Another possible family tree.[166]

The 250,000 to 400,000 species of flowering plants outnumber all other ground plants combined, and
are the dominant vegetation in most terrestrial ecosystems. There is fossil evidence that flowering plants
diversified rapidly in the Early Cretaceous, between 130 million years ago and 90 million years ago,[165]
[166] and that their rise was associated with that of pollinating insects.[166] Among modern flowering plants
Magnolias are thought to be close to the common ancestor of the group.[165] However paleontologists have
not succeeded in identifying the earliest stages in the evolution of flowering plants.[165][166]
[edit] Social insects

Main article: Social insects
The social insects are remarkable because the great majority of individuals in each colony are sterile.
This appears contrary to basic concepts of evolution such as natural selection and the selfish gene. In fact
there are very few eusocial insect species: only 15 out of approximately 2,600 living families of insects
contain eusocial species, and it seems that eusociality has evolved independently only 12 times among
arthropods, although some eusocial lineages have diversified into several families. Nevertheless social
insects have been spectacularly successful; for example although ants and termites account for only about
2% of known insect species, they form over 50% of the total mass of insects. Their ability to control a territory
appears to be the foundation of their success.[167]
These termite mounds have survived a bush fire.
The sacrifice of breeding opportunities by most individuals has long been explained as a
consequence of these species' unusual haplodiploid method of sex determination, which has the paradoxical
consequence that two sterile worker daughters of the same queen share more genes with each other than
they would with their offspring if they could breed.[168] However Wilson and Hölldobler argue that this
explanation is faulty: for example, it is based on kin selection, but there is no evidence of nepotism in
colonies that have multiple queens. Instead, they write, eusociality evolves only in species that are under
strong pressure from predators and competitors, but in environments where it is possible to build "fortresses";
after colonies have established this security, they gain other advantages though co-operative foraging. In
support of this explanation they cite the appearance of eusociality in bathyergid mole rats,[167] which are not
haplodiploid.[169]
The earliest fossils of insects have been found in Early Devonian rocks from about 400 million years
ago, which preserve only a few varieties of flightless insect. The Mazon Creek lagerstätten from the Late
Carboniferous, about 300 million years ago, include about 200 species, some gigantic by modern standards,
and indicate that insects had occupied their main modern ecological niches as herbivores, detritivores and
insectivores. Social termites and ants first appear in the Early Cretaceous, and advanced social bees have
been found in Late Cretaceous rocks but did not become abundant until the Mid Cenozoic.[170]
[edit] Humans
Main article: Human evolution
Modern humans evolved from a lineage of upright-walking apes that has been traced back over 6
million years ago to Sahelanthropus.[171] The first known stone tools were made about 2.5 million years ago,
apparently by Australopithecus garhi, and were found near animal bones that bear scratches made by these
tools.[172] The earliest hominines had chimp-sized brains, but there has been a fourfold increase in the last
3 million years; a statistical analysis suggests that hominine brain sizes depend almost completely on the
date of the fossils, while the species to which they are assigned has only slight influence.[173] There is a
long-running debate about whether modern humans evolved all over the world simultaneously from existing
advanced hominines or are descendants of a single small population in Africa, which then migrated all over
the world less than 200,000 years ago and replaced previous hominine species.[174] There is also debate
about whether anatomically-modern humans had
an intellectual, cultural and technological "Great
Leap Forward" under 100,000 years ago and, if so,
whether this was due to neurological changes that
are not visible in fossils.[175]
[edit] Mass extinctions

Main article: Mass extinction
Life on Earth has suffered occasional
mass extinctions at least since 542 million years
ago. Although they are disasters at the time, mass
extinctions have sometimes accelerated the
evolution of life on Earth. When dominance of
particular ecological niches passes from one
group of organisms to another, it is rarely because
the new dominant group is "superior" to the old
and usually because an extinction event
eliminates the old dominant group and makes way
for the new one.[176][177]
The fossil record appears to show that the
gaps between mass extinctions are becoming
Apparent extinction intensity, i.e. the fraction of

genera going extinct at any given time, as reconstructed
from the fossil record. (Graph not meant to include
recent epoch of Holocene extinction event)
longer and the average and background rates of extinction are decreasing. Both of these phenomena could
be explained in one or more ways:[178]
• The oceans may have become more hospitable to life over the last 500 million years and less
vulnerable to mass extinctions: dissolved oxygen became more widespread and penetrated to
greater depths; the development of life on land reduced the run-off of nutrients and hence the risk of
eutrophication and anoxic events; and marine ecosystems became more diversified so that food
chains were less likely to be disrupted.[179][180]
• Reasonably complete fossils are very rare, most extinct organisms are represented only by
partial fossils, and complete fossils are rarest in the oldest rocks. So paleontologists have mistakenly
assigned parts of the same organism to different genera which were often defined solely to
accommodate these finds – the story of Anomalocaris is an example of this. The risk of this mistake is
higher for older fossils because these are often unlike parts of any living organism. Many of the
"superfluous" genera are represented by fragments which are not found again and the "superfluous"
genera appear to become extinct very quickly.[178]
Biodiversity in the fossil record, which is
"the number of distinct genera
alive at any given time; that is, those
whose first occurrence predates and
whose last occurrence postdates that
time"[181]
shows a different trend: a fairly swift rise from
542 to 400 million years ago; a slight decline from
400 to 200 million years ago, in which the devastating
Permian–Triassic extinction event is an important factor;
and a swift rise from 200 million years ago to the
Thousands of genera
present.[181]
[edit] The present

Oxygenic photosynthesis accounts for virtually
all of the production of organic matter from non-organic
ingredients. Production is split about evenly between
land and marine plants, and phytoplankton are the
dominant marine producers.[182]
The processes that drive evolution are still
operating. Well-known examples include the changes in
coloration of the peppered moth over the last 200 years
and the more recent appearance of pathogens that are
Phanerozoic biodiversity as shown by the
fossil record
resistant to antibiotics.[183][184] There is even evidence that humans are still evolving, and possibly at an
accelerating rate over the last 40,000 years.[185]
[edit] See also

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[edit] Footnotes
1. ^ Name given as in Butterfield's paper "Bangiomorpha pubescens ..." (2000). A fossil fish,
also from China, has also been named Qingshania. The name of one of these will have to change.
2. ^ Myxozoa were thought to be an exception, but are now thought to be heavily modified
members of the Cnidaria: Jímenez-Guri, E., Philippe, H., Okamura, B. and Holland, P. W. H. (July
2007). "Buddenbrockia is a cnidarian worm". Science 317 (116): 116–118.
doi:10.1126/science.1142024. PMID 17615357.
http://www.sciencemag.org/cgi/content/abstract/317/5834/116. Retrieved 2008-09-03.
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of microbial mats". Proceedings of the Royal Society: Biology 266 (1436): 2375.
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G. et al. (2007). "A whiff of oxygen before the great oxidation event?". Science 317 (5846): 1903–
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237–240. doi:10.1126/science.281.5374.237. PMID 9657713.
http://www.sciencemag.org/cgi/content/full/sci;281/5374/237. Retrieved 2008-09-13.
183.^ Grant, B. S., and Wiseman, L. L. (2002). "Recent History of Melanism in American
Peppered Moths". Journal of Heredity 93 (2): 86–90. doi:10.1093/jhered/93.2.86. ISSN 1465-7333.
PMID 12140267. http://jhered.oxfordjournals.org/cgi/content/abstract/93/2/86. Retrieved 2008-09-11.
184.^ Levin, B. R., Perrot, V. and Walker, N. (March 1, 2000). "Compensatory Mutations,
Antibiotic Resistance and the Population Genetics of Adaptive Evolution in Bacteria". Genetics 154
(3): 985–997. PMID 10757748. PMC 1460977.
http://www.genetics.org/cgi/content/abstract/154/3/985. Retrieved 2008-09-11.
185.^ Hawks, J., Wang, E. T., Cochran, G. M., Harpending, H. C. and Moyzis, R. K. (December
2007). "Recent acceleration of human adaptive evolution". Proceedings of the National Academy of
Sciences 104 (52): 20753–20758. doi:10.1073/pnas.0707650104. PMID 18087044. PMC 2410101.
http://www.pnas.org/content/104/52/20753.full. Retrieved 2008-09-11.
[edit] Further reading
• Cowen, R. (2004). History of Life (4th ed.). Blackwell Publishing Limited. ISBN 978-
1405117562.
• The Ancestor's Tale, A Pilgrimage to the Dawn of Life . Boston: Houghton Mifflin Company.
2004. ISBN 0-618-00583-8.
• Richard Dawkins. (1990). The Selfish Gene. Oxford University Press. ISBN 0192860925.
• Smith, John Maynard; Eörs Szathmáry (1997). The Major Transitions in Evolution.
Oxfordshire: Oxford University Press. ISBN 0-198-50294-X.
[edit] External links

General information
• General information on evolution- Fossil Museum nav.
• Understanding Evolution from University of California, Berkeley
• National Academies Evolution Resources
• Evolution poster- PDF format "tree of life"
• Everything you wanted to know about evolution by New Scientist
• Howstuffworks.com — How Evolution Works
• Synthetic Theory Of Evolution: An Introduction to Modern Evolutionary Concepts and
Theories
History of evolutionary thought
• The Complete Work of Charles Darwin Online
• Understanding Evolution: History, Theory, Evidence, and Implications
Retrieved from "http://en.wikipedia.org/wiki/Evolutionary_history_of_life"
Categories: History by topic | Paleontology | Evolutionary biology

W000
Evolutionary history of plants


A late Silurian sporangium. Green: A spore tetrad. Blue: A spore bearing a trilete mark – the Y-
shaped scar. The spores are about 30-35 μm across
The evolution of plants has resulted in increasing levels of complexity, from the earliest algal mats,
through bryophytes, lycopods, ferns to the complex gymnosperms and angiosperms of today. While the
groups which appeared earlier continue to thrive, especially in the environments in which they evolved, each
new grade of organisation has eventually become more "successful" than its predecessors by most
measures.
Evidence suggests that an algal scum formed on the land 1,200 million years ago, but it was not until
the Ordovician Period, around 450 million years ago, that land plants appeared.[1] These began to diversify
in the late Silurian Period, around 420 million years ago, and the fruits of their diversification are displayed in
remarkable detail in an early Devonian fossil assemblage from the Rhynie chert. This chert preserved early
plants in cellular detail, petrified in volcanic springs. By the middle of the Devonian Period most of the
features recognised in plants today are present, including roots, leaves and secondary wood, and by late
Devonian times seeds had evolved.[2] Late Devonian plants had thereby reached a degree of sophistication
that allowed them to form forests of tall trees. Evolutionary innovation continued after the Devonian period.
Most plant groups were relatively unscathed by the Permo-Triassic extinction event, although the structures
of communities changed. This may have set the scene for the evolution of flowering plants in the Triassic
(~200 million years ago), which exploded in the Cretaceous and Tertiary. The latest major group of plants to
evolve were the grasses, which became important in the mid Tertiary, from around 40 million years ago. The
grasses, as well as many other groups, evolved new mechanisms of metabolism to survive the low CO 2 and
warm, dry conditions of the tropics over the last 10 million years.
Contents
[hide]
• 1 Colonisation of land
• 2 Changing life cycles
• 3 Water transport
• 4 Morphology
• 4.1 Meristems
• 4.1.1 Diversity in meristem architectures
• 4.1.2 Role of the KNOX-family genes
• 4.1.3 Architecture
• 4.2 Leaves
• 4.3 Tree form
• 4.4 Roots
• 4.4.1 Arbuscular mycorrhizae
• 4.5 Seeds
• 4.6 Flowers
• 5 Advances in metabolism
• 5.1 Concentrating carbon
• 5.2 Evolutionary record
• 5.3 When is C4 an advantage?
• 6 Evolutionary trends
• 7 See also
[edit] Colonisation of land
The Devonian period marks the beginning of extensive land colonization by plants, which through
their effects on erosion and sedimentation brought about significant climatic change.
Land plants evolved from chlorophyte algae, perhaps as early as 510 million years ago;[3] their
closest living relatives are the charophytes, specifically Charales. Assuming that the Charales' habit has
changed little since the divergence of lineages, this means that the land plants evolved from a branched,
filamentous, haplontic alga, dwelling in shallow fresh water,[4] perhaps at the edge of seasonally desiccating
pools.[3] Co-operative interactions with fungi may have helped early plants adapt to the stresses of the
terrestrial realm.[5]
Plants were not the first photosynthesisers on land, though: consideration of weathering rates
suggests that organisms were already living on the land 1,200 million years ago,[3] and microbial fossils have
been found in freshwater lake deposits from 1,000 million years ago,[6] but the carbon isotope record
suggests that they were too scarce to impact the atmospheric composition until around 850 million years ago.
[7] These organisms were probably small and simple, forming little more than an "algal scum".[3]
The first evidence of plants on land comes from spores of Mid-Ordovician age (early Llanvirn, ~470
million years ago).[8] [9] These spores, known as cryptospores, were produced either singly (monads), in
pairs (diade) or groups of four (tetrads), and their microstructure resembles that of modern liverwort spores,
suggesting they share an equivalent grade of organisation.[10] It could be that atmospheric 'poisoning'
prevented eukaryotes from colonising the land prior to this,[11] or it could simply have taken a great time for
the necessary complexity to evolve.[12]
Trilete spores similar to those of vascular plants appear soon afterwards, in Upper Ordovician rocks.
[13] Depending exactly when the tetrad splits, each of the four spores may bear a "trilete mark", a Y-shape,
reflecting the points at which each cell squashed up against its neighbours.[8] However, this requires that the
spore walls be sturdy and resistant at an early stage. This resistance is closely associated with having a
desiccation-resistant outer wall—a trait only of use when spores must survive out of water. Indeed, even those
embryophytes that have returned to the water lack a resistant wall, thus don't bear trilete marks.[8] A close
examination of algal spores shows that none have trilete spores, either because their walls are not resistant
enough, or in those rare cases where it is, the spores disperse before they are squashed enough to develop
the mark, or don't fit into a tetrahedral tetrad.[8]
The earliest megafossils of land plants were thalloid organisms, which dwelt in fluvial wetlands and
are found to have covered most of an early Silurian flood plain. They could only survive when the land was
waterlogged.[14]
Once plants had reached the land, there were two approaches to dealing with desiccation. The
bryophytes avoid it or give in to it, restricting their ranges to moist settings, or drying out and putting their
metabolism "on hold" until more water arrives. Tracheophytes resist desiccation. They all bear a waterproof
outer cuticle layer wherever they are exposed to air (as do some bryophytes), to reduce water loss, but—since
a total covering would cut them off from CO2 in the atmosphere—they rapidly evolved stomata, small
openings to allow gas exchange. Tracheophytes also developed vascular tissue to aid in the movement of
water within the organisms (see below), and moved away from a gametophyte dominated life cycle (see
below). Vascular tissue also facilitated upright growth without the support of water and paved the way for the
evolution of larger plants on land.
The establishment of a land-based flora permitted the accumulation of oxygen in the atmosphere as
never before, as the new hordes of land plants pumped it out as a waste product. When this concentration
rose above 13%, it permitted the possibility of wildfire. This is first recorded in the early Silurian fossil record
by charcoalified plant fossils.[15] Apart from a controversial gap in the Late Devonian, charcoal is present
ever since.
Charcoalification is an important taphonomic mode. Wildfire drives off the volatile compounds,
leaving only a shell of pure carbon. This is not a viable food source for herbivores or detritovores, so is prone
to preservation; it is also robust, so can withstand pressure and display exquisite, sometimes sub-cellular,
detail.
[edit] Changing life cycles

Further information: Alternation of generations
Angiosperm life cycle
All multicellular plants have a life cycle comprising two generations or phases. One is termed the
gametophyte, has a single set of chromosomes (denoted 1N), and produces gametes (sperm and eggs). The
other is termed the sporophyte, has paired chromosomes (denoted 2N), and produces spores. The
gametophyte and sporophyte may appear identical – homomorphy – or may be very different – heteromorphy.
The pattern in plant evolution has been a shift from homomorphy to heteromorphy. The algal
ancestors to land plants were almost certainly haplobiontic, being haploid for all their life cycles, with a
unicellular zygote providing the 2N stage. All land plants (i.e. embryophytes) are diplobiontic – that is, both
the haploid and diploid stages are multicellular.[16] Two trends are apparent: bryophytes (liverworts, mosses
and hornworts) have developed the gametophyte, with the sporophyte becoming almost entirely dependent
on it; vascular plants have developed the sporophyte, with the gametophyte being particularly reduced in the
seed plants.
There are two competing theories to explain the appearance of a diplobiontic lifecycle.
The interpolation theory (also known as the antithetic or intercalary theory)[17] holds that the
sporophyte phase was a fundamentally new invention, caused by the mitotic division of a freshly germinated
zygote, continuing until meiosis produces spores. This theory implies that the first sporophytes bore a very
different morphology that the gametophyte they depended on.[17] This seems to fit well with what we know of
the bryophytes, in which a vegetative thalloid gametophyte is parasitised by simple sporophytes, which often
comprise no more than a sporangium on a stalk. Increasing complexity of the ancestrally simple sporophyte,
including the eventual acquisition of photosynthetic cells, would free it from its dependence on a
gametophyte, as we see in some hornworts (Anthoceros), and eventually result in the sporophyte developing
organs and vascular tissue, and becoming the dominant phase, as in the tracheophytes (vascular plants).[16]
This theory may be supported by observations that smaller Cooksonia individuals must have been supported
by a gametophyte generation. The observed appearance of larger axial sizes, with room for photosynthetic
tissue and thus self-sustainability, provides a possible route for the development of a self-sufficient
sporophyte phase.[17]
The alternative hypothesis is termed the transformation theory (or homologous theory). This posits
that the sporophyte appeared suddenly by a delay in the occurrence of meiosis after the zygote germinated.
Since the same genetic material would be employed, the haploid and diploid phases would look the same.
This explains the behaviour of some algae, which produce alternating phases of identical sporophytes and
gametophytes. Subsequent adaption to the desiccating land environment, which makes sexual reproduction
difficult, would result in the simplification of the sexually active gametophyte, and elaboration of the
sporophyte phase to better disperse the waterproof spores.[16] The tissue of sporophytes and gametophytes
preserved in the Rhynie chert is of similar complexity, which is taken to support this hypothesis.[17] [18] [19]
[edit] Water transport

To photosynthesise, plants must absorb CO2 from the atmosphere. However, this comes at a price:
while stomata are open to allow CO 2 to enter, water can evaporate.[20] Water is lost much faster than CO2 is
absorbed, so plants need to replace it, and have developed systems to transport water from the moist soil to
the site of photosynthesis.[20] Early plants sucked water between the walls of their cells, then evolved the
ability to control water loss (and CO2 acquisition) through the use of stomata. Specialised water transport
tissues soon evolved in the form of hydroids, tracheids, then secondary xylem, followed by an endodermis
and ultimately vessels.[20]
The high CO2 levels of Silurian-Devonian times, when plants were first colonising land, meant that
the need for water was relatively low . As CO2 was withdrawn from the atmosphere by plants, more water
was lost in its capture, and more elegant transport mechanisms evolved.[20] As water transport mechanisms,
and waterproof cuticles, evolved, plants could survive without being continually covered by a film of water.
This transition from poikilohydry to homoiohydry opened up new potential for colonisation.[20] Plants were
then faced with a balance, between transporting water as efficiently as possible and preventing transporting
vessels to implode and cavitate.
During the Silurian, CO2 was readily available, so little water needed expending to acquire it. By the
end of the Carboniferous, when CO 2 levels had lowered to something approaching today's, around 17 times
more water was lost per unit of CO2 uptake.[20] However, even in these "easy" early days, water was at a
premium, and had to be transported to parts of the plant from the wet soil to avoid desiccation. This early
water transport took advantage of the cohesion-tension mechanism inherent in water. Water has a tendency
to diffuse to areas that are drier, and this process is accelerated when water can be wicked along a fabric
with small spaces. In small passages, such as that between the plant cell walls (or in tracheids), a column of
water behaves like rubber – when molecules evaporate from one end, they literally pull the molecules behind
them along the channels. Therefore transpiration alone provided the driving force for water transport in early
plants.[20] However, without dedicated transport vessels, the cohesion-tension mechanism cannot transport
water more than about 2 cm, severely limiting the size of the earliest plants.[20] This process demands a
steady supply of water from one end, to maintain the chains; to avoid exhausing it, plants developed a
waterproof cuticle. Early cuticle may not have had pores but did not cover the entire plant surface, so that gas
exchange could continue.[20] However, dehydration at times was inevitable; early plants cope with this by
having a lot of water stored between their cell walls, and when it comes to it sticking out the tough times by
putting life "on hold" until more water is supplied.[20]
A banded tube from the late Silurian/early Devonian. The bands are difficult to see on this specimen,
as an opaque carbonaceous coating conceals much of the tube. Bands are just visible in places on the left
half of the image – click on the image for a larger view. Scale bar: 20 μm
To be free from the constraints of small size and constant moisture that the parenchymatic transport
system inflicted, plants needed a more efficient water transport system. During the early Silurian, they
developed specialized cells, which were lignified (or bore similar chemical compounds)[20] to avoid
implosion; this process coincided with cell death, allowing their innards to be emptied and water to be passed
through them.[20] These wider, dead, empty cells were a million times more conductive than the inter-cell
method, giving the potential for transport over longer distances, and higher CO 2 diffusion rates.
The first macrofossils to bear water-transport tubes in situ are the early Devonian pretracheophytes
Aglaophyton and Horneophyton, which have structures very similar to the hydroids of modern mosses. Plants
continued to innovate new ways of reducing the resistance to flow within their cells, thereby increasing the
efficiency of their water transport. Bands on the walls of tubes, in fact apparent from the early Silurian
onwards,[21] are an early improvisation to aid the easy flow of water.[22] Banded tubes, as well as tubes with
pitted ornamentation on their walls, were lignified[23] and, when they form single celled conduits, are
considered to be tracheids. These, the "next generation" of transport cell design, have a more rigid structure
than hydroids, allowing them to cope with higher levels of water pressure.[20] Tracheids may have a single
evolutionary origin, possibly within the hornworts,[24] uniting all tracheophytes (but they may have evolved
more than once).[20]
Water transport requires regulation, and dynamic control is provided by stomata.[25] By adjusting the
amount of gas exchange, they can restrict the amount of water lost through transpiration. This is an important
role where water supply is not constant, and indeed stomata appear to have evolved before tracheids, being
present in the non-vascular hornworts.[20]
An endodermis probably evolved during the Silu-Devonian, but the first fossil evidence for such a
structure is Carboniferous.[20] This structure in the roots covers the water transport tissue and regulates ion
exchange (and prevents unwanted pathogens etc. from entering the water transport system). The
endodermis can also provide an upwards pressure, forcing water out of the roots when transpiration is not
enough of a driver.
Once plants had evolved this level of controlled water transport, they were truly homoiohydric, able to
extract water from their environment through root-like organs rather than relying on a film of surface moisture,
enabling them to grow to much greater size.[20] As a result of their independence from their surroundings,
they lost their ability to survive desiccation – a costly trait to retain.[20]
During the Devonian, maximum xylem diameter increased with time, with the minimum diameter
remaining pretty constant.[22] By the middle Devonian, the tracheid diameter of some plant lineages[26] had
plateaued.[22] Wider tracheids allow water to be transported faster, but the overall transport rate depends
also on the overall cross-sectional area of the xylem bundle itself.[22] The increase in vascular bundle
thickness further seems to correlate with the width of plant axes, and plant height; it is also closely related to
the appearance of leaves[22] and increased stomatal density, both of which would increase the demand for
water.[20]
While wider tracheids with robust walls make it possible to achieve higher water transport pressures,
this increases the problem of cavitation.[20] Cavitation occurs when a bubble of air forms within a vessel,
breaking the bonds between chains of water molecules and preventing them from pulling more water up with
their cohesive tension. A tracheid, once cavitated, cannot have its embolism removed and return to service
(except in a few advanced angiosperms[ verification needed] which have developed a mechanism of doing
so). Therefore it is well worth plants' while to avoid cavitation occurring. For this reason, pits in tracheid walls
have very small diameters, to prevent air entering and allowing bubbles to nucleate.[20] Freeze-thaw cycles
are a major cause of cavitation.[20] Damage to a tracheid's wall almost inevitably leads to air leaking in and
cavitation, hence the importance of many tracheids working in parallel.[20]
Cavitation is hard to avoid, but once it has occurred plants have a range of mechanisms to contain
the damage.[20] Small pits link adjacent conduits to allow fluid to flow between them, but not air – although
ironically these pits, which prevent the spread of embolisms, are also a major cause of them.[20] These
pitted surfaces further reduce the flow of water through the xylem by as much as 30%.[20] Conifers, by the
Jurassic, developed an ingenious improvement, using valve-like structures to isolate cavitated elements.
These torus-margo structures have a blob floating in the middle of a donut; when one side depressurises the
blob is sucked into the torus and blocks further flow.[20] Other plants simply accept cavitation; for instance,
oaks grow a ring of wide vessels at the start of each spring, none of which survive the winter frosts. Maples
use root pressure each spring to force sap upwards from the roots, squeezing out any air bubbles.
Growing to height also employed another trait of tracheids – the support offered by their lignified
walls. Defunct tracheids were retained to form a strong, woody stem, produced in most instances by a
secondary xylem. However, in early plants, tracheids were too mechanically vulnerable, and retained a
central position, with a layer of tough sclerenchyma on the outer rim of the stems.[20] Even when tracheids
do take a structural role, they are supported by sclerenchymatic tissue.
Tracheids end with walls, which impose a great deal of resistance on flow;[22] vessel members have
perforated end walls, and are arranged in series to operate as if they were one continuous vessel.[22] The
function of end walls, which were the default state in the Devonian, was probably to avoid embolisms. An
embolism is where an air bubble is created in a tracheid. This may happen as a result of freezing, or by
gases dissolving out of solution. Once an embolism is formed, it usually cannot be removed (but see later);
the affected cell cannot pull water up, and is rendered useless.
End walls excluded, the tracheids of prevascular plants were able to operate under the same
hydraulic conductivity as those of the first vascular plant, Cooksonia.[22]
The size of tracheids is limited as they comprise a single cell; this limits their length, which in turn
limits their maximum useful diameter to 80 μm.[20] Conductivity grows with the fourth power of diameter, so
increased diameter has huge rewards; vessel elements, consisting of a number of cells, joined at their ends,
overcame this limit and allowed larger tubes to form, reaching diameters of up to 500 μm, and lengths of up
to 10 m.[20]
Vessels first evolved during the dry, low CO2 periods of the late Permian, in the horsetails, ferns and
Selaginellales independently, and later appeared in the mid Cretaceous in angiosperms and gnetophytes.
[20] Vessels allow the same cross-sectional area of wood to transport around a hundred times more water
than tracheids![20] This allowed plants to fill more of their stems with structural fibres, and also opened a new
niche to vines, which could transport water without being as thick as the tree they grew on.[20] Despite these
advantages, tracheid-based wood is a lot lighter, thus cheaper to make, as vessels need to be much more
reinforced to avoid cavitation.[20]
[edit] Morphology
[edit] Meristems
The meristematic cells give rise to various organs of the plant, and keep the plant growing. The
Shoot Apical Meristem (SAM) gives rise to organs like the leaves and flowers. The cells of the apical
meristems - SAM and RAM (Root Apical Meristem)- divide rapidly and are considered to be indeterminate, in
that they do not possess any defined end fate. In that sense, the meristematic cells are frequently compared
to the stem cells in animals, that have an analogous behavior and function.
[edit] Diversity in meristem architectures

Is the mechanism of being indeterminate conserved in the SAM's of the plant world? The SAM
contains a population of stem cells that also produce the lateral meristems while the stem elongates. It turns
out that the mechanism of regulation of the stem cell number might indeed be evolutionarily conserved. The
CLAVATA gene CLV2 responsible for maintaining the stem cell population in Arabidopsis thaliana is very
closely related to the Maize gene FASCIATED EAR 2(FEA2) also involved in the same function.[27]
Similarly, in Rice, the FON1-FON2 system seems to bear a close relationship with the CLV signaling system
in Arabidopsis thaliana.[28] These studies suggest that the regulation of stem cell number, identity and
differentiation might be an evolutionarily conserved mechanism in monocots, if not in angiosperms. Rice also
contains another genetic system distinct from FON1-FON2, that is involved in regulating stem cell number.
[28] This example underlines the innovation that goes about in the living world all the time.
[edit] Role of the KNOX-family genes

Note the long spur of the above flower. Spurs attract pollinators and confer pollinator specificity.
(Flower:Linaria dalmatica)
Complex leaves of C. hirsuta are a result of KNOX gene expression

Genetic screens have identified genes belonging to the KNOX family in this function. These genes
essentially maintain the stem cells in an undifferentiated state. The KNOX family has undergone quite a bit of
evolutionary diversification, while keeping the overall mechanism more or less similar. Members of the KNOX
family have been found in plants as diverse as Arabidopsis thaliana, rice, barley and tomato. KNOX-like
genes are also present in some algae, mosses, ferns and gymnosperms. Misexpression of these genes
leads to formation of interesting morphological features. For example, among members of Antirrhinae, only
the species of genus Antirrhinum lack a structure called spur in the floral region. A spur is considered an
evolutionary innovation because it defines pollinator specificity and attraction. Researchers carried out
transposon mutagenesis in Antirrhinum majus, and saw that some insertions led to formation of spurs that
were very similar to the other members of Antirrhinae[29], indicating that the loss of spur in wild Antirrhinum
majus populations could probably be an evolutionary innovation.
The KNOX family has also been implicated in leaf shape evolution (See below for a more detailed
discussion). One study looked at the pattern of KNOX gene expression in A. thaliana, that has simple leaves
and Cardamine hirsuta, a plant having complex leaves. In A. thaliana, the KNOX genes are completely
turned off in leaves, but in C.hirsuta, the expression continued, generating complex leaves.[30] Also, it has
been proposed that the mechanism of KNOX gene action is conserved across all vascular plants, because
there is a tight correlation between KNOX expression and a complex leaf morphology.[31]
[edit] Architecture
The meristem architectures do differ between angiosperms, gymnosperms and pteridophytes. The
gymnosperm vegetative meristem lacks organization into distinct tunica and corpus layers. They possess
large cells called Central Mother Cells in the meristem. In angiosperms, the outermost layer of cells divides
anticlinally to generate the new cells, while in gymnosperms, the plane of division in the meristem differs for
different cells. However, the apical cells do contain organelles like large vacuoles and starch grains, like the
angiosperm meristematic cells.
Pteridophytes, like fern, on the other hand, do not possess a multicellular apical meristem. They
possess a tetrahedral apical cell, which goes on to form the plant body. Any somatic mutation in this cell can
lead to hereditary transmission of that mutation.[32] The earliest meristem-like organization is seen in an
algal organism from group Charales that has a single dividing cell at the tip, much like the pteridophytes, yet
simpler. One can thus see a clear pattern in evolution of the meristematic tissue, from pteridophytes to
angiosperms. Pteridophytes, with a single meristematic cell; gymnosperms with a multicellular, but less
defined organization and finally, angiosperms, with the highest degree of organization. The genetic
innovations that contributed to this evolution are yet not clearly known.
[edit] Leaves
The lycopod Isoetes bears microphylls with a single vascular trace.
The branching pattern of megaphyll veins may belie their origin as webbed, dichotomising branches.
Leaves today are, in almost all instances, an adaptation to increase the amount of sunlight that can
be captured for photosynthesis. Leaves certainly evolved more than once, and probably originated as spiny
outgrowths to protect early plants from herbivory.
The rhyniophytes of the Rhynie chert comprised nothing more than slender, unornamented axes.
The early to middle Devonian trimerophytes, therefore, are the first evidence we have of anything that could
be considered leafy. This group of vascular plants are recognisable by their masses of terminal sporangia,
which adorn the ends of axes which may bifurcate or trifurcate.[25] Some organisms, such as Psilophyton,
bore enations. These are small, spiny outgrowths of the stem, lacking their own vascular supply.
Around the same time, the zosterophyllophytes were becoming important. This group is recognisable
by their kidney-shaped sporangia, which grew on short lateral branches close to the main axes. They
sometimes branched in a distinctive H-shape.[25] The majority of this group bore pronounced spines on their
axes. However, none of these had a vascular trace, and the first evidence of vascularised enations occurs in
the Rhynie genus Asteroxylon. The spines of Asteroxylon had a primitive vasuclar supply – at the very least,
leaf traces could be seen departing from the central protostele towards each individual "leaf". A fossil known
as Baragwanathia appears in the fossil record slightly earlier, in the late Silurian.[33] In this organism, these
leaf traces continue into the leaf to form their mid-vein.[34] One theory, the "enation theory", holds that the
leaves developed by outgrowths of the protostele connecting with existing enations, but it is also possible
that microphylls evolved by a branching axis forming "webbing".[25]
Asteroxylon[35] and Baragwanathia are widely regarded as primitive lycopods.[25] The lycopods are
still extant today, familiar as the quillwort Isoetes and the club mosses. Lycopods bear distinctive microphylls
– leaves with a single vascular trace. Microphylls could grow to some size – the Lepidodendrales boasted
microphylls over a meter in length – but almost all just bear the one vascular bundle. (An exception is the
branching Selaginella).
The more familiar leaves, megaphylls, are thought to have separate origins – indeed, they appeared
four times independently, in the ferns, horsetails, progymnosperms, and seed plants.[36] They appear to
have originated from dichotomising branches, which first overlapped (or "overtopped") one another, and
eventually developed "webbing" and evolved into gradually more leaf-like structures.[34] So megaphylls, by
this "teleome theory", are composed of a group of webbed branches[34] – hence the "leaf gap" left where the
leaf's vascular bundle leaves that of the main branch resembles two axes splitting.[34] In each of the four
groups to evolve megaphylls, their leaves first evolved during the late Devonian to early Carboniferous,
diversifying rapidly until the designs settled down in the mid Carboniferous.[36]
The cessation of further diversification can be attributed to developmental constraints,[36] but why
did it take so long for leaves to evolve in the first place? Plants had been on the land for at least 50 million
years before megaphylls became significant. However, small, rare mesophylls are known from the early
Devonian genus Eophyllophyton – so development could not have been a barrier to their appearance.[37]
The best explanation so far incorporates observations that atmospheric CO 2 was declining rapidly during this
time – falling by around 90% during the Devonian.[38] This corresponded with an increase in stomatal density
by 100 times. Stomata allow water to evaporate from leaves, which causes them to curve. It appears that the
low stomatal density in the early Devonian meant that evaporation was limited, and leaves would overheat if
they grew to any size. The stomatal density could not increase, as the primitive steles and limited root
systems would not be able to supply water quickly enough to match the rate of transpiration.[39]
Clearly, leaves are not always beneficial, as illustrated by the frequent occurrence of secondary loss
of leaves, famously exemplified by cacti and the "whisk fern" Psilotum.
Secondary evolution can also disguise the true evolutionary origin of some leaves. Some genera of
ferns display complex leaves which are attached to the pseudostele by an outgrowth of the vascular bundle,
leaving no leaf gap.[34] Further, horsetail (Equisetum) leaves bear only a single vein, and appear for all the
world to be microphyllous; however, in the light of the fossil record and molecular evidence, we conclude that
their forbears bore leaves with complex venation, and the current state is a result of secondary simplification.
[40]
Deciduous trees deal with another disadvantage to having leaves. The popular belief that plants shed
their leaves when the days get too short is misguided; evergreens prospered in the Arctic circle during the
most recent greenhouse earth.[41] The generally accepted reason for shedding leaves during winter is to
cope with the weather – the force of wind and weight of snow are much more comfortably weathered without
leaves to increase surface area. Seasonal leaf loss has evolved independently several times and is exhibited
in the ginkgoales, pinophyta and angiosperms.[42] Leaf loss may also have arisen as a response to pressure
from insects; it may have been less costly to lose leaves entirely during the winter or dry season than to
continue investing resources in their repair.[43]
[edit] Tree form
The trunk of early tree fern Psaronius, showing internal structure. The top of the plant would have
been to the left of the image
External mold of Lepidodendron trunk showing leaf scars from the Upper Carboniferous of Ohio
The early Devonian landscape was devoid of vegetation taller than waist height. Without the
evolution of a robust vascular system, taller heights could not be attained. There was, however, a constant
evolutionary pressure to attain greater height. The most obvious advantage is the harvesting of more sunlight
for photosynthesis – by overshadowing competitors – but a further advantage is present in spore distribution,
as spores (and, later, seeds) can be blown greater distances if they start higher. This may be demonstrated
by Prototaxites, thought to be a late Silurian fungus reaching eight metres in height.[44]
To attain arborescence, early plants had to develop woody tissue that provided support and water
transport. To understand wood, we must know a little of vascular behaviour. The stele of plants undergoing
"secondary growth" is surrounded by the vascular cambium, a ring of cells which produces more xylem (on
the inside) and phloem (on the outside). Since xylem cells comprise dead, lignified tissue, subsequent rings
of xylem are added to those already present, forming wood.
The first plants to develop this secondary growth, and a woody habit, were apparently the ferns, and
as early as the middle Devonian one species, Wattieza, had already reached heights of 8 m and a tree-like
habit.[45]
Other clades did not take long to develop a tree-like stature; the late Devonian Archaeopteris, a
precursor to gymnosperms which evolved from the trimerophytes,[46] reached 30 m in height. These
progymnosperms were the first plants to develop true wood, grown from a bifacial cambium, of which the first
appearance is in the mid Devonian Rellimia.[47] True wood is only thought to have evolved once, giving rise
to the concept of a "lignophyte" clade.
These Archaeopteris forests were soon supplemented by lycopods, in the form of lepidodendrales,
which topped 50m in height and 2m across at the base. These lycopods rose to dominate late Devonian and
Carboniferous coal deposits.[48] Lepidodendrales differ from modern trees in exhibiting determinate growth:
after building up a reserve of nutrients at a low height, the plants would "bolt" to a genetically determined
height, branch at that level, spread their spores and die.[49] They consisted of "cheap" wood to allow their
rapid growth, with at least half of their stems comprising a pith-filled cavity.[25] Their wood was also
generated by a unifacial vascular cambium – it did not produce new phloem, meaning that the trunks could
not grow wider over time.[verification needed]
The horsetail Calamites was next on the scene, appearing in the Carboniferous. Unlike the modern
horsetail Equisetum, Calamites had a unifacial vascular cambium, allowing them to develop wood and grow
to heights in excess of 10 m. They also branched multiple times.
While the form of early trees was similar to that of today's, the groups containing all modern trees
had yet to evolve.
The dominant groups today are the gymnosperms, which include the coniferous trees, and the
angiosperms, which contain all fruiting and flowering trees. It was long thought that the angiosperms arose
from within the gymnosperms, but recent molecular evidence suggests that their living representatives form
two distinct groups.[50][51][52] It must be noted that the
molecular data has yet to be fully reconciled with
morphological data,[53][54][55] but it is becoming
accepted that the morphological support for paraphyly is
not especially strong.[56] This would lead to the
conclusion that both groups arose from within the
pteridosperms, probably as early as the Permian.[56]
The angiosperms and their ancestors played a
very small role until they diversified during the
Cretaceous. They started out as small, damp-loving
organisms in the understory, and have been diversifying
ever since the mid[verification needed]-Cretaceous, to
become the dominant member of non-boreal forests
today.
[edit] Roots
Roots are important to plants for two main
reasons: Firstly, they provide anchorage to the substrate;
more importantly, they provide a source of water and
nutrients from the soil. Roots allowed plants to grow taller
and faster.
The roots (bottom image) of
lepidodendrales are thought to be functionally
equivalent to the stems (top), as the similar
appearance of "leaf scars" and "root scars" on
these specimens from different species
demonstrates.
The onset of roots also had effects on a global scale. By disturbing the soil, and promoting its
acidification (by taking up nutrients such as nitrate and phosphate[ verification needed]), they enabled it to
weather more deeply, promoting the draw-down of CO 2[57] with huge implications for climate.[58] These
effects may have been so profound they led to a mass extinction.[59]
But how and when did roots evolve in the first place? While there are traces of root-like impressions
in fossil soils in the late Silurian,[60] body fossils show the earliest plants to be devoid of roots. Many had
tendrils which sprawled along or beneath the ground, with upright axes or thalli dotted here and there, and
some even had non-photosynthetic subterranean branches which lacked stomata. The distinction between
root and specialised branch is developmental; true roots follow a different developmental trajectory to stems.
Further, roots differ in their branching pattern, and in possession of a root cap.[3] So while Silu-Devonian
plants such as Rhynia and Horneophyton possessed the physiological equivalent of roots, roots – defined as
organs differentiated from stems – did not arrive until later.[3] Unfortunately, roots are rarely preserved in the
fossil record, and our understanding of their evolutionary origin is sparse.[3]
Rhizoids – small structures performing the same role as roots, usually a cell in diameter – probably
evolved very early, perhaps even before plants colonised the land; they are recognised in the Characeae, an
algal sister group to land plants.[3] That said, rhizoids probably evolved more than once; the rhizines of
lichens, for example, perform a similar role. Even some animals (Lamellibrachia) have root-like structures![3]
More advanced structures are common in the Rhynie chert, and many other fossils of comparable
early Devonian age bear structures that look like, and acted like, roots.[3] The rhyniophytes bore fine
rhizoids, and the trimerophytes and herbaceous lycopods of the chert bore root-like structure penetrating a
few centimetres into the soil.[61] However, none of these fossils display all the features borne by modern
roots.[3] Roots and root-like structures became increasingly more common and deeper penetrating during
the Devonian period, with lycopod trees forming roots around 20 cm long during the Eifelian and Givetian.
These were joined by progymnosperms, which rooted up to about a metre deep, during the ensuing Frasnian
stage.[61] True gymnosperms and zygopterid ferns also formed shallow rooting systems during the
Famennian period.[61]
The rhizomorphs of the lycopods provide a slightly different approach to rooting. They were
equivalent to stems, with organs equivalent to leaves performing the role of rootlets.[3] A similar construction
is observed in the extant lycopod Isoetes, and this appears to be evidence that roots evolved independently
at least twice, in the lycophytes and other plants.[3]
A vascular system is indispensable to a rooted plants, as non-photosynthesising roots need a supply
of sugars, and a vascular system is required to transport water and nutrients from the roots to the rest of the
plant.[4] These plants are little more advanced than their Silurian forbears, without a dedicated root system;
however, the flat-lying axes can be clearly seen to have growths similar to the rhizoids of bryophytes today.
[62]
By the mid-to-late Devonian, most groups of plants had independently developed a rooting system of
some nature.[62] As roots became larger, they could support larger trees, and the soil was weathered to a
greater depth.[59] This deeper weathering had effects not only on the aforementioned drawdown of CO 2, but
also opened up new habitats for colonisation by fungi and animals.[61]
Roots today have developed to the physical limits. They penetrate many[ quantify] metres of soil to
tap the water table.[verification needed] The narrowest roots are a mere 40 μm in diameter, and could not
physically transport water if they were any narrower.[3] The earliest fossil roots recovered, by contrast,
narrowed from 3 mm to under 700 μm in diameter; of course, taphonomy is the ultimate control of what
thickness we can see.[3]
[edit] Arbuscular mycorrhizae
The efficiency of many plants' roots is increased via a symbiotic relationship with a fungal partner.
The most common are arbuscular mycorrhizae (AM), literally "tree-like fungal roots". These comprise fungi
which invade some root cells, filling the cell membrane with their hyphae. They feed on the plant's sugars,
but return nutrients generated or extracted from the soil (especially phosphate), which the plant would
otherwise have no access to.
This symbiosis appears to have evolved early in plant history. AM are found in all plant groups, and
80% of extant vascular plants,[63] suggesting an early ancestry; a "plant"-fungus symbiosis may even have
been the step that enabled them to colonise the land,[64] and indeed AM are abundant in the Rhynie chert;
[65] the association occurred even before there were true roots to colonise, and some have suggested that
roots evolved to provide a more comfortable habitat for mycorrhizal fungi.[66]
[edit] Seeds
The fossil seed Trigonocarpus

Early land plants reproduced in the fashion of ferns: spores germinated into small gametophytes,
which produced sperm. These would swim across moist soils to find the female organs (archegonia) on the
same or another gametophyte, where they would fuse with an ovule to produce an embryo, which would
germinate into a sporophyte.[61]
This mode of reproduction restricted early plants to damp environments, moist enough that the
sperm could swim to their destination. Therefore, early land plants were constrained to the lowlands, near
shores and streams. The development of heterospory freed them from this constraint.
Heterosporic organisms, as their name suggests, bear spores of two sizes – microspores and
megaspores. These would germinate to form microgametophytes and megagametophytes, respectively. This
system paved the way for seeds: taken to the extreme, the megasporangia could bear only a single
megaspore tetrad, and to complete the transition to true seeds, three of the megaspores in the original tetrad
could be aborted, leaving one megaspore per megasporangium.
The transition to seeds continued with this megaspore being "boxed in" to its sporangium while it
germinates. Then, the megagametophyte is contained within a waterproof integument, which forms the bulk
of the seed. The microgametophyte – a pollen grain which has germinated from a microspore – is employed
for dispersal, only releasing its desiccation-prone sperm when it reaches a receptive megagametophyte.[25]
Lycopods go a fair way down the path to seeds without ever crossing the threshold. Fossil lycopod
megaspores reaching 1 cm in diameter, and surrounded by vegetative tissue, are known – these even
germinate into a megagametophyte in situ. However, they fall short of being seeds, since the nucellus, an
inner spore-covering layer, does not completely enclose the spore. A very small slit remains, meaning that
the seed is still exposed to the atmosphere. This has two consequences – firstly, it means it is not fully
resistant to desiccation, and secondly, sperm do not have to "burrow" to access the archegonia of the
megaspore.[25]
The first "spermatophytes" (literally:seed plants) – that is, the first plants to bear true seeds – are
called pteridosperms: literally, "seed ferns", so called because their foliage consisted of fern-like fronds,
although they were not closely related to ferns. The oldest fossil evidence of seed plants is of Late Devonian
age and they appear to have evolved out of an earlier group known as the progymnosperms. These early
seed plants ranged from trees to small, rambling shrubs; like most early progymnosperms, they were woody
plants with fern-like foliage. They all bore ovules, but no cones, fruit or similar. While it is difficult to track the
early evolution of seeds, we can trace the lineage of the seed ferns from the simple trimerophytes through
homosporous Aneurophytes.[25]
This seed model is shared by basically all gymnosperms (literally: "naked seeds"), most of which
encase their seeds in a woody or fleshy (the yew, for example) cone, but none of which fully enclose their
seeds. The angiosperms ("vessel seeds") are the only group to fully enclose the seed, in a carpel.
Fully enclosed seeds opened up a new pathway for plants to follow: that of seed dormancy. The
embryo, completely isolated from the external atmosphere and hence protected from desiccation, could
survive some years of drought before germinating. Gymnosperm seeds from the late Carboniferous have
been found to contain embryos, suggesting a lengthy gap between fertilisation and germination.[67] This
period is associated with the entry into a greenhouse earth period, with an associated increase in aridity. This
suggests that dormancy arose as a response to drier climatic conditions, where it became advantageous to
wait for a moist period before germinating.[67] This evolutionary breakthrough appears to have opened a
floodgate: previously inhospitable areas, such as dry mountain slopes, could now be tolerated, as were soon
covered by trees.[67]
Seeds offered further advantages to their bearers: they increased the success rate of fertilised
gametophytes, and because a nutrient store could be "packaged" in with the embryo, the seeds could
germinate rapidly in inhospitable environments, reaching a size where it could fend for itself more quickly.[61]
For example, without an endosperm, seedlings growing in arid environments would not have the reserves to
grow roots deep enough to reach the water table before they expired.[61] Likewise, seeds germinating in a
gloomy understory require an additional reserve of energy to quickly grow high enough to capture sufficient
light for self-sustenance.[61] A combination of these advantages gave seed plants the ecological edge over
the previously dominant genus Archaeopteris, this increasing the biodiversity of early forests.[61]
[edit] Flowers
The pollen bearing organs of the early "flower" Crossotheca

The evolution of syncarps.
a: sporangia borne at tips of leaf
b: Leaf curls up to protect sporangia
c: leaf curls to form enclosed roll
d: grouping of three rolls into a syncarp
Flowers are modified leaves possessed only by the group known as the angiosperms, which are
relatively late to appear in the fossil record. Colourful and/or pungent structures surround the cones of plants
such as cycads and gnetales, making a strict definition of the term "flower" elusive.[55]
The flowering plants have long been assumed to have evolved from within the gymnosperms;
according to the traditional morphological view, they are closely allied to the gnetales. However, as noted
above, recent molecular evidence is at odds to this hypothesis,[51][52] and further suggests that gnetales are
more closely related to some gymnosperm groups than angiosperms,[50] and that extant gymnosperms form
a distinct clade to the angiosperms,[51][52][50] the two clades diverging some 300 million years ago.[68]
Phylogeny of anthophytes and gymnosperms, from [69]
Cycads
Ginkgo
Conifers
Anthophytes
Bennettitales
Gnetales
Angiosperms
Angiosperms
Gymnosperms
Cycads
Bennettitales
Ginkgo
Conifers
Gnetales
Traditional view Modern view
Further information: Gnetophyta#Classification
The relationship of stem groups to the angiosperms is of utmost importance in determining the
evolution of flowers; stem groups provide an insight into the state of earlier "forks" on the path to the current
state. If we identify an unrelated group as a stem group, then we will gain an incorrect image of the lineages'
history. The traditional view that flowers arose by modification of a structure similar to that of the gnetales, for
example, no longer bears weight in the light of the molecular data.
Convergence increases our chances of misidentifying stem groups. Since the protection of the
megagametophyte is evolutionarily desirable, it would be unsurprising if many separate groups stumbled
upon protective encasements independently. Distinguishing ancestry in such a situation, especially where we
usually only have fossils to go on, is tricky – to say the least.[ clarification needed]
In flowers, this protection is offered by the carpel, an organ believed to represent an adapted leaf,
recruited into a protective role, shielding the ovules. These ovules are further protected by a double-walled
integument.
Penetration of these protective layers needs something more that a free-floating microgametophyte.
Angiosperms have pollen grains comprising just three cells. One cell is responsible for drilling down through
the integuments, and creating a conduit for the two sperm cells to flow down. The megagametophyte has just
seven cells; of these, one fuses with a sperm cell, forming the nucleus of the egg itself, and another other
joins with the other sperm, and dedicates itself to forming a nutrient-rich endosperm. The other cells take
auxiliary roles.[clarification needed] This process of "double fertilisation" is unique and common to all
angiosperms.
The inflorescences of the Bennettitales are strikingly similar to flowers

In the fossil record, there are three intriguing groups which bore flower-like structures. The first is the
Permian pteridosperm Glossopteris, which already bore recurved leaves resembling carpels. The Triassic
Caytonia is more flower-like still, with enclosed ovules – but only a single integument. Further, details of their
pollen and stamens set them apart from true flowering plants.
The Bennettitales bore remarkably flower-like organs, protected by whorls of bracts which may have
played a similar role to the petals and sepals of true flowers; however, these flower-like structures evolved
independently, as the Bennettitales are more closely related to cycads and ginkgos than to the angiosperms.
[69]
However, no true flowers are found in any groups save those extant today. Most morphological and
molecular analyses place Amborella, the nymphaeales and Austrobaileyaceae in a basal clade dubbed
"ANA". This clade appear to have diverged in the early Cretaceous, around 130 million years ago – around
the same time as the earliest fossil angiosperm,[70][71] and just after the first angiosperm-like pollen, 136
million years ago.[56] The magnoliids diverged soon after, and a rapid radiation had produced eudicots and
monocots by 125 million years ago.[56] By the end of the Cretaceous 65.5 million years ago, over 50% of
today's angiosperm orders had evolved, and the clade accounted for 70% of global species.[72] It was
around this time that flowering trees became dominant over conifers [73]
The features of the basal "ANA" groups suggest that angiosperms originated in dark, damp,
frequently disturbed areas.[74] It appears that the angiosperms remained constrained to such habitats
throughout the Cretaceous – occupying the niche of small herbs early in the successional series.[72] This
may have restricted their initial significance, but given them the flexibility that accounted for the rapidity of
their later diversifications in other habitats.[74]
[edit] Advances in metabolism
The C4 carbon concentrating mechanism
The most recent major innovation by the plants is the development of the C4 metabolic pathway.
Photosynthesis is not quite as simple as adding water to CO2 to produce sugars and oxygen. A
complex chemical pathway is involved, facilitated along the way by a range of enzymes and co-enzymes.
The enzyme RuBisCO is responsible for "fixing" CO2 – that is, it attaches it to a carbon-based molecule to
form a sugar, which can be used by the plant, releasing an oxygen molecule along the way. However, the
enzyme is notoriously inefficient, and just as effectively will also fix oxygen instead of CO 2 in a process called
photorespiration. This is energetically costly as the plant has to use energy to turn the products of
photorepsiration back into a form that can react with CO 2.
[edit] Concentrating carbon

To work around this inefficiency, C4 plants evolved carbon concentrating mechanisms. These work
by increasing the concentration of CO 2 around RuBisCO, thereby increasing the amount of photosynthesis
and decreasing photorespiration. The process of concentrating CO 2 around RuBisCO requires more energy
than allowing gases to diffuse, but under certain conditions – i.e. warm temperatures (>25°C), low CO 2
concentrations, or high oxygen concentrations – pays off in terms of the decreased loss of sugars through
photorespiration.
One, C4 metabolism, employs a so-called Kranz anatomy. This transports CO2 through an outer
mesophyll layer, via a range of organic molecules, to the central bundle sheath cells, where the CO 2 is
released. In this way, CO2 is concentrated near the site of RuBisCO operation. Because RuBisCO is
operating in an environment with much more CO 2 than it otherwise would be, it performs more efficiently.
A second method, CAM photosynthesis, temporally separates photosynthesis from the action of
RuBisCO. RuBisCO only operates during the day, when stomata are sealed and CO 2 is provided by the
breakdown of the chemical malate. More CO2 is then harvested from the atmosphere when stomata open,
during the cool, moist nights, reducing water loss.
[edit] Evolutionary record

These two pathways, with the same effect on RuBisCO, evolved a number of times independently –
indeed, C4 alone arose in 18 different plant families. The C4 construction is most famously used by a subset
of grasses, while CAM is employed by many succulents and cacti. The trait appears to have emerged during
the Oligocene, around 25 to 32 million years ago;[75] however, they did not become ecologically significant
until the Miocene, -1 million years ago.[76] Remarkably, some charcoalified fossils preserve tissue organised
into the Kranz anatomy, with intact bundle sheath cells,[77] allowing the presence C4 metabolism to be
identified without doubt at this time. In deducing their distribution and significance, we resort to the use of
isotopic markers. C3 plants preferentially use the lighter of two isotopes of carbon in the atmosphere, 12C,
which is more readily involved in the chemical pathways involved in its fixation. Because C 4 metabolism
involves a further chemical step, this effect is accentuated. Plant material can be analysed to deduce the
ratio of the heavier 13C to 12C. This ratio is denoted δ13C. C3 plants are on average around 14‰ (parts per
thousand) lighter than the atmospheric ratio, while C 4 plants are about 28‰ lighter. The δ13C of CAM plants
depends on the percentage of carbon fixed at night relative to what is fixed in the day, being closer to C 3
plants if they fix most carbon in the day and closer to C 4 plants if they fix all their carbon at night.[78]
It's troublesome procuring original fossil material in sufficient quantity to analyse the grass itself, but
fortunately we have a good proxy: horses. Horses were globally widespread in the period of interest, and
browsed almost exclusively on grasses. There's an old phrase in isotope palæontology, "you are what you
eat (plus a little bit)" – this refers to the fact that organisms reflect the isotopic composition of whatever they
eat, plus a small adjustment factor. There is a good record of horse teeth throughout the globe, and their
δ13C has been measured. The record shows a sharp negative inflection around -1 million years ago, during
the Messinian, and this is interpreted as the rise of C 4 plants on a global scale.[76]
[edit] When is C4 an advantage?

While C4 enhances the efficiency of RuBisCO, the concentration of carbon is highly energy intensive.
This means that C4 plants only have an advantage over C3 organisms in certain conditions: namely, high
temperatures and low rainfall. C4 plants also need high levels of sunlight to thrive.[79] Models suggest that
without wildfires removing shade-casting trees and shrubs, there would be no space for C 4 plants.[80] But
wildfires have occurred for 400 million years – why did C 4 take so long to arise, and then appear
independently so many times? The Carboniferous period (~300 million years ago) had notoriously high
oxygen levels – almost enough to allow spontaneous combustion[81] – and very low CO2, but there is no C4
isotopic signature to be found. And there doesn't seem to be a sudden trigger for the Miocene rise.
During the Micoene, the atmosphere and climate was relatively stable. If anything, CO 2 increased
gradually from 14 to 9 million years ago before settling down to concentrations similar to the Holocene.[82]
This suggests that it did not have a key role in invoking C 4 evolution.[75] Grasses themselves (the group
which would give rise to the most occurrences of C4) had probably been around for 60 million years or more,
so had had plenty of time to evolve C4,[83][84] which in any case is present in a diverse range of groups and
thus evolved independently. There is a strong signal of climate change in South Asia;[75] increasing aridity –
hence increasing fire frequency and intensity – may have led to an increase in the importance of grasslands.
[85] However, this is difficult to reconcile with the North American record.[75] It is possible that the signal is
entirely biological, forced by the fire- (and elephant?)-[86] driven acceleration of grass evolution – which, both
by increasing weathering and incorporating more carbon into sediments, reduced atmospheric CO 2 levels.
[86] Finally, there is evidence that the onset of C4 from 9 to 7 million years ago is a biased signal, which only
holds true for North America, from where most samples originate; emerging evidence suggests that
grasslands evolved to a dominant state at least 15Ma earlier in South America.
[edit] Evolutionary trends

The process of evolution works slightly differently in plants than animals. Differences in plant
physiology and reproduction mean that while the same evolutionary principles of natural selection apply, the
finer nuances of their effect are radically different.
One major difference is the ability of plants to reproduce clonally, and the totipotent nature of their
cells, allowing them to reproduce asexually much more easily than most animals. They are also capable of
polyploidy – where more than two chromosome sets are inherited from parents. This allows relatively fast
bursts of evolution to occur. The long periods of dormancy that seed plants can employ also makes them less
vulnerable to extinction, as they can "sit out" the tough periods and wait until more clement times to leap back
to life.
The effect of these differences is most profoundly seen during extinction events. These events, which
wiped out between 6 and 62% of terrestrial animal families, had "negligible" effect on plant families.[87]
However, the ecosystem structure is significantly rearranged, with the abundances and distributions of
different groups of plants changing profoundly.[87] These effects are perhaps due to the higher diversity
within families, as extinction – which was common at the species level – was very selective. For example,
wind-pollinated species survived better than insect-pollinated taxa, and specialised species generally lost
out.[87] In general, the surviving taxa were rare before the extinction, suggesting that they were generalists
who were poor competitors when times were easy, but prospered when specialised groups went extinct and
left ecological niches vacant.[87]
[edit] See also

Plants
• Plant
• Flora
General evolution
• Evolutionary history of life
• Timeline of plant evolution
Study of plants
• Paleobotany
• Plant evolutionary developmental biology
• Cryptospores
Plant interactions
• Evolution of herbivory
[edit] References
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3. ^ a b c d e f g h i j k l m n o Raven, J.A.; Edwards, D. (2001). "Roots: evolutionary origins and
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http://jxb.oxfordjournals.org/cgi/content/full/52/suppl_1/381.
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A cladistic study. Smithsonian Institution Press, Washington & London . Washington: Smithsonian
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Terrestrial Communities in Lacustrine Phosphate One Billion Years Ago". In Smith, Martin R.;
O'Brien, Lorna J.; Caron, Jean-Bernard. International Conference on the Cambrian Explosion
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[hide]v · d · e Basic topics in evolutionary biology
Processes of
Population genetic
Evolutionary
developmental
concepts

Modes of

Retrieved from "http://en.wikipedia.org/wiki/Evolutionary_history_of_plants"
Categories: Plants | Prehistoric plants | Evolution by taxon

W000
Lemur evolutionary history


Mouse lemurs, the smallest primates in the world, evolved in isolation along with other lemurs on the
island of Madagascar.
The evolutionary history of lemurs occurred in isolation from other primates on the island of
Madagascar for at least 40 million years. Lemurs are prosimian primates belonging to the suborder
Strepsirrhini, which branched off from other primates less than 63 mya (million years ago). They share some
traits with the most basal primates, and thus are often confused as being ancestral to modern monkeys,
apes, and humans. Instead, they merely resemble ancestral primates.
Lemurs are thought to have evolved during the Eocene or earlier, sharing a closest common
ancestor with lorisiforms. Fossils from Africa and tests of nuclear DNA suggest that lemurs made their way to
Madagascar between 40 and 52 mya. Molecular tests offer an alternative date range of 62 to 65 mya. An
ancestral lemur population is thought to have inadvertently rafted to the island on a floating mat of vegetation,
although hypotheses for land bridges and island hopping have also been proposed. The timing and number
of hypothesized colonizations has traditionally hinged on the phylogenetic affinities of the Aye-aye, the most
basal member of the lemur clade.
Having undergone their own independent evolution on Madagascar, lemurs have diversified to fill
many niches normally filled by other types of mammals. They include the smallest primates in the world, and
once included some of the largest. Since the arrival of humans approximately 2,000 years ago, they are now
restricted to 10% of the island, or approximately 60,000 square kilometres (23,000 sq mi), and many are
facing extinction. For this reason, researchers have been trying to identify and assess every species. Over
the last 10 to 20 years, there has been a steep increase in the number of recognized lemur species and
subspecies, both through the discovery of new species and the elevation of existing subspecies to full
species status. Currently there are approximately 100 or more recognized species or subspecies of living
lemur, which are divided into five families and 15 genera. If the extinct subfossil lemurs are included, an
additional three families, eight genera, and 17 species would be included. The recent rise in species numbers
is due to both improved genetic analysis and a push in conservation to encourage the protection of isolated
and distinct lemur populations. Not everyone in the scientific community supports these taxonomic changes,
with some preferring instead an estimate of 50 living species.
Contents
[hide]
• 1 Evolutionary history
• 1.1 Colonization of Madagascar
• 1.2 Diversification
• 2 Distribution and diversity
• 3 Taxonomic and phylogenetic classification
• 3.1 Suprageneric classification
• 3.2 Genus-level classification
• 3.3 Species-level classification
• 4 Notes
• 5 References
[edit] Evolutionary history

Lemurs are prosimian primates belonging to the suborder Strepsirrhini. Like other strepsirrhine
primates, such as lorises, pottos, and galagos, they share ancestral traits with early primates. In this regard,
lemurs are popularly confused with ancestral primates; however, lemurs did not give rise to monkeys and
apes, but evolved independently on Madagascar.[1]
Fossil evidence for the evolution of the toothcomb, a trait shared by lemurs with their closest
relatives, the lorises, weighs heavily on the evolutionary history of strepsirrhines and the lemur colonization
of Madagascar.
Primates first evolved sometime between the Middle Cretaceous and the early Paleocene periods on
either the supercontinent of Laurasia or in Africa.[2] According to molecular clock studies, the last common
ancestor of all primates dates to around 79.6 mya,[3] although the earliest known fossil primates are only 54–
55 mya old.[4] The closest relatives of primates are the extinct plesiadapiforms, the modern colugos
(commonly and inaccurately named "flying lemurs"), and treeshrews.[3] Some of the earliest known true
primates are represented by the fossil groups Omomyidae, Eosimiidae, and Adapiformes.
The relationship between known fossil primate families remains unclear. A conservative estimate for
the divergence of haplorrhines (tarsiers, monkeys, apes, and humans) and strepsirrhines is 58 to 63 mya,[5]
and a consensus is emerging that places tarsiers close to omomyids, while eosimids gave rise to the simians
(non-tarsier haplorrhines) and the adapiforms gave rise to modern strepsirrhines, including lemurs.[6] In
2009, a highly publicized and scientifically criticized publication proclaimed that a 47 million-year-old
adapiform fossil, Darwinius masillae, demonstrated both adapiform and simian traits, making it a transitional
form between the prosimian and simian lineages.[7] Media sources inaccurately dubbed the fossil as a
"missing link" between lemurs and humans.[8]
Lemurs are currently thought to have evolved during the Eocene (37 to 55 mya),[5][9] although
molecular tests suggest the Paleocene (65 to 56 mya) or later.[9] Until recently, they were thought to have
descended directly from the diverse group of adapiforms due to several shared postcranial traits,[10] as well
as long snouts and small brains. Although adapiforms also had lemur-like auditory bullae, a prosimian
characteristic,[11] they had smaller brains and longer snouts than lemurs.[12] There are also several other
morphological differences. Most noticeably, adapiforms lack a key derived trait, the strepsirrhine toothcomb,
and possibly the toilet-claw, found not only in strepsirrhines but also in tarsiers. Unlike lemurs, adapiforms
exhibited a fused mandibular symphysis (a characteristic of simians) and also possessed four premolars,
instead of three or two.[13]
Comparative studies of the cytochrome b gene, which are frequently used to determine phylogenetic
relationships among mammals—particularly within families and genera[14]—have been used to show that
lemurs share common ancestry with lorisiforms.[13][15] This conclusion is also corroborated by the shared
strepsirhine toothcomb, an unusual trait that is unlikely to have evolved twice.[16] If adapiforms were the
ancestors of the living strepsirrhines, then the first strepsirrhines would have to predate the early Eocene, a
view supported by molecular phylogenetic studies,[13] which show that lemurs split from lorises
approximately 62 to 65 mya.[17] The molecular studies also show that lemuriforms diversified before the
modern lorisiforms.[13] Using only nuclear genes, another study dated the split between lemurs and lorises
at 60 mya, lemur diversification at 50 mya, and the lemur colonization of Madagascar somewhere between
these two approximate dates.[18]
The fossil record tells a different story. Although it can only show the earliest possible date for the
appearance of a taxonomic group, other concerns have arisen about these vastly earlier divergence dates
predicted independently of the fossil record. First, palaeontologists have expressed concerns that if primates
have been around for significantly more than 65 million years, then the first one-third of the primate fossil
record is missing. Another problem is that some of the these molecular dates have overestimated the
divergence of other mammalian orders, such as Rodentia, suggesting primate divergence might also be
overestimated. Currently the oldest known strepsirrhine, Djebelemur, dates from the early Eocene in northern
Africa and lacks a fully differentiated toothcomb. Based on fossils and other genetic tests, a more
conservative estimate dates the divergence between lemurs and lorises to around 50 to 55 mya.[5]
To complicate the ancestry puzzle, no terrestrial Eocene or Paleocene fossils have been found on
Madagascar,[19][20] and the fossil record from both Africa and Asia around this time is not much better.[13]
Fossil sites in Madagascar are restricted to only five windows in time, which omit most of the Cenozoic, from
65 mya to ~26,000 years ago. What little fossil-bearing rock exists from this vast span of time is dominated
by marine strata along the west coast.[21] The oldest lemur fossils on Madagascar are actually subfossils
dating to the Late Pleistocene.[10]
[edit] Colonization of Madagascar

Once part of the supercontinent Gondwana, Madagascar broke away from eastern Africa, the likely
source of the ancestral lemur population, about 160 mya and then from Antarctica between 80 and 130 mya.
Initially, the island drifted south from where it split from Africa (around modern Somalia) until it reached its
current position between 80 and 90 mya. Around that time, it split with India, leaving it isolated in the Indian
Ocean and separated from nearby Africa by the Mozambique Channel,[22][23] a deep channel with a
minimum width of approximately 560 kilometers (350 mi).[13] These separation dates and the estimated age
of the primate lineage preclude any possibility that lemurs could have been on the island before the
Madagascar pulled away from Africa.[24] In support of this, mammalian fossils on Madagascar from the
Cretaceous (see Mesozoic mammals of Madagascar) include gondwanatheres and other mammalian groups
that would not have been ancestral to lemurs or the other endemic mammals present on the island today.[13]
A reconstructed map of the Earth during the Early Paleocene, approximately 65 million years ago,
around the time that lemurs evolved and colonized Madagascar
With Madagascar already geographically isolated by the Paleocene and lemur diversification dating
to the same time, an explanation was needed for how lemurs had made it to the island. In the 19th century,
prior to the theory of continental drift, scientists including Philip Sclater, Étienne Geoffroy Saint-Hilaire, and
Ernst Haeckel suggested that Madagascar and India were once part of a southern continent—named Lemuria
by Sclater—that has since disappeared under the Indian Ocean.[25][26] By the early 20th century, oceanic
dispersal emerged as the most popular explanation for how lemurs reached the island.[21][23][27] The idea
first took shape under the anti-plate tectonics movement of the early 1900s, when renowned paleontologist
William Diller Matthew proposed the idea in his influential article "Climate and Evolution" in 1915. In the
article, Matthew could only account for the presence of lemurs in Madagascar by "rafting".[28] Although
unlikely, over long periods of time terrestrial animals can occasionally raft to remote islands on floating mats
of tangled vegetation, which get flushed out to sea from major rivers by floodwaters.[13][28][29] In the 1940s,
American paleontologist George Gaylord Simpson coined the term "sweepstakes dispersal" for such unlikely
events. Today, a rafting event remains the most accepted explanation for the lemur colonization of
Madagascar.[30]
Any extended ocean voyage without fresh water or food would prove difficult for a large, warm-
blooded (or "homeothermic") mammal, but today many small, nocturnal species of lemur exhibit
heterothermy, which allows them to lower their metabolism and become dormant while living off fat reserves.
Such a trait in a small, nocturnal lemur ancestor would have facilitated the ocean voyage and could have
been passed on to its descendants.[29] However, this trait has not been observed in the closely-related
lorisiforms studied to date, and could have evolved on Madagascar in response to the island's harsh
environmental conditions.[13]
Because only five terrestrial orders of mammals have made it to the island, each likely to have
derived from a single colonization,[24] and since these colonizations date to either the early Cenozoic or the
early Miocene, the conditions for oceanic dispersal to Madagascar seem to have been better during two
separate periods in the past.[13] A report published in January 2010 supported this assumption by
demonstrating that both Madagascar and Africa were 1,650 km (1,030 mi) south of their present-day
positions around 60 mya, placing them in a different ocean gyre and reversing the strong current that
presently flows away from Madagascar. The currents were even shown to be stronger than they are today,
shortening the rafting time to approximately 30 days or less, making the crossing much easier for a small
mammal. Over time, as the continental plates drifted northward, the currents gradually changed, and by
20 mya the window for oceanic dispersal had closed.[31]
Since the 1970s, the rafting hypothesis has been called into question by claims that lemur family
Cheirogaleidae might be more closely related to the other Afro-Asian strepsirrhines than to the rest of the
lemurs. This idea was initially based on similarities in behavior and molar morphology, although it gained
support with the 2001 discovery of 30 million-year-old Bugtilemur in Pakistan and the 2003 discovery of
40 million-year-old Karanisia in Egypt. Karanisia is the oldest fossil found that bears a toothcomb, whereas
Bugtilemur was thought to have a toothcomb, but also had even more similar molar morphology to
Cheirogaleus (dwarf lemurs). If these relationships had been correct, the dates of these fossils would have
had implications on the colonization of Madagascar, requiring two separate events. The most parsimonious
explanation, given the genetic evidence and the absence of toothcombed primates in European fossil sites,
[13] is that stem strepsirrhines evolved on the Afro-Arabian landmass, dispersing to Madagascar and more
recently from Africa to Asia.[32] More recently, the structure and general presence of the toothcomb in
Bugtilemur has been questioned, as well as many other dental features, suggesting it is most likely an
adapiform.[5]
An alternative form of oceanic dispersal that had been considered was island hopping, where the
lemur ancestors might have made it to Madagascar in small steps by colonizing exposed seamounts during
times of low sea level.[12][21] However, this is unlikely since the only seamounts found along the Davie
Ridge would have been too small in a such a wide channel. Even though the Comoros Islands between
Africa and Madagascar are significantly larger, they are too young, having been formed by volcanic activity
only around 8 mya.[21] A land bridge between Madagascar and Africa has also been proposed, but a land
bridge would have facilitated the migration of a much greater sampling of Africa's mammalian fauna than is
endemic to the island. Furthermore, deep trenches separate Madagascar from the mainland, and prior to the
Oligocene, sea level was significantly higher than today.[33]
Despite these issues, one variant of the land bridge hypothesis may best explain both how a land
bridge could have formed, and how it other mammalian orders failed to cross it.[5] Geological studies have
shown that following the collision of India and Asia, the Davie Fracture Zone had been pushed up by tectonic
forces, possibly high enough to create a land bridge. Indeed, core samples along the Davie Fracture Zone
suggest that at least parts of the Mozambique Channel were above sea level between 45 and 26 mya,[34] or
possibly as early as 55 mya.[5] Following the Indian-Asian collision, the fault type changed from a strike-slip
fault to a normal fault, and seafloor spreading created compression along the Davie Fracture Zone, causing it
to rise. By the early Miocene, the East African Rift created tension along the fault, causing it to subside
beneath the ocean. Coinciding with this, the divergence dates of nearly all the Malagasy mammalian orders
fall within this window. Old World monkeys, dogs, and cats did not diverge or arrive in Africa until later in the
Miocene.[34] Still, more recent dating of divergence of the Malagasy mammalian clades falls outside of this
land bridge window, and a much greater diversity of mammal groups would be expected on Madagascar had
the land bridge been present during that stretch of time.[18]
The dating of the lemur colonization is controversial for the same reasons as that of strepsirrhine
evolution. Using molecular testing, the colonization has been estimated at 62 to 65 mya based on the split
between the Aye-aye and the rest of the lemurs.[17] On the other hand, the sparse fossil record and
estimates based on nuclear genes support a more conservative estimate of 40 to 52 mya.[5] Once safely
established on Madagascar, with its limited mammalian population, the lemurs were protected from the
increasing competition from evolving arboreal mammalian groups.[19] Monkeys had evolved by the
Oligocene, and their intelligence, aggression, and deceptiveness may have given them the advantage in
exploiting the environment over the diurnal adapiform primates in Africa and Asia, ultimately driving them to
extinction, leaving only the nocturnal lorisiforms.[12][35]
[edit] Diversification
A reconstructed map of the Earth during the Early Oligocene, approximately 35 million years ago, at
a time when lemurs were diversifying.
The ancestral lemur that colonized Madagascar is thought to have been small and nocturnal.[36]
More specifically, it is thought to have had adapiform-like cranial anatomy—particularly the cranial foramina
and the middle ear—comparable to that of lemurids, while being similar to cheirogaleids in dentition and
postcranial anatomy.[5] Lemurs have since diversified greatly, starting with the Aye-aye and its extinct
relations, whose divergence is popularly thought to have occurred shortly after the lemur colonization of
Madagascar.[17] According to molecular studies, there have since been two major episodes of
diversification, from which all other known extant and extinct family lineages emerged. The remaining
families diverged during a 10 to 12 million-year window, between the Late Eocene (42 mya) and into the
Oligocene (30 mya).[17][27] In fact, these dates for this divergence window coincided with the Eocene-
Oligocene extinction event, during which time climate cooling took place and changes in ocean currents
altered weather patterns.[17][5] Outside of Madagascar, these dates also coincide with the divergence of the
lorisiform primates and five major clades of squirrels, all occupying similar niches as lemurs.[17] The dates
do not suggest that increased predation drove family-level divergence since the first carnivores arrived on the
island between 24 and 18 mya.[36]
The second major episode of diversification occurred during the Late Miocene, approximately 8 to
12 mya, and included the true lemurs (Eulemur) and the mouse lemurs (Microcebus).[17][27] In both the true
lemurs and mouse lemurs, their populations were thought to have diverged due to habitat fragmentation
caused when humans arrived on the island roughly 2,000 years ago.[10] Only recently has molecular
research started to show a more distant split in these genera.[37] Most surprising were the mouse lemurs, a
group which is now thought to contain cryptic species, meaning they are indistinguishable from each other
based solely on appearance. In contrast, true lemurs are easier to distinguish and exhibit sexual
dichromatism.[17] Studies in karyology, molecular genetics, and biogeographic patterns have also assisted in
understanding their phylogeny and diversification.[37] Although the divergence times for these two genera
are imprecise, this estimated divergence time would overlap with a change to a wetter climate in
Madagascar, as new weather patterns generated monsoons and likely influenced the plant life.[17][27]
This difference in evolutionary divergence between the two genera may be due to differences in their
activity patterns. True lemurs are often diurnal, allowing sexual partners to distinguish each other as well as
other related species visually. Mouse lemurs, on the other hand, are nocturnal, reducing their ability to use
visual signals for mate selection. Instead, they use olfactory and auditory signaling. For these reasons, true
lemurs may have evolved sexual dichromatism while mouse lemurs evolved to be cryptic species.[17]
[edit] Distribution and diversity

See also: Subfossil lemur
Since their arrival on Madagascar, lemurs have diversified both in behavior and morphology. Their
diversity rivals that of the monkeys and apes found throughout the rest of the world, especially when the
recently extinct subfossil lemurs are considered.[35] Ranging in size from the 30 g (1.1 oz) Madame Berthe's
Mouse Lemur, the world's smallest primate,[38] to the extinct 160–200 kg (350–440 lb) Archaeoindris
fontoynonti,[39] lemurs evolved diverse forms of locomotion, varying levels of social complexity, and unique
adaptations to the local climate. Until recently, they had gone on to fill many niches normally occupied by
monkeys, squirrels, woodpeckers, and large grazing ungulates.[12][19] In addition to the incredible diversity
between lemur families, there has also been great diversification among closely-related lemurs. Yet despite
separation by geographical barriers or by niche differentiation in sympatry, occasionally hybridization can
occur.[35] Lemur diversification has also created generalist species, such as the true lemurs of northern
Madagascar, which are very adaptable, mostly nondescript, and found throughout most of the island's
forests.[10]
The Diademed Sifaka (Propithecus diadema) is one of the largest of the living lemurs, comparable in
size to the Indri. It lives in the rainforests of Madagascar and eats a varied diet of leaves and fruit.
Most of the 99 living lemur taxa are found only on Madagascar. Two species, the Common Brown
Lemur (Eulemur fulvus) and the Mongoose Lemur (Eulemur mongoz), can also be found on the Comoro
Islands, although it is assumed that both species were introduced to the islands from northwestern
Madagascar by humans within the last few hundred years.[40][41] Molecular studies on Eulemur fulvus
fulvus (from the mainland) and E. f. mayottensis (from the Comoro Islands)[24] and on Comoro and mainland
Mongoose Lemurs have supported this assumption by showing no genetic differences between the two
populations.[41] Because all lemurs, including these two brown lemur species, are only native to the island of
Madagascar, they are considered to be endemic.
Historically, lemurs ranged across the entire island inhabiting a wide variety of habitats, including dry
deciduous forests, lowland forests, spiny thickets, subhumid forests, montane forest, and mangrove. Today,
their collective range is restricted to 10% of the island, or approximately 60,000 km2 (23,000 sq mi).[42] Most
of the remaining forests and lemurs are found along the periphery of the island. The center of the island, the
Hauts-Plateaux, was converted by early settlers to rice paddies and grassland through slash-and-burn
agriculture, known locally as tavy. As erosion depleted the soil, the cyclical forest regrowth and burning
ended as the forest gradually failed to return.[43] Today, the level of floral diversity increases with
precipitation, from the dry southern forests to the wetter norther forests to the rainforests along the east
coast. Increased foliage corresponds to increased faunal diversity, including the diversity and complexity of
lemur communities.[10]
Having evolved in Madagascar's challenging environment, replete with poor soils, extreme shifts in
poor, seasonal plant productivity, and devastating climatic events such as extended droughts and annual
cyclones,[9] lemurs have adopted unique combinations of unusual traits to survive, distinguishing them
significantly from other primates. In response to limited, seasonal resources, lemurs may exhibit seasonal fat
storage, hypometabolism (including torpor and hibernation in some cheirogaleids), small group sizes, low
encephalization (relative brain size), cathemerality (activity both day and night), and/or strict breeding
seasons.[9][44] Secondarily, extreme resource limitations and seasonal breeding are thought to have
resulted in three other relatively common lemur traits: female dominance, sexual monomorphism (lack of size
differences between the sexes), and male-male competition for mates involving low levels of agonism
(conflict), such as sperm competition.[45]
The arrival of humans on the island 1,500 to 2,000 years ago has taken a significant toll, not only on
the size of lemur populations, but also on their diversity.[19] Due to habitat destruction and hunting, at least
17 species and 8 genera have gone extinct and the populations of all species have decreased.[39][46] A
couple of species once thought to have gone extinct have since been rediscovered. The Hairy-eared Dwarf
Lemur (Allocebus trichotis) was only known from five museum specimens, most collected in the late 19th
century and one in 1965. It was rediscovered in 1989[47] and has since been identified in five national parks,
although it is very rare within its range.[38] Likewise, the Greater Bamboo Lemur (Prolemur simus) was
thought to be extinct as recently as the late 1970s, but a population was located near Ranomafana National
Park in the late 1980s.[48] Historically, it had a much wider geographic distribution, shown by subfossil
remains, but today it remains one of the world's 25 most endangered primates.[48][49][50][51] One distinctive
morph (possibly a species or subspecies) of sifaka,[N 1] has not been so fortunate, having been extirpated
from all known localities.[54] Unless trends change, extinctions are likely to continue.[55]
A life restoration of Palaeopropithecus ingens, a giant sloth lemur that went extinct less than a
thousand years ago.
Until recently, giant species of lemur existed on Madagascar. Now represented only by recent or
subfossil remains, they were modern forms and are counted as part of the rich lemur diversity that has
evolved in isolation. Some of their adaptations were unlike those seen in lemurs today.[19] All 17 extinct
lemurs were larger than the extant forms, some weighing as much as 200 kg (440 lb),[35] and are thought to
have been active during the day.[56] Not only were they unlike the living lemurs in both size and appearance,
they also filled ecological niches that no longer exist or are now left unoccupied.[19] Large parts of
Madagascar, which are now devoid of forests and lemurs, once hosted diverse primate communities that
included more than 20 species covering the full range of lemur sizes.[57]
[edit] Taxonomic and phylogenetic classification

For a more comprehensive list, see List of lemur species
Lemur taxonomy is controversial, and not all experts agree, particularly with the recent increase in
the number of recognized species.[58][59][60] According to Russell Mittermeier, the president of
Conservation International (CI), taxonomist Colin Groves, and others, there are currently 101 recognized
species or subspecies of extant lemur, divided into five families and 15 genera.[61] Conversely, other experts
in the field label this as a possible example of taxonomic inflation,[60] and prefer instead an estimate of at
least 50 species.[58] All sides generally agree that the recently extinct subfossil lemurs should be classified
in three families, eight genera, and 17 species.[39][46]
Closest lemur relations[3]
Euarchonta
Scandentia
(treeshrews)
Dermoptera
(colugos)
†Plesiadapiforme
s
Primates
Haplorrhini
(tarsiers, monkeys,
apes)
Strepsirrhini
Lorises,
pottos, and
galagos
Lemurs
Since the first taxonomic classification of lemurs in 1758 by Carl Linnaeus, many changes have been
made to lemur taxonomy. Within the order Primates, treeshrews (order Scandentia) were considered basal,
prosimian primates—close relatives of lemurs—until the 1980s.[62] Colugos, also incorrectly referred to as
"flying lemurs", were once considered lemur-like primates, but were reclassified as close relatives of bats,[63]
and more recently as close relatives of primates within their own order, Dermoptera.[3] Primates, together
with their closest relatives, the treeshrews, colugos, and long-extinct plesiadapiforms, form the taxonomically
unranked Euarchonta clade within the Euarchontoglires. Also, all lorisids originally placed in the genus
Lemur by Carl Linnaeus have since been moved into either their own infraorder (Lorisiformes) or their own
superfamily (Lorisoidea) within Lemuriformes.[64][5]
For the Malagasy primate fauna, taxonomic nomenclature proliferated during the 1800s, with the aid
of museum systematists, such as Albert Günther and John Edward Gray, as well as naturalists and
explorers, such as Alfred Grandidier and Alphonse Milne-Edwards.[65][66] The taxonomic nomenclature of
lemurs was not sorted out until decades later, when Ernst Schwarz standardized it in 1931.[65][66][67] It was
not until the 1990s that this nomenclature started to see a new wave of taxonomic change.[60]
[edit] Suprageneric classification

Since the 19th century, the classification of lemurs above the genus level has seen many changes.
Early taxonomists proposed a variety of classifications for lemurs, but generally separated indriids from other
lemurs and placed the Aye-aye in a major group of its own; some classified the dwarf and mouse lemurs with
the galagos.[68] In 1915, William King Gregory published a classification[69] that remained generally
accepted over the next decades. He placed all the lemurs together in a "series" Lemuriformes and
recognized three families: Daubentoniidae, Indriidae, and Lemuridae (including the current Cheirogaleidae
and Lepilemuridae).[68] George Gaylord Simpson's influential 1945 classification of mammals placed the
treeshrews and the fossil Anagale (both now classified outside Primates) inside Lemuriformes and classified
the fossil families Plesiadapidae and Adapidae in a superfamily Lemuroidea with most of the lemurs.[70]
Although treeshrews, plesiadapids, and the like are now no longer considered to be closely related to
lemurs, disagreements persist over the classification of lemurs and related groups, resulting in two
competing arrangements of the infraorders and superfamilies within Strepsirrhini. Colin Groves, in the 2005
third edition of Mammal Species of the World, classifies living strepsirrhines under three infraorders and two
superfamilies. This places the Aye-aye within its own infraorder, separate from both lemurs (divided into two
superfamilies) and lorises.[64] Since the publication of Mammal Species of the World, there has been little
support in the academic literature for placing the Aye-aye in its own infraorder, and more recently
Mittermeier, Groves, and other editors have ignored this taxonomic level.[71] An alternative classification
draws the lines for infraorders and superfamilies differently, though using the same general phylogenetic
tree. It classifies all living strepsirrhines under one infraorder, with the lorises and lemurs in separate
superfamilies.[5]
Two alternative lemur classifications at the infraorder and superfamily levels
3 infraorders, 2 superfamilies[64] 1 infraorder, 2 superfamilies[5
• Order Primates • Order Primates

• Suborder Strepsirrhini: non-tarsier • Suborder Strepsirrhini: non-ta
prosimians • Infraorder Lemuriform
• Infraorder Chiromyiformes: Aye- • Superfamily L
aye • Family
• Infraorder Lemuriformes monkey or bab
• Family Cheirogaleidae: • Family
dwarf and mouse lemurs dwarf and mou
• Family • Family
†Archaeolemuridae: monkey or Aye-aye
baboon lemurs • Family
• Family Indriidae: woolly lemurs, sifaka
lemurs, sifakas, and allies • Family
• Family Lemuridae: brown lemurs and all
lemurs and allies • Family
• Family Lepilemuridae: sportive lemur
sportive lemurs • Family
• Family †Megaladapidae: koala lemurs
koala lemurs • Family
• Family †Palaeopropit
†Palaeopropithecidae: sloth
lemurs
• Superfamily L
• Infraorder Lorisiformes: galagos
and lorises
and lorises
• Suborder Haplorrhini: tarsiers,
• Suborder Haplorrhini: tarsiers, monkeys,
and apes
The classification of several lemur taxa has elicited particular debate. Most significantly, the
placement of the Aye-aye has been controversial since its introduction to Western science in 1782, and it has
been a topic of debate up until very recently.[35][73][68] Hinged upon morphological traits and molecular
data, it has had profound implications on scientific theories.[73] Arguing against Darwin's theory of natural
selection, Richard Owen claimed in 1863 that the Aye-aye's distinct characteristics, including its ever-
growing incisors and unique, highly flexible middle finger, are so perfectly adapted for their uses in extractive
foraging that they could not have evolved gradually through natural selection.[73] More recently, the Aye-
aye's placement within the order Primates has posed problems for the rafting hypothesis for the primate
colonization of Madagascar. If this species does not form a monophyletic group with the rest of the lemurs,
then multiple colonization events would have had to occur to explain the present-day distribution of non-
human primates on Madagascar.[35]
The Aye-aye has traditionally been difficult to classify due to its unique physical traits.
Until Richard Owen published a definitive anatomical study in 1866, early naturalists were uncertain
whether the Aye-aye (genus Daubentonia) was a primate, rodent, or marsupial.[73][74][75] In the late
eighteenth century, for example, the Aye-aye was classified under the squirrel genus Sciurus.[76] By
emphasizing its primate features, such as its postorbital bar, stereoscopic vision, and opposable hallux, over
its rodent-like teeth, Owen demonstrated its affinity with other primates.[73][77] In 1996, Ankel-Simons
demonstrated that the shape and arrangement of the Aye-aye's diminutive deciduous incisors indicate that
this genus has a shared ancestry with the toothcombed primates.[16] However, the placement of the Aye-aye
within the order Primates remained problematic until very recently. The karyotype of the Aye-aye is
noticeably different from that of its closest relatives, the lorises and the rest of the lemurs, with a diploid
chromosome count of 2n=30.[78] Based on its anatomy, researchers have found support for classifying the
genus Daubentonia as a specialized indriid, a monotypic sister group to all strepsirrhines, and an
indeterminate taxon within the order Primates.[15] In 1931, Schwarz labeled the Aye-aye as an offshoot of
Indriidae, claiming that all lemurs were monophyletic, whereas Reginald Innes Pocock had previously placed
the Aye-aye outside of the lemurs.[67] In that same year, Anthony and Coupin classified the Aye-aye under
infraorder Chiromyiformes, a sister group to the other strepsirrhines. Colin Groves upheld this classification in
2005 because he was not entirely convinced the Aye-aye formed a clade with the rest of the Malagasy
lemurs,[79] despite molecular tests that had shown Daubentoniidae was basal to all Lemuriformes.[15][80] In
2008, Russell Mittermeier, Colin Groves, and others ignored addressing higher-level taxonomy by defining
lemurs as monophyletic and containing five living families, including Daubentoniidae.[71]
Another interpretation of the Aye-aye's origins has once again called into question the single origins
of the lemurs. Comparisons have been made between the Aye-aye and a fossil strepsirrhine primate from
Africa, Plesiopithecus. Similarities in the shape of the skull and the morphology of the lower jaw have raised
the question of whether or not this could be an Aye-aye ancestor. However, the placement of an Aye-aye
ancestor in Africa would require multiple colonizations by strepsirrhine primates. Molecular tests may offer
support since they show that the Aye-aye was the first to diverge in the lemur clade and that the other lemur
families did not diverge until much later.[5]
Often classified with the galagos by early students, the cheirogaleids were placed with the other
lemurs from Gregory's 1915 classification until the early 1970s, when several anthropologists proposed that
they are more closely related to lorisiforms, based on morphological data.[68][81] However, relevant genetic
studies nearly unanimously place cheirogaleids within the lemuriform clade and Groves, who had promoted
the cheirogaleid-lorisiform relationship in a 1974 paper, by 2001 himself regarded the idea as refuted.[15][80]
[81]
Lemur phylogeny[27][57][82]
Strepsirrhini
Lorisiform clade
Galagidae
Lorisidae
Lemur clade
Daubentonia
Lemuridae
†Megaladapidae
Lepilemuridae
Cheirogaleidae
†Archaeolemurida
e
†Palaeopropithecida
e
Indriidae
Classifications in the first half of the 20th century divided lemurs into three families—Daubentoniidae,
Indriidae, and Lemuridae, with the latter including the current Cheirogaleidae and Lepilemuridae.[68]
Because of concerns that Lemuridae might not be monophyletic, the family was later split; in 1982, for
example, Tattersall separated the Cheirogaleidae for the dwarf lemurs, mouse lemurs, and relatives and the
Lepilemuridae for the sportive lemurs and bamboo lemurs (including the Greater Bamboo Lemur).[83] This
classification is still used, except that the bamboo lemurs are placed in Lemuridae.[84][71]
From the 1970s to the 1990s, there have been suggestions that the ruffed lemurs might be related to
indriids or a sister group to Lemuridae and Indriidae and that the bamboo lemurs are related to the sportive
lemurs,[85] but neither view is supported by molecular phylogeny.[27] The sportive lemurs and the extinct
koala lemurs (Megaladapidae) both lack upper incisors in the permanent dentition,[74] and in 1981, Groves
placed both together in the family Megaladapidae, which he renamed Lepilemuridae in 2005 because that
older name takes precedence.[86] Genetic research does not support a close relationship between the
sportive and koala lemurs and instead places the koala lemurs as a sister group to Lemuridae; therefore, the
two are now placed in separate families (Lepilemuridae for the sportive lemurs and Megaladapidae for the
koala lemurs).[87][88][82] The sloth lemurs (Palaeopropithecidae) and monkey lemurs (Archaeolemuridae)
were classified as subfamilies within Indriidae as late as 1982,[83] but are now recognized as separate
families.[82]
The relationships among the families of lemurs have been problematic and have yet to be definitively
resolved. Two competing phylogenies exist based on genetic and molecular data. One approach (Horvath,
et al.) looks at a larger number of genes, but among fewer species. This results in Lemuridae being a sister
group to Lepilemuridae, Cheirogaleidae, and Indriidae.[27] The other approach (Orlando, et al.) looks at
fewer genes, but more lemur species. Using this analysis, Lepilemuridae becomes the sister group to
Lemuridae, Cheirogaleidae, and Indriidae.[82] Both phylogenies agree that the Malagasy primates are
monophyletic and that Daubentoniidae (the Aye-aye) is basal to the lemuriform clade, having split off
significantly earlier than the other families.[15][27][82] However, two problems create complications for both
approaches. First, the four most closely related lemur families diverged within a narrow window of
approximately 10 million years, making it much harder to distinguish the splits with molecular evidence.
Second, the divergence occurred approximately 42 mya;[17] such distant splits create a lot of noise for
molecular techniques.
[edit] Genus-level classification
Early classifications of the genera of lemurs differed in a number of ways from current taxonomy. For
example, the fork-marked lemurs were initially placed in the genus Lemur and then in Microcebus with the
mouse lemurs before being placed in their own genus Phaner;[47][89][67][90] and Charles Immanuel Forsyth
Major split the Cheirogaleus medius species group of the dwarf lemurs into a separate genus Opolemur, but
this was not accepted.[90][67] Genus-level taxonomy was largely stabilized by Schwarz in 1931,[67] but a
number of changes have become accepted:
• The Ring-tailed Lemur, ruffed lemurs, and true lemurs were once grouped together in the
genus Lemur due to a host of morphological similarities. For instance, the skeletons of the Ring-
tailed Lemur and the true lemurs are nearly indistinguishable.[50] However, ruffed lemurs were
reassigned to the genus Varecia in 1962,[91] and due to similarities between the Ring-tailed Lemur
and the bamboo lemurs, particularly in regards to molecular evidence and scent glands similarities,
the true lemurs were moved to the genus Eulemur in 1988.[49][50][92] The genus Lemur is now
monotypic, containing only the Ring-tailed Lemur.
• In 2001, Colin Groves concluded that despite similarities, the Greater Bamboo Lemur was
sufficiently distinct from the bamboo lemurs of the genus Hapalemur to merit its own monotypic
genus, Prolemur.[50][93] This follows Schwarz's 1931 opposition to Pocock's decision to separate
Prolemur from Hapalemur.[67]
• Originally placed in the genus Microcebus (mouse lemurs), the Giant Mouse Lemur was
moved to its own genus, Mirza, in 1985 due to its larger size, morphological differences, dental
characteristics, and behavior.[47][94]
• The Hairy-eared Dwarf Lemur was first placed in the genus Cheirogaleus (dwarf lemurs) in
1875 and was later found to have closer affinities with Microcebus. However, its dentition and
cranium structure were sufficiently distinct to merit elevation to its own genus, Allocebus.[47][95]
[edit] Species-level classification

Over the past two decades, the number of recognized lemur species has more than doubled
according to some experts. In 1994, 32 distinct species were named in the first edition of Conservation
International's field guide, Lemurs of Madagascar, and 68 were described in the 2nd edition, published in
2006.[58][96] In December 2008, Russell Mittermeier, Colin Groves, and other experts co-wrote an article in
the International Journal of Primatology classifying 99 species and subspecies.[71] In late 2010, the 3rd
edition of Lemurs of Madagascar listed 101 taxa.[61] The number of lemur species is likely to continue
growing in the coming years, as field studies, cytogenetic and molecular genetic research continues,
particularly on cryptic species, such as mouse lemurs, which cannot be distinguished visually.[58]
The Sahamalaza Sportive Lemur (Lepilemur sahamalazensis) was identified as a distinct species as
recently as 2006.
This threefold increase in nearly 15 years has not had universal support among taxonomists and
lemur researchers. In many cases, classifications ultimately depends upon which species concept is used.
Due to the critical condition that most Malagasy primate populations are in, taxonomists and conservationists
sometimes favor splitting them into separate species to develop an effective strategy for the conservation of
the full range of lemur diversity.[58][71] Implicitly, this means that full species status will help grant genetically
distinct populations added environmental protection.[58]
The first large wave of new lemur species descriptions came in 2001 when Colin Groves elevated the
Red Ruffed Lemur (Varecia rubra),[50][97] five subspecies of brown lemur (Eulemur albifrons, E. albocollaris,
E. collaris, E. rufus and E. sanfordi),[98] and four subspecies of sifaka (Propithecus coquereli, P. deckenii,
P. edwardsi, and P. perrieri) to full species status.[99] Additional elevations of all remaining subspecies within
the Eulemur and Propithecus genera were made in the years that followed.[49][50][100] These and
subsequent changes in taxonomy were largely due to a shift to the phylogenetic species concept,[101] yet
are not universally endorsed.[60]
By far the most explosive growth in species numbers has been in the genera Microcebus and
Lepilemur. In 2006, 15 new species of Lepilemur were described, with three new species reported in
February,[102] one species in June,[103] and 11 in September.[104] Since then, two additional species have
been described.[71] Genetic and morphological differences seem to suggest that they are cryptic species,
but there is still debate whether these merit full species status or should be regarded as subspecies of
previously identified, "core" species.[60][87]
In true lemurs and mouse lemurs, both groups were initially divided into a small number of species,
either with no distinguishable subspecies (in the case of mouse lemurs) or with several distinguishable
subspecies (in the case of true lemurs).[10][37] With molecular research suggesting a more distant split in
both genera, these subspecies or undistinguished populations have been promoted to species status.[37]
In the case of mouse lemurs, the rise in species numbers has been only slightly less sudden and
dramatic. Classified as one species by Ernst Schwarz in 1931 (excluding one, Coquerel's Giant Mouse
Lemur, that is no longer classified in Microcebus),[67] the genus was revised to contain two species, the
Gray Mouse Lemur (Microcebus murinus) and the Brown Mouse Lemur (M. rufus), after an extensive field
study in 1972 showed both living in sympatry in southeastern Madagascar.[105] At the time, the Gray Mouse
Lemur was known in the drier parts of the north, west, and south, while the Brown Mouse Lemur inhabited
the humid rainforest regions of the east. However, we now know the species diversity and distribution to be
significantly more complex.[47] Revisions throughout the 1990s and 2000s identified numerous new species
through genetic testing using mitochondrial DNA, demonstrating that the genus is represented by a multitude
of cryptic species.[71][106][107][108] Many, but not all of these defined species have been supported by
nuclear DNA tests.[109]
However, there are still concerns that species are being identified prematurely. Ian Tattersall, an
anthropologist who recognized 42 species of lemur in 1982,[110] has expressed concern that the
geographically organized variety in lemur populations is being recognized with full species status while the
number of subspecies in lemur genera has virtually disappeared. He has argued that taxonomists are
confusing differentiation and speciation, two processes that are often unrelated, while denying the role of
microevolution in evolutionary processes.[60] Still, other researchers who emphasize the framework of the
"general lineage concept of species" contend that lineage divergence or differentiation demarcates the
beginning of a new species.[109]
New species have been identified due to differences in morphology, karyotypes, cytochrome b
sequences, and other genetic tests, as well as several combinations of these.[60] When nuclear DNA
(nDNA) was tested in conjunction with mitochondrial DNA (mtDNA) in mouse lemurs, a few species, such as
Claire's Mouse Lemur (Microcebus mamiratra) were demonstrated to be indistinguishable from other closely
related species. In such cases, nDNA did not vary, but the mtDNA that had been used to define it as a
species was still distinct. Differences in results between nDNA, which is inherited from both parents, and
mtDNA, which is inherited from the mother, was attributed to female philopatry, where females remain within
or close to the home range into which they were born while males disperse. Since the isolated population
known as Claire's Mouse Lemur has distinct mtDNA, but not nDNA, it is likely to contain a population
descended from a related group of females, but which still disperses and interbreeds with nearby
populations.[109]
Traditionally, karyology has been considered when determining species status. From the lemurs
studied so far, the diploid number of chromosomes in lemurs varies between 2n=20 and 2n=66. In the case
of the true lemurs, the diploid number ranges from 2n=48 to 2n=60 while the individual chromosome sizes
vary considerably.[78]
Sometimes distinctions are made due to very slight differences in pelage coloration. For instance,
three distinctly-colored types of mouse lemur were discovered in a multi-year study in Beza Mahafaly
Reserve in southern Madagascar, but rather than being separate species, DNA tests revealed that they all
belonged to a single species, the Reddish-gray Mouse Lemur (Microcebus griseorufus).[111] For this reason,
further research is needed to confirm or deny the recent species splits. Only through detailed studies of
morphology, ecology, behavior, and genetics can the true number of lemur species be determined.[58]
Lemur species and subspecies count by year and genus
1982 1994 2005
Tattersall[110] Mittermeier et al.[112] Groves[84] Mitterm
species subspecies species subspecies species subspecies specie

1 0 1 0 1 0 1
1 2 2 0 3 0 4
2 0 2 0 7 0 7
1 0 1 0 1 0 1
– – 5 8 11 2 10
2 3 3 3 4 2 5
1 0 1 0 1 2 1
4 7 1 0 1 0 1
1 6 7 0 8 0 8
2 0 3 0 8 0 12
1 0 1 0 1 0 2
1 0 1 4 4 0 4
– – – – 1 0 1
2 9 3 8 7 4 9
1 2 1 2 1 4 2
20 29 32 25 59 14 68
42 50 67
[edit] Notes
1. ^ Propithecus diadema holomelas was once considered one of five subspecies of Diademed
Sifaka. In 1986, Ian Tattersall subsumed it as a synonym for what is now known as Milne-Edwards'
Sifaka (known then as Propithecus diadema edwardsi). Both subspecies had only slight color
variations and were known to be sympatric with each other in at least one forest.[52] Since it was
extirpated, the taxonomic status of Propithecus diadema holomelas has been questioned, but
nothing definitive has been published.[53]
2. ^ a b In 1931, Allocebus was not recognized as a separate genus, but was lumped under the
genus Cheirogaleus.
3. ^ a b In 1931 and 1982, Eulemur was not recognized as a separate genus, but was lumped
under the genus Lemur.
4. ^ a b The genus Prolemur was not distinguished until 2001. Prior to this, the Greater Bamboo
Lemur was placed in the genus Hapalemur.
5. ^ a b In 1931, Varecia was not recognized as a separate genus, but was lumped under the
genus Lemur.
6. ^ a b In 1931, Mirza was not recognized as a separate genus, but was lumped under
Microcebus.
7. ^ The grand total of species and subspecies for each year will not equal the sum of the
species total and the subspecies total since each subspecies group already counts as one species.
For example, in 2010 there were three subspecies for Varecia variegata recognized: V. v. variegata,
V. v. editorum, and V. v. subcincta. Together, they count as one species, which is already included in
the species total for that year. The same applies that year for Hapalemur. For this reason there are
101 species and subspecies (97 − 2 + 6 = 101), not 103. In 1994, there were seven species with
subspecies listed, so the grand total is: 32 − 7 + 25 = 50
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78.^ a b Ankel-Simons 2007, pp. 533–559
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84.^ a b Groves, C. (2005). Wilson, D. E., & Reeder, D. M, eds. ed. Mammal Species of the
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85.^ Groves 2001, pp. 54–55
86.^ Groves, C. (2005). "Lepilemuridae". In Wilson, D. E., & Reeder, D. M, eds. Mammal
87.^ a b Garbutt 2007, pp. 115–136
88.^ Karanth, K. P.; Delefosse, T.; Rakotosamimanana, B.; Parsons, T. J.; Yoder, A. D. (2005).
"Ancient DNA from giant extinct lemurs confirms single origin of Malagasy primates" (PDF).
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89.^ Groves, C. (2005). "Phaner". In Wilson, D. E., & Reeder, D. M, eds. Mammal Species of
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90.^ a b Groves 2001, pp. 65–66
91.^ Groves, C. (2005). "Varecia". In Wilson, D. E., & Reeder, D. M, eds. Mammal Species of
92.^ Groves, C. (2005). "Eulemur". In Wilson, D. E., & Reeder, D. M, eds. Mammal Species of
93.^ Groves, C. (2005). "Prolemur". In Wilson, D. E., & Reeder, D. M, eds. Mammal Species of
94.^ Groves, C. (2005). "Mirza". In Wilson, D. E., & Reeder, D. M, eds. Mammal Species of the
World (3rd ed.). Baltimore: Johns Hopkins University Press. ISBN 0-801-88221-4. OCLC 62265494.
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95.^ Groves, C. (2005). "Allocebus". In Wilson, D. E., & Reeder, D. M, eds. Mammal Species of
97.^ Groves, C. (2005). "Varecia rubra". In Wilson, D. E., & Reeder, D. M, eds. Mammal
98.^ Groves, C. (2005). "Eulemur fulvus". In Wilson, D. E., & Reeder, D. M, eds. Mammal
99.^ Groves, C. (2005). "Propithecus". In Wilson, D. E., & Reeder, D. M, eds. Mammal Species
of the World (3rd ed.). Baltimore: Johns Hopkins University Press. ISBN 0-801-88221-4.
100.^ Garbutt 2007, pp. 176–204
101.^ Thalmann, U. (December 2006). "Lemurs – Ambassadors for Madagascar". Madagascar
Conservation & Development 1: 4–8. ISSN 1662-2510. http://www.mwc-
info.net/en/services/Journal_PDF%27s/Issue1/Flagshipspecies.pdf.
102.^ Andriaholinirina, N.; Fausser, J. L.; Roos, C.; Zinner, D.; Thalmann, U.; Rabarivola, C.;
Ravoarimanana, I.; Ganzhorn, J. U. et al. (2006). "Molecular phylogeny and taxonomic revision of the
sportive lemurs (Lepilemur, Primates)" (PDF). BMC Evolutionary Biology 6: 17. doi:10.1186/1471-
2148-6-17. PMID 16504080. PMC 1397877. http://www.biomedcentral.com/content/pdf/1471-2148-
6-17.pdf. edit
103.^ Rabarivola, C.; Zaramody, A.; Fausser, J.; Andriaholinirina, N.; Roos, C.; Zinner, D.;
Marcel, H.; Rumpler, Y. (2006). "Cytogenetic and molecular characteristics of a new species of
sportive lemur from Northern Madagascar". Lemur News 11: 45–49. ISSN 0343-3528.
http://www.primate-sg.org/PDF/LN11.pdf.
104.^ Louis, Jr., E.E.; Engberg, S.E.; Lei, R.; Geng, H.; Sommer, J.A.; Randriamamapionona,
R.; Randriamanana, J.C.; Zaonarivelo, J.R. et al. (2006). "Molecular and morphological analyses of
the sportive lemurs (Family Megaladapidae: Genus Lepilemur) reveals 11 previously unrecognized
species". Texas Tech University Special Publications (49): 1–49. ISSN 0169-0237.
http://www.nsrl.ttu.edu/publications/opapers/specpubs/SP49.pdf.
105.^ Kappeler & Rasoloarison 2003, pp. 1310–1315
106.^ Braune, P.; Schmidt, S.; Zimmermann, E. (2008). "Acoustic divergence in the
communication of cryptic species of nocturnal primates (Microcebus ssp.)". BMC Biology 6 (19).
doi:10.1186/1741-7007-6-19. PMID 18462484. Lay summary – ScienceDaily (14 May 2008). edit
107.^ Olivieri, G.; Zimmermann, E.; Randrianambinina, B.; Rasoloharijaona, S.;
Rakotondravony, D.; Guschanski, K.; Radespiel, U. (2007). "The ever-increasing diversity in mouse
lemurs: Three new species in north and northwestern Madagascar". Molecular Phylogenetics and
Evolution 43 (1): 309–327. doi:10.1016/j.ympev.2006.10.026. PMID 17197200. edit
108.^ Louis, E. E.; Engberg, S. E.; McGuire, S. M.; McCormick, M. J.; Randriamampionona, R.;
Ranaivoarisoa, J. F.; Bailey, C. A.; Mittermeier, R. A. et al. (2008). "Revision of the Mouse Lemurs,
Microcebus (Primates, Lemuriformes), of Northern and Northwestern Madagascar with Descriptions
of Two New Species at Montagne d'Ambre National Park and Antafondro Classified Forest" (PDF).
Primate Conservation 23: 19–38. doi:10.1896/052.023.0103. http://www.primate-
sg.org/PDF/PC23.new.microcebus.V3.pdf. re edit
109.^ a b c Weisrock, D. W.; Rasoloarison, R. M.; Fiorentino, I.; Ralison, J. M.; Goodman, S. M.;
Kappeler, P. M.; Yoder, A. D. (2010). "Delimiting Species without Nuclear Monophyly in
Madagascar's Mouse Lemurs". PLoS ONE 5 (3): e9883. doi:10.1371/journal.pone.0009883.
PMID 20360988. edit
110.^ a b Tattersall 1982
111.^ Heckman, K. L.; Rasoazanabary, E.; MacHlin, E.; Godfrey, L. R.; Yoder, A. D. (2006).
"Incongruence between genetic and morphological diversity in Microcebus griseorufus of Beza
Mahafaly" (PDF). BMC Evolutionary Biology 6 (98). doi:10.1186/1471-2148-6-98. PMID 17109740.
PMC 1657033. http://www.biology.duke.edu/yoderlab/reprints/2006heckman_etal.BMCEB.pdf. Lay
summary – ScienceDaily (16 November 2006). edit
112.^ Mittermeier, R.A.; Tattersall, I.; Konstant, W.R.; Meyers, D.M.; Mast, R.B. (1994). Lemurs
of Madagascar. Illustrated by S.D. Nash (1st ed.). Conservation International. pp. 80–82. ISBN 1-
881173-08-9.
Books cited
• Ankel-Simons, F. (2007). Primate Anatomy (3rd ed.). Academic Press. ISBN 0-12-372576-3.
• Garbutt, N. (2007). Mammals of Madagascar, A Complete Guide. A&C Black Publishers.
ISBN 978-0-300-12550-4.
• Goodman, S.M.; Benstead, J.P., eds (2003). The Natural History of Madagascar. University
of Chicago Press. ISBN 0-226-30306-3.
Chapter 2 - Geology and Soils
• Flynn, J.J.; Wyss, A.R. (2003). "Mesozoic Terrestrial Vertebrate Faunas: The Early
History of Madagascar's Vertebrate Diversity". pp. 34–40.
• Krause, D.W. (2003). "Late Cretaceous Vertebrates of Madagascar: A Window into
Gondwanan Biogeography at the End of the Age of Dinosaurs". pp. 40–47.
• Burney, D.A. (2003). "Madagascar's Prehistoric Ecosystems". pp. 47–51.
Chapter 13 - Mammals
• Goodman, S.M.; Ganzhorn, J.U.; Rakotondravony, D. (2003). "Introduction to the
Mammals". pp. 1159–1186.
• Yoder, A.D. (2003). "Phylogeny of the Lemurs". pp. 1242–1247.
• Godfrey, L.R.; Jungers, W.L. (2003). "Subfossil Lemurs". pp. 1247–1252.
• Kappeler, P.M.; Rasoloarison, R.M. (2003). "Microcebus, Mouse Lemurs, Tsidy".
pp. 1310–1315.
• Mutschler, T.; Tan, C.L. (2003). "Hapalemur, Bamboo or Gentle Lemur". pp. 1324–
1329.
• Goodman, S.M.; Patterson, B.D., eds (1997). Natural Change and Human Impact in
Madagascar. Smithsonian Institution Press. ISBN 978-1560986829.
• Krause, D.W.; Hartman, J.H.; Wells, N.A. (1997). "Chapter 1: Late Cretaceous
Vertebrates from Madagascar: Implications for Biotic Change in Deep Time". pp. 3–43.
• Simons, E.L. (1997). "Chapter 6: Lemurs: Old and New". pp. 142–166.
• Gould, L.; Sauther, M.L., eds (2006). Lemurs: Ecology and Adaptation. Springer. ISBN 978-
0387-34585-7.
• Gould, L.; Sauther, M.L. (2006). "Preface". pp. vii–xiii.
• Tattersall, I. (2006). "Chapter 1: Origin of the Malagasy Strepsirhine Primates". pp. 3–
18.
• Jolly, A.; Sussman, R.W. (2006). "Chapter 2: Notes on the History of Ecological
Studies of Malagasy Lemurs". pp. 19–40.
• Godfrey, L.R.; Jungers, W.L.; Schwartz, G.T. (2006). "Chapter 3: Ecology and
Extinction of Madagascar's Subfossil Lemurs". pp. 41–64.
• Freed, B.Z. (2006). "Chapter 6: Polyspecific Associations of Crowned Lemurs and
Sanford's Lemurs in Madagascar". pp. 111–132.
• Curtis, D.J. (2006). "Chapter 7: Cathemerality in Lemurs". pp. 133–158.
• Sterling, E.J.; McCreless, E.E. (2006). "Chapter 8: Adaptations in the Aye-aye: A
Review". pp. 159–184.
• Johnson, S.E. (2006). "Chapter 9: Evolutionary Divergence in the Brown Lemur
Species Complex". pp. 187–210.
• Irwin, M.T. (2006). "Chapter 14: Ecologically Enigmatic Lemurs: The Sifakas of the
Eastern Forests (Propithecus candidus, P. diadema, P. edwardsi, P. perrieri, and P.
tattersalli)". pp. 305–326.
• Groves, C.P. (2001). Primate Taxonomy. Washington, D.C.: Smithsonian Institution Press.
ISBN 978-1560988724.
• Mittermeier, R.A.; Louis, E.E.; Richardson, M.; Schwitzer, C.; Langrand, O.; Rylands, A.B.;
Hawkins, F.; Rajaobelina, S. et al. (2010). Lemurs of Madagascar. Illustrated by S.D. Nash (3rd ed.).
Conservation International. ISBN 978-1-934151-23-5.
• Mittermeier, R.A.; Konstant, W.R.; Hawkins, F.; Louis, E.E.; Langrand, O.; Ratsimbazafy, J.;
Rasoloarison, R.; Ganzhorn, J.U. et al. (2006). Lemurs of Madagascar. Illustrated by S.D. Nash (2nd
ed.). Conservation International. ISBN 1-881173-88-7.
• Platt, M.; Ghazanfar, A., eds (2010). Primate Neuroethology. Oxford University Press.
ISBN 978-0195-32659-8.
• Cartmill, M. (2010). "Chapter 2: Primate Classification and Diversity". pp. 10–30.
id=hv28p1tCnnEC&pg=PA10&dq=lemuridae+cladogram#v=onepage&q&f=false.
• Preston-Mafham, K. (1991). Madagascar: A Natural History. Facts on File. ISBN 978-
0816024032.
• Rowe, N. (1996). The Pictorial Guide to the Living Primates . Pogonias Press. ISBN 978-
0964882515.
• Sussman, R.W. (2003). Primate Ecology and Social Structure . Pearson Custom Publishing.
ISBN 978-0536743633.
• Tattersall, I. (1982). Primates of Madagascar. Columbia University Press. ISBN 978-
0231047043.
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Evolutionary history of cephalopods


The cephalopods have a long geological history, with the first nautiloids found in late Cambrian
strata,[1] and purported stem-group representatives present in the earliest Cambrian lagerstätten.[2]
The class developed during the middle Cambrian, and underwent pulses of diversification during the
Ordovician period[3] to become diverse and dominant in the Paleozoic and Mesozoic seas. Small shelly
fossils such as Tommotia were once interpreted as early cephalopods, but today these tiny fossils are
recognized as sclerites of larger animals,[4] and the earliest accepted cephalopods date to the Middle
Cambrian Period. During the Cambrian, cephalopods are most common in shallow near-shore environments,
but they have been found in deeper waters too.[5] Cephalopods were thought to have "undoubtedly" arisen
from within the tryblidiid monoplacophoran clade.[6] However genetic studies suggest that they are more
basal, forming a sister group to the Scaphopoda but otherwise basal to all other major mollusc classes.[7]
The internal phylogeny of Mollusca, however, is wide open to interpretation – see mollusc phylogeny.
Contents
[hide]
• 1 Traditional view of origins
• 2 Early shelly record
• 2.1 Cambrian cephalopods
• 2.1.1 Knightoconus
• 2.1.2 Plectronoceras
• 2.1.3 Yochelcionellids
• 2.1.4 Ellesmerocerida
• 2.2 Early Ordovician diversity
• 2.3 Fossils mistaken for cephalopods
• 2.3.1 Volborthella
• 2.3.2 Shelbyoceras
• 2.3.3 Kirengellids
• 3 Coleoidea
• 4 Organ origins
• 5 Exceptional preservation
• 6 Notes
• 7 References
• 8 Further reading
[edit] Traditional view of origins
The cephalopods were once thought to have evolved from a monoplacophoran-like ancestor[8] with
a curved, tapering shell,[9] and to be closely related to the gastropods (snails).[10] The similarity of the early
shelled cephalopod Plectronoceras to some gastropods was used in support of this view. The development
of a siphuncle would have allowed the shells of these early forms to become gas-filled (thus buoyant) in order
to support them and keep the shells upright while the animal crawled along the floor, and separated the true
cephalopods from putative ancestors such as Knightoconus, which lacked a siphuncle.[10] Negative
buoyancy (i.e. the ability to float) would have come later, followed by swimming in the Plectronocerida and
eventually jet propulsion in more derived cephalopods.[11] However, because chambered shells are found in
a range of molluscs – monoplacophora and gastropods as well as cephalopods – a siphuncle is essential to
ally a fossil shell conclusively to the cephalopoda.[9]:57 Chambered gastropods can be distinguished from
cephalopod shells by the absence of a siphuncle, the irregular spacing of septa, the layering of the shell and
(in younger or unmetamorphosed rocks) its microstructure, and the relatively thick width of the shell.[12] The
earliest such shells do not have the muscle scars which would be expected if they truly had a
monoplacophoran affinity.[9]:57 Additionally, the discovery that Nectocaris pteryx, which did not have a shell
and appeared to possess jet propulsion in the manner of "derived" cephalopods, complicated the question of
the order in which cephalopod features developed[2] – provided it is in fact a cephalopod and not an
arthropod.[13]
[edit] Early shelly record
Fossil orthoconic nautiloid from the Ordovician of Kentucky; an internal mold showing siphuncle and
half-filled camerae, both encrusted.
Understanding of early cephalopod origins is by necessity biased by the available fossil material,
which on the whole consists of shelly fossils. Critical fossils are detailed below; since their stratigraphic age
has guided the interpretation of the fossils,[14] they are listed in descending order of age.
[edit] Cambrian cephalopods

With the exception of the shelly genera Ectenolites and Eoclarkoceras, none of the 30+ Cambrian
cephalopod genera are known to have survived into the Ordovician.[15] Cambrian cephalopods differ from
their descendants by account of their small size (a few centimetres in length); long, tapering shells; smooth
shell surfaces; closely-spaced septa; and lack of deposits in their body chamber; several more specific
features are also only seen in certain groups of Cambrian cephalopod.[15]
[edit] Knightoconus
Knightoconus is a Cambrian monoplacophoran[16] thought to represent an ancestor to the
cephalopods. It had a chambered, conical shell, but lacked a siphuncle.[10] Although earlier molluscan
fossils are also septate, Knightoconus is the latest septate mollusc before the first sipunculate cephalopods –
a point that has been taken to prove its relevance to the Cephalopoda.[14] The absence of this siphuncle has
been taken as evidence against cephalopod ancestry – how, it is argued, could a siphuncle evolve to
penetrate existing septa? The prevailing argument suggests that a strand of tissue remained attached to the
previous septum as the mollusc moved forwards and deposited its next septum, producing an obstacle to the
complete closure of the septum and becoming mineralised itself.[17] 10 or more septa are found in mature
individuals, occupying around a third of the shell – septa form very early and have been found in specimens
as small as 2 mm in length.[14] Septa are uniformly spaced,[14] which is inconsistent with a gastropod
affinity.[12] Unlike monoplacophoran fossils, there is no evidence of muscle scarring in Knightoconus fossils.
[14]
[edit] Plectronoceras
Plectronoceras is arguably the earliest known crown-group cephalopod, dating to the Upper
Cambrian.[1] Its 14 known specimens hail from the basal Fengshan Formation (north-east China) of the
earliest Fengshanian stage.[6] None of the fossils are complete, and none show the tip or opening of the
shell.[6] Approximately half of its shell was filled with septa; 7 were recorded in a 2 cm shell.[14] Its shell
contains transverse septa separated by about half a millimetre, with a siphuncle on its concave side.[6] Its
morphology matches closely to that hypothesised for the last common ancestor of all cephalopods,[6] and
the Plectronocerida have been said to be the ancestors of the Ellesmerocerids, the first "true cephalopods".
[edit] Yochelcionellids
The Yochelcionellids have given rise to the "snorkel hypothesis". These fossils are aseptate
helcionellids with a snorkel-like tube on one surface. The snorkel has been seized upon as characteristic of a
cephalopod-like water circulatory system, or perhaps as a precursor to the siphuncle. However, neither of
these theories have been borne out.[14]
[edit] Ellesmerocerida
Sketch of the soft-part anatomy of early ellesmeroceridans

The earliest true cephalopod order to emerge was the Ellesmerocerida, which were quite small
organisms; their shells were slightly curved, and the internal chambers were closely spaced. The siphuncle
penetrated the septa with meniscus-like holes.[3] This marks an important difference from the earlier
cephalopods, whose siphuncle was positioned at the edge of the septum and against the shell wall.[14] On
the basis of muscle scars preserved in such genera as Paradakeoceras and Levisoceras, these animals are
reconstructed with a straight body and dorsal shell, with the head at the anterior, concave surface of the
shell, and the funnel – consisting of a pair of folds in the foot – at the rear — not juxtaposed with the head as in
later, oncocerid-like forms.[11]
[edit] Early Ordovician diversity

The Ellesmerocerids were the only shelled cephalopods that survived the end-Cambrian extinction,
and all subsequent cephalopods, which diversified throughout the Ordovician period, are thus thought to be
derived from these forms.
Early cephalopods had fine shells that could not cope with the pressures of deep water.[3] In the mid
Tremadoc, these were supplemented by larger shells around 20 cm in length; these larger forms included
straight and coiled shells, and fall into the orders Endocerida (with wide siphuncles) and Tarphycerida (with
narrow siphuncles).[3]
By the mid Ordovician these orders are joined by the Orthocerids, whose first chambers are small
and spherical, and Lituitids, whose siphuncles are thin. The Oncocerids also appear during this time; they are
restricted to shallow water and have short exogastric conchs.[3] The mid Ordovician saw the first
cephalopods with septa strong enough to cope with the pressures associated with deeper water, and could
inhabit depths greater than 100–200 m.[3] The wide-siphuncled Actinocerida and the Discocerida both
emerged during the Darriwilian.[3] The direction of coiling would prove to be crucial to the future success of
the lineages; endogastric coiling would only permit large size to be attained with a straight shell, whereas
exogastric coiling – initially rather rare – permitted the spirals familiar from the fossil record to develop, with
their corresponding large size and diversity.[17] (Endogastric mean the shell is curved so as the ventral or
lower side is longitudinally concave (belly in); exogastric means the shell is curved so as the ventral side is
longitudinally convex (belly out) allowing the funnel to be pointed backwards beneath the shell.)[17]
Curved shells brought a number of benefits. Firstly, minerals are not required in as large quantities,
as each successive whorl builds on the one before. Also, the organism is more stable (its centre of mass
coincides with its centre of buoyancy) and more manoeuvrable.[18]
Early cephalopods were likely predators near the top of the food chain.[19] In the Early Palaeozoic,
their range was far more restricted than today; they were mainly constrained to sub-littoral regions of shallow
shelves of the low latitudes, and usually occur in association with thrombolites.[20] A more pelagic habit was
gradually adopted as the Ordovician progressed.[20] Deep-water cephalopods, whilst rare, have been found
in the Lower Ordovician – but only in high-latitude waters.[20]
[edit] Fossils mistaken for cephalopods

A number of fossils have historically been considered to represent components of the cephalopods'
history, but been reinterpreted on the basis of additional material.
[edit] Volborthella
When it was discovered in 1888,[14] it was thought that the early Cambrian Volborthella was a
cephalopod. However discoveries of more detailed fossils showed that Volborthella’s small, conical shell was
not secreted but built from grains of the mineral silicon dioxide (silica); neither was it septate.[21] This illusion
was a result of the laminated texture of the organisms' tests.[22] Therefore, Volborthella’s classification is
now uncertain.[23]
[edit] Shelbyoceras
Because the characters differentiating monoplacophora from cephalopods are few, several
monoplacophora have been mistaken for cephalopod ancestors. One such genus is Shelbyoceras, which
was reclassified based on a depressed groove that forms a band around the shell, which is similar to a
feature seen in Hypseloconus.[24] The septa in this genus are either closely or irregularly spaced.[24]
[edit] Kirengellids
The Kirengellids are a group of shells that, whilst originally aligned to the monoplacophoran ancestry
of the cephalopods, have been reinterpreted as brachiopods.[25]
[edit] Coleoidea
An ammonitic ammonoid with the body chamber missing, showing the septal surface (especially at
right) with its undulating lobes and saddles.
The ancestors of coleoids (including most modern cephalopods) and the ancestors of the modern
nautilus, had diverged by the Floian Age of the Early Ordovician Period, over 470 million years ago. We know
this because the orthocerids were the first known representatives of the neocephalopoda,[26] were ultimately
the ancestors of ammonoids and coleoids, and had appeared by the Floian.[20] It is widely held that the
Bactritida, an Silurian–Triassic group of orthocones, are paraphyletic to the coleoids and ammonoids – that is,
the latter groups arose from within the Bactritida.[27]:393 An increase in the diversity of the coleoids and
ammonoids is observed around the start of the Devonian period, and corresponds with a profound increase
in fish diversity. This could represent the origin of the two derived groups.[27]
Unlike most modern cephalopods, most ancient varieties had protective shells. These shells at first
were conical but later developed into curved nautiloid shapes seen in modern nautilus species. It is thought
that competitive pressure from fish forced the shelled forms into deeper water, which provided an
evolutionary pressure towards shell loss and gave rise to the modern coleoids, a change which led to greater
metabolic costs associated with the loss of buoyancy, but which allowed them to recolonise shallow waters.
[10]:36 However, some of the straight-shelled nautiloids evolved into belemnites, out of which some evolved
into squid and cuttlefish.[verification needed] The loss of the shell may also have resulted from evolutionary
pressure to increase manoeuvrability, resulting in a more fish-like habit.[28]:289 This pressure may have
increased as a result of the increased complexity of fish in the late Palaeozoic, increasing the competitive
pressure.[28]:289 Internal shells still exist in many non-shelled living cephalopod groups but most truly
shelled cephalopods, such as the ammonites, became extinct at the end of the Cretaceous.
[edit] Organ origins
The tentacles of the ancestral cephalopod developed from the mollusc's foot;[29] the ancestral state
is thought to have had five pairs of tentacles which surrounded the mouth.[29] Smell-detecting organs
evolved very early in the cephalopod lineage.[29]
The earliest cephalopods,[note 1] like Nautilus and some coeloids, appeared to be able to propel
themselves forwards by directing their jet backwards.[28]:289 Because they had an external shell, they would
not have been able to generate their jets by contracting their mantle, so must have used alternate methods:
such as by contracting their funnels or moving the head in and out of the chamber.[28]:289
[edit] Exceptional preservation

The preservation of cephalopod soft parts is not entirely unusual; soft-bodied fossils, especially of
coeloids (squid), are relatively widespread in the Jurassic,[30] but phosphatized remains are unknown before
this period.[31] On the other hand, soft parts – including a possible ink sac — are known from the Paleozoic
Hunsrück Slate and Francis Creek shale.[32]
[edit] Notes
1. ^ Ordovician orthocone nautiloids are the first for which trace fossil evidence is available
[edit] References
1. ^ a b Dzik, J. (1981). "Origin of the cephalopoda" (PDF). Acta Palaeontologica Polonica 26
(2): 161–191. http://www.paleo.pan.pl/people/Dzik/Publications/Cephalopoda.pdf.
2. ^ a b Smith, M. R.; Caron, J. B. (2010). "Primitive soft-bodied cephalopods from the
Cambrian". Nature 465 (7297): 469. doi:10.1038/nature09068. PMID 20505727.
Bibcode: 2010Natur.465..469S. http://individual.utoronto.ca/martinsmith/nectocaris.html. edit
3. ^ a b c d e f g Kröger, B.; Yun-bai, Y. B. (2009). "Pulsed cephalopod diversification during the
Ordovician". Palaeogeography Palaeoclimatology Palaeoecology 273: 174–201.
doi:10.1016/j.palaeo.2008.12.015. edit
4. ^ Begtson, Stefan (1970). "The Lower Cambrian fossil Tommotia". Lethaia 3 (4): 363–392.
doi:10.1111/j.1502-3931.1970.tb00829.x.
5. ^ Landing, Ed; Kröger, Björn (2009). "The Oldest Cephalopods from East Laurentia". Journal
of Paleontology 83: 123–127. doi:10.1666/08-078R.1.
6. ^ a b c d e Clarke, M.R.; Trueman, E.R., ed (1988). "Main features of cephalopod evolution".
The Mollusca. 12: Palaeontology and Neontology of Cephalopods. Orlando, Fla.: Acad. Pr..
ISBN 0127514120.
7. ^ Giribet; Okusu, A; Lindgren, A.R.; Huff, S.W.; Schrödl, M; Nishiguchi, M.K. (May 2006).
"Evidence for a clade composed of molluscs with serially repeated structures: monoplacophorans are
related to chitons" (Free full text). Proceedings of the National Academy of Sciences of the United
States of America 103 (20): 7723–7728. doi:10.1073/pnas.0602578103. PMID 16675549.
PMC 1472512. http://www.pnas.org/cgi/pmidlookup?view=long&pmid=16675549. edit
8. ^ Lemche, H; Wingstrand, K.G. (1959). "The anatomy of Neopilina galatheae Lemche, 1957
(Mollusca, Tryblidiacea)." (Link to free full text + plates). Galathea Rep. 3: 9–73.
http://www.zmuc.dk/inverweb/Galathea/index.html.
9. ^ a b c Wingstrand, KG (1985). "On the anatomy and relationships of Recent
Monoplacophora" (Link to free full text + plates). Galathea Rep. 16: 7–94.
http://www.zmuc.dk/inverweb/Galathea/Galathea_p5.html.
10.^ a b c d "Origin and Evolution". Cephalopods – ecology and fisheries. 2005. pp. 36.
doi:10.1002/9780470995310.ch3. ISBN 0632060484.
11.^ a b Kroger, B. (2007). "Some Lesser Known Features of the Ancient Cephalopod Order
Ellesmerocerida (nautiloidea, Cephalopoda)". Palaeontology 50 (3): 565–572. doi:10.1111/j.1475-
4983.2007.00644.x.
12.^ a b Otto P. Majewske (1974). Recognition of Invertebrate Fossil Fragments in Rocks and
Thin Sections. Leiden, Netherlands: E.J. Brill. p. 76. http://books.google.com/?
id=m8gUAAAAIAAJ&pg=PA76.
13.^ Mazurek, D.; Zatoń, M. (2011). "Is Nectocaris pteryx a cephalopod?". Lethaia.
doi:10.1111/j.1502-3931.2010.00253.x. edit
14.^ a b c d e f g h i Webers, G. F.; Yochelson, E. L. (1989). "Late Cambrian molluscan faunas
and the origin of the Cephalopoda". In Crame, J. A.. Origins and Evolution of the Antarctic Biota . 47.
Geological Society, London: Special Publications. pp. 29. doi:10.1144/GSL.SP.1989.047.01.04. edit
15.^ a b Chen, J. Y.; Teichert, C. (1983). "Cambrian cephalopods". Geology 11: 647-650.
doi:10.1130/0091-7613(1983)11<647:CC>2.0.CO;2. ISSN 0091-7613. edit
16.^ doi:10.1111/j.1502-3931.1973.tb01199.x
17.^ a b c Holland, C. H. (1987). "The nautiloid cephalopods: a strange success: President's
anniversary address 1986". Journal of the Geological Society 144: 1–0.
doi:10.1144/gsjgs.144.1.0001. edit
18.^ Wells, M.J.; O'Dor, R.K.. "Jet Propulsion and the Evolution of the Cephalopods". Bulletin of
Marine Science, Volume 49, Number 1, July 1991 , pp. 419-432(14).
19.^ Boyle, Peter; Rodhouse, Paul (2004). Cephalopods : ecology and fisheries. Ames, Iowa:
Blackwell. doi:10.1002/9780470995310.ch2. ISBN 0632060484. http://books.google.com/?
id=4UtCi2B4VnoC.
20.^ a b c d Kröger, B. R.; Servais, T.; Zhang, Y.; Kosnik, M. (2009). "The Origin and Initial Rise
of Pelagic Cephalopods in the Ordovician". PLoS ONE 4 (9): e7262.
http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0007262. Retrieved 2009-
10-30. edit
21.^ Lipps, J. H.; Sylvester, A. G. (1 March 1968). "The Enigmatic Cambrian Fossil Volborthella
and Its Occurrence in California". Journal of Paleontology (Journal of Paleontology, Vol. 42, No. 2) 42
(2): 329–336. ISSN 00223360. http://jstor.org/stable/1302218. edit
22.^ "Lower Cambrian fossil Volborthella: the whole truth or just a piece of the beast?". Geology
21: 805. 1993. doi:10.1130/0091-7613(1993)021<0805:LCFVTW>2.3.CO;2. edit
23.^ Hagadorn, J.W.; Waggoner, B.M. (2002). "The Early Cambrian problematic fossil
Volborthella: New insights from the Basin and Range". In Corsetti, F.A.. Proterozoic-Cambrian of the
Great Basin and Beyond, Pacific Section SEPM Book 93 . SEPM (Society for Sedimentary Geology).
pp. 135–150. http://www3.amherst.edu/~jwhagadorn/publications/volb.pdf
24.^ a b Stinchcomb, B. L. (1980). "New Information on Late Cambrian Monoplacophora
Hypseloconus and Shelbyoceras (Mollusca)". Journal of Paleontology (Paleontological Society) 54
(1): 45–49. http://jstor.org/stable/1304159. edit
25.^ Dzik, Jerzy (2010). "BRACHIOPOD IDENTITY OF THE ALLEGED MONOPLACOPHORAN
ANCESTORS OF CEPHALOPODS". Malacologia %V 52 %N 1 %P 97-113.
http://www.paleo.pan.pl/people/Dzik/Publications/Angarella.pdf.
26.^ Kröger, Björn (2006). "Early growth-stages and classification of orthoceridan Cephalopods
of the Darriwillian (Middle Ordovician) of Baltoscandia". Lethaia 39 (2): 129–139.
doi:10.1080/00241160600623749. http://www3.interscience.wiley.com/journal/119918421/abstract.
27.^ a b Young, R.E.; Vecchione, M.; Donovan, D.T.. "The evolution of coleoid cephalopods and
their present biodivesity and ecology". In Payne, AIL; Lipin'ski, M.R.; Clarke, M.R.; Roeleveld, M.A.C.
Cephalopod biodiversity, ecology & evolution. South Afriocan journal of Marine Sciences. 20.
pp. 393–420.
28.^ a b c d Wilbur, Karl M.; Trueman, E.R.; Clarke, M.R., eds. (1985), The Mollusca, 11. Form
and Function, New York: Academic Press, ISBN 0-12-728702-7
29.^ a b c Shigeno, S.; Sasaki, T.; Moritaki, T.; Kasugai, T.; Vecchione, M.; Agata, K. (2008).
"Evolution of the cephalopod head complex by assembly of multiple molluscan body parts: Evidence
from Nautilus Embryonic Development". Journal of Morphology 269 (1): 1–17.
doi:10.1002/jmor.10564. PMID 17654542.
30.^ Kear, A.J.; Briggs, D.E.G.; Donovan, D.T. (1995). "Decay and fossilization of non-
mineralized tissue in coleoid cephalopods". Palaeontology 38 (1): 105–132.
http://palaeontology.palass-pubs.org/pdf/Vol%2038/Pages%20105-131.pdf. Retrieved 2009-04-21.
31.^ Briggs, D. E. G.; Kear, A. J.; Martill, D. M.; Wilby, P. R. (1993). "Phosphatization of soft-
tissue in experiments and fossils". Journal of the Geological Society 150: 1035–1033.
doi:10.1144/gsjgs.150.6.1035. edit
32.^ Allison, P. A. (1987). "A new cephalopod with soft parts from the Upper Carboniferous
Francis Creek Shale of Illinois, USA". Lethaia 20 (78): 117–121. doi:10.1111/j.1502-
3931.1987.tb02028.x. http://www3.interscience.wiley.com/journal/120019493/abstract. edit

• Chen, J. Y.; Teichert, C. (1983). "Cambrian cephalopods". Geology 11: 647-650.
doi:10.1130/0091-7613(1983)11<647:CC>2.0.CO;2. ISSN 0091-7613. edit
[hide]v · d · eNotable fossil cephalopods (listed by first occurrence)

Cenozoic The belemnite-cuttlefish-like Belosaepia
• advanced nautiloids; Nautilaceae • true

ammonites;
Mesozoic • early coleoids: Proteroctopus, Styletoctopus,
Keuppia, Palaeoctopus, Paleocirroteuthis
• ceratitid ammonoids; • late Nautilida Evolutionary history of
cephalopods
• Long-lasting lineages: Belemnites • Goniatite

and ceratite ammonoids • Probable coleoid ancestor
Bactritida
• Early forms: the earliest cephalopod group
Palaeozoic Ellesmerocerida • Orthocones: Endocerida,
Orthocerida, Actinocerida • Brevicones: Ascocerida,
Oncocerida • Earliest coiled cephalopods:
Tarphycerida • Spirulid?: Shimanskya
• Octopus: Pohlsepia
Cambrian • earliest unambiguous cephalopods;

Plectronocerida • Plectronoceras
• Monoplacophoran-like ancestral forms: •
Knightoconus
• possible, nude, "stem-group cephalopods":
Nectocarididae
Retrieved from "http://en.wikipedia.org/wiki/Evolutionary_history_of_cephalopods"
Categories: Extinct cephalopods | Evolution by taxon | Cephalopods

W000
Evolutionary history of brachiopods


The origin of the brachiopods is uncertain; they either arose from reduction of a multi-plated tubular
organism, or from the folding of a slug-like organism with a protective shell on either end. Since their
Cambrian origin, the phylum rose to a Palaeozoic dominance, but dwindled during the Mesozoic.
Contents
[hide]
• 1 Origins
• 1.1 Brachiopod fold hypothesis
• 1.2 Tommotiids
• 1.3 Crown group
• 2 Mineralization
• 3 Palaeozoic dominance
• 4 Mesozoic decline
• 5 References
[edit] Origins
[edit] Brachiopod fold hypothesis
The long-standing hypothesis of brachiopod origins, which has recently come under fire,[1] suggests
that the brachiopods arose by the folding of a Halkieria-like organism, which bore two protective shells at
either end of a scaled body.[2] Under this hypothesis, the Phoronid worms share a similar evolutionary
history; molecular data also appear to indicate their membership of Brachiopoda.[3]
Under the Brachiopod Fold Hypothesis, the "dorsal" and "ventral" valves would in fact represent an
anterior and posterior shell. This would make the axes of symmetry consistent with that of other bilaterian
phyla[3] and appears to be consistent with the embryological development, in which the body axis folds to
bring the shells from the dorsal surface to their mature position.[3] However the axis of folding varies
somewhat between phoronids and brachiopods, and in articulate brachiopods the folding is reflected by the
reversal of the mantle in embryos.[3] Nevertheless, the embryological origins of homologous features often
appear significantly different, simply as a result of changes in developmental timing.[4]
[edit] Tommotiids
An alternative to the BFH suggests that brachiopods arose through the shortening of a tube-like
organism consisting of many shell plates. It is possible that they arose from within the tommotiid group in this
fashion.[5] The more derived tommotiid Paterimitra has a pair of brachiopod-like shells at its rear, in just the
arrangement one would expect of a brachiopod.[1] This is supported by the similarities in mineralogy
between the Tommotiids and the earliest brachiopods.[6]
[edit] Crown group

The earliest unequivocal brachiopod fossils appeared in the early Cambrian Period.[7][8] The oldest
known brachiopod is the Aldanotreta sunnaginensis from the lowest Tommotian Stage, early Cambrian of the
Siberia was confidently identified as a paterinid linguliforms.
[edit] Mineralization
The Lingiliformea brachiopods have apatite shells, which contrasts with the calcitic exoskeleta of the
other two brachiopod subphyla. This split occurred very early - the earliest brachiopod assemblages, from the
Tommotian, already contain both apatite- and calcite-secreting organisms.[6] Since the minerals used to form
exoskeleta rarely change,[9] one might expect these two forms to represent two discrete lineages - but in
fact, early brachiopods used a wide range of techniques and materials in shell construction, drawing from
phosphatic, calcitic and organic building blocks, and sometimes employing all three.[6] Deducing the original
method of mineralisation is tricky; however, it appears that the tommotiids - probably the closest stem group
to the brachiopoda, assuming that the Brachiopod Fold Hypothesis is false - produced the same shell
microstructre as the earliest known brachiopods. Their shells had a relatively high concentration of
phosphate and organic material, though this decreased over time.[6]
[edit] Palaeozoic dominance

Brachiopods are extremely common fossils throughout the Palaeozoic. During the Ordovician and
Silurian periods, brachiopods became adapted to life in most marine environments and became particularly
numerous in shallow water habitats, in some cases forming whole banks in much the same way as bivalves
(such as mussels) do today. In some places, large sections of limestone strata and reef deposits are
composed largely of their shells.
The major shift came with the Permian extinction, as a result of the Mesozoic marine revolution.
Before the extinction event, brachiopods were more numerous and diverse than bivalve mollusks.
Afterwards, in the Mesozoic, their diversity and numbers were drastically reduced and they were largely
replaced by bivalve molluscs. Molluscs continue to dominate today, and the remaining orders of brachiopods
survive largely in fringe environments.
[edit] Mesozoic decline

Throughout their long geological history, the brachiopods have gone through several major
proliferations and diversifications, and have also suffered from major extinctions as well.
It has been suggested that the slow decline of the brachiopods over the last 100 million years or so is
a direct result of the rise in diversity of filter feeding bivalves, which have ousted the brachiopods from their
former habitats; however, the bivalves have undergone a steady rise in diversity from the mid-Paleozoic
onwards, and their abundance is unrelated to that of the brachiopods; further, many bivalves occupy niches
(e.g. burrowing) which brachiopods never inhabited.[10]
Alternative possibilities for their demise include the increasing disturbance of sediments by roving
deposit feeders (including many burrowing bivalves); the increased intensity and variety of shell-crushing
predation; or even chance demise - they were hard hit in the End-P extinction and may simply never have
recovered.
[edit] References
1. ^ a b "The scleritome of Paterimitra: an Early Cambrian stem group brachiopod from South
Australia". Proceedings of the Royal Society B: Biological Sciences 276 (1662): 1651–1656. May
2009. doi:10.1098/rspb.2008.1655. ISSN 0962-8452. PMID 19203919. edit
2. ^ Sigwart; Sutton, M. D. (Oct 2007). "Deep molluscan phylogeny: synthesis of
palaeontological and neontological data". Proceedings of the Royal Society B: Biological sciences
274 (1624): 2413–2419. doi:10.1098/rspb.2007.0701. PMID 17652065. For a summary, see "The
Mollusca". University of California Museum of Paleontology.
http://www.ucmp.berkeley.edu/taxa/inverts/mollusca/mollusca.php. Retrieved 2008-10-02. edit
3. ^ a b c d Cohen, B. L.; Holmer, L. E.; Luter, C. (2003). "The brachiopod fold: a neglected body
plan hypothesis". Palaeontology 46: 59. doi:10.1111/1475-4983.00287. edit
4. ^ Budd, G. E.; Jensen, S. (2000). "A critical reappraisal of the fossil record of the bilaterian
phyla". Biological reviews of the Cambridge Philosophical Society 75 (2): 253–95.
doi:10.1017/S000632310000548X. PMID 10881389. edit
5. ^ Skovsted, C. B.; Brock, G. A.; Paterson, J. R.; Holmer, L. E.; Budd, G. E. (2008). "The
scleritome of Eccentrotheca from the Lower Cambrian of South Australia: Lophophorate affinities and
implications for tommotiid phylogeny". Geology 36: 171. doi:10.1130/G24385A.1. edit
6. ^ a b c d Balthasar, Uwe (August 2009). "The Evolution of Shell Composition in Brachiopods".
In Smith, Martin R.; O'Brien, Lorna J.; Caron, Jean-Bernard. International Conference on the
Cambrian Explosion (Walcott 2009). Abstract Volume. Toronto, Ontario, Canada: The Burgess Shale
Consortium. 31st July 2009. ISBN 978-0-9812885-1-2. http://burgess-shale.info/abstract/balthasar.
7. ^ Alwyn Williams, Leonid E. Popov, Lars E. Holmer, Maggie Cusack (1998). "The diversity
and phylogeny of the paterinate Brachiopods". Palaeontology 41 (2): 241–262.
http://palaeontology.palass-pubs.org/pdf/Vol%2041/Pages%20221-262.pdf.
8. ^ Valentine, James W. (2004). On the origin of phyla. Chicago: University of Chicago Press.
pp. 638. ISBN 0-226-84548-6. http://books.google.com/?id=DMBkmHm5fe4C&pg=PA354.
9. ^ Porter, S. M. (Jun 2007). "Seawater chemistry and early carbonate biomineralization".
Science 316 (5829): 1302–1301. doi:10.1126/science.1137284. ISSN 0036-8075. PMID 17540895.
edit
10.^ Gould, S. J.; Calloway, C. B. (1 October 1980). "Clams and Brachiopods — Ships that Pass
in the Night". Paleobiology (Paleobiology, Vol. 6, No. 4) 6 (4): 383–396. ISSN 00948373.
http://jstor.org/stable/2400538. edit
Retrieved from "http://en.wikipedia.org/wiki/Evolutionary_history_of_brachiopods"
Categories: Brachiopods
W000
Timeline of evolution

For the history of evolutionary biology, see History of evolutionary thought.
This article is about the timeline of evolution in a general aspect. See also the timeline of human
evolution.
This timeline of the evolution of life outlines the major events in the development of life on the planet
Earth (See Organism). For a thorough explanatory context, see the history of Earth, and geologic time scale.
The dates given in this article are estimates based on scientific evidence.
In biology, evolution is the process by which populations of organisms acquire and pass on novel
traits from generation to generation. Its occurrence over large stretches of time explains the origin of new
species and ultimately the vast diversity of the biological world. Contemporary species are related to each
other through common descent, products of evolution and speciation over billions of years.
Contents
[hide]
• 1 Basic timeline
• 2 Detailed timeline
• 2.1 Hadean Eon
• 2.2 Archean Eon
• 2.3 Proterozoic Eon
• 2.4 Phanerozoic Eon
• 2.4.1 Paleozoic Era
• 2.4.2 Mesozoic Era
• 2.4.3 Cenozoic Era
• 3 See also
• 5 References
• 6 External links
[edit] Basic timeline
Life on Earth
view • discuss • edit
-4500 —
–
-4000 —
–
-3500 —
–
-3000 —
–
-2500 —
–
-2000 —
–
-1500 —
–
-1000 —
–
-500 —
–
0—
Life?
Photosynthesis?
Eukaryotes
Complex multicellular life
Animals
Land plants
"Dinosaurs"
Mammals
Flowers
←
Formation
of Earth
←
Meteorite bombardment
←
Atmospheric oxygen
←
Ediacara biota
←
Cambrian explosion
←
Modern-looking
humans
Axis scale: millions of years ago.
Dates prior to 1 billion years ago are speculative.
The basic timeline is a 4.5 billion year old Earth, with (very approximate) dates:
• 3.8 billion years of simple cells (prokaryotes),
• 3 billion years of photosynthesis,
• 2 billion years of complex cells (eukaryotes),
• 1 billion years of multicellular life,
• 600 million years of simple animals,
• 570 million years of arthropods (ancestors of insects, arachnids and crustaceans),
• 550 million years of complex animals,
• 500 million years of fish and proto-amphibians,
• 475 million years of land plants,
• 400 million years of insects and seeds,
• 360 million years of amphibians,
• 300 million years of reptiles,
• 200 million years of mammals,
• 150 million years of birds,
• 130 million years of flowers,
• 65 million years since the non-avian dinosaurs died out,
• 2.5 million years since the appearance of the genus Homo,
• 200,000 years since humans started looking like they do today,
• 25,000 years since Neanderthals died out.
[edit] Detailed timeline
Ma, ("megaannum") means "million years ago". ka means "thousand years ago" and ya
means "years ago"
[edit] Hadean Eon

3800 Ma and earlier.
Date Event
4600 Ma The planet Earth forms from the accretion disc

revolving around the young Sun.
4500 Ma According to one plausible theory, the planet Earth

and the planet Theia collide, sending a very large number
of moonlets into orbit around the young Earth. These
moonlets eventually coalesce to form the Moon.[1] The
gravitational pull of the new Moon stabilises the Earth's
fluctuating axis of rotation and sets up the conditions in
which life formed.[2]
4100 Ma The surface of the Earth cools enough for the crust
to solidify. The atmosphere and the oceans form.[3] PAH
infall,[4] and iron sulfide synthesis along deep ocean
platelet boundaries, may have led to the RNA world of
competing organic compounds.
Between 4500 and 3500 Ma The earliest life appears, possibly derived from
self-reproducing RNA molecules.[5][6] The replication of
these organisms requires resources like energy, space,
and smaller building blocks, which soon become limited,
resulting in competition, with natural selection favouring
those molecules which are more efficient at replication.
DNA molecules then take over as the main replicators and
these archaic genomes soon develop inside enclosing
membranes which provide a stable physical and chemical
environment conducive to their replication: proto-cells.[7]
[8][9]
3900 Ma Late Heavy Bombardment: peak rate of impact

events upon the inner planets by meteoroids. This
constant disturbance may have obliterated any life that
had evolved to that point, or possibly not, as some early
microbes could have survived in hydrothermal vents below
the Earth's surface;[10] or life might have been transported
to Earth by a meteoroid.[11]
Somewhere between 3900 and 2500 Ma Cells resembling prokaryotes appear.[12] These
first organisms are chemoautotrophs: they use carbon
dioxide as a carbon source and oxidize inorganic materials
to extract energy. Later, prokaryotes evolve glycolysis, a
set of chemical reactions that free the energy of organic
molecules such as glucose and store it in the chemical
bonds of ATP. Glycolysis (and ATP) continue to be used in
almost all organisms, unchanged, to this day.[13][14]
[edit] Archean Eon

3800 Ma – 2500 Ma
Date Event
3500 Ma Lifetime of the last universal ancestor;[15][16] the split between bacteria and
archaea occurs.[17]
Bacteria develop primitive forms of photosynthesis which at first do not produce
oxygen.[18] These organisms generate ATP by exploiting a proton gradient, a mechanism
still used in virtually all organisms.
3000 Ma Photosynthesizing cyanobacteria evolve; they use water as a reducing agent,

thereby producing oxygen as waste product.[19] More recent research, however, suggests
a later time of 2700 Ma. The oxygen initially oxidizes dissolved iron in the oceans, creating
iron ore. The oxygen concentration in the atmosphere subsequently rises, acting as a
poison for many bacteria. The moon is still very close to the earth and causes tides
1,000 feet (305 m) high. The earth is continually wracked by hurricane force winds. These
extreme mixing influences are thought to stimulate evolutionary processes. (See Oxygen
catastrophe)
2700 Ma Timeframe of cyanobacteria evolution suggested by more recent research.
[edit] Proterozoic Eon

2500 Ma – 542 Ma
Date Event
By 1850 Ma Eukaryotic cells appear.[20][21] Eukaryotes contain membrane-bound

organelles with diverse functions, probably derived from prokaryotes engulfing each
other via phagocytosis. (See Endosymbiosis)
By 1200 Ma Sexual reproduction first appears, increasing the rate of evolution.[22]
1200 Ma Simple multicellular organisms evolve, mostly consisting of cell colonies of

limited complexity.
850–630 Ma A global glaciation may have occurred.[23][24] Opinion is divided on whether it

increased or decreased biodiversity or the rate of evolution.[25][26][27]
580–542 Ma The Ediacaran biota represent the first large, complex multicellular organisms -
although their affinities remain a subject of debate.[28]
580–500 Ma Most modern phyla of animals begin to appear in the fossil record during the
Cambrian explosion.[29][30]
580–540 Ma The accumulation of atmospheric oxygen allows the formation of an ozone

layer.[31] This blocks ultraviolet radiation, permitting the colonisation of the land.[31]
560 Ma Earliest fungi
[edit] Phanerozoic Eon

542 Ma – present
The Phanerozoic Eon, literally the "period of well-displayed life", marks the appearance in the fossil
record of abundant, shell-forming and/or trace-making organisms. It is subdivided into three eras, the
Paleozoic, Mesozoic and Cenozoic, which are divided by major mass extinctions.
[edit] Paleozoic Era

542 Ma – 251.0 Ma
Date Event
535 Ma Major diversification of living things in the oceans: chordates, arthropods (e.g.
trilobites, crustaceans), echinoderms, mollusks, brachiopods, foraminifers and
radiolarians, etc.
530 Ma The first known footprints on land date to 530 Ma, indicating that early animal
explorations may have predated the development of terrestrial plants.[32]
525 Ma Earliest graptolites.
510 Ma First cephalopods (Nautiloids) and chitons.
505 Ma Fossilization of the Burgess Shale.
485 Ma First vertebrates with true bones (jawless fishes).
450 Ma Land arthropod burrows (millipedes) appear, along with the first complete
conodonts and echinoids.
440 Ma First agnathan fishes: Heterostraci, Galeaspida, and Pituriaspida.
434 Ma The first primitive plants move onto land,[33][citation needed] having evolved from
green algae living along the edges of lakes.[34] They are accompanied by fungi[citation
needed], which may have aided the colonization of land through symbiosis.
420 Ma Earliest ray-finned fishes, trigonotarbid arachnids, and land scorpions.
410 Ma First signs of teeth in fish. Earliest nautiid nautiloids, lycophytes, and
trimerophytes.
395 Ma First lichens, stoneworts. Earliest harvestman, mites, hexapods (springtails), and
ammonoids.
363 Ma By the start of the Carboniferous Period, the Earth begins to be recognisable.
Insects roamed the land and would soon take to the skies; sharks swam the oceans as
top predators,[35] and vegetation covered the land, with seed-bearing plants and forests
soon to flourish.
Four-limbed tetrapods gradually gain adaptations which will help them occupy a
terrestrial life-habit.
360 Ma First crabs and ferns. Land flora dominated by seed ferns.
350 Ma First large sharks, ratfishes, and hagfish.
340 Ma Diversification of amphibians.
330 Ma First amniote vertebrates (Paleothyris).
305 Ma Earliest diapsid reptiles (e.g. Petrolacosaurus).
280 Ma Earliest beetles, seed plants and conifers diversify while lepidodendrids and
sphenopsids decrease. Terrestrial temnospondyl amphibians and pelycosaurs (e.g.
Dimetrodon) diversify in species.
251.4 Ma The Permian-Triassic extinction event eliminates over 90-95% of marine species.
Terrestrial organisms were not as seriously affected as the marine biota. This "clearing of
the slate" may have led to an ensuing diversification, but life on land took 30M years to
completely recover.[36]
[edit] Mesozoic Era
Date Event
From 251.4 Ma The Mesozoic Marine Revolution begins: increasingly well-adapted and
diverse predators pressurise sessile marine groups; the "balance of power" in the
oceans shifts dramatically as some groups of prey adapt more rapidly and
effectively than others.
245 Ma Earliest ichthyosaurs.
240 Ma Increase in diversity of gomphodont cynodonts and rhynchosaurs.
225 Ma Earliest dinosaurs (prosauropods), first cardiid bivalves, diversity in cycads,

bennettitaleans, and conifers. First teleost fishes.
215 Ma First mammals (e.g. Eozostrodon), minor vertebrate extinctions occur
220 Ma
Eoraptor, among the earliest dinosaurs, appeared in the fossil record 230
million years ago.
Gymnosperm forests dominate the land; herbivores grow to huge sizes in
order to accommodate the large guts necessary to digest the nutrient-poor plants.
[citation needed], first flies and turtles (Odontochelys). First Coelophysoid dinosaurs
200 Ma The first accepted evidence for viruses (at least, the group Geminiviridae)
exists.[37] Viruses are still poorly understood and may have arisen before "life"
itself, or may be a more recent phenomenon.
Major extinctions in terrestrial vertebrates and large amphibians. Earliest
examples of Ankylosaurian dinosaurs
195 Ma First pterosaurs with specialized feeding (Dorygnathus). First sauropod

dinosaurs. Diversification in small, ornithischian dinosaurs: heterodontosaurids,
fabrosaurids, and scelidosaurids.
190 Ma Pliosaurs appear in the fossil record. First lepidopteran insects

(Archaeolepis), hermit crabs, modern starfish, irregular echinoids, corbulid bivalves,
and tubulipore bryozoans. Extensive development of sponge reefs.
176 Ma First members of the Stegosauria group of dinosaurs
170 Ma Earliest salamanders, newts, cryptoclidid & elasmosaurid plesiosaurs, and

cladotherian mammals. Cynodonts become extinct while sauropod dinosaurs
diversify.
165 Ma First rays and glycymeridid bivalves.
161 Ma Ceratopsian dinosaurs appear in the fossil record (Yinlong)
155 Ma First blood-sucking insects (ceratopogonids), rudist bivalves, and

cheilosome bryozoans. Archaeopteryx, a possible ancestor to the birds, appears in
the fossil record, along with triconodontid and symmetrodont mammals. Diversity in
stegosaurian and theropod dinosaurs.
130 Ma The rise of the Angiosperms: These flowering plants boast structures that
attract insects and other animals to spread pollen. This innovation causes a major
burst of animal evolution through co-evolution. First freshwater pelomedusid turtles.
120 Ma Oldest fossils of heterokonts, including both marine diatoms and

silicoflagellates.
115 Ma First monotreme mammals.
110 Ma First hesperornithes, toothed diving birds. Earliest limopsid, verticordiid, and
thyasirid bivalves.
106 Ma Spinosaurus, the largest theropod dinosaur, appears in the fossil record.
100 Ma Earliest bees.
90 Ma Extinction of ichthyosaurs. Earliest snakes and nuculanid bivalves. Large
diversification in angiosperms: magnoliids, rosids, hamamelidids, monocots, and
ginger. Earliest examples of ticks.
80 Ma First ants and termites.
70 Ma Multituberculate mammals increase in diversity. First yoldiid bivalves.
68 Ma Tyrannosaurus, the largest terrestrial predator of North America appears in

the fossil record. First species of Triceratops.
[edit] Cenozoic Era

65.5 Ma – present
Date Event
65.5 Ma The Cretaceous–Tertiary extinction event eradicates about half of all animal
species, including mosasaurs, pterosaurs, plesiosaurs, ammonites, belemnites,
rudist and inoceramid bivalves, most planktic foraminifers, and all of the dinosaurs
excluding their descendants the birds [38]
From 65 Ma Rapid dominance of conifers and ginkgos in high latitudes, along with
mammals becoming the dominant species. First psammobiid bivalves. Rapid
diversification in ants.
63 Ma Evolution of the creodonts, an important group of carnivorous mammals.
60 Ma Diversification of large, flightless birds. Earliest true primates, along with the
first semelid bivalves, edentates, carnivorous and lipotyphlan mammals, and owls.
The ancestors of the carnivorous mammals (miacids) were alive.
56 Ma Gastornis, a large, flightless bird appears in the fossil record, becoming an

apex predator at the time.
55 Ma Modern bird groups diversify (first song birds, parrots, loons, swifts,
woodpeckers), first whale (Himalayacetus), earliest rodents, lagomorphs, armadillos,
appearance of sirenians, proboscideans, perissodactyl and artiodactyl mammals in
the fossil record. Angiosperms diversify. The ancestor (according to theory) of the
species in Carcharodon, the early mako shark Isurus hastalis, is alive.
52 Ma First bats appear (Onychonycteris).
50 Ma Peak diversity of dinoflagellates and nanofossils, increase in diversity of

anomalodesmatan and heteroconch bivalves, brontotheres, tapirs, rhinoceroses, and
camels appear in the fossil record, diversification of primates.
40 Ma Modern type butterflies and moths appear. Extinction of Gastornis.

Basilosaurus, one of the first of the giant whales, appeared in the fossil record.
37 Ma First Nimravid carnivores ("False Saber-toothed Cats") - these species are
unrelated to modern-type felines
35 Ma Grasses evolve from among the angiosperms; grasslands begin to expand.
Slight increase in diversity of cold-tolerant ostracods and foraminifers, along with
major extinctions of gastropods, reptiles, and amphibians. Many modern mammal
groups begin to appear: first glyptodonts, ground sloths, dogs, peccaries, and the first
eagles and hawks. Diversity in toothed and baleen whales.
33 Ma Evolution of the thylacinid marsupials (Badjcinus).
30 Ma First balanids and eucalypts, extinction of embrithopod and brontothere

mammals, earliest pigs and cats.
28 Ma Paraceratherium appears in the fossil record, the largest terrestrial mammal

that ever lived.
25 Ma First deer.
20 Ma First giraffes and giant anteaters, increase in bird diversity.
15 Ma Mammut appears in the fossil record, first bovids and kangaroos, diversity in
Australian megafauna.
10 Ma Grasslands and savannas are established, diversity in insects, especially

ants and termites, horses increase in body size and develop high-crowned teeth,
major diversification in grassland mammals and snakes.
6.5 Ma First hominin (Sahelanthropus).

6 Ma Australopithecines diversify (Orrorin, Ardipithecus)
5 Ma First tree sloths and hippopotami, diversification of grazing herbivores, large

carnivorous mammals, burrowing rodents, kangaroos, birds, and small carnivores,
vultures increase in size, decrease in the number of perissodactyl mammals.
Extinction of Nimravid carnivores
4.8 Ma Mammoths appear in the fossil record.
4 Ma Evolution of Australopithecus, Stupendemys appears in the fossil record as

the largest freshwater turtle.
3 Ma The Great American Interchange, where various land and freshwater faunas
migrated between North and South America. Armadillos, opossums, hummingbirds,
and vampire bats traveled to North America while horses, tapirs, saber-toothed cats,
and deer entered South America. The first short-faced bears ( Arctodus) appear.
2.7 Ma Evolution of Paranthropus
2.5 Ma The earliest species of Smilodon evolve
2 Ma First members of the genus Homo appear in the fossil record. Diversification
of conifers in high latitudes. The eventual ancestor of cattle, Bos primigenius evolves
in India
1.7 Ma Extinction of australopithecines.

1.2 Ma Evolution of Homo antecessor. The last members of Paranthropus die out.
600 ka Evolution of Homo heidelbergensis
350 ka Evolution of Neanderthals
300 ka Gigantopithecus, a giant relative of the orangutan dies out from Asia
200 ka Anatomically modern humans appear in Africa.[39][40][41] Around 50,000
years before present they start colonising the other continents, replacing the
Neanderthals in Europe and other hominins in Asia.
40 ka The last of the giant monitor lizards ( Megalania) die out
30 ka Extinction of Neanderthals
15 ka The last Woolly rhinoceros (Coelodonta) are believed to have gone extinct
11 ka The giant short-faced bears (Arctodus) vanish from North America, with the
last Giant Ground Sloths dying out. All Equidae become extinct in North America
10 ka The Holocene Epoch starts 10,000[42] years ago after the Late Glacial
Maximum. The last mainland species of Woolly mammoth (Mammuthus primigenius)
die out, as does the last Smilodon species
6 ka Small populations of American Mastodon die off in places like Utah and
Michigan
4500 ya The last members of a dwarf race of Woolly Mammoths vanish from Wrangel
Island near Alaska
384 ya (1627) The last recorded wild Aurochs die out
75 ya (1936) The Thylacine goes extinct in a Tasmanian zoo, the last member of the family
Thylacinidae
[edit] See also

Book:Evolution

• Extinction events
• Geologic time scale
• History of Earth
• Natural history
• Sociocultural evolution
• Timeline of human evolution
• The Ancestor's Tale by Richard Dawkins, for a list of ancestors common to humans and
other living species
[edit] References
1. ^ Planetary Science Institute page on the Giant Impact Hypothesis. Hartmann and Davis
belonged to the PSI. This page also contains several paintings of the impact by Hartmann himself.
2. ^ "Because the Moon helps stabilize the tilt of the Earth's rotation, it prevents the Earth from
wobbling between climatic extremes. Without the Moon, seasonal shifts would likely outpace even
the most adaptable forms of life." Making the Moon Astrobiology Magazine. (URL accessed on
August 7, 2010)
3. ^ "However, once the Earth cooled sufficiently, sometime in the first 700 million years of its
existence, clouds began to form in the atmosphere, and the Earth entered a new phase of
development." How the Oceans Formed (URL accessed on January 9, 2005)
4. ^ *The 'PAH World'
5. ^ Gilbert, Walter (February 1986). "The RNA World". Nature 319: 618.
doi:10.1038/319618a0.
6. ^ Joyce, G.F. (2002). "The antiquity of RNA-based evolution". Nature 418 (6894): 214–21.
doi:10.1038/418214a. PMID 12110897.
7. ^ Hoenigsberg, H. (December 2003). "Evolution without speciation but with selection: LUCA,
the Last Universal Common Ancestor in Gilbert’s RNA world". Genetic and Molecular Research 2 (4):
366–375. PMID 15011140. http://www.funpecrp.com.br/gmr/year2003/vol4-2/gmr0070_full_text.htm.
Retrieved 2008-08-30. (also available as PDF)
8. ^ Trevors, J. T. and Abel, D. L. (2004). "Chance and necessity do not explain the origin of
life". Cell Biol. Int. 28 (11): 729–39. doi:10.1016/j.cellbi.2004.06.006. PMID 15563395.
9. ^ Forterre, P., Benachenhou-Lahfa, N., Confalonieri, F., Duguet, M., Elie, C. and Labedan, B.
(1992). "The nature of the last universal ancestor and the root of the tree of life, still open questions".
BioSystems 28 (1-3): 15–32. doi:10.1016/0303-2647(92)90004-I. PMID 1337989.
10.^ Steenhuysen, Julie (May 21, 2009). "Study turns back clock on origins of life on Earth".
Reuters.com. Reuters. http://www.reuters.com/article/scienceNews/idUSTRE54J5PX20090520?
pageNumber=1&virtualBrandChannel=0. Retrieved May 21, 2009.
11.^ " Between about 3.8 billion and 4.5 billion years ago, no place in the solar system was safe
from the huge arsenal of asteroids and comets left over from the formation of the planets. Sleep and
Zahnle calculate that Earth was probably hit repeatedly by objects up to 500 kilometers across "
Geophysicist Sleep: Martian underground may have harbored early life (URL accessed on January 9,
2005)
12.^ Carl Woese, J Peter Gogarten, "When did eukaryotic cells (cells with nuclei and other
internal organelles) first evolve? What do we know about how they evolved from earlier life-forms?"
Scientific American, October 21, 1999.
13.^ Romano AH, Conway T. (1996). "Evolution of carbohydrate metabolic pathways". Res
Microbiol 147 (6-7): 448–55. doi:10.1016/0923-2508(96)83998-2. PMID 9084754.
14.^ Knowles JR (1980). "Enzyme-catalyzed phosphoryl transfer reactions". Annu. Rev.
Biochem. 49: 877–919. doi:10.1146/annurev.bi.49.070180.004305. PMID 6250450.
15.^ Doolittle, W. Ford (February, 2000). Uprooting the tree of life. Scientific American 282 (6):
90–95.
16.^ Nicolas Glansdorff, Ying Xu & Bernard Labedan: The Last Universal Common Ancestor :
emergence, constitution and genetic legacy of an elusive forerunner. Biology Direct 2008, 3:29.
17.^ Hahn, Jürgen; Pat Haug (1986). "Traces of Archaebacteria in ancient sediments". System
Applied Microbiology 7 (Archaebacteria '85 Proceedings): 178–83.
18.^ Olson JM (May 2006). "Photosynthesis in the Archean era". Photosyn. Res. 88 (2): 109–17.
doi:10.1007/s11120-006-9040-5. PMID 16453059.
19.^ Buick R (August 2008). "When did oxygenic photosynthesis evolve?". Philos. Trans. R.
Soc. Lond., B, Biol. Sci. 363 (1504): 2731–43. doi:10.1098/rstb.2008.0041. PMID 18468984.
20.^ Knoll, Andrew H.; Javaux, E.J, Hewitt, D. and Cohen, P. (2006). "Eukaryotic organisms in
Proterozoic oceans". Philosophical Transactions of the Royal Society of London, Part B 361 (1470):
1023–38. doi:10.1098/rstb.2006.1843. PMID 16754612.
21.^ Fedonkin, M. A. (March 2003). "The origin of the Metazoa in the light of the Proterozoic
fossil record" (PDF). Paleontological Research 7 (1): 9–41. doi:10.2517/prpsj.7.9.
22.^ Nicholas J. Butterfield, "Bangiomorpha pubescens n. gen., n. sp.: implications for the
evolution of sex, multicellularity, and the Mesoproterozoic/Neoproterozoic radiation of eukaryotes"
23.^ Hoffman, P.F.; Kaufman, A.J., Halverson, G.P., Schrag, D.P. (1998-08-28). "A
Neoproterozoic Snowball Earth". Science 281 (5381): 1342–6. doi:10.1126/science.281.5381.1342.
PMID 9721097. http://www.sciencemag.org/cgi/content/full/281/5381/1342?
ijkey=48d78da67bab492803c333f50c0dd84fbbef109c. Retrieved 2007-05-04. Full online article (pdf
260 Kb)
24.^ Kirschvink, J.L. (1992). "Late Proterozoic low-latitude global glaciation: The snowball
Earth". In Schopf, JW, and Klein, C. (PDF). The Proterozoic Biosphere: A Multidisciplinary Study.
Cambridge University Press, Cambridge. pp. 51–52.
http://www.gps.caltech.edu/~jkirschvink/pdfs/firstsnowball.pdf.
25.^ http://researchpages.net/media/resources/2007/06/21/richtimhywelfinal.pdf
26.^ Corsetti, F.A.; Awramik, S.M.; Pierce, D. (2003-04-15). "A complex microbiota from
snowball Earth times: Microfossils from the Neoproterozoic Kingston Peak Formation, Death Valley,
USA". Proceedings of the National Academy of Sciences 100 (8): 4399–4404.
doi:10.1073/pnas.0730560100. PMID 12682298. PMC 153566.
http://www.pnas.org/cgi/content/abstract/100/8/4399. Retrieved 2007-06-28.
27.^ Corsetti, F.A.; Olcott, A.N.; Bakermans, C. (2006). "The biotic response to Neoproterozoic
Snowball Earth". Palaeogeography, Palaeoclimatology, Palaeoecology 232 (232): 114–130.
doi:10.1016/j.palaeo.2005.10.030.
28.^ Narbonne, Guy (June 2006). "The Origin and Early Evolution of Animals". Department of
Geological Sciences and Geological Engineering, Queen's University.
http://geol.queensu.ca/people/narbonne/recent_pubs1.html. Retrieved 2007-03-10.
29.^ The Cambrian Period
30.^ The Cambrian Explosion – Timing
31.^ a b Formation of the Ozone Layer
32.^ "The oldest fossils of footprints ever found on land hint that animals may have beaten
plants out of the primordial seas. Lobster-sized, centipede-like or slug like animals such as
Protichnites and Climactichnites made the prints wading out of the ocean and scuttling over sand
dunes about 530 million years ago. Previous fossils indicated that animals didn't take this step until
40 million years later." Oldest fossil footprints on land
33.^ "The oldest fossils reveal evolution of non-vascular plants by the middle to late Ordovician
Period (~450–440 Ma) on the basis of fossil spores" Transition of plants to land
34.^ "The land plants evolved from the algae, more specifically green algae, as suggested by
certain common biochemical traits" The first land plants
35.^ "The ancestry of sharks dates back more than 200 million years before the earliest known
dinosaur. Introduction to shark evolution, geologic time and age determination
36.^ Sahney, S. and Benton, M.J. (2008). "Recovery from the most profound mass extinction of
all time" (PDF). Proceedings of the Royal Society: Biological 275 (1636): 759–65.
doi:10.1098/rspb.2007.1370. PMID 18198148. PMC 2596898.
37.^ "Viruses of nearly all the major classes of organisms—animals, plants, fungi and
bacteria/archaea—probably evolved with their hosts in the seas, given that most of the evolution of
life on this planet has occurred there. This means that viruses also probably emerged from the
waters with their different hosts, during the successive waves of colonisation of the terrestrial
environment." Origins of Viruses (URL accessed on January 9, 2005)
38.^ Chiappe, Luis M., & Dyke, Gareth J. (2002). "The Mesozoic Radiation of Birds". Annual
Review of Ecology & Systematics 33: 91–124. doi:10.1146/annurev.ecolsys.33.010802.150517.
39.^ The Oldest Homo Sapiens: - URL retrieved May 15, 2009
40.^ Alemseged, Z., Coppens, Y., Geraads, D. (2002). "Hominid cranium from Homo:
Description and taxonomy of Homo-323-1976-896". Am J Phys Anthropol 117 (2): 103–12.
doi:10.1002/ajpa.10032. PMID 11815945.
41.^ Stoneking, Mark; Soodyall, Himla (1996). "Human evolution and the mitochondrial
genome". Current Opinion in Genetics & Development 6 (6): 731–6. doi:10.1016/S0959-
437X(96)80028-1.
42.^ "International Stratigraphic Chart". International Commission on Stratigraphy.
http://www.stratigraphy.org/cheu.pdf. Retrieved 2009-02-03. [dead link]

• Berkeley Evolution
• Tree of Life Web Project - explore complete phylogenetic tree interactively
• A more compact timeline at the TalkOrigins Archive
• Palaeos - The Trace of Life on Earth
• University of Waikato - Sequence of Plant Evolution
• University of Waikato - Sequence of Animal Evolution
• Graphical Timeline of evolution
• History of Life on Earth
• Exploring Time from Planck Time to the lifespan of the universe
• Interactive Plant Evolution Timeline - from the University of Cambridge Ensemble Project
Processes of
Population genetic
Evolutionary Canalisation · Inversion · Modularity · Phenotypic plasticity

developmental
biology (Evo-devo)
concepts

Modes of


[hide]v · d · eEarth-related topics


Retrieved from "http://en.wikipedia.org/wiki/Timeline_of_evolution"
Categories: Biology timelines | Evolutionary biology

W000
ry of evolutionary thought
W000
History of Evolutionary Thought

This article is about the history of evolutionary thought in biology. For the history of evolutionary
thought in the social sciences, see Sociocultural evolution. For the history of religious discussions, see
History of the creation-evolution controversy.
The Tree of Life as depicted by Ernst Haeckel in The Evolution of Man (1879) illustrates the 19th-
century view that evolution was a progressive process leading towards man.
Evolutionary thought, the conception that species change over time, has roots in antiquity, in the
ideas of the ancient Greeks, Romans, and Chinese as well as in medieval Islamic science. However, until the
18th century, Western biological thinking was dominated by essentialism, the belief that every species has
essential characteristics that are unalterable. This began to change during the Enlightenment when
evolutionary cosmology and the mechanical philosophy spread from the physical sciences to natural history.
Naturalists began to focus on the variability of species; the emergence of paleontology with the concept of
extinction further undermined the static view of nature. In the early 19th century, Jean-Baptiste Lamarck
proposed his theory of the transmutation of species, the first fully formed scientific theory of evolution.
In 1858, Charles Darwin and Alfred Russel Wallace published a new evolutionary theory that was
explained in detail in Darwin's On the Origin of Species (1859). Unlike Lamarck, Darwin proposed common
descent and a branching tree of life. The theory was based on the idea of natural selection, and it
synthesized a broad range of evidence from animal husbandry, biogeography, geology, morphology, and
embryology.
The debate over Darwin's work led to the rapid acceptance of the general concept of evolution, but
the specific mechanism he proposed, natural selection, was not widely accepted until it was revived by
developments in biology that occurred during 1920s through the 1940s. Before that time most biologists
argued that other factors were responsible for evolution. Alternatives to natural selection suggested during
the eclipse of Darwinism included inheritance of acquired characteristics (neo-Lamarckism), an innate drive
for change (orthogenesis), and sudden large mutations (saltationism). The synthesis of natural selection with
Mendelian genetics during the 1920s and 1930s founded the new discipline of population genetics.
Throughout the 1930s and 1940s, population genetics became integrated with other biological fields,
resulting in a widely applicable theory of evolution that encompassed much of biology—the modern
evolutionary synthesis.
Following the establishment of evolutionary biology, studies of mutation and variation in natural
populations, combined with biogeography and systematics, led to sophisticated mathematical and causal
models of evolution. Paleontology and comparative anatomy allowed more detailed reconstructions of the
history of life. After the rise of molecular genetics in the 1950s, the field of molecular evolution developed,
based on protein sequences and immunological tests, and later incorporating RNA and DNA studies. The
gene-centered view of evolution rose to prominence in the 1960s, followed by the neutral theory of molecular
evolution, sparking debates over adaptationism, the units of selection, and the relative importance of genetic
drift versus natural selection. In the late 20th century, DNA sequencing led to molecular phylogenetics and
the reorganization of the tree of life into the three-domain system. In addition, the newly recognized factors of
symbiogenesis and horizontal gene transfer introduced yet more complexity into evolutionary history.
Evolution
Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation
Introduction
Evidence
History
Level of support
Modern synthesis
Social effect
Theory and fact
Cladistics
Ecological genetics
Molecular evolution
Phylogenetics
Population genetics
Systematics
Contents
[hide]
• 1 Antiquity
• 1.1 Greeks
• 1.2 Chinese
• 1.3 Romans
• 1.4 Augustine of Hippo
• 2 Middle Ages
• 2.1 Islamic philosophy and the struggle for
existence
• 2.2 Christian philosophy and the great chain of
being
• 2.3 Thomas Aquinas on creation and natural
processes
• 3 Renaissance and Enlightenment
• 4 Early 19th century
• 4.1 Paleontology and geology
• 4.2 Transmutation of species
• 4.3 Anticipations of natural selection
• 4.4 Natural selection
• 5 1859–1930s: Darwin and his legacy
• 5.1 Application to humans
[edit] Antiquity
[edit] Greeks
Several ancient Greek philosophers discussed ideas that involved change in living organisms over
time. Anaximander (c.610–546 BC) proposed that the first animals lived in water and animals that live on land
were generated from them.[1] Empedocles (c. 490–430 BC) wrote of a non-supernatural origin for living
things,[2] suggesting that adaptation did not require an organizer or final cause. Aristotle summarized his
idea: "Wherever then all the parts came about just what they would have been if they had come to be for an
end, such things survived, being organized spontaneously in a fitting way; whereas those which grew
otherwise perished and continue to perish ..." although Aristotle himself rejected this view.[3]
Plato (left) and Aristotle (right), a detail of The School of Athens
Plato (c. 428–348 BC) was, in the words of biologist and historian Ernst Mayr, "the great antihero of
evolutionism",[4] as he established the philosophy of essentialism, which he called the Theory of Forms. This
theory holds that objects observed in the real world are only reflections of a limited number of essences
(eide). Variation merely results from an imperfect reflection of these constant essences. In his Timaeus, Plato
set forth the idea that the Demiurge had created the cosmos and everything in it because, being good, and
hence, "... free from jealousy, He desired that all things should be as like Himself as they could be". The
creator created all conceivable forms of life, since "... without them the universe will be incomplete, for it will
not contain every kind of animal which it ought to contain, if it is to be perfect". This "plenitude principle"—the
idea that all potential forms of life are essential to a perfect creation—greatly influenced Christian thought.[5]
Aristotle (384–322 BC), one of the most influential of the Greek philosophers, is the earliest natural
historian whose work has been preserved in any real detail. His writings on biology resulted from his
research into natural history on and around the isle of Lesbos, and have survived in the form of four books,
usually known by their Latin names, De anima (on the essence of life), Historia animalium (inquiries about
animals), De generatione animalium (reproduction), and De partibus animalium (anatomy). Aristotle's works
contain some remarkably astute observations and interpretations—along with sundry myths and mistakes—
reflecting the uneven state of knowledge during his time.[6] However, for Charles Singer, "Nothing is more
remarkable than [Aristotle's] efforts to [exhibit] the relationships of living things as a scala naturæ."[6] This
scala naturæ, described in Historia animalium, classified organisms in relation to a hierarchical "Ladder of
Life" or "Chain of Being", placing them according to their complexity of structure and function, with organisms
that showed greater vitality and ability to move described as "higher organisms".[5]
[edit] Chinese
Ancient Chinese thinkers such as Zhuangzi (Chuang Tzu), a Taoist philosopher who lived around the
4th century BC, expressed ideas on changing biologic species. According to Joseph Needham, Taoism
explicitly denies the fixity of biological species and Taoist philosophers speculated that species had
developed differing attributes in response to differing environments.[7] Taoism regards humans, nature and
the heavens as existing in a state of "constant transformation" known as the Tao, in contrast with the more
static view of nature typical of Western thought.[8]
[edit] Romans
Titus Lucretius Carus (d. 50 BC), the Roman philosopher and atomist, wrote the poem On the Nature
of Things (De rerum natura), which provides the best surviving explanation of the ideas of the Greek
Epicurean philosophers. It describes the development of the cosmos, the Earth, living things, and human
society through purely naturalistic mechanisms, without any reference to supernatural involvement. On the
Nature of things would influence the cosmological and evolutionary speculations of philosophers and
scientists during and after the Renaissance.[9][10]
[edit] Augustine of Hippo

In line with earlier Greek thought, the 4th century bishop and theologian, St. Augustine of Hippo,
wrote that the creation story in Genesis should not be read too literally. In his book De Genesi ad litteram
("On The Literal Interpretation of Genesis"), he stated that in some cases new creatures may have come
about through the "decomposition" of earlier forms of life.[11] For Augustine, "plant, fowl and animal life are
not perfect ... but created in a state of potentiality", unlike what he considered the theologically perfect forms
of angels, the firmament and the human soul.[12] Augustine's idea 'that forms of life had been transformed
"slowly over time"' prompted Father Giuseppe Tanzella-Nitti, Professor of Theology at the Pontifical Santa
Croce University in Rome, to claim that Augustine had suggested a form of evolution.[13][14]
[edit] Middle Ages

[edit] Islamic philosophy and the struggle for existence
See also: Early Islamic philosophy and Science in medieval Islam
Although Greek and Roman evolutionary ideas died out in Europe after the fall of the Roman Empire,
they were not lost to Islamic philosophers and scientists. In the Islamic Golden Age of the 8th to the 13th
centuries, philosophers explored ideas about natural history. These ideas included transmutation from non-
living to living: "from mineral to plant, from plant to animal, and from animal to man".[15]
The first Muslim biologist and philosopher to publish detailed speculations about natural history, the
Afro-Arab writer al-Jahiz, wrote in the 9th century. In the Book of Animals, he considered the effects of the
environment on an animal's chances for survival, and described the struggle for existence.[16] Al-Jahiz also
wrote descriptions of food chains.[17] Al-Jahiz speculated on the influence of the environment on animals
and considered the effects of the environment on the likelihood of an animal to survive. For example, Al-
Jahiz's wrote in his Book of Animals: "All animals, in short, can not exist without food, neither can the hunting
animal escape being hunted in his turn. Every weak animal devours those weaker than itself. Strong animals
cannot escape being devoured by other animals stronger than they".[16]
[edit] Christian philosophy and the great chain of being
Drawing of the great chain of being from Rhetorica Christiana (1579) by Diego Valades
During the Early Middle Ages, Greek classical learning was all but lost to the West. However, contact
with the Islamic world, where Greek manuscripts were preserved and expanded, soon led to a massive spate
of Latin translations in the 12th century. Europeans were re-introduced to the works of Plato and Aristotle, as
well as to Islamic thought. Christian thinkers of the scholastic school, in particular Abelard and Thomas
Aquinas, combined Aristotelian classification with Plato's ideas of the goodness of God, and of all potential
life forms being present in a perfect creation, to organize all inanimate, animate, and spiritual beings into a
huge interconnected system: the scala naturæ, or great chain of being.[5][18]
Within this system, everything that existed could be placed in order, from "lowest" to "highest", with
Hell at the bottom and God at the top—below God, an angelic hierarchy marked by the orbits of the planets,
mankind in an intermediate position, and worms the lowest of the animals. As the universe was ultimately
perfect, the great chain was also perfect. There were no empty links in the chain, and no link was
represented by more than one species. Therefore no species could ever move from one position to another.
Thus, in this Christianized version of Plato's perfect universe, species could never change, but remained
forever fixed, in accordance with the text of Genesis. For humans to forget their position was seen as sinful,
whether they behaved like lower animals or aspired to a higher station than was given them by their Creator.
[5]
Creatures on adjacent steps were expected to closely resemble each other, an idea expressed in the
saying: natura non facit saltum ("nature does not make leaps").[5] This basic concept of the great chain of
being greatly influenced the thinking of Western civilization for centuries (and still has an influence today). It
formed a part of the argument from design presented by natural theology. As a classification system, it
became the major organizing principle and foundation of the emerging science of biology in the 17th and
18th centuries.[5]
[edit] Thomas Aquinas on creation and natural processes
While the development of the great chain of being and the argument from design by Christian
theologians contributed to the view that the natural world fit into an unchanging designed hierarchy, some
theologians were more open to the possibility that the world might have developed through natural
processes. Thomas Aquinas went even farther than Augustine of Hippo in arguing that scriptural texts like
Genesis should not be interpreted in a literal way that conflicted with or constrained what natural
philosophers learned about the workings of the natural world. He felt that the autonomy of nature was a sign
of God's goodness and that there was no conflict between the concept of a divinely created universe, and the
idea that the universe may have evolved over time through natural mechanisms.[19] However, Aquinas
disputed the views of those like the ancient Greek philosopher Empodocles who held that such natural
processes showed that the universe could have developed without an underlying purpose. Rather holding
that: "Hence, it is clear that nature is nothing but a certain kind of art, i.e., the divine art, impressed upon
things, by which these things are moved to a determinate end. It is as if the shipbuilder were able to give to
timbers that by which they would move themselves to take the form of a ship."[20]
[edit] Renaissance and Enlightenment

Main article: Evolutionary ideas of the Renaissance and Enlightenment
Pierre Belon compared the skeletons of birds and humans in his Book of Birds (1555).
In the first half of the 17th century, René Descartes's mechanical philosophy encouraged the use of
the metaphor of the universe as a machine, a concept that would come to characterise the scientific
revolution.[21] Between 1650 and 1800, some evolutionist theories supported the view that the universe,
including life on Earth, had developed mechanically, entirely without divine guidance. In contrast, most
contemporary theories of evolution, such of those of Gottfried Leibniz and J. G. Herder, held that evolution
was a fundamentally spiritual process.[22] In 1751, Pierre Louis Maupertuis veered toward more materialist
ground. He wrote of natural modifications occurring during reproduction and accumulating over the course of
many generations, producing races and even new species, a description that anticipated in general terms the
concept of natural selection.[23]
Later in the 18th century, the French philosopher G. L. L. Buffon, one of the leading 18th century
naturalists, suggested that what most people referred to as species were really just well-marked varieties,
modified from an original form by environmental factors. For example, he believed that lions, tigers, leopards
and house cats might all have a common ancestor. He further speculated that the 200 or so species of
mammals then known might have descended from as few as 38 original animal forms. Buffon's evolutionary
ideas were limited; he believed each of the original forms had arisen through spontaneous generation and
that each was shaped by "internal moulds" that limited the amount of change. Buffon's works, Natural History
and The Epochs of Nature, containing well developed theories about a completely materialistic origin for the
Earth and his ideas questioning the fixity of species, were extremely influential.[24][25]
Between 1767 and 1792, James Burnett, Lord Monboddo included in his writings not only the
concept that man had descended from primates, but also that, in response to the environment, creatures had
found methods of transforming their characteristics over long time intervals.[26] Charles Darwin's
grandfather, Erasmus Darwin, published Zoönomia in 1796, which suggested that "all warm-blooded animals
have arisen from one living filament".[27] In his 1802 poem Temple of Nature, he described the rise of life
from minute organisms living in mud to all of its modern diversity.[28]
[edit] Early 19th century
Diagram of the geologic timescale from an 1861 book by Richard Owen showing the appearance of
major animal types
[edit] Paleontology and geology

See also: History of paleontology
In 1796, Georges Cuvier published his findings on the differences between living elephants and
those found in the fossil record. His analysis demonstrated that mammoths and mastodons were distinct
species, different from any living animal, effectively ending a long-running debate over whether the extinction
of a species was possible.[29] In 1788, James Hutton described gradual geological processes operating
continuously over deep time.[30] In the 1790s William Smith began the process of ordering rock strata by
examining fossils in the layers while he worked on his geologic map of England. Independently, in 1811,
Georges Cuvier and Alexandre Brongniart published an influential study of the geologic history of the region
around Paris, based on the stratigraphic succession of rock layers. These works helped establish the
antiquity of the Earth.[31] Cuvier advocated catastrophism to explain the patterns of extinction and faunal
succession revealed by the fossil record.
Knowledge of the fossil record continued to advance rapidly during the first few decades of the 19th
century. By the 1840s, the outlines of the geologic timescale were becoming clear, and in 1841 John Phillips
named three major eras, based on the predominant fauna of each: the Paleozoic, dominated by marine
invertebrates and fish, the Mesozoic, the age of reptiles, and the current Cenozoic age of mammals. This
progressive picture of the history of life was accepted even by conservative English geologists like Adam
Sedgwick and William Buckland; however, like Cuvier, they attributed the progression to repeated
catastrophic episodes of extinction followed by new episodes of creation.[32] Unlike Cuvier, Buckland and
some other advocates of natural theology among British geologists made efforts to explicitly link the last
catastrophic episode proposed by Cuvier to the biblical flood.[33][34]
From 1830 to 1833, Charles Lyell published his multi-volume work Principles of Geology, which,
building on Hutton's ideas, advocated a uniformitarian alternative to the catastrophic theory of geology. Lyell
claimed that, rather than being the products of cataclysmic (and possibly supernatural) events, the geologic
features of the Earth are better explained as the result of the same gradual geologic forces observable in the
present day—but acting over immensely long periods of time. Although Lyell opposed evolutionary ideas
(even questioning the consensus that the fossil record demonstrates a true progression), his concept that the
Earth was shaped by forces working gradually over an extended period, and the immense age of the Earth
assumed by his theories, would strongly influence future evolutionary thinkers such as Charles Darwin.[35]
[edit] Transmutation of species

Main article: Transmutation of species
Diagram from Vestiges of the Natural History of Creation (1844) by Robert Chambers shows a model
of development where fish (F), reptiles (R), and birds (B) represent branches from a path leading to
mammals (M).
Jean-Baptiste Lamarck proposed, in his Philosophie Zoologique of 1809, a theory of the
transmutation of species. Lamarck did not believe that all living things shared a common ancestor but rather
that simple forms of life were created continuously by spontaneous generation. He also believed that an
innate life force drove species to become more complex over time, advancing up a linear ladder of
complexity that was related to the great chain of being. Lamarck recognized that species were adapted to
their environment. He explained this by saying that the same innate force driving increasing complexity
caused the organs of an animal (or a plant) to change based on the use or disuse of those organs, just as
muscles are affected by exercise. He argued that these changes would be inherited by the next generation
and produce slow adaptation to the environment. It was this secondary mechanism of adaptation through the
inheritance of acquired characteristics that would become known as Lamarckism and would influence
discussions of evolution into the 20th century.[36][37]
A radical British school of comparative anatomy that included the anatomist Robert Grant was closely
in touch with Lamarck's French school of Transformationism. One of the French scientists who influenced
Grant was the anatomist Étienne Geoffroy Saint-Hilaire, whose ideas on the unity of various animal body
plans and the homology of certain anatomical structures would be widely influential and lead to intense
debate with his colleague Georges Cuvier. Grant became an authority on the anatomy and reproduction of
marine invertebrates. He developed Lamarck's and Erasmus Darwin's ideas of transmutation and
evolutionism, and investigated homology, even proposing that plants and animals had a common
evolutionary starting point. As a young student Charles Darwin joined Grant in investigations of the life cycle
of marine animals. In 1826 an anonymous paper, probably written by Robert Jameson, praised Lamarck for
explaining how higher animals had "evolved" from the simplest worms; this was the first use of the word
"evolved" in a modern sense.[38][39]
In 1844, the Scottish publisher Robert Chambers anonymously published an extremely controversial
but widely read book entitled Vestiges of the Natural History of Creation. This book proposed an evolutionary
scenario for the origins of the Solar System and life on Earth. It claimed that the fossil record showed a
progressive ascent of animals with current animals being branches off a main line that leads progressively to
humanity. It implied that the transmutations lead to the unfolding of a preordained plan that had been woven
into the laws that governed the universe. In this sense it was less completely materialistic than the ideas of
radicals like Robert Grant, but its implication that humans were only the last step in the ascent of animal life
incensed many conservative thinkers. The high profile of the public debate over Vestiges, with its depiction of
evolution as a progressive process, would greatly influence the perception of Darwin's theory a decade later.
[40][41]
Ideas about the transmutation of species were associated with the radical materialism of the
Enlightenment and were attacked by more conservative thinkers. Georges Cuvier attacked the ideas of
Lamarck and Geoffroy Saint-Hilaire, agreeing with Aristotle that species were immutable. Cuvier believed
that the individual parts of an animal were too closely correlated with one another to allow for one part of the
anatomy to change in isolation from the others, and argued that the fossil record showed patterns of
catastrophic extinctions followed by re-population, rather than gradual change over time. He also noted that
drawings of animals and animal mummies from Egypt, which were thousands of years old, showed no signs
of change when compared with modern animals. The strength of Cuvier's arguments and his scientific
reputation helped keep transmutational ideas out of the mainstream for decades.[42]
This 1847 diagram by Richard Owen shows his conceptual archetype for all vertebrates.
In Britain the philosophy of natural theology remained influential. William Paley's 1802 book Natural
Theology with its famous watchmaker analogy had been written at least in part as a response to the
transmutational ideas of Erasmus Darwin.[43] Geologists influenced by natural theology, such as Buckland
and Sedgwick, made a regular practice of attacking the evolutionary ideas of Lamarck, Grant, and The
Vestiges of the Natural History of Creation .[44][45] Although the geologist Charles Lyell opposed scriptural
geology, he also believed in the immutability of species, and in his Principles of Geology (1830–1833), he
criticized Lamarck's theories of development.[35] Idealists such as Louis Agassiz and Richard Owen believed
that each species was fixed and unchangeable because it represented an idea in the mind of the creator.
They believed that relationships between species could be discerned from developmental patterns in
embryology, as well as in the fossil record, but that these relationships represented an underlying pattern of
divine thought, with progressive creation leading to increasing complexity and culminating in humanity. Owen
developed the idea of "archetypes" in the Divine mind that would produce a sequence of species related by
anatomical homologies, such as vertebrate limbs. Owen led a public campaign that successfully
marginalized Robert Grant in the scientific community. Darwin would make good use of the homologies
analyzed by Owen in his own theory, but the harsh treatment of Grant, and the controversy surrounding
Vestiges, showed him the need to ensure that his own ideas were scientifically sound.[39][46][47]
[edit] Anticipations of natural selection
Several writers anticipated aspects of Darwin's theory, and in the third edition of On the Origin of
Species published in 1861 Darwin named those he knew about in an introductory appendix, An Historical
Sketch of the Recent Progress of Opinion on the Origin of Species , which he expanded in later editions.[48]
In 1813, William Charles Wells read before the Royal Society essays assuming that there had been
evolution of humans, and recognising the principle of natural selection. Charles Darwin and Alfred Russel
Wallace were unaware of this work when they jointly published the theory in 1858, but Darwin later
acknowledged that Wells had recognised the principle before them, writing that the paper "An Account of a
White Female, part of whose Skin resembles that of a Negro" was published in 1818, and "he distinctly
recognises the principle of natural selection, and this is the first recognition which has been indicated; but he
applies it only to the races of man, and to certain characters alone."[49] When Darwin was developing his
theory, he was influenced by Augustin de Candolle's natural system of classification, which laid emphasis on
the war between competing species.[50][51]
Patrick Matthew wrote in the obscure book Naval Timber & Arboriculture (1831) of "continual
balancing of life to circumstance. ... [The] progeny of the same parents, under great differences of
circumstance, might, in several generations, even become distinct species, incapable of co-
reproduction."[52] Charles Darwin discovered this work after the initial publication of the Origin. In the brief
historical sketch that Darwin included in the 3rd edition he says "Unfortunately the view was given by Mr.
Matthew very briefly in an Appendix to a work on a different subject ... He clearly saw, however, the full force
of the principle of natural selection."[53]
It is possible to look through the history of biology from the ancient Greeks onwards and discover
anticipations of almost all of Darwin's key ideas. However, as historian of science Peter J. Bowler says,
"Through a combination of bold theorizing and comprehensive evaluation, Darwin came up with a concept of
evolution that was unique for the time." Bowler goes on to say that simple priority alone is not enough to
secure a place in the history of science; someone has to develop an idea and convince others of its
importance to have a real impact.[54]
T. H. Huxley said in his essay on the reception of the Origin of Species:
The suggestion that new species may result from the selective action of external conditions
upon the variations from their specific type which individuals present and which we call
spontaneous because we are ignorant of their causation is as wholly unknown to the historian of
scientific ideas as it was to biological specialists before 1858. But that suggestion is the central
idea of the Origin of Species, and contains the quintessence of Darwinism.[55]
Darwin's first sketch of an evolutionary tree from his First Notebook on Transmutation of Species
(1837)
[edit] Natural selection

Main articles: Inception of Darwin's theory, Development of Darwin's theory, and Publication of
Darwin's theory
The biogeographical patterns Charles Darwin observed in places such as the Galapagos islands
during the voyage of the Beagle caused him to doubt the fixity of species, and in 1837 Darwin started the first
of a series of secret notebooks on transmutation. Darwin's observations led him to view transmutation as a
process of divergence and branching, rather than the ladder-like progression envisioned by Lamarck and
others. In 1838 he read the new 6th edition of An Essay on the Principle of Population, written in the late 18th
century by Thomas Malthus. Malthus' idea of population growth leading to a struggle for survival combined
with Darwin's knowledge on how breeders selected traits, led to the inception of Darwin's theory of natural
selection. Darwin did not publish his ideas on evolution for 20 years. However he did share them with certain
other naturalists and friends, starting with Joseph Hooker, with whom he discussed his unpublished 1844
essay on natural selection. During this period he used the time he could spare from his other scientific work
to slowly refine his ideas and, aware of the intense controversy around transmutation, amass evidence to
support them. In September 1854 he began full time work on writing his book on natural selection.[47][56][57]
Unlike Darwin, Alfred Russel Wallace, influenced by the book Vestiges of the Natural History of
Creation, already suspected that transmutation of species occurred when he began his career as a naturalist.
By 1855 his biogeographical observations during his field work in South America and the Malay Archipelago
made him confident enough in a branching pattern of evolution to publish a paper stating that every species
originated in close proximity to an already existing closely allied species. Like Darwin, it was Wallace's
consideration of how the ideas of Malthus might apply to animal populations that led him to conclusions very
similar to those reached by Darwin about the role of natural selection. In February 1858 Wallace, unaware of
Darwin's unpublished ideas, composed his thoughts into an essay and mailed them to Darwin, asking for his
opinion. The result was the joint publication in July of an extract from Darwin's 1844 essay along with
Wallace's letter. Darwin also began work on a short abstract summarising his theory, which he would publish
in 1859 as On the Origin of Species.[58]
Diagram by O.C. Marsh of the evolution of horse feet and teeth over time as reproduced in T.H
Huxley's 1876 book Professor Huxley in America
[edit] 1859–1930s: Darwin and his legacy
See also: Reaction to Darwin's theory
By the 1850s whether or not species evolved was a subject of intense debate, with prominent
scientists arguing both sides of the issue.[59] However, it was the publication of Charles Darwin's On the
Origin of Species (1859) that fundamentally transformed the discussion over biological origins.[60] Darwin
argued that his branching version of evolution explained a wealth of facts in biogeography, anatomy,
embryology, and other fields of biology. He also provided the first cogent mechanism by which evolutionary
change could persist: his theory of natural selection.[61]
One of the first and most important naturalists to be convinced by Origin of the reality of evolution
was the British anatomist Thomas Henry Huxley. Huxley recognized that unlike the earlier transmutational
ideas of Lamarck and Vestiges, Darwin's theory provided a mechanism for evolution without supernatural
involvement, even if Huxley himself was not completely convinced that natural selection was the key
evolutionary mechanism. Huxley would make advocacy of evolution a cornerstone of the program of the X
Club to reform and professionalise science by displacing natural theology with naturalism and to end the
domination of British natural science by the clergy. By the early 1870s in English-speaking countries, thanks
partly to these efforts, evolution had become the mainstream scientific explanation for the origin of species.
[61] In his campaign for public and scientific acceptance of Darwin's theory, Huxley made extensive use of
new evidence for evolution from paleontology. This included evidence that birds had evolved from reptiles,
including the discovery of Archaeopteryx in Europe, and a number of fossils of primitive birds with teeth found
in North America. Another important line of evidence was the finding of fossils that helped trace the evolution
of the horse from its small five-toed ancestors.[62] However, acceptance of evolution among scientists in
non-English speaking nations such as France, and the countries of southern Europe and Latin America was
slower. An exception to this was Germany, where both August Weismann and Ernst Haeckel championed
this idea: Haeckel used evolution to challenge the established tradition of metaphysical idealism in German
biology, much as Huxley used it to challenge natural theology in Britain.[63] Haeckel and other German
scientists would take the lead in launching an ambitious programme to reconstruct the evolutionary history of
life based on morphology and embryology.[64]
Darwin's theory succeeded in profoundly altering scientific opinion regarding the development of life
and in producing a small philosophical revolution.[65] However, this theory could not explain several critical
components of the evolutionary process. Specifically, Darwin was unable to explain the source of variation in
traits within a species, and could not identify a mechanism that could pass traits faithfully from one
generation to the next. Darwin's hypothesis of pangenesis, while relying in part on the inheritance of acquired
characteristics, proved to be useful for statistical models of evolution that were developed by his cousin
Francis Galton and the "biometric" school of evolutionary thought. However, this idea proved to be of little
use to other biologists.[66]
[edit] Application to humans

This illustration was the frontispiece of Thomas Henry Huxley's book Evidence as to Man's Place in
Nature (1863).
Charles Darwin was aware of the severe reaction in some parts of the scientific community against
the suggestion made in Vestiges of the Natural History of Creation that humans had arisen from animals by a
process of transmutation. Therefore he almost completely ignored the topic of human evolution in The Origin
of Species. Despite this precaution, the issue featured prominently in the debate that followed the book's
publication. For most of the first half of the 19th century, the scientific community believed that, although
geology had shown that the Earth and life were very old, human beings had appeared suddenly just a few
thousand years before the present. However, a series of archaeological discoveries in the 1840s and 1850s
showed stone tools associated with the remains of extinct animals. By the early 1860s, as summarized in
Charles Lyell's 1863 book Geological Evidences of the Antiquity of Man, it had become widely accepted that
humans had existed during a prehistoric period – which stretched many thousands of years before the start of
written history. This view of human history was more compatible with an evolutionary origin for humanity than
was the older view. On the other hand, at that time there was no fossil evidence to demonstrate human
evolution. The only human fossils found before the discovery of Java man in the 1890s were either of
anatomically modern humans or of Neanderthals that were too close, especially in the critical characteristic
of cranial capacity, to modern humans for them to be convincing intermediates between humans and other
primates.[67]
Therefore the debate that immediately followed the publication of The Origin of Species centered on
the similarities and differences between humans and modern apes. Carolus Linnaeus had been criticised in
the 18th century for grouping humans and apes together as primates in his ground breaking classification
system.[68] Richard Owen vigorously defended the classification suggested by Cuvier and Johann Friedrich
Blumenbach that placed humans in a separate order from any of the other mammals, which by the early 19th
century had become the orthodox view. On the other hand, Thomas Henry Huxley sought to demonstrate a
close anatomical relationship between humans and apes. In one famous incident, Huxley showed that Owen
was mistaken in claiming that the brains of gorillas lacked a structure present in human brains. Huxley
summarized his argument in his highly influential 1863 book Evidence as to Man's Place in Nature. Another
viewpoint was advocated by Charles Lyell and Alfred Russel Wallace. They agreed that humans shared a
common ancestor with apes, but questioned whether any purely materialistic mechanism could account for
all the differences between humans and apes, especially some aspects of the human mind.[67]
In 1871, Darwin published The Descent of Man, and Selection in Relation to Sex , which contained
his views on human evolution. Darwin argued that the differences between the human mind and the minds of
the higher animals were a matter of degree rather than of kind. For example, he viewed morality as a natural
outgrowth of instincts that were beneficial to animals living in social groups. He argued that all the differences
between humans and apes were explained by a combination of the selective pressures that came from our
ancestors moving from the trees to the plains, and sexual selection. The debate over human origins, and
over the degree of human uniqueness continued well into the 20th century.[67]
[edit] Alternatives to natural selection

Main article: The eclipse of Darwinism
This photo from Henry Fairfield Osborn's 1918 book Origin and Evolution of Life shows models
depicting the evolution of Titanothere horns over time, which Osborn claimed was an example of an
orthogenic trend in evolution.
The concept of evolution was widely accepted in scientific circles within a few years of the publication
of Origin, but the acceptance of natural selection as its driving mechanism was much less widespread. The
four major alternatives to natural selection in the late 19th century were theistic evolution, neo-Lamarckism,
orthogenesis, and saltationism. Theistic evolution (a term promoted by Darwin's greatest American advocate
Asa Gray) was the idea that God intervened in the process of evolution to guide it in such a way that the
living world could still be considered to be designed. However, this idea gradually fell out of favor among
scientists, as they became more and more committed to the idea of methodological naturalism and came to
believe that direct appeals to supernatural involvement were scientifically unproductive. By 1900, theistic
evolution had largely disappeared from professional scientific discussions, although it retained a strong
popular following.[69][70]
In the late 19th century, the term neo-Lamarckism came to be associated with the position of
naturalists who viewed the inheritance of acquired characteristics as the most important evolutionary
mechanism. Advocates of this position included the British writer and Darwin critic Samuel Butler, the
German biologist Ernst Haeckel, and the American paleontologist Edward Drinker Cope. They considered
Lamarckism to be philosophically superior to Darwin's idea of selection acting on random variation. Cope
looked for, and thought he found, patterns of linear progression in the fossil record. Inheritance of acquired
characteristics was part of Haeckel's recapitulation theory of evolution, which held that the embryological
development of an organism repeats its evolutionary history.[69][70] Critics of neo-Lamarckism, such as the
German biologist August Weismann and Alfred Russel Wallace, pointed out that no one had ever produced
solid evidence for the inheritance of acquired characteristics. Despite these criticisms, neo-Lamarckism
remained the most popular alternative to natural selection at the end of the 19th century, and would remain
the position of some naturalists well into the 20th century.[69][70]
Orthogenesis was the hypothesis that life has an innate tendency to change, in a unilinear fashion,
towards ever-greater perfection. It had a significant following in the 19th century, and its proponents included
the Russian biologist Leo Berg and the American paleontologist Henry Fairfield Osborn. Orthogenesis was
popular among some paleontologists, who believed that the fossil record showed a gradual and constant
unidirectional change. Saltationism was the idea that new species arise as a result of large mutations. It was
seen as a much faster alternative to the Darwinian concept of a gradual process of small random variations
being acted on by natural selection, and was popular with early geneticists such as Hugo de Vries, William
Bateson, and early in his career, T. H. Morgan. It became the basis of the mutation theory of evolution.[69]
[70]
Diagram from T.H. Morgan's 1919 book The Physical Basis of Heredity, showing the sex-linked
inheritance of the white-eyed mutation in Drosophila melanogaster
[edit] Mendelian genetics, biometrics, and mutation

The so-called rediscovery of Gregor Mendel's laws of inheritance in 1900 ignited a fierce debate
between two camps of biologists. In one camp were the Mendelians, who were focused on discrete variations
and the laws of inheritance. They were led by William Bateson (who coined the word genetics) and Hugo de
Vries (who coined the word mutation). Their opponents were the biometricians, who were interested in the
continuous variation of characteristics within populations. Their leaders, Karl Pearson and Walter Frank
Raphael Weldon, followed in the tradition of Francis Galton, who had focused on measurement and statistical
analysis of variation within a population. The biometricians rejected Mendelian genetics on the basis that
discrete units of heredity, such as genes, could not explain the continuous range of variation seen in real
populations. Weldon's work with crabs and snails provided evidence that selection pressure from the
environment could shift the range of variation in wild populations, but the Mendelians maintained that the
variations measured by biometricians were too insignificant to account for the evolution of new species.[71]
[72]
When T. H. Morgan began experimenting with breeding the fruit fly Drosophila melanogaster, he was
a saltationist who hoped to demonstrate that a new species could be created in the lab by mutation alone.
Instead, the work at his lab between 1910 and 1915 reconfirmed Mendelian genetics and provided solid
experimental evidence linking it to chromosomal inheritance. His work also demonstrated that most
mutations had relatively small effects, such as a change in eye color, and that rather than creating a new
species in a single step, mutations served to increase variation within the existing population.[71][72]
[edit] 1920s–1940s
See also: Modern evolutionary synthesis
Biston betularia f. typica is the white-bodied form of the peppered moth.
Biston betularia f. carbonaria is the black-bodied form of the peppered moth.
[edit] Population genetics

The Mendelian and biometrician models were eventually reconciled with the development of
population genetics. A key step was the work of the British biologist and statistician R.A. Fisher. In a series of
papers starting in 1918 and culminating in his 1930 book The Genetical Theory of Natural Selection , Fisher
showed that the continuous variation measured by the biometricians could be produced by the combined
action of many discrete genes, and that natural selection could change gene frequencies in a population,
resulting in evolution. In a series of papers beginning in 1924, another British geneticist, J.B.S. Haldane,
applied statistical analysis to real-world examples of natural selection, such as the evolution of industrial
melanism in peppered moths, and showed that natural selection worked at an even faster rate than Fisher
assumed.[73][74]
The American biologist Sewall Wright, who had a background in animal breeding experiments,
focused on combinations of interacting genes, and the effects of inbreeding on small, relatively isolated
populations that exhibited genetic drift. In 1932, Wright introduced the concept of an adaptive landscape and
argued that genetic drift and inbreeding could drive a small, isolated sub-population away from an adaptive
peak, allowing natural selection to drive it towards different adaptive peaks. The work of Fisher, Haldane and
Wright founded the discipline of population genetics. This integrated natural selection with Mendelian
genetics, which was the critical first step in developing a unified theory of how evolution worked.[73][74]
[edit] Modern evolutionary synthesis

In the first few decades of the 20th century, most field naturalists continued to believe that
Lamarckian and orthogenic mechanisms of evolution provided the best explanation for the complexity they
observed in the living world. But as the field of genetics continued to develop, those views became less
tenable.[75] Theodosius Dobzhansky, a postdoctoral worker in T. H. Morgan's lab, had been influenced by
the work on genetic diversity by Russian geneticists such as Sergei Chetverikov. He helped to bridge the
divide between the foundations of microevolution developed by the population geneticists and the patterns of
macroevolution observed by field biologists, with his 1937 book Genetics and the Origin of Species.
Dobzhansky examined the genetic diversity of wild populations and showed that, contrary to the assumptions
of the population geneticists, these populations had large amounts of genetic diversity, with marked
differences between sub-populations. The book also took the highly mathematical work of the population
geneticists and put it into a more accessible form. In Great Britain E.B. Ford, the pioneer of ecological
genetics, continued throughout the 1930s and 1940s to demonstrate the power of selection due to ecological
factors including the ability to maintain genetic diversity through genetic polymorphisms such as human
blood types. Ford's work would contribute to a shift in emphasis during the course of the modern synthesis
towards natural selection over genetic drift.[73][74][76][77]
Evolutionary biologist Ernst Mayr was influenced by the work of the German biologist Bernhard
Rensch showing the influence of local environmental factors on the geographic distribution of sub-species
and closely related species. Mayr followed up on Dobzhansky's work with the 1942 book Systematics and
the Origin of Species, which emphasized the importance of allopatric speciation in the formation of new
species. This form of speciation occurs when the geographical isolation of a sub-population is followed by the
development of mechanisms for reproductive isolation. Mayr also formulated the biological species concept
that defined a species as a group of interbreeding or potentially interbreeding populations that were
reproductively isolated from all other populations.[73][74][78]
In the 1944 book Tempo and Mode in Evolution, George Gaylord Simpson showed that the fossil
record was consistent with the irregular non-directional pattern predicted by the developing evolutionary
synthesis, and that the linear trends that earlier paleontologists had claimed supported orthogenesis and
neo-Lamarckism did not hold up to closer examination. In 1950, G. Ledyard Stebbins published Variation
and Evolution in Plants, which helped to integrate botany into the synthesis. The emerging cross-disciplinary
consensus on the workings of evolution would be known as the modern evolutionary synthesis. It received its
name from the book Evolution: The Modern Synthesis by Julian Huxley.[73][74]
The evolutionary synthesis provided a conceptual core—in particular, natural selection and Mendelian
population genetics—that tied together many, but not all, biological disciplines. It helped establish the
legitimacy of evolutionary biology, a primarily historical science, in a scientific climate that favored
experimental methods over historical ones.[79] The synthesis also resulted in a considerable narrowing of
the range of mainstream evolutionary thought (what Stephen Jay Gould called the "hardening of the
synthesis"): by the 1950s, natural selection acting on genetic variation was virtually the only acceptable
mechanism of evolutionary change (panselectionism), and macroevolution was simply considered the result
of extensive microevolution.[80][81]
[edit] 1940s–1960s: Molecular biology and evolution

Main article: History of molecular evolution
The middle decades of the 20th century saw the rise of molecular biology, and with it an
understanding of the chemical nature of genes as sequences of DNA and their relationship, through the
genetic code, to protein sequences. At the same time, increasingly powerful techniques for analyzing
proteins, such as protein electrophoresis and sequencing, brought biochemical phenomena into realm of the
synthetic theory of evolution. In the early 1960s, biochemists Linus Pauling and Emile Zuckerkandl proposed
the molecular clock hypothesis: that sequence differences between homologous proteins could be used to
calculate the time since two species diverged. By 1969, Motoo Kimura and others provided a theoretical
basis for the molecular clock, arguing that—at the molecular level at least—most genetic mutations are neither
harmful nor helpful and that genetic drift (rather than natural selection) causes a large portion of genetic
change: the neutral theory of molecular evolution.[82] Studies of protein differences within species also
brought molecular data to bear on population genetics by providing estimates of the level of heterozygosity in
natural populations.[83]
From the early 1960s, molecular biology was increasingly seen as a threat to the traditional core of
evolutionary biology. Established evolutionary biologists—particularly Ernst Mayr, Theodosius Dobzhansky
and G. G. Simpson, three of the architects of the modern synthesis—were extremely skeptical of molecular
approaches, especially when it came to the connection (or lack thereof) to natural selection. The molecular-
clock hypothesis and the neutral theory were particularly controversial, spawning the neutralist-selectionist
debate over the relative importance of drift and selection, which continued into the 1980s without a clear
resolution.[84][85]
[edit] Late 20th century

[edit] Gene-centered view
In the mid-1960s, George C. Williams strongly critiqued explanations of adaptations worded in terms
of "survival of the species" (group selection arguments). Such explanations were largely replaced by a gene-
centered view of evolution, epitomized by the kin selection arguments of W. D. Hamilton, George R. Price
and John Maynard Smith.[86] This viewpoint would be summarized and popularized in the influential 1976
book The Selfish Gene by Richard Dawkins.[87] Models of the period showed that group selection was
severely limited in its strength; though newer models do admit the possibility of significant multi-level
selection.[88]
In 1973, Leigh Van Valen proposed the term "Red Queen", which he took from Through the Looking-
Glass by Lewis Carroll, to describe a scenario where a species involved in one or more evolutionary arms
races would have to constantly change just to keep pace with the species with which it was co-evolving.
Hamilton, Williams and others suggested that this idea might explain the evolution of sexual reproduction: the
increased genetic diversity caused by sexual reproduction would help maintain resistance against rapidly
evolving parasites, thus making sexual reproduction common, despite the tremendous cost from the gene-
centric point of view of a system where only half of an organism's genome is passed on during reproduction.
[89][90] The gene-centric view has also led to an increased interest in Darwin's old idea of sexual selection,
[91] and more recently in topics such as sexual conflict and intragenomic conflict.
[edit] Sociobiology
W. D. Hamilton's work on kin selection contributed to the emergence of the discipline of sociobiology.
The existence of altruistic behaviors has been a difficult problem for evolutionary theorists from the
beginning.[92] Significant progress was made in 1964 when Hamilton formulated the inequality in kin
selection known as Hamilton's rule, which showed how eusociality in insects (the existence of sterile worker
classes) and many other examples of altruistic behavior could have evolved through kin selection. Other
theories followed, some derived from game theory, such as reciprocal altruism.[93] In 1975, E.O. Wilson
published the influential and highly controversial book Sociobiology: The New Synthesis which claimed
evolutionary theory could help explain many aspects of animal, including human, behavior. Critics of
sociobiology, including Stephen Jay Gould and Richard Lewontin, claimed that sociobiology greatly
overstated the degree to which complex human behaviors could be determined by genetic factors. They also
claimed that the theories of sociobiologists often reflected their own ideological biases. Despite these
criticisms, work has continued in sociobiology and the related discipline of evolutionary psychology, including
work on other aspects of the altruism problem.[94][95]
A phylogenetic tree showing the three-domain system. Eukaryotes are colored red, Archaea green,
and Bacteria blue.
[edit] Evolutionary paths and processes

One of the most prominent debates arising during the 1970s was over the theory of punctuated
equilibrium. Niles Eldredge and Stephen Jay Gould proposed that there was a pattern of fossil species that
remained largely unchanged for long periods (what they termed stasis), interspersed with relatively brief
periods of rapid change during speciation.[96][97] Improvements in sequencing methods resulted in a large
increase of sequenced genomes, allowing the testing and refining of evolutionary theories using this huge
amount of genome data.[98] Comparisons between these genomes provide insights into the molecular
mechanisms of speciation and adaptation.[99][100] These genomic analyses have produced fundamental
changes in the understanding of the evolutionary history of life, such as the proposal of the three-domain
system by Carl Woese.[101] Advances in computational hardware and software allow the testing and
extrapolation of increasingly advanced evolutionary models and the development of the field of systems
biology.[102] One of the results has been an exchange of ideas between theories of biological evolution and
the field of computer science known as evolutionary computation, which attempts to mimic biological
evolution for the purpose of developing new computer algorithms. Discoveries in biotechnology now allow the
modification of entire genomes, advancing evolutionary studies to the level where future experiments may
involve the creation of entirely synthetic organisms.[103]
[edit] Microbiology, horizontal gene transfer, and endosymbiosis

Main article: Horizontal gene transfer
Microbiology was largely ignored by early evolutionary theory. This was due to the paucity of
morphological traits and the lack of a species concept in microbiology, particularly amongst prokaryotes.[104]
Now, evolutionary researchers are taking advantage of their improved understanding of microbial physiology
and ecology, produced by the comparative ease of microbial genomics, to explore the taxonomy and
evolution of these organisms.[105] These studies are revealing unanticipated levels of diversity amongst
microbes.[106][107]
One particularly important outcome from studies on microbial evolution was the discovery in Japan of
horizontal gene transfer in 1959.[108] This transfer of genetic material between different species of bacteria
was first recognized because it played a major role in the spread of antibiotic resistance.[109] More recently,
as knowledge of genomes has continued to expand, it has been suggested that lateral transfer of genetic
material has played an important role in the evolution of all organisms.[110] These high levels of horizontal
gene transfer have led to suggestions that the family tree of today's organisms, the so-called "tree of life", is
more similar to an interconnected web or net.[111][112]
Indeed, as part of the endosymbiotic theory for the origin of organelles, a form of horizontal gene
transfer has been a critical step in the evolution of eukaryotes such as fungi, plants, and animals.[113][114]
The endosymbiotic theory holds that organelles within the cells of eukorytes such as mitochondria and
chloroplasts, had descended from independent bacteria that came to live symbiotically within other cells. It
had been suggested in the late 19th century when similarities between mitochondria and bacteria were
noted, but largely dismissed until it was revived and championed by Lynn Margulis in the 1960s and 70s;
Margulis was able to make use of new evidence that such organelles had their own DNA that was inherited
independently from that in the cell's nucleus.[115]
[edit] Evolutionary developmental biology

Main article: Evolutionary developmental biology
In the 1980s and 1990s the tenets of the modern evolutionary synthesis came under increasing
scrutiny. There was a renewal of structuralist themes in evolutionary biology in the work of biologists such as
Brian Goodwin and Stuart Kauffman, which incorporated ideas from cybernetics and systems theory, and
emphasized the self-organizing processes of development as factors directing the course of evolution. The
evolutionary biologist Stephen Jay Gould revived earlier ideas of heterochrony, alterations in the relative
rates of developmental processes over the course of evolution, to account for the generation of novel forms,
and, with the evolutionary biologist Richard Lewontin, wrote an influential paper in 1979 suggesting that a
change in one biological structure, or even a structural novelty, could arise incidentally as an accidental
result of selection on another structure, rather than through direct selection for that particular adaptation.
They called such incidental structural changes "spandrels" after an architectural feature.[116] Later, Gould
and Vrba discussed the acquisition of new functions by novel structures arising in this fashion, calling them
"exaptations".[117]
Molecular data regarding the mechanisms underlying development accumulated rapidly during the
1980s and '90s. It became clear that the diversity of animal morphology was not the result of different sets of
proteins regulating the development of different animals, but from changes in the deployment of a small set
of proteins that were common to all animals.[118] These proteins became known as the "developmental
toolkit".[119] Such perspectives influenced the disciplines of phylogenetics, paleontology and comparative
developmental biology, and spawned the new discipline of evolutionary developmental biology.[120]
More recent work in this field by Mary Jane West-Eberhard has emphasized phenotypic and
developmental plasticity.[121] It has been suggested, for example, that the rapid emergence of basic animal
body plans in the Cambrian explosion was due in part to changes in the environment acting on inherent
material properties of cell aggregates, such as differential cell adhesion and biochemical oscillation. The
resulting forms were later stabilized by natural selection.[122] Experimental and theoretical research on
these and related ideas have been presented in the multi-authored volume Origination of Organismal Form.
[edit] 21st century

[edit] Epigenetic inheritance
Main article: Epigenetics
Yet another area where developmental biology has led to the questioning of some tenets of the
modern evolutionary synthesis is in the field of epigenetics, the study of how environmental factors affect the
way genes express themselves during development. By the first decade of the 21st century it had become
accepted that in some cases such environmental factors could affect the expression of genes in subsequent
generations even though the offspring were not exposed to the same environmental factors, and there had
been no genetic changes. This shows that in some cases non genetic changes to an organism can be
inherited and it has been suggested that such inheritance can help with adaptation to local conditions and
affect evolution.[123][124] Some have suggested that in some cases a form of Lamarckian evolution may
occur.[125]
[edit] Unconventional evolutionary theory

[edit] Omega Point
Pierre Teilhard de Chardin's metaphysical Omega Point Theory describes the gradual development
of the universe from subatomic particles to human society, which he viewed as its final stage and goal.
[edit] Gaia hypothesis

Teilhard de Chardin's ideas have been seen as being connected to the more specific Gaia theory by
James Lovelock, who proposed that the living and nonliving parts of Earth can be viewed as a complex
interacting system with similarities to a single organism.[126] The Gaia hypothesis has also been viewed by
Lynn Margulis[127] and others as an extension of endosymbiosis and exosymbiosis.[128] This modified
hypothesis postulates that all living things have a regulatory effect on the Earth's environment that promotes
life overall.
[edit] Transhumanism
Futurists have often viewed scientific and technological progress as a continuation of biological
evolution. Among these, transhumanists often view such technological evolution itself as a goal in their
philosophy, possibly in the form of a technological singularity.
[edit] See also
Book:Evolution

• Faith and rationality
• Galápagos Islands
• The Voyage of the Beagle
[edit] Notes
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11.
6. ^ a b Singer 1931
7. ^ Needham & Ronan 1995, p. 101
8. ^ Miller, James (January 8, 2008). "Daoism and Nature" (PDF). Royal Asiatic Society.
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9. ^ Sedley, David (August 4, 2004). "Lucretius". Stanford Encyclopedia of Philosophy.
http://plato.stanford.edu/entries/lucretius/. Retrieved 2008-07-24.
10.^ Simpson, David (2006). "Lucretius". The Internet Encyclopedia of Philosophy.
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11.^ Augustine 1982, pp. 89–90
12.^ Gill 2005, p. 251
13.^ Owen, Richard (2009-02-11). "Vatican buries the hatchet with Charles Darwin". London:
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14.^ Irvine, Chris (2009-02-12). "The Vatican claims Darwin's theory of evolution is compatible
with Christianity". London: Telegraph.co.uk.
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22.^ Schelling, System of Transcendental Idealism, 1800
23.^ Bowler 2003, pp. 73–75
24.^ Bowler 2003, pp. 75–80
25.^ Larson 2004, pp. 14–15
26.^ Henderson 2000
27.^ Darwin 1818, Vol I section XXXIX
28.^ Darwin 1825, p. 15
29.^ Larson 2004, p. 7
30.^ American Museum of Natural History (2000). "James Hutton: The Founder of Modern
Geology". Earth: Inside and Out.
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31.^ Bowler 2003, p. 113
32.^ Larson 2004, pp. 29–38
33.^ Bowler 2003, pp. 115–116
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36.^ Bowler 2003, pp. 86–94
37.^ Larson 2004, pp. 38–41
38.^ Desmon & Moore 1993, p. 40
39.^ a b Bowler 2003, pp. 120–129
40.^ Bowler 2003, pp. 134–138
41.^ Bowler & Morus 2005, pp. 142–143
42.^ Larson 2004, pp. 5–24
43.^ Bowler 2003, pp. 103–104
44.^ Larson 2004, pp. 37–38
45.^ Bowler 2003, p. 138
46.^ Larson 2004, pp. 42–46
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49.^ Darwin 1866, p. xiv
50.^ Bowler 2003, p. 151
51.^ Darwin 1859, pp. 62
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57.^ Larson 2004, pp. 55–71
58.^ Bowler 2003, pp. 173–176
59.^ Larson 2004, p. 50
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62.^ Larson 2004, pp. 139–40
63.^ Larson 2004, pp. 109–110
64.^ Bowler 2003, pp. 190–191
65.^ Bowler 2003, pp. 177–223
66.^ Larson 2004, pp. 121–123, 152–157
67.^ a b c Bowler 2003, pp. 207–216
68.^ Bowler 2003, pp. 49–51
69.^ a b c d Larson 2004, pp. 105–129
70.^ a b c d Bowler 2003, pp. 196–253
71.^ a b Bowler 2003, pp. 256–273
72.^ a b Larson 2004, pp. 153–174
73.^ a b c d e Bowler 2003, pp. 325–339
74.^ a b c d e Larson 2004, pp. 221–243
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[edit] References
• Augustine (1982). The Literal Meaning of Genesis. trans. John Hammond Taylor. The
Newman Press. ISBN 0809103265, 9780809103263. http://books.google.com/?id=_s0kIgD0nCcC.
• Bowler, Peter J. (2003). Evolution: The History of an Idea (3rd ed.). University of California
Press. ISBN 0-52023693-9.
• Bowler, Peter J.; Morus, Iwan Rhys (2005). Making Modern Science. The University of
Chicago Press. ISBN 0-226-06861-7.
• Darlington, Cyril (1959). Darwin's place in history. Blackwell, Oxford, p85.
• Darwin, Charles (1859). On the Origin of Species by Means of Natural Selection, or the
Preservation of Favoured Races in the Struggle for Life (1st ed.). John Murray, London.
Preservation of Favoured Races in the Struggle for Life (3rd ed.). John Murray, London.
Preservation of Favoured Races in the Struggle for Life (4th ed.). John Murray, London.
Preservation of Favoured Races in the Struggle for Life (6th ed.). John Murray, London.
• Darwin, Erasmus (1825). The Temple of Nature, or The Origin of Society. Jones & Company.
http://books.google.com/?id=oAl9y-0FSJQC&dq=Erasmus+Darwin+Temple.
• Darwin, Erasmus (1818). Zoonomia. http://www.gutenberg.org/files/15707/15707-h/15707-
h.htm.
• Desmond, Adrian; Moore, James (1994). Darwin: The Life of a Tormented Evolutionist. W.
W. Norton & Company. ISBN 0393311503.
• Gill, Meredith Jane (2005). Augustine in the Italian Renaissance: Art and Philosophy from
Petrarch to Michelangelo. Cambridge University Press. ISBN 0521832144, ISBN 9780521832144.
http://books.google.com/?id=AksE4jKI8yQC.
• Gould, Stephen Jay (2002). The Structure of Evolutionary Theory. Belknap Press of Harvard
University Press. ISBN 0-674-00613-5.
• Henderson, Jan-Andrew (2000). The Emperor's Kilt: The Two Secret Histories of Scotland .
Mainstream Publishing.
• Larson, Edward J. (2004). Evolution: The Remarkable History of a Scientific Theory. Modern
Library. ISBN 0-679-64288-9.
• Lovejoy, Arthur (1936). The Great Chain of Being: A Study of the History of an Idea . Harvard
• Mayr, Ernst (1982). The Growth of Biological Thought: Diversity, Evolution, and Inheritance .
The Belknap Press of Harvard University Press. ISBN 0-674-36445-7.
• Mayr, Ernst; Provine, W. B., eds (1998). The Evolutionary Synthesis: Perspectives on the
Unification of Biology. Harvard University Press. ISBN 0-674-27225-0.
• Needham, Joseph; Ronan, Colin Alistair (1995). The Shorter Science and Civilisation in
China: An Abridgement of Joseph Needham's Original Text, Vol. 1. Cambridge University Press.
ISBN 0521292867.
• Sapp, Jan (2003). Genesis: The Evolution of Biology. Oxford University Press. ISBN 0-19-
515618-8.
• Secord, James A. (2000). Victorian Sensation: The Extraordinary Publication, Reception,
and Secret Authorship of Vestiges of the Natural History of Creation . University of Chicago Press.
ISBN 0-226-74410-8.
• Singer, Charles (1931). A Short History of Biology. Clarendon Press.
• Smocovitis, Vassiliki Betty (1996). Unifying Biology: The Evolutionary Synthesis and
Evolutionary Biology. Princeton University Press. ISBN 0-691-03343-9.
• This article incorporates text from a publication now in the public domain: Chisholm,
Hugh, ed (1911). Encyclopædia Britannica (Eleventh ed.). Cambridge University Press.

• UC Berkeley's History of Evolutionary Thought
• Darwin's precursors and influences by John Wilkins . Part of the Talk.Origins Archive.
• The Alfred Russel Wallace Page
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Evolution

This article is about evolution in biology. For other uses, see Evolution (disambiguation).
For a generally accessible and less technical introduction to the topic, see Introduction to evolution.
Evolution
Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation
Introduction
Evidence
History
Level of support
Modern synthesis
Social effect
Theory and fact
Cladistics
Ecological genetics
Molecular evolution
Phylogenetics
Population genetics
Systematics
Evolution (also known as biological or organic evolution) is the change over time in the proportion of
individual organisms differing in one or more inherited traits.[1] A trait is a particular characteristic—
anatomical, biochemical or behavioural—that is the result of gene–environment interaction. Evolution may
occur when there is variation of inherited traits within a population. The major sources of such variation are
mutation, genetic recombination and gene flow.[2][3][4][5]
Two processes are generally distinguished as common causes of evolution. One is natural selection,
a process in which there is differential survival and/or reproduction of organisms that differ in one or more
inherited traits.[1] Another cause is genetic drift, a process in which there are random changes to the
proportions of two or more inherited traits within a population.[6][7]
A notable result of evolution is speciation, in which a single ancestral species splits and diversifies
into two or more different species. Speciation is visible in anatomical, genetic and other similarities between
groups of organisms, geographical distribution of related species, the fossil record and the recorded genetic
changes in living organisms over many generations. Speciation stretches back over 3.5 billion years during
which life has existed on earth.[8][9][10][11] It is thought to occur in multiple ways such as slowly, steadily
and gradually over time (see gradualism) or rapidly from one long static state to another (see punctuated
equilibrium).
The scientific study of evolution began in the mid-nineteenth century, when research into the fossil
record and the diversity of living organisms convinced most scientists that species evolve.[12] The
mechanism driving these changes remained unclear until the theory of natural selection was independently
proposed by Charles Darwin and Alfred Wallace in 1858. In the early 20th century, Darwinian theories of
evolution were combined with genetics, palaeontology, and systematics, which culminated into a union of
ideas known as the modern evolutionary synthesis.[13] The synthesis became a major principle of biology as
it provided a coherent and unifying explanation for the history and diversity of life on Earth.[14][15][16]
Evolution is currently applied and studied in various areas within biology such as conservation
biology, developmental biology, ecology, physiology, paleontology and medicine. Moreover, it has also made
an impact on traditionally non-biological disciplines such as agriculture, anthropology, philosophy and
psychology.
Contents
[hide]
• 1 History of evolutionary thought
• 2 Heredity
• 3 Variation
• 3.1 Mutation
• 3.2 Sex and recombination
• 3.3 Population genetics
• 3.4 Gene flow
• 4 Mechanisms
• 4.1 Natural selection
• 4.2 Genetic drift
• 5 Outcomes
• 5.1 Adaptation
• 5.2 Co-evolution
• 5.3 Co-operation
• 5.4 Speciation
• 5.5 Extinction
• 6 Evolutionary history of life
• 6.1 Origin of life
• 6.2 Common descent
• 6.3 Evolution of life
For more details on this topic, see History of evolutionary thought.
Around 1854 Charles Darwin began writing out what became On the Origin of Species.
The roots of naturalistic thinking on biology can be dated to at least the 6th century BCE, with the
Greek philosopher Anaximander.[17]
Early Christian Church Fathers and Medieval European scholars treated the Genesis creation myth
as allegory and believed that natural organisms were unstable and capricious, but the Protestant
Reformation inspired Biblical literalism and a natural theology in which species were static and fixed.
As emerging science explored mechanical philosophy in the 18th century, proto-evolutionary ideas
were set out by a few natural philosophers such as Pierre Maupertuis in 1745 and Erasmus Darwin in 1796.
[18]
The word evolution itself (from the Latin evolutio, meaning "to unroll like a scroll") was initially used to
refer to embryological development; its first use in relation to development of species came in 1762, when
Charles Bonnet used it for his concept of "pre-formation", in which females carried a miniature form of all
future generations. The term gradually gained a more general meaning of growth or progressive
development.[19] The first published modern use of the word has been attributed to the Edinburgh New
Philosophical Journal in 1826, edited by Robert Jameson but arguably authored by Robert Edmond Grant.
[20]
The Bible-based Ussher chronology of the 1650s had calculated creation at 4004 BC, but by the
1780s geologists assumed a much older world. Wernerians thought strata were deposits from shrinking seas,
but James Hutton proposed a self-maintaining infinite cycle. Georges Cuvier's paleontological work in the
1790s established the reality of extinction, which he explained by local catastrophes, followed by
repopulation of the affected areas by other species. He held that species were fixed, and marginalised the
ideas of the biologist Jean-Baptiste Lamarck about transmutation of species which were only taken up by
radicals.
Geologists such as Adam Sedgwick adapted Cuvier's catastrophism to show repeated worldwide
annihilation and creation of new fixed species adapted to a changed environment, initially identifying the
most recent catastrophe as the biblical flood. In opposition to this view, Charles Lyell adapted Hutton's
concept into a stricter uniformitarianism which strongly influenced the young geologist Charles Darwin during
the Beagle expedition. Darwin initially followed Lyell's idea of repeated "centres of creation" of fixed species,
but questioned Lyell's views and in 1836, near the end of the voyage, he expressed doubts that species were
fixed. Darwin formulated his idea of natural selection in 1838 and was still developing his theory in 1858
when Alfred Russel Wallace sent him a similar theory, and both were presented to the Linnean Society of
London in separate papers.[21] At the end of 1859, Darwin's publication of On the Origin of Species
explained natural selection in detail and presented evidence leading to increasingly wide acceptance of the
occurrence of evolution.
Debate about the mechanisms of evolution continued, and Darwin could not explain the source of the
heritable variations which would be acted on by natural selection.[22] Like Lamarck, he still thought that
parents passed on adaptations acquired during their lifetimes,[23] a theory which was subsequently dubbed
Lamarckism.[24] In the 1880s, August Weismann's experiments indicated that changes from use and disuse
were not heritable, and Lamarckism gradually fell from favour.[25][26] More significantly, Darwin could not
account for how traits were passed down from generation to generation. In 1865 Gregor Mendel found that
traits were inherited in a predictable manner.[27] When Mendel's work was rediscovered in the 1900s,
disagreements over the rate of evolution predicted by early geneticists and biometricians led to a rift between
the Mendelian and Darwinian models of evolution.
Yet it was the rediscovery of Gregor Mendel's pioneering work on the fundamentals of genetics (of
which Darwin and Wallace were unaware) by Hugo de Vries and others in the early 1900s that provided the
impetus for a better understanding of how variation occurs in plant and animal traits. That variation is the
main fuel used by natural selection to shape the wide variety of adaptive traits observed in organic life. Even
though Hugo de Vries and other early geneticists rejected gradual natural selection, their rediscovery of and
subsequent work on genetics eventually provided a solid basis on which the theory of evolution stood even
more convincingly than when it was originally proposed.[28]
The apparent contradiction between Darwin's theory of evolution by natural selection and Mendel's
work was reconciled in the 1920s and 1930s by evolutionary biologists such as J.B.S. Haldane, Sewall
Wright, and particularly Ronald Fisher, who set the foundations for the establishment of the field of population
genetics. The end result was a combination of evolution by natural selection and Mendelian inheritance, the
modern evolutionary synthesis.[29] In the 1940s, the identification of DNA as the genetic material by Oswald
Avery and colleagues and the subsequent publication of the structure of DNA by James Watson and Francis
Crick in 1953, demonstrated the physical basis for inheritance. Since then, genetics and molecular biology
have become core parts of evolutionary biology and have revolutionised the field of phylogenetics.[13]
In its early history, evolutionary biology primarily drew in scientists from traditional taxonomically
oriented disciplines, whose specialist training in particular organisms addressed general questions in
evolution. As evolutionary biology expanded as an academic discipline, particularly after the development of
the modern evolutionary synthesis, it began to draw more widely from the biological sciences.[13] Currently
the study of evolutionary biology involves scientists from fields as diverse as biochemistry, ecology, genetics
and physiology, and evolutionary concepts are used in even more distant disciplines such as psychology,
medicine, philosophy and computer science.
In the 21st century, current research in evolutionary biology deals with several areas where the
modern evolutionary synthesis may need modification or extension, such as assessing the relative
importance of various ideas on the unit of selection and evolvability and how to fully incorporate the findings
of evolutionary developmental biology.[30][31]
Heredity
Further information: Introduction to genetics, Genetics, and Heredity
DNA structure. Bases are in the centre, surrounded by phosphate–sugar chains in a double helix.
Evolution in organisms occurs through changes in heritable traits – particular characteristics of an
organism. In humans, for example, eye colour is an inherited characteristic and an individual might inherit the
"brown-eye trait" from one of their parents.[32] Inherited traits are controlled by genes and the complete set
of genes within an organism's genome is called its genotype.[33]
The complete set of observable traits that make up the structure and behaviour of an organism is
called its phenotype. These traits come from the interaction of its genotype with the environment.[34] As a
result, many aspects of an organism's phenotype are not inherited. For example, suntanned skin comes from
the interaction between a person's genotype and sunlight; thus, suntans are not passed on to people's
children. However, some people tan more easily than others, due to differences in their genotype; a striking
example are people with the inherited trait of albinism, who do not tan at all and are very sensitive to
sunburn.[35]
Heritable traits are known to be passed from one generation to the next via DNA, a molecule that
encodes genetic information.[33] DNA is a long polymer composed of four types of bases. The sequence of
bases along a particular DNA molecule specify the genetic information, in a manner similar to a sequence of
letters spelling out a sentence. Before a cell divides, the DNA is copied, so that each of the resulting two cells
will inherit the DNA sequence. Portions of a DNA molecule that specify a single functional unit are called
genes; different genes have different sequences of bases. Within cells, the long strands of DNA form
condensed structures called chromosomes. The specific location of a DNA sequence within a chromosome is
known as a locus. If the DNA sequence at a locus varies between individuals, the different forms of this
sequence are called alleles. DNA sequences can change through mutations, producing new alleles. If a
mutation occurs within a gene, the new allele may affect the trait that the gene controls, altering the
phenotype of the organism.[1]
However, while this simple correspondence between an allele and a trait works in some cases, most
traits are more complex and are controlled by multiple interacting genes within and among organisms.[36]
[37] Developmental biologists suggest that complex interactions in genetic networks and communication
among cells can lead to heritable variations that may underlay some of the mechanics in developmental
plasticity and canalization.[38] Recent findings have confirmed important examples of heritable changes that
cannot be explained by direct agency of the DNA molecule. These phenomena are classed as epigenetic
inheritance systems that are causally or independently evolving over genes.
Research into modes and mechanisms of epigenetic inheritance is still in its scientific infancy,
however, this area of research has attracted much recent activity as it broadens the scope of heritability and
evolutionary biology in general.[39] DNA methylation marking chromatin, self-sustaining metabolic loops,
gene silencing by RNA interference, and the three dimensional conformation of proteins (such as prions) are
areas where epigenetic inheritance systems have been discovered at the organismic level.[40][41]
Heritability may also occur at even larger scales. For example, ecological inheritance through the process of
niche construction is defined by the regular and repeated activities of organisms in their environment. This
generates a legacy of effect that modifies and feeds back into the selection regime of subsequent
generations. Descendants inherit genes plus environmental characteristics generated by the ecological
actions of ancestors.[42] Other examples of heritability in evolution that are not under the direct control of
genes include the inheritance of cultural traits, group heritability, and symbiogenesis.[43][44][45] These
examples of heritability that operate above the gene are covered broadly under the title of multilevel or
hierarchical selection, which has been a subject of intense debate in the history of evolutionary science.[44]
[46]
Variation
Further information: Genetic diversity and Population genetics
An individual organism's phenotype results from both its genotype and the influence from the
environment it has lived in. A substantial part of the variation in phenotypes in a population is caused by the
differences between their genotypes.[37] The modern evolutionary synthesis defines evolution as the change
over time in this genetic variation. The frequency of one particular allele will fluctuate, becoming more or less
prevalent relative to other forms of that gene. Evolutionary forces act by driving these changes in allele
frequency in one direction or another. Variation disappears when a new allele reaches the point of fixation —
when it either disappears from the population or replaces the ancestral allele entirely.[47]
Variation comes from mutations in genetic material, migration between populations (gene flow), and
the reshuffling of genes through sexual reproduction. Variation also comes from exchanges of genes
between different species; for example, through horizontal gene transfer in bacteria, and hybridisation in
plants.[48] Despite the constant introduction of variation through these processes, most of the genome of a
species is identical in all individuals of that species.[49] However, even relatively small changes in genotype
can lead to dramatic changes in phenotype: for example, chimpanzees and humans differ in only about 5%
of their genomes.[50]
Mutation
Further information: Mutation and Molecular evolution
Duplication of part of a chromosome.
Random mutations constantly occur in the genomes of organisms; these mutations create genetic
variation. Mutations are changes in the DNA sequence of a cell's genome and are caused by radiation,
viruses, transposons and mutagenic chemicals, as well as errors that occur during meiosis or DNA
replication.[51][52][53] These mutations involve several different types of change in DNA sequences; these
can either have no effect, alter the product of a gene, or prevent the gene from functioning. Studies in the fly
Drosophila melanogaster suggest that if a mutation changes a protein produced by a gene, this will probably
be harmful, with about 70% of these mutations having damaging effects, and the remainder being either
neutral or weakly beneficial.[54]
Due to the damaging effects that mutations can have on cells, organisms have evolved mechanisms
such as DNA repair to remove mutations.[51] Therefore, the optimal mutation rate for a species is a trade-off
between costs of a high mutation rate, such as deleterious mutations, and the metabolic costs of maintaining
systems to reduce the mutation rate, such as DNA repair enzymes.[55] Viruses that use RNA as their genetic
material have rapid mutation rates,[56] which can be an advantage since these viruses will evolve constantly
and rapidly, and thus evade the defensive responses of e.g. the human immune system.[57]
Mutations can involve large sections of a chromosome becoming duplicated (usually by genetic
recombination), which can introduce extra copies of a gene into a genome.[58] Extra copies of genes are a
major source of the raw material needed for new genes to evolve.[59] This is important because most new
genes evolve within gene families from pre-existing genes that share common ancestors.[60] For example,
the human eye uses four genes to make structures that sense light: three for colour vision and one for night
vision; all four are descended from a single ancestral gene.[61]
New genes can be created from an ancestral gene when a duplicate copy mutates and acquires a
new function. This process is easier once a gene has been duplicated because it increases the redundancy
of the system; one gene in the pair can acquire a new function while the other copy continues to perform its
original function.[62][63] Other types of mutation can even create entirely new genes from previously
noncoding DNA.[64][65]
The creation of new genes can also involve small parts of several genes being duplicated, with these
fragments then recombining to form new combinations with new functions.[66][67] When new genes are
assembled from shuffling pre-existing parts, domains act as modules with simple independent functions,
which can be mixed together creating new combinations with new and complex functions.[68] For example,
polyketide synthases are large enzymes that make antibiotics; they contain up to one hundred independent
domains that each catalyze one step in the overall process, like a step in an assembly line.[69]
Changes in chromosome number may involve even larger mutations, where segments of the DNA
within chromosomes break and then rearrange. For example, two chromosomes in the Homo genus fused to
produce human chromosome 2; this fusion did not occur in the lineage of the other apes, and they retain
these separate chromosomes.[70] In evolution, the most important role of such chromosomal
rearrangements may be to accelerate the divergence of a population into new species by making populations
less likely to interbreed, and thereby preserving genetic differences between these populations.[71]
Sequences of DNA that can move about the genome, such as transposons, make up a major fraction
of the genetic material of plants and animals, and may have been important in the evolution of genomes.[72]
For example, more than a million copies of the Alu sequence are present in the human genome, and these
sequences have now been recruited to perform functions such as regulating gene expression.[73] Another
effect of these mobile DNA sequences is that when they move within a genome, they can mutate or delete
existing genes and thereby produce genetic diversity.[52]
Sex and recombination

Further information: Genetic recombination and Sexual reproduction
In asexual organisms, genes are inherited together, or linked, as they cannot mix with genes of other
organisms during reproduction. In contrast, the offspring of sexual organisms contain random mixtures of
their parents' chromosomes that are produced through independent assortment. In a related process called
homologous recombination, sexual organisms exchange DNA between two matching chromosomes.[74]
Recombination and reassortment do not alter allele frequencies, but instead change which alleles are
associated with each other, producing offspring with new combinations of alleles.[75] Sex usually increases
genetic variation and may increase the rate of evolution.[76][77] However, asexuality is advantageous in
some environments as it can evolve in previously sexual animals.[78] Here, asexuality might allow the two
sets of alleles in their genome to diverge and gain different functions.[79]
Recombination allows even alleles that are close together in a strand of DNA to be inherited
independently. However, the rate of recombination is low (approximately two events per chromosome per
generation). As a result, genes close together on a chromosome may not always be shuffled away from each
other, and genes that are close together tend to be inherited together, a phenomenon known as linkage.[80]
This tendency is measured by finding how often two alleles occur together on a single chromosome, which is
called their linkage disequilibrium. A set of alleles that is usually inherited in a group is called a haplotype.
This can be important when one allele in a particular haplotype is strongly beneficial: natural selection can
drive a selective sweep that will also cause the other alleles in the haplotype to become more common in the
population; this effect is called genetic hitchhiking.[81]
When alleles cannot be separated by recombination – such as in mammalian Y chromosomes, which
pass intact from fathers to sons – harmful mutations accumulate.[82][83] By breaking up allele combinations,
sexual reproduction allows the removal of harmful mutations and the retention of beneficial mutations.[84] In
addition, recombination and reassortment can produce individuals with new and advantageous gene
combinations. These positive effects are balanced by the fact that sex reduces an organism's reproductive
rate, can cause mutations and may separate beneficial combinations of genes.[84] The reasons for the
evolution of sexual reproduction are therefore unclear and this question is still an active area of research in
evolutionary biology,[85][86] that has prompted ideas such as the Red Queen hypothesis.[87]
Population genetics
White peppered moth
Black morph in peppered moth evolution

Further information: Population genetics
From a genetic viewpoint, evolution is a generation-to-generation change in the frequencies of
alleles within a population that shares a common gene pool .[88] A population is a localised group of
individuals belonging to the same species. For example, all of the moths of the same species living in an
isolated forest represent a population. A single gene in this population may have several alternate forms,
which account for variations between the phenotypes of the organisms. An example might be a gene for
colouration in moths that has two alleles: black and white.
A gene pool is the complete set of alleles for a gene in a single population; the allele frequency
measures the fraction of the gene pool composed of a single allele (for example, what fraction of moth
colouration genes are the black allele). Evolution occurs when there are changes in the frequencies of alleles
within a population of interbreeding organisms; for example, the allele for black colour in a population of
moths becoming more common.
To understand the mechanisms that cause a population to evolve, it is useful to consider what
conditions are required for a population not to evolve. The Hardy-Weinberg principle states that the
frequencies of alleles (variations in a gene) in a sufficiently large population will remain constant if the only
forces acting on that population are the random reshuffling of alleles during the formation of the sperm or
egg, and the random combination of the alleles in these sex cells during fertilisation.[89] Such a population is
said to be in Hardy-Weinberg equilibrium; it is not evolving.[90]
Gene flow
Further information: Gene flow, Hybrid (biology), and Horizontal gene transfer
When they mature, male lions leave the pride where they were born and take over a new pride to
mate, causing gene flow between prides.[91]
Gene flow is the exchange of genes between populations, which are usually of the same species.[92]
Examples of gene flow within a species include the migration and then breeding of organisms, or the
exchange of pollen. Gene transfer between species includes the formation of hybrid organisms and
horizontal gene transfer.
Migration into or out of a population can change allele frequencies, as well as introducing genetic
variation into a population. Immigration may add new genetic material to the established gene pool of a
population. Conversely, emigration may remove genetic material. As barriers to reproduction between two
diverging populations are required for the populations to become new species, gene flow may slow this
process by spreading genetic differences between the populations. Gene flow is hindered by mountain
ranges, oceans and deserts or even man-made structures such as the Great Wall of China, which has
hindered the flow of plant genes.[93]
Depending on how far two species have diverged since their most recent common ancestor, it may
still be possible for them to produce offspring, as with horses and donkeys mating to produce mules.[94]
Such hybrids are generally infertile, due to the two different sets of chromosomes being unable to pair up
during meiosis. In this case, closely related species may regularly interbreed, but hybrids will be selected
against and the species will remain distinct. However, viable hybrids are occasionally formed and these new
species can either have properties intermediate between their parent species, or possess a totally new
phenotype.[95] The importance of hybridisation in creating new species of animals is unclear, although cases
have been seen in many types of animals,[96] with the gray tree frog being a particularly well-studied
example.[97]
Hybridisation is, however, an important means of speciation in plants, since polyploidy (having more
than two copies of each chromosome) is tolerated in plants more readily than in animals.[98][99] Polyploidy is
important in hybrids as it allows reproduction, with the two different sets of chromosomes each being able to
pair with an identical partner during meiosis.[100] Polyploids also have more genetic diversity, which allows
them to avoid inbreeding depression in small populations.[101]
Horizontal gene transfer is the transfer of genetic material from one organism to another organism
that is not its offspring; this is most common among bacteria.[102] In medicine, this contributes to the spread
of antibiotic resistance, as when one bacteria acquires resistance genes it can rapidly transfer them to other
species.[103] Horizontal transfer of genes from bacteria to eukaryotes such as the yeast Saccharomyces
cerevisiae and the adzuki bean beetle Callosobruchus chinensis may also have occurred.[104][105] An
example of larger-scale transfers are the eukaryotic bdelloid rotifers, which appear to have received a range
of genes from bacteria, fungi, and plants.[106] Viruses can also carry DNA between organisms, allowing
transfer of genes even across biological domains.[107] Large-scale gene transfer has also occurred between
the ancestors of eukaryotic cells and prokaryotes, during the acquisition of chloroplasts and mitochondria.
[108]
Mechanisms
The two main mechanisms that produce evolution are natural selection and genetic drift. Natural
selection is the process which favours genes that aid survival and reproduction. Genetic drift is the random
change in the frequency of alleles, caused by the random sampling of a generation's genes during
reproduction. The relative importance of natural selection and genetic drift in a population varies depending
on the strength of the selection and the effective population size, which is the number of individuals capable
of breeding.[109] Natural selection usually predominates in large populations, whereas genetic drift
dominates in small populations. The dominance of genetic drift in small populations can even lead to the
fixation of slightly deleterious mutations.[110] As a result, changing population size can dramatically influence
the course of evolution. Population bottlenecks, where the population shrinks temporarily and therefore loses
genetic variation, result in a more uniform population.[47]
Natural selection
Further information: Natural selection and Fitness (biology)
Natural selection of a population for dark colouration.
Natural selection is the process by which genetic mutations that enhance reproduction become, and
remain, more common in successive generations of a population. It has often been called a "self-evident"
mechanism because it necessarily follows from three simple facts:
• Heritable variation exists within populations of organisms.
• Organisms produce more offspring than can survive.
• These offspring vary in their ability to survive and reproduce.
These conditions produce competition between organisms for survival and reproduction.
Consequently, organisms with traits that give them an advantage over their competitors pass these
advantageous traits on, while traits that do not confer an advantage are not passed on to the next generation.
[111]
The central concept of natural selection is the evolutionary fitness of an organism.[112] Fitness is
measured by an organism's ability to survive and reproduce, which determines the size of its genetic
contribution to the next generation.[112] However, fitness is not the same as the total number of offspring:
instead fitness is indicated by the proportion of subsequent generations that carry an organism's genes.[113]
For example, if an organism could survive well and reproduce rapidly, but its offspring were all too small and
weak to survive, this organism would make little genetic contribution to future generations and would thus
have low fitness.[112]
If an allele increases fitness more than the other alleles of that gene, then with each generation this
allele will become more common within the population. These traits are said to be "selected for". Examples of
traits that can increase fitness are enhanced survival, and increased fecundity. Conversely, the lower fitness
caused by having a less beneficial or deleterious allele results in this allele becoming rarer — they are
"selected against".[114] Importantly, the fitness of an allele is not a fixed characteristic; if the environment
changes, previously neutral or harmful traits may become beneficial and previously beneficial traits become
harmful.[1] However, even if the direction of selection does reverse in this way, traits that were lost in the past
may not re-evolve in an identical form (see Dollo's law).[115][116]
A chart showing three types of selection. 1.Disruptive selection 2.Stabilizing selection 3.Directional
selection
Natural selection within a population for a trait that can vary across a range of values, such as height,
can be categorised into three different types. The first is directional selection, which is a shift in the average
value of a trait over time — for example, organisms slowly getting taller.[117] Secondly, disruptive selection is
selection for extreme trait values and often results in two different values becoming most common, with
selection against the average value. This would be when either short or tall organisms had an advantage, but
not those of medium height. Finally, in stabilizing selection there is selection against extreme trait values on
both ends, which causes a decrease in variance around the average value and less diversity.[111][118] This
would, for example, cause organisms to slowly become all the same height.
A special case of natural selection is sexual selection, which is selection for any trait that increases
mating success by increasing the attractiveness of an organism to potential mates.[119] Traits that evolved
through sexual selection are particularly prominent in males of some animal species, despite traits such as
cumbersome antlers, mating calls or bright colours that attract predators, decreasing the survival of individual
males.[120] This survival disadvantage is balanced by higher reproductive success in males that show these
hard to fake, sexually selected traits.[121]
Natural selection most generally makes nature the measure against which individuals, and individual
traits, are more or less likely to survive. "Nature" in this sense refers to an ecosystem, that is, a system in
which organisms interact with every other element, physical as well as biological, in their local environment.
Eugene Odum, a founder of ecology, defined an ecosystem as: "Any unit that includes all of the
organisms...in a given area interacting with the physical environment so that a flow of energy leads to clearly
defined trophic structure, biotic diversity, and material cycles (ie: exchange of materials between living and
nonliving parts) within the system."[122] Each population within an ecosystem occupies a distinct niche, or
position, with distinct relationships to other parts of the system. These relationships involve the life history of
the organism, its position in the food chain, and its geographic range. This broad understanding of nature
enables scientists to delineate specific forces which, together, comprise natural selection.
An active area of research is the unit of selection, with natural selection being proposed to work at
the level of genes, cells, individual organisms, groups of organisms and species.[123][124] None of these are
mutually exclusive and selection can act on multiple levels simultaneously.[125] An example of selection
occurring below the level of the individual organism are genes called transposons, which can replicate and
spread throughout a genome.[126] Selection at a level above the individual, such as group selection, may
allow the evolution of co-operation, as discussed below.[127]
Genetic drift
Further information: Genetic drift and Effective population size
Simulation of genetic drift of 20 unlinked alleles in populations of 10 (top) and 100 (bottom). Drift to
fixation is more rapid in the smaller population.
Genetic drift is the change in allele frequency from one generation to the next that occurs because
alleles in offspring are a random sample of those in the parents, as well as from the role that chance plays in
determining whether a given individual will survive and reproduce. In mathematical terms, alleles are subject
to sampling error. As a result, when selective forces are absent or relatively weak, allele frequencies tend to
"drift" upward or downward randomly (in a random walk). This drift halts when an allele eventually becomes
fixed, either by disappearing from the population, or replacing the other alleles entirely. Genetic drift may
therefore eliminate some alleles from a population due to chance alone. Even in the absence of selective
forces, genetic drift can cause two separate populations that began with the same genetic structure to drift
apart into two divergent populations with different sets of alleles.[128]
The time for an allele to become fixed by genetic drift depends on population size, with fixation
occurring more rapidly in smaller populations.[129] The precise measure of population that is important is
called the effective population size. The effective population is always smaller than the total population since
it takes into account factors such as the level of inbreeding, the number of animals that are too old or young
to breed, and the lower probability of animals that live far apart managing to mate with each other.[130]
An example of when genetic drift is probably of central importance in determining a trait is the loss of
pigments from animals that live in caves, a change that produces no obvious advantage or disadvantage in
complete darkness.[131] However, it is usually difficult to measure the relative importance of selection and
drift,[132] so the comparative importance of these two forces in driving evolutionary change is an area of
current research.[133] These investigations were prompted by the neutral theory of molecular evolution,
which proposed that most evolutionary changes are the result of the fixation of neutral mutations that do not
have any immediate effects on the fitness of an organism.[134] Hence, in this model, most genetic changes
in a population are the result of constant mutation pressure and genetic drift.[135] This form of the neutral
theory is now largely abandoned, since it does not seem to fit the genetic variation seen in nature.[136][137]
However, a more recent and better-supported version of this model is the nearly neutral theory, where most
mutations only have small effects on fitness.[111]
Outcomes
Evolution influences every aspect of the form and behaviour of organisms. Most prominent are the
specific behavioural and physical adaptations that are the outcome of natural selection. These adaptations
increase fitness by aiding activities such as finding food, avoiding predators or attracting mates. Organisms
can also respond to selection by co-operating with each other, usually by aiding their relatives or engaging in
mutually beneficial symbiosis. In the longer term, evolution produces new species through splitting ancestral
populations of organisms into new groups that cannot or will not interbreed[ citation needed].
These outcomes of evolution are sometimes divided into macroevolution, which is evolution that
occurs at or above the level of species, such as extinction and speciation, and microevolution, which is
smaller evolutionary changes, such as adaptations, within a species or population.[138] In general,
macroevolution is regarded as the outcome of long periods of microevolution.[139] Thus, the distinction
between micro- and macroevolution is not a fundamental one – the difference is simply the time involved.
[140] However, in macroevolution, the traits of the entire species may be important. For instance, a large
amount of variation among individuals allows a species to rapidly adapt to new habitats, lessening the
chance of it going extinct, while a wide geographic range increases the chance of speciation, by making it
more likely that part of the population will become isolated. In this sense, microevolution and macroevolution
might involve selection at different levels – with microevolution acting on genes and organisms, versus
macroevolutionary processes such as species selection acting on entire species and affecting their rates of
speciation and extinction.[141][142][143]
A common misconception is that evolution has goals or long-term plans; realistically however,
evolution has no long-term goal and does not necessarily produce greater complexity.[144][145] Although
complex species have evolved, they occur as a side effect of the overall number of organisms increasing,
and simple forms of life still remain more common in the biosphere.[146] For example, the overwhelming
majority of species are microscopic prokaryotes, which form about half the world's biomass despite their
small size,[147] and constitute the vast majority of Earth's biodiversity.[148] Simple organisms have therefore
been the dominant form of life on Earth throughout its history and continue to be the main form of life up to
the present day, with complex life only appearing more diverse because it is more noticeable.[149] Indeed,
the evolution of microorganisms is particularly important to modern evolutionary research, since their rapid
reproduction allows the study of experimental evolution and the observation of evolution and adaptation in
real time.[150][151]
Adaptation
For more details on this topic, see Adaptation.
Adaptation is one of the basic phenomena of biology,[152] and is the process whereby an organism
becomes better suited to its habitat.[153][154] Also, the term adaptation may refer to a trait that is important
for an organism's survival. For example, the adaptation of horses' teeth to the grinding of grass, or the ability
of horses to run fast and escape predators. By using the term adaptation for the evolutionary process, and
adaptive trait for the product (the bodily part or function), the two senses of the word may be distinguished.
Adaptations are produced by natural selection.[155] The following definitions are due to Theodosius
Dobzhansky.
1. Adaptation is the evolutionary process whereby an organism becomes better able to live in
its habitat or habitats.[156]
2. Adaptedness is the state of being adapted: the degree to which an organism is able to live
and reproduce in a given set of habitats.[157]
3. An adaptive trait is an aspect of the developmental pattern of the organism which enables or
enhances the probability of that organism surviving and reproducing.[158]
Adaptation may cause either the gain of a new feature, or the loss of an ancestral feature. An
example that shows both types of change is bacterial adaptation to antibiotic selection, with genetic changes
causing antibiotic resistance by both modifying the target of the drug, or increasing the activity of transporters
that pump the drug out of the cell.[159] Other striking examples are the bacteria Escherichia coli evolving the
ability to use citric acid as a nutrient in a long-term laboratory experiment,[160] Flavobacterium evolving a
novel enzyme that allows these bacteria to grow on the by-products of nylon manufacturing,[161][162] and
the soil bacterium Sphingobium evolving an entirely new metabolic pathway that degrades the synthetic
pesticide pentachlorophenol.[163][164] An interesting but still controversial idea is that some adaptations
might increase the ability of organisms to generate genetic diversity and adapt by natural selection
(increasing organisms' evolvability).[165][166]
A baleen whale skeleton, a and b label flipper bones, which were adapted from front leg bones: while
c indicates vestigial leg bones, suggesting an adaptation from land to sea.[167]
Adaptation occurs through the gradual modification of existing structures. Consequently, structures
with similar internal organisation may have different functions in related organisms. This is the result of a
single ancestral structure being adapted to function in different ways. The bones within bat wings, for
example, are very similar to those in mice feet and primate hands, due to the descent of all these structures
from a common mammalian ancestor.[168] However, since all living organisms are related to some extent,
[169] even organs that appear to have little or no structural similarity, such as arthropod, squid and vertebrate
eyes, or the limbs and wings of arthropods and vertebrates, can depend on a common set of homologous
genes that control their assembly and function; this is called deep homology.[170][171]
During adaptation, some structures may lose their original function and become vestigial structures.
[172] Such structures may have little or no function in a current species, yet have a clear function in ancestral
species, or other closely related species. Examples include pseudogenes,[173] the non-functional remains of
eyes in blind cave-dwelling fish,[174] wings in flightless birds,[175] and the presence of hip bones in whales
and snakes.[167] Examples of vestigial structures in humans include wisdom teeth,[176] the coccyx,[172] the
vermiform appendix,[172] and other behavioural vestiges such as goose bumps,[177] and primitive reflexes.
[178][179][180][181]
However, many traits that appear to be simple adaptations are in fact exaptations: structures
originally adapted for one function, but which coincidentally became somewhat useful for some other function
in the process.[182] One example is the African lizard Holaspis guentheri, which developed an extremely flat
head for hiding in crevices, as can be seen by looking at its near relatives. However, in this species, the head
has become so flattened that it assists in gliding from tree to tree—an exaptation.[182] Within cells, molecular
machines such as the bacterial flagella[183] and protein sorting machinery[184] evolved by the recruitment of
several pre-existing proteins that previously had different functions.[138] Another example is the recruitment
of enzymes from glycolysis and xenobiotic metabolism to serve as structural proteins called crystallins within
the lenses of organisms' eyes.[185][186]
A critical principle of ecology is that of competitive exclusion: no two species can occupy the same
niche in the same environment for a long time.[187] Consequently, natural selection will tend to force species
to adapt to different ecological niches. This may mean that, for example, two species of cichlid fish adapt to
live in different habitats, which will minimise the competition between them for food.[188]
An area of current investigation in evolutionary developmental biology is the developmental basis of
adaptations and exaptations.[189] This research addresses the origin and evolution of embryonic
development and how modifications of development and developmental processes produce novel features.
[190] These studies have shown that evolution can alter development to create new structures, such as
embryonic bone structures that develop into the jaw in other animals instead forming part of the middle ear in
mammals.[191] It is also possible for structures that have been lost in evolution to reappear due to changes
in developmental genes, such as a mutation in chickens causing embryos to grow teeth similar to those of
crocodiles.[192] It is now becoming clear that most alterations in the form of organisms are due to changes in
a small set of conserved genes.[193]
Co-evolution
Common garter snake (Thamnophis sirtalis sirtalis) which has evolved resistance to tetrodotoxin in
its amphibian prey.
Further information: Co-evolution
Interactions between organisms can produce both conflict and co-operation. When the interaction is
between pairs of species, such as a pathogen and a host, or a predator and its prey, these species can
develop matched sets of adaptations. Here, the evolution of one species causes adaptations in a second
species. These changes in the second species then, in turn, cause new adaptations in the first species. This
cycle of selection and response is called co-evolution.[194] An example is the production of tetrodotoxin in
the rough-skinned newt and the evolution of tetrodotoxin resistance in its predator, the common garter snake.
In this predator-prey pair, an evolutionary arms race has produced high levels of toxin in the newt and
correspondingly high levels of toxin resistance in the snake.[195]
Co-operation
Further information: Co-operation (evolution)
However, not all interactions between species involve conflict.[196] Many cases of mutually
beneficial interactions have evolved. For instance, an extreme cooperation exists between plants and the
mycorrhizal fungi that grow on their roots and aid the plant in absorbing nutrients from the soil.[197] This is a
reciprocal relationship as the plants provide the fungi with sugars from photosynthesis. Here, the fungi
actually grow inside plant cells, allowing them to exchange nutrients with their hosts, while sending signals
that suppress the plant immune system.[198]
Coalitions between organisms of the same species have also evolved. An extreme case is the
eusociality found in social insects, such as bees, termites and ants, where sterile insects feed and guard the
small number of organisms in a colony that are able to reproduce. On an even smaller scale, the somatic
cells that make up the body of an animal limit their reproduction so they can maintain a stable organism,
which then supports a small number of the animal's germ cells to produce offspring. Here, somatic cells
respond to specific signals that instruct them whether to grow, remain as they are, or die. If cells ignore these
signals and multiply inappropriately, their uncontrolled growth causes cancer.[51]
Such cooperation within species may have evolved through the process of kin selection, which is
where one organism acts to help raise a relative's offspring.[199] This activity is selected for because if the
helping individual contains alleles which promote the helping activity, it is likely that its kin will also contain
these alleles and thus those alleles will be passed on.[200] Other processes that may promote cooperation
include group selection, where cooperation provides benefits to a group of organisms.[201]
Speciation
Further information: Speciation
The four mechanisms of speciation.
Speciation is the process where a species diverges into two or more descendant species.[202]
Evolutionary scientists view the concept of "species" as a statistical phenomenon; this view is counterintuitive
since the classical idea of species is still widely held, with a species seen as a class of organisms exemplified
by a "type specimen" that bears all the traits common to this species.[203][204] Instead, a species is now
defined as a separately evolving lineage that forms a single gene pool. Although properties such as genetics
and morphology are used to help separate closely related lineages, this definition has fuzzy boundaries.[205]
Species are defined in two ways, taxonomically and categorically.[206] Species are partitioned
taxonmically into operational units for the practical application of framing evolutionary hypotheses in
systematics. Systematicists study and analyze the morphological or genetic characters from different
lineages and use parsimonious methods, such as cladistics or other statistical means to locate the position of
the taxon in the Linnean taxonomic hierarchy or biological classification. These methods create evolutionary
trees that are used to infer, illustrate, test, or explain evolutionary relations, historical patterns, and
phylogenetic transitions.[207][208] "Systematics is one of the oldest scientific disciplines and, from its
beginning, one of its central concepts has been the concept of species. Systematics can be characterized
generally as the branch of science devoted to the study of the different kinds of organisms (biological
diversity, in contemporary terms), and the term 'species' is Latin for 'kind.'"[206]:6600 When a new species is
discovered a type specimen and holotype specimens are usually deposited into a recognized or accredited
academic institution, such as a museum, that serves as a taxonomic reference point.
Species are also defined categorically by critical natural forces that best explain the evolutionary
mechanisms that are responsible for the crossing of the speciation threshold, from one species into two. In
this context, the exact definition of a "species" is still controversial, particularly in prokaryotes,[209] and this is
called the species problem.[206]
There is much diversity in life and varied biological reasons for speciation, which has resulted in
more than twenty different kinds of species concepts to facilitate the diverse modes, mechanisms, and
evolutionary processes. The concept that is used is a pragmatic choice that depends on the particularities of
the species concerned.[206] For example, some species concepts may apply more readily toward sexually
reproducing organisms and some lend themselves better toward asexual organisms. The various concepts,
however, can be placed into one of three general philosophical approaches: 1) the interbreeding, 2) the
ecological, and 3) the phylogenetic.[210] The biological species concept (BSC) is a classic example of the
interbreeding approach. Introduced by Ernst Mayr in 1942, the BSC states that "species are groups of
actually or potentially interbreeding natural populations, which are reproductively isolated from other such
groups"[211]:120. Despite its wide and long-term use, the BSC like others is not without controversy. Some
researchers have attempted a unifying monistic definition of species, while others adopt a pluralistic
approach and suggest that there may be a variety of ways to logically interpret what a species is.[206][210]
Speciation has been observed multiple times under both controlled laboratory conditions and in
nature.[212] In sexually reproducing organisms, speciation results from reproductive isolation followed by
genealogical divergence. There are four mechanisms for speciation. The most common in animals is
allopatric speciation, which occurs in populations initially isolated geographically, such as by habitat
fragmentation or migration. Selection under these conditions can produce very rapid changes in the
appearance and behaviour of organisms.[213][214] As selection and drift act independently on populations
isolated from the rest of their species, separation may eventually produce organisms that cannot interbreed.
[215]
The second mechanism of speciation is peripatric speciation, which occurs when small populations
of organisms become isolated in a new environment. This differs from allopatric speciation in that the isolated
populations are numerically much smaller than the parental population. Here, the founder effect causes rapid
speciation through both rapid genetic drift and selection on a small gene pool.[216]
The third mechanism of speciation is parapatric speciation. This is similar to peripatric speciation in
that a small population enters a new habitat, but differs in that there is no physical separation between these
two populations. Instead, speciation results from the evolution of mechanisms that reduce gene flow between
the two populations.[202] Generally this occurs when there has been a drastic change in the environment
within the parental species' habitat. One example is the grass Anthoxanthum odoratum, which can undergo
parapatric speciation in response to localised metal pollution from mines.[217] Here, plants evolve that have
resistance to high levels of metals in the soil. Selection against interbreeding with the metal-sensitive
parental population produced a gradual change in the flowering time of the metal-resistant plants, which
eventually produced complete reproductive isolation. Selection against hybrids between the two populations
may cause reinforcement, which is the evolution of traits that promote mating within a species, as well as
character displacement, which is when two species become more distinct in appearance.[218]
Geographical isolation of finches on the Galápagos Islands produced over a dozen new species.
Finally, in sympatric speciation species diverge without geographic isolation or changes in habitat.
This form is rare since even a small amount of gene flow may remove genetic differences between parts of a
population.[219] Generally, sympatric speciation in animals requires the evolution of both genetic differences
and non-random mating, to allow reproductive isolation to evolve.[220]
One type of sympatric speciation involves cross-breeding of two related species to produce a new
hybrid species. This is not common in animals as animal hybrids are usually sterile. This is because during
meiosis the homologous chromosomes from each parent are from different species and cannot successfully
pair. However, it is more common in plants because plants often double their number of chromosomes, to
form polyploids.[221] This allows the chromosomes from each parental species to form matching pairs during
meiosis, since each parent's chromosomes are represented by a pair already.[222] An example of such a
speciation event is when the plant species Arabidopsis thaliana and Arabidopsis arenosa cross-bred to give
the new species Arabidopsis suecica.[223] This happened about 20,000 years ago,[224] and the speciation
process has been repeated in the laboratory, which allows the study of the genetic mechanisms involved in
this process.[225] Indeed, chromosome doubling within a species may be a common cause of reproductive
isolation, as half the doubled chromosomes will be unmatched when breeding with undoubled organisms.[99]
Speciation events are important in the theory of punctuated equilibrium, which accounts for the
pattern in the fossil record of short "bursts" of evolution interspersed with relatively long periods of stasis,
where species remain relatively unchanged.[226] In this theory, speciation and rapid evolution are linked,
with natural selection and genetic drift acting most strongly on organisms undergoing speciation in novel
habitats or small populations. As a result, the periods of stasis in the fossil record correspond to the parental
population, and the organisms undergoing speciation and rapid evolution are found in small populations or
geographically restricted habitats, and therefore rarely being preserved as fossils.[227]
Extinction
Further information: Extinction
Tyrannosaurus rex. Non-avian dinosaurs died out in the Cretaceous–Tertiary extinction event at the
end of the Cretaceous period.
Extinction is the disappearance of an entire species. Extinction is not an unusual event, as species
regularly appear through speciation, and disappear through extinction.[228] Nearly all animal and plant
species that have lived on Earth are now extinct,[229] and extinction appears to be the ultimate fate of all
species.[230] These extinctions have happened continuously throughout the history of life, although the rate
of extinction spikes in occasional mass extinction events.[231] The Cretaceous–Tertiary extinction event,
during which the non-avian dinosaurs went extinct, is the most well-known, but the earlier Permian–Triassic
extinction event was even more severe, with approximately 96% of species driven to extinction.[231] The
Holocene extinction event is an ongoing mass extinction associated with humanity's expansion across the
globe over the past few thousand years. Present-day extinction rates are 100–1000 times greater than the
background rate, and up to 30% of species may be extinct by the mid 21st century.[232] Human activities are
now the primary cause of the ongoing extinction event;[233] global warming may further accelerate it in the
future.[234]
The role of extinction in evolution is not very well understood and may depend on which type of
extinction is considered.[231] The causes of the continuous "low-level" extinction events, which form the
majority of extinctions, may be the result of competition between species for limited resources (competitive
exclusion).[13] If one species can out-compete another, this could produce species selection, with the fitter
species surviving and the other species being driven to extinction.[123] The intermittent mass extinctions are
also important, but instead of acting as a selective force, they drastically reduce diversity in a nonspecific
manner and promote bursts of rapid evolution and speciation in survivors.[235]

Main article: Evolutionary history of life
See also: Timeline of evolution and Timeline of human evolution
Origin of life
Further information: Abiogenesis and RNA world hypothesis
The origin of life is a necessary precursor for biological evolution, but understanding that evolution
occurred once organisms appeared and investigating how this happens does not depend on understanding
exactly how life began.[236] The current scientific consensus is that the complex biochemistry that makes up
life came from simpler chemical reactions, but it is unclear how this occurred.[237] Not much is certain about
the earliest developments in life, the structure of the first living things, or the identity and nature of any last
universal common ancestor or ancestral gene pool.[238][239] Consequently, there is no scientific consensus
on how life began, but proposals include self-replicating molecules such as RNA,[240] and the assembly of
simple cells.[241]
Common descent
Further information: Common descent, Evidence of common descent, and Homology (biology)
The hominoids are descendants of a common ancestor.

All organisms on Earth are descended from a common ancestor or ancestral gene pool.[169][242]
Current species are a stage in the process of evolution, with their diversity the product of a long series of
speciation and extinction events.[243] The common descent of organisms was first deduced from four simple
facts about organisms: First, they have geographic distributions that cannot be explained by local adaptation.
Second, the diversity of life is not a set of completely unique organisms, but organisms that share
morphological similarities. Third, vestigial traits with no clear purpose resemble functional ancestral traits,
and finally, that organisms can be classified using these similarities into a hierarchy of nested groups – similar
to a family tree.[244] However, modern research has suggested that, due to horizontal gene transfer, this
"tree of life" may be more complicated than a simple branching tree since some genes have spread
independently between distantly related species.[245][246]
Past species have also left records of their evolutionary history. Fossils, along with the comparative
anatomy of present-day organisms, constitute the morphological, or anatomical, record.[247] By comparing
the anatomies of both modern and extinct species, paleontologists can infer the lineages of those species.
However, this approach is most successful for organisms that had hard body parts, such as shells, bones or
teeth. Further, as prokaryotes such as bacteria and archaea share a limited set of common morphologies,
their fossils do not provide information on their ancestry.
More recently, evidence for common descent has come from the study of biochemical similarities
between organisms. For example, all living cells use the same basic set of nucleotides and amino acids.[248]
The development of molecular genetics has revealed the record of evolution left in organisms' genomes:
dating when species diverged through the molecular clock produced by mutations.[249] For example, these
DNA sequence comparisons have revealed that humans and chimpanzees share 96% of their genomes and
analyzing the few areas where they differ helps shed light on when the common ancestor of these species
existed.[250]
Evolution of life
For more details on this topic, see Timeline of evolution.
Evolutionary tree showing the divergence of modern species from their common ancestor in the
centre.[251] The three domains are coloured, with bacteria blue, archaea green, and eukaryotes red.
Prokaryotes inhabited the Earth from approximately 3–4 billion years ago.[252][253] No obvious
changes in morphology or cellular organisation occurred in these organisms over the next few billion years.
[254]
The eukaryotic cells emerged between 1.6 – 2.7 billion years ago. The next major change in cell
structure came when bacteria were engulfed by eukaryotic cells, in a cooperative association called
endosymbiosis.[108][255] The engulfed bacteria and the host cell then underwent co-evolution, with the
bacteria evolving into either mitochondria or hydrogenosomes.[256] Another engulfment of cyanobacterial-
like organisms led to the formation of chloroplasts in algae and plants.[257]
The history of life was that of the unicellular eukaryotes, prokaryotes, and archaea until about 610
million years ago when multicellular organisms began to appear in the oceans in the Ediacaran period.[252]
[258] The evolution of multicellularity occurred in multiple independent events, in organisms as diverse as
sponges, brown algae, cyanobacteria, slime moulds and myxobacteria.[259]
Soon after the emergence of these first multicellular organisms, a remarkable amount of biological
diversity appeared over approximately 10 million years, in an event called the Cambrian explosion. Here, the
majority of types of modern animals appeared in the fossil record, as well as unique lineages that
subsequently became extinct.[260] Various triggers for the Cambrian explosion have been proposed,
including the accumulation of oxygen in the atmosphere from photosynthesis.[261]
About 500 million years ago, plants and fungi colonised the land, and were soon followed by
arthropods and other animals.[262] Insects were particularly successful and even today make up the majority
of animal species.[263] Amphibians first appeared around 300 million years ago, followed by early amniotes,
then mammals around 200 million years ago and birds around 100 million years ago (both from "reptile"-like
lineages). However, despite the evolution of these large animals, smaller organisms similar to the types that
evolved early in this process continue to be highly successful and dominate the Earth, with the majority of
both biomass and species being prokaryotes.[148]
Applications
Further information: Artificial selection and Evolutionary computation
Evolutionary biology, and in particular the understanding of how organisms evolve through natural
selection, is an area of science with many practical applications.[264] A major technological application of
evolution is artificial selection, which is the intentional selection of certain traits in a population of organisms.
Humans have used artificial selection for thousands of years in the domestication of plants and animals.[265]
More recently, such selection has become a vital part of genetic engineering, with selectable markers such
as antibiotic resistance genes being used to manipulate DNA in molecular biology. It is also possible to use
repeated rounds of mutation and selection to evolve proteins with particular properties, such as modified
enzymes or new antibodies, in a process called directed evolution.[266]
Understanding the changes that have occurred during organism's evolution can reveal the genes
needed to construct parts of the body, genes which may be involved in human genetic disorders.[267] For
example, the Mexican tetra is an albino cavefish that lost its eyesight during evolution. Breeding together
different populations of this blind fish produced some offspring with functional eyes, since different mutations
had occurred in the isolated populations that had evolved in different caves.[268] This helped identify genes
required for vision and pigmentation, such as crystallins and the melanocortin 1 receptor.[269] Similarly,
comparing the genome of the Antarctic icefish, which lacks red blood cells, to close relatives such as the
Antarctic rockcod revealed genes needed to make these blood cells.[270]
As evolution can produce highly optimised processes and networks, it has many applications in
computer science. Here, simulations of evolution using evolutionary algorithms and artificial life started with
the work of Nils Aall Barricelli in the 1960s, and was extended by Alex Fraser, who published a series of
papers on simulation of artificial selection.[271] Artificial evolution became a widely recognised optimisation
method as a result of the work of Ingo Rechenberg in the 1960s and early 1970s, who used evolution
strategies to solve complex engineering problems.[272] Genetic algorithms in particular became popular
through the writing of John Holland.[273] As academic interest grew, dramatic increases in the power of
computers allowed practical applications, including the automatic evolution of computer programs.[274]
Evolutionary algorithms are now used to solve multi-dimensional problems more efficiently than software
produced by human designers, and also to optimise the design of systems.[275]
Social and cultural responses

Further information: Social effect of evolutionary theory and Objections to evolution
As evolution became widely accepted in the 1870s, caricatures of Charles Darwin with an ape or
monkey body symbolised evolution.[276]
In the 19th century, particularly after the publication of On the Origin of Species in 1859, the idea that
life had evolved was an active source of academic debate centred on the philosophical, social and religious
implications of evolution. Nowadays, the fact that organisms evolve is uncontested in the scientific literature
and the modern evolutionary synthesis is widely accepted by scientists.[13] However, evolution remains a
contentious concept for some theists.[277]
While various religions and denominations have reconciled their beliefs with evolution through
concepts such as theistic evolution, there are creationists who believe that evolution is contradicted by the
creation myths found in their respective religions and who raise various objections to evolution.[138][278]
[279] As had been demonstrated by responses to the publication of Vestiges of the Natural History of
Creation in 1844, the most controversial aspect of evolutionary biology is the implication of human evolution
that human mental and moral faculties, which had been thought purely spiritual, are not distinctly separated
from those of other animals.[12] In some countries, notably the United States, these tensions between
science and religion have fuelled the current creation-evolution controversy, a religious conflict focusing on
politics and public education.[280] While other scientific fields such as cosmology[281] and Earth
science[282] also conflict with literal interpretations of many religious texts, evolutionary biology experiences
significantly more opposition from religious literalists.
The teaching of evolution in American secondary school biology classes was uncommon in most of
the first half of the 20th century. The Scopes Trial decision of 1925 caused the subject to become very rare in
American secondary biology textbooks for a generation, but it was gradually re-introduced about a generation
later and legally protected with the 1968 Epperson v. Arkansas decision. Since then, the competing religious
belief of creationism was legally disallowed in secondary school curricula in various decisions in the 1970s
and 1980s, but it returned in the form of intelligent design, to be excluded once again in the 2005 Kitzmiller v.
Dover Area School District case.[283]
Another example associated with evolutionary theory that is now widely regarded as unwarranted is
"Social Darwinism", a derogatory term associated with the 19th century Malthusian theory developed by
Whig philosopher Herbert Spencer. It was later expanded by others into ideas about "survival of the fittest" in
commerce and human societies as a whole, and led to claims that social inequality, sexism, racism and
imperialism were justified.[284] However, these ideas contradict Darwin's own views, and contemporary
scientists and philosophers consider these ideas to be neither mandated by evolutionary theory nor
supported by data.[285][286][287]
See also
Book:Evolution

• Current research in evolutionary biology
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p. 62. ISBN 978-0415340779.
286.^ Allhoff F (2003). "Evolutionary ethics from Darwin to Moore". History and philosophy of the
life sciences 25 (1): 51–79. doi:10.1080/03919710312331272945. PMID 15293515.
287.^ Gowaty, Patricia Adair (1997). Feminism and evolutionary biology: boundaries,
intersections, and frontiers. London: Chapman & Hall. ISBN 0-412-07361-7.
Further reading
Introductory reading
• Carroll, S. (2005). Endless Forms Most Beautiful. New York: W.W. Norton. ISBN 0-393-
06016-0.
• Charlesworth, C.B. and Charlesworth, D. (2003). Evolution. Oxfordshire: Oxford University
Press. ISBN 0-192-80251-8.
• Dawkins, R. (2006). The Selfish Gene: 30th Anniversary Edition . Oxford University Press.
ISBN 0199291152.
• Gould, S.J. (1989). Wonderful Life: The Burgess Shale and the Nature of History . New York:
W.W. Norton. ISBN 0-393-30700-X.
• Jones, S. (2001). Almost Like a Whale: The Origin of Species Updated. (American title:
Darwin's Ghost). New York: Ballantine Books. ISBN 0-345-42277-5.
• Mader, Sylvia S. (2007). Biology. Murray P. Pendarvis (9th ed.). McGraw Hill.
ISBN 9780073258393.
• Maynard Smith, J. (1993). The Theory of Evolution: Canto Edition . Cambridge University
Press. ISBN 0-521-45128-0.
• Pallen, M.J. (2009). The Rough Guide to Evolution . Rough Guides. ISBN 978-1-85828-946-
5.
• Smith, C.B. and Sullivan, C. (2007). The Top 10 Myths about Evolution. Prometheus Books.
ISBN 978-1-59102-479-8.
• Darwin, Charles (1859). On the Origin of Species (1st ed.). London: John Murray.
ISBN 0801413192. http://darwin-online.org.uk/content/frameset?
• Larson, E.J. (2004). Evolution: The Remarkable History of a Scientific Theory . New York:
Modern Library. ISBN 0-679-64288-9.
• Zimmer, C. (2001). Evolution: The Triumph of an Idea . London: HarperCollins. ISBN 0-060-
19906-7.
Advanced reading
• Barton, N.H., Briggs, D.E.G., Eisen, J.A., Goldstein, D.B. and Patel, N.H. (2007). Evolution.
Cold Spring Harbor Laboratory Press. ISBN 0-879-69684-2.
• Coyne, J.A. and Orr, H.A. (2004). Speciation. Sunderland: Sinauer Associates. ISBN 0-878-
93089-2.
• Futuyma, D.J. (2005). Evolution. Sunderland: Sinauer Associates. ISBN 0-878-93187-2.
• Gould, S.J. (2002). The Structure of Evolutionary Theory. Cambridge: Belknap Press
(Harvard University Press). ISBN 0-674-00613-5.
• Maynard Smith, J. and Szathmáry, E. (1997). The Major Transitions in Evolution.
Oxfordshire: Oxford University Press. ISBN 0-198-50294-X.
• Mayr, E. (2001). What Evolution Is. New York: Basic Books. ISBN 0-465-04426-3.
• Olson, Wendy; Hall, Brian Keith (2003). Keywords and concepts in evolutionary
developmental biology. Cambridge: Harvard University Press. ISBN 0-674-02240-8.
External links
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General information
• Evolution on In Our Time at the BBC. (listen now)
• Everything you wanted to know about evolution by New Scientist
• Synthetic Theory Of Evolution: An Introduction to Modern Evolutionary Concepts and
Theories
• Understanding Evolution from University of California, Berkeley
• Evolution of Evolution – 150 Years of Darwin's "On the Origin of Species"
• The Complete Work of Charles Darwin Online
• Understanding Evolution: History, Theory, Evidence, and Implications
On-line lectures
• What Genomes Can Tell Us About the Past – lecture by Sydney Brenner
• The Origin of Vertebrates – lecture by Marc Kirschner
• The Making of the Fittest – lecture by Sean B. Carroll
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Categories: Articles with separate introductions | Evolution | Biology theories | Evolutionary biology
W000
DNA

For a non-technical introduction to the topic, see Introduction to genetics.
For other uses, see DNA (disambiguation).
The structure of part of a DNA double helix
Deoxyribonucleic acid (/diˌɒksiˌraɪbɵ.njuːˌkleɪ.ɨk ˈæsɪd/ ( listen)), or DNA, is a nucleic acid that
contains the genetic instructions used in the development and functioning of all known living organisms (with
the exception of RNA viruses). The main role of DNA molecules is the long-term storage of information. DNA
is often compared to a set of blueprints, like a recipe or a code, since it contains the instructions needed to
construct other components of cells, such as proteins and RNA molecules. The DNA segments that carry this
genetic information are called genes, but other DNA sequences have structural purposes, or are involved in
regulating the use of this genetic information.
DNA consists of two long polymers of simple units called nucleotides, with backbones made of
sugars and phosphate groups joined by ester bonds. These two strands run in opposite directions to each
other and are therefore anti-parallel. Attached to each sugar is one of four types of molecules called bases. It
is the sequence of these four bases along the backbone that encodes information. This information is read
using the genetic code, which specifies the sequence of the amino acids within proteins. The code is read by
copying stretches of DNA into the related nucleic acid RNA, in a process called transcription.
Within cells, DNA is organized into long structures called chromosomes. These chromosomes are
duplicated before cells divide, in a process called DNA replication. Eukaryotic organisms (animals, plants,
fungi, and protists) store most of their DNA inside the cell nucleus and some of their DNA in organelles, such
as mitochondria or chloroplasts.[1] In contrast, prokaryotes (bacteria and archaea) store their DNA only in the
cytoplasm. Within the chromosomes, chromatin proteins such as histones compact and organize DNA.
These compact structures guide the interactions between DNA and other proteins, helping control which
parts of the DNA are transcribed.
Contents
[hide]
• 1 Properties
• 1.1 Grooves
• 1.2 Base pairing
• 1.3 Sense and antisense
• 1.4 Supercoiling
• 1.5 Alternate DNA structures
• 1.6 Alternate DNA chemistry
• 1.7 Quadruplex structures
• 1.8 Branched DNA
• 1.9 Vibration
• 2 Chemical modifications
• 2.1 Base modifications
• 2.2 Damage
• 3 Biological functions
• 3.1 Genes and genomes
• 3.2 Transcription and translation
• 3.3 Replication
• 4 Interactions with proteins
• 4.1 DNA-binding proteins
• 4.2 DNA-modifying enzymes
Properties
Chemical structure of DNA. Hydrogen bonds shown as dotted lines.
DNA is a long polymer made from repeating units called nucleotides.[2][3][4] As first discovered by
James D. Watson and Francis Crick, the structure of DNA of all species comprises two helical chains each
coiled round the same axis, and each with a pitch of 34 Ångströms (3.4 nanometres) and a radius of
10 Ångströms (1.0 nanometres).[5] According to another study, when measured in a particular solution, the
DNA chain measured 22 to 26 Ångströms wide (2.2 to 2.6 nanometres), and one nucleotide unit measured
3.3 Å (0.33 nm) long.[6] Although each individual repeating unit is very small, DNA polymers can be very
large molecules containing millions of nucleotides. For instance, the largest human chromosome,
chromosome number 1, is approximately 220 million base pairs long.[7]
In living organisms, DNA does not usually exist as a single molecule, but instead as a pair of
molecules that are held tightly together.[5][8] These two long strands entwine like vines, in the shape of a
double helix. The nucleotide repeats contain both the segment of the backbone of the molecule, which holds
the chain together, and a base, which interacts with the other DNA strand in the helix. A base linked to a
sugar is called a nucleoside and a base linked to a sugar and one or more phosphate groups is called a
nucleotide. If multiple nucleotides are linked together, as in DNA, this polymer is called a polynucleotide.[9]
The backbone of the DNA strand is made from alternating phosphate and sugar residues.[10] The
sugar in DNA is 2-deoxyribose, which is a pentose (five-carbon) sugar. The sugars are joined together by
phosphate groups that form phosphodiester bonds between the third and fifth carbon atoms of adjacent
sugar rings. These asymmetric bonds mean a strand of DNA has a direction. In a double helix the direction of
the nucleotides in one strand is opposite to their direction in the other strand: the strands are antiparallel. The
asymmetric ends of DNA strands are called the 5′ (five prime) and 3′ (three prime) ends, with the 5' end
having a terminal phosphate group and the 3' end a terminal hydroxyl group. One major difference between
DNA and RNA is the sugar, with the 2-deoxyribose in DNA being replaced by the alternative pentose sugar
ribose in RNA.[8]
A section of DNA. The bases lie horizontally between the two spiraling strands.[11] Animated version
at File:DNA orbit animated.gif.
The DNA double helix is stabilized by hydrogen bonds between the bases attached to the two
strands. The four bases found in DNA are adenine (abbreviated A), cytosine (C), guanine (G) and thymine
(T). These four bases are attached to the sugar/phosphate to form the complete nucleotide, as shown for
adenosine monophosphate.
These bases are classified into two types; adenine and guanine are fused five- and six-membered
heterocyclic compounds called purines, while cytosine and thymine are six-membered rings called
pyrimidines.[8] A fifth pyrimidine base, called uracil (U), usually takes the place of thymine in RNA and differs
from thymine by lacking a methyl group on its ring. Uracil is not usually found in DNA, occurring only as a
breakdown product of cytosine. In addition to RNA and DNA, a large number of artificial nucleic acid
analogues have also been created to study the proprieties of nucleic acids, or for use in biotechnology.[12]
Major and minor grooves of DNA. Minor groove is a binding site for the dye Hoechst 33258.
Grooves
Twin helical strands form the DNA backbone. Another double helix may be found by tracing the
spaces, or grooves, between the strands. These voids are adjacent to the base pairs and may provide a
binding site. As the strands are not directly opposite each other, the grooves are unequally sized. One
groove, the major groove, is 22 Å wide and the other, the minor groove, is 12 Å wide.[13] The narrowness of
the minor groove means that the edges of the bases are more accessible in the major groove. As a result,
proteins like transcription factors that can bind to specific sequences in double-stranded DNA usually make
contacts to the sides of the bases exposed in the major groove.[14] This situation varies in unusual
conformations of DNA within the cell (see below), but the major and minor grooves are always named to
reflect the differences in size that would be seen if the DNA is twisted back into the ordinary B form.
Base pairing
Further information: Base pair
Each type of base on one strand forms a bond with just one type of base on the other strand. This is
called complementary base pairing. Here, purines form hydrogen bonds to pyrimidines, with A bonding only
to T, and C bonding only to G. This arrangement of two nucleotides binding together across the double helix
is called a base pair. As hydrogen bonds are not covalent, they can be broken and rejoined relatively easily.
The two strands of DNA in a double helix can therefore be pulled apart like a zipper, either by a mechanical
force or high temperature.[15] As a result of this complementarity, all the information in the double-stranded
sequence of a DNA helix is duplicated on each strand, which is vital in DNA replication. Indeed, this
reversible and specific interaction between complementary base pairs is critical for all the functions of DNA in
living organisms.[3]
Top, a GC base pair with three hydrogen bonds. Bottom, an AT base pair with two hydrogen bonds.
Non-covalent hydrogen bonds between the pairs are shown as dashed lines.
The two types of base pairs form different numbers of hydrogen bonds, AT forming two hydrogen
bonds, and GC forming three hydrogen bonds (see figures, left). DNA with high GC-content is more stable
than DNA with low GC-content, but contrary to popular belief, this is not due to the extra hydrogen bond of a
GC base pair but rather the contribution of stacking interactions (hydrogen bonding merely provides
specificity of the pairing, not stability).[16]
As a result, it is both the percentage of GC base pairs and the overall length of a DNA double helix
that determine the strength of the association between the two strands of DNA. Long DNA helices with a high
GC content have stronger-interacting strands, while short helices with high AT content have weaker-
interacting strands.[17] In biology, parts of the DNA double helix that need to separate easily, such as the
TATAAT Pribnow box in some promoters, tend to have a high AT content, making the strands easier to pull
apart.[18]
In the laboratory, the strength of this interaction can be measured by finding the temperature required
to break the hydrogen bonds, their melting temperature (also called Tm value). When all the base pairs in a
DNA double helix melt, the strands separate and exist in solution as two entirely independent molecules.
These single-stranded DNA molecules (ssDNA) have no single common shape, but some conformations are
more stable than others.[19]
Sense and antisense

Further information: Sense (molecular biology)
A DNA sequence is called "sense" if its sequence is the same as that of a messenger RNA copy that
is translated into protein.[20] The sequence on the opposite strand is called the "antisense" sequence. Both
sense and antisense sequences can exist on different parts of the same strand of DNA (i.e. both strands
contain both sense and antisense sequences). In both prokaryotes and eukaryotes, antisense RNA
sequences are produced, but the functions of these RNAs are not entirely clear.[21] One proposal is that
antisense RNAs are involved in regulating gene expression through RNA-RNA base pairing.[22]
A few DNA sequences in prokaryotes and eukaryotes, and more in plasmids and viruses, blur the
distinction between sense and antisense strands by having overlapping genes.[23] In these cases, some
DNA sequences do double duty, encoding one protein when read along one strand, and a second protein
when read in the opposite direction along the other strand. In bacteria, this overlap may be involved in the
regulation of gene transcription,[24] while in viruses, overlapping genes increase the amount of information
that can be encoded within the small viral genome.[25]
Supercoiling
Further information: DNA supercoil
DNA can be twisted like a rope in a process called DNA supercoiling. With DNA in its "relaxed" state,
a strand usually circles the axis of the double helix once every 10.4 base pairs, but if the DNA is twisted the
strands become more tightly or more loosely wound.[26] If the DNA is twisted in the direction of the helix, this
is positive supercoiling, and the bases are held more tightly together. If they are twisted in the opposite
direction, this is negative supercoiling, and the bases come apart more easily. In nature, most DNA has slight
negative supercoiling that is introduced by enzymes called topoisomerases.[27] These enzymes are also
needed to relieve the twisting stresses introduced into DNA strands during processes such as transcription
and DNA replication.[28]
From left to right, the structures of A, B and Z DNA
Alternate DNA structures

Further information: A Structure for DNA, Molecular models of DNA, and DNA structure
DNA exists in many possible conformations that include A-DNA, B-DNA, and Z-DNA forms, although,
only B-DNA and Z-DNA have been directly observed in functional organisms.[10] The conformation that DNA
adopts depends on the hydration level, DNA sequence, the amount and direction of supercoiling, chemical
modifications of the bases, the type and concentration of metal ions, as well as the presence of polyamines
in solution.[29]
The first published reports of A-DNA X-ray diffraction patterns— and also B-DNA used analyses
based on Patterson transforms that provided only a limited amount of structural information for oriented fibers
of DNA.[30][31] An alternate analysis was then proposed by Wilkins et al., in 1953, for the in vivo B-DNA X-
ray diffraction/scattering patterns of highly hydrated DNA fibers in terms of squares of Bessel functions.[32] In
the same journal, James D. Watson and Francis Crick presented their molecular modeling analysis of the
DNA X-ray diffraction patterns to suggest that the structure was a double-helix.[5]
Although the `B-DNA form' is most common under the conditions found in cells,[33] it is not a well-
defined conformation but a family of related DNA conformations[34] that occur at the high hydration levels
present in living cells. Their corresponding X-ray diffraction and scattering patterns are characteristic of
molecular paracrystals with a significant degree of disorder.[35][36]
Compared to B-DNA, the A-DNA form is a wider right-handed spiral, with a shallow, wide minor
groove and a narrower, deeper major groove. The A form occurs under non-physiological conditions in
partially dehydrated samples of DNA, while in the cell it may be produced in hybrid pairings of DNA and RNA
strands, as well as in enzyme-DNA complexes.[37][38] Segments of DNA where the bases have been
chemically modified by methylation may undergo a larger change in conformation and adopt the Z form.
Here, the strands turn about the helical axis in a left-handed spiral, the opposite of the more common B form.
[39] These unusual structures can be recognized by specific Z-DNA binding proteins and may be involved in
the regulation of transcription.[40]
Alternate DNA chemistry

For a number of years exobiologists have proposed the existence of a shadow biosphere, a
postulated microbial biosphere of Earth that uses radically different biochemical and molecular processes
than currently known life. One of the proposals was the existence of lifeforms that use arsenic instead of
phosphorus in DNA.
A December 2010 NASA press conference revealed that the bacterium GFAJ-1, which has evolved
in an arsenic-rich environment, is the first terrestrial lifeform found which may have this ability. The bacterium
was found in Mono Lake, east of Yosemite National Park. GFAJ-1 is a rod-shaped extremophile bacterium in
the family Halomonadaceae that, when starved of phosphorus, may be capable of incorporating the usually
poisonous element arsenic in its DNA.[41] This discovery lends weight to the long-standing idea that
extraterrestrial life could have a different chemical makeup from life on Earth.[41][42] The research was
carried out by a team led by Felisa Wolfe-Simon, a geomicrobiologist and geobiochemist, a Postdoctoral
Fellow of the NASA Astrobiology Institute with Arizona State University.
Quadruplex structures
Further information: G-quadruplex
At the ends of the linear chromosomes are specialized regions of DNA called telomeres. The main
function of these regions is to allow the cell to replicate chromosome ends using the enzyme telomerase, as
the enzymes that normally replicate DNA cannot copy the extreme 3′ ends of chromosomes.[43] These
specialized chromosome caps also help protect the DNA ends, and stop the DNA repair systems in the cell
from treating them as damage to be corrected.[44] In human cells, telomeres are usually lengths of single-
stranded DNA containing several thousand repeats of a simple TTAGGG sequence.[45]
DNA quadruplex formed by telomere repeats. The looped conformation of the DNA backbone is very
different from the typical DNA helix.[46]
These guanine-rich sequences may stabilize chromosome ends by forming structures of stacked
sets of four-base units, rather than the usual base pairs found in other DNA molecules. Here, four guanine
bases form a flat plate and these flat four-base units then stack on top of each other, to form a stable G-
quadruplex structure.[47] These structures are stabilized by hydrogen bonding between the edges of the
bases and chelation of a metal ion in the centre of each four-base unit.[48] Other structures can also be
formed, with the central set of four bases coming from either a single strand folded around the bases, or
several different parallel strands, each contributing one base to the central structure.
In addition to these stacked structures, telomeres also form large loop structures called telomere
loops, or T-loops. Here, the single-stranded DNA curls around in a long circle stabilized by telomere-binding
proteins.[49] At the very end of the T-loop, the single-stranded telomere DNA is held onto a region of double-
stranded DNA by the telomere strand disrupting the double-helical DNA and base pairing to one of the two
strands. This triple-stranded structure is called a displacement loop or D-loop.[47]
Single Multiple
branch branches
Branched DNA can form networks containing multiple branches.
Branched DNA
Further information: Branched DNA and DNA nanotechnology
In DNA fraying occurs when non-complementary regions exist at the end of an otherwise
complementary double-strand of DNA. However, branched DNA can occur if a third strand of DNA is
introduced and contains adjoining regions able to hybridize with the frayed regions of the pre-existing double-
strand. Although the simplest example of branched DNA involves only three strands of DNA, complexes
involving additional strands and multiple branches are also possible.[50] Branched DNA can be used in
nanotechnology to construct geometric shapes, see the section on uses in technology below.
Vibration
DNA may carry out low-frequency collective motion as observed by the Raman spectroscopy[51][52]
and analyzed with a quasi-continuum model.[53][54]
Chemical modifications
5-
cytosine thymine
methylcytosine
Structure of cytosine with and without the 5-methyl group. Deamination converts 5-methylcytosine
into thymine.
Base modifications
Further information: DNA methylation
The expression of genes is influenced by how the DNA is packaged in chromosomes, in a structure
called chromatin. Base modifications can be involved in packaging, with regions that have low or no gene
expression usually containing high levels of methylation of cytosine bases. For example, cytosine
methylation, produces 5-methylcytosine, which is important for X-chromosome inactivation.[55] The average
level of methylation varies between organisms - the worm Caenorhabditis elegans lacks cytosine
methylation, while vertebrates have higher levels, with up to 1% of their DNA containing 5-methylcytosine.
[56] Despite the importance of 5-methylcytosine, it can deaminate to leave a thymine base, so methylated
cytosines are particularly prone to mutations.[57] Other base modifications include adenine methylation in
bacteria, the presence of 5-hydroxymethylcytosine in the brain,[58] and the glycosylation of uracil to produce
the "J-base" in kinetoplastids.[59][60]
Damage
Further information: Mutation
A covalent adduct between a metabolically activated form of benzo[a]pyrene, the major mutagen in
tobacco smoke, and DNA[61]
DNA can be damaged by many sorts of mutagens, which change the DNA sequence. Mutagens
include oxidizing agents, alkylating agents and also high-energy electromagnetic radiation such as ultraviolet
light and X-rays. The type of DNA damage produced depends on the type of mutagen. For example, UV light
can damage DNA by producing thymine dimers, which are cross-links between pyrimidine bases.[62] On the
other hand, oxidants such as free radicals or hydrogen peroxide produce multiple forms of damage, including
base modifications, particularly of guanosine, and double-strand breaks.[63] A typical human cell contains
about 150,000 bases that have suffered oxidative damage.[64] Of these oxidative lesions, the most
dangerous are double-strand breaks, as these are difficult to repair and can produce point mutations,
insertions and deletions from the DNA sequence, as well as chromosomal translocations.[65]
Many mutagens fit into the space between two adjacent base pairs, this is called intercalation. Most
intercalators are aromatic and planar molecules; examples include ethidium bromide, daunomycin, and
doxorubicin. In order for an intercalator to fit between base pairs, the bases must separate, distorting the
DNA strands by unwinding of the double helix. This inhibits both transcription and DNA replication, causing
toxicity and mutations. As a result, DNA intercalators are often carcinogens, and benzo[a]pyrene diol
epoxide, acridines, aflatoxin and ethidium bromide are well-known examples.[66][67][68] Nevertheless, due
to their ability to inhibit DNA transcription and replication, other similar toxins are also used in chemotherapy
to inhibit rapidly growing cancer cells.[69]
Biological functions
DNA usually occurs as linear chromosomes in eukaryotes, and circular chromosomes in prokaryotes.
The set of chromosomes in a cell makes up its genome; the human genome has approximately 3 billion base
pairs of DNA arranged into 46 chromosomes.[70] The information carried by DNA is held in the sequence of
pieces of DNA called genes. Transmission of genetic information in genes is achieved via complementary
base pairing. For example, in transcription, when a cell uses the information in a gene, the DNA sequence is
copied into a complementary RNA sequence through the attraction between the DNA and the correct RNA
nucleotides. Usually, this RNA copy is then used to make a matching protein sequence in a process called
translation, which depends on the same interaction between RNA nucleotides. In alternative fashion, a cell
may simply copy its genetic information in a process called DNA replication. The details of these functions
are covered in other articles; here we focus on the interactions between DNA and other molecules that
mediate the function of the genome.
Genes and genomes

Further information: Cell nucleus, Chromatin, Chromosome, Gene, Noncoding DNA
Genomic DNA is tightly and orderly packed in the process called DNA condensation to fit the small
available volumes of the cell. In eukaryotes, DNA is located in the cell nucleus, as well as small amounts in
mitochondria and chloroplasts. In prokaryotes, the DNA is held within an irregularly shaped body in the
cytoplasm called the nucleoid.[71] The genetic information in a genome is held within genes, and the
complete set of this information in an organism is called its genotype. A gene is a unit of heredity and is a
region of DNA that influences a particular characteristic in an organism. Genes contain an open reading
frame that can be transcribed, as well as regulatory sequences such as promoters and enhancers, which
control the transcription of the open reading frame.
In many species, only a small fraction of the total sequence of the genome encodes protein. For
example, only about 1.5% of the human genome consists of protein-coding exons, with over 50% of human
DNA consisting of non-coding repetitive sequences.[72] The reasons for the presence of so much non-coding
DNA in eukaryotic genomes and the extraordinary differences in genome size, or C-value, among species
represent a long-standing puzzle known as the "C-value enigma".[73] However, DNA sequences that do not
code protein may still encode functional non-coding RNA molecules, which are involved in the regulation of
gene expression.[74]
T7 RNA polymerase (blue) producing a mRNA (green) from a DNA template (orange).[75]
Some non-coding DNA sequences play structural roles in chromosomes. Telomeres and
centromeres typically contain few genes, but are important for the function and stability of chromosomes.[44]
[76] An abundant form of non-coding DNA in humans are pseudogenes, which are copies of genes that have
been disabled by mutation.[77] These sequences are usually just molecular fossils, although they can
occasionally serve as raw genetic material for the creation of new genes through the process of gene
duplication and divergence.[78]
Transcription and translation

Further information: Genetic code, Transcription (genetics), Protein biosynthesis
A gene is a sequence of DNA that contains genetic information and can influence the phenotype of
an organism. Within a gene, the sequence of bases along a DNA strand defines a messenger RNA
sequence, which then defines one or more protein sequences. The relationship between the nucleotide
sequences of genes and the amino-acid sequences of proteins is determined by the rules of translation,
known collectively as the genetic code. The genetic code consists of three-letter 'words' called codons
formed from a sequence of three nucleotides (e.g. ACT, CAG, TTT).
In transcription, the codons of a gene are copied into messenger RNA by RNA polymerase. This
RNA copy is then decoded by a ribosome that reads the RNA sequence by base-pairing the messenger RNA
to transfer RNA, which carries amino acids. Since there are 4 bases in 3-letter combinations, there are 64
possible codons (43 combinations). These encode the twenty standard amino acids, giving most amino acids
more than one possible codon. There are also three 'stop' or 'nonsense' codons signifying the end of the
coding region; these are the TAA, TGA and TAG codons.
DNA replication. The double helix is unwound by a helicase and topoisomerase. Next, one DNA
polymerase produces the leading strand copy. Another DNA polymerase binds to the lagging strand. This
enzyme makes discontinuous segments (called Okazaki fragments) before DNA ligase joins them together.
Replication
Further information: DNA replication
Cell division is essential for an organism to grow, but, when a cell divides, it must replicate the DNA
in its genome so that the two daughter cells have the same genetic information as their parent. The double-
stranded structure of DNA provides a simple mechanism for DNA replication. Here, the two strands are
separated and then each strand's complementary DNA sequence is recreated by an enzyme called DNA
polymerase. This enzyme makes the complementary strand by finding the correct base through
complementary base pairing, and bonding it onto the original strand. As DNA polymerases can only extend a
DNA strand in a 5′ to 3′ direction, different mechanisms are used to copy the antiparallel strands of the
double helix.[79] In this way, the base on the old strand dictates which base appears on the new strand, and
the cell ends up with a perfect copy of its DNA.
Interactions with proteins

All the functions of DNA depend on interactions with proteins. These protein interactions can be non-
specific, or the protein can bind specifically to a single DNA sequence. Enzymes can also bind to DNA and of
these, the polymerases that copy the DNA base sequence in transcription and DNA replication are
particularly important.
DNA-binding proteins
Further information: DNA-binding protein
Interaction of DNA with histones (shown in white, top). These proteins' basic amino acids (below left,
blue) bind to the acidic phosphate groups on DNA (below right, red).
Structural proteins that bind DNA are well-understood examples of non-specific DNA-protein
interactions. Within chromosomes, DNA is held in complexes with structural proteins. These proteins
organize the DNA into a compact structure called chromatin. In eukaryotes this structure involves DNA
binding to a complex of small basic proteins called histones, while in prokaryotes multiple types of proteins
are involved.[80][81] The histones form a disk-shaped complex called a nucleosome, which contains two
complete turns of double-stranded DNA wrapped around its surface. These non-specific interactions are
formed through basic residues in the histones making ionic bonds to the acidic sugar-phosphate backbone of
the DNA, and are therefore largely independent of the base sequence.[82] Chemical modifications of these
basic amino acid residues include methylation, phosphorylation and acetylation.[83] These chemical
changes alter the strength of the interaction between the DNA and the histones, making the DNA more or
less accessible to transcription factors and changing the rate of transcription.[84] Other non-specific DNA-
binding proteins in chromatin include the high-mobility group proteins, which bind to bent or distorted DNA.
[85] These proteins are important in bending arrays of nucleosomes and arranging them into the larger
structures that make up chromosomes.[86]
A distinct group of DNA-binding proteins are the DNA-binding proteins that specifically bind single-
stranded DNA. In humans, replication protein A is the best-understood member of this family and is used in
processes where the double helix is separated, including DNA replication, recombination and DNA repair.[87]
These binding proteins seem to stabilize single-stranded DNA and protect it from forming stem-loops or
being degraded by nucleases.
The lambda repressor helix-turn-helix transcription factor bound to its DNA target[88]
In contrast, other proteins have evolved to bind to particular DNA sequences. The most intensively
studied of these are the various transcription factors, which are proteins that regulate transcription. Each
transcription factor binds to one particular set of DNA sequences and activates or inhibits the transcription of
genes that have these sequences close to their promoters. The transcription factors do this in two ways.
Firstly, they can bind the RNA polymerase responsible for transcription, either directly or through other
mediator proteins; this locates the polymerase at the promoter and allows it to begin transcription.[89]
Alternatively, transcription factors can bind enzymes that modify the histones at the promoter; this will change
the accessibility of the DNA template to the polymerase.[90]
As these DNA targets can occur throughout an organism's genome, changes in the activity of one
type of transcription factor can affect thousands of genes.[91] Consequently, these proteins are often the
targets of the signal transduction processes that control responses to environmental changes or cellular
differentiation and development. The specificity of these transcription factors' interactions with DNA come
from the proteins making multiple contacts to the edges of the DNA bases, allowing them to "read" the DNA
sequence. Most of these base-interactions are made in the major groove, where the bases are most
accessible.[92]
The restriction enzyme EcoRV (green) in a complex with its substrate DNA[93]
DNA-modifying enzymes
Nucleases and ligases

Nucleases are enzymes that cut DNA strands by catalyzing the hydrolysis of the phosphodiester
bonds. Nucleases that hydrolyse nucleotides from the ends of DNA strands are called exonucleases, while
endonucleases cut within strands. The most frequently used nucleases in molecular biology are the
restriction endonucleases, which cut DNA at specific sequences. For instance, the EcoRV enzyme shown to
the left recognizes the 6-base sequence 5′-GAT|ATC-3′ and makes a cut at the vertical line. In nature, these
enzymes protect bacteria against phage infection by digesting the phage DNA when it enters the bacterial
cell, acting as part of the restriction modification system.[94] In technology, these sequence-specific
nucleases are used in molecular cloning and DNA fingerprinting.
Enzymes called DNA ligases can rejoin cut or broken DNA strands.[95] Ligases are particularly
important in lagging strand DNA replication, as they join together the short segments of DNA produced at the
replication fork into a complete copy of the DNA template. They are also used in DNA repair and genetic
recombination.[95]
Topoisomerases and helicases

Topoisomerases are enzymes with both nuclease and ligase activity. These proteins change the
amount of supercoiling in DNA. Some of these enzymes work by cutting the DNA helix and allowing one
section to rotate, thereby reducing its level of supercoiling; the enzyme then seals the DNA break.[27] Other
types of these enzymes are capable of cutting one DNA helix and then passing a second strand of DNA
through this break, before rejoining the helix.[96] Topoisomerases are required for many processes involving
DNA, such as DNA replication and transcription.[28]
Helicases are proteins that are a type of molecular motor. They use the chemical energy in
nucleoside triphosphates, predominantly ATP, to break hydrogen bonds between bases and unwind the DNA
double helix into single strands.[97] These enzymes are essential for most processes where enzymes need
to access the DNA bases.
Polymerases
Polymerases are enzymes that synthesize polynucleotide chains from nucleoside triphosphates. The
sequence of their products are copies of existing polynucleotide chains - which are called templates. These
enzymes function by adding nucleotides onto the 3′ hydroxyl group of the previous nucleotide in a DNA
strand. As a consequence, all polymerases work in a 5′ to 3′ direction.[98] In the active site of these
enzymes, the incoming nucleoside triphosphate base-pairs to the template: this allows polymerases to
accurately synthesize the complementary strand of their template. Polymerases are classified according to
the type of template that they use.
In DNA replication, a DNA-dependent DNA polymerase makes a copy of a DNA sequence. Accuracy
is vital in this process, so many of these polymerases have a proofreading activity. Here, the polymerase
recognizes the occasional mistakes in the synthesis reaction by the lack of base pairing between the
mismatched nucleotides. If a mismatch is detected, a 3′ to 5′ exonuclease activity is activated and the
incorrect base removed.[99] In most organisms, DNA polymerases function in a large complex called the
replisome that contains multiple accessory subunits, such as the DNA clamp or helicases.[100]
RNA-dependent DNA polymerases are a specialized class of polymerases that copy the sequence of
an RNA strand into DNA. They include reverse transcriptase, which is a viral enzyme involved in the infection
of cells by retroviruses, and telomerase, which is required for the replication of telomeres.[43][101]
Telomerase is an unusual polymerase because it contains its own RNA template as part of its structure.[44]
Transcription is carried out by a DNA-dependent RNA polymerase that copies the sequence of a
DNA strand into RNA. To begin transcribing a gene, the RNA polymerase binds to a sequence of DNA called
a promoter and separates the DNA strands. It then copies the gene sequence into a messenger RNA
transcript until it reaches a region of DNA called the terminator, where it halts and detaches from the DNA. As
with human DNA-dependent DNA polymerases, RNA polymerase II, the enzyme that transcribes most of the
genes in the human genome, operates as part of a large protein complex with multiple regulatory and
accessory subunits.[102]
Genetic recombination
Structure of the Holliday junction intermediate in genetic recombination. The four separate DNA
strands are coloured red, blue, green and yellow.[103]
Further information: Genetic recombination
Recombination involves the breakage and rejoining of two chromosomes (M and F) to produce two
re-arranged chromosomes (C1 and C2).
A DNA helix usually does not interact with other segments of DNA, and in human cells the different
chromosomes even occupy separate areas in the nucleus called "chromosome territories".[104] This physical
separation of different chromosomes is important for the ability of DNA to function as a stable repository for
information, as one of the few times chromosomes interact is during chromosomal crossover when they
recombine. Chromosomal crossover is when two DNA helices break, swap a section and then rejoin.
Recombination allows chromosomes to exchange genetic information and produces new
combinations of genes, which increases the efficiency of natural selection and can be important in the rapid
evolution of new proteins.[105] Genetic recombination can also be involved in DNA repair, particularly in the
cell's response to double-strand breaks.[106]
The most common form of chromosomal crossover is homologous recombination, where the two
chromosomes involved share very similar sequences. Non-homologous recombination can be damaging to
cells, as it can produce chromosomal translocations and genetic abnormalities. The recombination reaction is
catalyzed by enzymes known as recombinases, such as RAD51.[107] The first step in recombination is a
double-stranded break either caused by an endonuclease or damage to the DNA.[108] A series of steps
catalyzed in part by the recombinase then leads to joining of the two helices by at least one Holliday junction,
in which a segment of a single strand in each helix is annealed to the complementary strand in the other
helix. The Holliday junction is a tetrahedral junction structure that can be moved along the pair of
chromosomes, swapping one strand for another. The recombination reaction is then halted by cleavage of
the junction and re-ligation of the released DNA.[109]
Evolution
Further information: RNA world hypothesis
DNA contains the genetic information that allows all modern living things to function, grow and
reproduce. However, it is unclear how long in the 4-billion-year history of life DNA has performed this
function, as it has been proposed that the earliest forms of life may have used RNA as their genetic material.
[98][110] RNA may have acted as the central part of early cell metabolism as it can both transmit genetic
information and carry out catalysis as part of ribozymes.[111] This ancient RNA world where nucleic acid
would have been used for both catalysis and genetics may have influenced the evolution of the current
genetic code based on four nucleotide bases. This would occur, since the number of different bases in such
an organism is a trade-off between a small number of bases increasing replication accuracy and a large
number of bases increasing the catalytic efficiency of ribozymes.[112]
However, there is no direct evidence of ancient genetic systems, as recovery of DNA from most
fossils is impossible. This is because DNA will survive in the environment for less than one million years and
slowly degrades into short fragments in solution.[113] Claims for older DNA have been made, most notably a
report of the isolation of a viable bacterium from a salt crystal 250 million years old,[114] but these claims are
controversial.[115][116]
Uses in technology
Genetic engineering
Further information: Molecular biology, nucleic acid methods and genetic engineering
Methods have been developed to purify DNA from organisms, such as phenol-chloroform extraction,
and to manipulate it in the laboratory, such as restriction digests and the polymerase chain reaction. Modern
biology and biochemistry make intensive use of these techniques in recombinant DNA technology.
Recombinant DNA is a man-made DNA sequence that has been assembled from other DNA sequences.
They can be transformed into organisms in the form of plasmids or in the appropriate format, by using a viral
vector.[117] The genetically modified organisms produced can be used to produce products such as
recombinant proteins, used in medical research,[118] or be grown in agriculture.[119][120]
Forensics
Further information: DNA profiling
Forensic scientists can use DNA in blood, semen, skin, saliva or hair found at a crime scene to
identify a matching DNA of an individual, such as a perpetrator. This process is formally termed DNA
profiling, but may also be called "genetic fingerprinting". In DNA profiling, the lengths of variable sections of
repetitive DNA, such as short tandem repeats and minisatellites, are compared between people. This method
is usually an extremely reliable technique for identifying a matching DNA.[121] However, identification can be
complicated if the scene is contaminated with DNA from several people.[122] DNA profiling was developed in
1984 by British geneticist Sir Alec Jeffreys,[123] and first used in forensic science to convict Colin Pitchfork in
the 1988 Enderby murders case.[124]
People convicted of certain types of crimes may be required to provide a sample of DNA for a
database. This has helped investigators solve old cases where only a DNA sample was obtained from the
scene. DNA profiling can also be used to identify victims of mass casualty incidents.[125] On the other hand,
many convicted people have been released from prison on the basis of DNA techniques, which were not
available when a crime had originally been committed.
Bioinformatics
Further information: Bioinformatics
Bioinformatics involves the manipulation, searching, and data mining of biological data, and this
includes DNA sequence data. The development of techniques to store and search DNA sequences have led
to widely applied advances in computer science, especially string searching algorithms, machine learning
and database theory.[126] String searching or matching algorithms, which find an occurrence of a sequence
of letters inside a larger sequence of letters, were developed to search for specific sequences of nucleotides.
[127] The DNA sequenced may be aligned with other DNA sequences to identify homologous sequences and
locate the specific mutations that make them distinct. These techniques, especially multiple sequence
alignment, are used in studying phylogenetic relationships and protein function.[128] Data sets representing
entire genomes' worth of DNA sequences, such as those produced by the Human Genome Project, are
difficult to use without the annotations that identify the locations of genes and regulatory elements on each
chromosome. Regions of DNA sequence that have the characteristic patterns associated with protein- or
RNA-coding genes can be identified by gene finding algorithms, which allow researchers to predict the
presence of particular gene products and their possible functions in an organism even before they have been
isolated experimentally.[129] Entire genomes may also be compared which can shed light on the
evolutionary history of particular organism and permit the examination of complex evolutionary events.
DNA nanotechnology
The DNA structure at left (schematic shown) will self-assemble into the structure visualized by atomic
force microscopy at right. DNA nanotechnology is the field that seeks to design nanoscale structures using
the molecular recognition properties of DNA molecules. Image from Strong, 2004.
Further information: DNA nanotechnology
DNA nanotechnology uses the unique molecular recognition properties of DNA and other nucleic
acids to create self-assembling branched DNA complexes with useful properties.[130] DNA is thus used as a
structural material rather than as a carrier of biological information. This has led to the creation of two-
dimensional periodic lattices (both tile-based as well as using the "DNA origami" method) as well as three-
dimensional structures in the shapes of polyhedra.[131] Nanomechanical devices and algorithmic self-
assembly have also been demonstrated,[132] and these DNA structures have been used to template the
arrangement of other molecules such as gold nanoparticles and streptavidin proteins.[133]
History and anthropology

Further information: Phylogenetics and Genetic genealogy
Because DNA collects mutations over time, which are then inherited, it contains historical
information, and, by comparing DNA sequences, geneticists can infer the evolutionary history of organisms,
their phylogeny.[134] This field of phylogenetics is a powerful tool in evolutionary biology. If DNA sequences
within a species are compared, population geneticists can learn the history of particular populations. This can
be used in studies ranging from ecological genetics to anthropology; For example, DNA evidence is being
used to try to identify the Ten Lost Tribes of Israel.[135][136]
DNA has also been used to look at modern family relationships, such as establishing family
relationships between the descendants of Sally Hemings and Thomas Jefferson. This usage is closely
related to the use of DNA in criminal investigations detailed above. Indeed, some criminal investigations have
been solved when DNA from crime scenes has matched relatives of the guilty individual.[137]
History of DNA research

James D. Watson and Francis Crick (right), co-originators of the double-helix model, with Maclyn
McCarty (left).
Rosalind Franklin, co-creator of the single X-ray diffraction image

Raymond Gosling, co-creator of the single X-ray diffraction image
Further information: History of molecular biology
DNA was first isolated by the Swiss physician Friedrich Miescher who, in 1869, discovered a
microscopic substance in the pus of discarded surgical bandages. As it resided in the nuclei of cells, he
called it "nuclein".[138] In 1919, Phoebus Levene identified the base, sugar and phosphate nucleotide unit.
[139] Levene suggested that DNA consisted of a string of nucleotide units linked together through the
phosphate groups. However, Levene thought the chain was short and the bases repeated in a fixed order. In
1937 William Astbury produced the first X-ray diffraction patterns that showed that DNA had a regular
structure.[140]
In 1928, Frederick Griffith discovered that traits of the "smooth" form of the Pneumococcus could be
transferred to the "rough" form of the same bacteria by mixing killed "smooth" bacteria with the live "rough"
form.[141] This system provided the first clear suggestion that DNA carries genetic information—the Avery–
MacLeod–McCarty experiment—when Oswald Avery, along with coworkers Colin MacLeod and Maclyn
McCarty, identified DNA as the transforming principle in 1943.[142] DNA's role in heredity was confirmed in
1952, when Alfred Hershey and Martha Chase in the Hershey–Chase experiment showed that DNA is the
genetic material of the T2 phage.[143]
In 1953, James D. Watson and Francis Crick suggested what is now accepted as the first correct
double-helix model of DNA structure in the journal Nature.[5] Their double-helix, molecular model of DNA
was then based on a single X-ray diffraction image (labeled as "Photo 51")[144] taken by Rosalind Franklin
and Raymond Gosling in May 1952, as well as the information that the DNA bases are paired — also obtained
through private communications from Erwin Chargaff in the previous years. Chargaff's rules played a very
important role in establishing double-helix configurations for B-DNA as well as A-DNA.
Experimental evidence supporting the Watson and Crick model were published in a series of five
articles in the same issue of Nature.[145] Of these, Franklin and Gosling's paper was the first publication of
their own X-ray diffraction data and original analysis method that partially supported the Watson and Crick
model;[31][146] this issue also contained an article on DNA structure by Maurice Wilkins and two of his
colleagues, whose analysis and in vivo B-DNA X-ray patterns also supported the presence in vivo of the
double-helical DNA configurations as proposed by Crick and Watson for their double-helix molecular model
of DNA in the previous two pages of Nature.[32] In 1962, after Franklin's death, Watson, Crick, and Wilkins
jointly received the Nobel Prize in Physiology or Medicine.[147] However, Nobel rules of the time allowed
only living recipients, but a vigorous debate continues on who should receive credit for the discovery.[148]
In an influential presentation in 1957, Crick laid out the central dogma of molecular biology, which
foretold the relationship between DNA, RNA, and proteins, and articulated the "adaptor hypothesis".[149]
Final confirmation of the replication mechanism that was implied by the double-helical structure followed in
1958 through the Meselson–Stahl experiment.[150] Further work by Crick and coworkers showed that the
genetic code was based on non-overlapping triplets of bases, called codons, allowing Har Gobind Khorana,
Robert W. Holley and Marshall Warren Nirenberg to decipher the genetic code.[151] These findings
represent the birth of molecular biology.
See also
Molecular and Cellular Biology portal
• Crystallography
• DNA microarray
• DNA sequencing
• Genetic disorder
• Junk DNA
• Molecular models of DNA
• Molecular Structure of Nucleic Acids: A Structure for Deoxyribose Nucleic Acid
• Nucleic acid analogues
• Nucleic acid methods
• Nucleic acid modeling
• Nucleic acid notation
• Paracrystal model and theory
• X-ray crystallography
• X-ray scattering
• Phosphoramidite
• Plasmid
• Polymerase chain reaction
• Proteopedia DNA
• Southern blot
• Triple-stranded DNA
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Further reading
• Calladine, Chris R.; Drew, Horace R.; Luisi, Ben F. and Travers, Andrew A. (2003).
Understanding DNA: the molecule & how it works. Amsterdam: Elsevier Academic Press. ISBN 0-12-
155089-3.
• Dennis, Carina; Julie Clayton (2003). 50 years of DNA. Basingstoke: Palgrave Macmillan.
ISBN 1-4039-1479-6.
• Judson, Horace Freeland (1996). The eighth day of creation: makers of the revolution in
biology. Plainview, N.Y: CSHL Press. ISBN 0-87969-478-5.
• Olby, Robert C. (1994). The path to the double helix: the discovery of DNA . New York: Dover
Publications. ISBN 0-486-68117-3. , first published in October 1974 by MacMillan, with foreword by
Francis Crick;the definitive DNA textbook,revised in 1994 with a 9 page postscript.
• Olby, Robert C. (2009). Francis Crick: A Biography. Plainview, N.Y: Cold Spring Harbor
Laboratory Press. ISBN 0-87969-798-9.
• Ridley, Matt (2006). Francis Crick: discoverer of the genetic code. [Ashland, OH: Eminent
Lives, Atlas Books. ISBN 0-06-082333-X.
• Berry, Andrew; Watson, James D. (2003). DNA: the secret of life. New York: Alfred A. Knopf.
ISBN 0-375-41546-7.
• Stent, Gunther Siegmund; Watson, James D. (1980). The double helix: a personal account
of the discovery of the structure of DNA. New York: Norton. ISBN 0-393-95075-1.
• Wilkins, Maurice (2003). The third man of the double helix the autobiography of Maurice
Wilkins. Cambridge, Eng: University Press. ISBN 0-19-860665-6.
External links
This audio file was created from a revision of DNA dated 2007-02-12, and does not reflect subsequent edits to the article. (Audio help)
Wikimedia Commons has media related to: DNA
• DNA at the Open Directory Project

• DNA binding site prediction on protein
• DNA coiling to form chromosomes
• DNA from the Beginning Another DNA Learning Center site on DNA, genes, and heredity
from Mendel to the human genome project.
• DNA Lab, demonstrates how to extract DNA from wheat using readily available equipment
and supplies.
• DNA the Double Helix Game From the official Nobel Prize web site
• DNA under electron microscope
• Dolan DNA Learning Center
• Double Helix: 50 years of DNA, Nature
• Double Helix 1953–2003 National Centre for Biotechnology Education
• Francis Crick and James Watson talking on the BBC in 1962, 1972, and 1974
• Genetic Education Modules for Teachers —DNA from the Beginning Study Guide
• Guide to DNA cloning
• Olby R (January 2003). "Quiet debut for the double helix". Nature 421 (6921): 402–5.
doi:10.1038/nature01397. PMID 12540907. http://chem-faculty.ucsd.edu/joseph/CHEM13/DNA1.pdf.
• PDB Molecule of the Month pdb23_1
• Rosalind Franklin's contributions to the study of DNA
• The Register of Francis Crick Personal Papers 1938 - 2007 at Mandeville Special Collections
Library, Geisel Library, University of California, San Diego
• U.S. National DNA Day—watch videos and participate in real-time chat with top scientists
• "Clue to chemistry of heredity found" . The New York Times. Saturday, June 13, 1953.
http://www.nytimes.com/packages/pdf/science/dna-article.pdf. The first American newspaper
coverage of the discovery of the DNA structure.
• An Introduction to DNA and Chromosomes from HOPES: Huntington's Disease Outreach
Project for Education at Stanford
[hide]v · d · eGene expression
General flow: DNA > RNA > Protein

Introduction to
genetics special transfers (RNA > RNA, RNA > DNA, Protein > Protein)
Genetic code
(Transcription factors, RNA Polymerase,promoter) Prokaryotic /
Archaeal / Eukaryotic
Transcription
post-transcriptional modification (hnRNA,5'
capping,Splicing,Polyadenylation)
(Ribosome,tRNA) Prokaryotic / Archaeal / Eukaryotic

Translation
post-translational modification (functional groups, peptides, structural
changes)
epigenetic regulation (Genomic imprinting)

transcriptional regulation
Gene regulation post-transcriptional regulation (sequestration, alternative splicing,
miRNA)
translational regulation
post-translational regulation (reversible, irreversible)
[hide]v · d · eTypes of nucleic acids
Constituents Nucleobases · Nucleosides · Nucleotides · Deoxynucleotides

translation: mRNA (pre-mRNA/hnRNA) · tRNA · rRNA · tmRNA
Ribonucleic acids regulatory: miRNA · siRNA · piRNA · aRNA
(coding and non-coding)
RNA processing: snRNA · snoRNA
other/ungrouped: gRNA · shRNA · stRNA · ta-siRNA
Deoxyribonucleic acids cDNA · cpDNA · gDNA · msDNA · mtDNA
Nucleic acid analogues GNA · LNA · BNA · PNA · TNA · morpholino
Cloning vectors phagemid · plasmid · lambda phage · cosmid · fosmid · PAC · BAC ·
YAC · HAC
biochemical families: prot · nucl · carb (glpr, alco, glys) · lipd (fata/i, phld, strd, gllp, eico) · amac/i ·
ncbs/i · ttpy/i
[hide]v · d · eGenetics
Introduction · History · Related topics · List of organizations
Key components Chromosome · DNA · Nucleotide · RNA · Genome
Classical genetics · Conservation genetics · Ecological genetics ·

Fields of genetics Immunogenetics · Molecular genetics · Population genetics · Quantitative
genetics
Archaeogenetics
of... Americas · British Isles · Europe · Italy · Near East · South Asia
Related topics Geneticist · Genomics · Genetic code · Medical genetics · Molecular

evolution · Reverse genetics · Genetic engineering · Genetic diversity · Heredity
Retrieved from "http://en.wikipedia.org/wiki/DNA"
Categories: DNA | Genetics | Helices | Scattering

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Recapitulation theory

The theory of recapitulation, also called the biogenetic law or embryological parallelism - and often
expressed as "ontogeny recapitulates phylogeny" - is a hypothesis that in developing from embryo to adult,
animals go through stages resembling or representing successive stages in the evolution of their remote
ancestors. With different formulations, such ideas have been applied to several fields, including biology,
anthropology[1] and education theory.[2] In biology, there are several examples of embryonic stages showing
features of ancestral organisms, but a "strong" formulation of the concept has been discredited.
The concept originated in the 1790s among the German Natural philosophers[3] and, as proposed by
Étienne Serres in 1824–26, became known as the "Meckel-Serres Law". In 1866, the German zoologist Ernst
Haeckel proposed that the embryonic development of an individual organism (its ontogeny) followed the
same path as the evolutionary history of its species (its phylogeny).
Contents
[hide]
• 1 Origins
• 2 Haeckel
• 3 Historical influence
• 4 Modern observations
• 5 Notes
• 6 References
[edit] Origins
The general agreement among historians is that the concept originated in the 1790s among the
German Natural philosophers.[3] The first formal formulation was proposed by Étienne Serres in 1824–26 as
what became known as the "Meckel-Serres Law", it attempted to provide a link between comparative
embryology and a "pattern of unification" in the organic world. It was supported by Étienne Geoffroy Saint-
Hilaire and became a prominent part of his ideas which suggested that past transformations of life could have
had environmental causes working on the embryo, rather than on the adult as in Lamarckism. These
naturalistic ideas led to disagreements with Georges Cuvier. It was widely supported in the Edinburgh and
London schools of higher anatomy around 1830, notably by Robert Edmond Grant, but was opposed by Karl
Ernst von Baer's ideas of divergence, and attacked by Richard Owen in the 1830s.[4]
[edit] Haeckel
Romanes's 1892 copy of Ernst Haeckel's controversial embryo drawings (this version of the figure is
often attributed incorrectly to Haeckel).[5]
Haeckel attempted to synthesise the ideas of Lamarckism and Goethe's Naturphilosophie with
Charles Darwin's concepts. While often seen as rejecting Darwin's theory of branching evolution for a more
linear Lamarckian "biogenic law" of progressive evolution, this is not accurate: Haeckel used the Lamarckian
picture to describe the ontogenic and phylogenic history of the individual species, but agreed with Darwin
about the branching nature of all species from one, or a few, original ancestors.[6] Since around the start of
the twentieth century, Haeckel's "biogenic law" has been refuted on many fronts.[7]
Haeckel formulated his theory as "Ontogeny recapitulates phylogeny". The notion later became
simply known as the recapitulation (OED: 'a summing up or brief repetition') theory. Ontogeny is the growth
(size change) and development (shape change) of an individual organism; phylogeny is the evolutionary
history of a species. Haeckel's recapitulation theory claims that the development of advanced species passes
through stages represented by adult organisms of more primitive species.[7] Otherwise put, each successive
stage in the development of an individual represents one of the adult forms that appeared in its evolutionary
history.
For example, Haeckel proposed that the pharyngeal slits of the pharyngeal arches in the neck of the
human embryo resembled gill slits of fish, thus representing an adult "fishlike" developmental stage as well
as signifying a fishlike ancestor. Embryonic pharyngeal slits, formed when the thin branchial plates
separating pharyngeal pouches and ectodermal grooves perforate, open the pharynx to the outside.
Pharyngeal pouches appear in all tetrapod animal embryos: in mammals, the first pharyngeal pouch
develops into the lower jaw (Meckel's cartilage), the malleus and the stapes. At a later stage, all pharyngeal
slits close, only the ear remaining open.[8] But these embryonic pharyngeal arches, pouches, and slits could
not at any stage carry out the same function as the gills of an adult fish.
Haeckel produced several embryo drawings that often overemphasized similarities between embryos
of related species. These found their ways into many biology textbooks, and into popular knowledge. Modern
biology rejects the literal and universal form of Haeckel's theory.[9]
Although humans share ancestors with other taxa, stages of human embryonic development are not
functionally equivalent to the adults of these shared common ancestors. In other words, no cleanly defined
and functional "fish", "reptile" and "mammal" stages of human embryonal development can be discerned.
Moreover, development is nonlinear. For example, during kidney development, at one given time, the anterior
region of the kidney is less developed (nephridium) than the posterior region (nephron).
Darwin's view, that early embryonic stages are similar to the same embryonic stage of related
species but not to the adult stages of these species, has been confirmed by modern evolutionary
developmental biology.
[edit] Historical influence

Although Haeckel's specific form of recapitulation theory is now discredited among biologists, it had a
strong influence on social and educational theories of the late 19th century.
English philosopher Herbert Spencer was one of the most energetic promoters of evolutionary ideas
to explain many phenomena. He compactly expressed the basis for a cultural recapitulation theory of
education in the following claim, published in 1861, five years before Haeckel first published on the subject:
[2]
If there be an order in which the human race has mastered its various kinds of knowledge, there
will arise in every child an aptitude to acquire these kinds of knowledge in the same order....
Education is a repetition of civilization in little.[10]
– Herbert Spencer
The maturationist theory of G. Stanley Hall was based on the premise that growing children would
recapitulate evolutionary stages of development as they grew up and that there was a one-to-one
correspondence between childhood stages and evolutionary history, and that it was counterproductive to
push a child ahead of its development stage. The whole notion fit nicely with other social Darwinist concepts,
such as the idea that "primitive" societies needed guidance by more advanced societies, i.e. Europe and
North America, which were considered by social Darwinists as the pinnacle of evolution.[ citation needed]
The Austrian neurologist Sigmund Freud also held a favorable position towards Haeckel's doctrine.
He was trained as a biologist under the influence of recapitulation theory at the time of its domination, and
retained a Lamarckian outlook with justification from the recapitulation theory.[11] He also distinguished
between physical and mental recapitulation, in which the differences would become an essential argument
for his theory of neuroses.[11]
[edit] Modern observations

This section does not cite any references or sources.
Please help improve this article by adding citations to reliable sources. Unsourced material
may be challenged and removed. (August 2009)
Generally, if a structure pre-dates another structure in evolutionary terms, then it also appears earlier
than the other in the embryo. Species which have an evolutionary relationship typically share the early stages
of embryonal development and differ in later stages. Examples include:
• The backbone, the common structure among all vertebrates such as fish, reptiles and
mammals, appears as one of the earliest structures laid out in all vertebrate embryos.
• The cerebrum in humans, the most sophisticated part of the brain, develops last.
If a structure vanished in an evolutionary sequence, then one can often observe a corresponding
structure appearing at one stage during embryonic development, only to disappear or become modified in a
later stage. Examples include:
• Whales, which have evolved from land mammals, don't have legs, but tiny remnant leg
bones lie buried deep in their bodies. During embryonal development, leg extremities first occur, then
recede. Similarly, whale embryos have hair at one stage (like all mammalian embryos), but lose most
of it later.
• The common ancestor of humans and monkeys had a tail, and human embryos also have a
tail at one point; it later recedes to form the coccyx.
• The swim bladder in fish presumably evolved from a sac connected to the gut, allowing the
fish to gulp air. In most modern fish, this connection to the gut has disappeared. In the embryonal
development of these fish, the swim bladder originates as an outpocketing of the gut, and is later
disconnected from the gut.
• In bird embryos, very briefly fingers start to develop. After a short time, they partly disappear
again, partly are fused with the handbones to form the carpometacarpus.
But this rule-of-thumb is not universal. Modern birds, for example, though descended from toothed
animals, never grow teeth even as embryos. However, they still possess the genes required to do so, and
tooth formation has been experimentally induced in chickens.
[edit] Notes
1. ^ Carneiro, (1981) Robert L. Herbert Spencer as an Anthropologist Journal of Libertarian
Studies, vol. 5, 1981, pp.156-60
2. ^ a b Kieran Egan, The Educated Mind: How Cognitive Tools Shape Our Understanding.,
p.27 (University of Chicago Press, 1997, Chicago. ISBN 0-226-19036-6)
3. ^ a b Mayr (1994) [1] in The Quarterly Review of Biology
4. ^ Desmond 1989, pp. 52–53, 86–88, 337–340
5. ^ Richardson and Keuck, "Haeckel’s ABC of evolution and development," p. 516
6. ^ Richards, Robert J. 2008. The Tragic Sense of Life: Ernst Haeckel and the Struggle Over
Evolutionary Thought. Chicago: University of Chicago Press. Pp. 136-142
7. ^ a b Scott F Gilbert (2006). "Ernst Haeckel and the Biogenetic Law". Developmental Biology,
8th edition. Sinauer Associates. http://8e.devbio.com/article.php?id=219. Retrieved 2008-05-03.
"Eventually, the Biogenetic Law had become scientifically untenable."
8. ^ Toby White. "The Gill Arches: Meckel's Cartilage". paleos.com.
http://www.palaeos.com/Vertebrates/Bones/Gill_Arches/Meckelian.html. Retrieved 2008-04-30.
9. ^ Gerhard Medicus (1992). "The Inapplicability of the Biogenetic Rule to Behavioral
Development" (PDF). Human Development 35 (1): 1–8. ISSN 0018-716X/92/0351/0001-0008.
http://homepage.uibk.ac.at/~c720126/humanethologie/ws/medicus/block6/HumanDevelopment.pdf.
Retrieved 2008-04-30. "The present interdisciplinary article offers cogent reasons why the biogenetic
rule has no relevance for behavioral ontogeny. ... In contrast to anatomical ontogeny, in the case of
behavioral ontogeny there are no empirical indications of 'behavioral interphenes, that developed
phylogenetically from (primordial) behavioral metaphenes. ... These facts lead to the conclusion that
attempts to establish a psychological theory on the basis of the biogenetic rule will not be fruitful.".
10.^ Herbert Spencer (1861). Education. pp. 5. http://www.questia.com/PM.qst?
a=o&d=98953755.
11.^ a b Gould 1977, pp. 156–158
[edit] References
• Division of Biology and Medicine, Brown University. "Evolution and Development I: Size and
shape". http://biomed.brown.edu/Courses/BIO48/30.S&S.HTML.
• Haeckel, E (1899). "Riddle of the Universe at the Close of the Nineteenth Century".
http://www.ucmp.berkeley.edu/history/haeckel.html.
• Richardson, M., et al. (1997). "There is no highly conserved stage in the vertebrates:
implications for current theories of evolution and development". Anatomy and Embryology 196 (2):
91–106. doi:10.1007/s004290050082. PMID 9278154.
• Gould, Stephen Jay (1977). Ontogeny and Phylogeny. Cambridge Mass.: Belknap Press of
Harvard University Press. ISBN 0-674-63941-3.
• Desmond, Adrian J. (1989). The politics of evolution: morphology, medicine, and reform in
radical London. Chicago: University of Chicago Press. ISBN 0-226-14374-0.

Look up ontogeny recapitulates phylogeny in Wiktionary, the free dictionary.
• Of Parts and Wholes: Self-similarity and Synecdoche in Science, Culture and Literature
• "Ontology Recapitulates Philology", a parody phrase
Retrieved from "http://en.wikipedia.org/wiki/Recapitulation_theory"
Categories: Biology theories | History of evolutionary biology | Obsolete scientific theories

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Species

For other uses, see Species (disambiguation).
The hierarchy of biological classification's eight major taxonomic ranks, which is an example of
definition by genus and differentia. A genus contains one or more species. Intermediate minor rankings are
not shown.
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A
species is often defined as a group of organisms capable of interbreeding and producing fertile offspring.
While in many cases this definition is adequate, more precise or differing measures are often used, such as
similarity of DNA, morphology or ecological niche. Presence of specific locally adapted traits may further
subdivide species into subspecies.
The commonly used names for plant and animal taxa sometimes correspond to species: for example,
"lion," "walrus," and "Camphor tree" – each refers to a species. In other cases common names do not: for
example, "deer" refers to a family of 34 species, including Eld's Deer, Red Deer and Elk (Wapiti). The last
two species were once considered a single species, illustrating how species boundaries may change with
increased scientific knowledge.
Each species is placed within a single genus. This is a hypothesis that the species is more closely
related to other species within its genus than to species of other genera. All species are given a binomial
name consisting of the generic name and specific name (or specific epithet). For example, Boa constrictor,
which is commonly called by its bionomial name, and is one of five species of the Boa genus.
A usable definition of the word "species" and reliable methods of identifying particular species are
essential for stating and testing biological theories and for measuring biodiversity. Traditionally, multiple
examples of a proposed species must be studied for unifying characters before it can be regarded as a
species. Extinct species known only from fossils are generally difficult to assign precise taxonomic rankings.
Because of the difficulties with both defining and tallying the total numbers of different species in the
world, it is estimated that there are anywhere between 2 and 100 million different species.[1]
Contents
[hide]
• 1 Biologists' working definition
• 1.1 Common names and species
• 1.2 Placement within genera
• 1.3 Abbreviated names
• 2 Difficulty of defining "species" and identifying particular
species
• 3 Definitions of species
• 4 Numbers of species
• 5 Importance in biological classification
• 6 Implications of assignment of species status
• 7 Historical development of the species concept
• 8 Species as taxa
• 9 See also
• 10 Notes and references
[edit] Biologists' working definition
A usable definition of the word "species" and reliable methods of identifying particular species is
essential for stating and testing biological theories and for measuring biodiversity. Traditionally, multiple
examples of a proposed species must be studied for unifying characters before it can be regarded as a
species. It is generally difficult to give precise taxonomic rankings to extinct species known only from fossils.
Some biologists may view species as statistical phenomena, as opposed to the traditional idea, with
a species seen as a class of organisms. In that case, a species is defined as a separately evolving lineage
that forms a single gene pool. Although properties such as DNA-sequences and morphology are used to help
separate closely related lineages,[2] this definition has fuzzy boundaries.[3] However, the exact definition of
the term "species" is still controversial, particularly in prokaryotes,[4] and this is called the species problem.
[5] Biologists have proposed a range of more precise definitions, but the definition used is a pragmatic choice
that depends on the particularities of the species of concern.[5]
[edit] Common names and species

The commonly used names for plant and animal taxa sometimes correspond to species: for example,
"lion", "walrus", and "Camphor tree" – each refers to a species. In other cases common names do not: for
example, "deer" refers to a family of 34 species, including Eld's Deer, Red Deer and Elk (Wapiti). The last
two species were once considered a single species, illustrating how species boundaries may change with
increased scientific knowledge.
Because of the difficulties with both defining and tallying the total numbers of different species in the
world, it is estimated that there are anywhere between 2 and 100 million different species.[1]
[edit] Placement within genera
Ideally, a species is given a formal, scientific name, although in practice there are very many
unnamed species (which have only been described, not named). When a species is named, it is placed
within a genus. From a scientific point of view this can be regarded as a hypothesis that the species is more
closely related to other species within its genus (if any) than to species of other genera. Species and genus
are usually defined as part of a larger taxonomic hierarchy. The best-known taxonomic ranks are, in order:
life, domain, kingdom, phylum, class, order, family, genus, and species. This assignment to a genus is not
immutable; later a different (or the same) taxonomist may assign it to a different genus, in which case the
name will also change.
In biological nomenclature, the name for a species is a two-part name (a binomial name), treated as
Latin, although roots from any language can be used as well as names of locales or individuals. The generic
name is listed first (with its leading letter capitalized), followed by a second term, the specific name (or
specific epithet). For example, the species commonly known as the Longleaf Pine is Pinus palustris; gray
wolves belong to the species Canis lupus, coyotes to Canis latrans, golden jackals to Canis aureus, etc., and
all of those belong to the genus Canis (which also contains many other species). The name of the species is
the whole binomial, not just the second term (which may be called the specific name for animals).
This binomial naming convention, later formalized in the biological codes of nomenclature, was first
used by Leonhart Fuchs and introduced as the standard by Carolus Linnaeus in his 1753, Species Plantarum
(followed by his, 1758 Systema Naturae, 10th edition). At that time, the chief biological theory was that
species represented independent acts of creation by God and were therefore considered objectively real and
immutable, so the hypothesis of common descent did not apply.
[edit] Abbreviated names
Books and articles sometimes intentionally do not identify species fully and use the abbreviation "sp."
in the singular or "spp." in the plural in place of the specific epithet: for example, Canis sp. This commonly
occurs in the following types of situations:
• The authors are confident that some individuals belong to a particular genus but are not sure
to which exact species they belong. This is particularly common in paleontology.
• The authors use "spp." as a short way of saying that something applies to many species
within a genus, but do not wish to say that it applies to all species within that genus. If scientists
mean that something applies to all species within a genus, they use the genus name without the
specific epithet.
In books and articles, genus and species names are usually printed in italics. If using "sp." and
"spp.", these should not be italicized.
[edit] Difficulty of defining "species" and identifying particular species

Main article: Species problem
The Greenish Warbler demonstrates the concept of a ring species.
It is surprisingly difficult to define the word "species" in a way that applies to all naturally occurring
organisms, and the debate among biologists about how to define "species" and how to identify actual species
is called the species problem. Over two dozen distinct definitions of "species" are in use amongst biologists.
[6]
Most textbooks follow Ernst Mayr's definition of a species as "groups of actually or potentially
interbreeding natural populations, which are reproductively isolated from other such groups".[5]
Various parts of this definition serve to exclude some unusual or artificial matings:
• Those that occur only in captivity (when the animal's normal mating partners may not be
available) or as a result of deliberate human action
• Animals that may be physically and physiologically capable of mating but, for various
reasons, do not normally do so in the wild
The typical textbook definition above works well for most multi-celled organisms, but there are
several types of situations in which it breaks down:
• By definition it applies only to organisms that reproduce sexually. So it does not work for
asexually reproducing single-celled organisms and for the relatively few parthenogenetic multi-celled
organisms. The term "phylotype" is often applied to such organisms.
• Biologists frequently do not know whether two morphologically similar groups of organisms
are "potentially" capable of interbreeding.
• There is considerable variation in the degree to which hybridization may succeed under
natural conditions, or even in the degree to which some organisms use sexual reproduction between
individuals to breed.
• In ring species, members of adjacent populations interbreed successfully but members of
some non-adjacent populations do not.
• In a few cases it may be physically impossible for animals that are members of the same
species to mate. However, these are cases in which human intervention has caused gross
morphological changes, and are therefore excluded by the biological species concept.
Horizontal gene transfer makes it even more difficult to define the word "species". There is strong
evidence of horizontal gene transfer between very dissimilar groups of prokaryotes, and at least occasionally
between dissimilar groups of eukaryotes; and Williamson[7] argues that there is evidence for it in some
crustaceans and echinoderms. All definitions of the word "species" assume that an organism gets all its
genes from one or two parents that are very like that organism, but horizontal gene transfer makes that
assumption false.
[edit] Definitions of species
See also: Species problem
The question of how best to define "species" is one that has occupied biologists for centuries, and
the debate itself has become known as the species problem. Darwin wrote in chapter II of On the Origin of
Species:
No one definition has satisfied all naturalists; yet every naturalist knows vaguely what he
means when he speaks of a species. Generally the term includes the unknown element of a distinct
act of creation.[8]
But later, in The Descent of Man, when addressing "The question whether mankind consists of one
or several species", Darwin revised his opinion to say:
it is a hopeless endeavour to decide this point on sound grounds, until some definition of the
term "species" is generally accepted; and the definition must not include an element that cannot
possibly be ascertained, such as an act of creation.[9]
The modern theory of evolution depends on a fundamental redefinition of "species". Prior to Darwin,
naturalists viewed species as ideal or general types, which could be exemplified by an ideal specimen
bearing all the traits general to the species. Darwin's theories shifted attention from uniformity to variation
and from the general to the particular. According to intellectual historian Louis Menand,
Once our attention is redirected to the individual, we need another way of making
generalizations. We are no longer interested in the conformity of an individual to an ideal type; we are
now interested in the relation of an individual to the other individuals with which it interacts. To
generalize about groups of interacting individuals, we need to drop the language of types and
essences, which is prescriptive (telling us what finches should be), and adopt the language of statistics
and probability, which is predictive (telling us what the average finch, under specified conditions, is
likely to do). Relations will be more important than categories; functions, which are variable, will be
more important than purposes; transitions will be more important than boundaries; sequences will be
more important than hierarchies.
This shift results in a new approach to "species"; Darwin
concluded that species are what they appear to be: ideas, which are provisionally useful for
naming groups of interacting individuals. "I look at the term species", he wrote, "as one arbitrarily given
for the sake of convenience to a set of individuals closely resembling each other ... It does not
essentially differ from the word variety, which is given to less distinct and more fluctuating forms. The
term variety, again, in comparison with mere individual differences, is also applied arbitrarily, and for
convenience sake." [10]
Practically, biologists define species as populations of organisms that have a high level of genetic
similarity. This may reflect an adaptation to the same niche, and the transfer of genetic material from one
individual to others, through a variety of possible means. The exact level of similarity used in such a definition
is arbitrary, but this is the most common definition used for organisms that reproduce asexually (asexual
reproduction), such as some plants and microorganisms.
This lack of any clear species concept in microbiology has led to some authors arguing that the term
"species" is not useful when studying bacterial evolution. Instead they see genes as moving freely between
even distantly related bacteria, with the entire bacterial domain being a single gene pool. Nevertheless, a
kind of rule of thumb has been established, saying that species of Bacteria or Archaea with 16S rRNA gene
sequences more similar than 97% to each other need to be checked by DNA-DNA Hybridization if they
belong to the same species or not.[11] This concept has been updated recently, saying that the border of
97% was too low and can be raised to 98.7%.[12]
In the study of sexually reproducing organisms, where genetic material is shared through the process
of reproduction, the ability of two organisms to interbreed and produce fertile offspring of both sexes is
generally accepted as a simple indicator that the organisms share enough genes to be considered members
of the same species. Thus a "species" is a group of interbreeding organisms.
This definition can be extended to say that a species is a group of organisms that could potentially
interbreed – fish could still be classed as the same species even if they live in different lakes, as long as they
could still interbreed were they ever to come into contact with each other. On the other hand, there are many
examples of series of three or more distinct populations, where individuals of the population in the middle can
interbreed with the populations to either side, but individuals of the populations on either side cannot
interbreed. Thus, one could argue that these populations constitute a single species, or two distinct species.
This is not a paradox; it is evidence that species are defined by gene frequencies, and thus have fuzzy
boundaries.
Consequently, any single, universal definition of "species" is necessarily arbitrary. Instead, biologists
have proposed a range of definitions; which definition a biologists uses is a pragmatic choice, depending on
the particularities of that biologist's research.
Typological species
A group of organisms in which individuals are members of the species if they sufficiently
conform to certain fixed properties or "rights of passage". The clusters of variations or phenotypes
within specimens (i.e. longer or shorter tails) would differentiate the species. This method was used as
a "classical" method of determining species, such as with Linnaeus early in evolutionary theory.
However, we now know that different phenotypes do not always constitute different species (e.g.: a 4-
winged Drosophila born to a 2-winged mother is not a different species). Species named in this
manner are called morphospecies[13]
Morphological species
A population or group of populations that differs morphologically from other populations. For
example, we can distinguish between a chicken and a duck because they have different shaped bills
and the duck has webbed feet. Species have been defined in this way since well before the beginning
of recorded history. This species concept is highly criticized because more recent genetic data reveal
that genetically distinct populations may look very similar and, contrarily, large morphological
differences sometimes exist between very closely related populations. Nonetheless, most species
known have been described solely from morphology.
Biological / Isolation species
A set of actually or potentially interbreeding populations. This is generally a useful formulation
for scientists working with living examples of the higher taxa like mammals, fish, and birds, but more
problematic for organisms that do not reproduce sexually. The results of breeding experiments done in
artificial conditions may or may not reflect what would happen if the same organisms encountered
each other in the wild, making it difficult to gauge whether or not the results of such experiments are
meaningful in reference to natural populations.
Biological / reproductive species
Two organisms that are able to reproduce naturally to produce fertile offspring of both sexes.
Organisms that can reproduce but almost always make infertile hybrids of at least one sex, such as a
mule, hinny or F1 male cattalo are not considered to be the same species.
Recognition species
Based on shared reproductive systems, including mating behavior. The Recognition concept
of species has been introduced by Hugh E. H. Paterson.
Mate-recognition species
A group of organisms that are known to recognize one another as potential mates. Like the
isolation species concept above, it applies only to organisms that reproduce sexually. Unlike the
isolation species concept, it focuses specifically on pre-mating reproductive isolation.
Evolutionary / Darwinian species
A group of organisms that shares an ancestor; a lineage that maintains its integrity with
respect to other lineages through both time and space. At some point in the progress of such a group,
some members may diverge from the main population and evolve into a subspecies, a process that
eventually will lead to the formation of a new full species if isolation (geographical or ecological) is
maintained.
Phylogenetic (Cladistic)[verification needed]
A group of organisms that shares an ancestor; a lineage that maintains its integrity with
respect to other lineages through both time and space. At some point in the progress of such a group,
members may diverge from one another: when such a divergence becomes sufficiently clear, the two
populations are regarded as separate species. This differs from evolutionary species in that the parent
species goes extinct taxonomically when a new species evolve, the mother and daughter populations
now forming two new species. Subspecies as such are not recognized under this approach; either a
population is a phylogenetic species or it is not taxonomically distinguishable.
Ecological species
A set of organisms adapted to a particular set of resources, called a niche, in the environment.
According to this concept, populations form the discrete phonetic clusters that we recognize as species
because the ecological and evolutionary processes controlling how resources are divided up tend to
produce those clusters.
Genetic species
Based on similarity of DNA of individuals or populations. Techniques to compare similarity of
DNA include DNA-DNA hybridization, and genetic fingerprinting (or DNA barcoding).
Phenetic species
Based on phenotypes.[verification needed]
Microspecies
Species that reproduce without meiosis or fertilization so that each generation is genetically
identical to the previous generation. See also apomixis.
Cohesion species
Most inclusive population of individuals having the potential for phenotypic cohesion through
intrinsic cohesion mechanisms. This is an expansion of the mate-recognition species concept to allow
for post-mating isolation mechanisms; no matter whether populations can hybridize successfully, they
are still distinct cohesion species if the amount of hybridization is insufficient to completely mix their
respective gene pools.
Evolutionarily Significant Unit (ESU)
An evolutionarily significant unit is a population of organisms that is considered distinct for
purposes of conservation. Often referred to as a species or a wildlife species, an ESU also has several
possible definitions, which coincide with definitions of species.
In practice, these definitions often coincide, and the differences between them are more a matter of
emphasis than of outright contradiction. Nevertheless, no species concept yet proposed is entirely objective,
or can be applied in all cases without resorting to judgment. Given the complexity of life, some have argued
that such an objective definition is in all likelihood impossible, and biologists should settle for the most
practical definition.
For most vertebrates, this is the biological species concept (BSC), and to a lesser extent (or for
different purposes) the phylogenetic species concept (PSC). Many BSC subspecies are considered species
under the PSC; the difference between the BSC and the PSC can be summed up insofar as that the BSC
defines a species as a consequence of manifest evolutionary history, while the PSC defines a species as a
consequence of manifest evolutionary potential. Thus, a PSC species is "made" as soon as an evolutionary
lineage has started to separate, while a BSC species starts to exist only when the lineage separation is
complete. Accordingly, there can be considerable conflict between alternative classifications based upon the
PSC versus BSC, as they differ completely in their treatment of taxa that would be considered subspecies
under the latter model (e.g., the numerous subspecies of honey bees).
[edit] Numbers of species
Undiscovered and discovered species[verification needed][citation needed]

Bearing in mind the aforementioned problems with categorising species, the following numbers are
only a soft guide. In 2007, they broke down as follows:[14]
Total number of species (estimated): 7–100 millions (identified and unidentified), including:
• 5–10 million bacteria;[15]
• 74,000–120,000 fungi;[16]
Of the identified eukaryote species we have:[14]
• 1.6 million, including:
• 297,326 plants, including:
• 15,000 mosses,
• 13,025 Ferns and horsetails,
• 980 gymnosperms,
• 258,650 angiosperms,
• 199,350 dicotyledons,
• 59,300 monocotyledons,
• 9,671 Red and green algae,
• 28,849 fungi & other non-animals, including:
• 10,000 lichens,
• 16,000 mushrooms,
• 2,849 brown algae,
• 1,250,000 animals, including:
• 1,203,375 invertebrates:
• 950,000 insects,
• 81,000 mollusks,
• 40,000 crustaceans,
• 2,175 corals,
• 130,200 others;
• 59,811 vertebrates:
• 29,300 fish,
• 6,199 amphibians,
• 8,240 reptiles,
• 9,956 birds,
• 5,416 mammals
At present, organisations such as the Global Taxonomy Initiative, the European Distributed Institute
of Taxonomy and the Census of Marine Life[17] (the latter only for marine organisms) are trying to improve
taxonomy and implement previously undiscovered species to the taxonomy system. Because we know but a
portion of the organisms in the biosphere, we do not have a complete understanding of the workings of our
environment. To make matters worse, despite the discovery of new species, according to professor James
Mallet, we are wiping out these species at an unprecedented rate.[18] This means that even before a new
species has had the chance of being studied and classified, it may already be extinct.
[edit] Importance in biological classification

The idea of species has a long history. It is one of the most important levels of classification, for
several reasons:
• It often corresponds to what lay people treat as the different basic kinds of organism – dogs
are one species, cats another.
• It is the standard binomial nomenclature (or trinomial nomenclature) by which scientists
typically refer to organisms.
• It is the highest taxonomic level that cannot be made more or less inclusionary.
After years of use, the concept remains central to biology and a host of related fields, and yet also
remains at times ill-defined.
[edit] Implications of assignment of species status

The naming of a particular species may be regarded as a hypothesis about the evolutionary
relationships and distinguishability of that group of organisms. As further information comes to hand, the
hypothesis may be confirmed or refuted. Sometimes, especially in the past when communication was more
difficult, taxonomists working in isolation have given two distinct names to individual organisms later
identified as the same species. When two named species are discovered to be of the same species, the older
species name is usually retained, and the newer species name dropped, a process called synonymization, or
colloquially, as lumping. Dividing a taxon into multiple, often new, taxons is called splitting. Taxonomists are
often referred to as "lumpers" or "splitters" by their colleagues, depending on their personal approach to
recognizing differences or commonalities between organisms (see lumpers and splitters).
Traditionally, researchers relied on observations of anatomical differences, and on observations of
whether different populations were able to interbreed successfully, to distinguish species; both anatomy and
breeding behavior are still important to assigning species status. As a result of the revolutionary (and still
ongoing) advance in microbiological research techniques, including DNA analysis, in the last few decades, a
great deal of additional knowledge about the differences and similarities between species has become
available. Many populations formerly regarded as separate species are now considered a single taxon, and
many formerly grouped populations have been split. Any taxonomic level (species, genus, family, etc.) can
be synonymized or split, and at higher taxonomic levels, these revisions have been still more profound.
From a taxonomical point of view, groups within a species can be defined as being of a taxon
hierarchically lower than a species. In zoology only the subspecies is used, while in botany the variety,
subvariety, and form are used as well. In conservation biology, the concept of evolutionary significant units
(ESU) is used, which may be define either species or smaller distinct population segments. Identifying and
naming species is the providence of alpha taxonomy.
[edit] Historical development of the species concept

The following text needs to be harmonized with text in Species problem.
This section does not cite any references or sources.

may be challenged and removed. (September 2009)
Linnaeus believed in the fixity of species.
In the earliest works of science, a species was simply an individual organism that represented a
group of similar or nearly identical organisms. No other relationships beyond that group were implied.
Aristotle used the words genus and species to mean generic and specific categories. Aristotle and other pre-
Darwinian scientists took the species to be distinct and unchanging, with an "essence", like the chemical
elements. When early observers began to develop systems of organization for living things, they began to
place formerly isolated species into a context. Many of these early delineation schemes would now be
considered whimsical and these included consanguinity based on color (all plants with yellow flowers) or
behavior (snakes, scorpions and certain biting ants).
In the 18th century Swedish scientist Carolus Linnaeus classified organisms according to differences
in the form of reproductive apparatus. Although his system of classification sorts organisms according to
degrees of similarity, it made no claims about the relationship between similar species. At that time, it was
still widely believed that there was no organic connection between species, no matter how similar they
appeared. This approach also suggested a type of idealism: the notion that each species existed as an "ideal
form". Although there are always differences (although sometimes minute) between individual organisms,
Linnaeus considered such variation problematic. He strove to identify individual organisms that were
exemplary of the species, and considered other non-exemplary organisms to be deviant and imperfect.
By the 19th century most naturalists understood that species could change form over time, and that
the history of the planet provided enough time for major changes. Jean-Baptiste Lamarck, in his 1809
Zoological Philosophy, offered one of the first logical arguments against creationism. The new emphasis was
on determining how a species could change over time. Lamarck suggested that an organism could pass on
an acquired trait to its offspring, i.e., the giraffe's long neck was attributed to generations of giraffes stretching
to reach the leaves of higher treetops (this well-known and simplistic example, however, does not do justice
to the breadth and subtlety of Lamarck's ideas). With the acceptance of the natural selection idea of Charles
Darwin in the 1860s, however, Lamarck's view of goal-oriented evolution, also known as a teleological
process, was eclipsed. Recent interest in inheritance of acquired characteristics centers around epigenetic
processes, e.g. methylation, that do not affect DNA sequences, but instead alter expression in an inheritable
manner. Thus, neo-lamarckism, as it is sometimes termed, is not a challenge to the theory of evolution by
natural selection.
Charles Darwin and Alfred Wallace provided what scientists now consider as the most powerful and
compelling theory of evolution. Darwin argued that it was populations that evolved, not individuals. His
argument relied on a radical shift in perspective from that of Linnaeus: rather than defining species in ideal
terms (and searching for an ideal representative and rejecting deviations), Darwin considered variation
among individuals to be natural. He further argued that variation, far from being problematic, actually
provides the explanation for the existence of distinct species.
Darwin's work drew on Thomas Malthus' insight that the rate of growth of a biological population will
always outpace the rate of growth of the resources in the environment, such as the food supply. As a result,
Darwin argued, not all the members of a population will be able to survive and reproduce. Those that did will,
on average, be the ones possessing variations—however slight—that make them slightly better adapted to the
environment. If these variable traits are heritable, then the offspring of the survivors will also possess them.
Thus, over many generations, adaptive variations will accumulate in the population, while counter-adaptive
traits will tend to be eliminated.
Whether a variation is adaptive or non-adaptive depends on the environment: different environments
favor different traits. Since the environment effectively selects which organisms live to reproduce, it is the
environment (the "fight for existence") that selects the traits to be passed on. This is the theory of evolution
by natural selection. In this model, the length of a giraffe's neck would be explained by positing that proto-
giraffes with longer necks would have had a significant reproductive advantage to those with shorter necks.
Over many generations, the entire population would be a species of long-necked animals.
In 1859, when Darwin published his theory of natural selection, the mechanism behind the
inheritance of individual traits was unknown. Although Darwin made some speculations on how traits are
inherited (pangenesis), his theory relies only on the fact that inheritable traits exist, and are variable (which
makes his accomplishment even more remarkable.) Although Gregor Mendel's paper on genetics was
published in 1866, its significance was not recognized. It was not until 1900 that his work was rediscovered
by Hugo de Vries, Carl Correns and Erich von Tschermak, who realised that the "inheritable traits" in
Darwin's theory are genes.
The theory of the evolution of species through natural selection has two important implications for
discussions of species—consequences that fundamentally challenge the assumptions behind Linnaeus'
taxonomy. First, it suggests that species are not just similar, they may actually be related. Some students of
Darwin argue that all species are descended from a common ancestor. Second, it supposes that "species"
are not homogeneous, fixed, permanent things; members of a species are all different, and over time species
change. This suggests that species do not have any clear boundaries but are rather momentary statistical
effects of constantly changing gene-frequencies. One may still use Linnaeus' taxonomy to identify individual
plants and animals, but one can no longer think of species as independent and immutable.
The rise of a new species from a parental line is called speciation. There is no clear line demarcating
the ancestral species from the descendant species.
Although the current scientific understanding of species suggests that there is no rigorous and
comprehensive way to distinguish between different species in all cases, biologists continue to seek concrete
ways to operationalize the idea. One of the most popular biological definitions of species is in terms of
reproductive isolation; if two creatures cannot reproduce to produce fertile offspring of both sexes, then they
are in different species. This definition captures a number of intuitive species boundaries, but it remains
imperfect. It has nothing to say about species that reproduce asexually, for example, and it is very difficult to
apply to extinct species. Moreover, boundaries between species are often fuzzy: there are examples where
members of one population can produce fertile offspring of both sexes with a second population, and
members of the second population can produce fertile offspring of both sexes with members of a third
population, but members of the first and third population cannot produce fertile offspring, or can only produce
fertile offspring of the homozygous sex. Consequently, some people reject this definition of a species.
Richard Dawkins defines two organisms as conspecific if and only if they have the same number of
chromosomes and, for each chromosome, both organisms have the same number of nucleotides (The Blind
Watchmaker, p. 118). However, most if not all taxonomists would strongly disagree[citation needed]. For
example, in many amphibians, most notably in New Zealand's Leiopelma frogs, the genome consists of
"core" chromosomes that are mostly invariable and accessory chromosomes, of which exist a number of
possible combinations. Even though the chromosome numbers are highly variable between populations,
these can interbreed successfully and form a single evolutionary unit. In plants, polyploidy is extremely
commonplace with few restrictions on interbreeding; as individuals with an odd number of chromosome sets
are usually sterile, depending on the actual number of chromosome sets present, this results in the odd
situation where some individuals of the same evolutionary unit can interbreed with certain others and some
cannot, with all populations being eventually linked as to form a common gene pool.
The classification of species has been profoundly affected by technological advances that have
allowed researchers to determine relatedness based on molecular markers, starting with the comparatively
crude blood plasma precipitation assays in the mid-20th century to Charles Sibley's ground-breaking DNA-
DNA hybridization studies in the 1970s leading to DNA sequencing techniques. The results of these
techniques caused revolutionary changes in the higher taxonomic categories (such as phyla and classes),
resulting in the reordering of many branches of the phylogenetic tree (see also: molecular phylogeny). For
taxonomic categories below genera, the results have been mixed so far; the pace of evolutionary change on
the molecular level is rather slow, yielding clear differences only after considerable periods of reproductive
separation. DNA-DNA hybridization results have led to misleading conclusions, the Pomarine Skua – Great
Skua phenomenon being a famous example. Turtles have been determined to evolve with just one-eighth of
the speed of other reptiles on the molecular level, and the rate of molecular evolution in albatrosses is half of
what is found in the rather closely related storm-petrels. The hybridization technique is now obsolete and is
replaced by more reliable computational approaches for sequence comparison. Molecular taxonomy is not
directly based on the evolutionary processes, but rather on the overall change brought upon by these
processes. The processes that lead to the generation and maintenance of variation such as mutation,
crossover and selection are not uniform (see also molecular clock). DNA is only extremely rarely a direct
target of natural selection rather than changes in the DNA sequence enduring over generations being a result
of the latter; for example, silent transition-transversion combinations would alter the melting point of the DNA
sequence, but not the sequence of the encoded proteins and thus are a possible example where, for
example in microorganisms, a mutation confers a change in fitness all by itself.
[edit] Species as taxa

The scientific name of a species (often of Latin or Greek origin) in the binominal nomenclature
introduced by Carl Linnaeus in 1753 is composed of two parts, which are written in italic font and for which
there are different expressions in botany and zoology. The first part of that name is spelled upper case and is
in both disciplines known as the genus name (also called the generic name). The second part is always
spelled lower case and in botany it is called the specific epithet.
• Example: in the European beech (Fagus sylvatica) the component Fagus refers to the genus,
sylvatica is the specific epithet.
In zoology, the second part is called the specific name.
• Example: in the lion (Panthera leo) the component Panthera refers to the genus, leo is the
specific name.
The scientific name is completed when authors, years and parentheses are added.
In botany the name of the author is usually abbreviated, for example "L." stands for "Linnaeus".
• Example: shiitake Lentinula edodes (Berk.) Pegler
M. J. Berkeley was the first to describe the species, D. Pegler has placed it into the currently used
system.
In zoology, the addition of author and year are optional, Panthera leo is thus an entirely correct
name. The International Code of Zoological Nomenclature prescribes how to add author(s) (if possible not
abbreviated) and year (or author alone without year). If the species is cited in different genus than the one in
which it was originally described, author(s) and year are given in parentheses. Between author and year
often a comma is set (but not required).
• Example: lion Panthera leo (Linnaeus, 1758)
Carl Nilsson Linnæus described the lion first and as Felis leo. The person who first placed the lion in
the genus Panthera Oken, 1816 is not relevant in zoology. Instead of Linnæus, usually Linnaeus is
spelled.
[edit] See also

• Cline
• Cryptic species complex
• Encyclopedia of Life
• Endangered species
• Genetic pollution
• Genetic erosion
• Ring species
• Species problem
• Systematics
[edit] Notes and references

1. ^ a b "Just How Many Species Are There, Anyway?". 2003-05-26.
http://www.sciencedaily.com/releases/2003/05/030526103731.htm. Retrieved 2008-01-15
2. ^ Koch, H. 2010. Combining morphology and DNA barcoding resolves the taxonomy of
Western Malagasy Liotrigona Moure, 1961. African Invertebrates 51 (2): 413-421.[1]
3. ^ De Queiroz K (December 2007). "Species concepts and species delimitation". Syst. Biol.
56 (6): 879–86. doi:10.1080/10635150701701083. PMID 18027281.
4. ^ Fraser C, Alm EJ, Polz MF, Spratt BG, Hanage WP (February 2009). "The bacterial
species challenge: making sense of genetic and ecological diversity". Science (journal) 323 (5915):
741–6. doi:10.1126/science.1159388. PMID 19197054.
5. ^ a b c de Queiroz K (May 2005). "Ernst Mayr and the modern concept of species". Proc. Natl.
6. ^ Wilkins, John (2010-10-20). "How many species concepts are there?". The Guardian.
http://www.guardian.co.uk/science/punctuated-equilibrium/2010/oct/20/3. Retrieved 2010-10-19.
7. ^ David I. Williamson (2003). The Origins of Larvae. Kluwer. ISBN 1-4020-1514-3.
8. ^ Darwin 1859 p.59
9. ^ Darwin 1871 p. 24
10.^ Louis Menand (2001) The Metaphysical Club New York: Farrar, Straus and Giroux 123–124
11.^ Stackebrandt E, Goebel BM (1994). "Taxonomic note: a place for DNA-DNA reassociation
and 16S rRNA sequence analysis in the present species definition in bacteriology". Int. J. Syst.
Bacteriol. 44: 846–9. doi:10.1099/00207713-44-4-846.
12.^ Stackebrandt E, Ebers J (2006). "Taxonomic parameters revisited: tarnished gold
standards". Microbiol. Today 33: 152–5.
13.^ Michael Ruse (August 1969). "Definitions of Species in Biology". The British Journal for the
Philosophy of Science (Oxford University Press) 20 (2): 97–119. doi:10.1093/bjps/20.2.97.
http://www.jstor.org/stable/686173.
14.^ a b "Number of Species on Earth". Current Results. 2007-01-01.
http://www.currentresults.com/Environment-Facts/Plants-Animals/number-species.php. Retrieved
2010-04-23.
15.^ Sogin ML, Morrison HG, Huber JA, et al. (August 2006). "Microbial diversity in the deep sea
and the underexplored "rare biosphere"". Proc. Natl. Acad. Sci. U.S.A. 103 (32): 12115–20.
doi:10.1073/pnas.0605127103. PMID 16880384. PMC 1524930.
http://www.pnas.org/cgi/pmidlookup?view=long&pmid=16880384. Cheung L (Monday, 31 July 200).
"Thousands of microbes in one gulp". BBC. http://news.bbc.co.uk/1/hi/sci/tech/5232928.stm.
16.^ David L. Hawksworth (2001). "The magnitude of fungal diversity: the 1•5 million species
estimate revisited". Mycological Research 105 (12): 1422–1432. doi:10.1017/S0953756201004725.
17.^ "Census of marine life". Coml.org. http://www.coml.org/. Retrieved 2010-04-23.
18.^ Robin McKie and Zoe Corbyn (2005-09-25). "Discovery of new species and extermination
at high rate". London: Guardian.
http://www.guardian.co.uk/science/2005/sep/25/taxonomy.conservationandendangeredspecies.
Look up species in Wiktionary, the free dictionary.
Wikispecies has information related to: Species
• Stanford Encyclopedia of Philosophy entry

• Barcoding of species
• European Species Names in Linnaean, Czech, English, German and French
• Catalogue of Life
• VisualTaxa
• Other Species Concepts Berkeley University
• "Gone", Mother Jones, May/June 2007.
• Speciation
• 2003-12-31, ScienceDaily: Working On The 'Porsche Of Its Time': New Model For Species
Determination Offered Quote: "...two species of dinosaur that are members of the same genera
varied from each other by just 2.2%. Translation of the percentage into an actual number results in
an average of just three skeletal differences out of the total 338 bones in the body. Amazingly, 58%
of these differences occurred in the skull alone. "This is a lot less variation than I'd expected", said
Novak..."
• 2003-08-08, ScienceDaily: Cross-species Mating May Be Evolutionarily Important And Lead
To Rapid Change, Say Indiana University Researchers Quote: "...the sudden mixing of closely
related species may occasionally provide the energy to impel rapid evolutionary change..."
• 2004-01-09 ScienceDaily: Mayo Researchers Observe Genetic Fusion Of Human, Animal
Cells; May Help Explain Origin Of AIDS Quote: "...The researchers have discovered conditions in
which pig cells and human cells can fuse together in the body to yield hybrid cells that contain
genetic material from both species... "What we found was completely unexpected", says Jeffrey Platt,
M.D."
• 2000-09-18, ScienceDaily: Scientists Unravel Ancient Evolutionary History Of
Photosynthesis Quote: "...gene-swapping was common among ancient bacteria early in evolution..."
[hide]v · d · eSpeciation
Basic concepts Species · Cline · Chronospecies · Speciation
Modes of Allopatric · Heteropatric · Peripatric · Parapatric · Sympatric ·

speciation Polyploidy · Paleopolyploidy
Auxiliary
mechanisms Sexual selection · Assortative mating · Punctuated equilibrium
Intermediate stages Hybrid · Species complex · Ring species · Haldane's rule

[hide]v · d · e Taxonomic ranks
Magnorder
Domain/Super Superphylum/
Superclass Superorder Superfam
kingdom Superdivision
Phylum/Divisi
Kingdom Class Legion Order Family
on
Subkingdom Subphylum Subclass Cohort Suborder Subfamily
Infrakingdom/B
Infraphylum Infraclass Infraorder
ranch
Microphylum Parvclass Parvorder
Retrieved from "http://en.wikipedia.org/wiki/Species"
Categories: Biology | Scientific classification | Zoological nomenclature

W000
Mammal

For other uses, see Mammal (disambiguation).
Mammal
Mammals
Temporal range: Late Triassic – Recent, 220–0 Ma
PreЄ
g
Clockwise from the upper left: koala, White-winged vampire bat, Bottlenose Dolphin,
echidna
Scientific classification [ e ]
Kingdom: Animalia
Phylum: Chordata
Superclass: Tetrapoda
(unranked): Mammaliaformes
Mammalia
Class:
Linnaeus, 1758
Subgroups
• Subclass †Allotheria*
• Subclass Prototheria
• Subclass Theria
• Infraclass †Trituberculata
• Infraclass Metatheria
• Infraclass Eutheria
This article needs additional citations for verification.

Please help improve this article by adding reliable references. Unsourced material may be
challenged and removed. (August 2009)
Mammals (formally Mammalia) are a class of air-breathing vertebrate animals characterized by the
possession of hair, three middle ear bones, a neocortex, and mammary glands functional in mothers with
young. Most mammals also possess sweat glands and specialized teeth, and the largest group of mammals,
the placentals, have a placenta which feeds the offspring during gestation. The mammalian brain regulates
endothermic and circulatory systems, including a four-chambered heart. Mammals range in size from the 30–
40 millimeter (1- to 1.5-inch) Bumblebee Bat to the 33-meter (108-foot) Blue Whale.
Depending on classification scheme, there are approximately 5,500 species (5,490, according to the
IUCN Red List) of mammals, distributed in about 1,200 genera, 153 families, 29 orders[1] divided into three
infraclasses in two subclasses: the monotremes of the sublclass Prototheria, and the marsupials and
placentals or the subclass Theria. Except for the five species of monotremes (which lay eggs), all mammal
species give birth to live young. Most mammals, including the six largest orders, belong to the placental
group. The three largest orders, in descending order, are Rodentia (mice, rats, porcupines, beavers,
capybaras, and other gnawing mammals), Chiroptera (bats), and Soricomorpha (shrews, moles and
solenodons). The next three largest orders include the Carnivora (dogs, cats, weasels, bears, seals, and their
relatives), the Cetartiodactyla (including the even-toed hoofed mammals and the whales) and the Primates to
which the human species belongs.
The early synapsid mammalian ancestors, a group which included pelycosaurs such as Dimetrodon,
diverged from the amniote line that would lead to reptiles at the end of the Carboniferous period. Although
they were preceded by many diverse groups of non-mammalian synapsids (sometimes misleadingly referred
to as mammal-like reptiles), the first true mammals appeared in the Triassic period. Modern mammalian
orders appeared in the Palaeocene and Eocene epochs of the Palaeogene period. Phylogenetically, the
clade Mammalia is defined as all descendants of the most recent common ancestor of monotremes (e.g.,
echidnas and platypuses) and therian mammals (marsupials and placentals).[2] This means that some
extinct groups of "mammals" are not members of the crown group Mammalia, even though most of them
have all the characteristics that traditionally would have classified them as mammals.[3] These "mammals"
are now usually placed in the unranked clade Mammaliaformes.
Contents
[hide]
• 1 Distinguishing features
• 2 Classification
• 2.1 Standardized textbook classification
• 2.2 McKenna/Bell classification
• 2.3 Molecular classification of placentals
• 3.1 Evolution
• 3.2 Earliest appearances of features
• 4 Anatomy and morphology
• 4.1 Skeletal system
• 4.2 Respiratory system
• 4.3 Nervous system
• 4.4 Integumentary system
• 4.5 Reproductive system
• 5 Physiology
• 5.1 Endothermy
• 5.2 Intelligence
• 5.3 Social structure
• 5.4 Locomotion
• 5.4.1 Terrestrial
[edit] Distinguishing features
Living mammal species can be identified by the presence of sweat glands, including those that are
specialized to produce milk. However, other features are required when classifying fossils, since soft tissue
glands and some other features are not visible in fossils. Paleontologists use a distinguishing feature that is
shared by all living mammals (including monotremes), but is not present in any of the early Triassic
synapsids: mammals use two bones for hearing that were used for eating by their ancestors. The earliest
synapsids had a jaw joint composed of the articular (a small bone at the back of the lower jaw) and the
quadrate (a small bone at the back of the upper jaw). Most reptiles including lizards, crocodilians, dinosaurs
(and their descendants the birds) use this system, as did non-mammalian synapsids such as therapsids.
Mammals have a different jaw joint, however, composed only of the dentary (the lower jaw bone which
carries the teeth) and the squamosal (another small skull bone). In mammals the quadrate and articular
bones have become the incus and malleus bones in the middle ear.
Mammals also have a double occipital condyle: they have two knobs at the base of the skull which fit
into the topmost neck vertebra, and other vertebrates have a single occipital condyle. Paleontologists use
only the jaw joint and middle ear as criteria for identifying fossil mammals, since it would be confusing if they
found a fossil that had one feature, but not the other.
[edit] Classification
Main article: Mammal classification
Over 70% of mammal species are in the orders Rodentia (blue), Chiroptera (red), and Soricomorpha
(yellow)
George Gaylord Simpson's "Principles of Classification and a Classification of Mammals" (AMNH
Bulletin v. 85, 1945) was the original source for the taxonomy listed here. Simpson laid out a systematics of
mammal origins and relationships that was universally taught until the end of the 20th century. Since
Simpson's classification, the paleontological record has been recalibrated, and the intervening years have
seen much debate and progress concerning the theoretical underpinnings of systematization itself, partly
through the new concept of cladistics. Though field work gradually made Simpson's classification outdated, it
remained the closest thing to an official classification of mammals.
[edit] Standardized textbook classification

A somewhat standardized classification system has been adopted by most current mammalogy
classroom textbooks. The following taxonomy of extant and recently extinct mammals is from Vaughan et al.
(2000).
Class Mammalia
• Subclass Prototheria: monotremes: platypuses and echidnas
• Subclass Theria: live-bearing mammals
• Infraclass Metatheria: marsupials
• Infraclass Eutheria: placentals
[edit] McKenna/Bell classification

In 1997, the mammals were comprehensively revised by Malcolm C. McKenna and Susan K. Bell,
which has resulted in the "McKenna/Bell classification".
McKenna and Bell, Classification of Mammals: Above the species level , (1997) is the most
comprehensive work to date on the systematics, relationships, and occurrences of all mammal taxa, living
and extinct, down through the rank of genus. The new McKenna/Bell classification was quickly accepted by
paleontologists, though recent molecular genetic data challenge several of the higher level groupings. The
authors work together as paleontologists at the American Museum of Natural History, New York. McKenna
inherited the project from Simpson and, with Bell, constructed a completely updated hierarchical system,
covering living and extinct taxa that reflects the historical genealogy of Mammalia.
The McKenna/Bell hierarchical listing of all of the terms used for mammal groups above the species
includes extinct mammals as well as modern groups, and introduces some fine distinctions such as legions
and sublegions (ranks which fall between classes and orders) that are likely to be glossed over by the
nonprofessionals.
The published re-classification forms both a comprehensive and authoritative record of approved
names and classifications and a list of invalid names.
Extinct groups are represented by a cross (†).
Class Mammalia
• Subclass Prototheria: monotremes: echidnas and the Platypus
• Subclass Theriiformes: live-bearing mammals and their prehistoric relatives
• Infraclass †Allotheria: multituberculates
• Infraclass †Triconodonta: triconodonts
• Infraclass Holotheria: modern live-bearing mammals and their prehistoric relatives
• Supercohort Theria: live-bearing mammals
• Cohort Marsupialia: marsupials
• Magnorder Australidelphia: Australian marsupials and the
Monito del Monte
• Magnorder Ameridelphia: New World marsupials
• Cohort Placentalia: placentals
• Magnorder Xenarthra: xenarthrans
• Magnorder Epitheria: epitheres
• Grandorder Anagalida: lagomorphs, rodents, and
elephant shrews
• Grandorder Ferae: carnivorans, pangolins,
†creodonts, and relatives
• Grandorder Lipotyphla: insectivorans
• Grandorder Archonta: bats, primates, colugos, and
treeshrews
• Grandorder Ungulata: ungulates
• Order Tubulidentata incertae sedis: aardvark
• Mirorder Eparctocyona: †condylarths,
whales, and artiodactyls (even-toed ungulates)
• Mirorder †Meridiungulata: South American
ungulates
• Mirorder Altungulata: perissodactyls (odd-
toed ungulates), elephants, manatees, and hyraxes
[edit] Molecular classification of placentals

Molecular studies based on DNA analysis have suggested new relationships among mammal
families over the last few years. Most of these findings have been independently validated by retrotransposon
presence/absence data. The most recent classification systems based on molecular studies have proposed
four groups or lineages of placental mammals. Molecular clocks suggest that these clades diverged from
early common ancestors in the Cretaceous, but fossils have not yet been found to corroborate this
hypothesis. These molecular findings are consistent with mammal zoogeography:
Following molecular DNA sequence analyses, the first divergence was that of the Afrotheria 110–100
million years ago. The Afrotheria proceeded to evolve and diversify in the isolation of the African-Arabian
continent. The Xenarthra, isolated in South America, diverged from the Boreoeutheria approximately 100–95
million years ago. According to an alternative view, the Xenarthra has the Afrotheria as closest allies, forming
the Atlantogenata as sistergroup to Boreoeutheria. The Boreoeutheria split into the Laurasiatheria and
Euarchontoglires between 95 and 85 mya; both of these groups evolved on the northern continent of
Laurasia. After tens of millions of years of relative isolation, Africa-Arabia collided with Eurasia, exchanging
Afrotheria and Boreoeutheria. The formation of the Isthmus of Panama linked South America and North
America, which facilitated the exchange of mammal species in the Great American Interchange. The
traditional view that no placental mammals reached Australasia until about 5 million years ago when bats and
murine rodents arrived has been challenged by recent evidence and may need to be reassessed. These
molecular results are still controversial because they are not reflected by morphological data, and thus not
accepted by many systematists. Further there is some indication from retrotransposon presence/absence
data that the traditional Epitheria hypothesis, suggesting Xenarthra as the first divergence, might be true.
With the old order Insectivora shown to be polyphylectic and more properly subdivided (as Afrosoricida,
Erinaceomorpha, and Soricomorpha), the following classification for placental mammals contains 21 orders:
• Clade Atlantogenata
• Group I: Afrotheria
• Clade Afroinsectiphilia
• Order Macroscelidea: elephant shrews (Africa)
• Order Afrosoricida: tenrecs and golden moles (Africa)
• Order Tubulidentata: aardvark (Africa south of the Sahara)
• Clade Paenungulata
• Order Hyracoidea: hyraxes or dassies (Africa, Arabia)
• Order Proboscidea: elephants (Africa, Southeast Asia)
• Order Sirenia: dugong and manatees (cosmopolitan tropical)
• Group II: Xenarthra
• Order Pilosa: sloths and anteaters (Neotropical)
• Order Cingulata: armadillos (Americas)
• Clade Boreoeutheria
• Group III: Euarchontoglires (Supraprimates)
• Superorder Euarchonta
• Order Scandentia: treeshrews (Southeast Asia).
• Order Dermoptera: flying lemurs or colugos (Southeast Asia)
• Order Primates: lemurs, bushbabies, monkeys, apes (cosmopolitan),
humans
• Superorder Glires
• Order Lagomorpha: pikas, rabbits, hares (Eurasia, Africa, Americas)
• Order Rodentia: rodents (cosmopolitan)
• Group IV: Laurasiatheria
• Order Erinaceomorpha: hedgehogs
• Order Soricomorpha: moles, shrews, solenodons
• Clade Ferungulata
• Clade Cetartiodactyla
• Order Cetacea: whales, dolphins and porpoises
• Order Artiodactyla: even-toed ungulates, including pigs,
hippopotamus, camels, giraffe, deer, antelope, cattle, sheep, goats
• Clade Pegasoferae
• Order Chiroptera: bats (cosmopolitan)
• Clade Zooamata
• Order Perissodactyla: odd-toed ungulates, including
horses, donkeys, zebras, tapirs, and rhinoceroses
• Clade Ferae
• Order Pholidota: pangolins or scaly
anteaters (Africa, South Asia)
• Order Carnivora: carnivores (cosmopolitan)

For more details on this topic, see Evolution of mammals.
Mammaliaformes
Adelobasileus
void
Sinocodon
void
Morganucodon
void
Docodonta
void
––
Hadrocodium
––Mammalia
Synapsida, the group which contains mammals and their extinct relatives, originated during the
Pennsylvanian epoch, when they split from the lineage that led to reptiles and birds. Non-mammalian
synapsids were once called "mammal-like reptiles", although they are usually no longer considered reptiles.
Mammals evolved from non-mammalian synapsids during the Early Jurassic.
[edit] Evolution
The original synapsid skull structure contains one temporal opening behind the orbitals, in a fairly low
position on the skull (lower right in this image). This might have assisted in the containing the jaw muscles of
these organisms that could have increased their biting strength.
The first fully terrestrial vertebrates were amniotes. Like amphibians, they have lungs and limbs.
Amniotes' eggs, however, have internal membranes which allow the developing embryo to breathe but keep
water in. Hence amniotes can lay eggs on dry land, while amphibians generally need to lay their eggs in
water.
The first amniotes apparently arose in the late Carboniferous. They descended from earlier
tetrapods,[4] which lived on land already inhabited by insects, and other invertebrates, and by ferns, mosses,
and other plants. Within a few million years two important amniote lineages became distinct: the synapsids,
which include mammals; and the sauropsids, which include lizards, snakes, crocodilians, dinosaurs and
birds.[5] Synapsids have a single hole (temporal fenestra) low on each side of the skull.
One synapsid group, the pelycosaurs, were the most common land vertebrates of the early Permian
and included the largest land animals of the time.[6]
Therapsids descended from pelycosaurs in the middle Permian, about 260M years ago, and took
over their position as the dominant land vertebrates. They differ from pelycosaurs in several features of the
skull and jaws, including: larger temporal fenestrae and incisors which are equal in size.[7] The therapsids
went through a series of stages, beginning with animals which were very like their pelycosaur ancestors and
ending with the Triassic cynodonts, some of which could easily be mistaken for mammals. Those stages
were characterized by:
• gradual development of a bony secondary palate.[8]
• progress towards an erect limb posture, which would increase the animals' stamina by
avoiding Carrier's constraint. But this process was slow and erratic – for example: all herbivorous
non-mammaliaform therapsids retained sprawling limbs (some late forms may have had semi-erect
hind limbs); Permian carnivorous therapsids had sprawling forelimbs, and some late Permian ones
also had semi-sprawling hindlimbs. In fact modern monotremes still have semi-sprawling limbs.
• the dentary gradually becoming the main bone of the lower jaw; and in the Triassic, progress
towards the fully mammalian jaw (the lower consisting only of the dentary) and middle ear (which is
constructed by the bones that were previously used to construct the jaws of Reptiles)
• there is possible evidence of hair in Triassic therapsids, but none for Permian therapsids.
• some scientists have argued that some Triassic therapsids show signs of lactation.
The Permian–Triassic extinction event ended the dominance of the therapsids, and in the Early
Triassic all the medium to large land animal niches were taken over by early archosaurs, which were the
ancestors of crocodilians, pterosaurs, dinosaurs and birds. After this "Triassic Takeover" the cynodonts and
their descendants could only survive as small, mainly nocturnal insectivores.[9]
The first true mammals appeared in the Late Triassic (ca. 200 million years ago), over 70 million
years after the first therapsids and approximately 30 million years after the first mammaliaformes.
Hadrocodium appears to be in the middle of the transition to true mammal status — it had a mammalian jaw
joint (formed by the dentary and squamosal bones), but there is some debate about whether its middle ear
was fully mammalian.[10] The majority of the mammal species that existed in the Mesozoic Era were
characterized by Multituberculates.
The earliest known monotreme is Teinolophos, which lived about 123M years ago in Australia.
Monotremes have some features which may be inherited from the original amniotes:
• they use the same orifice to urinate, defecate and reproduce ("monotreme" means "one
hole") – as lizards and birds also do.
• they lay eggs which are leathery and uncalcified, like those of lizards, turtles and
crocodilians.
Unlike other mammals, female monotremes do not have nipples and feed their young by "sweating"
milk from patches on their bellies.
The oldest known marsupial is Sinodelphys, found in 125M-year old early Cretaceous shale in
China's northeastern Liaoning Province. The fossil is nearly complete and includes tufts of fur and imprints of
soft tissues.[11]
Reconstruction based on Megalonyx jeffersonii, Iowa Museum of Natural History, University of Iowa.
The living Eutheria ("true beasts") are all placentals. But the earliest known eutherian, Eomaia, found
in China and dated to 125M years ago, obtained some features which are more like those of marsupials,
which suggested it was perhaps a transitional fossil that eventually give rise to the placental lineage (the
surviving metatherians):[12]
• Epipubic bones extending forwards from the pelvis, which are not found in any modern
placental, but are found in marsupials, monotremes and mammaliformes such as multituberculates.
In other words, they appear to be an ancestral feature which subsequently disappeared in the
placental lineage. These epipubic bones seem to function by stiffening the muscles of these animals
during locomotion, reducing the amount of space being presented, which placentals require to
contain their fetus during gestation periods.
• A narrow pelvic outlet, which indicates that the young were very small at birth and therefore
pregnancy was short, as in modern marsupials. This suggests that the placenta was a later
development.
It is not certain when true placental mammals evolved – the earliest undisputed fossils of placentals
come from the early Paleocene, after the extinction of the dinosaurs.[13]
Mammals and near-mammals expanded out of their nocturnal insectivore niche from the mid
Jurassic onwards – for example Castorocauda had adaptations for swimming, digging and catching fish.[14]
The traditional view is that: mammals only took over the medium- to large-sized ecological niches in
the Cenozoic, after the extinction of the dinosaurs; but then they diversified very quickly; for example the
earliest known bat dates from about 50M years ago, only 15M years after the extinction of the dinosaurs.[15]
On the other hand recent molecular phylogenetic studies suggest that most placental orders diverged
about 100M to 85M years ago, but that modern families first appeared in the late Eocene and early
Miocene[16] But paleontologists object that no placental fossils have been found from before the end of the
Cretaceous.[13]
During the Cenozoic several groups of mammals appeared which were much larger than their
nearest modern equivalents – but none was even close to the size of the largest dinosaurs with similar
feeding habits.
[edit] Earliest appearances of features

Hadrocodium, whose fossils date from the early Jurassic (approx. 195 million years ago), provides
the first clear evidence of fully mammalian jaw joints.
It has been suggested that the original function of lactation (milk production) was to keep eggs moist.
Much of the argument is based on monotremes (egg-laying mammals):[17][18][19]
The earliest clear evidence of hair or fur is in fossils of Castorocauda, from 164M years ago in the
mid Jurassic. From 1955 onwards some scientists have interpreted the foramina (passages) in the maxillae
(upper jaws) and premaxillae (small bones in front of the maxillae) of cynodonts as channels which supplied
blood vessels and nerves to vibrissae (whiskers), and suggested that this was evidence of hair or fur.[20][21]
But foramina do not necessarily show that an animal had vibrissae – for example the modern lizard
Tupinambis has foramina which are almost identical to those found in the non-mammalian cynodont
Thrinaxodon.[22][23]
American Lion was one of the abundant Pleistocene megafauna, a wide variety of very large
mammals that lived during the Pleistocene and went extinct about 10,000 years ago.[24]
The evolution of erect limbs in mammals is incomplete — living and fossil monotremes have sprawling
limbs. In fact some scientists think that the parasagittal (non-sprawling) limb posture is a synapomorphy
(distinguishing characteristic) of the Boreosphenida, a group which contains the Theria and therefore
includes the last common ancestor of modern marsupial and placentals – and therefore that all earlier
mammals had sprawling limbs.[25] Sinodelphys (the earliest known marsupial) and Eomaia (the earliest
known eutherian) lived about 125M years ago, so erect limbs must have evolved before then.
It is currently very difficult to be confident when endothermy first appeared in the evolution of
mammals. Modern monotremes have a lower body temperature and more variable metabolic rate than
marsupials and placentals.[26] So the main question is when a monotreme-like metabolism evolved in
mammals. The evidence found so far suggests Triassic cynodonts may have had fairly high metabolic rates,
but is not conclusive. In particular it is difficult to see how small animals can maintain a high and stable body
temperature without fur.
[edit] Anatomy and morphology
[edit] Skeletal system
The majority of mammals have seven cervical vertebrae (bones in the neck); this includes bats,
giraffes, whales, and humans. The few exceptions include the manatee and the two-toed sloth, which have
only six cervical vertebrae, and the three-toed sloth with nine cervical vertebrae.[27]
[edit] Respiratory system

The lungs of mammals have a spongy texture and are honeycombed with epithelium having a much
larger surface area in total than the outer surface area of the lung itself. The lungs of humans are typical of
this type of lung.
Breathing is largely driven by the muscular diaphragm which divides the thorax from the abdominal
cavity, forming a dome with its convexity towards the thorax. Contraction of the diaphragm flattens the dome
increasing the volume of the cavity in which the lung is enclosed. Air enters through the oral and nasal
cavities; it flows through the larynx, trachea and bronchi and expands the alveoli. Relaxation of the
diaphragm has the opposite effect, passively recoiling during normal breathing. During exercise, the
abdominal wall contracts, increasing visceral pressure on the diaphragm, thus forcing the air out more quickly
and forcefully. The rib cage itself also is able to expand and contract the thoracic cavity to some degree,
through the action of other respiratory and accessory respiratory muscles. As a result, air is sucked into or
expelled out of the lungs, always moving down its pressure gradient. This type of lung is known as a bellows
lung as it resembles a blacksmith's bellows.
[edit] Nervous system
All mammalian brains possess a neocortex, a brain region that is unique to mammals. Placental
mammals have a corpus callosum unlike monotremes and marsupials. The size and number of cortical areas
(Brodmann's areas) is least in monotremes (about 8-10) and most in placentals (up to 50).
[edit] Integumentary system

The integumentary system is made up of three layers: the outermost epidermis, the dermis, and the
hypodermis.
The epidermis is typically ten to thirty cells thick, its main function being to provide a waterproof layer.
Its outermost cells are constantly lost; its bottommost cells are constantly dividing and pushing upward. The
middle layer, the dermis, is fifteen to forty times thicker than the epidermis. The dermis is made up of many
components such as bony structures and blood vessels. The hypodermis is made up of adipose tissue. Its
job is to store lipids, and to provide cushioning and insulation. The thickness of this layer varies widely from
species to species.
Although mammals and other animals have cilia that superficially may resemble it, no other animals
except mammals have hair. It is a definitive characteristic of the class. Some mammals have very little, but
nonetheless, careful examination reveals the characteristic, often in obscure parts of their bodies. None are
known to have hair that naturally is blue or green in color[ citation needed] although some cetaceans, along
with the mandrills appear to have shades of blue skin. Many mammals are indicated as having blue hair or
fur, but in all known cases, it has been found to be a shade of gray. The two-toed sloth and the polar bear
may seem to have green fur, but this color is caused by algae growths.
[edit] Reproductive system
Goat kids will stay with their mother until they are weaned
Most mammals give birth to live young (vivipary), but a few, namely the monotremes, lay eggs. The
platypus and the echidna present a particular sex determination system that is different from other
vertebrates.[28]
Certain glands of mammals known as mammary glands are specialized to produce milk, a liquid
used by newborns as their primary source of nutrition. The monotremes branched early from other mammals
and do not have the nipples seen in most mammals, but they do have mammary glands.
Viviparous mammals are classified into the subclass Theria and are divided into two infraclasses:
Metatheria (of which only the Marsupialia survive), and Eutheria. Marsupialia, or marsupials, have short
gestation periods and give birth to undeveloped young which are contained within a pouch-like sac
(marsupium) located in front of the mothers' abdomen. Eutherians, commonly known as placentals, are
mammals that give birth to complete and fully developed young. This is usually characterized by long
gestation periods. The majority of mammal species are classified as eutherians.
[edit] Physiology
[edit] Endothermy
Nearly all mammals are endothermic ("warm-blooded"). Most mammals also have hair to help keep
them warm. Like birds, mammals can forage or hunt in cold weather and climates where non-avian reptiles
and large insects cannot.
Endothermy requires plenty of food energy, so pound for pound mammals eat more food than most
reptiles. Small insectivorous mammals eat prodigious amounts for their size.
A rare exception, the naked mole rat produces little metabolic heat, so it is considered an operational
poikilotherm . Birds are also endothermic, so endothermy is not a defining mammalian feature.
[edit] Intelligence
In intelligent mammals, such as primates, the cerebrum is larger relative to the rest of the brain.
Intelligence itself is not easy to define, but indications of intelligence include the ability to learn, matched with
behavioral flexibility. Rats, for example, are considered to be highly intelligent as they can learn and perform
new tasks, an ability that may be important when they first colonize a fresh habitat. In some mammals, food
gathering appears to be related to intelligence: a deer feeding on plants has a brain smaller than a cat, which
must think to outwit its prey.[29]
[edit] Social structure
This section is empty. You can help by adding to it.
[edit] Locomotion
See also: Animal locomotion
Mammals evolved from four-legged ancestors. They use their limbs to walk, climb, swim, and fly.
Some land mammals have toes that produce claws and hooves for climbing and running. Aquatic mammals
such as whales and dolphins have flippers which evolved from legs.
[edit] Terrestrial
See also: Terrestrial locomotion
The fastest land animal, the cheetah can reach 120 kmh (75 mph)
[edit] Arboreal
See also: Arboreal locomotion
Sloths travel slowly along branches rather than swinging energetically like monkeys.
[edit] Aquatic
Buoyed by their aquatic environment, whales have evolved into the largest mammals and indeed the
largest animals ever.
[edit] Aerial
Townsend's Big-eared Bat, Corynorhinus townsendii

See also: Aerial locomotion
[edit] Feeding
To maintain a high constant body temperature is energy expensive – mammals therefore need a
nutritious and plentiful diet. While the earliest mammals were probably predators, different species have
since adapted to meet their dietary requirements in a variety of ways. Some eat other animals – this is a
carnivorous diet (and includes insectivorous diets). Other mammals, called herbivores, eat plants. A
herbivorous diet includes sub-types such as fruit-eating and grass-eating. An omnivore eats both prey and
plants. Carnivorous mammals have a simple digestive tract, because the proteins, lipids, and minerals found
in meat require little in the way of specialized digestion. Plants, on the other hand, contain complex
carbohydrates, such as cellulose. The digestive tract of an herbivore is therefore host to bacteria that ferment
these substances, and make them available for digestion. The bacteria are either housed in the multi-
chambered stomach or in a large cecum. The size of an animal is also a factor in determining diet type. Since
small mammals have a high ratio of heat-losing surface area to heat-generating volume, they tend to have
high-energy requirements and a high metabolic rate. Mammals that weigh less than about 18 oz (500 g) are
mostly insectivorous because they cannot tolerate the slow, complex digestive process of a herbivore. Larger
animals on the other hand generate more heat and less of this heat is lost. They can therefore tolerate either
a slower collection process (those that prey on larger vertebrates) or a slower digestive process (herbivores).
Furthermore, mammals that weigh more than 18 oz (500 g) usually cannot collect enough insects during their
waking hours to sustain themselves. The only large insectivorous mammals are those that feed on huge
colonies of insects (ants or termites).[29]
Specializations in herbivory include: Granivory "seed eating", folivory "leaf eating", fruivory "fruit
eating", nectivory "nectar eating", gumivory "gum eating", and mycophagy "fungus eating".
[edit] See also

Mammals portal
• List of African mammals

• List of extinct mammals
• List of Indian mammals
• List of mammalogists
• List of mammals
• Lists of mammals by region
• List of prehistoric mammals
• Mammal classification
• Mammals discovered in the 2000s
• Prehistoric mammals
[edit] References
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Baltimore: Johns Hopkins University Press, 2 vols. (2142 pp.). ISBN 978-0-8018-8221-0.
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2. ^ Rose, Kenneth D. (2006). The beginning of the age of mammals. Baltimore: Johns Hopkins
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3. ^ McKenna, Malcolm C.; Bell, Susan Groag. Classification of Mammals. Columbia University
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mammals-dinos.html. Retrieved 2009-03-08.
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Grenyer, Richard; Price, Samantha A.; Vos, Rutger A. et al. (2007). "The delayed rise of present-day
mammals". Nature 446 (446): 507–511. doi:10.1038/nature05634. PMID 17392779.
http://scienceblogs.com/pharyngula/2007/03/dont_blame_the_dinosaurs.php.
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eggs=Journal of Mammary Gland Biology and Neoplasia. 7. pp. 253–266.
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39.
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&q&f=false
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Ferguson-Smith, M.A., and Graves, J.A.M. (December 2007). Sex determination in platypus and
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1185-3.
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• Vaughan, Terry A., James M. Ryan, and Nicholas J. Capzaplewski. 2000. Mammalogy:
Fourth Edition. Saunders College Publishing, 565 pp. ISBN 0-03-025034-X (Brooks Cole, 1999)
• Jan Ole Kriegs, Gennady Churakov, Martin Kiefmann, Ursula Jordan, Juergen Brosius,
Juergen Schmitz. (2006) Retroposed Elements as Archives for the Evolutionary History of Placental
Mammals. PLoS Biol 4(4): e91."PLoS Biology – Retroposed Elements as Archives for the
Evolutionary History of Placental Mammals". Biology.plosjournals.org.
doi:10.1371/journal.pbio.0040091. http://biology.plosjournals.org/perlserv/?request=get-
document&doi=10.1371/journal.pbio.0040091. Retrieved 2009-03-08.
• David MacDonald, Sasha Norris. 2006. The Encyclopedia of Mammals, 3rd edition. Printed
in China, 930 pp. ISBN 0-681-45659-0.

Find more about Mammal on Wikipedia's sister projects:

Wikispecies has information related to: Mammalia
The Wikibook Dichotomous Key has a page on the topic of

Mammalia
External identifiers for Mammalia
NCBI Taxonomy 40674

ITIS 179913
Encyclopedia of Life 1642
uBio NameBank 2478620
Also found in: Wikispecies, Arctos

• BBC Wildlife Finder – video clips from the BBC's natural history archive
• GlobalTwitcher.com – All species in the world with distribution maps and images
• Paleocene Mammals, a site covering the rise of the mammals, paleocene-mammals.de
• Evolution of Mammals, a brief introduction to early mammals, enchantedlearning.com
• Tree of Life poster – Shows mammals' evolutionary relation to other organisms,
tellapallet.com
• The Evolution of Mesozoic Mammals, a Rough Sketch , an informal introduction,
home.arcor.de
• Carnegie Museum of Natural History, some discoveries of early mammal fossils,
carnegiemnh.org
• High-Resolution Images of various Mammalian Brains, brainmaps.org
• Mammal Species, collection of information sheets about various mammal species,
learnanimals.com
• Summary of molecular support for Epitheria , biology.plosjournals.org
• Mikko's Phylogeny Archive, fmnh.helsinki.fi
• European Mammal Atlas EMMA from Societas Europaea Mammalogica, european-
mammals.org
• Marine Mammals of the World—An overview of all marine mammals, including descriptions,
multimedia and a key, eti.uva.nl
• Mammalogy.org The American Society of Mammalogists was established in 1919 for the
purpose of promoting the study of mammals, and this website includes a mammal image library
[hide]v · d · eExtant Chordata classes by subphylum
Kingdom Animalia · Subkingdom Eumetazoa · (unranked) Bilateria · Superphylum Deuterostomia
Urochordata Ascidiacea (Ascidians) · Thaliacea · Appendicularia (Larvaceans) ·

(Tunicates) Sorberacea
Cephalochordata
(Lancelets) Leptocardii
Myxini (Hagfish) · Hyperoartia (Lampreys) · Chondrichthyes

Craniata (Cartilaginous fish) · Actinopterygii (Ray-finned fish) · Sarcopterygii (Lobe-
finned fish) · Amphibia (Amphibians) · Sauropsida (Reptiles) · Aves (Birds) ·
Mammalia (Mammals)
[hide]v · d · eExtant mammal orders by infraclass
Kingdom Animalia · Phylum Chordata · Subphylum Vertebrata · (unranked) Amniota
Australosphenida Monotremata (Platypus and echidnas)
Paucituberculata (Shrew opossums) ·

Ameridelphia
Didelphimorphia (Opossums)
Metatheria
(Marsupial inclusive) Microbiotheria (Monito del Monte) ·
Notoryctemorphia (Marsupial moles) · Dasyuromorphia
Australidelphia
(Quolls and dunnarts) · Peramelemorphia (Bilbies and
bandicoots) · Diprotodontia (Kangaroos and relatives)
Eutheria Cingulata (Armadillos) · Pilosa (Anteaters and

(Placental inclusive) Xenarthra
sloths)
Afrosoricida (Tenrecs and golden moles) ·

Macroscelidea (Elephant shrews) · Tubulidentata
Afrotheria
(Aardvark) · Hyracoidea (Hyraxes) · Proboscidea
(Elephants) · Sirenia (Dugongs and manatees)
Soricomorpha (Shrews and moles) ·
Erinaceomorpha (Hedgehogs and relatives) ·
Chiroptera (Bats) · Pholidota (Pangolins) · Carnivora ·
Laurasiatheria
Perissodactyla (Odd-toed ungulates) · Artiodactyla
(Even-toed ungulates) · Cetacea (Whales and
dolphins)
Rodentia (Rodents) · Lagomorpha (Rabbits

Euarchontoglires and relatives) · Scandentia (Treeshrews) ·
Dermoptera (Colugos) · Primates
Retrieved from "http://en.wikipedia.org/wiki/Mammal"
Categories: Mammals
W000
Evolution of Mollusca

Diagram of primitive mollusc viewed from left side. Hypothetical mollusc ancestor. From
Encyclopædia Britannica Eleventh Edition.
The current understanding of molluscan phylogeny is shown below.
Horizontal cladogram (source:[1]):
Mollusca
Aplacophora
Solenogastres
Caudofoveata
Testaria
Polyplacophora
Palaeoloricata
Neoloricata
Conchifera
Monoplacophora
(unnamed)
Cyrtosoma
Gastropoda
Cephalopoda
Diasoma
Rostroconchia
(unnamed)
Scaphopoda
Bivalvia
Basal Mollusca phylogeny.
Early organisms which have been compared to molluscs include Kimberella and Odontogriphus.
[edit] References
1. ^ Zoology 250 Phylogenetic Trees (2006)
http://www.biology.ualberta.ca/courses.hp/zool250/Clades/clade08-Mollusca.htm
• From sea to land and beyond – New insights into the evolution of euthyneuran Gastropoda
(Mollusca)
Processes of
Population genetic
Evolutionary
developmental
concepts

Modes of

This mollusc-related article is a stub. You can help Wikipedia by expanding it.v · d
·e
Retrieved from "http://en.wikipedia.org/wiki/Evolution_of_Mollusca"
Categories: Evolution by taxon | Molluscs | Mollusc stubs

W000
Cambrian explosion
The Cambrian explosion or Cambrian radiation was the relatively rapid appearance, over a period of
many million years, of most major Phyla around 530 million years ago, as found in the fossil record.[1][2] This
was accompanied by a major diversification of other organisms, including animals, phytoplankton, and
calcimicrobes.[3] Before about 580 million years ago, most organisms were simple, composed of individual
cells occasionally organized into colonies. Over the following 70 or 80 million years the rate of evolution
accelerated by an order of magnitude (as defined in terms of the extinction and origination rate of species[4])
and the diversity of life began to resemble today’s.[5]
The Cambrian explosion has generated extensive scientific debate. The seemingly rapid appearance
of fossils in the “Primordial Strata” was noted as early as the mid 19th century,[6] and Charles Darwin saw it
as one of the main objections that could be made against his theory of evolution by natural selection.[7]
The long-running puzzlement about the appearance of the Cambrian fauna, seemingly abruptly and
from nowhere, centers on three key points: whether there really was a mass diversification of complex
organisms over a relatively short period of time during the early Cambrian; what might have caused such
rapid change; and what it would imply about the origin and evolution of animals. Interpretation is difficult due
to a limited supply of evidence, based mainly on an incomplete fossil record and chemical signatures left in
Cambrian rocks.
Key Cambrian explosion events

-580 —
–
-570 —
–
-560 —
–
-550 —
–
-540 —
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-530 —
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-520 —
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-510 —
–
-500 —
–
-490 —
N
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P
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Ediacaran
C
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Ordovician
Cryogenian
T
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S
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2
S
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F
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U
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M
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d
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L
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Nemakit-
Daldynian
Tommotian
Atdabanian
Botomian
Toyonian
Stage 5
Drumian
Guzhangian
Paibian
Stage 9
Stage 10
←
Orsten Fauna
←
Burgess Shale
←
Kaili biota
←
Archaeocyatha extinction
←
Emu Bay Shale
←
Chengjiang and Sirius Passet biotas
←
First Trilobites
←
SSF diversification, first brachiopods & archaeocyatha
←
First halkieriids, mollusсs, hyoliths SSF
←
Treptichnus pedum trace
←
First Treptichnus sp. trace
←
First Cloudina & Namacalathus mineral skeletal fossils
←
Claimed bilaterian trace fossils
←
First Ediacara biota members (Charnia antecedens, C. wardae, Thectardis)
←
Glaskiers glaciation
←
Doushantuo "embryos"
←
End Marinoan glaciation
-635 —
One possible timescale for events around the Cambrian/Precambrian boundary.
Stratigraphic scale of the ICS with Russian Lower Cambrian subdivision and Precambrian/Cambrian
boundary.
Contents
[hide]
• 1 History and significance
• 2 Types of evidence
• 2.1 Dating the Cambrian
• 2.2 Body fossils
• 2.3 Trace fossils
• 2.4 Geochemical observations
• 2.5 Phylogenetic techniques
• 3 Explanation of a few scientific terms
• 4 Precambrian life
• 4.1 Evidence of animals around 1 billion years ago
• 4.2 Fossils of the Doushantuo formation
• 4.3 Burrows
• 4.4 Ediacaran organisms
• 5 Ediacaran–Early Cambrian skeletalization
• 6 Cambrian life
• 6.1 Small shelly fauna
• 6.2 Early Cambrian trilobites and echinoderms
• 6.3 Burgess shale type faunas
• 6.4 Early Cambrian crustaceans
• 6.5 Early Ordovician radiation
[edit] History and significance
Geologists as long ago as Buckland (1784–1856) realised that a dramatic step-change in the fossil
record occurred around the base of what we now call the Cambrian.[6] Charles Darwin considered this
sudden appearance of many animal groups with few or no antecedents to be the greatest single objection to
his theory of evolution. He had even devoted a substantial chapter of The Origin of Species to solving this
problem.[7]
American palæontologist Charles Walcott proposed that an interval of time, the “Lipalian”, was not
represented in the fossil record or did not preserve fossils, and that the ancestors of the Cambrian animals
evolved during this time.[8]
More recently it was discovered that the history of life on earth goes back at least 3,450 million years:
[9] rocks of that age at Warrawoona in Australia contain fossils of stromatolites, stubby pillars that are formed
by colonies of micro-organisms. Fossils (Grypania) of more complex eukaryotic cells, from which all animals,
plants and fungi are built, have been found in rocks from 1,400 million years ago, in China and Montana.
Rocks dating from 565 to 543 million years ago contain fossils of the Ediacara biota, organisms so large that
they must have been multi-celled, but very unlike any modern organism.[10] P. E. Cloud argued in 1948 that
there was a period of "eruptive" evolution in the Early Cambrian,[11] but as recently as the 1970s there was
no sign of how the relatively modern-looking organisms of the Middle and Late Cambrian arose.[10]
Opabinia made the largest single contribution to modern interest in the Cambrian explosion.
The intense modern interest in this "Cambrian explosion" was sparked by the work of Harry B.
Whittington and colleagues, who in the 1970s re-analysed many fossils from the Burgess Shale (see below)
and concluded that several were complex but different from any living animals.[12][13] The most common
organism, Marrella, was clearly an arthropod, but not a member of any known arthropod class. Organisms
such as the five-eyed Opabinia and spiny slug-like Wiwaxia were so different from anything else known that
Whittington's team assumed they must represent different phyla, only distantly related to anything known
today. Stephen Jay Gould’s popular 1989 account of this work, Wonderful Life,[14] brought the matter into
the public eye and raised questions about what the explosion represented. While differing significantly in
details, both Whittington and Gould proposed that all modern animal phyla had appeared rather suddenly.
This view was influenced by the theory of punctuated equilibrium, which Eldredge and Gould developed in
the early 1970s and which views evolution as long intervals of near-stasis "punctuated" by short periods of
rapid change.[15]
Other analyses, some more recent and some dating back to the 1970s, argue that complex animals
similar to modern types evolved well before the start of the Cambrian.[16][17][18] There has also been
intense debate whether there was a genuine "explosion" of modern forms in the Cambrian and, to the extent
that there was, how it happened and why it happened then.[19]
[edit] Types of evidence

Deducing the events of half a billion years ago is difficult, as evidence comes exclusively from
biological and chemical signatures in rocks and very sparse fossils.
[edit] Dating the Cambrian

Accurate absolute radiometric dates for much of the Cambrian, obtained by detailed analysis of
radioactive elements contained within rocks, have only rather recently become available, and for only a few
regions.[20]
Relative dating (A was before B) is often sufficient for studying processes of evolution, but this too
has been difficult, because of the problems involved in matching up rocks of the same age across different
continents.[21]
Therefore dates or descriptions of sequences of events should be regarded with some caution until
better data become available.
[edit] Body fossils
Fossils of organisms' bodies are usually the most informative type of evidence. Fossilisation is a rare
event, and most fossils are destroyed by erosion or metamorphism before they can be observed. Hence the
fossil record is very incomplete, increasingly so, further back in time. Despite this, they are often adequate to
illustrate the broader patterns of life's history.[22] There are also biases in the fossil record: different
environments are more favourable to the preservation of different types of organism or parts of organisms.
[23] Further, only the parts of organisms that were already mineralised are usually preserved, such as the
shells of molluscs. Since most animal species are soft-bodied, they decay before they can become fossilised.
As a result, although there are 30-plus phyla of living animals, two-thirds have never been found as fossils.
[10]
This Marrella specimen illustrates how clear and detailed the fossils from the Burgess Shale
lagerstätte are.
The Cambrian fossil record includes an unusually high number of lagerstätten, which preserve soft
tissues. These allow palæontologists to examine the internal anatomy of animals which in other sediments
are only represented by shells, spines, claws, etc. – if they are preserved at all. The most significant
Cambrian lagerstätten are the early Cambrian Maotianshan shale beds of Chengjiang (Yunnan, China) and
Sirius Passet (Greenland);[24] the middle Cambrian Burgess Shale (British Columbia, Canada);[25] and the
late Cambrian Orsten (Sweden) fossil beds.
While lagerstätten preserve far more than the conventional fossil record, they are far from complete.
Because lagerstätten are restricted to a narrow range of environments (where soft-bodied organisms can be
preserved very quickly, e.g. by mudslides), most animals are probably not represented; further, the
exceptional conditions that create lagerstätten probably do not represent normal living conditions.[26] In
addition, the known Cambrian lagerstätten are rare and difficult to date, while Precambrian lagerstätten have
yet to be studied in detail.
The sparseness of the fossil record means that organisms usually exist long before they are found in
the fossil record – this is known as the Signor-Lipps effect.[27]
[edit] Trace fossils
Rusophycus and other trace fossils from the Gog Formation, Middle Cambrian, Lake Louise, Alberta,
Canada.
Trace fossils consist mainly of tracks and burrows, but also include coprolites (fossil feces) and
marks left by feeding.[28][29] Trace fossils are particularly significant because they represent a data source
that is not limited to animals with easily-fossilized hard parts, and which reflects organisms' behaviour. Also
many traces date from significantly earlier than the body fossils of animals that are thought to have been
capable of making them.[30] Whilst exact assignment of trace fossils to their makers is generally impossible,
traces may for example provide the earliest physical evidence of the appearance of moderately complex
animals (comparable to earthworms).[29]
[edit] Geochemical observations
Main article: Early Cambrian geochemical fluctuations
Several chemical markers indicate a drastic change in the environment around the start of the
Cambrian. The markers are consistent with a mass extinction,[31][32] or with a massive warming resulting
from the release of methane ice. [19] Such changes may reflect a cause of the Cambrian explosion, although
they may also have resulted from an increased level of biological activity – a possible result of the explosion.
[19] Despite these uncertainties, the geochemical evidence helps by making scientists focus on theories that
are consistent with at least one of the likely environmental changes.
[edit] Phylogenetic techniques

Cladistics is a technique for working out the “family tree” of a set of organisms. It works by the logic
that, if groups B and C have more similarities to each other than either has to group A, then B and C are
more closely related to each other than either is to A. Characteristics which are compared may be
anatomical, such as the presence of a notochord, or molecular, by comparing sequences of DNA or protein.
The result of a successful analysis is a hierarchy of clades – groups whose members are believed to share a
common ancestor. The cladistic technique is sometimes fallible, as some features, such as wings or camera
eyes, evolved more than once, convergently – this must be taken into account in analyses.
From the relationships, it may be possible to constrain the date that lineages first appeared. For
instance, if fossils of B or C date to X million years ago and the calculated "family tree" says A was an
ancestor of B and C, then A must have evolved more than X million years ago.
It is also possible to estimate how long ago two living clades diverged – i.e. approximately how long
ago their last common ancestor must have lived – by assuming that DNA mutations accumulate at a constant
rate. These "molecular clocks", however, are fallible, and provide only a very approximate timing: they are
not sufficiently precise and reliable for estimating when the groups that feature in the Cambrian explosion first
evolved,[33] and estimates produced by different techniques vary by a factor of two.[34] However, the clocks
can give an indication of branching rate, and when combined with the constraints of the fossil record, recent
clocks suggest a sustained period of diversification through the Ediacaran and Cambrian.[35]
[edit] Explanation of a few scientific terms

A phylum is the highest level in the Linnean system for
classifying organisms. Phyla can be thought of as groupings of animals
based on general body plan.[37] Despite the seemingly different external
appearances of organisms, they are classified into phyla based on their
internal and developmental organizations.[38] For example, despite their
obvious differences, spiders and barnacles both belong to the phylum
Arthropoda; but earthworms and tapeworms, although similar in shape,
belong to different phyla.
A phylum is not a fundamental division of nature, such as the
difference between electrons and protons. It is simply a very high-level = Crown group
grouping in a classification system created to describe all currently living = Stem group
organisms. This system is imperfect, even for modern animals: different
books quote different numbers of phyla, mainly because they disagree
about the classification of a huge number of worm-like species. As it is
based on living organisms, it accommodates extinct organisms poorly, if
at all.[10][39]
Stem groups.[36]
The concept of stem groups was introduced to cover
evolutionary "aunts" and "cousins" of living groups. A crown
group is a group of closely-related living animals plus their last
common ancestor plus all its descendants. A stem group is a set
of offshoots from the lineage at a point earlier than the last
common ancestor of the crown group; it is a relative concept, for
example tardigrades are living animals which form a crown group
in their own right, but Budd (1996) regarded them also as being a
stem group relative to the arthropods.[36][40]
Triploblastic means consisting of 3 layers, which are
formed in the embryo, quite early in the animal's development
from a single-celled egg to a larva or juvenile form. The innermost
layer forms the digestive tract (gut); the outermost forms skin;
and the middle one forms muscles and all the internal organs
except the digestive system. Most types of living animal are
triploblastic – the best-known exceptions are Porifera (sponges)
and Cnidaria (jellyfish, sea anemones, etc.).
The bilaterians are animals which have right and left
sides at some point in their life history. This implies that they have
top and bottom surfaces and, importantly, distinct front and back
ends. All known bilaterian animals are triploblastic, and all known
triploblastic animals are bilaterian. Living Echinoderms (sea
stars, sea urchins, sea cucumbers, etc.) look radially symmetrical
(like wheels) rather than bilaterian, but their larvae exhibit
A coelomate animal is basically a

set of concentric tubes, with a gap
between the gut and the outer tubes.
bilateral symmetry and some of the earliest echinoderms may have been bilaterally symmetrical.[41] Porifera
and Cnidaria are radially symmetrical, non-bilaterian and non-triploblastic.
Coelomate means having a body cavity (coelom) which contains the internal organs. Most of the
phyla featured in the debate about the Cambrian explosion are coelomates: arthropods, annelid worms,
molluscs, echinoderms and chordates – the non-coelomate priapulids are an important exception. All known
coelomate animals are triploblastic bilaterians, but some triploblastic bilaterian animals do not have a
coelom – for example flatworms, whose organs are surrounded by unspecialized tissues.
[edit] Precambrian life

Our understanding of the Cambrian explosion relies upon knowing what was there beforehand – did
the event herald the sudden appearance of a wide range of animals and behaviours, or did such things exist
beforehand?
[edit] Evidence of animals around 1 billion years ago

For further information, see Acritarch and Stromatolite
Stromatolites (Pika Formation, Middle Cambrian) near Helen Lake, Banff National Park, Canada.
Modern stromatolites in Hamelin Pool Marine Nature Reserve, Western Australia.
Changes in the abundance and diversity of some types of fossil have been interpreted as evidence
for "attacks" by animals or other organisms. Stromatolites, stubby pillars built by colonies of microorganisms,
are a major constituent of the fossil record from about 2,700 million years ago, but their abundance and
diversity declined steeply after about 1,250 million years ago. This decline has been attributed to disruption
by grazing and burrowing animals.[16][17][42]
Precambrian marine diversity was dominated by small fossils known as acritarchs. This term
describes almost any small organic walled fossil–from the egg cases of small metazoans to resting cysts of
many different kinds of green algae. After appearing around 2,000 million years ago, acritarchs underwent a
boom around 1,000 million years ago, increasing in abundance, diversity, size, complexity of shape and
especially size and number of spines. Their increasingly spiny forms in the last 1 billion years may indicate
an increased need for defence against predation. Other groups of small organisms from the Neoproterozoic
era also show signs of anti-predator defenses.[42] A consideration of taxon longevity appears to support an
increase in predation pressure around this time,[43] However, in general, the rate of evolution in the
Precambrian was very slow, with many cyanobacterial species persisting unchanged for billions of years.[4]
If these predatory organisms really were metazoans, this means that Cambrian animals did not
appear "from no-where" at the base of the Cambrian; their predecessors had existed for hundreds of millions
of years.
[edit] Fossils of the Doushantuo formation

Main article: Doushantuo formation
The 580 million year old[44] Doushantuo formation harbours microscopic fossils which may
represent early bilaterians. Some have been described as animal embryos and eggs, although some of these
may represent the remains of giant bacteria.[45] Another fossil, Vernanimalcula, has been interpreted as a
coelomate bilaterian,[46] but may simply be an infilled bubble.[47]
These fossils form the earliest hard-and-fast evidence of animals, as opposed to other predators.[45]
[48]
[edit] Burrows
Main article: Cambrian substrate revolution
An Ediacaran trace fossil, made when an organism burrowed below a microbial mat.
The traces of organisms moving on and directly underneath the microbial mats that covered the
Ediacaran sea floor are preserved from the Ediacaran period, about 565 million years ago. They were
probably made by organisms resembling earthworms in shape, size, and how they moved. The burrow-
makers have never been found preserved, but because they would need a head and a tail, the burrowers
probably had bilateral symmetry – which would in all probability make them bilaterian animals.[49] They fed
above the sediment surface, but were forced to burrow to avoid predators.[50]
Around the start of the Cambrian (about 542 million years ago) many new types of traces first
appear, including well-known vertical burrows such as Diplocraterion and Skolithos, and traces normally
attributed to arthropods, such as Cruziana and Rusophycus. The vertical burrows indicate that worm-like
animals acquired new behaviours, and possibly new physical capabilities. Some Cambrian trace fossils
indicate that their makers possessed hard exoskeletons, although they were not necessarily mineralised.[51]
Burrows provide firm evidence of complex organisms; they are also much more readily preserved
than body fossils, to the extent that the absence of trace fossils has been used to imply the genuine absence
of large, motile bottom-dwelling organisms.[ citation needed] They provide a further line of evidence to show
that the Cambrian explosion represents a real diversification, and is not a preservational artefact.[52]
Indeed, as burrowing became established, it allowed an explosion of its own, for as burrowers
disturbed the sea floor, they aerated it, mixing oxygen into the toxic muds. This made the bottom sediments
more hospitable, and allowed a wider range of organisms to inhabit them – creating new niches and the
scope for higher diversity.[52]
[edit] Ediacaran organisms
Dickinsonia costata, an Ediacaran organism of unknown affinity, with a quilted appearance.

Main articles: Ediacara biota, Cloudinid, Kimberella, and Spriggina
At the start of the Ediacaran period, much of the acritarch fauna, which had remained relatively
unchanged for hundreds of millions of years, became extinct, to be replaced with a range of new, larger
species which would prove far more ephemeral.[4] This radiation, the first in the fossil record,[4] is followed
soon after by an array of unfamiliar, large, fossils dubbed the Ediacara biota,[53] which flourished for 40
million years until the start of the Cambrian.[54] Most of this "Ediacara biota" were at least a few centimeters
long, significantly larger than any earlier fossils. The organisms form three distinct assemblages, increasing
in size and complexity as time progresses.[55]
Many of these organisms were quite unlike anything that appeared before or since, resembling discs,
mud-filled bags, or quilted mattresses – one palæontologist proposed that the strangest organisms should be
classified as a separate kingdom, Vendozoa.[56]
Fossil of Kimberella, a triploblastic bilaterian , and possibly a mollusc.

At least some may have been early forms of the phyla at the heart of the "Cambrian explosion"
debate, having been interpreted as early molluscs ( Kimberella),[18][57] echinoderms (Arkarua);[58] and
arthropods (Spriggina,[59] Parvancorina[60]). There is still debate about the classification of these
specimens, mainly because the diagnostic features which allow taxonomists to classify more recent
organisms, such as similarities to living organisms, are generally absent in the Ediacarans.[61] However
there seems little doubt that Kimberella was at least a triploblastic bilaterian animal.[61] These organisms are
central to the debate about how abrupt the Cambrian explosion was. If some were early members of the
animal phyla seen today, the "explosion" looks a lot less sudden than if all these organisms represent an
unrelated "experiment", and were replaced by the animal kingdom fairly soon thereafter (40M years is "soon"
by evolutionary and geological standards).
[edit] Ediacaran–Early Cambrian skeletalization

The first Ediacaran and lowest Cambrian (Nemakit-Daldynian) skeletal fossils represent tubes and
problematic sponge spicules. The oldest sponge spicules are monaxon siliceous, aged around 580 million
years ago, known from the Doushantou Formation in the China and from deposits of the same age in
Mongolia. In the late Ediacaran-lowest Cambrian, numerous tube dwellings of enigmatic organisms
appeared. It was organic-walled tubes (e.g. Saarina) and chitinous tubes of the sabelliditids (e.g. Sokoloviina,
Sabellidites, Paleolina) [62][63] which prospered up to the beginning of the Tommotian. The mineralized
tubes of Cloudina, Namacalathus, Sinotubulites and a dozen more of the other organisms from carbonate
rocks formed near the end of the Ediacaran period from 549 to 542 million years ago, as well as the triradially
symmetrical mineralized tubes of anabaritids (e.g. Anabarites, Cambrotubulus) from uppermost Ediacaran
and lower Cambrian.[64] It is interesting to notice that Ediacaran mineralized tubes are often found in
carbonates of the stromatolite reefs and thrombolites,[65][66] i.e. they could live in environment adverse for
the majority of animals.
Although they are as hard to classify as most other Ediacaran organisms, they are important in two
other ways. First, they are the earliest known calcifying organisms (organisms that built shells out of calcium
carbonate).[66][67][68] Secondly, these tubes are a device to rise over a substrate and competitors for
effective feeding and, to a lesser degree, they serve as armor for protection against predators and adverse
conditions of environment. Some Cloudina fossils show small holes in shells. It is possible that the holes are
evidence of boring by predators sufficiently advanced to penetrate shells.[69] A possible "evolutionary arms
race" between predators and prey is one of the theories that attempt to explain the Cambrian explosion.[42]
In the lowest Cambrian occurs the extinction of stromatolites. This allowed animals to begin
colonization of warm-water pools with carbonate sedimentation. At first it was anabaritids and Protohertzina
(the fossilized grasping spines of chaetognaths) fossils. Such mineral skeletons as shells, sclerites, thorns
and plates appeared in uppermost Nemakit-Daldynian; they were the earliest species of halkierids,
gastropods, hyoliths and other rare organisms. The beginning of the Tommotian has historically been
understood to mark an explosive increase of the number and variety of fossils of mollusks, hyoliths and
sponges, along with a rich complex of skeletal elements of unknown animals, the first archaeocyathids,
brachiopods, tommotiids and others.[70][71][72][73] This sudden increase is partially an artefact of missing
strata at the Tommotian type section, and most of this fauna in fact began to diversify in a series of pulses
through the Nemakit-Daldynian and into the Tommotian.[74]
Some animals may already have had sclerites, thorns and plates in the Ediacaran (e.g. Kimberella
had hard sclerites, probably of carbonate), but thin carbonate skeletons cannot be fossilized in siliciclastic
deposits.[75]
[edit] Cambrian life

[edit] Small shelly fauna
Main article: Small shelly fauna
Fossils known as “small shelly fauna” have been found in many parts on the world, and date from just
before the Cambrian to about 10 million years after the start of the Cambrian (the Nemakit-Daldynian and
Tommotian ages; see timeline). These are a very mixed collection of fossils: spines, sclerites (armor plates),
tubes, archeocyathids (sponge-like animals) and small shells very like those of brachiopods and snail-like
molluscs – but all tiny, mostly 1 to 2 mm long.[76]
While small, these fossils are far more common than complete fossils of the organisms that produced
them; crucially, they cover the window from the start of the Cambrian to the first lagerstatten: a period of time
that is otherwise lacking in fossils. Hence they supplement the conventional fossil record, and allow the fossil
ranges of many groups to be extended.
[edit] Early Cambrian trilobites and echinoderms
A fossilized trilobite, an ancient type of arthropod. This specimen, from the Burgess shale, preserves
"soft parts" – the antennae and legs.
The earliest Cambrian trilobite fossils are about 530 million years old, but the class was already quite
diverse and worldwide, suggesting that they had been around for quite some time.[77] It is important to
remember that the fossil record of trilobites begins from the time of appearance of trilobites with mineral
exoskeleton - not from the time of their origin.
The earliest generally-accepted echinoderm fossils appeared a little bit later, in the Late Atdabanian;
unlike modern echinoderms, these early Cambrian echinoderms were not all radially symmetrical.[78]
These provide firm data points for the "end" of the explosion, or at least indications that the crown
groups of modern phyla were represented.
[edit] Burgess shale type faunas

Main article: Burgess shale type preservation
The Burgess shale and similar lagerstatten preserve the soft parts of organisms, which provides a
wealth of data to aid in the classification of enigmatic fossils. It often preserved complete specimens of
organisms only otherwise known from dispersed parts, such as loose scales or isolated mouthparts. Further,
the majority of organisms and taxa in these horizons are entirely soft bodied – hence absent from the rest of
the fossil record.[79] Since a large part of the ecosystem is preserved, the ecology of the community can also
be tentatively reconstructed.[verification needed] However, the assemblages may represent a "museum": a
deep water ecosystem that is evolutionarily "behind" the rapidly diversifying faunas of shallower waters.[80]
Because the lagerstatten provide a mode and quality of preservation that's virtually absent outside of
the Cambrian, lots of organisms appear completely different to anything known from the conventional fossil
record. This led early workers in the field to attempt to shoehorn the organisms into extant phyla; the
shortcomings of this approach led them to erect a multitude of new phyla to accommodate all the oddballs. It
has since been realised that most oddballs diverged from lineages before they established the phyla we
know today[clarification needed] – slightly different designs, which were fated to perish rather than flourish
into phyla, as their cousin lineages did.
The preservational mode is rare in the preceding Ediacaran period, but those assemblages known
show no trace of animal life – perhaps implying a genuine absence of macroscopic metazoans.[81]
[edit] Early Cambrian crustaceans

For more details on this topic, see Orsten.
Crustaceans, one of the four great modern groups of arthropods, are very rare throughout the
Cambrian. Convincing crustaceans were once thought to be common in Burgess shale-type biotas, but none
of these individuals can be shown to fall into the crown group of "true crustaceans".[82] The Cambrian record
of crown group crustaceans comes from microfossils. The Swedish Orsten horizons contain later Cambrian
crustacea, but only organisms smaller than 2 mm are preserved. This restricts the data set to juveniles and
miniaturised adults.
A more informative data source is the organic microfossils of the Mount Cap formation, Canada. This
late Early Cambrian assemblage (510 to 515 million years ago) consists of microscopic fragments of
arthropods' cuticle, which is left behind when the rock is dissolved with a strong acid. The diversity of this
assemblage is similar to that of modern crustacean faunas. Most interestingly, analysis of fragments of
feeding machinery found in the formation shows that it was adapted to feed in a very precise and refined
fashion. This contrasts with most other early Cambrian arthropods, which fed messily by shovelling anything
they could get their feeding appendages on into their mouths. This sophisticated and specialised feeding
machinery belonged to a large (~30 cm)[83] organism, and would have provided great potential for
diversification: specialised feeding apparatus allows a number of different approaches to feeding and
development, and creates a number of different approaches to avoid being eaten[82]
[edit] Early Ordovician radiation

Main article: Ordovician radiation
After a mass extinction at the Cambrian-Ordovician boundary, another radiation occurred, which
established the taxa which would dominate the Palaeozoic.[84]
During this radiation, the total number of orders doubled, and families tripled,[84] increasing marine
diversity to levels typical of the Palaeozoic,[19] and disparity to levels approximately equivalent to today's.[5]
[edit] How real was the explosion?

The fossil record as Darwin knew it seemed to suggest that the major metazoan groups appeared in
a few million years of the early to mid-Cambrian, and even in the 1980s this still appeared to be the case.[13]
[14]
However, evidence of Precambrian metazoa is gradually accumulating. If the Ediacaran Kimberella
was a mollusc-like protostome (one of the two main groups of coelomates),[18][57] the protostome and
deuterostome lineages must have split significantly before 550 million years ago (deuterostomes are the
other main group of coelomates).[85] Even if it is not a protostome, it is widely accepted as a bilaterian.[61]
[85] Since fossils of rather modern-looking Cnidarians (jellyfish-like organisms) have been found in the
Doushantuo lagerstätte, the Cnidarian and bilaterian lineages must have diverged well over 580 million years
ago.[85]
Trace fossils[55] and predatory borings in Cloudina shells provide further evidence of Ediacaran
animals.[86] Some fossils from the Doushantuo formation have been interpreted as embryos and one
(Vernanimalcula) as a bilaterian coelomate, although these interpretations are not universally accepted.[46]
[47][87] Earlier still, predatory pressure has acted on stromatolites and acritarchs since around 1,250 million
years ago.[42]
The presence of Precambrian animals somewhat dampens the "bang" of the explosion: not only was
the appearance of animals gradual, but their evolutionary radiation ("diversification") may also not have been
as rapid as once thought. Indeed, statistical analysis shows that the Cambrian explosion was no faster than
any of the other radiations in animals' history.[4] However, it does seem that some innovations linked to the
explosion — such as resistant armour — only evolved once in the animal lineage; this makes a lengthy
Precambrian animal lineage harder to defend.[88] Further, the conventional view that all the phyla arose in
the Cambrian is flawed; whilst the phyla may have diversified in this time period, representatives of the
crown-groups of many phyla do not appear until much later in the Phanerozoic.[89] Further, the mineralized
phyla that form the basis of the fossil record may not be representative of other phyla, since most mineralized
phyla originated in a benthic setting. The fossil record is consistent with a Cambrian Explosion that was
limited to the benthos, with pelagic phyla evolving much later.[89]
There is little doubt that disparity – that is, the range of different organism "designs" or "ways of life" –
rose sharply in the early Cambrian.[5] However, recent research has overthrown the once-popular idea that
disparity was exceptionally high throughout the Cambrian, before subsequently decreasing.[90] In fact,
disparity remains relatively low throughout the Cambrian, with modern levels of disparity only attained after
the early Ordovician radiation.[5]
The diversity of many Cambrian assemblages is similar to today's.[91][82]
[edit] Possible causes of the “explosion”
Despite the evidence that moderately complex animals (triploblastic bilaterians) existed before and
possibly long before the start of the Cambrian, it seems that the pace of evolution was exceptionally fast in
the early Cambrian. Possible explanations for this fall into three broad categories: environmental,
developmental, and ecological changes. Any explanation must explain the timing and magnitude of the
explosion.
[edit] Changes in the environment
[edit] Increase in oxygen levels

Earth’s earliest atmosphere contained no free oxygen; the oxygen that animals breathe today, both in
the air and dissolved in water, is the product of billions of years of photosynthesis. As a general trend, the
concentration of oxygen in the atmosphere has risen gradually over about the last 2.5 billion years.[10]
Shortage of oxygen might well have prevented the rise of large, complex animals. The amount of
oxygen an animal can absorb is largely determined by the area of its oxygen-absorbing surfaces (lungs and
gills in the most complex animals; the skin in less complex ones); but the amount needed is determined by its
volume, which grows faster than the oxygen-absorbing area if an animal’s size increases equally in all
directions. An increase in the concentration of oxygen in air or water would increase the size to which an
organism could grow without its tissues becoming starved of oxygen. However, members of the Ediacara
biota reached metres in length; clearly oxygen did not limit their growth.[31] Other metabolic functions may
have been inhibited by lack of oxygen, for example the construction of tissue such as collagen, required for
the construction of complex structures,[92] or to form molecules for the construction of a hard exoskeleton.
[93] However, animals are not affected when similar oceanographic conditions occur in the Phanerozoic;
there is no convincing correlation between oxygen levels and evolution, so oxygen may have been no more a
prerequisite to complex life than liquid water or primary productivity.[94]
[edit] Snowball Earths

Main article: Snowball Earth
In the late Neoproterozoic (extending into the early Ediacaran period), the Earth suffered massive
glaciations in which most of its surface was covered by ice. This may have caused a mass extinction,
creating a genetic bottleneck; the resulting diversification may have given rise to the Ediacara biota, which
appears soon after the last "Snowball Earth" episode.[95] However, the snowball episodes occurred a long
time before the start of the Cambrian, and it is hard to see how so much diversity could have been caused by
even a series of bottlenecks;[19] the cold periods may even have delayed the evolution of large size.[42]
[edit] Developmental explanations

A range of theories are based on the concept that minor modifications to animals' development as
they grow from embryo to adult may have been able to cause very large changes in the final adult form. The
hox genes, for example, control which organs individual regions of an embryo will develop into. For instance,
if a certain hox gene is expressed, a region will develop into a limb; if a different hox gene is expressed in
that region (a minor change), it could develop into an eye instead (a phenotypically major change).
Such a system allows a large range of disparity to appear from a limited set of genes, but such
theories linking this with the explosion struggle to explain why the origin of such a development system
should by itself lead to increased diversity or disparity. Evidence of Precambrian metazoans[19] combines
with molecular data[96] to show that much of the genetic architecture that could feasibly have played a role in
the explosion was already well established by the Cambrian.
This apparent paradox is addressed in a theory that focuses on the physics of development. It is
proposed that the emergence of simple multicellular forms provided a changed context and spatial scale in
which novel physical processes and effects were mobilized by the products of genes that had previously
evolved to serve unicellular functions. Morphological complexity (layers, segments, lumens, appendages)
arose, in this view, by self-organization.[97]
[edit] Ecological explanations

These focus on the interactions between different types of organism. Some of these hypotheses deal
with changes in the food chain; some suggest arms races between predators and prey, and others focus on
the more general mechanisms of coevolution. Such theories are well suited to explaining why there was a
rapid increase in both disparity and diversity, but they must explain why the "explosion" happened when it
did.[19]
[edit] End-Ediacaran mass extinction

Main article: End-Ediacaran extinction
Evidence for such an extinction includes the disappearance from the fossil record of the Ediacara
biota and shelly fossils such as Cloudina, and the accompanying perturbation in the δ13C record. Mass
extinctions are often followed by adaptive radiations as existing clades expand to occupy the ecospace
emptied by the extinction. However, once the dust had settled, overall disparity and diversity returned to the
pre-extinction level in each of the Phanerozoic extinctions.[19]
[edit] Evolution of eyes
Main article: Evolution of the eye
Andrew Parker has proposed that predator-prey relationships changed dramatically after eyesight
evolved. Prior to that time hunting and evading were both close-range affairs – smell, vibration, and touch
were the only senses used. When predators could see their prey from a distance, new defensive strategies
were needed. Armor, spines, and similar defenses may also have evolved in response to vision. He further
observed that where animals lose vision in unlighted environments such as caves, diversity of animal forms
tends to decrease.[98] Nevertheless many scientists doubt that vision could have caused the explosion. Eyes
may well have evolved long before the start of the Cambrian.[99] It is also difficult to understand why the
evolution of eyesight would have caused an explosion, since other senses such as smell and pressure
detection can detect things further away than they can be seen under the sea, but the appearance of these
other senses apparently did not cause an evolutionary explosion.[19]
[edit] Arms races between predators and prey

The ability to avoid or recover from predation often makes the difference between life and death, and
is therefore one of the strongest components of natural selection. The pressure to adapt is stronger on the
prey than on the predator: if the predator fails to win a contest, it loses its lunch; if the prey is the loser, it
loses its life.[100]
But there is evidence that predation was rife long before the start of the Cambrian, for example in the
increasingly spiny forms of acritarchs, the holes drilled in Cloudina shells, and traces of burrowing to avoid
predators. Hence it is unlikely that the appearance of predation was the trigger for the Cambrian "explosion",
although it may well have exhibited a strong influence on the body forms that the "explosion" produced.[42]
However, the intensity of predation does appear to have increased dramatically during the Cambrian as new
predatory "tactics" (such as shell-crushing) emerged.[101]
[edit] Increase in size and diversity of planktonic animals

Geochemical evidence strongly indicates that the total mass of plankton has been similar to modern
levels since early in the Proterozoic. Before the start of the Cambrian, their corpses and droppings were too
small to fall quickly towards the seabed, since their drag was about the same as their weight. This meant they
were destroyed by scavengers or by chemical processes before they reached the sea floor.[26]
Mesozooplankton are plankton of a larger size, and early Cambrian specimens filtered microscopic
plankton from the seawater. These larger organisms would have produced droppings and corpses that were
large enough to fall fairly quickly. This provided a new supply of energy and nutrients to the mid-levels and
bottoms of the seas, which opened up a huge range of new possible ways of life. If any of these remains
sank uneaten to the sea floor they could be buried; this would have taken some carbon out of circulation,
resulting in an increase in the concentration of breathable oxygen in the seas (carbon readily combines with
oxygen).[26]
The initial herbivorous mesozooplankton were probably larvae of benthic (seafloor) animals. A larval
stage was probably an evolutionary innovation driven by the increasing level of predation at the seafloor
during the Ediacaran period.[3][102]
Metazoans have an amazing ability to increase diversity through co-evolution.[4] This means that a
trait of one organism can cause another to evolve in response; a number of responses are possible, and a
different species can potentially emerge for each. As a simple example, the evolution of predation may have
caused one organism to develop defence while another developed motion to flee. This would cause the
predator lineage to split into two species: one that was good at chasing prey, and another that was good at
breaking through defences. Actual co-evolution is somewhat more subtle, but in this fashion, great diversity
can arise: three quarters of living species are animals, and most of the rest have formed by co-evolution with
animals.[4]
[edit] Mis-dating of species

The results of an article published in Nature in 2010,[103] have shown that eukaryotic multicellularity,
which had been thought to evolve with the beginning of Cambrian Period, might date back to 2.1 billion years
ago, which is approximately 1.55 billion years earlier than the date indicated by currently dominating scientific
evidence.[104]
[edit] Discredited hypotheses

Main article: Discredited hypotheses for the Cambrian explosion
As our understanding of the events of the Cambrian becomes clearer, data has accumulated to make
some hypotheses look improbable. Causes that have been proposed but are now discounted include the
evolution of herbivory, vast changes in the speed of tectonic plate movement or of the cyclic changes in the
Earth's orbital motion, or the operation of different evolutionary mechanisms from those that are seen in the
rest of the Phanerozoic eon.[citation needed]
[edit] No explanation required

The explosion may not have been a significant evolutionary event. It may represent a threshold being
crossed: for example a threshold in genetic complexity that allowed a vast range of morphological forms to
be employed.[105]
[edit] Uniqueness of the explosion
The "Cambrian explosion" can be viewed as two waves of metazoan expansion into empty niches:
first, a co-evolutionary rise in diversity as animals explored niches on the Ediacaran sea floor, followed by a
second expansion in the early Cambrian as they became established in the water column.[4] The rate of
diversification seen in the Cambrian phase of the explosion is unparalleled among marine animals: it affected
all metazoan clades of which Cambrian fossils have been found. Later radiations, such as those of fish in the
Silurian and Devonian periods, involved fewer taxa, mainly with very similar body plans.[10] Although the
recovery from the Permian-Triassic extinction started with about as few animal species as the Cambrian
explosion, the recovery produced far fewer significantly new types of animals.[106]
Whatever triggered the early Cambrian diversification opened up an exceptionally wide range of
previously-unavailable ecological niches. When these were all occupied, there was little room for such wide-
ranging diversifications to occur again, because there was strong competition in all niches and incumbents
usually had the advantage. If there had continued to be a wide range of empty niches, clades would be able
to continue diversifying and become disparate enough for us to recognise them as different phyla; when
niches are filled, lineages will continue to resemble one another long after they diverge, as there is limited
opportunity for them to change their life-styles and forms.[107]
There were two similar explosions in the evolution of land plants: after a cryptic history beginning
about 450 million years ago, land plants underwent a uniquely rapid adaptive radiation during the Devonian
period, about 400 million years ago.[10] Furthermore, Angiosperms (flowering plants) originated and rapidly
diversified during the Cretaceous period.
[edit] Notes
^1 Older marks found in billion-year old rocks[108] have since been recognised as non-biogenic.[51]
[89]
^2 The analysis considered the bioprovinciality of trilobite lineages, as well as their evolutionary rate.
[109]
[edit] References
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Part of a series on
The Cambrian explosion
[show]Fossil localities
• Burgess shale
• Chengjiang
• Sirius Passet
• Doushantuo
[show]Key organisms
[show]Ediacara biota
• Kimberella
• Vernanimalcu
la
[show]Burgess-type
• Marrella
• Anomalocari
dids
• Halwaxiids
• Opabinia
• Odontogriph
us
[show]Small shelly fauna
• Helcionellids
[show]Evolutionary concepts
Trends
• Cambrian substrate
revolution
Themes
• Cladistics
• Convergent evolution
• Stem and crown
groups
v · d · e• taskforce
• Budd, G. E. & Jensen, J. (2000). "A critical reappraisal of the fossil record of the bilaterian
phyla". Biological Reviews 75 (2): 253–295. doi:10.1017/S000632310000548X. PMID 10881389.
• Collins, Allen G. “Metazoa: Fossil record”. Retrieved Dec. 14, 2005.
• Conway Morris, S. (1997). The Crucible of Creation: the Burgess Shale and the rise of
animals. Oxford University Press. ISBN 0–19–286202–2.
• Conway Morris, S. (June 2006). "Darwin’s dilemma: the realities of the Cambrian
‘explosion’". Philosophical Transactions of the Royal Society B: Biological Sciences 361 (1470):
1069–1083. doi:10.1098/rstb.2006.1846. ISSN 0962-8436. PMID 16754615. An enjoyable account.
• Gould, S.J. (1989). Wonderful Life: The Burgess Shale and the Nature of History. W.W.
Norton & Company.
• Kennedy, M., M. Droser, L. Mayer., D. Pevear, and D. Mrofka (2006). "Clay and Atmospheric
Oxygen". Science 311 (5766): 1341. doi:10.1126/science.311.5766.1341c.
• Knoll, A.H.; Carroll, S.B. (1999-06-25). "Early Animal Evolution: Emerging Views from
Comparative Biology and Geology". Science 284 (5423): 2129. doi:10.1126/science.284.5423.2129.
PMID 10381872.
• Alexander V. Markov, and Andrey V. Korotayev (2007) “Phanerozoic marine biodiversity
follows a hyperbolic trend” Palaeoworld 16(4): pp. 311–318.
• "Montenari, M. & Leppig, U. (2003): The Acritarcha: their classification morphology,
ultrastructure and palaeoecological/palaeogeographical distribution.". Paläontologische Zeitschrift
77: 173–194. 2003.
• Wang, D. Y.-C., S. Kumar and S. B. Hedges (January 1999). "Divergence time estimates for
the early history of animal phyla and the origin of plants, animals and fungi". Proceedings of the
Royal Society of London, Series B, Biological Sciences 266 (1415): 163–71.
doi:10.1098/rspb.1999.0617. ISSN 0962-8452. PMID 10097391.
• Xiao, S., Y. Zhang, and A. Knoll (January 1998). "Three-dimensional preservation of algae
and animal embryos in a Neoproterozoic phosphorite". Nature 391 (1): 553–58. doi:10.1038/35318.
ISSN 0090-9556. Bibcode: 1998Natur.391..553X.
Timeline References:
• Martin, M.W; Grazhdankin, D.V; Bowring, S.A; Evans, D.A.D; Fedonkin, M.A; Kirschvink, J.L
(2000). "Age of Neoproterozoic Bilaterian Body and Trace Fossils, White Sea, Russia: Implications
for Metazoan Evolution". Science 288 (5467): 841–845. doi:10.1126/science.288.5467.841.
PMID 10797002.

• The Cambrian “explosion” of metazoans and molecular biology: would Darwin be satisfied?
• On embryos and ancestors by Stephen Jay Gould
• "The Cambrian “explosion”: Slow-fuse or megatonnage?" (Scholar search).
http://www.pnas.org/content/97/9/4426.full.
• The Cambrian Explosion – In Our Time, BBC Radio 4 broadcast, 17 February 2005
• Utah's Cambrian life – new (2008) website with good images of a range of Burgess-shale-
type and other Cambrian fossils.
Retrieved from "http://en.wikipedia.org/wiki/Cambrian_explosion"
Categories: Paleontology | Cambrian | Fossils

W000
Plant

For other uses, see Plant (disambiguation).
Plants
Temporal range:
Early Cambrian to recent, but see text, 520–0 Ma
PreЄ
g
Scientific classification
Domain: Eukaryota
(unranked): Archaeplastida
Plantae
Kingdom:
Haeckel, 1866[1]
Divisions
Green algae
• Chlorophyta
• Charophyta
Land plants (embryophytes)
• Non-vascular land plants (bryophytes)
• Marchantiophyta—liverworts
• Anthocerotophyta—hornworts
• Bryophyta—mosses
• †Horneophytopsida
• Vascular plants (tracheophytes)
• †Rhyniophyta—rhyniophytes
• †Zosterophyllophyta—zosterophylls
• Lycopodiophyta—clubmosses
• †Trimerophytophyta—trimerophytes
• Pteridophyta—ferns and horsetails
• †Progymnospermophyta
• Seed plants (spermatophytes)
• †Pteridospermatophyta—
seed ferns
• Pinophyta—conifers
• Cycadophyta—cycads
• Ginkgophyta—ginkgo
• Gnetophyta—gnetae
• Magnoliophyta—flowering
plants
†Nematophytes
Plants are living organisms belonging to the kingdom Plantae. They include familiar organisms such
as trees, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The scientific study of plants,
known as botany, has identified about 350,000 extant species of plants, defined as seed plants, bryophytes,
ferns and fern allies. As of 2004, some 287,655 species had been identified, of which 258,650 are flowering
and 18,000 bryophytes (see table below). Green plants, sometimes called Viridiplantae, obtain most of their
energy from sunlight via a process called photosynthesis.
Contents
[hide]
• 1 Definition
• 1.1 Current definitions of Plantae
• 1.2 Algae
• 1.3 Fungi
• 2 Diversity
• 2.1 Phylogeny
• 2.2 Embryophytes
• 2.3 Fossils
• 3 Structure, growth, and development
• 3.1 Factors affecting growth
• 3.2 Plant cell
• 4 Physiology
• 4.1 Photosynthesis
• 4.2 Immune system
• 4.3 Internal distribution
• 5 Ecology
• 5.1 Distribution
• 5.2 Ecological relationships
• 6 Importance
• 6.1 Food
Definition
Aristotle divided all living things between plants (which generally do not move), and animals (which
often are mobile to catch their food). In Linnaeus' system, these became the Kingdoms Vegetabilia (later
Metaphyta or Plantae) and Animalia (also called Metazoa). Since then, it has become clear that the Plantae
as originally defined included several unrelated groups, and the fungi and several groups of algae were
removed to new kingdoms. However, these are still often considered plants in many contexts, both technical
and popular.
Current definitions of Plantae

When the name Plantae or plants is applied to a specific taxon, it is usually referring to one of three
concepts. From smallest to largest in inclusiveness, these three groupings are:
Name(s) Scope Description
Land plants, also

Plantae As the narrowest of plant categories, this is
known as Embryophyta or
sensu strictissimo further delineated below.
Metaphyta.
Comprise the above Embryophytes, Charophyta

Green plants - also (i.e., primitive stoneworts), and Chlorophyta (i.e., green
Plantae algae such as sea lettuce). Viridiplantae encompasses a
known as Viridiplantae,
sensu stricto group of organisms that possess chlorophyll a and b,
Viridiphyta or Chlorobionta
have plastids that are bound by only two membranes,
are capable of storing starch, and have cellulose in their
cell walls. It is this clade which is mainly the subject of
this article.
Comprises the green plants above, as well as

Rhodophyta (red algae) and Glaucophyta (simple
Archaeplastida, Plantae glaucophyte algae). As the broadest plant clade, this
Plastida or Primoplantae sensu lato comprises most of the eukaryotes that eons ago
acquired their chloroplasts directly by engulfing
cyanobacteria.
Outside of formal scientific contexts, the term "plant" implies an association with certain traits, such
as multicellularity, cellulose, and photosynthesis.[2][3] Many of the classification controversies involve
organisms that are rarely encountered and are of minimal apparent economic significance, but are crucial in
developing an understanding of the evolution of modern flora.
Algae
Green algae from Ernst Haeckel's Kunstformen der Natur, 1904.

Main article: Algae
Most algae are no longer classified within the Kingdom Plantae.[4][5] The algae comprise several
different groups of organisms that produce energy through photosynthesis, each of which arose
independently from separate non-photosynthetic ancestors. Most conspicuous among the algae are the
seaweeds, multicellular algae that may roughly resemble terrestrial plants, but are classified among the
green, red, and brown algae. Each of these algal groups also includes various microscopic and single-celled
organisms.
The two groups of green algae are the closest relatives of land plants (embryophytes). The first of
these groups is the Charophyta (desmids and stoneworts), from which the embryophytes developed.[6][7][8]
The sister group to the combined embryophytes and charophytes is the other group of green algae,
Chlorophyta, and this more inclusive group is collectively referred to as the green plants or Viridiplantae. The
Kingdom Plantae is often taken to mean this monophyletic grouping. With a few exceptions among the green
algae, all such forms have cell walls containing cellulose, have chloroplasts containing chlorophylls a and b,
and store food in the form of starch. They undergo closed mitosis without centrioles, and typically have
mitochondria with flat cristae.
The chloroplasts of green plants are surrounded by two membranes, suggesting they originated
directly from endosymbiotic cyanobacteria. The same is true of two additional groups of algae: the
Rhodophyta (red algae) and Glaucophyta. All three groups together are generally believed to have a
common origin, and so are classified together in the taxon Archaeplastida. In contrast, most other algae (e.g.
heterokonts, haptophytes, dinoflagellates, and euglenids) have chloroplasts with three or four surrounding
membranes. They are not close relatives of the green plants, presumably acquiring chloroplasts separately
from ingested or symbiotic green and red algae.
Fungi
Main article: Fungi
The classification of fungi has been controversial until quite recently in the history of biology.
Linnaeus' original classification placed the fungi within the Plantae, since they were unquestionably not
animalian; this being the only other alternative. With later developments in microbiology, in the 19th century
Ernst Haeckel felt that a third kingdom was required to classify newly discovered micro-organisms. The
introduction of the new kingdom Protista as an alternative to Animalia, led to uncertainty as to whether fungi
truly were best placed in the Plantae or whether they ought to be reclassified as protists. Haeckel himself
found it difficult to decide and it was not until 1969 that a solution was found whereby Robert Whittaker
proposed the creation of the kingdom Fungi. Molecular evidence has since shown that the concestor (last
common ancestor) of the Fungi was probably more similar to that of the Animalia than of any other kingdom,
including the Plantae.
Whittaker's original reclassification was based on the fundamental difference in nutrition between the
Fungi and the Plantae. Unlike plants, which are generally autotrophic multicellular phototrophs which gain
carbon through photosynthesis, fungi are generally heterotrophic uni- or multi-cellular saprotrophs, obtaining
carbon by breaking down and absorbing surrounding materials. In addition, the substructure of multicellular
fungi takes the form of many chitinous microscopic strands called hyphae, which may be further subdivided
into cells or may form a syncytium containing many eukaryotic nuclei. Fruiting bodies, of which mushrooms
are most familiar example, are the reproductive structures of fungi.
Diversity
About 350,000 species of plants, defined as seed plants, bryophytes, ferns and fern allies, are
estimated to exist currently. As of 2004, some 287,655 species had been identified, of which 258,650 are
flowering plants, 16,000 bryophytes, 11,000 ferns and 8,000 green algae.
Diversity of living plant divisions
No. of living
Informal group Division name Common name
species
Green algae green algae

Chlorophyta 3,800 [9]
(chlorophytes)
green algae (desmids & 4,000 - 6,000

Charophyta
charophytes) [10]
Bryophytes 6,000 - 8,000

Marchantiophyta liverworts
[11]
Anthocerotophyta hornworts 100 - 200 [12]
Bryophyta mosses 12,000 [13]
Pteridophytes Lycopodiophyta club mosses 1,200 [5]

ferns, whisk ferns &
Pteridophyta 11,000 [5]
horsetails
Seed plants Cycadophyta cycads 160 [14]
Ginkgophyta ginkgo 1 [15]
Pinophyta conifers 630 [5]
Gnetophyta gnetophytes 70 [5]
Magnoliophyta flowering plants 258,650 [16]
The naming of plants is governed by the International Code of Botanical Nomenclature and International
Code of Nomenclature for Cultivated Plants (see cultivated plant taxonomy).
Phylogeny
This article may be confusing or unclear to readers. Please help clarify the article;
suggestions may be found on the talk page. (March 2009)
A proposed phylogenetic tree of Plantae, after Kenrick and Crane,[17] is as follows, with modification
to the Pteridophyta from Smith et al.[18] The Prasinophyceae may be a paraphyletic basal group to all green
plants.
Streptobionta
Embryophytes
Stomatophytes
Polysporangiates
Tracheophytes
Eutracheophytes
Euphyllophytina
Lignophytia
Spermatophytes (seed
plants)
Progymnospermophyta †
Pteridophyta
Pteridopsida
(true ferns)
Marattiopsida
Equisetopsida
(horsetails)
Psilotopsida
(whisk ferns & adders'-
tongues)
Cladoxylopsida †
Lycophytina
Lycopodiophyta
Zosterophyllophyta †
Rhyniophyta †
Aglaophyton †
Horneophytopsida †
Chlorophyta
Trebouxiophyceae (Pleurastrophyceae)
Chlorophyceae
Ulvophyceae
Embryophytes
Main article: Embryophyte
Dicksonia antarctica, a species of tree fern
The plants that are likely most familiar to us are the multicellular land plants, called embryophytes.
They include the vascular plants, plants with full systems of leaves, stems, and roots. They also include a few
of their close relatives, often called bryophytes, of which mosses and liverworts are the most common.
All of these plants have eukaryotic cells with cell walls composed of cellulose, and most obtain their
energy through photosynthesis, using light and carbon dioxide to synthesize food. About three hundred plant
species do not photosynthesize but are parasites on other species of photosynthetic plants. Plants are
distinguished from green algae, which represent a mode of photosynthetic life similar to the kind modern
plants are believed to have evolved from, by having specialized reproductive organs protected by non-
reproductive tissues.
Bryophytes first appeared during the early Paleozoic. They can only survive where moisture is
available for significant periods, although some species are desiccation tolerant. Most species of bryophyte
remain small throughout their life-cycle. This involves an alternation between two generations: a haploid
stage, called the gametophyte, and a diploid stage, called the sporophyte. The sporophyte is short-lived and
remains dependent on its parent gametophyte.
Vascular plants first appeared during the Silurian period, and by the Devonian had diversified and
spread into many different land environments. They have a number of adaptations that allowed them to
overcome the limitations of the bryophytes. These include a cuticle resistant to desiccation, and vascular
tissues which transport water throughout the organism. In most the sporophyte acts as a separate individual,
while the gametophyte remains small.
The first primitive seed plants, Pteridosperms (seed ferns) and Cordaites, both groups now extinct,
appeared in the late Devonian and diversified through the Carboniferous, with further evolution through the
Permian and Triassic periods. In these the gametophyte stage is completely reduced, and the sporophyte
begins life inside an enclosure called a seed, which develops while on the parent plant, and with fertilisation
by means of pollen grains. Whereas other vascular plants, such as ferns, reproduce by means of spores and
so need moisture to develop, some seed plants can survive and reproduce in extremely arid conditions.
Early seed plants are referred to as gymnosperms (naked seeds), as the seed embryo is not
enclosed in a protective structure at pollination, with the pollen landing directly on the embryo. Four surviving
groups remain widespread now, particularly the conifers, which are dominant trees in several biomes. The
angiosperms, comprising the flowering plants, were the last major group of plants to appear, emerging from
within the gymnosperms during the Jurassic and diversifying rapidly during the Cretaceous. These differ in
that the seed embryo (angiosperm) is enclosed, so the pollen has to grow a tube to penetrate the protective
seed coat; they are the predominant group of flora in most biomes today.
Fossils
Main articles: Paleobotany and Evolutionary history of plants
A petrified log in Petrified Forest National Park.
Plant fossils include roots, wood, leaves, seeds, fruit, pollen, spores, phytoliths, and amber (the
fossilized resin produced by some plants). Fossil land plants are recorded in terrestrial, lacustrine, fluvial and
nearshore marine sediments. Pollen, spores and algae (dinoflagellates and acritarchs) are used for dating
sedimentary rock sequences. The remains of fossil plants are not as common as fossil animals, although
plant fossils are locally abundant in many regions worldwide.
The earliest fossils clearly assignable to Kingdom Plantae are fossil green algae from the Cambrian.
These fossils resemble calcified multicellular members of the Dasycladales. Earlier Precambrian fossils are
known which resemble single-cell green algae, but definitive identity with that group of algae is uncertain.
The oldest known fossils of embryophytes date from the Ordovician, though such fossils are
fragmentary. By the Silurian, fossils of whole plants are preserved, including the lycophyte Baragwanathia
longifolia. From the Devonian, detailed fossils of rhyniophytes have been found. Early fossils of these ancient
plants show the individual cells within the plant tissue. The Devonian period also saw the evolution of what
many believe to be the first modern tree, Archaeopteris. This fern-like tree combined a woody trunk with the
fronds of a fern, but produced no seeds.
The Coal measures are a major source of Paleozoic plant fossils, with many groups of plants in
existence at this time. The spoil heaps of coal mines are the best places to collect; coal itself is the remains
of fossilised plants, though structural detail of the plant fossils is rarely visible in coal. In the Fossil Forest at
Victoria Park in Glasgow, Scotland, the stumps of Lepidodendron trees are found in their original growth
positions.
The fossilized remains of conifer and angiosperm roots, stems and branches may be locally
abundant in lake and inshore sedimentary rocks from the Mesozoic and Cenozoic eras. Sequoia and its
allies, magnolia, oak, and palms are often found.
Petrified wood is common in some parts of the world, and is most frequently found in arid or desert
areas where it is more readily exposed by erosion. Petrified wood is often heavily silicified (the organic
material replaced by silicon dioxide), and the impregnated tissue is often preserved in fine detail. Such
specimens may be cut and polished using lapidary equipment. Fossil forests of petrified wood have been
found in all continents.
Fossils of seed ferns such as Glossopteris are widely distributed throughout several continents of the
Southern Hemisphere, a fact that gave support to Alfred Wegener's early ideas regarding Continental drift
theory.
Structure, growth, and development

Further information: Plant morphology
Most of the solid material in a plant is taken from the atmosphere. Through a process known as
photosynthesis, most plants use the energy in sunlight to convert carbon dioxide from the atmosphere, plus
water, into simple sugars. Parasitic plants, on the other hand, use the resources of its host to grow. These
sugars are then used as building blocks and form the main structural component of the plant. Chlorophyll, a
green-colored, magnesium-containing pigment is essential to this process; it is generally present in plant
leaves, and often in other plant parts as well.
Plants usually rely on soil primarily for support and water (in quantitative terms), but also obtain
compounds of nitrogen, phosphorus, and other crucial elemental nutrients. Epiphytic and lithophytic plants
often depend on rainwater or other sources for nutrients and carnivorous plants supplement their nutrient
requirements with insect prey that they capture. For the majority of plants to grow successfully they also
require oxygen in the atmosphere and around their roots for respiration. However, some plants grow as
submerged aquatics, using oxygen dissolved in the surrounding water, and a few specialized vascular plants,
such as mangroves, can grow with their roots in anoxic conditions.
The leaf is usually the primary site of photosynthesis in plants.

There is no photosynthesis in deciduous leaves in autumn.
Factors affecting growth

The genotype of a plant affects its growth. For example, selected varieties of wheat grow rapidly,
maturing within 110 days, whereas others, in the same environmental conditions, grow more slowly and
mature within 155 days.[19]
Growth is also determined by environmental factors, such as temperature, available water, available
light, and available nutrients in the soil. Any change in the availability of these external conditions will be
reflected in the plants growth.
Biotic factors are also capable of affecting plant growth. Plants compete with other plants for space,
water, light and nutrients. Plants can be so crowded that no single individual produces normal growth,
causing etiolation and chlorosis. Optimal plant growth can be hampered by grazing animals, suboptimal soil
composition, lack of mycorrhizal fungi, and attacks by insects or plant diseases, including those caused by
bacteria, fungi, viruses, and nematodes.[19]
Simple plants like algae may have short life spans as individuals, but their populations are commonly
seasonal. Other plants may be organized according to their seasonal growth pattern: annual plants live and
reproduce within one growing season, biennial plants live for two growing seasons and usually reproduce in
second year, and perennial plants live for many growing seasons and continue to reproduce once they are
mature. These designations often depend on climate and other environmental factors; plants that are annual
in alpine or temperate regions can be biennial or perennial in warmer climates. Among the vascular plants,
perennials include both evergreens that keep their leaves the entire year, and deciduous plants which lose
their leaves for some part of it. In temperate and boreal climates, they generally lose their leaves during the
winter; many tropical plants lose their leaves during the dry season.
The growth rate of plants is extremely variable. Some mosses grow less than 0.001 millimeters per
hour (mm/h), while most trees grow 0.025-0.250 mm/h. Some climbing species, such as kudzu, which do not
need to produce thick supportive tissue, may grow up to 12.5 mm/h.
Dried dead plants
Plants protect themselves from frost and dehydration stress with antifreeze proteins, heat-shock
proteins and sugars (sucrose is common). LEA (Late Embryogenesis Abundant) protein expression is
induced by stresses and protects other proteins from aggregation as a result of desiccation and freezing.[20]
Plant cell
Plant cell structure

Main article: Plant cell
Plant cells are typically distinguished by their large water-filled central vacuole, chloroplasts, and rigid
cell walls that are made up of cellulose, hemicellulose, and pectin. Cell division is also characterized by the
development of a phragmoplast for the construction of a cell plate in the late stages of cytokinesis. Just as in
animals, plant cells differentiate and develop into multiple cell types. Totipotent meristematic cells can
differentiate into vascular, storage, protective (e.g. epidermal layer), or reproductive tissues, with more
primitive plants lacking some tissue types.[21]
Physiology
Main article: Plant physiology
Photosynthesis
Main articles: Photosynthesis and Biological pigment
Plants are photosynthetic, which means that they manufacture their own food molecules using
energy obtained from light. The primary mechanism plants have for capturing light energy is the pigment
chlorophyll. All green plants contain two forms of chlorophyll, chlorophyll a and chlorophyll b. The latter of
these pigments is not found in red or brown algae.
Immune system
By means of cells that behave like nerves, plants receive and distribute within their systems
information about incident light intensity and quality. Incident light which stimulates a chemical reaction in
one leaf, will cause a chain reaction of signals to the entire plant via a type of cell termed a "bundle sheath
cell". Researchers from the Warsaw University of Life Sciences in Poland, found that plants have a specific
memory for varying light conditions which prepares their immune systems against seasonal pathogens.[22]
Internal distribution
Main article: Vascular tissue
Vascular plants differ from other plants in that they transport nutrients between different parts through
specialized structures, called xylem and phloem. They also have roots for taking up water and minerals. The
xylem moves water and minerals from the root to the rest of the plant, and the phloem provides the roots with
sugars and other nutrient produced by the leaves.[21]
Ecology
Main article: Plant ecology
The photosynthesis conducted by land plants and algae is the ultimate source of energy and organic
material in nearly all ecosystems. Photosynthesis radically changed the composition of the early Earth's
atmosphere, which as a result is now 21% oxygen. Animals and most other organisms are aerobic, relying on
oxygen; those that do not are confined to relatively rare anaerobic environments. Plants are the primary
producers in most terrestrial ecosystems and form the basis of the food web in those ecosystems. Many
animals rely on plants for shelter as well as oxygen and food.
Land plants are key components of the water cycle and several other biogeochemical cycles. Some
plants have coevolved with nitrogen fixing bacteria, making plants an important part of the nitrogen cycle.
Plant roots play an essential role in soil development and prevention of soil erosion.
Distribution
This section requires expansion.
Plants are distributed worldwide in varying numbers. While they inhabit a multitude of biomes and
ecoregions, few can be found beyond the tundras at the northernmost regions of continental shelves. At the
southern extremes, plants have adapted tenaciously to the prevailing conditions. (See Antarctic flora.)
Plants are often the dominant physical and structural component of habitats where they occur. Many
of the Earth's biomes are named for the type of vegetation because plants are the dominant organisms in
those biomes, such as grasslands and forests.
Ecological relationships
The Venus flytrap, a species of carnivorous plant.

Numerous animals have coevolved with plants. Many animals pollinate flowers in exchange for food
in the form of pollen or nectar. Many animals disperse seeds, often by eating fruit and passing the seeds in
their feces. Myrmecophytes are plants that have coevolved with ants. The plant provides a home, and
sometimes food, for the ants. In exchange, the ants defend the plant from herbivores and sometimes
competing plants. Ant wastes provide organic fertilizer.
The majority of plant species have various kinds of fungi associated with their root systems in a kind
of mutualistic symbiosis known as mycorrhiza. The fungi help the plants gain water and mineral nutrients
from the soil, while the plant gives the fungi carbohydrates manufactured in photosynthesis. Some plants
serve as homes for endophytic fungi that protect the plant from herbivores by producing toxins. The fungal
endophyte, Neotyphodium coenophialum, in tall fescue (Festuca arundinacea) does tremendous economic
damage to the cattle industry in the U.S.
Various forms of parasitism are also fairly common among plants, from the semi-parasitic mistletoe
that merely takes some nutrients from its host, but still has photosynthetic leaves, to the fully parasitic
broomrape and toothwort that acquire all their nutrients through connections to the roots of other plants, and
so have no chlorophyll. Some plants, known as myco-heterotrophs, parasitize mycorrhizal fungi, and hence
act as epiparasites on other plants.
Many plants are epiphytes, meaning they grow on other plants, usually trees, without parasitizing
them. Epiphytes may indirectly harm their host plant by intercepting mineral nutrients and light that the host
would otherwise receive. The weight of large numbers of epiphytes may break tree limbs. Hemiepiphytes like
the strangler fig begin as epiphytes but eventually set their own roots and overpower and kill their host. Many
orchids, bromeliads, ferns and mosses often grow as epiphytes. Bromeliad epiphytes accumulate water in
leaf axils to form phytotelmata, complex aquatic food webs.[23]
Approximately 630 plants are carnivorous, such as the Venus Flytrap (Dionaea muscipula) and
sundew (Drosera species). They trap small animals and digest them to obtain mineral nutrients, especially
nitrogen and phosphorus.[24]
Importance
Potato plant. Potatoes spread to the rest of the world after European contact with the Americas in the
late 15th and early 16th centuries and have since become an important field crop.
Timber in storage for later processing at a sawmill.

A section of a Yew branch showing 27 annual growth rings, pale sapwood and dark heartwood, and
pith (centre dark spot). The dark radial lines are longitudinal sections of small branches which became
included by growth of the tree.
The study of plant uses by people is termed economic botany or ethnobotany; some consider
economic botany to focus on modern cultivated plants, while ethnobotany focuses on indigenous plants
cultivated and used by native peoples. Human cultivation of plants is part of agriculture, which is the basis of
human civilization. Plant agriculture is subdivided into agronomy, horticulture and forestry.
Food
Much of human nutrition depends on land plants, either directly or indirectly.
Human nutrition depends to a large extent on cereals, especially maize (or corn), wheat and rice. Other
staple crops include potato, cassava, and legumes. Human food also includes vegetables, spices, and
certain fruits, nuts, herbs, and edible flowers.
Beverages produced from plants include coffee, tea, wine, beer and alcohol.
Sugar is obtained mainly from sugar cane and sugar beet.
Cooking oils and margarine come from maize, soybean, rapeseed, safflower, sunflower, olive and others.
Food additives include gum arabic, guar gum, locust bean gum, starch and pectin.
Livestock animals including cows, pigs, sheep, and goats are all herbivores; and feed primarily or entirely on
cereal plants, particularly grasses.
Nonfood products
Wood is used for buildings, furniture, paper, cardboard, musical instruments and sports equipment.
Cloth is often made from cotton, flax or synthetic fibers derived from cellulose, such as rayon and acetate.
Renewable fuels from plants include firewood, peat and many other biofuels. Coal and petroleum are fossil
fuels derived from plants. Medicines derived from plants include aspirin, taxol, morphine, quinine, reserpine,
colchicine, digitalis and vincristine. There are hundreds of herbal supplements such as ginkgo, Echinacea,
feverfew, and Saint John's wort. Pesticides derived from plants include nicotine, rotenone, strychnine and
pyrethrins. Drugs obtained from plants include opium, cocaine and marijuana. Poisons from plants include
ricin, hemlock and curare. Plants are the source of many natural products such as fibers, essential oils,
natural dyes, pigments, waxes, tannins, latex, gums, resins, alkaloids, amber and cork. Products derived
from plants include soaps, paints, shampoos, perfumes, cosmetics, turpentine, rubber, varnish, lubricants,
linoleum, plastics, inks, chewing gum and hemp rope. Plants are also a primary source of basic chemicals for
the industrial synthesis of a vast array of organic chemicals. These chemicals are used in a vast variety of
studies and experiments.
Aesthetic uses
Thousands of plant species are cultivated for aesthetic purposes as well as to provide shade, modify
temperatures, reduce wind, abate noise, provide privacy, and prevent soil erosion. People use cut flowers,
dried flowers and houseplants indoors or in greenhouses. In outdoor gardens, lawn grasses, shade trees,
ornamental trees, shrubs, vines, herbaceous perennials and bedding plants are used. Images of plants are
often used in art, architecture, humor, language, and photography and on textiles, money, stamps, flags and
coats of arms. Living plant art forms include topiary, bonsai, ikebana and espalier. Ornamental plants have
sometimes changed the course of history, as in tulipomania. Plants are the basis of a multi-billion dollar per
year tourism industry which includes travel to arboretums, botanical gardens, historic gardens, national
parks, tulip festivals, rainforests, forests with colorful autumn leaves and the National Cherry Blossom
Festival. Venus Flytrap, sensitive plant and resurrection plant are examples of plants sold as novelties.
Scientific and cultural uses

Tree rings are an important method of dating in archeology and serve as a record of past climates.
Basic biological research has often been done with plants, such as the pea plants used to derive Gregor
Mendel's laws of genetics. Space stations or space colonies may one day rely on plants for life support.
Plants are used as national and state emblems, including state trees and state flowers. Ancient trees are
revered and many are famous. Numerous world records are held by plants. Plants are often used as
memorials, gifts and to mark special occasions such as births, deaths, weddings and holidays. Plants figure
prominently in mythology, religion and literature. The field of ethnobotany studies plant use by indigenous
cultures which helps to conserve endangered species as well as discover new medicinal plants. Gardening is
the most popular leisure activity in the U.S. Working with plants or horticulture therapy is beneficial for
rehabilitating people with disabilities. Certain plants contain psychotropic chemicals which are extracted and
ingested, including tobacco, cannabis (marijuana), and opium.
Negative effects
Weeds are plants that grow where people do not want them. People have spread plants beyond their
native ranges and some of these introduced plants become invasive, damaging existing ecosystems by
displacing native species. Invasive plants cause billions of dollars in crop losses annually by displacing crop
plants, they increase the cost of production and the use of chemical means to control them affects the
environment.
Plants may cause harm to people and animals. Plants that produce windblown pollen invoke allergic
reactions in people who suffer from hay fever. A wide variety of plants are poisonous to people and/or
animals. Toxalbumins are plant poisons fatal to most mammals and act as a serious deterrent to
consumption. Several plants cause skin irritations when touched, such as poison ivy. Certain plants contain
psychotropic chemicals, which are extracted and ingested or smoked, including tobacco, cannabis
(marijuana), cocaine and opium. Smoking causes damage to health or even death, while some drugs may
also be harmful or fatal to people[25][26]. Both illegal and legal drugs derived from plants may have negative
effects on the economy, affecting worker productivity and law enforcement costs.[27][28] Some plants cause
allergic reactions in people and animals when ingested, while other plants cause food intolerances that
negatively affect health.
See also
Plants portal
• The Plant List

• Biosphere
• Leaf sensor
• Plant defense against herbivory
• Plant perception (paranormal)
• Plant perception (physiology)
• Plant identification
• Rapid plant movement
• Germination
References
1. ^ Haeckel G (1866). Generale Morphologie der Organismen. Berlin: Verlag von Georg
Reimer. pp. vol.1: i–xxxii, 1–574, pls I–II; vol. 2: i–clx, 1–462, pls I–VIII.
2. ^ "plant[2 - Definition from the Merriam-Webster Online Dictionary"]. http://www.merriam-
webster.com/dictionary/plant%5B2%5D. Retrieved 2009-03-25.
3. ^ "plant (life form) -- Britannica Online Encyclopedia".
http://www.britannica.com/EBchecked/topic/463192/plant. Retrieved 2009-03-25.
4. ^ Margulis, L. (1974). "Five-kingdom classification and the origin and evolution of cells".
Evolutionary Biology 7: 45–78.
5. ^ a b c d e Raven, Peter H., Ray F. Evert, & Susan E. Eichhorn, 2005. Biology of Plants, 7th
edition. (New York: W. H. Freeman and Company). ISBN 0-7167-1007-2.
6. ^ Bremer, K. (1985). "Summary of green plant phylogeny and classification". Cladistics 1:
369–385. doi:10.1111/j.1096-0031.1985.tb00434.x.
7. ^ Mishler, Brent D.; S. P. Churchill (1985). "Transition to a land flora: phylogenetic
relationships of the green algae and bryophytes". Cladistics 1: 305–328. doi:10.1111/j.1096-
0031.1985.tb00431.x.
8. ^ Mishler, Brent D.; Louise A. Lewis; Mark A. Buchheim; Karen S. Renzaglia; D. J. Garbary;
Carl F. Delwiche; F. W. Zechman; T. S. Kantz; & Ron L. Chapman (1994). "Phylogenetic
relationships of the "green algae" and "bryophytes"". Annals of the Missouri Botanical Garden
(Annals of the Missouri Botanical Garden, Vol. 81, No. 3) 81 (3): 451–483. doi:10.2307/2399900.
9. ^ Van den Hoek, C., D. G. Mann, & H. M. Jahns, 1995. Algae: An Introduction to Phycology .
pages 343, 350, 392, 413, 425, 439, & 448 (Cambridge: Cambridge University Press). ISBN 0-521-
30419-9
10.^ Van den Hoek, C., D. G. Mann, & H. M. Jahns, 1995. Algae: An Introduction to Phycology .
pages 457, 463, & 476. (Cambridge: Cambridge University Press). ISBN 0-521-30419-9
11.^ Crandall-Stotler, Barbara. & Stotler, Raymond E., 2000. "Morphology and classification of
the Marchantiophyta". page 21 in A. Jonathan Shaw & Bernard Goffinet (Eds.), Bryophyte Biology.
(Cambridge: Cambridge University Press). ISBN 0-521-66097-1
12.^ Schuster, Rudolf M., The Hepaticae and Anthocerotae of North America , volume VI, pages
712-713. (Chicago: Field Museum of Natural History, 1992). ISBN 0-914-86821-7.
13.^ Goffinet, Bernard; William R. Buck (2004). "Systematics of the Bryophyta (Mosses): From
molecules to a revised classification". Monographs in Systematic Botany (Missouri Botanical Garden
Press) 98: 205–239.
14.^ Gifford, Ernest M. & Adriance S. Foster, 1988. Morphology and Evolution of Vascular
Plants, 3rd edition, page 358. (New York: W. H. Freeman and Company). ISBN 0-7167-1946-0.
15.^ Taylor, Thomas N. & Edith L. Taylor, 1993. The Biology and Evolution of Fossil Plants,
page 636. (New Jersey: Prentice-Hall). ISBN 0-13-651589-4.
16.^ International Union for Conservation of Nature and Natural Resources, 2006. IUCN Red
List of Threatened Species:Summary Statistics
17.^ Kenrick, Paul & Peter R. Crane. 1997. The Origin and Early Diversification of Land Plants:
A Cladistic Study. (Washington, D.C.: Smithsonian Institution Press). ISBN 1-56098-730-8.
18.^ Smith, Alan R., Kathleen M. Pryer, E. Schuettpelz, P. Korall, H. Schneider, & Paul G. Wolf.
(2006). "A classification for extant ferns". Taxon 55(3): 705-731.
19.^ a b Robbins, W.W., Weier, T.E., et al., Botany:Plant Science, 3rd edition , Wiley
International, New York, 1965.
20.^ Goyal, K., Walton, L. J., & Tunnacliffe, A. (2005). "LEA proteins prevent protein aggregation
due to water stress". Biochemical Journal 388 (Part 1): 151 – 157. doi:10.1042/BJ20041931.
PMID 15631617. PMC 1186703. Archived from the original on 2009-08-03.
http://www.webcitation.org/5il9QhYT0.
21.^ a b Campbell, Reece, Biology, 7th edition, Pearson/Benjamin Cummings, 2005.
22.^ BBC Report
23.^ Howard Frank, Bromeliad Phytotelmata, October 2000
24.^ Barthlott, W., S. Porembski, R. Seine, and I. Theisen. 2007. The Curious World of
Carnivorous Plants: A Comprehensive Guide to Their Biology and Cultivation. Timber Press:
Portland, Oregon.
25.^ "cocaine/crack". http://www.urban75.com/Drugs/drugcoke.html.
26.^ "Deaths related to cocaine". http://ar2005.emcdda.europa.eu/en/page050-en.html.
27.^ "Illegal drugs drain $160 billion a year from American economy". Archived from the original
on 2008-02-15.
http://web.archive.org/web/20080215071055/http://www.whitehousedrugpolicy.gov/NEWS/press02/0
12302.html.
28.^ "The social cost of illegal drug consumption in Spain".
http://www.ingentaconnect.com/content/bsc/add/2002/00000097/00000009/art00012.
Further reading
General
• Evans, L. T. (1998). Feeding the Ten Billion - Plants and Population Growth. Cambridge
University Press. Paperback, 247 pages. ISBN 0-521-64685-5.
• Kenrick, Paul & Crane, Peter R. (1997). The Origin and Early Diversification of Land Plants:
A Cladistic Study. Washington, D. C.: Smithsonian Institution Press. ISBN 1-56098-730-8.
• Raven, Peter H., Evert, Ray F., & Eichhorn, Susan E. (2005). Biology of Plants (7th ed.).
New York: W. H. Freeman and Company. ISBN 0-7167-1007-2.
• Taylor, Thomas N. & Taylor, Edith L. (1993). The Biology and Evolution of Fossil Plants.
Englewood Cliffs, NJ: Prentice Hall. ISBN 0-13-651589-4.
• Trewavas, A. (2003). Aspects of Plant Intelligence, Annals of Botany 92: 1-20.
Species estimates and counts
• International Union for Conservation of Nature and Natural Resources (IUCN) Species
Survival Commission (2004). IUCN Red List [1].
• Prance, G. T. (2001). Discovering the Plant World. Taxon 50: 345-359.
External links
Find more about Plants on Wikipedia's sister projects:
Species directories Plantae from Wikispecies

Plantae
• Chaw, S.-M. et al. (1997). "Molecular Phylogeny of Extant Gymnosperms and Seed Plant
Evolution: Analysis of Nuclear 18s rRNA Sequences". Molec. Biol. Evol. 14 (1): 56–68.
PMID 9000754. http://mbe.library.arizona.edu/data/1997/1401/7chaw.pdf.
• Index Nominum Algarum
• Interactive Cronquist classification
• Plant Photo Gallery of Japan - Flavon's Wild herb and Alpine plants
• Plant Picture Gallery
• Plant Resources of Tropical Africa
• www.prota.org - PROTA’s mission
• Tree of Life
Botanical and vegetation databases
• African Plants Initiative database
• Australia
• Chilean plants at Chilebosque
• Dave's garden plenty of information mostly about garden plants
• e-Floras (Flora of China, Flora of North America and others)
• Flora Europaea
• Flora of Central Europe (German)
• Flora of North America
• List of Japanese Wild Plants Online
• Meet the Plants-National Tropical Botanical Garden
• Lady Bird Johnson Wildflower Center - Native Plant Information Network at University of
Texas, Austin
• The Plant List
• United States Department of Agriculture not limited to continental US species
[hide]v · d · eBotany
Subdisciplines of Ethnobotany · Paleobotany · Plant anatomy ·

botany Plant ecology · Plant evo-devo · Plant morphology ·
Plant physiology
Plants Evolutionary history of plants · Algae ·

Bryophyte · Pteridophyte · Gymnosperm · Angiosperm
Plant parts Flower · Fruit · Leaf · Meristem · Root · Stem ·

Stoma · Vascular tissue · Wood
Cell wall · Chlorophyll · Chloroplast ·

Plant cells Photosynthesis · Plant hormone · Plastid ·
Transpiration
Plant Alternation of generations · Gametophyte ·

reproduction Plant sexuality · Pollen · Pollination · Seed · Spore ·
Sporophyte
Plant taxonomy Botanical name · Botanical nomenclature ·

Herbarium · IAPT · ICBN · Species Plantarum
Glossaries Glossary of botanical terms · Glossary of plant

morphology terms
Category · Portal
[hide]v · d · eClassification of Archaeplastida / Plantae sensu lato
Rhodophyta/ Cyanidiophyceae · Porphyridiophyceae · Compsopogonophyceae · Stylonematophy

Rhodophyceae Bangiophyceae · Florideophyceae (Hildenbrandiales, Acrochaetiales, Nemaliales, Batracho
Gelidiales, Gracilariales, Ceramiales)
Glaucocystophyceae Glaucocystis · Cyanophora · Gloeochaete
Viridiplantae/ Chlorophyta/ Prasinophyceae

Plantae GA
sensu stricto UTC clade: Ulvophyceae · Trebouxiophyceae · Chlorophycea
Charophyta/G Charales · Coleochaetales · Desmidia
A Mesostigmatales · Zygnematales
Bryophytes Marchantiophy
(non-vascular) Bryophyta "Moss" · Ho
Lycopodiophy
Streptophyta
Embryophyta/
Plantae
sensu strictissimo
Tracheophyta
Euphyllophyta
See also: list of plant orders
[hide]v · d · eEukaryota
Domain : Archaea · Bacteria · Eukaryota
Archaeplastida, or Plantae Viridiplantae/Plantae sens

sensu lato Rhodophyta · Glaucocystophyceae
AH
Hacrobia, or non-SAR
Haptophyta · Cryptophyta
chromalveolata
AH/SAR Heterokont
Bikonta Ochrophyta · Bigyra
("S")
Halvaria
SAR Ciliates · Myzozoa (A
Alveolata
Dinoflagellata)
Rhizaria Cercozoa · Retaria (Foraminifera, Radiolaria)
Excavata Discoba (Euglenozoa, Percolozoa) · Metamonad · Malawimonas

Unikonta Apusomonadida (Apusomonas, Amastigomonas) · Ancyromonadida (Anc
Apusozoa
Hemimastigida (Hemimastix, Spironema, Stereonema)
Amoebozoa Lobosea · Conosa · Phalansterium · Breviata
Opisthokonta Mesomycetozoea Dermocystida · Ichthyop
Filasterea
Minis
Holozoa Choanoflagellatea
Filozoa
(Bila
Metazoa
Cten
or "Animalia"
Mes
(Plac
Holomycota Dikarya (Ascomycota, Basidio

Fungi Glomeromycota · Zygomycota · Blast
Chytridiomycota/Neocallimastigomyc
Nuclearia · Micronuclearia · R
Nucleariidae
Pinaciophora · Pompholyxophrys · Fo
[hide]v · d · eElements of nature

Category · Portal
Retrieved from "http://en.wikipedia.org/wiki/Plant"
Categories: Plants | Plant taxonomy

W000
Paleontology

"Palaeontology" redirects here. For the scientific journal, see Palaeontology (journal).
Paleontology investigates the entire history of life on Earth.
A paleontologist at work at John Day Fossil Beds National Monument

The preparation of the fossilized bones of Europasaurus holgeri
Paleontology ( /ˌpælɪɒnˈtɒlədʒi/; British: palaeontology)[note 1] is the study of prehistoric life,
including organisms' evolution and interactions with each other and their environments (their paleoecology).
As a "historical science" it tries to explain causes rather than conduct experiments to observe effects.
Paleontological observations have been documented as far back as the 5th century BC. The science became
established in the 18th century as a result of Georges Cuvier's work on comparative anatomy, and developed
rapidly in the 19th century. Fossils found in China since the 1990s have provided new information about the
earliest evolution of animals, early fish, dinosaurs and the evolution of birds and mammals. Paleontology lies
on the border between biology and geology, and shares with archaeology a border that is difficult to define. It
now uses techniques drawn from a wide range of sciences, including biochemistry, mathematics and
engineering. As knowledge has increased, paleontology has developed specialized subdivisions, some of
which focus on different types of fossil organisms while others study ecological and environmental history,
such as ancient climates.
Body fossils and trace fossils are the principal types of evidence about ancient life, and geochemical
evidence has helped to decipher the evolution of life before there were organisms large enough to leave
fossils. Estimating the dates of these remains is essential but difficult: sometimes adjacent rock layers allow
radiometric dating, which provides absolute dates that are accurate to within 0.5%, but more often
paleontologists have to rely on relative dating by solving the "jigsaw puzzles" of biostratigraphy. Classifying
ancient organisms is also difficult, as many do not fit well into the Linnean taxonomy that is commonly used
for classifying living organisms, and paleontologists more often use cladistics to draw up evolutionary "family
trees". The final quarter of the 20th century saw the development of molecular phylogenetics, which
investigates how closely organisms are related by measuring how similar the DNA is in their genomes.
Molecular phylogenetics has also been used to estimate the dates when species diverged, but there is
controversy about the reliability of the molecular clock on which such estimates depend.
Use of all these techniques has enabled paleontologists to discover much of the evolutionary history
of life, almost all the way back to when Earth became capable of supporting life, about 3,800 million years
ago. For about half of that time the only life was single-celled micro-organisms, mostly in microbial mats that
formed ecosystems only a few millimeters thick. Earth's atmosphere originally contained virtually no oxygen,
and its oxygenation began about 2,400 million years ago. This may have caused an accelerating increase in
the diversity and complexity of life, and early multicellular plants and fungi have been found in rocks dated
from 1,700 to 1,200 million years ago. The earliest multicellular animal fossils are much later, from about 580
million years ago, but animals diversified very rapidly and there is a lively debate about whether most of this
happened in a relatively short Cambrian explosion or started earlier but has been hidden by lack of fossils. All
of these organisms lived in water, but plants and invertebrates started colonizing land from about 490 million
years ago and vertebrates followed them about 370 million years ago. The first dinosaurs appeared about
230 million years ago and birds evolved from one dinosaur group about 150 million years ago. During the
time of the dinosaurs, mammals' ancestors survived only as small, mainly nocturnal insectivores, but after
the non-avian dinosaurs became extinct in the Cretaceous–Tertiary extinction event 65 million years ago
mammals diversified rapidly. Flowering plants appeared and rapidly diversified between 130 million years
ago and 90 million years ago, possibly helped by coevolution with pollinating insects. Social insects
appeared around the same time and, although they have relatively few species, now form over 50% of the
total mass of all insects. Humans evolved from a lineage of upright-walking apes whose earliest fossils date
from over 6 million years ago, and anatomically modern humans appeared under 200,000 years ago. The
course of evolution has been changed several times by mass extinctions that wiped out previously dominant
groups and allowed other to rise from obscurity to become major components of ecosystems.
Contents
[hide]
• 1 Definition
• 1.1 A historical science
• 1.2 Related sciences
• 1.3 Subdivisions
• 2 Sources of evidence
• 2.1 Body fossils
• 2.2 Trace fossils
• 2.3 Geochemical observations
• 3 Classifying ancient organisms
• 4 Estimating the dates of organisms
• 5 Overview of the history of life
• 6 Mass extinctions
• 7 History of paleontology
• 8 See also
• 9 Notes
• 10 References
[edit] Definition
A paleontologist carefully chips rock from a column of dinosaur vertebrae.

The simplest definition is "the study of ancient life".[1] Paleontology seeks information about several
aspects of past organisms: "their identity and origin, their environment and evolution, and what they can tell
us about the Earth's organic and inorganic past".[2]
[edit] A historical science

Paleontology is one of the historical sciences, along with archaeology, geology, biology, astronomy,
cosmology, philology and history itself.[3] This means that it aims to describe phenomena of the past and
reconstruct their causes.[4] Hence it has three main elements: description of the phenomena; developing a
general theory about the causes of various types of change; and applying those theories to specific facts.[3]
When trying to explain past phenomena, paleontologists and other historical scientists often
construct a set of hypotheses about the causes and then look for a "smoking gun", a piece of evidence which
indicates that one of the hypotheses is a better explanation than the others. Sometimes the "smoking gun" is
discovered by a fortunate accident during other research, for example the discovery by Luis Alvarez and
Walter Alvarez of an iridium-rich layer at the Cretaceous–Tertiary boundary made asteroid impact and
volcanism the most favored explanations for the Cretaceous–Tertiary extinction event.[4]
The other main type of science is experimental science, which is often said to work by conducting
experiments to disprove hypotheses about the workings and causes of natural phenomena – note that this
approach cannot prove a hypothesis is correct, since some later experiment may disprove it. However, when
confronted with totally unexpected phenomena, such as the first evidence for invisible radiation, experimental
scientists often use the same approach as historical scientists: construct a set of hypotheses about the
causes and then look for a "smoking gun".[4]
[edit] Related sciences
Paleontology lies on the boundary between biology and geology since paleontology focuses on the
record of past life but its main source of evidence is fossils, which are found in rocks.[5] For historical
reasons paleontology is part of the geology departments of many universities, because in the 19th century
and early 20th century geology departments found paleontological evidence important for estimating the
ages of rocks while biology departments showed little interest.[6]
Paleontology also has some overlap with archaeology, which primarily works with objects made by
humans and with human remains, while paleontologists are interested in the characteristics and evolution of
humans as organisms. When dealing with evidence about humans, archaeologists and paleontologists may
work together – for example paleontologists might identify animal or plant fossils around an archaeological
site, to discover what the people who lived there ate; or they might analyze the climate at the time when the
site was inhabited by humans.[7]
Analyses using engineering techniques show that Tyrannosaurus had a devastating bite, but raise
doubts about how fast it could move.
In addition paleontology often uses techniques derived from other sciences, including biology,
ecology, chemistry, physics and mathematics.[1] For example geochemical signatures from rocks may help
to discover when life first arose on Earth,[8] and analyses of carbon isotope ratios may help to identify
climate changes and even to explain major transitions such as the Permian–Triassic extinction event.[9] A
relatively recent discipline, molecular phylogenetics, often helps by using comparisons of different modern
organisms' DNA and RNA to re-construct evolutionary "family trees"; it has also been used to estimate the
dates of important evolutionary developments, although this approach is controversial because of doubts
about the reliability of the "molecular clock".[10] Techniques developed in engineering have been used to
analyse how ancient organisms might have worked, for example how fast Tyrannosaurus could move and
how powerful its bite was.[11][12]
Paleontology even contributes to astrobiology, the investigation of possible life on other planets, by
developing models of how life may have arisen and by providing techniques for detecting evidence of life.[13]
[edit] Subdivisions
As knowledge has increased, paleontology has developed specialised subdivisons.[14] Vertebrate
paleontology concentrates on fossils of vertebrates, from the earliest fish to the immediate ancestors of
modern mammals. Invertebrate paleontology deals with fossils of invertebrates such as molluscs,
arthropods, annelid worms and echinoderms. Paleobotany focuses on the study of fossil plants, but
traditionally includes the study of fossil algae and fungi. Palynology, the study of pollen and spores produced
by land plants and protists, straddles the border between paleontology and botany, as it deals with both living
and fossil organisms. Micropaleontology deals with all microscopic fossil organisms, regardless of the group
to which they belong.[15]
In the Carboniferous period, the continents were not in the same places as they are today, and there
was extensive glaciation.
Instead of focusing on individual organisms, paleoecology examines the interactions between
different organisms, such as their places in food chains, and the two-way interaction between organisms and
their environment[16] – for example the development of oxygenic photosynthesis by bacteria hugely
increased the productivity and diversity of ecosystems,[17] and also caused the oxygenation of the
atmosphere, which in turn was a prerequisite for the evolution of the most complex eucaryotic cells, from
which all multicellular organisms are built.[18] Paleoclimatology, although sometimes treated as part of
paleoecology,[15] focuses more on the history of Earth's climate and the mechanisms which have changed
it[19] – which have sometimes included evolutionary developments, for example the rapid expansion of land
plants in the Devonian period removed more carbon dioxide from the atmosphere, reducing the greenhouse
effect and thus helping to cause an ice age in the Carboniferous period.[20]
Biostratigraphy, the use of fossils to work out the chronological order in which rocks were formed, is
useful to both paleontologists and geologists.[21] Biogeography studies the spatial distribution of organisms,
and is also linked to geology, which explains how Earth's geography has changed over time.[22]
[edit] Sources of evidence
[edit] Body fossils
This Marrella specimen illustrates how clear and detailed the fossils from the Burgess Shale
lagerstätte are.
Fossils of organisms' bodies are usually the most informative type of evidence. The most common
types are wood, bones, and shells.[23] Fossilisation is a rare event, and most fossils are destroyed by
erosion or metamorphism before they can be observed. Hence the fossil record is very incomplete,
increasingly so further back in time. Despite this, it is often adequate to illustrate the broader patterns of life's
history.[24] There are also biases in the fossil record: different environments are more favorable to the
preservation of different types of organism or parts of organisms.[25] Further, only the parts of organisms that
were already mineralised are usually preserved, such as the shells of molluscs. Since most animal species
are soft-bodied, they decay before they can become fossilised. As a result, although there are 30-plus phyla
of living animals, two-thirds have never been found as fossils.[26]
Occasionally, unusual environments may preserve soft tissues. These lagerstätten allow
paleontologists to examine the internal anatomy of animals that in other sediments are represented only by
shells, spines, claws, etc. – if they are preserved at all. However, even lagerstätten present an incomplete
picture of life at the time. The majority of organisms living at the time are probably not represented because
lagerstätten are restricted to a narrow range of environments, e.g. where soft-bodied organisms can be
preserved very quickly by events such as mudslides; and the exceptional events that cause quick burial
make it difficult to study the normal environments of the animals.[27] The sparseness of the fossil record
means that organisms are expected to exist long before and after they are found in the fossil record – this is
known as the Signor-Lipps effect.[28]
[edit] Trace fossils
Cambrian trace fossils including Rusophycus, made by a trilobite

Main article: Trace fossil
Trace fossils consist mainly of tracks and burrows, but also include coprolites (fossil feces) and
marks left by feeding.[23][29] Trace fossils are particularly significant because they represent a data source
that is not limited to animals with easily-fossilized hard parts, and which reflects organisms' behaviour. Also
many traces date from significantly earlier than the body fossils of animals that are thought to have been
capable of making them.[30] Whilst exact assignment of trace fossils to their makers is generally impossible,
traces may for example provide the earliest physical evidence of the appearance of moderately complex
animals (comparable to earthworms).[29]
[edit] Geochemical observations
Main article: Geochemistry
Geochemical observations may help to deduce the global level of biological activity, or the affinity of
a certain fossil. For example geochemical features of rocks may reveal when life first arose on Earth,[8] and
may provide evidence of the presence of eucaryotic cells, the type from which all multicellular organisms are
built.[31] Analyses of carbon isotope ratios may help to explain major transitions such as the Permian–
Triassic extinction event.[9]
[edit] Classifying ancient organisms

Main articles: Biological classification , Cladistics , Phylogenetic nomenclature , and Evolutionary
taxonomy
Tetrapods
Amphibians
Amniotes
Synapsids
Extinct Synapsids
Mammals
Reptiles
Extinct
reptiles
Lizards and
snakes
Archosaurs
?
Extinct
Archosaurs
Crocodilians
Dinosaurs
?
Extinct
Dinosaurs
? Birds
Simple example cladogram

Warm-bloodedness evolved somewhere in the
synapsid–mammal transition.
? Warm-bloodedness must also have evolved at one of
these points – an example of convergent evolution.[32]
Levels in the Linnean taxonomy
Naming groups of organisms in a way that is clear and widely agreed is important, as some disputes
in palaeontology have been based just on misunderstandings over names.[33] Linnean taxonomy is
commonly used for classifying living organisms, but runs into difficulties when dealing with newly-discovered
organisms that are significantly different from known ones. For example: it is hard to decide at what level to
place a new higher-level grouping, e.g. genus or family or order; this is important since the Linnean rules for
naming groups are tied to their levels, and hence if a group is moved to a different level it has to be renamed.
[34]
Paleontologists generally use approaches based on cladistics, a technique for working out the
evolutionary "family tree" of a set of organisms.[33] It works by the logic that, if groups B and C have more
similarities to each other than either has to group A, then B and C are more closely related to each other than
either is to A. Characters that are compared may be anatomical, such as the presence of a notochord, or
molecular, by comparing sequences of DNA or proteins. The result of a successful analysis is a hierarchy of
clades – groups that share a common ancestor. Ideally the "family tree" has only two branches leading from
each node ("junction"), but sometimes there is too little information to achieve this and paleontologists have
to make do with junctions that have several branches. The cladistic technique is sometimes fallible, as some
features, such as wings or camera eyes, evolved more than once, convergently – this must be taken into
account in analyses.[32]
Evolutionary developmental biology, commonly abbreviated to "Evo Devo", also helps
paleontologists to produce "family trees". For example the embryological development of some modern
brachiopods suggests that brachiopods may be descendants of the halkieriids, which became extinct in the
Cambrian period.[35]
[edit] Estimating the dates of
organisms
Paleontology seeks to map out how living
things have changed through time. A substantial
hurdle to this aim is the difficulty of working out
how old fossils are. Beds which preserve fossils
typically lack the radioactive elements needed for
radiometric dating. This technique is our only
means of giving rocks greater than about 50
million years old an absolute age, and can be
accurate to within 0.5% or better.[36] Although
radiometric dating requires very careful laboratory
work, its basic principle is simple: the rates at Permian
which various radioactive elements decay are
known, and so the ratio of the radioactive element Missis-
to the element into which it decays shows how sippian
long ago the radioactive element was incorporated
into the rock. Radioactive elements are common Pennsyl-
only in rocks with a volcanic origin, and so the vanian
only fossil-bearing rocks that can be dated
Devo-
radiometrically are a few volcanic ash layers.[36]
nian
Consequently, paleontologists must
usually rely on stratigraphy to date fossils. Silurian
Ordo-
vician
Camb-
rian
Pecten gibbus
Stratigraphy is the science of deciphering the "layer-cake" that is the sedimentary record, and has been
compared to a jigsaw puzzle.[37] Rocks normally form relatively horizontal layers, with each layer younger
than the one underneath it. If a fossil is found between two layers whose ages are known, the fossil's age
must lie between the two known ages.[38] Because rock sequences are not continuous, but may be broken
up by faults or periods of erosion, it is very difficult to match up rock beds that are not directly next to one
another. However, fossils of species that survived for a relatively short time can be used to link up isolated
rocks: this technique is called biostratigraphy. For instance, the conodont Eoplacognathus pseudoplanus has
a short range in the Middle Ordovician period.[39] If rocks of unknown age are found to have traces of E.
pseudoplanus, they must have a mid-Ordovician age. Such index fossils must be distinctive, be globally
distributed and have a short time range to be useful. However, misleading results are produced if the index
fossils turn out to have longer fossil ranges than first thought.[40] Stratigraphy and biostratigraphy can in
general provide only relative dating (A was before B), which is often sufficient for studying evolution.
However, this is difficult for some time periods, because of the problems involved in matching up rocks of the
same age across different continents.[41]
Family-tree relationships may also help to narrow down the date when lineages first appeared. For
instance, if fossils of B or C date to X million years ago and the calculated "family tree" says A was an
ancestor of B and C, then A must have evolved more than X million years ago.
It is also possible to estimate how long ago two living clades diverged – i.e. approximately how long
ago their last common ancestor must have lived – by assuming that DNA mutations accumulate at a constant
rate. These "molecular clocks", however, are fallible, and provide only a very approximate timing: for
example, they are not sufficiently precise and reliable for estimating when the groups that feature in the
Cambrian explosion first evolved,[42] and estimates produced by different techniques may vary by a factor of
two.[10]
[edit] Overview of the history of life
The evolutionary history of life stretches back to over 3,000 million years ago, possibly as far as
3,800 million years ago. Earth formed about 4,540 million years ago and, after a collision that formed the
Moon about 40 million years later, may have cooled quickly enough to have oceans and an atmosphere
about 4,440 million years ago.[43] However there is evidence on the Moon of a Late Heavy Bombardment
from 4,000 to 3,800 million years ago. If, as seem likely, such a bombardment struck Earth at the same time,
the first atmosphere and oceans may have been stripped away.[44] The oldest clear evidence of life on Earth
dates to 3,000 million years ago, although there have been reports, often disputed, of fossil bacteria from
3,400 million years ago and of geochemical evidence for the presence of life 3,800 million years ago.[8][45]
Even the simplest modern organisms are too complex to have emerged directly from non-living materials.[46]
Some scientists have proposed that life on Earth was "seeded" from elsewhere,[47] but most research
concentrates on various explanations of how life could have arisen independently on Earth.[48]
This wrinkled "elephant skin" texture is a trace fossil of a non-stromatolite microbial mat.
The image shows the location, in the Burgsvik beds of Sweden, where the texture was first identified as
evidence of a microbial mat.[49]
For about 2,000 million years microbial mats, multi-layered colonies of different types of bacteria,
were the dominant life on Earth.[50] The evolution of oxygenic photosynthesis enabled them to play the
major role in the oxygenation of the atmosphere[51] from about 2,400 million years ago. This change in the
atmosphere increased their effectiveness as nurseries of evolution.[52] While eukaryotes, cells with complex
internal structures, may have been present earlier, their evolution speeded up when they acquired the ability
to transform oxygen from a poison to a powerful source of energy in their metabolism. This innovation may
have come from primitive eukaryotes capturing oxygen-powered bacteria as endosymbionts and
transforming them into organelles called mitochondria.[53] The earliest evidence of complex eukaryotes with
organelles such as mitochondria, dates from 1,850 million years ago.[18]
Multicellular life is composed only of eukaryotic cells, and the earliest evidence for it is the
Francevillian Group Fossils from 2,100 million years ago,[54] although specialization of cells for different
functions first appears between 1,430 million years ago (a possible fungus) and 1,200 million years ago (a
probable red alga). Sexual reproduction may be a prerequisite for specialization of cells, as an asexual
multicellular organism might be at risk of being taken over by rogue cells that retain the ability to reproduce.
[55][56]
Opabinia made the largest single contribution to modern interest in the Cambrian explosion.
The earliest known animals are cnidarians from about 580 million years ago, but these are so
modern-looking that the earliest animals must have appeared before then.[57] Early fossils of animals are
rare because they did not develop mineralized hard parts that fossilize easily until about 548 million years
ago.[58] The earliest modern-looking bilaterian animals appear in the Early Cambrian, along with several
"weird wonders" that bear little obvious resemblance to any modern animals. There is a long-running debate
about whether this Cambrian explosion was truly a very rapid period of evolutionary experimentation;
alternative views are that modern-looking animals began evolving earlier but fossils of their precursors have
not yet been found, or that the "weird wonders" are evolutionary "aunts" and "cousins" of modern groups.[59]
Vertebrates remained an obscure group until the first fish with jaws appeared in the Late Ordovician.[60][61]
The spread of life from water to land required organisms to solve several problems, including
protection against drying out and supporting themselves against gravity.[62][63] The earliest evidence of land
plants and land invertebrates date back to about 476 million years ago and 490 million years ago
respectively.[63][64] The lineage that produced land vertebrates evolved later but very rapidly between 370
million years ago and 360 million years ago;[65] recent discoveries have overturned earlier ideas about the
history and driving forces behind their evolution.[66] Land plants were so successful that they caused an
ecological crisis in the Late Devonian, until the evolution and spread of fungi that could digest dead wood.[20]
At about 13 centimetres (5.1 in) the Early

Cretaceous Yanoconodon was longer than the average
mammal of the time.[67]
Birds are the last surviving dinosaurs.

[68]
During the Permian period synapsids, including the ancestors of mammals, may have dominated
land environments,[69] but the Permian–Triassic extinction event 251 million years ago came very close to
wiping out complex life.[70] During the slow recovery from this catastrophe a previously obscure group,
archosaurs, became the most abundant and diverse terrestrial vertebrates. One archosaur group, the
dinosaurs, were the dominant land vertebrates for the rest of the Mesozoic,[71] and birds evolved from one
group of dinosaurs.[68] During this time mammals' ancestors survived only as small, mainly nocturnal
insectivores, but this apparent set-back may have accelerated the development of mammalian traits such as
endothermy and hair.[72] After the Cretaceous–Tertiary extinction event 65 million years ago killed off the
non-avian dinosaurs – birds are the only surviving dinosaurs – mammals increased rapidly in size and
diversity, and some took to the air and the sea.[73][74][75]
A modern social insect collects pollen from a modern flowering plant.

Fossil evidence indicates that flowering plants appeared and rapidly diversified in the Early
Cretaceous, between 130 million years ago and 90 million years ago.[76] Their rapid rise to dominance of
terrestrial ecosystems is thought to have been propelled by coevolution with pollinating insects.[77] Social
insects appeared around the same time and, although they account for only small parts of the insect "family
tree", now form over 50% of the total mass of all insects.[78]
Humans evolved from a lineage of upright-walking apes whose earliest fossils date from over 6
million years ago.[79] Although early members of this lineage had chimp-sized brains, about 25% as big as
modern humans', there are signs of a steady increase in brain size after about 3 million years ago.[80] There
is a long-running debate about whether modern humans are descendants of a single small population in
Africa, which then migrated all over the world less than 200,000 years ago and replaced previous hominine
species, or arose worldwide at the same time as a result of interbreeding.[81]

Main article: Mass extinction
Life on earth has suffered occasional
mass extinctions at least since 542 million years
ago. Although they are disasters at the time, mass
extinctions have sometimes accelerated the
evolution of life on earth. When dominance of
particular ecological niches passes from one
group of organisms to another, it is rarely because
the new dominant group is "superior" to the old
and usually because an extinction event
eliminates the old dominant group and makes way
for the new one.[82][83]
The fossil record appears to show that the
rate of extinction is slowing down, with both the
gaps between mass extinctions becoming longer
and the average and background rates of
extinction decreasing. However, it is not certain
Apparent extinction intensity, i.e. the fraction of

genera going extinct at any given time, as reconstructed
from the fossil record (graph not meant to include recent
epoch of Holocene extinction event)
whether the actual rate of extinction has altered, since both of these observations could be explained in
several ways:[84]
• The oceans may have become more hospitable to life over the last 500 million years and less
vulnerable to mass extinctions: dissolved oxygen became more widespread and penetrated to
greater depths; the development of life on land reduced the run-off of nutrients and hence the risk of
eutrophication and anoxic events; marine ecosystems became more diversified so that food chains
were less likely to be disrupted.[85][86]
• Reasonably complete fossils are very rare, most extinct organisms are represented only by
partial fossils, and complete fossils are rarest in the oldest rocks. So paleontologists have mistakenly
assigned parts of the same organism to different genera which were often defined solely to
accommodate these finds – the story of Anomalocaris is an example of this.[87] The risk of this
mistake is higher for older fossils because these are often unlike parts of any living organism. Many
of the "superfluous" genera are represented by fragments which are not found again and the
"superfluous" genera appear to become extinct very quickly.[84]
Biodiversity in the fossil record, which is
"the number of distinct genera
alive at any given time; that is, those
whose first occurrence predates and
whose last occurrence postdates that
time"[88]
shows a different trend: a fairly swift rise from
542 to 400 million years ago, a slight decline from
400 to 200 million years ago, in which the devastating
Permian–Triassic extinction event is an important factor,
and a swift rise from 200 million years ago to the
Thousands of genera
present.[88]
Phanerozoic biodiversity as shown by the

fossil record
This illustration of an Indian elephant jaw and a mammoth jaw (top) is from Cuvier's 1796 paper on
living and fossil elephants.
[edit] History of paleontology

Main article: History of paleontology
Although paleontology became established around 1800, earlier thinkers had noticed aspects of the
fossil record. The ancient Greek philosopher Xenophanes (570–480 BC) concluded from fossil sea shells that
some areas of land were once under water.[89] During the Middle Ages the Persian naturalist Ibn Sina,
known as Avicenna in Europe, discussed fossils and proposed a theory of petrifying fluids on which Albert of
Saxony elaborated in the 14th century.[90] The Chinese naturalist Shen Kuo (1031–1095) proposed a theory
of climate change based on the presence of petrified bamboo in regions that in his time were too dry for
bamboo.[91]
In early modern Europe, the systematic study of fossils emerged as an integral part of the changes in
natural philosophy that occurred during the Age of Reason. At the end of the 18th century Georges Cuvier's
work established comparative anatomy as a scientific discipline and, by proving that some fossil animals
resembled no living ones, demonstrated that animals could become extinct, leading to the emergence of
paleontology.[92] The expanding knowledge of the fossil record also played an increasing role in the
development of geology, particularly stratigraphy.[93]
The first half of the 19th century saw geological and paleontological activity become increasingly well
organized with the growth of geologic societies and museums[94][95] and an increasing number of
professional geologists and fossil specialists. Interest increased for reasons that were not purely scientific, as
geology and paleontology helped industrialists to find and exploit natural resources such as coal.[96]
This contributed to a rapid increase in knowledge about the history of life on Earth and to progress in
the definition of the geologic time scale, largely based on fossil evidence. In 1822 Henri Marie Ducrotay de
Blanville, editor of Journal de Phisique, coined the word "paleontology" to refer to the study of ancient living
organisms through fossils.[97] As knowledge of life's history continued to improve, it became increasingly
obvious that there had been some kind of successive order to the development of life. This encouraged early
evolutionary theories on the transmutation of species.[98] After Charles Darwin published Origin of Species
in 1859, much of the focus of paleontology shifted to understanding evolutionary paths, including human
evolution, and evolutionary theory.[98]
Haikouichthys, from about 518 million years ago in China, may be the earliest known fish.[99]
The last half of the 19th century saw a tremendous expansion in paleontological activity, especially in
North America.[100] The trend continued in the 20th century with additional regions of the Earth being
opened to systematic fossil collection. Fossils found in China near the end of the 20th century have been
particularly important as they have provided new information about the earliest evolution of animals, early
fish, dinosaurs and the evolution of birds.[101] The last few decades of the 20th century saw a renewed
interest in mass extinctions and their role in the evolution of life on Earth.[102] There was also a renewed
interest in the Cambrian explosion that apparently saw the development of the body plans of most animal
phyla. The discovery of fossils of the Ediacaran biota and developments in paleobiology extended knowledge
about the history of life back far before the Cambrian.[59]
Increasing awareness of Gregor Mendel's pioneering work in genetics led first to the development of
population genetics and then in the mid-20th century to the modern evolutionary synthesis, which explains
evolution as the outcome of events such as mutations and horizontal gene transfer which provide genetic
variation, with genetic drift and natural selection driving changes in this variation over time.[103] Within the
next few years the role and operation of DNA in genetic inheritance were discovered, leading to what is now
known as the "Central Dogma" of molecular biology.[104] In the 1960s molecular phylogenetics, the
investigation of evolutionary "family trees" by techniques derived from biochemistry, began to make an
impact, particularly when it was proposed that the human lineage had diverged from apes much more
recently than was generally thought at the time.[105] Although this early study compared proteins from apes
and humans, most molecular phylogenetics research is now based on comparisons of RNA and DNA.[106]
[edit] See also

Paleontology portal
• Archaeobiology
• Dinosaurs
• Fossil collecting
• Fossils
• Geology
• History of paleontology
• Important publications in paleontology
• List of fossil sites (with link directory)
• List of notable fossils
• List of transitional fossils
• Radiometric dating
• Timeline of paleontology
[edit] Notes
1. ^ from Greek: παλαιός (palaeos) "old, ancient", ὄν, ὀντ- (on, ont-) "being, creature", and
λόγος (logos) "speech, thought"
[edit] References
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W000
Phylogenetic tree

"ptree" redirects here. For Patricia tree, see Radix tree.
Fig. 1: A speculatively rooted tree for rRNA genes
A phylogenetic tree or evolutionary tree is a branching diagram or "tree" showing the inferred
evolutionary relationships among various biological species or other entities based upon similarities and
differences in their physical and/or genetic characteristics. The taxa joined together in the tree are implied to
have descended from a common ancestor. In a rooted phylogenetic tree, each node with descendants
represents the inferred most recent common ancestor of the descendants, and the edge lengths in some
trees may be interpreted as time estimates. Each node is called a taxonomic unit. Internal nodes are
generally called hypothetical taxonomic units (HTUs) as they cannot be directly observed. Trees are useful in
fields of biology such as bioinformatics,systematics and comparative phylogenetics.
Contents
[hide]
• 1 History
• 2 Formal Definition
• 3 Types
• 4 Construction
• 5 Limitations
• 6 See also
• 6.1 The "tree of life"
• 6.2 Fields of study
• 7 References
• 9.1 Images
• 9.2 General
[edit] History
The idea of a "tree of life" arose from ancient notions of a ladder-like progression from lower to higher
forms of life (such as in the Great Chain of Being). Early representations of branching phylogenetic trees
include a "Paleontological chart" showing the geological relationships among plants and animals in the book
Elementary Geology, by Edward Hitchcock (first edition: 1840).
Charles Darwin (1859) also produced one of the first illustrations and crucially popularized the notion
of an evolutionary "tree" in his seminal book The Origin of Species. Over a century later, evolutionary
biologists still use tree diagrams to depict evolution because such diagrams effectively convey the concept
that speciation occurs through the adaptive and random splitting of lineages. Over time, species classification
has become less static and more dynamic.
[edit] Formal Definition

In the analysis of genetic evolution, we are given a set X of k sequences, each stands for an extant
species. Let Y be a set of hypothetical sequences, where (usually, each sequence in Y could
represent an extinct species). An evolutionary tree TX,Y for X is a weighted (sometimes rooted) tree of
nodes, where each node is associated with a unique sequence in . The cost of an edge
is the edit distance between the two sequence associated with the ends of the edge. The cost c(TX,Y) of the
tree TX,Y is the total cost of all the edges in TX,Y. [1]
[edit] Types
Fig. 1: Unrooted tree of the myosin supergene family[2]

Fig. 2: A highly resolved, automatically generated Tree Of Life, based on completely sequenced
genomes[3][4].
A phylogenetic tree, showing how Eukaryota and Archaea are more closely related to each other
than to Bacteria, based on Cavalier-Smith's theory of bacterial evolution. (Cf. LUCA, Neomura.)
A rooted phylogenetic tree is a directed tree with a unique node corresponding to the (usually
imputed) most recent common ancestor of all the entities at the leaves of the tree. The most common method
for rooting trees is the use of an uncontroversial outgroup — close enough to allow inference from sequence
or trait data, but far enough to be a clear outgroup.
Unrooted trees illustrate the relatedness of the leaf nodes without making assumptions about
ancestry at all. While unrooted trees can always be generated from rooted ones by simply omitting the root, a
root cannot be inferred from an unrooted tree without some means of identifying ancestry; this is normally
done by including an outgroup in the input data or introducing additional assumptions about the relative rates
of evolution on each branch, such as an application of the molecular clock hypothesis. Figure 1 depicts an
unrooted phylogenetic tree for myosin, a superfamily of proteins.[5]
Both rooted and unrooted phylogenetic trees can be either bifurcating or multifurcating, and either
labeled or unlabeled. A rooted bifurcating tree has exactly two descendants arising from each interior node
(that is, it forms a binary tree), and an unrooted bifurcating tree takes the form of an unrooted binary tree, a
free tree with exactly three neighbors at each internal node. In contrast, a rooted multifurcating tree may have
more than two children at some nodes and an unrooted multifurcating tree may have more than three
neighbors at some nodes. A labeled tree has specific values assigned to its leaves, while an unlabeled tree,
sometimes called a tree shape, defines a topology only. The number of possible trees for a given number of
leaf nodes depends on the specific type of tree, but there are always more multifurcating than bifurcating
trees, more labeled than unlabeled trees, and more rooted than unrooted trees. The last distinction is the
most biologically relevant; it arises because there are many places on an unrooted tree to put the root. For
labeled bifurcating trees, there are
total rooted trees and

total unrooted trees, where n represents the number of leaf nodes. Among labeled bifurcating trees,
the number of unrooted trees with n leaves is equal to the number of rooted trees with n − 1 leaves.[6]
A dendrogram is a broad term for the diagrammatic representation of a phylogenetic tree.
A cladogram is a tree formed using cladistic methods. This type of tree only represents a branching
pattern, i.e., its branch lengths do not represent time.
A phylogram is a phylogenetic tree that explicitly represents number of character changes through its
branch lengths.
A chronogram is a phylogenetic tree that explicitly represents evolutionary time through its branch
lengths.
[edit] Construction
Main article: Computational phylogenetics
Phylogenetic trees among a nontrivial number of input sequences are constructed using
computational phylogenetics methods. Distance-matrix methods such as neighbor-joining or UPGMA, which
calculate genetic distance from multiple sequence alignments, are simplest to implement, but do not invoke
an evolutionary model. Many sequence alignment methods such as ClustalW also create trees by using the
simpler algorithms (i.e. those based on distance) of tree construction. Maximum parsimony is another simple
method of estimating phylogenetic trees, but implies an implicit model of evolution (i.e. parsimony). More
advanced methods use the optimality criterion of maximum likelihood, often within a Bayesian Framework,
and apply an explicit model of evolution to phylogenetic tree estimation.[6] Identifying the optimal tree using
many of these techniques is NP-hard[6], so heuristic search and optimization methods are used in
combination with tree-scoring functions to identify a reasonably good tree that fits the data.
Tree-building methods can be assessed on the basis of several criteria:[7]
• efficiency (how long does it take to compute the answer, how much memory does it need?)
• power (does it make good use of the data, or is information being wasted?)
• consistency (will it converge on the same answer repeatedly, if each time given different data
for the same model problem?)
• robustness (does it cope well with violations of the assumptions of the underlying model?)
• falsifiability (does it alert us when it is not good to use, i.e. when assumptions are violated?)
Tree-building techniques have also gained the attention of mathematicians. Trees can also be built
using T-theory.[8]
[edit] Limitations
Although phylogenetic trees produced on the basis of sequenced genes or genomic data in different
species can provide evolutionary insight, they have important limitations. They do not necessarily accurately
represent the species evolutionary history. The data on which they are based is noisy; the analysis can be
confounded by horizontal gene transfer[9], hybridisation between species that were not nearest neighbors on
the tree before hybridisation takes place, convergent evolution, and conserved sequences.
Also, there are problems in basing the analysis on a single type of character, such as a single gene
or protein or only on morphological analysis, because such trees constructed from another unrelated data
source often differ from the first, and therefore great care is needed in inferring phylogenetic relationships
among species. This is most true of genetic material that is subject to lateral gene transfer and
recombination, where different haplotype blocks can have different histories. In general, the output tree of a
phylogenetic analysis is an estimate of the character's phylogeny (i.e. a gene tree) and not the phylogeny of
the taxa (i.e. species tree) from which these characters were sampled, though ideally, both should be very
close. For this reason, serious phylogenetic studies generally use a combination of genes that come from
different genomic sources (e.g., from mitochondrial or plastid vs. nuclear genomes), or genes that would be
expected to evolve under different selective regimes, so that homoplasy (false homology) would be unlikely
to result from natural selection.
When extinct species are included in a tree, they are terminal nodes, as it is unlikely that they are
direct ancestors of any extant species. Scepticism must apply when extinct species are included in trees that
are wholly or partly based on DNA sequence data, due to the fact that little useful "ancient DNA" is preserved
for longer than 100,000 years, and except in the most unusual circumstances no DNA sequences long
enough for use in phylogenetic analyses have yet been recovered from material over 1 million years old.
In some organisms, endosymbionts have an independent genetic history from the host.
Phylogenetic networks are used when bifurcating trees are not suitable, due to these complications
which suggest a more reticulate evolutionary history of the organisms sampled..
[edit] See also

[edit] The "tree of life"
• Evolutionary history of life - An overview of the major time periods of life on earth
• Life - The top level for Wikipedia articles on living species, reflecting a diversity of
classification systems.
• Three-domain system (cell types)
• Wikispecies - An external Wikimedia Foundation project to construct a "tree of life"
appropriate for use by scientists
[edit] Fields of study

• Archaeopteryx
• Cladistics
• Comparative phylogenetics
• Computational phylogenetics
• Evolutionary biology
• Generalized tree alignment
• Phylogenetics
[edit] References
1. ^ "On the complexity of multiple sequence alignment", J Comput Biol 1 (4): 337–348, 1994
2. ^ Hodge T, Cope M (1 October 2000). "A myosin family tree". J Cell Sci 113 Pt 19 (19): 3353–
4. PMID 10984423. http://jcs.biologists.org/cgi/content/full/113/19/3353.
3. ^ Letunic, I; Bork, P (2007). "Interactive Tree Of Life (iTOL): an online tool for phylogenetic
tree display and annotation." (Pubmed). Bioinformatics 23 (1): 127–8.
doi:10.1093/bioinformatics/btl529. PMID 17050570.
4. ^ Ciccarelli, FD; Doerks, T; Von Mering, C; Creevey, CJ; Snel, B; Bork, P (2006). "Toward
automatic reconstruction of a highly resolved tree of life." (Pubmed). Science 311 (5765): 1283–7.
doi:10.1126/science.1123061. PMID 16513982.
5. ^ Maher BA (2002). "Uprooting the Tree of Life". The Scientist 16: 18. http://www.the-
scientist.com/yr2002/sep/research1_020916.html.
6. ^ a b c Felsenstein J. (2004). Inferring Phylogenies Sinauer Associates: Sunderland, MA.
7. ^ Penny, D., Hendy, M. D. & M. A. Steel. 1992. Progress with methods for constructing
evolutionary trees. Trends in Ecology and Evolution 7: 73-79.
8. ^ A. Dress, K. T. Huber, and V. Moulton. 2001. Metric Spaces in Pure and Applied
Mathematics. Documenta Mathematica LSU 2001: 121-139
9. ^ Woese C (2002). "On the evolution of cells". Proc Natl Acad Sci USA 99 (13): 8742–7.
doi:10.1073/pnas.132266999. PMID 12077305.

• Schuh, R. T. and A. V. Z. Brower. 2009. Biological Systematics: principles and applications
(2nd edn.) ISBN 978-0-8014-4799-0
• MEGA, a free software to draw phylogenetic tress.
Wikimedia Commons has media related to: Phylogenetic tree
[edit] Images
• Phylogenetic Trees Based on 16s rDNA
• Poster-sized tree of life illustration
• A 3D View
• Human Y-Chromosome 2002 Phylogenetic Tree
• In 2003, the Science journal dedicated a special issue to the tree of life, including an online
version of a tree of life.
• iTOL: Interactive Tree Of Life
• Phylogenetic Tree of Artificial Organisms Evolved on Computers
• Miyamoto and Goodman's Phylogram of Eutherian Mammals
[edit] General
• Discover Life An interactive tree based on the U.S. National Science Foundation's
Assembling the Tree of Life Project
• PhyloCode
• A Multiple Alignment of 139 Myosin Sequences and a Phylogenetic Tree
• Tree of Life Web Project
• Aisee.com, detailed and comprehensive family tree of dinosaurs yet available
• SplitsTree
• Dendroscope
• Phylogenetic inferring on the T-REX server
• NCBI's Taxonomy Database[1]
• ETE: A Python Environment for Tree Exploration This is a programming library to analyze,
manipulate and visualize phylogenetic trees. Ref.
[hide]v · d · eTopics in phylogenetics
Relevant
fields Computational phylogenetics · Molecular phylogenetics · Cladistics
Basic Phylogenetic tree · Phylogenetic network · Long branch attraction · Clade vs

concepts Grade · Ghost lineage
Inference Maximum parsimony · Maximum likelihood · Neighbor-joining · UPGMA ·

methods Bayesian inference · Least squares · Three-taxon analysis
Current topics PhyloCode · DNA barcoding

-morphy Symplesiomorphy · Apomorphy · Synapomorphy · Autapomorphy
-phyly Monophyly/Holophyly · Paraphyly · Polyphyly
List of evolutionary biology topics
[hide]v · d · eOrigin of life
Quasispecies model · Protobiont · Universal common descent · Last universal ancestor · RNA world
hypothesis · Iron–sulfur world theory · PAH world hypothesis · Miller–Urey experiment · Panspermia
Retrieved from "http://en.wikipedia.org/wiki/Phylogenetic_tree"
Categories: Phylogenetics | Tree of life | Trees (structure)

W000
The Ancestor's Tale


The Ancestor's Tale
Author Richard Dawkins
Subject(s) Evolutionary biology
Publisher Boston: Houghton Mifflin
Publication date 2004
Pages 673
ISBN 0618005838
OCLC Number 56617123
Dewey Decimal 576.8 22
LC Classification QH361 .D39 2004
Preceded by A Devil's Chaplain

Followed by The God Delusion
The Ancestor's Tale (subtitled A Pilgrimage to the Dawn of Life ) is a 2004 popular science book by
Richard Dawkins, with contributions from Dawkins' research assistant Yan Wong. It follows the path of
humans backwards through evolutionary history, meeting humanity's cousins as they converge on common
ancestors. The book was nominated for the 2005 Aventis Prize for Science Books.
Contents
[hide]
• 1 Synopsis
• 2 List of rendezvous points
• 2.1 Prologue
• 2.2 Primates
• 2.3 Non-primate mammals
• 2.4 Non-mammal chordates
• 2.5 Non-chordate animals
• 2.6 Non-animal eukaryotes
• 2.7 Prokaryotes
• 3 Editions
• 4 Translations
• 5 See also
• 6 References
[edit] Synopsis
Cladogram showing relationship between mammalian species as recounted in the book

The narrative is structured as a pilgrimage, with all modern animals following their own path through
history to the origin of life. Humans meet their evolutionary cousins at rendezvous points along the way, the
points at which the lineage diverged. At each point Dawkins attempts to infer, from molecular and fossil
evidence, the probable form of the most recent common ancestor and describes the modern animals that join
humanity's growing travelling party. This structure is inspired by Geoffrey Chaucer's The Canterbury Tales.
The pilgrimage visits a total of 40 "rendezvous points" from rendezvous zero, the most recent
common ancestor of all of humanity, to rendezvous 39, eubacteria, the ancestor of all surviving organisms.
Though Dawkins is confident of the essential shape of this phylogenetic taxonomy, he enters caveats on a
small number of branch points where a compelling weight of evidence had not been assembled at the time of
writing.
At each rendezvous point, Dawkins recounts interesting tales concerning the cousin animals which
are about to join the band of pilgrims. Every newly recruited species, genus or family has its own peculiar
features, often ones that are relevant to human anatomy or otherwise interesting for humans. For instance,
Dawkins discusses why the axolotl never needs to grow up, how new species come about, how hard it is to
classify animals, and why our fish-like ancestors moved to the land. These peculiar features are studied and
analyzed using a newly introduced tool or method from evolutionary biology, carefully woven into a tale to
illustrate how the Darwinian theory of evolution explains all diversity in nature.
A highly resolved Tree Of Life, based on completely sequenced genomes.[1][2]
Even though the book is best read sequentially, every chapter can also be read independently as a
self-contained tale with an emphasis on a particular aspect of modern biology. As a whole, the book
elaborates on all major topics in evolution.
Dawkins also tells personal stories about his childhood and time at university. He talks with fondness
about a tiny bushbaby he kept as a child in Malawi (Nyasaland). He described his surprise when he learned
that the closest living relatives to the hippos are the whales.
The book was produced in two hardback versions: a British one with extensive colour illustrations (by
Weidenfeld & Nicolson), and an American one with a reduced number of black-and-white illustrations (by
Houghton Mifflin). Paperback versions and an abridged audio version (narrated by Dawkins and his wife Lalla
Ward) have also been published.
The book is dedicated to Dawkins' friend and mentor, population geneticist John Maynard Smith, who
died shortly before the book went to press.
[edit] List of rendezvous points

Dawkins uses the term concestor—coined by Nicky Warren—for the most recent common ancestor at
each rendezvous point. At each rendezvous point, we meet the concestor of ourselves and the listed species
or collection of species. This does not mean that the concestor was much like those creatures; after the
"rendezvous", our fellow "pilgrims" have had as much time to evolve and change as we have. Only creatures
alive at the time of the book's writing join us at each rendezvous point. Except for a few special cases,
numerous extinct species and families such as the dinosaurs are excluded from the pilgrimage.
[edit] Prologue
Rendezvous Significant
Time Story
point event
0.01 Neolithic The Farmer's Tale describes the

n/a
mya Revolution Neolithic Revolution
0.04 Great Leap The Cro-Magnon's Tale describes

n/a
mya Forward the Great Leap Forward.
[edit] Primates
Rendezvous
Time Joining party Story
point
0 All Humankind The Tasmanian's Tale illustrates the

identical ancestors point starting from which
all living people trace exactly the same set of
ancestors back in time.
Eve's Tale touches upon coalescent

theory, Mitochondrial Eve, Y-chromosomal
Adam and polymorphism. The story ends
with a speculation that the ABO blood group
system in humans and chimps are examples
of trans-specific polymorphism; a type-B
human may actually be more closely related
to type-B chimp than type-B human is
related to type-A human, from the
perspective of the genes (or alleles)
responsible for the antigens.
The Ergast's Tale recounts how a

mutated form of the FOXP2 gene could have
allowed Homo ergaster to acquire language.
The Handyman's Tale explains how

Homo habilis acquired high 'brain to body
mass ratio', at the same time introducing
logarithmic scale and scatterplot as tools for
scientific studies.
Little Foot's Tale examines how

hominid first learned to walk on two legs.
Human pilgrims join their

6 Chimpanzees
1 evolutionary cousins, chimpanzees and
mya (Pan)
bonobos.
2 7 Gorillas (Gorilla) The Gorilla's Tale considers

mya human's changing attitude towards the great
apes, ending with a discussion on racism,
speciesism and the Great Ape Project.
The Orangutan's Tale introduces the

principle of parsimony and its use in
14 Orangutans
3 construction of family tree (cladogram) of
mya (Pongo)
species. Orangutan is the last of the great
apes to join the pilgrimage.
4 18 Gibbons The Gibbon's Tale further elaborates

mya (Hylobatidae) on neighbor-joining, parsimony and textual
criticism techniques used to construct
cladograms. When simple principle of
parsimony proves inadequate to handle
'long branch attraction' problems caused by
convergence and reversion, the
phylogenetic tree (phylogram) and
computational phylogenetic methods such
as maximum likelihood analysis are
introduced. The tale ends with yet another
example of trans-specific polymorphism:
sexual dimorphism; the sex-determining
SRY has never been in female bodies long
since gibbons and humans diverged. This
serves to highlight the fact that different
phylogenetic trees can be created by tracing
different sets of genes; the one mainstream
'species tree' is nothing more than a
summary of multitude of gene trees, a
'majority vote' among gene trees. Gibbon is
the last ape to join the pilgrimage.
Old World monkeys, being in the

Old World same Catarrhini clade as apes, are closer
25
5 Monkeys cousins to apes than to New World
mya
(Cercopithecidae) monkeys. Old World monkeys are
sometimes called the 'tailed apes'.
6 40 New World The Howler Monkey's Tale calls

mya Monkeys (Platyrrhini) attention to the critical role of gene
duplication in evolution. While our remote
vertebrate ancestors possessed trichromatic
vision, our nocturnal, warm-blooded,
mammalian ancestors lost one of three
cones in the retina at the time of dinosaurs.
This is why fish, reptiles and birds are
trichromatic while all mammals with the
exception of apes and New World monkeys
are strictly handicapped dichromats.
Because color vision is of paramount
importance to diurnal animals that eat ripe
fruits, apes and New World monkeys
regained tri-color vision independently via
chromosomal translocation. In apes,
trichromacy resulted from true duplication of
the opsin gene. New World monkeys first
achieved trichromacy in its female
population by producing two alleles (green
and red) for the same locus for the opsin
gene on the X-chromosome, an example of
polymorphism. Its males, with only one copy
of the X-chromosome, remained dichromats
with either a green or a red opsin, an
example of heterozygote advantage. Howler
monkeys, a type of New World monkey, took
this one step further and achieved
trichromacy for both sexes when its X-
chromosome gained two loci to house both
the green allele and the red allele. New
World monkeys are the last simians (also
known as 'higher primates' or anthropoids)
to join the pilgrimage.
7 58 Tarsiers (Tarsius) Tarsier is the last haplorrhine to join

mya the pilgrimage. A nocturnal animals, the
tarsier has two enormous eyes each as
large as its brain. Unlike other nocturnal
mammals, however, tarsier eyes do not
contain tapetum lucidum which reflects light
from the back of the eye for a second
exposure on the retina to maximize light
capture. From this we can infer that the
common ancestor of all haplorrhine must
have been a diurnal animal which shed the
tapetum lucidum to eliminate blurry images
caused by reflected light. When the tarsier
became a nocturnal animal, it enlarged its
eyes to compensate for the lack of tapetum
lucidum.
8 63 Lemurs, The pilgrimage meets with the rest

mya Bushbabies and their kin of the prosimian cousins: the lemurs, pottos,
(Strepsirrhini) bushbabies, and lorises. The Aye-Aye's Tale
showcases the strange lemurs which are
only found on the island of Madagascar.
Madagascar was originally part of the
Gondwana supercontinent which included
present Africa continent and Indian
subcontinent. Gondwana broke off into
drifting blocks of land, some of which
became Africa, India and Madagascar. As
an estranged island, Madagascar became a
speciation hotbed. For instance a small
founding population of strepsirrhine primates
(possibly rafted in from neighboring
continent) flourished and diversified into all
niches of the ecosystem, in the absence of
monkeys. The story reminds us how
Madagascar, with a land mass 1/1000 of
Earth's total land area, ends up housing
unique species that account for 4% of all
species of animals and plants. Lemurs and
their kin are the last of the primates to join
the pilgrimage.
[edit] Non-primate mammals

Rendezvous
point
9 70 Colugos and The Cretaceous–Tertiary extinction

mya Tree Shrews event occurred at 65 million years ago, due to
(Dermoptera and both large scale volcanic activities in the
Scandentia) Deccan traps over a period of time, and the
final asteroid impact event which created the
Chicxulub Crater. The sudden temperature
change and sunlight reduction caused
massive disruptions to Earth's ecosystem. As
a result, all dinosaurs except the birds, as well
as numerous other species went extinct. The
disappearance of dinosaurs made it possible
for many different species of shrew-like,
nocturnal insectivores to evolve into hippos,
lions, elephants, etc. to fill the new ecological
voids, an example of evolutionary radiation.
One of these shrew-like creatures was the
concestor of the current pilgrimage party and
the new joiners, the colugos and tree shrews.
The tree shrews resemble the

squirrels. The colugos resemble flying
squirrels. In both cases, the resemblance is
only superficial, due to convergence; the
squirrels are rodents and will meet us further
down the pilgrimage. At the present, scientists
are not yet sure about the exact relationships
among the tree shrews, the colugos and us.
Dawkins provisionally accepts the view that
they join forces first, before meeting the
pilgrimage party. This would place our
concestor 9 at a time before K-T boundary
which marked the extinction of the dinosaurs.
The Colugo's Tale warns us that even though
the general structure of the family tree is
sound, some of the details could change as
more evidences become known.
10 75 Rodents and Rodents comprise the largest number

mya Rabbitkind (Glires) of species in mammalia, more than 40 percent
of all mammalian species. Members include
rats, mice, lemmings, beavers, squirrels, etc.
The Mouse's Tale explains how mammals
possess similar and relatively small genomes
in the order of 30,000 genes, yet each animal
exhibits distinct features and surprising
complexities. Dawkins debunks the popular
description of genome as blueprints which
give rise to the misconception that the more
complex the animal, the more complex the
blueprint ought to be. Instead, genes in a
genome should be thought of as words or
sentences in a language, and embryonic
development over time is akin to 'order' of
words and sentences in a book. While the
number of genes are limited, the endless
number of 'orders' by which similar genes in
mice and humans are deployed during
embryonic development can generate
astonishing complexity and distinguish a
mouse from a man.
The Beavers's Tale revisits the key

insights that Dawkins contributed to the field of
evolution in his book The Extended
Phenotype. A beaver's body is known as a
phenotype, an external and visible
manifestation of the internal and hidden
genotype. In the same way the beaver body is
regarded as an expression of its genes,
beaver dams or beaver lakes can be
considered 'extended phenotypes' of the same
beaver genes. Better beaver genes make
better beaver bodies, beaver dams and
beaver lakes. In other words, beaver genes
are selected not only by the fitness of beaver
bodies, but also by the effectiveness of beaver
dams and beaver lakes they produce.
11 85 Laurasiatheres An extremely diverse group of animals

join the pilgrimage, including Carnivora (dogs,
mya (Laurasiatheria) cats, bears and seals), Perissodactyla
(horses, tapirs and rhinos), Cetartiodactyla
(deer, cattle, pigs and hippos), Chiroptera
(bats), Insectivora (moles and shrews), etc.
Some of them fly, others swim, and yet many
of them gallop. Half of them are predators
which hunt the other half of the group. The
only thing they share in common is that they
join up with one another before the group joins
us to meet concestor 11. This group of
animals belong to the Laurasiatheria clade as
all of them originated from the supercontinent
of Laurasia.
The Hippo's Tale is really the whale's

tale. All cetaceans, including whales, dolphins
and porpoises, are descendants of land-living
mammals of the Artiodactyl order (even-toed
ungulate animals). Both cetaceans and
artiodactyl are now classified under the super-
order Cetartiodactyla which includes both
whales and hippos. In fact, whales are the
closest living relatives of hippos; they evolved
from a common ancestor at around 54 million
years ago. This story illustrates how a species
can flip into evolutionary overdrive when it
enters into a new environment, while its
closest relatives remain unchanged for long
time in their static environment.
The Seal's Tale illustrates how a sex

ratio of 50:50 (males to females) is found in
most sexually reproducing animals from both
monogamous and polygamous species. In a
harem-based (polygynous) system such as
that of elephant seals where 4 percent of
males account for 88 percent of all
copulations, the actual sex ratio of 50:50
seems to produce an excess of males who
consume resources but end up leaving no
offsprings. This puzzle is solved by the
concept of Fisher's principle (named 'parental
expenditure') proposed by R.A. Fisher. This
led to further work by Robert Trivers on
parental investment to elucidate sexual
selection. More importantly, the elephant seal
typifies sexual dimorphism, as a bull elephant
seal can grow to be three times the size of a
cow seal, thanks to sex-limited genes which
exist in both male and female bodies, but
remain turned off in females. The degree of
sexual dimorphism is correlated with the
harem size, which allows us to draw
inferences about our immediate human
ancestors: they were probably mildly
polygynous.
12 95 Xenarthrans The Armadillo's Tale reminds us of the

mya (Xenarthra) aye-aye's tale, except that instead of
Madagascar, the speciation hotbed is the
continent of South America. This continent
broke off from Gondwana in Early Cretaceous
period, then joined North America which broke
off from Laurasia. During its long period of
isolation, South America was host to
marsupials which flourished and took up all
carnivorous niches. The placental mammals
(including armadillo) and now-extinct
ungulates evolved to fill the rest of the
ecosystem. When South America joined North
America during the Great American
Interchange at 3 million years ago, animals
and plants cross the Isthmus of Panama in
both directions, introducing new species to
new land and driving some local species to
extinction. Jaguars and other carnivorous
placental mammals were introduced to South
America, while armadillos migrated to North
America.
The pilgrimage party is joined by the

last of the placental mammals: elephants,
elephant shrews, dugongs, manatees,
hyraxes, aardvarks, etc. They all hail from
105 Afrotheres
13 Africa, as hinted by the name of their clade,
mya (Afrotheria)
Afrotheria. The concestor we greet at this
point, as well as those we met earlier at
rendezvous 12 and 11, all look like
insectivorous shrews.
14 140 Marsupials The entire band of placental mammals

mya (Metatheria) meet up with the other great group of
mammals, the marsupials. Even though
present-day marsupials are mostly found in
Australia and New Guinea, they originally
flourished and diversified for a period of time
in South America. Evidence points to the
migration of a single species of opossum-like
marsupial from South America to Australia
before 55 million years ago, when it was still
possible to make the journey through
Antarctica before Australia pulled too far away
from Gondwana. Once settled in the isolated
Australia, the founding marsupials quickly
evolved into distinct species and, for the next
40 million years, took up the entire range of
'trades' previously occupied by dinosaurs, in
the absence of any placental mammals.
The Marsupial Mole's Tale again

highlights the wonders that convergent
evolution can create. Despite great
evolutionary distance between marsupial
moles in Australia and the golden moles in
Africa, they are remarkably similar in terms of
phenotypes, with the exception that the
marsupial moles sport a pouch as all
marsupials do. There are also marsupial mice
(Dasyuridae), marsupial flying squirrels (Sugar
Glider) and marsupial wolf (Thylacine), not to
mention the equivalent of antelopes and
gazelles, the kangaroos and wallabies which
despite great differences in shape, cover the
same range of diet and way of life as their
African counterparts.
15 180 Monotremes The monotremes are the last of the
mya (Monotremata) mammals to join us, and we meet a concestor
for the first time in the then-contiguous
supercontinent of Pangea. The monotremes
consititutes only a few genera: Platypus,
Short-beaked Echidna and Long-beaked
echidna. They are mammals and have typical
mammalian features such as warm-
bloodedness, hair and milk production. But
they resemble reptiles and birds in their
possession of the cloaca and their egg-laying
mode of reproduction.
The Duckbill's Tale warns us about

the fallacy of labeling a half-mammal and half-
reptile animal such as the duckbill platypus as
primitive. The platypus has exactly the same
time to evolve as the rest of mammals, even if
it does resemble our concestor 15 on the
surface. In fact, it has evolved a highly
developed form of electroreception served by
40,000 electric sensors, and 60,000
mechanical push rods, both on its large bill to
aid it in search of crustaceans in the mud. In
human, the brain dedicates disproportionally
large amount of cells to the two hands as
illustrated by the Penfield brain map, or
Penfield homunculus. When the same
somatotopic map is drawn for platypus brain,
the bill is served by equally prominent
percentage of the brain.
[edit] Non-mammal chordates

Rendezvous
point
16 310 Sauropsids The pilgrims are about to join their

mya (Sauropsida) reptile cousins, after marching for 130 million
unbroken years from the last mammal
concestor 15 who looks like a shrew to the
reptile concestor 16 who looks like a lizard.
In these 130 million years, mammal-like
reptiles flourished, even before dinosaurs
roamed the Earth. But like the 99 percent of
all species that ever existed, all branches of
mammal-like reptiles are now extinct, so they
cannot join us in our pilgrimage.
The term reptile is not a true clade

name, as it fails to include birds which share
a common ancestry. The terms reptile and
fish are known as grades which only make
sense scientifically when used in the now-
discredited theory of progressive evolution
(Orthogenesis). Progressive evolution
proposes that species evolve independently,
in a parallel, progressive direction from fish
grade through amphibian grade via reptile
grade towards mammal grade. From a
cladistic point of view, turtles, lizards,
snakes, crocodiles, dinosaurs and birds are
all members of the clade Sauropsids which is
what Dawkins adapts instead of the term
reptile. Dinosaurs, unfortunately as extinct
species, cannot join us. But their only
surviving descendants, the birds, take their
place in the pilgrimage.
The Galapagos Finch's Tale

addresses a surprising question: why doesn't
evolution go much faster than it does?
Studies on Galapagos finches show that the
Medium Ground-finch (Geospiza fortis) could
grow to be as large as the Large Ground-
finch (Geospiza magnirostris), if 23
consecutive drought years put pressure on
birds to grow larger beaks to better handle
bigger and tougher seeds. But such extreme
evolutionary speed is not observed in nature,
given the geological time life has had on
Earth. This is mostly because the rate of
evolution follows major trends over
geological timescale, while short-term
pressures tend to cancel out one another.
The Peacock's Tale is the

quintessential illustration of sexual selection;
the peacock's tail is the ultimate example of
non-utilitarian phenotype which appears to
be an anomaly in natural selection, as it is a
hindrance to peacock's survival in its natural
habitat. These arbitrary spurts of evolution
can be explained by sexual selection, a
special case of natural selection. The
runaway explosion of the extravagant tail is
created by lockstep dimorphic selections of
male's genes for pretty tails and of female's
genes for preferring such tails. The
preference for pretty tails, in turns, is rooted
in their use by males as tokens of underlying
fitness; an advertisement of health. Sexual
selection often complements other natural
selection forces, and helps explain why
human became bipedal to free the two hands
for tool making and wielding, attained a
larger brain with artistic abilities, and shed
body hair to advertise lack of ectoparasites
during the course of human evolution.
The Dodo's Tale illustrates how

evolution optimizes genes for the present
environment, how it has no foresight, and
how it marches blindly sometimes to the
detriment of the species. The dodo, originally
a flying pigeon related to the Rodrigues
Solitaire, reached the remote island of
Mauritius and shed its flying powers due to
the lack of competition and natural predators
to become a Flightless bird. The reallocation
of resources away from building of massive
breast muscles for flying allowed the dodo to
flourish, but ultimately lead to their extinction
when European sailors and their carnivorous
pets finally arrived.
The Elephant Bird's Tale
demonstrates how enigmatic distributions of
genetically close species in completely
separate continents can be explained and
corroborated by evidences of continental drift
and seafloor spreading. The tale recounts
the diaspora of a large group of flightless
ratite birds from the then unbroken
Gondwana; Moa ended up in New Zealand,
rhea in South America, emu in Australia,
cassowary in New Guinea, kiwi in New
Zealand by island hopping, and ostrich in
Africa by way of Asia and Europe.
Radioactive dating of and magnetic striping
studies on continuously formed crust around
rifts such as the Mid-Atlantic Ridge allow
Paleobiogeographers to piece back a
coherent story of these birds' dispersion
based on both phylogenetic tree and plate
tectonics.
17 340 Amphibians Mammals and reptiles (the amniotes)

mya (Amphibia) join the amphibians to meet the ancestor of
all land vertebrates with four feet, the
tetrapod. Amphibians include frogs, toads,
salamanders, newts and caecilians. While
amniotes either give live births or lay
waterproof eggs, the amphibians retain the
ancestral practice of laying eggs in water.
Unlike the waterproof skin of amniotes, the
amphibian skin allows body water to
evaporate through it, restricting amphibians
to land areas with access to fresh water.
The Salamander's Tale uses

examples of ring species to illustrate how a
continuous series of interbreeding animals in
the spatial dimension is conceptually
equivalent to that in the time dimension. The
Ensatina salamanders in the Central Valley
in California form a continuous ring (actually
a horseshoe shape) around the valley. Any
two neighboring population of Ensatina
around the horseshoe can interbreed, but the
plain Ensatina eschscholtzii on the western
end of the horseshoe cannot interbreed with
the large blotched Ensatina klauberi on the
eastern end. Larus gulls form another ring
species which starts at Herring Gull in Great
Britain and ends at Lesser Black-backed Gull
in north-western Europe. Dawkins likens
both ring species in space to the ring in time
that unites humans and chimpanzees via
generations of ancestors over 6 million
years, with concestor 1 in the midpoint.
The Narrowmouth's Tale shows how

speciation may still continue via parapatric
speciation, when two closely related toad
species meet again after initial geographical
isolation. Gastrophryne olivacea (Great
Plains narrowmouth toad) and Gastrophryne
carolinensis (Eastern narrowmouth toad) are
closely related and can interbreed when their
habitats overlap. But reinforcement, a
selection process which increases
reproductive isolation via character
displacement, causes both species to
differentiate their mating calls from each
other by shifting pitch and duration in
opposite directions; the more the two
populations overlap, the more distinct their
mating calls become.
The Axolotl's Tale is about

metamorphosis, a biological process which
turns juveniles or larvae into drastically
dissimilar adult forms for reproduction, and
about pedomorphosis, another process
which enables juveniles of some species to
become sexually-mature without ever
developing into their usual adult forms.
Species which undergo metamorphosis
include butterflies, barnacles and
salamanders. Species which exhibit neoteny,
a type of pedomorphosis, include human,
ostrich, pekingese and axolotl. A text book
example of neoteny, the axolotls are
members of the Tiger Salamander complex,
yet they become sexually-mature in larva
form, remaining aquatic and gilled. With a
treatment of thyroxine, it is possible to induce
an axolotl to develop into a salamander,
demonstrating that axolotl genome still
retains information on its lost adult form. On
the other hand, newt, a type of salamander,
first develops from tadpole into land-based
salamander, but later reverts back to its
juvenile tadpole form, and returns to the
water to reproduce. The axolotl's tale
reminds us that paedomorphosis often
allows species to break out of an
evolutionary dead end by sudden changes.
417 The Queensland lungfish (Australian

18 Lungfish (Dipnoi)
mya lungfish) and Coelacanth are two of the most
famous living fossils; they resemble ancient
19 425 Coelacanths fossils and unlike most species, seemingly
mya (Latimeria) refused to continue to evolve for the past 400
million years. The lungfish joins the
pilgrimage to meet concestor 18, before the
coelacanth joins at rendezvous point 19.
Instead of looking like members from the
pilgrimage, the lungfish actually resembles
coelacanth and concestor 19, the lobe-finned
fish Sarcopterygii. Despite their
morphological similarities, however, the
lungfish and coelacanth are very different
genetically, as expected of species which
lived separately for more than 400 million
years. Because genes do not stop evolving,
the molecular DNA of these two species
show greater evolutionary distance from
each other than to DNA of the rest of the
pilgrimage. The Lungfish's Tale reminds us
that the rate of morphological changes is not
always obviously correlated with that of
20 440 Ray-Finned Fish The current pilgrimage consisting of
mya (Actinopterygii) all descendants of lobe-finned fish is joined
by the equally successful ray-finned fishes
which includes sturgeon, paddlefish, eel,
herring, carp, salmon, trout, seahorse, cod,
etc. to meet concestor 19, the bony fish. Of
all ray-finned fishes, most belong to the large
infraclass teleostei.
Some teleost fishes evolved unfishy

shapes to cope with their chosen ecological
niches. The leafy sea dragon, for instance,
abandons the typical streamline fish shape
which works so well for the majority of fishes.
Instead, it adopts a leafy shape to hang
motionless in kelp forest, pretending to be a
piece of seaweed. The razorfish takes up an
elongated, laterally compressed body,
together with a long, flattened snout. It swims
in a head-down vertical stance, allowing it to
hide amongst tall spines of a sea urchin. The
snipe eel is ridiculously thin, while the gulper
eel sports jaws which look disproportionally
large for its body. Lastly, the ocean sunfish
resembles a huge, two-ton disc or millstone,
as its Latin name, Mola mola, suggests. The
Leafy Sea Dragon's Tale demonstrates how
animal shapes are malleable, ever changing
to meet the requirements of each animal's
way of life.
The Pike's Tale highlights a special

organ which gives teleost fishes superior
buoyancy control, the swim bladder.
Contrary to common assumptions, swim
bladder is not a precursor to lungs in human
and other lobe-finned fishes. Instead, the
bony fish ancestor possessed a primitive
lung which was co-opted by teleost fishes for
buoyancy control, and in some cases as ear
drum for hearing. The teleost fishes rely on
gills for breathing underwater. They
repurposed the primitive lung, turning its
ability to absorb from and release gas into
the blood stream into a volume-changing
mechanism, thus allowing teleost fishes to
move vertically in a water column without the
use of fins.
The Mudskipper's Tale shows how

animals rediscover long lost faculties and
reenact ancient evolutionary events via
completely different biological mechanisms.
Having forgone air-breathing by repurposing
the lung for buoyancy control, some teleost
fishes, such as the Siamese fighting fish
(Betta splendens), reinvent air-breathing by
gulping air and locally oxygenerating water in
the gill chamber. The mudskipper
Periophthalmus not only takes air into its
moist gill chamber, but can also breathe
through its skin. Both the gill chamber and
the skin must be wet at all times, and this
distinguishes the new type of air-breathing
from breathing through lungs. Re-equipped
with air-breathing apparatus, the mudskipper
emerges onto land, replaying ancient
lobefin's conquest of the land.
The rapid speciation of

haplochromine cichlid fishes endemic to
Lake Victoria, Lake Malawi and Lake
Tanganyika exemplifies adaptive radiation
and species flock. The Cichlid's Tale
recounts how, by constructing an "unrooted
haplotype network" using phylogenetic
analysis on mitochondrial DNA of living
species from regional rivers and lakes,
researchers were able to infer the time and
the location of each major speciation event in
the evolutionary history of these cichlids. The
haplotype network differs from normal
phylogenetic tree in that each node
represents a haplotype, not a species, and
the node size is determined by number of
species in which the haplotype is found. By
analyzing genetic relationships between
haplotypes, relative prevalence of each
haplotype, and locations where species
currently live, it is possible to trace past
waves of adaptive radiation originating from
a small founding species, as rivers and lakes
rose and fell in level.
The Blind Cave Fish's Tale illustrates

how normal organs can degenerate into
vestigial organs. Different populations of
Mexican tetra (Astyanax mexicanus) have
ventured into dark caves separately, and
have separately evolved regressive traits
such as white skin coloration and regressive
or blind eyes. This is partially explained by
the opportunity cost theory; resources
wasted on building the eye in a pitch-black
cave deprives the fish of other traits useful
for such environment. But more importantly,
without evolutionary pressure to weed out
bad mutations on the multitude of genes
which together build the eye, any random
change is more likely to disrupt the dedicate
process of building the eye than to enhance
it. There is no need to revert precisely the
sets of genes carefully shaped by millions of
years of evolution to get back to a blind
creature. In other words, there are many
more random ways of building a blind fish
than of building a sighted one. And this is the
essence of the Dollo's Law as Dawkins
interprets it - that evolution cannot be
precisely and exactly reversed.
The Flounder's Tale is a tale of
imperfection. The flounder's contorted head
and eyes allow it to lie on its side on the
ocean floor, but they betray the lack of an
intelligent designer. As expounded in The
Blind Watchmaker and Climbing Mount
Improbable, evolution does not 'design' every
creature anew on a drawing board. Instead,
natural selection works without foresight and
makes gradual improvements on existing
body plans from generation to generation.
Because each creature at every step of the
process must remain fit for its environment,
evolution cannot make sudden and drastic
changes to build a better future organism at
the expense of current generation.
21 460 Sharks and Their Cartilaginous fishes chondrichthyan

mya Kin (Chondrichthyes) including sharks, rays and chimaeras join the
pilgrimage in the Middle Ordovician. The
newcomers have no bones. Instead, they are
supported by a cartilaginous skeleton that
never ossifies, in contrast to bony fishes.
Their skin is covered in dermal denticles, tiny
scale-like protrusions, from which teeth may
have evolved. Interestingly, sharks lack a
swim bladder for buoyancy, and instead rely
on swimming constantly, retaining urea in
their blood, and having large livers with
plenty of oil to remain afloat. The
Carcharocles megalodon from the Miocene
is described as a predator more terrifying
than the Great White Shark, as it was three
times the size. The strange Chimaera has
strange gill covers, has no dermal denticles,
and swims using their pectoral fins. Dawkins
explains that concestor 21 is ancestor to all
gnathostomes, animals with lower jaws, a
structure which evolved from the gill arches.
22 530 Lampreys and Jawless and limbless fishes, the

mya Hagfish lampreys and hagfish, join the pilgrimage to
(Cyclostomata) meet the concestor of all vertebrates. The
jawless fish and the concestor 22 are
borderline vertebrates. Unlike the rest of
vertebrates, they retain the notochord, a
stiffening cartilage rod running along the
back of an animal, well into adulthood. In all
other vertebrates, the vestigial notochord
appears in the embryo briefly and is replaced
by segmented, articulate backbones in
adults. On the other hand, both the jawless
fish and the jawed fish share characteristics
common to all members of the phylum
Chordata at some time in their life cycle,
including the notochord, pharyngeal slit, and
the post-anal tail.
The Lamprey's Tale further develops

the gene's eye view of ancestry and
pedigree that earlier tales, The Eve's Tale
and The Gibbon's Tale, alluded to. In human,
four haemoglobin genes are known to be
cousin genes of each other. An ancestor
globin gene from an ancient vertebrate split
into two genes, alpha and beta, which ended
up in two different chromosomes and
continued to evolve independently. Both
alpha and beta further split into more
independently evolving genes. All jawed fish
show such alpha/beta split as predicated by
evolution. However, lampreys and hagfish
are ancient enough that they predate this
gene split. In fact, jawless fishes, whenever
investigated, do not possess split globin
genes. As Dawkins explained in the chapter
'All Africa and her progenies' in his book
River out of Eden, there are two ways to
trace ancestry: via animals and via individual
genes. The two mechanisms produce very
different results. Ancestry of animals form a
family tree (more correctly, a graph because
sexually reproducing animals may share
female and male parents). On the other
hand, Ancestry of an individual gene is
always a single chain going back to the first
self-replicating RNA, since a gene is either a
faithful copy or a mutated form of its single
parent gene. The Ancestor's Tale is written
from an animal's perspective, following the
family tree of human backward in time. But
the book could have been written from the
gene's point of view. Starting from any gene
(e.g. the alpha haemoglobin), each gene
gene duplication event could become a
rendezvous point where pilgrimage of genes
join their cousin genes.
23 560 Lancelets Lancelets are text book examples of

mya (Amphioxiformes) a chordate. Equipped with a notochord, a
nerve tube on the dorsal side and gill slits,
they typify the phylum Chordata. But
lancelets are not primitive nor our remote
ancestor. They are as modern as all other
members in the pilgrimage. The Lancelet's
Tale continues to develop the theme
introduced in The Duckbill's Tale, that all
living animals have had equal time to evolve
since the first concestor, and that no living
animal should be described as either lower
or more primitive. Dawkins extends this
concept to apply to fossils as well. Even
though it is tempting to label fossils as our
remote ancestor, they are more accurately
described as our distant cousins who have
been frozen in time.
24 565 Sea Squirts A Sea squirt resemble a sedentary

mya (Urochordata) bag of sea water anchored to a rock. It feeds
on food particles strained from water.
Anatomically, the sea squirt looks very
different to the joining pilgrimage of all
vertebrates and protochordates, that is, until
its larvae are examined. The sea squirt larva
looks and swims like a tadpole. it possesses
a notochord and a dorsal nerve tube, and
moves by undulating its post-anal tail from
side to side. Vertebrates may have branched
off from ancient sea squirt larvae via
neoteny, in a process reminiscent of The
Axolotl's Tale. But recent DNA analysis on
larvacea favors Darwin's initial interpretation,
that one branch of ancient tadpole-like
protochordates evolved a new
metamorphosis stage to turn into sedentary
sea squirts.
[edit] Non-chordate animals

Note: From the Lancelets onward, Dawkins only provides dates under duress stating that, "dating
becomes so difficult and controversial that my courage fails me".
Rendezvous
point
570 Ambulacrarians
25
mya (Ambulacraria)
26 590 Protostomes The current pilgrimage, known as

deuterostomes, is joined by a much larger
mya (Protostomia) group of animals, the protostomes, to meet
the ancestor of almost all organisms in the
kingdom Animalia, a worm. Just a single class
in the joining sub-kingdom of protostomia, the
Insecta, represents three quarter of all animal
species on Earth. The great divide between
protostomes (meaning 'mouth first') and
deuterostomes (meaning 'mouth second') was
devised by comparative embryologists based
on the way animal embryos diverge after
gastrulation where the blastula (a hollow ball
of cells) indents to form a cup. In the sub-
kingdom of protostomia, the indentation
eventually becomes the mouth. In
deuterostomia which includes humans, the
indentation eventually becomes the anus; the
mouth is formed later. An extremely large
variety of animal phyla constitute protostomia,
including annelid worms (e.g. garden
earthworms), flatworms (e.g. tapeworms and
flukes), molluscs (e.g. snails, oysters,
ammonites and octopuses), and arthropods
(e.g. insects, crustaceans, spiders and
centipedes). Unlike the species, classes and
genera of animals from the pilgrimage prior to
this rendezvous point, joining animals from
different phyla have no obvious relationship to
one another based on traditional anatomy. But
modern molecular rangefinding has allowed
molecular taxonomists to organize all phyla in
the pilgrimage into a hierarchy, with the worm
as the concestor of all animals in Bilateria who
are bilaterally symmetrical, with left and right
side, a dorsal and a ventral side, and a head
and a tail end.
The Ragworm's Tale talks about the

evolution of left-right symmetry in bilaterians.
The Brine Shrimp's Tale discusses

the possibility of chordates having a back-
swimming ancestor.
The Leaf Cutter's Tale discusses town

like ant socities and their agricultural use of
fungi.
The Grasshopper's Tale talks about

the futility of discriminating between races.
The Fruit Fly's Tale introduces Hox

genes.
The Rotifer's Tale talks about the

outstanding paradox of sexual and asexual
reproduction.
The Barnacle's Tale talks about

palaentology and the deceptiveness of weird
looking organisms.
The Velvet Worm's Tale talks about

the Cambrian explosion.
Acoelomorph
630
27 Flatworms
mya
(Acoelomorpha)
680 Cnidarians The Jellyfish's Tale

28
mya (Cnidaria) The Polypifer's Tale
730 Ctenophores
29
mya (Ctenophora)
780 Placozoans
30
mya (Placozoa)
800 Sponges
31 The Sponge's Tale
mya (Porifera)
[edit] Non-animal eukaryotes

Rendezvous
point
The Choanoflagellate's Tale is

about the evolution of multicellularity.
Choanoflagellates are the closest living
relatives of the multicellular animals, and
900 Choanoflagellates can form temporary colonies from a free-
32
mya (Choanoflagellatea) living unicellular stage. Sponges have
choanocytes, cells that resemble single-
celled choanoflagellates, providing an
indication about how multicellularity may
have evolved.
DRIPs
33 (?)
(Mesomycetozoea)
34 (?) Fungi
35 (?) Amoebozoans
(Amoebozoa)
The Cauliflower's Tale tells the

story about how geometrical
considerations of constructing the most
efficient supply tube network in tissues
dictate a scaling exponent of 3/4 for such
36 (?) Plants Plantae different structures as a cauliflower and
our brain.
The Redwood's Tale introduces

methods of dating historical artifacts and
fossils.
37 (?) Uncertain The Mixotrich's Tale

In what Dawkins calls the "Great Historic Rendezvous", he describes the significantly important event
of endosymbiosis, which results in the beginnings of eukaryotic cells. In his estimates, this occurred in two or
three steps, roughly two billion years ago. Firstly, bacteria, perhaps related to Rickettsia, entered proto-
protozoan cells. For one reason or another, the bacteria were not digested and did not kill the cell. The cell
offered protection to the bacteria, and the bacteria provided energy to the cell, resulting in a mutualistic
symbiotic relationship. This is the speculated origin of mitochondria. Subsequently, photosynthetic bacteria
(thought to be related to cyanobacteria) entered some, but not all, of these mitochondria-containing cells.
These ancient bacteria evolved to become chloroplasts, and the cells became the Plant and Algal lineages.
Meanwhile, the cells which this second endosymbiotic relationship did not occur in went on to form the
Kingdoms Fungi and Animalia, as well as various Protozoa.
Chloroplasts and mitochondria have their own genomes, and they replicate independent of the cell in
which they live. Dawkins acknowledges how the endosymbiotic theory proposed by Lynn Margulis is now
virtually universally accepted.
[edit] Prokaryotes
Rendezvous point Time Joining party Story
38 (?) Archaea
The Rhizobium's Tale

39 (?) Eubacteria
Taq's Tale
[edit] Editions
• (2004) US hardcover ISBN 0-618-00583-8
• (2004) UK hardcover ISBN 0297825038
• (2005) US paperback ISBN 061861916X
• (2005) UK paperback ISBN 0753819961
• (2005) UK Audio ISBN 0-7528-7321-0
[edit] Translations
• Czech translation: Příběh předka
• Dutch translation: Het verhaal van onze voorouders
• French translation: Il était une fois nos ancêtres
• German translation: Geschichten vom Ursprung des Lebens
• Italian translation: Il racconto dell'antenato
• Korean translation: 조상 이야기
• Portuguese translation: A grande história da evolução
• Spanish translation: El cuento del antepasado
• Turkish translation: Ataların hikâyesi
[edit] See also

• Phylogenetic tree
[edit] References
1. ^ Letunic, I; Bork, P (2007). "Interactive Tree Of Life (iTOL): an online tool for phylogenetic
tree display and annotation" (Pubmed). Bioinformatics 23 (1): 127–8.
doi:10.1093/bioinformatics/btl529. PMID 17050570.
2. ^ Ciccarelli, FD; Doerks, T; Von Mering, C; Creevey, CJ; Snel, B; Bork, P (2006). "Toward
automatic reconstruction of a highly resolved tree of life" (Pubmed). Science 311 (5765): 1283–7.
doi:10.1126/science.1123061. PMID 16513982.
• Video introduction by Richard Dawkins
• Richard Dawkins talks to Ira Flatow on "Science Friday"
• Family and kid's experiential programs based on Ancestors Tale
[hide]v · d · eRichard Dawkins
The Selfish Gene (1976) · The Extended Phenotype (1982) · The Blind
Watchmaker (1986) · River Out of Eden (1995) · Climbing Mount Improbable
Books (1996) · Unweaving the Rainbow (1998) · A Devil's Chaplain (2003) · The
Ancestor's Tale (2004) · The God Delusion (2006) · The Greatest Show on Earth:
The Evidence for Evolution (2009) · List of publications
Growing Up in the Universe (1991) · Dawkins vs. Gould (2001) · Beyond

Related works Belief (2006) · Richard Dawkins: How a Scientist Changed the Way We Think
(2006) · The Oxford Book of Modern Science Writing (2008)
Nice Guys Finish First (1987) · Break the Science Barrier (1996) · The
Documentaries Atheism Tapes (2004) · The Root of All Evil? (2006) · The Enemies of Reason
(2007) · The Genius of Charles Darwin (2008) · Faith School Menace (2010)
See also Meme · Out Campaign · Gerin oil · Foundation for Reason and Science ·
Ultimate Boeing 747 gambit · Lalla Ward · Frameshift · Weasel program · Marian
Stamp Dawkins · Middle World · Go God Go · The Courtier's Reply
Book:Richard Dawkins · Category:Richard Dawkins · Portal:Biography
Retrieved from "http://en.wikipedia.org/wiki/The_Ancestor%27s_Tale"
Categories: 2004 books | Books by Richard Dawkins | Biological evolution | Human evolution books |
Biology books
W000
Paleobotany

Paleobotany, also spelled as palaeobotany (from the Greek words paleon = old and "botany", study
of plants), is the branch of paleontology or paleobiology dealing with the recovery and identification of plant
remains from geological contexts, and their use for the biological reconstruction of past environments
(paleogeography), and both the evolutionary history of plants, with a bearing upon the evolution of life in
general. A synonym is paleophytology. Paleobotany includes the study of terrestrial plant fossils, as well as
the study of prehistoric marine photoautotrophs, such as photosynthetic algae, seaweeds or kelp. A closely-
related field is palynology, which is the study of fossilized and extant spores and pollen.
Paleobotany is important in the reconstruction of ancient ecological systems and climate, known as
paleoecology and paleoclimatology respectively; and is fundamental to the study of green plant development
and evolution. Paleobotany has also become important to the field of archaeology, primarily for the use of
phytoliths in relative dating and in paleoethnobotany,
Contents
[hide]
• 1 Overview of the paleobotanical record
• 2 Plant fossils
• 2.1 Preservation of plant fossils
• 2.2 Fossil-taxa
• 3 Fossil groups of plants
• 4 Notable Paleobotanists
• 5 See also
• 6 References
• 6.1 Further reading
[edit] Overview of the paleobotanical record

Macroscopic remains of true vascular plants are first found in the fossil record during the Silurian
Period of the Paleozoic era. Some dispersed, fragmentary fossils of disputed affinity, primarily spores and
cuticles, have been found in rocks from the Ordovician Period in Oman, and are thought to derive from
liverwort- or moss-grade fossil plants (Wellman et al., 2003).
An unpolished hand sample of the Lower Devonian Rhynie Chert from Scotland.
An important early land plant fossil locality is the Rhynie Chert, found outside the village of Rhynie in
Scotland. The Rhynie chert is an Early Devonian sinter (hot spring) deposit composed primarily of silica. It is
exceptional due to its preservation of several different clades of plants, from mosses and lycopods to more
unusual, problematic forms. Many fossil animals, including arthropods and arachnids, are also found in the
Rhynie Chert, and it offers a unique window on the history of early terrestrial life.
Plant-derived macrofossils become abundant in the Late Devonian and include tree trunks, fronds,
and roots. The earliest tree is Archaeopteris, which bears simple, fern-like leaves spirally arranged on
branches atop a conifer-like trunk (Meyer-Berthaud et al., 1999).
Widespread coal swamp deposits across North America and Europe during the Carboniferous Period
contain a wealth of fossils containing arborescent lycopods up to 30 meters tall, abundant seed plants, such
as conifers and seed ferns, and countless smaller, herbaceous plants.
Angiosperms (flowering plants) evolved during the Mesozoic, and flowering plant pollen and leaves
first appear during the Early Cretaceous, approximately 130 million years ago.
[edit] Plant fossils

A plant fossil is any preserved part of a plant that has long since died. Such fossils may be prehistoric
impressions that are many millions of years old, or bits of charcoal that are only a few hundred years old.
Prehistoric plants are various groups of plants that lived before recorded history (before about 3500 BC).
[edit] Preservation of plant fossils
Ginkgoites huttonii, Middle Jurassic, Yorkshire, UK. Leaves preserved as compressions. Specimen
in Munich Palaeontological Museum, Germany. Photo by Ghedoghedo
Plant fossils can be preserved in a variety of ways, each of which can give different types of
information about the original parent plant. These modes of preservation are discussed in the general pages
on fossils but may be summarised in a palaeobotanical context as follows.
1. Adpressions (compressions - impressions). These are the most commonly found type of plant
fossil. They provide good morphological detail, especially of dorsiventral (flattened) plant parts such as
leaves. If the cuticle is preserved, they can also yield fine anatomical detail of the epidermis. Little other detail
of cellular anatomy is normally preserved.
Rhynia, Lower Devonian Rhynie Chert, Scotland, UK. Transverse section through a stem preserved
as a silica petrifaction. Photo by Plantsurfer
2. Petrifactions (permineralisations or anatomically preserved fossils). These provide fine detail of
the cell anatomy of the plant tissue. Morphological detail can also be determined by serial sectioning, but this
is both time consuming and difficult.
3. Moulds and casts. These only tend to preserve the more robust plant parts such as seeds or
woody stems . They can provide information about the three-dimensional form of the plant, and in the case of
casts of tree stumps can provide evidence of the density of the original vegetation. However, they rarely
preserve any fine morphological detail or cell anatomy. A subset of such fossils are pith casts, where the
centre of a stem is either hollow or has delicate pith. After death, sediment enters and forms a cast of the
central cavity of the stem. The best known examples of pith casts are in the Carboniferous Sphenophyta
(Calamites) and cordaites (Artisia).
Crossotheca hughesiana Kidston, Middle Pennsylvanian, Coseley, near Dudley, UK. A

lyginopteridalean pollen organ preserved as an authigenic mineralisation. Specimen in Sedgwick Museum,
Cambridge, UK. Photo by Verisimilus.
4. Authigenic mineralisations. These can provide very fine, three-dimensional morphological detail,
and have proved especially important in the study of reproductive structures that can be severely distorted in
adpressions. However, as they are formed in mineral nodules, such fossils can rarely be of large size.
5. Fusain. Fire normally destroys plant tissue but sometimes charcoalified remains can preserve fine
morphological detail that is lost in other modes of preservation; some of the best evidence of early flowers
has been preserved in fusain. Fusian fossils are delicate and often small, but because of their bouyancy can
often drift for long distances and can thus provide evidence of vegetation away from areas of sedimentation.
[edit] Fossil-taxa
Plant fossils almost always represent disarticulated parts of plants; even small herbaceous plants are
rarely preserved whole. Those few examples of plant fossils that appear to be the remains of whole plants in
fact are incomplete as the internal cellular tissue and fine micromorphological detail is normally lost during
fossilisation. An added complication is that, as explained above, plant remains can be preserved in a variety
of ways, each revealing different features of the original parent plant.
Because of these difficulties, palaeobotanists usually assign different taxonomic names to different
parts of the plant in different modes of preservation. For instance, in the subarborescent Palaeozoic
sphenophytes, an impression of a leaf might be assigned to the genus Annularia, a compression of a cone
assigned to Palaeostachya, and the stem assigned to either Calamites or Arthroxylon depending on whether
it is preserved as a cast or a petrifaction. All of these fossils may have originated from the same parent plant
but they are each given their own taxonomic name. This approach to naming plant fossils originated with the
work of Brongniart (1822) and has stood the test of time; although non-palaeobotanist may find it a confusing
system, it is generally regarded as the most practical way to overcome the special taphonomic difficulties
encountered with plant fossils.
For many years this approach to naming plant fossils was tacitly accepted by palaeobotanists but not
formaised within the International Rules of Botanical Nomenclature (e.g. Briquet, 1906). Eventually, Thomas
(1935) and Jongmans et al. (1935) proposed a set of formal provisions, the essence of which was introduced
into the first International Code of Botanical Nomenclature (Lanjou et al., 1952). These early provisions
allowed fossils representing particular parts of plants in a particular state of preservation to be referred to
organ-genera. In addition, a small subset of organ-genera, to be known as form-genera, were recognised
based on the artificial taxa introduced by Brongniart (1822) mainly for foliage fossils. Over the years, the
concepts and regulations surrounding organ- and form-genera became modified within successive
International Codes of Botanical Nomenclature , reflecting a failure of the palaeobotanical community to agree
on how this aspect of plant taxonomic nomenclature should work (a history reviewed by Cleal & Thomas,
2010). Eventually, the use of organ- and fossil-genera was abandoned with the St Louis Code (Greuter et al.,
2000).
The situation in the current Code (McNeill et al., 2006) is that any plant taxon whose type is a fossil is
referred to as a fossil-taxon. Such taxa can refer to a particular part of a plant preserved in a particular way,
as defined in the diagnosis of that taxon. Otherwise, the names of fossil-taxa are subject to essentially the
same regulations as control the nomenclature of living plants, notably that the names are fixed to a type
specimen, and that competing names are chosen mainly on the basis of chronological priority of first
publication. Although the name is always fixed to the type specimen, the circumscription (i.e. range of
specimens that may be included within the taxon) is defined by the diagnosis and can be changed by formal
emendation. Such emendation could result in an expansion of the range of plant parts and/or preservation
states that can be incorporated within the taxon. For instance, a fossil-genus originally based on
compressions of ovules could be emended so that it also included the multi-ovulate cupules within which the
ovules were originally borne. A complication can arise if, in this case, there was an already named fossil-
genus for these cupules. If palaeobotanists were confident that the type of the ovule fossil-genus and of the
cupule fossil-genus could be included within the newly emended genus, then the two names would compete
as to being the correct one for the newly emended genus. However, this only happens if the actual type
specimens (and not just specimens that are similar to the types) can be included within the newly revised
taxon.
The current Code also refers to a specific subset of fossil-taxa that are known as morphotaxa. These
differ from normal fossil-taxa in that they can only be used for fossils that represent the same part of the
parent plant and that are preserved in the same way as the type specimen. Morphotaxa were introduced to
try to overcome the issue of competing names that represented different plant parts and/or preservation
states. What would you do if the species-name of a pollen-organ was pre-dated by the species name of the
type of pollen produced by that pollen organ. It was argued that palaeobotanists would be unhappy if the
pollen organs were named using the taxonomic name whose type specimen is a pollen grain. As pointed out
by Cleal & Thomas (2010), however, the risk of the name of a pollen grain supplanting the name of a pollen
organ is most unlikely. Palaeobotanists would have to be totally confident that the type specimen of the
pollen species, which would normally be a dispersed grain, definitely came from the same plant as produced
the pollen organ. We know from modern plants that closely related but distinct species can produce virtually
indistinguishable pollen. It would seem that morphotaxa offer no real advantage to palaeobotanists over
normal fossil-taxa and the concept is best abandoned.
[edit] Fossil groups of plants
Stigmaria, a common fossil tree root. Upper Carboniferous of northeastern Ohio.

External mold of Lepidodendron from the Upper Carboniferous of Ohio.
Main articles: Evolutionary history of plants and List of extinct plants
Some plants have remained remarkedly unchanged throughout earth's geological time scale. Early
ferns had developed by the Mississippian, conifers by the Pennsylvanian. Some plants of prehistory are the
same ones around today and are thus living fossils, such as Ginkgo biloba and Sciadopitys verticillata. Other
plants have changed radically, or have gone extinct entirely.
Examples of prehistoric plants are:
• Araucaria mirabilis
• Archaeopteris
• Calamites
• Glossopteris
• Hymenaea protera
• Nelumbo aureavallis
• Protosalvinia
• Palaeoraphe
• Trochodendron nastae
• Dillhoffia
• Peltandra primaeva
[edit] Notable Paleobotanists

• Edward W. Berry (1875–1945), paleoecology and phytogeography
• Constantin von Ettingshausen (1826–1897), Tertiary floras
• Dunkinfield Henry Scott (1854–1934), analysis of the structures of fossil plants
• Kaspar Maria von Sternberg (1761–1838), the "father of paleobotany"
• Jack A. Wolfe (1936–2005) Tertiary paleoclimate of western North America.
[edit] See also

Paleontology portal

• Prehistoric life
• Cryptospores
[edit] References
• Brigitte Meyer-Berthaud, S.E. Scheckler, J. Wendt, " Archaeopteris is the Earliest Modern
Tree." Nature, 398, 700-701 (22 April 1999) | doi:10.1038/19516
• J. Briquet, 1906. Règles internationales de la nomenclature botanique adoptées par le
Congrès International de Botanique de Vienne 1905. Fischer, Jena.
• A. Brongniart, 1822. Sur la classification et la distribution des végétaux fossiles en général,
et sur ceux des terrains de sediment supérieur en particulier. Mém. Mus. Natl. Hist. Nat. 8: 203–240,
297–348.
• C. J. Cleal & B. A. Thomas, 2010. Botanical nomenclature and plant fossils. Taxon, 59, 261-
268.
• W. Greuter, J. McNeill, F. R. Barrie, H. M. Burdet, V. Demoulin, T. S. Filgueiras, D. H.
Nicolson, P. C. Silva, J. E. Skog, P. Trehane, N. J. Turland & D. L. Hawksworth, 2000. International
Code of Botanical Nomenclature (Saint Louis Code). Koeltz Scientific Books, Königstein.
• W. J. Jongmans, T. G. Halle & W. Gothan, 1935. Proposed editions to the International Rules
of Botanical Nomenclature adopted by the fifth International Botanical Congress Cambridge 1930.
Heerlen.
• J. Lanjouw, C. Baehni, E. D. Merrill, H. W. Rickett, W. Robyns, T. A. Sprague & F. A. Stafleu,
1952. International Code of Botanical Nomenclature: Adopted by the Seventh International Botanical
Congress; Stockholm, July 1950. Regnum Vegetabile 3. International Bureau for Plant Taxonomy of
the International Association for Plant Taxonomy, Utrecht.
• J. McNeill, F. R. Barrie, H. M. Burdet, V. Demoulin, D. L. Hawksworth, J. Marhold, D. H.
Nicolson, J. Prado, P. C. Silva, J. E. Skog, J. H. Wiersema & N. J. Turland, 2006. International Code
of Botanical Nomenclature (Vienna Code): Adopted by the Seventeenth International Botanical
Congress; Vienna, Austria, July 2005. Regnum Vegetabile 146. Gantner, Ruggell.
• H. H. Thomas, 1935. Proposed additions to the International Rules of Botanical
Nomenclature suggested by British palæobotanists. Journal of Botany, 73, 111.
• Charles H. Wellman, Peter L. Osterloff and Uzma Mohiuddin, "Fragments of the Earliest
Land Plants." Nature, 425, 282-285 (18 September 2003) | doi: 10.1038/nature01884
• Jurassic Park plants Plants that lived when dinosaurs roamed the earth.

• Stewart, W.N. and Rothwell, G.W. 1993. Paleobotany and the evolution of plants, Second
edition. Cambridge University Press, Cambridge, UK. ISBN 0-521-38294-7
• Taylor, T. N. and E. L. Taylor. 1993. The Biology and Evolution of Fossil Plants, Prentice-
Hall, Inc., Englewood Cliffs, New Jersey, USA. ISBN 0-13-651589-4

• International Organisation of Paleobotany
• Botanical Society of America - Paleobotanical Section
• Paleobotany Research Group, University Münster, Germany.
• The Biota of Early Terrestrial Ecosystems: The Rhynie Chert, University of Aberdeen, UK.
• Bibliography of Paleobotany
• The Sternberg Project
• Paleobotany - 1911 Encyclopaedia Britannica article
• PaleoNet - listservs and links related to paleontology

Plant physiology



Transpiration
Sporophyte


morphology terms
Category · Portal
Retrieved from "http://en.wikipedia.org/wiki/Paleobotany"
Categories: Prehistoric plants | Subfields of paleontology | Branches of botany

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Paleobiology

This article is about the discipline of Palæobiology. For the journal, see Paleobiology (journal).
Brachiopods and bryozoans in an Ordovician limestone, southern Minnesota.
Paleobiology (sometimes spelled palaeobiology) is a growing and comparatively new discipline
which combines the methods and findings of the natural science biology with the methods and findings of the
earth science paleontology. It is occasionally referred to as "geobiology."
Paleobiological research uses biological field research of current biota and of fossils millions of years
old to answer questions about the molecular evolution and the evolutionary history of life. In this scientific
quest, macrofossils, microfossils and trace fossils are typically analyzed. However, the 21st-century
biochemical analysis of DNA and RNA samples offers much promise, as does the biometric construction of
phylogenetic trees.
An investigator in this field is known as a paleobiologist.
Contents
[hide]
• 1 Important research areas
• 2 Paleobiologists
• 3 Paleobiologic journals
• 4 Footnotes
• 5 See also
• 6 References
[edit] Important research areas

• Paleobotany applies the principles and methods of paleobiology to flora, especially green
land plants, but also including the fungi and seaweeds (algae). See also mycology, phycology and
dendrochronology.
• Paleozoology uses the methods and principles of paleobiology to understand fauna, both
vertebrates and invertebrates. See also vertebrate and invertebrate paleontology, as well as
paleoanthropology.
• Micropaleontology applies paleobiologic principles and methods to archaea, bacteria, protists
and microscopic pollen/spores. See also microfossils and palynology.
• Paleobiochemistry uses the methods and principles of organic chemistry to detect and
analyze molecular-level evidence of ancient life, both microscopic and macroscopic.
• Paleoecology examines past ecosystems, climates, and geographies so as to better
comprehend prehistoric life.
• Taphonomy analyzes the post-mortem history (for example, decay and decomposition) of an
individual organism in order to gain insight on the behavior, death and environment of the fossilized
organism.
• Paleoichnology analyzes the tracks, borings, trails, burrows, impressions, and other trace
fossils left by ancient organisms in order to gain insight into their behavior and ecology.
• Stratigraphic paleobiology studies long-term secular changes, as well as the (short-term)
bed-by-bed sequence of changes, in organismal characteristics and behaviors. See also
stratification, sedimentary rocks and the geologic time scale.
• Evolutionary developmental paleobiology examines the evolutionary aspects of the modes
and trajectories of growth and development in the evolution of life – clades both extinct and extant.
See also adaptive radiation, cladistics, evolutionary biology, developmental biology and phylogenetic
tree.
[edit] Paleobiologists
Baron Nopcsa
The founder or "father" of modern paleobiology is said to be Baron Franz Nopcsa (1877 to 1933), a
turn-of-the-century Hungarian scientist. He is also known as Baron Nopcsa, Ferenc Nopcsa, and Franz
Nopcsa von Felsö-Szilvás. He initially termed the discipline "paleophysiology."
However, credit for coining the word paleobiology itself should go to Professor Charles Schuchert.
He proposed the term in 1904 so as to initiate "a broad new science" joining "traditional paleontology with the
evidence and insights of geology and isotopic chemistry."[1]
On the other hand, Charles Doolittle Walcott, a Smithsonian adventurer, has been cited as the
"founder of Precambrian paleobiology." Although best-known as the discoverer of the mid-Cambrian Burgess
shale animal fossils, in 1883 this American curator found the "first Precambrian fossil cells known to science"
– a stromatolite reef then known as Cryptozoon algae. In 1899 he discovered the first acritarch fossil cells, a
Precambrian algal phytoplankton he named Chuaria. Lastly, in 1914, Walcott reported "minute cells and
chains of cell-like bodies" belonging to Precambrian purple bacteria.[2]
Later 20th-century paleobiologists have also figured prominently in finding Archaean and Proterozoic
eon microfossils: In 1954, Stanley A. Tyler and Elso S. Barghoorn described 2.1 billion-year-old
cyanobacteria and fungi-like microflora at their Gunflint Chert fossil site. Eleven years later, Barghoorn and J.
William Schopf reported finely-preserved Precambrian microflora at their Bitter Springs site of the Amadeus
Basin, Central Australia.[3]
Finally, in 1993, Schopf discovered O2-producing blue-green bacteria at his 3.5 billion-year-old Apex
Chert site in Pilbara Craton, Marble Bar, in the northwestern part of Western Australia. So paleobiologists
were at last homing in on the origins of the Precambrian "Oxygen catastrophe."[4]
[edit] Paleobiologic journals

• Acta Palaeontologica Polonica
• Biology and Geology
• Historical Biology
• Palaios
• Palaeogeography, Palaeoclimatology, Palaeoecology
• Paleobiology (journal)
• Paleoceanography
[edit] Footnotes
1. ^ Schuchert is cited on page 170 of Cradle of Life: The Discovery of Earth's Earliest Fossils
(Princeton: Princeton University Press) by J. William Schopf (1999). ISBN 0-691-00230-4.
2. ^ Walcott's contributions are described by J. William Schopf (1999) on pages 23 to 31.
Another good source is E. L. Yochelson (1997), Charles Doolittle Walcott: Paleontologist (Kent, Ohio:
Kent State University Press).
3. ^ The paleobiologic discoveries of Tyler, Barghoorn and Schopf are related on pages 35 to
70 of Schopf (1999).
4. ^ The Apex chert microflora is related by Schopf (1999) himself on pages 71 to 100.
[edit] See also

Paleontology portal
• History of biology
• History of invertebrate paleozoology
[edit] References
• Derek E.G. Briggs and Peter R. Crowther, eds. (2003). Palaeobiology II. Malden,
Massachusetts: Blackwell Publishing. ISBN 0-632-05147-7 and ISBN 0-632-05149-3. The second
edition of an acclaimed British textbook.
• Robert L. Carroll (1998). Patterns and Processes of Vertebrate Evolution. Cambridge
Paleobiology Series. Cambridge, England: Cambridge University Press. ISBN 9788521478090 and
ISBN 052147809X. Applies paleobiology to the adaptive radiation of fishes and quadrapeds.
• Matthew T. Carrano, Timothy Gaudin, Richard Blob, and John Wible, eds. (2006). Amniote
Paleobiology: Perspectives on the Evolution of Mammals, Birds and Reptiles . Chicago: University of
Chicago Press. ISBN 0226094782 and ISBN 978-0226094786. This new book describes
paleobiological research into land vertebrates of the Mesozoic and Cenozoic eras.
• Robert B. Eckhardt (2000). Human Paleobiology. Cambridge Studies in Biology and
Evolutionary Anthropology. Cambridge, England: Cambridge University Press. ISBN 0521451604
and ISBN 9780521451604. This book connects paleoanthropology and archeology to the field of
paleobiology.
• Douglas H. Erwin (2006). Extinction: How Life on Earth Nearly Ended 250 Million Years Ago.
Princeton: Princeton University Press. ISBN 978-0-691-00524-9. An investigation by a paleobiologist
into the many theories as to what happened during the catastrophic Permian-Triassic transition.
• Brian Keith Hall and Wendy M. Olson, eds. (2003). Keywords and Concepts in Evolutionary
Biology. Cambridge, Massachusetts: Harvard University Press. ISBN 0674009045 and ISBN
9780674009042.
• David Jablonski, Douglas H. Erwin, and Jere H. Lipps (1996). Evolutionary Paleobiology.
Chicago: University of Chicago Press, 492 pages. ISBN 0226389111 and ISBN 0226389138. A fine
American textbook.
• Masatoshi Nei and Sudhir Kumar (2000). Molecular Evolution and Phylogenetics. Oxford,
England: Oxford University Press. ISBN 0195135857 and ISBN 978-0195-135855. This text links
DNA/RNA analysis to the evolutionary "tree of life" in paleobiology.
• Donald R. Prothero (2004). Bringing Fossils to Life: An Introduction to Paleobiology . New
York: McGraw Hill. ISBN 0073661708 and ISBN 978-007366-1704. An acclaimed book for the novice
fossil-hunter and young adults.
• Mark Ridley, ed. (2004). Evolution. Oxford, England: Oxford University Press. ISBN
0199267944 and ISBN 9781-405-103459. An anthology of analytical studies in paleobiology.
• Raymond Rogers, David Eberth, and Tony Fiorillo (2007). Bonebeds: Genesis, Analysis and
Paleobiological Significance. Chicago: University of Chicago Press. ISBN 0226723704 and ISBN
9780226723709. A new book regarding the fossils of vertebrates, especially tetrapods on land during
the Mesozoic and Cenozoic eras.
• Thomas J. M. Schopf, ed. (1972). Models in Paleobiology. San Francisco: Freeman, Cooper.
ISBN 0877353256 and ISBN 978-0877353256. A much-cited, seminal classic in the field discussing
methodology and quantitative analysis.
• Thomas J.M. Schopf (1980). Paleoceanography. Cambridge, Massachusetts: Harvard
University Press. ISBN 0674652150 and ISBN 9780674652156. A later book by the noted
paleobiologist. This text discusses ancient marine ecology.
• J. William Schopf (2001). Cradle of Life: The Discovery of Earth's Earliest Fossils . Princeton:
Princeton University Press. ISBN 0691088640. The use of biochemical and ultramicroscopic analysis
to analyze microfossils of bacteria and archaea.
• Paul Selden and John Nudds (2005). Evolution of Fossil Ecosystems. Chicago: University of
Chicago Press. ISBN 978-02267-46418 and ISBN 022-6746410. A recent analysis and discussion of
paleoecology.
• Paul Tasch (1980). Paleobiology of the Invertebrates. New York: John Wiley & Sons. ISBN
0471-052728 and ISBN 9780471-052722. Applies statistics to the evolution of sponges, cnidarians,
worms, brachiopods, bryozoa, mollusks, and arthropods.
• Shuhai Xiao and Alan J. Kaufman, eds. (2006). Neoproterozoic Geobiology and
Paleobiology. New York: Springer Science+Business Media. ISBN 978-1-4020-5201-9. This new
book describes research into the fossils of the earliest multicellular animals and plants, especially the
Ediacaran period invertebrates and algae.
• Bernard Ziegler and R. O. Muir (1983). Introduction to Palaeobiology. Chichester, England:
E. Horwood. ISBN 0470275529 and ISBN 9780470275528. A classic, British introductory textbook.
• Paleobiology website of the National Museum of Natural History (Smithsonian) in
Washington, D.C.
• The Paleobiology Database
Retrieved from "http://en.wikipedia.org/wiki/Paleobiology"
Categories: Paleobiology | Biology | Developmental biology | Evolutionary biology | Subfields of

paleontology
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Brain

This article is about the brains of all types of animals, including humans. For information specific to
the human brain, see Human brain. For other uses, see Brain (disambiguation).
A chimpanzee brain
The brain is the center of the nervous system in all vertebrate and most invertebrate animals.[1]
Some primitive animals such as jellyfish and starfish have a decentralized nervous system without a brain,
while sponges lack any nervous system at all. In vertebrates the brain is located in the head, protected by the
skull and close to the primary sensory apparatus of vision, hearing, balance, taste, and smell.
Brains can be extremely complex. The cerebral cortex of the human brain contains roughly 15–33
billion neurons, perhaps more, depending on gender and age,[2] linked with up to 10,000 synaptic
connections each. Each cubic millimeter of cerebral cortex contains roughly one billion synapses.[3] These
neurons communicate with one another by means of long protoplasmic fibers called axons, which carry trains
of signal pulses called action potentials to distant parts of the brain or body and target them to specific
recipient cells.
The brain controls the other organ systems of the body, either by activating muscles or by causing
secretion of chemicals such as hormones and neurotransmitters. This centralized control allows rapid and
coordinated responses to changes in the environment. Some basic types of responsiveness are possible
without a brain: even single-celled organisms may be capable of extracting information from the environment
and acting in response to it.[4] Sponges, which lack a central nervous system, are capable of coordinated
body contractions and even locomotion.[5] In vertebrates, the spinal cord by itself contains neural circuitry
capable of generating reflex responses as well as simple motor patterns such as swimming or walking.[6]
However, sophisticated control of behavior on the basis of complex sensory input requires the information-
integrating capabilities of a centralized brain.
Despite rapid scientific progress, much about how brains work remains a mystery. The operations of
individual neurons and synapses are now understood in considerable detail, but the way they cooperate in
ensembles of thousands or millions has been very difficult to decipher. Methods of observation such as EEG
recording and functional brain imaging tell us that brain operations are highly organized, while single unit
recording can resolve the activity of single neurons, but how individual cells give rise to complex operations
is unknown.[7]
Contents
[hide]
• 1 Macroscopic structure
• 1.1 Bilateria
• 1.2 Invertebrates
• 1.3 Vertebrates
• 1.3.1 Vertebrate brain regions
• 1.4 Mammals
• 1.5 Primates
• 2 Microscopic structure
• 3 Development
• 4 Functions
• 4.1 Functional subsystems
• 4.1.1 Neurotransmitter systems
• 4.1.2 Sensory systems
• 4.1.3 Motor systems
• 4.1.4 Arousal systems
• 4.2 Brain energy consumption
• 5 Brain and mind
• 6 Research
• 7 History
• 8 See also
Macroscopic structure
The brain is the most complex biological structure known,[8] and comparing the brains of different
species on the basis of appearance is often difficult. Nevertheless, there are common principles of brain
architecture that apply across a wide range of species. These are revealed mainly by three approaches. The
evolutionary approach compares brain structures of different species, and using the principle that features
found in all branches that have descended from a given ancient form were probably present in the common
ancestor as well. The developmental approach examines how the form of the brain changes during the
progression from embryonic to adult stages. The genetic approach analyzes gene expression in various parts
of the brain across a range of species. Each approach complements and informs the other two.
Bilateria
Nervous system of a bilaterian animal, in the form of a nerve cord with segmental enlargements, and
a "brain" at the front
With the exception of a few primitive forms such as sponges and jellyfish, all living animals are
bilateria, meaning animals with a bilaterally symmetric body shape (that is, left and right sides that are
approximate mirror images of each other).[9]
All bilateria are thought to have descended from a common ancestor that appeared early in the
Cambrian period, 550–600 million years ago.[9] This ancestor had the shape of a simple tube worm with a
segmented body, and at an abstract level, that worm-shape continues to be reflected in the body and
nervous system plans of all modern bilateria, including humans.[10] The fundamental bilateral body form is a
tube with a hollow gut cavity running from mouth to anus, and a nerve cord with an enlargement (a
"ganglion") for each body segment, with an especially large ganglion at the front, called the "brain".
Invertebrates
Drosophila
For invertebrates (e.g., insects, molluscs, worms, etc.) the components of the brain differ so greatly
from the vertebrate pattern that it is hard to make meaningful comparisons except on the basis of genetics.
Two groups of invertebrates have notably complex brains: arthropods (insects, crustaceans, arachnids, and
others), and cephalopods (octopuses, squids, and similar molluscs).[11] The brains of arthropods and
cephalopods arise from twin parallel nerve cords that extend through the body of the animal. Arthropods
have a central brain with three divisions and large optical lobes behind each eye for visual processing.[11]
Cephalopods have the largest brains of any invertebrates. The brain of the octopus in particular is highly
developed, comparable in complexity to the brains of some vertebrates.
There are a few invertebrates whose brains have been studied intensively. The large sea slug
Aplysia was chosen by Nobel Prize-winning neurophysiologist Eric Kandel, because of the simplicity and
accessibility of its nervous system, as a model for studying the cellular basis of learning and memory, and
subjected to hundreds of experiments.[12] The most thoroughly studied invertebrate brains, however, belong
to the fruit fly Drosophila and the tiny roundworm Caenorhabditis elegans (C. elegans).
Because of the large array of techniques available for studying their genetics, fruit flies have been a
natural subject for studying the role of genes in brain development.[13] Remarkably, many aspects of
Drosophila neurogenetics have turned out to be relevant to humans. The first biological clock genes, for
example, were identified by examining Drosophila mutants that showed disrupted daily activity cycles.[14] A
search in the genomes of vertebrates turned up a set of analogous genes, which were found to play similar
roles in the mouse biological clock—and therefore almost certainly in the human biological clock as well.[15]
Like Drosophila, the nematode worm C. elegans has been studied largely because of its importance
in genetics.[16] In the early 1970s, Sydney Brenner chose it as a model system for studying the way that
genes control development. One of the advantages of working with this worm is that the body plan is very
stereotyped: the nervous system of the hermaphrodite morph contains exactly 302 neurons, always in the
same places, making identical synaptic connections in every worm.[17] In a heroic project, Brenner's team
sliced worms into thousands of ultrathin sections and photographed every section under an electron
microscope, then visually matched fibers from section to section, in order to map out every neuron and
synapse in the entire body.[18] Nothing approaching this level of detail is available for any other organism,
and the information has been used to enable a multitude of studies that would not have been possible
without it.
Vertebrates
The brain of a shark

The brains of vertebrates are made of very soft tissue, with a texture that has been compared to Jell-
O.[19] Living brain tissue is pinkish on the outside and mostly white on the inside, with subtle variations in
color. Vertebrate brains are surrounded by a system of connective tissue membranes called meninges that
separate the skull from the brain.[20] This three-layered covering is composed of (from the outside in) the
dura mater ("hard mother"), arachnoid mater ("spidery mother"), and pia mater ("soft mother"). The arachnoid
and pia are physically connected and thus often considered as a single layer, the pia-arachnoid. Below the
arachnoid is the subarachnoid space which contains cerebrospinal fluid (CSF), which circulates in the narrow
spaces between cells and through cavities called ventricles, and serves to nourish, support, and protect the
brain tissue. Blood vessels enter the central nervous system through the perivascular space above the pia
mater. The cells in the blood vessel walls are joined tightly, forming the blood-brain barrier which protects the
brain from toxins that might enter through the blood.
The first vertebrates appeared over 500 million years ago (mya), during the Cambrian period, and
may have somewhat resembled the modern hagfish in form.[21] Sharks appeared about 450 mya,
amphibians about 400 mya, reptiles about 350 mya, and mammals about 200 mya. No modern species
should be described as more "primitive" than others, since all have an equally long evolutionary history, but
the brains of modern hagfishes, lampreys, sharks, amphibians, reptiles, and mammals show a gradient of
size and complexity that roughly follows the evolutionary sequence.[22] All of these brains contain the same
set of basic anatomical components, but many are rudimentary in hagfishes, whereas in mammals the
foremost parts are greatly elaborated and expanded.
All vertebrate brains share a common underlying form, which can most easily be appreciated by
examining how they develop.[23] The first appearance of the nervous system is as a thin strip of tissue
running along the back of the embryo. This strip thickens and then folds up to form a hollow tube. The front
end of the tube develops into the brain. In its earliest form, the brain appears as three swellings, which
eventually become the forebrain, midbrain, and hindbrain. In many classes of vertebrates these three parts
remain similar in size in the adult, but in mammals the forebrain becomes much larger than the other parts,
and the midbrain quite small.
The relationship between brain size, body size and other variables has been studied across a wide
range of vertebrate species. Brain size increases with body size but not proportionally. Averaging across all
orders of mammals, it follows a power law, with an exponent of about 0.75.[24] This formula applies to the
average brain of mammals but each family departs from it, reflecting their sophistication of behavior.[25] For
example, primates have brains 5 to 10 times as large as the formula predicts. Predators tend to have larger
brains. When the mammalian brain increases in size, not all parts increase at the same rate. The larger the
brain of a species, the greater the fraction taken up by the cortex.[26]
Vertebrate brain regions

Neuroanatomists usually consider the brain to consist of six main regions: the telencephalon
(cerebral hemispheres), diencephalon (thalamus and hypothalamus), mesencephalon (midbrain),
cerebellum, pons, and medulla oblongata.[27] Each of these areas in turn has a complex internal structure.
Some areas, such as the cortex and cerebellum, consist of layers, folded or convoluted to fit within the
available space. Other areas consist of clusters of many small nuclei. If fine distinctions are made on the
basis of neural structure, chemistry, and connectivity, thousands of distinguishable areas can be identified
within the vertebrate brain.
Some branches of vertebrate evolution have led to substantial changes in brain shape, especially in
the forebrain. The brain of a shark shows the basic components in a straightforward way, but in teleost fishes
(the great majority of modern species), the forebrain has become "everted", like a sock turned inside out. In
birds, also, there are major changes in shape.[28] One of the main structures in the avian forebrain, the
dorsal ventricular ridge, was long thought to correspond to the basal ganglia of mammals, but is now thought
to be more closely related to the neocortex.[29]
Main anatomical regions of the vertebrate brain
Several brain areas have maintained their identities across the whole range of vertebrates, from
hagfishes to humans.[1] Here is a list of some of the most important areas, along with a very brief description
of their functions as currently understood (but note that the functions of most of them are still disputed to
some degree):
• The medulla, along with the spinal cord, contains many small nuclei involved in a wide
variety of sensory and motor functions.[30]
• The hypothalamus is a small region at the base of the forebrain, whose complexity and
importance belies its size. It is composed of numerous small nuclei, each with distinct connections
and distinct neurochemistry. The hypothalamus is the central control station for sleep/wake cycles,
control of eating and drinking, control of hormone release, and many other critical biological
functions.[31]
• Like the hypothalamus, the thalamus is a collection of nuclei with diverse functions. Some of
them are involved in relaying information to and from the cerebral hemispheres. Others are involved
in motivation. The subthalamic area (zona incerta) seems to contain action-generating systems for
several types of "consummatory" behaviors, including eating, drinking, defecation, and copulation.
[32]
• The cerebellum modulates the outputs of other brain systems to make them more precise.
Removal of the cerebellum does not prevent an animal from doing anything in particular, but it makes
actions hesitant and clumsy. This precision is not built-in, but learned by trial and error. Learning how
to ride a bicycle is an example of a type of neural plasticity that may take place largely within the
cerebellum.[33]
• The tectum, often called "optic tectum", allows actions to be directed toward points in space.
In mammals it is called the "superior colliculus", and its best studied function is to direct eye
movements. It also directs reaching movements, though. It gets strong visual inputs, but also inputs
from other senses that are useful in directing actions, such as auditory input in owls, input from the
thermosensitive pit organs in snakes, etc. In some fishes, such as lampreys, it is the largest part of
the brain.[34]
• The pallium is a layer of gray matter that lies on the surface of the forebrain. In reptiles and
mammals it is called cortex instead. The pallium is involved in multiple functions, including olfaction
and spatial memory. In mammals, where it comes to dominate the brain, it subsumes functions from
many subcortical areas.[35]
• The hippocampus, strictly speaking, is found only in mammals. However, the area it derives
from, the medial pallium, has counterparts in all vertebrates. There is evidence that this part of the
brain is involved in spatial memory and navigation in fishes, birds, reptiles, and mammals.[36]
• The basal ganglia are a group of interconnected structures in the forebrain, of which our
understanding has increased enormously over the last few years. The primary function of the basal
ganglia seems to be action selection. They send inhibitory signals to all parts of the brain that can
generate actions, and in the right circumstances can release the inhibition, so that the action-
generating systems are able to execute their actions. Rewards and punishments exert their most
important neural effects within the basal ganglia.[37]
• The olfactory bulb is a special structure that processes olfactory sensory signals, and sends
its output to the olfactory part of the pallium. It is a major brain component in many vertebrates, but
much reduced in primates.[38]
Mammals
The cerebral cortex is the part of the brain that most strongly distinguishes mammals from other
vertebrates, primates from other mammals, and humans from other primates. The hindbrain and midbrain of
mammals are generally similar to those of other vertebrates, but dramatic differences appear in the forebrain,
which is not only greatly enlarged, but also altered in structure.[39] In non-mammalian vertebrates, the
surface of the cerebrum is lined with a comparatively simple layered structure called the pallium.[40] In
mammals, the pallium evolves into a complex 6-layered structure called neocortex or isocortex. In primates,
the neocortex is greatly enlarged, especially the part called the frontal lobes. In humans, this enlargement of
the frontal lobes is taken to an extreme, and other parts of the cortex also become quite large and complex.
The hippocampus of mammals also has a distinctive structure.
Scale model of bottlenose dolphin (Tursiops truncatus) brain (middle), compared with brains of wild
pig (Sus scrofa) (left), and man (Homo sapiens) (right)
Unfortunately, the evolutionary history of these mammalian features, especially the 6-layered cortex,
is difficult to trace.[41] This is largely because of a missing link problem. The ancestors of mammals, called
synapsids, split off from the ancestors of modern reptiles and birds about 350 million years ago. However,
the most recent branching that has left living results within the mammals was the split between monotremes
(the platypus and echidna), marsupials (opossum, kangaroo, etc.) and placentals (most living mammals),
which took place about 120 million years ago. The brains of monotremes and marsupials are distinctive from
those of placentals in some ways, but they have fully mammalian cortical and hippocampal structures. Thus,
these structures must have evolved between 350 and 120 million years ago, a period that has left no
evidence except fossils, which do not preserve tissue as soft as brain.
Primates
See also: Human brain
The brains of humans and other primates contain the same structures as the brains of other
mammals, but are considerably larger in proportion to body size.[26] Most of the enlargement comes from a
massive expansion of the cortex, focusing especially on the parts serving vision and forethought.[42] The
visual processing network of primates is very complex, including at least 30 distinguishable areas, with a
bewildering web of interconnections. Taking all of these together, visual processing makes use of more than
half of the primate neocortex.[43] The other part of the brain that is greatly enlarged is the prefrontal cortex,
whose functions are difficult to summarize succinctly, but relate to planning, working memory, motivation,
attention, and executive control.
Microscopic structure
Oligodendrocytes are glial cells that form myelin around the axons of neurons.
The brain is composed of two broad classes of cells: neurons and glia.[44] These two types are
equally numerous in the brain as a whole, although glial cells outnumber neurons roughly 4 to 1 in the
cerebral cortex.[45] Glia come in several types, which perform a number of critical functions, including
structural support, metabolic support, insulation, and guidance of development.
The property that makes neurons so important is that, unlike glia, they are capable of sending signals
to each other over long distances.[46] They send these signals by means of an axon, a thin protoplasmic
fiber that extends from the cell body and projects, usually with numerous branches, to other areas,
sometimes nearby, sometimes in distant parts of the brain or body. The extent of an axon can be
extraordinary: to take an example, if a pyramidal cell of the neocortex were magnified so that its cell body
became the size of a human, its axon, equally magnified, would become a cable a few centimeters in
diameter, extending farther than a kilometer. These axons transmit signals in the form of electrochemical
pulses called action potentials, lasting less than a thousandth of a second and traveling along the axon at
speeds of 1–100 meters per second. Some neurons emit action potentials constantly, at rates of 10–100 per
second, usually in irregular temporal patterns; other neurons are quiet most of the time, but occasionally emit
a burst of action potentials.
Neurons often have extensive networks of dendrites, which receive synaptic connections. Shown
here is photomicrograph of a pyramidal neuron from the human hippocampus, stained by the Golgi method.
Axons transmit signals to other neurons, or to non-neuronal cells, by means of specialized junctions
called synapses.[47] A single axon may make as many as several thousand synaptic connections. When an
action potential, traveling along an axon, arrives at a synapse, it causes a chemical called a neurotransmitter
to be released. The neurotransmitter binds to receptor molecules in the membrane of the target cell. Some
types of neuronal receptors are excitatory, meaning that they increase the rate of action potentials in the
target cell; other receptors are inhibitory, meaning that they decrease the rate of action potentials; others
have complex modulatory effects.
Axons actually fill most of the space in the brain.[48] Often large groups of them are bundled together
in what are called nerve fiber tracts. Many axons are wrapped in thick sheaths of a fatty substance called
myelin, which serves to greatly increase the speed of action potential propagation. Myelin is white, so parts of
the brain filled exclusively with nerve fibers appear as white matter, in contrast to the gray matter that marks
areas with high densities of neuron cell bodies. The total length of myelinated axons in an average adult
human brain is well over 100,000 kilometres (62,000 mi) .[49]
Development
Main article: Neural development
This diagram depicts the main subdivisions of the embryonic vertebrate brain. These regions will
later differentiate into forebrain, midbrain and hindbrain structures.
The brain does not simply grow; rather, it develops in an intricately orchestrated sequence of stages.
[50] Many neurons are created in special zones that contain stem cells, and then migrate through the tissue
to reach their ultimate locations.[51] In the cortex, for example, the first stage of development is the formation
of a "scaffold" by a special group of glial cells, called radial glia, which send fibers vertically across the cortex.
New cortical neurons are created at the bottom of the cortex, and then "climb" along the radial fibers until
they reach the layers they are destined to occupy in the adult.
In vertebrates, the early stages of neural development are similar for all species.[50] As the embryo
transforms from a round blob of cells into a wormlike structure, a narrow strip of ectoderm running along the
midline of the back is induced to become the neural plate, the precursor of the nervous system. The neural
plate invaginates to form the neural groove, and then the folds that line the groove merge to enclose the
neural tube, a hollow cord of cells with a fluid-filled ventricle at the center. At the front end, the ventricles and
cord swell to form three vesicles that are the precursors of the forebrain, midbrain, and hindbrain.[50] At the
next stage, the forebrain splits into two vesicles called the telencephalon (which will contain the cerebral
cortex, basal ganglia, and related structures) and the diencephalon (which will contain the thalamus and
hypothalamus). At about the same time, the hindbrain splits into the metencephalon (which will contain the
cerebellum and pons) and the myelencephalon (which will contain the medulla oblongata). Each of these
areas contains proliferative zones at which neurons and glia cells are generated; the resulting cells then
migrate, sometimes for long distances, to their final positions.
Once a neuron is in place, it begins to extend dendrites and an axon into the area around it.[52]
Axons, because they commonly extend a great distance from the cell body and need to make contact with
specific targets, grow in a particularly complex way. The tip of a growing axon consists of a blob of
protoplasm called a "growth cone", studded with chemical receptors. These receptors sense the local
environment, causing the growth cone to be attracted or repelled by various cellular elements, and thus to be
pulled in a particular direction at each point along its path. The result of this pathfinding process is that the
growth cone navigates through the brain until it reaches its destination area, where other chemical cues
cause it to begin generating synapses. Taking the entire brain into account, many thousands of genes give
rise to proteins that influence axonal pathfinding.
The synaptic network that finally emerges is only partly determined by genes, though. In many parts
of the brain, axons initially "overgrow", and then are "pruned" by mechanisms that depend on neural activity.
[53] In the projection from the eye to the midbrain, for example, the structure in the adult contains a very
precise mapping, connecting each point on the surface of the retina to a corresponding point in a midbrain
layer. In the first stages of development, each axon from the retina is guided to the right general vicinity in the
midbrain by chemical cues, but then branches very profusely and makes initial contact with a wide swath of
midbrain neurons. The retina, before birth, contains special mechanisms that cause it to generate waves of
activity that originate spontaneously at some point and then propagate slowly across the retinal layer.[54]
These waves are useful because they cause neighboring neurons to be active at the same time: that is, they
produce a neural activity pattern that contains information about the spatial arrangement of the neurons. This
information is exploited in the midbrain by a mechanism that causes synapses to weaken, and eventually
vanish, if activity in an axon is not followed by activity of the target cell. The result of this sophisticated
process is a gradual tuning and tightening of the map, leaving it finally in its precise adult form.
Similar things happen in other brain areas: an initial synaptic matrix is generated as a result of
genetically determined chemical guidance, but then gradually refined by activity-dependent mechanisms,
partly driven by internal dynamics, partly by external sensory inputs. In some cases, as with the retina-
midbrain system, activity patterns depend on mechanisms that operate only in the developing brain, and
apparently exist solely for the purpose of guiding development.[54]
In humans and many other mammals, new neurons are created mainly before birth, and the infant
brain actually contains substantially more neurons than the adult brain.[55] There are, however, a few areas
where new neurons continue to be generated throughout life. The two areas for which this is well established
are the olfactory bulb, which is involved in the sense of smell, and the dentate gyrus of the hippocampus,
where there is evidence that the new neurons play a role in storing newly acquired memories. With these
exceptions, however, the set of neurons that is present in early childhood is the set that is present for life.
Glial cells are different, however; as with most types of cells in the body, these are generated throughout the
lifespan.
Although the pool of neurons is largely in place by birth, the axonal connections continue to develop
for a long time afterward. In humans, full myelination is not completed until adolescence.[56]
There has long been debate about whether the qualities of mind, personality, and intelligence can
mainly be attributed to heredity or to upbringing; the nature versus nurture debate.[57] This is not just a
philosophical question: it has great practical relevance to parents and educators. Although many details
remain to be settled, neuroscience clearly shows that both factors are essential. Genes determine the
general form of the brain, and genes determine how the brain reacts to experience. Experience, however, is
required to refine the matrix of synaptic connections. In some respects it is mainly a matter of presence or
absence of experience during critical periods of development.[58] In other respects, the quantity and quality
of experience may be more relevant: for example, there is substantial evidence that animals raised in
enriched environments have thicker cortices, indicating a higher density of synaptic connections, than
animals whose levels of stimulation are restricted.[59]
Functions
From an organismic perspective, the primary function of a brain is to control the actions of an animal.
[60] To do this, it extracts enough relevant information from sense organs to refine actions. Sensory signals
may stimulate an immediate response as when the olfactory system of a deer detects the odor of a wolf; they
may modulate an ongoing pattern of activity as in the effect of light-dark cycles on an organism's sleep-wake
behavior; or their information may be stored in case of future relevance. The brain manages its complex task
by orchestrating functional subsystems, which can be categorized in a number of ways: anatomically,
chemically, and functionally.
Functional subsystems
The most straightforward way to categorize the parts of the brain is anatomically, but there are also
several ways to subdivide it functionally. One of the most important of these is on the basis of the chemical
neurotransmitters used by neurons to communicate with each other. Another is in terms of the way a brain
area contributes to information processing: sensory areas bring information into the brain and reformat it;
motor signals send information out of the brain to control muscles and glands; arousal systems modulate the
activity of the brain according to time of day and other factors.
Neurotransmitter systems
Main article: Neurotransmitter systems
With few exceptions, each neuron in the brain consistently releases the same chemical
neurotransmitter, or combination of neurotransmitters, at all of the synaptic connections it makes with other
neurons; this rule is known as Dale's principle.[61] Thus, a neuron can be characterized by the
neurotransmitters it releases. The two neurotransmitters that appear most frequently are glutamate, which is
almost always excitatory, and gamma-aminobutyric acid (GABA), which is almost always inhibitory. Neurons
using these transmitters can be found in nearly every part of the brain, making up a large percentage of the
brain's pool of synapses.[62]
Nevertheless, the great majority of psychoactive drugs exert their effects by altering neurotransmitter
systems not directly involving glutamatergic or GABAergic transmission.[63] Drugs such as caffeine, nicotine,
heroin, cocaine, Prozac, Thorazine, etc., act on other neurotransmitters. Many of these other transmitters
come from neurons that are localized in particular parts of the brain. Serotonin, for example—the primary
target of antidepressant drugs and many dietary aids—comes exclusively from a small brainstem area called
the Raphe nuclei. Norepinephrine, which is involved in arousal, comes exclusively from a nearby small area
called the locus ceruleus. Histamine, as a neurotransmitter, comes from a tiny part of the hypothalamus
called the tuberomammilary nucleus (histamine also has non-CNS functions, but the neurotransmitter
function is what causes antihistamines to have sedative effects). Other neurotransmitters such as
acetylcholine and dopamine have multiple sources in the brain, but are not as ubiquitously distributed as
glutamate and GABA.
Sensory systems
Main article: Sensory system
One of the primary functions of a brain is to extract biologically relevant information from sensory
inputs.[64] Even in the human brain, sensory processes go well beyond the classical five senses of sight,
sound, taste, touch, and smell: our brains are provided with information about temperature, balance, limb
position, and the chemical composition of the bloodstream, among other things. All of these modalities are
detected by specialized sensors that project signals into the brain. In other animals, additional senses may
be present, such as the infrared heat-sensors in the pit organs of snakes; or the "standard" senses may be
used in nonstandard ways, as in the auditory "sonar" of bats. Here are a few principles that apply to most
sensory systems, using the auditory system for specific examples.
Each sensory system begins with specialized "sensory receptor" cells. These are neurons, but unlike
most neurons, they are not controlled by synaptic input from other neurons: instead they are activated by
membrane-bound receptors that are sensitive to some physical modality, such as light, temperature, or
physical stretching. The axons of sensory receptor cells travel into the spinal cord or brain. For the sense of
hearing, the receptors are located in the inner ear, on the cochlea, and are activated by vibration.[65]
For most senses, there is a "primary nucleus" or set of nuclei, located in the brainstem, that gathers
signals from the sensory receptor cells. For the sense of hearing, these are the cochlear nuclei.[65] In many
cases, there are secondary subcortical areas that extract special information of some sort. For the sense of
hearing, the superior olivary area and inferior colliculus are involved in comparing the signals from the two
ears to extract information about the direction of the sound source, among other functions.[65] Each sensory
system also has a special part of the thalamus dedicated to it, which serves as a relay to the cortex. For the
sense of hearing, this is the medial geniculate nucleus.[65]
For each sensory system, there is a "primary" cortical area that receives direct input from the
thalamic relay area. For the auditory system this is the primary auditory cortex, located in the upper part of
the temporal lobe.[65] There are also usually a set of "higher level" cortical sensory areas, which analyze the
sensory input in specific ways. For the auditory system, there are areas that analyze sound quality, rhythm,
and temporal patterns of change, among other features.[65] Finally, there are multimodal areas that combine
inputs from different sensory modalities, for example auditory and visual. At this point, the signals have
reached parts of the brain that are best described as integrative rather than specifically sensory.[65]
All of these rules have exceptions. For example, the sense of touch (which is actually a set of at least
half-a-dozen distinct mechanical senses), the sensory inputs terminate mainly in the spinal cord, on neurons
that then project to the brainstem.[66] For the sense of smell, there is no relay in the thalamus; instead the
signals go directly from the primary brain area—the olfactory bulb—to the cortex.[67]
Motor systems
Main article: Motor system
Motor systems are areas of the brain that are more or less directly involved in producing body
movements, that is, in activating muscles. With the exception of the muscles that control the eye, all of the
voluntary muscles[68] in the body are directly innervated by motor neurons in the spinal cord, which therefore
are the final common path for the movement-generating system.[69] Spinal motor neurons are controlled
both by neural circuits intrinsic to the spinal cord, and by inputs that descend from the brain. The intrinsic
spinal circuits implement many reflex responses, and also contain pattern generators for rhythmic
movements such as walking or swimming.[70] The descending connections from the brain allow for more
sophisticated control.
The brain contains a number of areas that project directly to the spinal cord.[71] At the lowest level
are motor areas in the medulla and pons. At a higher level are areas in the midbrain, such as the red
nucleus, which is responsible for coordinating movements of the arms and legs. At a higher level yet is the
primary motor cortex, a strip of tissue located at the posterior edge of the frontal lobe. The primary motor
cortex sends projections to the subcortical motor areas, but also sends a massive projection directly to the
spinal cord, via the so-called pyramidal tract. This direct corticospinal projection allows for precise voluntary
control of the fine details of movements. Other "secondary" motor-related brain areas do not project directly
to the spinal cord, but instead act on the cortical or subcortical primary motor areas. Among the most
important secondary areas are the premotor cortex, basal ganglia, and cerebellum:
• The premotor cortex (which is actually a large complex of areas) adjoins the primary motor
cortex, and projects to it. Whereas elements of the primary motor cortex map to specific body areas,
elements of the premotor cortex are often involved in coordinated movements of multiple body parts.
[72]
• The basal ganglia are a set of structures in the base of the forebrain that project to many
other motor-related areas.[73] Their function has been difficult to understand, but one of the most
popular theories currently is that they play a key role in action selection.[74] Most of the time they
restrain actions by sending constant inhibitory signals to action-generating systems, but in the right
circumstances, they release this inhibition and therefore allow their targets to take control of
behavior.
• The cerebellum is a very distinctive structure attached to the back of the brain.[33] It does not
control or originate behaviors, but instead generates corrective signals to make movements more
precise. People with cerebellar damage are not paralyzed in any way, but their body movements
become erratic and uncoordinated.
In addition to all of the above, the brain and spinal cord contain extensive circuitry to control the
autonomic nervous system, which works by secreting hormones and by modulating the "smooth" muscles of
the gut.[75] The autonomic nervous system affects heart rate, digestion, respiration rate, salivation,
perspiration, urination, and sexual arousal—but most of its functions are not under direct voluntary control.
Arousal systems
See also: Sleep
Perhaps the most obvious aspect of the behavior of any animal is the daily cycle between sleeping
and waking. Arousal and alertness are also modulated on a finer time scale, though, by an extensive network
of brain areas.[76]
A key component of the arousal system is the suprachiasmatic nucleus (SCN), a tiny part of the
hypothalamus located directly above the point at which the optic nerves from the two eyes cross.[77] The
SCN contains the body's central biological clock. Neurons there show activity levels that rise and fall with a
period of about 24 hours, circadian rhythms: these activity fluctuations are driven by rhythmic changes in
expression of a set of "clock genes". The SCN continues to keep time even if it is excised from the brain and
placed in a dish of warm nutrient solution, but it ordinarily receives input from the optic nerves, through the
retinohypothalamic tract (RHT), that allow daily light-dark cycles to calibrate the clock.
The SCN projects to a set of areas in the hypothalamus, brainstem, and midbrain that are involved in
implementing sleep-wake cycles. An important component of the system is the so-called reticular formation,
a group of neuron-clusters scattered diffusely through the core of the lower brain.[76] Reticular neurons send
signals to the thalamus, which in turn sends activity-level-controlling signals to every part of the cortex.
Damage to the reticular formation can produce a permanent state of coma.
Sleep involves great changes in brain activity.[78] Until the 1950s it was generally believed that the
brain essentially shuts off during sleep,[79] but this is now known to be far from true: activity continues, but
patterns become very different. In fact, there are two types of sleep, REM sleep (with dreaming) and NREM
(non-REM, usually without dreaming) sleep, which repeat in slightly varying patterns throughout a sleep
episode. Three broad types of distinct brain activity patterns can be measured: REM, light NREM and deep
NREM. During deep NREM sleep, also called slow wave sleep, activity in the cortex takes the form of large
synchronized waves, where in the waking state it is noisy and desynchronized. Levels of the
neurotransmitters norepinephrine and serotonin drop during slow wave sleep, and fall almost to zero during
REM sleep; levels of acetylcholine show the reverse pattern.
Brain energy consumption
PET Image of the human brain showing energy consumption

Although the human brain represents only 2% of the body weight, it receives 15% of the cardiac
output, 20% of total body oxygen consumption, and 25% of total body glucose utilization.[80] The need to
limit body weight in order, for example, to fly, has led to selection for a reduction of brain size in some
species, such as bats.[81] The brain mostly utilizes glucose for energy, and deprivation of glucose, as can
happen in hypoglycemia, can result in loss of consciousness. The energy consumption of the brain does not
vary greatly over time, but active regions of the cortex consume somewhat more energy than inactive
regions: this fact forms the basis for the functional brain imaging methods PET and fMRI.[82] These are
nuclear medicine imaging techniques which produce a three-dimensional image of metabolic activity.
Brain and mind

See also: Philosophies of mind
Understanding the relationship between the brain and the mind is a challenging problem both
philosophically and scientifically.[83] The most straightforward scientific evidence that there is a strong
relationship between the physical brain matter and the mind is the impact physical alterations to the brain
have on the mind, such as with traumatic brain injury and psychoactive drug use.[84]
The mind-body problem is one of the central issues in the history of philosophy,[85] which asks us to
consider if the brain and the mind are identical, partially distinct, or related in some unknown way. There are
three major schools of thought concerning the answer: dualism, materialism, and idealism. Dualism holds
that the mind exists independently of the brain;[86] materialism holds that mental phenomena are identical to
neuronal phenomena;[87] and idealism holds that only mental phenomena exist.[87]
In addition to the philosophical questions, the relationship between mind and brain involves a high
number of scientific questions, including understanding the relationship between mental activity and brain
activity, the exact mechanisms by which drugs influence cognition, and the neural correlates of
consciousness.
Through most of history many philosophers found it inconceivable that cognition could be
implemented by a physical substance such as brain tissue (that is neurons and synapses).[88] Philosophers
such as Patricia Churchland posit that the drug-mind interaction is indicative of an intimate connection
between the brain and the mind, not that the two are the same entity.[89] Descartes, who thought extensively
about mind-brain relationships, found it possible to explain reflexes and other simple behaviors in
mechanistic terms, although he did not believe that complex thought, and language in particular, could be
explained by reference to the physical brain alone.[90]
Research
The field of neuroscience encompasses all approaches that seek to understand the brain and the
rest of the nervous system.[91] Psychology seeks to understand mind and behavior, and neurology is the
medical discipline that diagnoses and treats pathologies of the nervous system. The brain is also the most
important organ studied in psychiatry, the branch of medicine that works to study, prevent, and treat mental
disorders.[92] Cognitive science seeks to unify neuroscience and psychology with other fields that concern
themselves with the brain, such as computer science (artificial intelligence and similar fields) and philosophy.
The oldest method of studying the brain is anatomical, and until the middle of the 20th century, much
of the progress in neuroscience came from the development of better stains and better microscopes—the
neuroanatomist Floyd Bloom famously quipped that "the gain in brain is mainly in the stain."[93]
Neuroanatomists study the large-scale structure of the brain as well as the microscopic structure of neurons
and their components, especially synapses. Among other tools, they employ a plethora of stains that reveal
neural structure, chemistry, and connectivity. In recent years, the development of immunostaining techniques
has allowed staining of neurons that express specific sets of genes. Also, functional neuroanatomy uses
medical imaging techniques to correlate variations in human brain structure with differences in cognition or
behavior.
Neurophysiologists study the chemical, pharmacological, and electrical properties of the brain: their
primary tools are drugs and recording devices. Many thousands of experimentally developed drugs affect the
nervous system, some in highly specific ways. Recordings of brain activity can be made using electrodes,
either glued to the skull as in EEG studies, or implanted inside the brains of animals for extracellular
recordings, which can detect action potentials generated by individual neurons.[94] Because the brain does
not contain pain receptors, it is possible using these techniques to record from animals that are awake and
behaving without causing distress. The same techniques have occasionally been used to study brain activity
in human patients suffering from intractable epilepsy, in cases where there was a medical necessity to
implant electrodes in order to localize the brain area responsible for seizures.[95] It is also possible to study
brain activity noninvasively in humans using functional imaging techniques such as MRI—this field has
expanded enormously over the past two decades.
A different approach to brain function is to examine the consequences of damage to specific brain
areas. Even though it is protected by the skull and meninges, surrounded by cerebrospinal fluid, and isolated
from the bloodstream by the blood-brain barrier, the delicate nature of the brain makes it vulnerable to
numerous diseases and several types of damage. In humans, the effects of strokes and other types of brain
damage have been a key source of information about brain function.[96] Because there is no ability to
experimentally control the nature of the damage, however, this information is often difficult to interpret. In
animal studies, most commonly involving rats, it is possible to use electrodes or locally injected chemicals to
produce precise patterns of damage and then examine the consequences for behavior.
Computational neuroscience encompasses two approaches: first, the use of computers to study the
brain; second, the study of how brains perform computation.[97] On one hand, it is possible to write a
computer program to simulate the operation of a group of neurons by making use of systems of equations
that describe their electrochemical activity; such simulations are known as biologically realistic neural
networks. On the other hand, it is possible to study algorithms for neural computation by simulating, or
mathematically analyzing, the operations of simplified "units" that have some of the properties of neurons but
abstract out much of their biological complexity. The computational functions of the brain are studied both by
neuroscientists and computer scientists.
Recent years have seen the first applications of genetic engineering techniques to the study of the
brain.[98] The most common subjects are mice, because the technical tools are more advanced for this
species than for any other. It is now possible with relative ease to "knock out" or mutate a wide variety of
genes, and then examine the effects on brain function. More sophisticated approaches are also beginning to
be used: for example, using the Cre-Lox recombination method it is possible to activate or inactivate genes in
specific parts of the brain, at specific times.
History
See also: History of neuroscience
Early views were divided as to whether the seat of the soul lies in the brain or heart. On one hand, it
was impossible to miss the fact that awareness feels like it is localized in the head, and that blows to the
head can cause unconsciousness much more easily than blows to the chest, and that shaking the head
causes dizziness. On the other hand, the brain to a superficial examination seems inert, whereas the heart is
constantly beating. Cessation of the heartbeat means death; strong emotions produce changes in the
heartbeat; and emotional distress often produces a sensation of pain in the region of the heart ("heartache").
Aristotle favored the heart, and thought that the function of the brain was merely to cool the blood.
Democritus, the inventor of the atomic theory of matter, favored a three-part soul, with intellect in the head,
emotion in the heart, and lust in the vicinity of the liver.[99] Hippocrates, the "father of medicine", was entirely
in favor of the brain. In On the Sacred Disease, his account of epilepsy, he wrote:
Men ought to know that from nothing else but the brain come joys, delights, laughter and sports,
and sorrows, griefs, despondency, and lamentations. ... And by the same organ we become
mad and delirious, and fears and terrors assail us, some by night, and some by day, and dreams
and untimely wanderings, and cares that are not suitable, and ignorance of present
circumstances, desuetude, and unskilfulness. All these things we endure from the brain, when it
is not healthy...
—Hippocrates, On the Sacred Disease[100]
The famous Roman physician Galen also advocated the importance of the brain, and theorized in
some depth about how it might work. Even after physicians and philosophers had accepted the primacy of
the brain, though, the idea of the heart as seat of intelligence continued to survive in popular idioms, such as
"learning something by heart".[101] Galen did a masterful job of tracing out the anatomical relationships
among brain, nerves, and muscles, demonstrating that all muscles in the body are connected to the brain via
a branching network of nerves. He postulated that nerves activate muscles mechanically, by carrying a
mysterious substance he called pneumata psychikon, usually translated as "animal spirits". His ideas were
widely known during the Middle Ages, but not much further progress came until the Renaissance, when
detailed anatomical study resumed, combined with the theoretical speculations of Descartes and his
followers. Descartes, like Galen, thought of the nervous system in hydraulic terms. He believed that the
highest cognitive functions—language in particular—are carried out by a non-physical res cogitans, but that the
majority of behaviors of humans and animals could be explained mechanically. The first real progress toward
a modern understanding of nervous function, though, came from the investigations of Luigi Galvani, who
discovered that a shock of static electricity applied to an exposed nerve of a dead frog could cause its leg to
contract.
Drawing by Santiago Ramon y Cajal of two types of Golgi-stained neurons from the cerebellum of a
pigeon
Each major advance in understanding has followed more or less directly from the development of a
new method of investigation. Until the early years of the 20th century, the most important advances were
derived from new stains.[102] Particularly critical was the invention of the Golgi stain, which (when correctly
used) stains only a small, and apparently random, fraction of neurons, but stains them in their entirety,
including cell body, dendrites, and axon. Without such a stain, brain tissue under a microscope appears as
an impenetrable tangle of protoplasmic fibers, in which it is impossible to determine any structure. In the
hands of Camillo Golgi, and especially of the Spanish neuroanatomist Santiago Ramon y Cajal, the new stain
revealed hundreds of distinct types of neurons, each with its own unique dendritic structure and pattern of
connectivity.
In the 20th century, progress in electronics enabled investigation of the electrical properties of nerve
cells, culminating in the work by Alan Hodgkin, Andrew Huxley, and others on the biophysics of the action
potential, and the work of Bernard Katz and others on the electrochemistry of the synapse.[103] The earliest
studies used special preparations, such as the "fast escape response" system of the squid, which involves a
giant axon as thick as a pencil lead, and giant synapses connecting to this axon. Steady improvements in
electrodes and electronics allowed ever finer levels of resolution. These studies complemented the
anatomical picture with a conception of the brain as a dynamic entity. Reflecting the new understanding, in
1942 Charles Sherrington visualized the workings of the brain in action in somewhat breathless terms:
The great topmost sheet of the mass, that where hardly a light had twinkled or moved, becomes
now a sparkling field of rhythmic flashing points with trains of traveling sparks hurrying hither
and thither. ... It is as if the Milky Way entered upon some cosmic dance. Swiftly the head mass
becomes an enchanted loom where millions of flashing shuttles weave a dissolving pattern,
always a meaningful pattern though never an abiding one; a shifting harmony of subpatterns.
—Sherrington, 1942, Man on his Nature[104]
The 1990s were known in the US as the "Decade of the Brain" to commemorate advances made in
brain research, and to promote funding for such research.[105][106]
See also
Mind and Brain portal
Neuroscience portal
• Brain–computer interface
• Cephalization
• List of neuroscience databases
• Slice preparation
Notes
1. ^ a b Shepherd, Neurobiology
2. ^ Pelvig et al., 2008
3. ^ AlonsoNanclares et al., 2008
4. ^ Gehring, 2005
5. ^ Nickel, 2002
6. ^ Grillner & Wallén, 2002
7. ^ Sejnowski, 23 Problems in Systems Neuroscience
8. ^ Shepherd, Neurobiology, p 3
9. ^ a b Balavoine & Adoutte, 2003
10.^ Schmidt-Rhaesa, Evolution of Organ Systems, p 110
11.^ a b Butler, 2000
12.^ Kandel, In Search of Memory
13.^ Flybrain web site
14.^ Konopka & Benzer, 1971
15.^ Shin et al., 1985
16.^ WormBook web site
17.^ Hobert, WormBook
18.^ White et al., 1986
19.^ Principles of neural science, Ch. 17
20.^ Carpenter's Human Neuroanatomy, Ch. 1
21.^ Shu et al., 2003
22.^ Striedter, Principles of Brain Evolution
23.^ Principles of Neural Science, p 1019
24.^ Armstrong, 1983
25.^ Jerison, Evolution of the Brain and Intelligence
26.^ a b Finlay et al., 2001
27.^ Principles of Neural Science, Ch. 17
28.^ Northcutt, 2008
29.^ Reiner et al., 2005
30.^ Principles of Neural Science, Ch. 44, 45
31.^ Swaab et al., The Human Hypothalamus
32.^ Jones, The Thalamus
33.^ a b Principles of Neural Science, Ch. 42
34.^ Saitoh et al., 2007
35.^ Puelles, 2001
36.^ Salas et al., 2003
37.^ Grillner et al., 2005
38.^ Northcutt, 1981
39.^ Barton & Harvey, 2000
40.^ Aboitiz et al., 2003
41.^ Aboitiz, 2004
42.^ Calvin, How Brains Think
43.^ "Over half of the neocortex in non-human primates is occupied by visual areas. At least 25
visual areas beyond the primary visual cortex (V1) have been identified with a combination of
microelectrode mapping, tracer injections, histological stains, and functional studies (1)." see:
Sereno, MI; Dale, AM; Reppas, AM; Kwong, KK; Belliveau, JW; Brady, TJ; Rosen, BR; Tootell, RBH
(1995). "Borders of multiple visual areas in human revealed by functional magnetic resonance
imaging". Science (AAAS) 268 (5212): 889–893. doi:10.1126/science.7754376. PMID 7754376.
http://www.cogsci.ucsd.edu/~sereno/papers/HumanRetin95.pdf. and refs contained therein
44.^ Principles of Neural Science p 20
45.^ Azevedo et al., 2009
46.^ Principles of Neural Science, p 21
49.^ Marner et al., 2003
50.^ a b c Principles of Neural Development, Ch. 1
51.^ Principles of Neural Development, Ch. 4
52.^ Principles of Neural Development, Chs. 5, 7
54.^ a b Wong, 1999
56.^ Paus et al., 2001
57.^ Ridley, Nature vs Nurture
58.^ Wiesel, 1982
59.^ van Praag et al., 2000
60.^ Carew, Behavioral Neurobiology, Ch. 1
62.^ McGeer and McGeer, Basic Neurochemistry, pp. 311–332
63.^ Cooper et al., Biochemical Basis of Neuropharmacology
65.^ a b c d e f g Principles of Neural Science, Chs. 21, 30
68.^ See muscle
70.^ Principles of Neural Science, Chs. 36, 37
74.^ Gurney et al., 2004
76.^ a b Principles of Neural Science, Ch. 45
77.^ Antle & Silver, 2005
79.^ Kleitman, 1938
80.^ Clark & Sokoloff, 1999
81.^ Safi et al., 2005
82.^ Raichle & Gusnard, 2002
83.^ Churchland, Neurophilosophy
84.^ Boake and Diller, 2005
85.^ Neurophilosophy, Ch. 7
86.^ Hart, 1996
87.^ a b Lacey, 1996
90.^ Descartes, Description of the human body
92.^ Storrow, Outline of Clinical Psychiatry
93.^ Neuroscience: Exploring the Brain, Ch. 2
94.^ Dowling, Neurons and Networks, pages 15–24
95.^ Wyllie et al., Treatment of Epilepsy, Ch. 77
96.^ Fundamentals of Human Neuropsychology, Ch. 1
97.^ Abbott & Dayan, Theoretical Neuroscience
98.^ Tonegawa et al., 2003
99.^ Finger, Origins of Neuroscience, p 14
100.^ Hippocrates, On the Sacred Disease
101.^ Encyclopedia of Word and Phrase Origins
102.^ Bloom, 1972, p 211
103.^ Piccolino, 2002
104.^ Sherrington, Man on his Nature
105.^ Bush, George H.W. (July 1990). "Project on the Decade of the Brain".
http://www.loc.gov/loc/brain/proclaim.html. Retrieved 2009-09-30.
106.^ Jones, Edward G.; Mendell, Lorne M. (April 30, 1999). "Assessing the Decade of the
Brain". Science (American Association for the Advancement of Science) 284 (5415): 739.
doi:10.1126/science.284.5415.739. http://www.sciencemag.org/cgi/content/summary/284/5415/739.
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Further reading
• Bear, Mark F.; Barry W. Connors, Michael A. Paradiso (2006). Neuroscience. Philadelphia,
Pennsylvania: Lippincott Williams & Wilkins. ISBN 9780781760034. OCLC 62509134.
• Blackmore, Susan M. (2006). Conversations on Consciousness. Oxford; New York: Oxford
University Press. ISBN 9780195179583. OCLC 62555307.
• Buzsaki, Gyorgy (2006). Rhythms of the Brain. Oxford; New York: Oxford University Press.
ISBN 9780195301069. OCLC 63279497.
• Calvin, William H. (2001). The River That Flows Uphill: A Journey from the Big Bang to the
Big Brain. Lincoln, Nebraska: Iuniverse.com. ISBN 9780595167005. OCLC 48962546.
• Della Sala, Sergio (1999). Mind myths: Exploring popular assumptions about the mind and
brain. Chichester England; New York: J. Wiley & Sons. ISBN 0471983039. OCLC 39700332.
• Restak, Richard (2001). The Secret Life of the Brain. Washington, DC: Joseph Henry Press.
ISBN 9780309074353. OCLC 47863192.
• Shepherd, Gordon M. (2004). The Synaptic Organization of the Brain (Fifth ed.). Oxford; New
York: Oxford University Press. ISBN 9780195159561. OCLC 51769076.
Written for children 8 and older:
• Simon, Seymour (2000). The Brain. New York: Morrow Junior. ISBN 9780688170608.
OCLC 35686089.
External links
Wikimedia Commons has media related to: Brain
• The Society for Neuroscience

• IBRO (International Brain Research Organization)
• The HOPES Brain Tutorial at hopes.stanford.edu
• Comparative Mammalian Brain Collection
• Brain Research News from ScienceDaily
• BrainInfo for Neuroanatomy
• Neuroscience for kids
• BrainMaps.org, interactive high-resolution digital brain atlas based on scanned images of
serial sections of both primate and non-primate brains
• The Brain from Top to Bottom
• The Department of Neuroscience at Wikiversity
• The Secret Life of the Brain : History of the Brain from PBS
• University of Washington 3D animations of brain regions - click through from "Click for
copyright"
• 7 circuits of the Brain
• Cell Centered Database
[hide]v · d · eHuman systems and organs
Bone (Carpus · Collar bone (clavicle) · Thigh
bone (femur) · Fibula · Humerus · Mandible · Metacarpus ·
Skeletal system Metatarsus · Ossicles · Patella · Phalanges · Radius ·
Skull (cranium) · Tarsus · Tibia · Ulna · Rib · Vertebra ·
TA 2-4:
Pelvis · Sternum) · Cartilage
MS
Joints Fibrous joint · Cartilaginous joint · Synovial joint
Muscular system Muscle · Tendon · Diaphragm
URT (Nose,
mostly
Respiratory system Nasopharynx, Larynx) · LRT
Thoracic
(Trachea, Bronchus, Lung)
TA 5-11: Mouth (Salivary gland,

splanchnic/ Tongue) · upper GI (Oropharynx,
viscus Digestive Laryngopharynx, Esophagus,
mostly system+ Stomach) · lower GI (Small
Abdominopelvic adnexa intestine, Appendix, Colon,
Rectum, Anus) · accessory (Liver,
Biliary tract, Pancreas)
GU: Kidney · Ureter · Bladder ·
Urinary system Urethra
Female (Uterus, Vulva,

GU: Ovary, Placenta) · Male (Scrotum,
Reproductive system Penis, Prostate, Testicle, Seminal
vesicle)
Pituitary · Pineal · Thyroid · Parathyroid · Adrenal ·

Endocrine system
Islets of Langerhans
peripheral
Cardiovascular system (Artery, Vein, Lymph
vessel) · Heart
Circulatory system
primary (Bone
TA 12-16 marrow, Thymus) ·
Lymphatic system
secondary (Spleen,
Lymph node)
(Brain, Spinal cord, Nerve) · Sensory system

Nervous system
(Ear, Eye)
Skin · Subcutaneous tissue · Breast (Mammary
Integumentary system
gland)
Blood Myeloid Myeloid mmune system

(Non-TA)
Lymphoid Lymphoid mmune system
Retrieved from "http://en.wikipedia.org/wiki/Brain"
Categories: Brain | Central nervous system | Neuroanatomy | Organs

W000
Systematics

"Systematic" redirects here. For the band, see Systematic (band).
For other uses, see Systematics (disambiguation).
A comparison of phylogenetic and phenetic concepts
Biological systematics is the study of the diversification of life on the planet Earth, both past and
present, and the relationships among living things through time. Relationships are visualized as evolutionary
trees (synonyms: cladograms, phylogenetic trees, phylogenies). Phylogenies have two components,
branching order (showing group relationships) and branch length (showing amount of evolution).
Phylogenetic trees of species and higher taxa are used to study the evolution of traits (e.g., anatomical or
molecular characteristics) and the distribution of organisms (biogeography). Systematics, in other words, is
used to understand the evolutionary history of life on Earth.
"Systematic biology" and "taxonomy" (terms that are often confused and used interchangeably) were
defined in relationship to one another as follows:[1]
Systematic biology (hereafter called simply systematics) is the field that (a) provides scientific
names for organisms, (b) describes them, (c) preserves collections of them, (d) provides
classifications for the organisms, keys for their identification, and data on their distributions, (e)
investigates their evolutionary histories, and (f) considers their environmental adaptations. This
is a field with a long history that in recent years has experienced a notable renaissance,
principally with respect to theoretical content. Part of the theoretical material has to do with
evolutionary areas (topics e and f above), the rest relates especially to the problem of
classification. Taxonomy is that part of Systematics concerned with topics (a) to (d) above.
The term "systematics" is sometimes used synonymously with "taxonomy" and may be confused with
"scientific classification". However, Taxonomy is more specifically the identification, description, and naming
(i.e. nomenclature) of organisms[citation needed], while "classification" is focused on placing organisms
within hierarchical groups that show their relationships to other organisms. All of these biological disciplines
can be involved with extinct and extant organisms. However, systematics alone deals specifically with
relationships through time, and can be synonymous with phylogenetics, broadly dealing with the inferred
hierarchy of organisms.
Systematics uses taxonomy as a primary tool in understanding organisms, as nothing about an
organism's relationships with other living things can be understood without it first being properly studied and
described in sufficient detail to identify and classify it correctly. Scientific classifications are aids in recording
and reporting information to other scientists and to laymen. The systematist, a scientist who specializes in
systematics, must, therefore, be able to use existing classification systems, or at least know them well
enough to skillfully justify not using them.
Phenetic systematics was an attempt to determine the relationships of organisms through a measure
of similarity, considering plesiomorphies (ancestral traits) and apomorphies (derived traits) to be equally
informative. From the 20th century onwards, it was superseded by cladistics, which considers plesiomorphies
to be uninformative for an attempt to resolve the phylogeny of Earth's various organisms through time.
Today's systematists generally make extensive use of molecular biology and computer programs to study
organisms.
Systematics is fundamental to biology because it is the foundation for all studies of organisms, by
showing how any organism relates to other living things (ancestor-descendant relationships).
Systematics is also of major importance in understanding conservation issues because it attempts to
explain the Earth's biodiversity and could be used to assist in allocating limited means to preserve and
protect endangered species, by looking at, for example, the genetic diversity among various taxa of plants or
animals and deciding how much of that to preserve.
[edit] See also

• Cladistics - a popular methodology in systematics
• Phenetics - an obsolete methodology, predecessor to cladistics
• Phylogeny - what is analyzed in systematics and taxonomy
• 16S ribosomal RNA - the most intensively studied nucleic acid in phylogenetics
• Phylogenetic comparative methods - use of evolutionary trees to study biodiversity and
comparative biology
• Scientific classification - the result of research in systematics and taxonomy
• Taxonomy - a branch of the biological sciences related to systematics
[edit] References
[edit] Notes
1. ^ Michener, Charles D., John O. Corliss, Richard S. Cowan, Peter H. Raven, Curtis W.
Sabrosky, Donald S. Squires, and G. W. Wharton (1970). Systematics In Support of Biological
Research. Division of Biology and Agriculture, National Research Council. Washington, D.C. 25 pp.

• Schuh, Randall T. and Andrew V. Z. Brower. 2009. Biological Systematics: Principles and
Applications, 2nd edn. ISBN 978-0-8014-4799-0
• Simpson, Michael G. 2005. Plant Systematics. ISBN 978-0-12-644460-5

• Society of Australian Systematic Biologists
• Society of Systematic Biologists
• Willi Hennig Society
Processes of
Population genetic
Evolutionary
developmental
concepts

Modes of

Retrieved from "http://en.wikipedia.org/wiki/Systematics"
Categories: Evolutionary biology | Scientific classification

W000
Multiregional origin of modern humans


A graph detailing the evolution to modern humans using the Multiregional theory of human evolution.
The horizontal lines represent 'multiregional evolution' gene flow between regional lineages. In Weidenreich's
original graphic (which is more accurate than this one), there were also diagonal lines between the
populations, e.g. between African H. erectus and Archaic Asians and between Asian H. erectus and Archaic
Africans. This created a "trellis" (as Wolpoff called it) or a "network" that emphasized gene flow between
geographic regions and within time. It is important to remember that the populations on the chart are not
discrete - i.e., they do not represent different species, but are samples within a long lineage experiencing
extensive gene flow.
The multiregional hypothesis is a scientific model that provides an explanation for the pattern of
human evolution. The hypothesis holds that humans first arose near the beginning of the Pleistocene two
million years ago and subsequent human evolution has been within a single, continuous human species.
This species encompasses archaic human forms such as Homo erectus and Neanderthals as well as
modern forms, and evolved worldwide to the diverse populations of modern Homo sapiens sapiens. The
theory contends that humans evolve through a combination of adaptation within various regions of the world
and gene flow between those regions. Proponents of multiregional origin point to fossil and genomic
evidence as support for their hypothesis.
The primary alternative hypothesis is recent African origin of modern humans, which contends that
modern humans arose in Africa around 100-200,000 years ago, moving out of Africa around 50-60,000 years
ago to replace archaic human forms without interbreeding.
Contents
[hide]
• 1 Overview
• 2 Fossil evidence
• 2.1 Southeast Asia
• 2.2 China
• 2.3 Europe
• 3 Genetic evidence
• 3.1 Mitochondrial DNA
• 3.2 Nuclear DNA
• 3.3 Ancient DNA
• 4 Alternate hypotheses
• 4.1 Polygenism
• 4.2 Recent African replacement
• 5 See also
• 6 References
[edit] Overview
The idea of multiregional evolution goes back to Franz Weidenreich in the early 20th century, who
saw similarities between the archaic human Peking man fossils and modern humans in China.[1][2] The term
"multiregional hypothesis" was first applied to the theory in the early 1980s by Milford H. Wolpoff and
colleagues, who used the theory to explain these and other similarities between archaic humans and modern
humans in various regions, in what they regional continuity.[3][4]
Wolpoff rejected the proposal by Coon of parallel evolution,[4] and proposed that the mechanism of
clinal variation allowed for the necessary balance between both local selection and overall evolution as a
global species, with Homo erectus, Neanderthals, Homo sapiens and other human forms as subspecies. This
species arose in Africa two million years ago as H. erectus and then spread out over the world, developing
adaptations to regional conditions. Some populations became isolated for periods of time, developing in
different directions, but through continuous interbreeding, replacement, genetic drift and selection,
adaptations that were an advantage anywhere on earth would spread, keeping the development of the
species in the same overall direction while maintaining adaptations to regional factors. By these
mechanisms, surviving local varieties of the species evolved into modern humans, retaining some regional
adaptations but with many features common to all regions.[4]
[edit] Fossil evidence
Replica of Sangiran 17 Homo erectus skull from Indonesia showing obtuse face to vault angle
determined by fitting of bones at brow.
Cast of anatomically modern human Kow Swamp 1 skull from Australia with a face to vault angle
matching that of Sangiran 17.
Proponents of the multiregional hypothesis see regional continuity of certain morphological traits
from archaic humans to modern humans, demonstrating regional genetic continuity, even as changes in
other traits occur in parallel over time across all regions, demonstrating lateral genetic exchange.[5] For
example, in 2001 Wolpoff and colleagues published an analysis of character traits of the skulls of early
modern human fossils in Australia and central Europe. They concluded that the diversity of these recent
humans could not "result exclusively from a single late Pleistocene dispersal", and implied dual ancestry from
Javan Homo erectus for Australia and from Neanderthals for Central Europe.[6][7]
[edit] Southeast Asia
Alan Thorne held that there was regional continuity in the human fossils in southeast Asia. Wolpoff,
initially skeptical, became convinced when reconstructing the Sangiran 17 Homo erectus skull from
Indonesia, when he was surprised that the skull's face to vault angle matched that of the Australian modern
human Kow Swamp 1 skull. Wolpoff had expected the skull to match that of the Homo erectus specimens
from China like the Dali skull, but instead, the face to vault angle seemed to be retained regionally over time,
even while the fossils in the two regions showed parallel increases in brain case size and parallel reductions
in masticatory structures over the intervening approximately 750,000 years.[5]
[edit] China
Replica of Homo erectus ("Peking man") skull from China.

Franz Weidenreich, who oversaw the excavations of numerous "Peking man" Homo erectus fossils
at Zhoukoudian in the early 20th century, believed the fossil record demonstrated certain unique features
linking prehistoric and modern human populations in China. Many subsequent Chinese paleoanthropologists,
such as Wu Xinzhi, were also disposed to favor the multiregional hypothesis for the same reason.[2]
More recent finds provide additional support for regional human continuity in China. The Tianyuan 1
specimen unearthed in 2003 in Tianyuan Cave, Zhoukoudian, and Carbon 14 dated to 42-39 kya exhibits a
series of typical modern human features such as a distinct chin. However, the skeleton also has archaic traits
such as low anterior to posterior dental proportions indicating relatively large molars and certain leg bone
proportions typical of archaic forms such as Neanderthals. Shang et. al. conclude that this combination of
modern and archaic traits "implies that a simple spread of modern humans from Africa is unlikely."[8] A jaw
bone found in 2008 and dated to 110,000 kya may also exhibit a mixture of archaic and modern human traits.
[9]
[edit] Europe
Comparison of modern human and neanderthal skull.
Proponents of the multiregional hypothesis argue for regional continuity in Europe on the basis of
skeletal anatomy, morphology and genetics of speech, and the archaeology of the middle to upper paleolithic
transition, which they believe to be inconsistent with the possibility of complete replacement of the
Neanderthals in Europe without interbreeding.[10]
Some detractors of the theory have argued, in contrast, that the morphological differences between
Neanderthals and early and modern humans indicate that they are different species, based on skull
differences more distinct than between any subspecies pairs examined except for the two subspecies of
gorilla, implying limited or no interbreeding.[11][12]
Many of the multiregional claims regarding skeletal morphology in Europe center on forms with both
archaic Neanderthal traits and modern traits, to provide evidence of interbreeding rather than replacement.
Examples include the Lapedo child found in Portugal[13] and the Oase 1 mandible from Peştera cu Oase,
Romania,[14] though the Lapedo child example is disputed by some.[15] In a 2007 paper examining
numerous samples from European early modern humans, later European humans from the Gravettian
period, and the earlier Neanderthal and east African populations from whom the first two populations could
have descended, Erik Trikaus identified numerous features in the later European samples which were absent
from the African sample, but present in the Neanderthal sample. These features included various aspects of
skull and mandible shape, tooth shape and size, and shapes and proportions of other bones. Trinkaus
concluded that early modern Europeans had predominant African ancestry with a substantial degree of
admixture from the Neanderthals then indigenous to Europe.[16][17]
[edit] Genetic evidence
Genetic evidence from the late 1980s on the mitochondrial genome indicated that all living humans
had as an ancestor a single female living in African about 200,000 years ago. This led to a hypothesis that
around that time period, a small founder population of humans left Africa and eventually replaced all archaic
humans then living outside of Africa without interbreeding, contrary to the multiregional hypothesis. However,
as data on the far larger nuclear DNA genome started to became available, evidence mounted that genetic
contributions from archaic human populations from around the world, and not just from Africa, also persist in
modern humans. Recent analyses of DNA taken directly from Neanderthal and denisovan specimens
indicates that those nonafrican populations also contributed to the genome of living humans, as predicted by
the multiregional hypothesis.
[edit] Mitochondrial DNA
Human mitochondrial DNA tree. "Mitochondrial Eve" is near the top of the diagram, next to the
jagged arrow pointing to "Outgroup", and her distance from any nonafrican groups indicates that living human
mitochondrial lineages coalesce in Africa.
A 1987 analysis of mitochondrial DNA from 147 people from around the world indicated that their
mitochondrial lineages all coalesced in a common ancestor in Africa about 200,000 years ago. The analysis
suggested that this reflected the worldwide expansion of modern humans as a new species, replacing rather
than mixing with local archaic humans.[18] Later analysis of mitochondrial DNA from neanderthals and from
the denisova hominin indicated that those mitochondrial strains had diverged from the living human
mitochondrial line long before 200,000 years ago, consistent with lack of interbreeding between early modern
and archaic humans.[19][20]
The original mitochondrial DNA results and the resulting recent African replacement theory posed a
serious challenge to the multiregional hypothesis.[21] Mitochondrial DNA alone, however, could not entirely
rule out interbreeding between early modern and archaic humans, since archaic human mitochondrial strains
from such interbreeding could have been lost due to genetic drift or a selective sweep.[22][23] Indeed, later
analysis of autosomal nuclear DNA from both living and ancient humans was to show results very different
from those from mitochondrial DNA.
[edit] Nuclear DNA

Initial analysis of Y chromosome DNA, which like mitochondrial DNA is inherited from only one
parent, was consistent with a recent African replacement model. However, the mitochondrial and Y
chromosome data could not be explained by the same modern human expansion out of Africa; the Y
chromosome expansion would have involved genetic mixing that retained regionally local mitochondrial lines.
In addition, the Y chromosome data indicated a later expansion back into Africa from Asia, demonstrating
that gene flow between regions was not unidirectional.[24]
An early analysis of 15 noncoding sites on the X chromosome found additional inconsistencies with
the recent African replacement hypothesis. The analysis found a multimodal distribution of coalescence
times to the most recent common ancestor for those sites, contrary to the predictions for recent African
replacement; in particular, there were more coalescence times near 2 million years ago (Mya) than expected,
suggesting an ancient population split around the time humans first emerged from Africa as Homo erectus,
rather than more recently as suggested by the mitochondrial data. While most of these X chromosome sites
showed greater diversity in Africa, consistent with African origins, a few of the sites showed greater diversity
in Asia rather than Africa. For four of the 15 gene sites that did show greater diversity in Africa, the sites'
varying diversity by region could not be explained by simple expansion from Africa, as would be required by
the recent African replacement hypothesis.[25]
Later analyses of X chromosome and autosomal DNA continued to find sites with deep coalescence
times inconsistent with a single origin of modern humans,[26][27][28][29][30][31] diversity patterns
inconsistent with a recent expansion from Africa,[32] or both.[33][34] For example, analyses of a region of
RRM2P4 (ribonucleotide reductase M2 subunit pseudogene 4) showed a coalescence time of about 2 Mya,
with a clear root in Asia,[35][36] while the MAPT locus at 17q21.31 is split into two deep genetic lineages,
one of which is common in and largely confined to the present European population, suggesting inheritance
from Neanderthals.[37][38][39][40] In the case of the Microcephalin D allele, evidence for rapid recent
expansion indicated introgression from an archaic population.[41][42][43][44]
In a 2005 review and analysis of the genetic lineages of 25 chromosomal regions, Alan Templeton
found evidence of more than 34 occurrences of gene flow between Africa and Eurasia. Of these occurrences,
19 were associated with continuous restricted gene exchange through at least 1.46 million years ago; only 5
were associated with a recent expansion from Africa to Eurasia. Three were associated with the original
expansion of Homo erectus out of Africa around 2 million years ago, 7 with an intermediate expansion out of
Africa at a date consistent with the expansion of Acheulean tool technology, and a few others with other gene
flows such as an expansion out of Eurasia and back into Africa subsequent to the most recent expansion out
of Africa. Templeton rejected a hypothesis of recent African replacement with greater than 99% certainty (p <
10−17).[45]
[edit] Ancient DNA

By 2006, extraction of DNA directly from some archaic human samples was becoming possible. The
earliest analyses were of neanderthal DNA, and indicated that the neanderthal contribution to modern human
genetic diversity was no more than 20%, with a most likely value of 0%.[46] By 2010, however, detailed DNA
sequencing of the neanderthal specimens from Europe indicated that the contribution was nonzero, with
neanderthals sharing 1-4% more genetic variants with living nonafricans than with living humans in
subsaharan Africa, supporting regional continuity outside of Africa.[47][48] In late 2010, a recently discovered
nonneanderthal archaic human, the denisova hominin from southern siberia, was found to share 4-6% of its
genome with living melanesian humans and with no other living group, supporting lateral gene transfer
between two regions outside of Africa.[49][50] The combination of regional continuity inside and outside of
Africa and lateral gene transfer between various regions around the world supports the multiregional
hypothesis.
[edit] Alternate hypotheses
[edit] Polygenism
Main article: Polygenism
The polygenic hypothesis for human origins, popular in the 19th and early 20th centuries, proposed
that human populations had evolved independently and in parallel in the various regions of the world, without
gene interchange. This differs from multiregional evolution in that the multiregional model posits significant
lateral gene flow through migrations or interbreeding between populations, while polygenism envisions none.
The polygenic hypothesis is no longer considered a viable scientific hypothesis since separately evolving
species could not develop the interfertility of modern human populations.[ citation needed]
[edit] Recent African replacement

Main article: Recent African origin of modern humans
The primary competing scientific hypothesis is currently recent African origin of modern humans,
which proposes that modern humans arose as a new species in Africa around 100-200,000 years ago,
moving out of Africa around 50-60,000 years ago to replace existing human species such as Homo erectus
and the Neanderthals without interbreeding.[51][52][53][54] This differs from the multiregional hypothesis in
that the multiregional model predicts interbreeding with preexisting local human populations in any such
migration.[54]
[edit] See also
• Phyletic gradualism
• Neanderthal admixture theory
• Mitochondrial Eve
• Y-chromosomal Adam
[edit] References
1. ^ Wolpoff, MH; JN Spuhler, FH Smith, J Radovcic, G Pope, DW Frayer, R Eckhardt, and G
Clark (1988). "Modern human origins". Science 241 (4867): 772–774. doi:10.1126/science.3136545.
PMID 3136545. http://www.sciencemag.org/cgi/pdf_extract/241/4867/772.
2. ^ a b Wolpoff, Milford, and Caspari, Rachel (1997). Race and Human Evolution. Simon &
Schuster. pp. 28–29.
3. ^ Wolpoff, MH; Hawks J, Caspari R (2000). "Multiregional, not multiple origins". Am J Phys
Anthropol 112 (1): 129–36. doi:10.1002/(SICI)1096-8644(200005)112:1<129::AID-AJPA11>3.0.CO;2-
K. PMID 10766948. http://www3.interscience.wiley.com/journal/71008905/abstract.
4. ^ a b c Wolpoff, MH; JN Spuhler, FH Smith, J Radovcic, G Pope, DW Frayer, R Eckhardt, and
G Clark (1988). "Modern human origins". Science 241 (4867): 772–774.
doi:10.1126/science.3136545. PMID 3136545.
http://www.sciencemag.org/cgi/pdf_extract/241/4867/772.
5. ^ a b Wolpoff, Milford, and Caspari, Rachel (1997). Race and Human Evolution. Simon &
Schuster. pp. 25–26.
6. ^ Wolpoff, Milford H; John Hawks, David W Frayer, Keith Hunley (2001). "Modern Human
Ancestry at the Peripheries: A Test of the Replacement Theory". Science (AAAS) 291 (5502): 293–
297. doi:10.1126/science.291.5502.293. PMID 11209077.
7. ^ Analysis of the full brain case shape confirms the idea that dispersal from a single
population could not explain the early modern human variability, but does not confirm ties to regional
archaic humans. Gunz, P., et. al. (2009-04-14). "Early modern human diversity suggests subdivided
population structure and a complex out-of-Africa scenario". Proceedings of the National Academy of
Sciences of the United States of America 106 (15): 6094–6098. doi:10.1073/pnas.0808160106.
8. ^ Shang et al.; Tong, H; Zhang, S; Chen, F; Trinkaus, E (1999). "An early modern human
from Tianyuan Cave, Zhoukoudian, China" (Free full text). Proceedings of the National Academy of
ISSN 0027-8424. PMID 17416672. PMC 1871827. http://www.pnas.org/cgi/pmidlookup?
9. ^ Phil McKenna, "Chinese challenge to 'out of Africa' theory", New Scientist, Nov 2009
10.^ Wolpoff, Milford; Bruce Mannheim, Alan Mann, John Hawks, Rachel Caspari, Karen R.
Rosenberg, David W. Frayer, George W. Gill and Geoffrey Clark (2004). "Why not the Neandertals?".
World Archaeology 36: 527. doi:10.1080/0043824042000303700.
11.^ Harvati, Katerina; Stephen R Frost and Kieran P McNulty (2004). "Neanderthal taxonomy
reconsidered: Implications of 3D primate models of intra- and interspecific differences". PNAS 101
(5): 1147–1152. doi:10.1073/pnas.0308085100. PMID 14745010. PMC 337021.
http://www.pnas.org/content/101/5/1147.full.
12.^ Pearson, Osbjorn M (2004). "Has the Combination of Genetic and Fossil Evidence Solved
the Riddle of Modern Human Origins?". Evolutionary Anthropology 13: 145–159.
doi:10.1002/evan.20017.
13.^ Duarte C, 2. Maurício J, Pettitt P, Souto P, Trinkaus E, van der Plicht H, Zilhão J (1999).
"The early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern
human emergence in Iberia". Proc Natl Acad Sci USA 96 (13): 7604–7609.
doi:10.1073/pnas.96.13.7604. PMID 10377462. PMC 22133.
http://www.pnas.org/content/96/13/7604.abstract?
ijkey=9335ab52731624a02b5f7f426c4a8c2147934993&keytype2=tf_ipsecsha.
14.^ Trinkaus, E; Moldovan, O; Milota, S; Bîlgăr, A; Sarcina, L; Athreya, S; Bailey, Se; Rodrigo,
R; Mircea, G; Higham, T; Ramsey, Cb; Van, Der, Plicht, J (Sep 2003). "An early modern human from
the Peştera cu Oase, Romania" (free full text). Proceedings of the National Academy of Sciences of
the United States of America 100 (20): 11231–11236. doi:10.1073/pnas.2035108100. ISSN 0027-
8424. PMID 14504393. PMC 208740. http://www.pnas.org/cgi/pmidlookup?
view=long&pmid=14504393. "When multiple measurements are undertaken, the mean result can be
determined through averaging the activity ratios. For Oase 1, this provides a weighted average
activity ratio of 〈14a〉= 1.29 ± 0.15%, resulting in a combined OxA-GrA 14C age of 34,950, +990, and
–890 B.P.".
15.^ Tattersall, Ian, and Schwartz, Jeffrey H. (1999). "Hominids and hybrids: The place of
Neanderthals in human evolution". Proceedings of the National Academy of Sciences of the United
States of America 96 (13): 7117–7119. http://www.pnas.org/cgi/content/full/96/13/7117.
16.^ Trinkaus, E (May 2007). "European early modern humans and the fate of the Neandertals"
(Free full text). Proceedings of the National Academy of Sciences of the United States of America
104 (18): 7367–72. doi:10.1073/pnas.0702214104. ISSN 0027-8424. PMID 17452632.
17.^ http://www.sciencedaily.com/releases/2007/04/070423185434.htm The Emerging Fate Of
The Neandertals
18.^ Cann, Rebecca L., Stoneking, Mark, and Wilson, Allan C. (1987-01-01). "Mitochondrial
DNA and human evolution". nature 325: 31–36. doi:10.1038/325031a0. PMID 3025745.
http://artsci.wustl.edu/~landc/html/cann/.
19.^ Hodgson, JA; Disotell TR (2008). "No evidence of a Neanderthal contribution to modern
human diversity.". Genome Biology (BioMed Central) 9 (2): 206. doi:10.1186/gb-2008-9-2-206.
PMID 18304371. PMC 2374707. http://genomebiology.com/2008/9/2/206.
20.^ Krause, Johannes, et al (2010-03-24). "The complete mitochondrial DNA genome of an
unknown hominin from southern Siberia". Nature 464: 894–897. doi:10.1038/nature08976.
21.^ Wolpoff, Milford, and Caspari, Rachel (1997). Race and Human Evolution. Simon &
Schuster. pp. 30.
22.^ Relethford, JH (2008-03-05). "Genetic evidence and the modern human origins debate".
Heredity (Macmillan) 100 (6): 555–63. doi:10.1038/hdy.2008.14. PMID 18322457.
23.^ "Selection, nuclear genetic variation, and mtDNA". john hawks weblog.
http://johnhawks.net/weblog/reviews/neandertals/neandertal_dna/weaver_roseman_2005.html.
24.^ Hammer, M. F., et. al." (1998). "Out of Africa and Back Again: Nested Cladistic Analysis of
Human Y Chromosome Variation". Molecular Biology and Evolution 15 (4): 427–441.
http://mbe.oxfordjournals.org/content/15/4/427.short.
25.^ Hammer, Mf; Garrigan, D; Wood, E; Wilder, Ja; Mobasher, Z; Bigham, A; Krenz, Jg;
Nachman, Mw (Aug 2004). "Heterogeneous patterns of variation among multiple human x-linked
Loci: the possible role of diversity-reducing selection in non-africans" (Free full text). Genetics 167
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http://www.genetics.org/cgi/content/abstract/167/4/1841?
ijkey=cb14a3724516d1a584feb8454d2c49cd72e003ee&keytype2=tf_ipsecsha. Additional
discussion of these results is available in a video of a presentation given by Hammer at
http://www.youtube.com/watch?v=Ff0jwWaPlnU (video) from about 40 to 50 minutes into the video.
26.^ The CMP-N-acetylneuraminic acid hydroxylase CMAH pseudogene shows 2.9 Mya
coalescence time. Hayakawa, T; Aki, I; Varki, A; Satta, Y; Takahata, N (Feb 2006). "Fixation of the
human-specific CMP-N-acetylneuraminic acid hydroxylase pseudogene and implications of
haplotype diversity for human evolution". Genetics 172 (2): 1139–46.
doi:10.1534/genetics.105.046995. ISSN 0016-6731. PMID 16272417.
27.^ The ALMS1 site suggests ancient and complex evolutionary history with a coalescence
time of about 2 Mya. Scheinfeldt, Lb; Biswas, S; Madeoy, J; Connelly, Cf; Schadt, Ee; Akey, Jm (Jun
2009). "Population genomic analysis of ALMS1 in humans reveals a surprisingly complex
evolutionary history". Molecular biology and evolution 26 (6): 1357–67. doi:10.1093/molbev/msp045.
ISSN 0737-4038. PMID 19279085.
28.^ The PDHA1 (pyruvate dehydrogenase) locus on the X chromosome has an estimated
coalescence time of 1.86 Mya, inconsistent with a recent species origin, although the worldwide
lineage pattern is unlike other autosomal sites and may be consistent with recent dispersal from
Africa. Rosalind M. Harding (March 16, 1999). "More on the X files". Proceedings of the National
Academy of Sciences 96 (6): 2582–2584. doi:10.1073/pnas.96.6.2582.
29.^ A second group finds the same ancient origin for PDHA1, but finds no evidence of a recent
expansion, consistent with other autosomal and X chromosome sites and contrary to mitochondrial
data. Harris, E. E.; Jody Hey (1999). "X chromosome evidence for ancient human histories".
Proceedings of the National Academy of Sciences 96 (6): 3320–4. doi:10.1073/pnas.96.6.3320.
PMID 10077682. http://www.pnas.org/content/96/6/3320.full.
30.^ The ASAH1 gene has two recently differentiated lineages with a coalescence time 2.4±.4
Mya not explainable by balancing selection. The V lineage shows evidence of recent positive
selection. The lineage pattern may be the result of hybridization during a recent range expansion
from Africa with the V lineage tracing to archaic humans from outside Africa, though it is also
consistent with a mixture of two long isolated groups within Africa; it is not consistent with a recent
origination of a modern human species that replaced archaic forms without interbreeding. Kim, Hl;
Satta, Y (Mar 2008). "Population genetic analysis of the N-acylsphingosine amidohydrolase gene
associated with mental activity in humans" (Free full text). Genetics 178 (3): 1505–15.
doi:10.1534/genetics.107.083691. ISSN 0016-6731. PMID 18245333. PMC 2278054.
http://www.genetics.org/cgi/pmidlookup?view=long&pmid=18245333. ""It is speculated that, when
modern humans dispersed from Africa, admixture of the distinct V and M lineages occurred and the V
lineage has since spead in the entire population by possible positive selection."".
31.^ Daniel Garrigan, Zahra Mobasher, Sarah B. Kingan, Jason A. Wilder and Michael F.
Hammer (Aug 2005). "Deep Haplotype Divergence and Long-Range Linkage Disequilibrium at
Xp21.1 Provide Evidence That Humans Descend From a Structured Ancestral Population". Genetics
170 (4): 1849–1856. doi:10.1534/genetics.105.041095.
http://www.genetics.org/cgi/content/abstract/170/4/1849.
32.^ NAT2 SNPs lineages cluster in sub-Saharan Africa, Europe, and East Asia, with genetic
distances scaling with geographic distances. Sabbagh, A; Langaney, A; Darlu, P; Gérard, N;
Krishnamoorthy, R; Poloni, Es (Feb 2008). "Worldwide distribution of NAT2 diversity: implications for
NAT2 evolutionary history" (Free full text). BMC genetics 9: 21. doi:10.1186/1471-2156-9-21.
PMID 18304320. PMC 2292740. http://www.biomedcentral.com/1471-2156/9/21. Also see map; may
resize browser window.
33.^ The NAT1 lineage tree is rooted in Eurasia with a coalescence time of 2.0 Mya that cannot
be explained by balancing selection and with the NAT1*11A haplotype absent from subsaharan
Africa. Patin, E; Barreiro, Lb; Sabeti, Pc; Austerlitz, F; Luca, F; Sajantila, A; Behar, Dm; Semino, O;
Sakuntabhai, A; Guiso, N; Gicquel, B; Mcelreavey, K; Harding, Rm; Heyer, E; Quintana-Murci, L (Mar
2006). "Deciphering the ancient and complex evolutionary history of human arylamine N-
acetyltransferase genes". American journal of human genetics 78 (3): 423–36. doi:10.1086/500614.
ISSN 0002-9297. PMID 16416399.
34.^ johnhawks.net "Variation in NAT1 and NAT2". john hawks weblog.
http://johnhawks.net/weblog/reviews/genetics/diet/nat1_nat2_patin_2006_selection.html
johnhawks.net. Retrieved 2011-01.04.
35.^ Garrigan, D; Mobasher, Z; Severson, T; Wilder, Ja; Hammer, Mf (Feb 2005). "Evidence for
archaic Asian ancestry on the human X chromosome" (Free full text). Molecular biology and
evolution 22 (2): 189–92. doi:10.1093/molbev/msi013. ISSN 0737-4038. PMID 15483323.
http://mbe.oxfordjournals.org/cgi/pmidlookup?view=long&pmid=15483323.
36.^ Cox, Mp; Mendez, Fl; Karafet, Tm; Pilkington, Mm; Kingan, Sb; Destro-Bisol, G;
Strassmann, Bi; Hammer, Mf (Jan 2008). "Testing for archaic hominin admixture on the X
chromosome: model likelihoods for the modern human RRM2P4 region from summaries of
genealogical topology under the structured coalescent" (Free full text). Genetics 178 (1): 427–37.
doi:10.1534/genetics.107.080432. ISSN 0016-6731. PMID 18202385. PMC 2206091.
http://www.genetics.org/cgi/pmidlookup?view=long&pmid=18202385.
37.^ J. Hardy, A. Pittman, A. Myers, K. Gwinn-Hardy, H.C. Fung, R. de Silva, M. Hutton and J.
Duckworth (2005). "Evidence suggesting that Homo neanderthalensis contributed the H2 MAPT
haplotype to Homo sapiens". Biochemical Society Transactions 33, part 4: 582–585.
http://www.biochemsoctrans.org/bst/033/0582/0330582.pdf.
38.^ Zody, Mc; Jiang, Z; Fung, Hc; Antonacci, F; Hillier, Lw; Cardone, Mf; Graves, Ta; Kidd, Jm;
Cheng, Z; Abouelleil, A; Chen, L; Wallis, J; Glasscock, J; Wilson, Rk; Reily, Ad; Duckworth, J;
Ventura, M; Hardy, J; Warren, Wc; Eichler, Ee (Aug 2008). "Evolutionary toggling of the MAPT
17q21.31 inversion region". Nature genetics 40: 1076. doi:10.1038/ng.193. ISSN 1061-4036.
PMID 18690220.
39.^ Almos, Pz; Horváth, S; Czibula, A; Raskó, I; Sipos, B; Bihari, P; Béres, J; Juhász, A; Janka,
Z; Kálmán, J (Nov 2008). "H1 tau haplotype-related genomic variation at 17q21.3 as an Asian
heritage of the European Gypsy population". Heredity 101 (5): 416–9. doi:10.1038/hdy.2008.70.
ISSN 0018-067X. PMID 18648385.
40.^ Stefansson H, Helgason A, Thorleifsson G, Steinthorsdottir V, Masson G, Barnard J, Baker
A, Jonasdottir A, Ingason A, Gudnadottir VG, et. al. (2005-01-16). "A common inversion under
selection in Europeans". nature genetics 37: 129–137. doi:10.1038/ng1508.
http://www.genetics.org/cgi/content/abstract/170/4/1849.
41.^ Evans, Pd; Mekel-Bobrov, N; Vallender, Ej; Hudson, Rr; Lahn, Bt (Nov 2006). "Evidence
that the adaptive allele of the brain size gene microcephalin introgressed into Homo sapiens from an
archaic Homo lineage" (Free full text). Proceedings of the National Academy of Sciences of the
United States of America 103 (48): 18178–83. doi:10.1073/pnas.0606966103. ISSN 0027-8424.
42.^ Trinkaus, E (May 2007). "European early modern humans and the fate of the Neandertals"
(Free full text). Proceedings of the National Academy of Sciences of the United States of America
104 (18): 7367–72. doi:10.1073/pnas.0702214104. ISSN 0027-8424. PMID 17452632.
43.^ Evans, Pd; Gilbert, Sl; Mekel-Bobrov, N; Vallender, Ej; Anderson, Jr; Vaez-Azizi, Lm;
Tishkoff, Sa; Hudson, Rr; Lahn, Bt (Sep 2005). "Microcephalin, a gene regulating brain size,
continues to evolve adaptively in humans". Science 309 (5741): 1717–20.
doi:10.1126/science.1113722. ISSN 0036-8075. PMID 16151009.
44.^ "Introgression and microcephalin FAQ". john hawks weblog.
http://johnhawks.net/weblog/reviews/neandertals/neandertal_dna/introgression_faq_2006.html.
45.^ Templeton, Alan R. (2005). "Haplotype Trees and Modern Human Origins". Yearbook of
Physical Anthropology 48: 33–59. doi:10.1002/ajpa.20351. http://esa.ipb.pt/pdf/28.pdf.
46.^ Noonan, James P., et. al. (2006-11-17). "Sequencing and Analysis of Neanderthal
Genomic DNA". Science 314 (5802): 1113–1118. doi:10.1126/science.1131412.
http://www.sciencemag.org/content/328/5979/710.
47.^ Green, Richard E., et. al. (2010-05-07). "A Draft Sequence of the Neandertal Genome".
48.^ "NEANDERTALS LIVE!". john hawks weblog.
http://johnhawks.net/weblog/reviews/neandertals/neandertal_dna/neandertals-live-genome-
sequencing-2010.html. Retrieved 2010-12-31.
49.^ Reich, David, et al (2010-12-23). "Genetic history of an archaic hominin group from
Denisova Cave in Siberia". Nature 468: 1053–1060. doi:10.1038/nature09710.
http://www.nature.com/nature/journal/v468/n7327/full/nature09710.html.
50.^ "The Denisova genome FAQ". john hawks weblog.
http://johnhawks.net/weblog/reviews/neandertals/neandertal_dna/denisova-nuclear-genome-reich-
2010.html. Retrieved 2010-12-31.
51.^ Hua Liu, et al. (2006). "A Geographically Explicit Genetic Model of Worldwide Human-
Settlement History". American Journal of Human Genetics 79 (2): 230–237. doi:10.1086/505436.
PMID 16826514.
52.^ Weaver, Timothy D; Charles C. Roseman (2008). "New developments in the genetic
evidence for modern human origins". Evolutionary Anthropology: Issues, News, and Reviews (Wiley-
Liss) 17 (1): 69–80. doi:10.1002/evan.20161.
http://www3.interscience.wiley.com/journal/117921411/abstract.
53.^ Fagundes, NJ; Ray N, Beaumont M, Neuenschwander S, Salzano FM, Bonatto SL,
Excoffier L. (2007). "Statistical evaluation of alternative models of human evolution". Proc Natl Acad
Sci USA 104 (45): 17614–9. doi:10.1073/pnas.0708280104. PMID 17978179. PMC 2077041.
http://www.pnas.org/content/104/45/17614.long.
54.^ a b Wolpoff, Milford, and Caspari, Rachel (1997). Race and Human Evolution. Simon &
Schuster. p. 42.

• From www.conrante.com: Templeton's lattice diagram showing major gene flows graphically.
• From darwin.eeb.uconn.edu: notes on drift and migration with equations for calculating the
effects on allele frequencies of different populations.
[hide]v · d · ePart of the series on Human evolution
Hominini
Sahelanthropus tchadensis · Orrorin tugenensis · Ardipithecus · Kenyanthropus platyops
Australopithecines
Australopithecus: A. anamensis · A. afarensis · A. bahrelghazali · A. africanus · A. garhi · A. sediba

Paranthropus: P. aethiopicus · P. boisei · P. robustus
Humans and Proto-humans

Homo: H. gautengensis · H. habilis · H. rudolfensis · H. georgicus · H. ergaster · H. erectus
(H. e. erectus · H. e. lantianensis · H. e. palaeojavanicus · H. e. pekinensis · H. e. nankinensis ·
H. e. wushanensis · H. e. yuanmouensis · H. e. soloensis) · H. cepranensis · H. antecessor ·
H. heidelbergensis · Denisova hominin · H. neanderthalensis · H. rhodesiensis · H. floresiensis · Archaic
Homo sapiens · Anatomically modern humans (H. s. idaltu · H. s. sapiens)
Topics: Timeline of human evolution · List of human evolution fossils · Human evolutionary genetics
Models: Recent African origin · Multiregional origin
Retrieved from "http://en.wikipedia.org/wiki/Multiregional_origin_of_modern_humans"
Categories: Human evolution | Race

W000
Dinosaur

For other uses, see Dinosaur (disambiguation).
Dinosaur
Opisthokonta
Dinosaurs
Temporal range:
Late Triassic-Late Cretaceous, 231.4–65.5 Ma
PreЄ
g Descendant taxon Aves survives to present
Mounted skeletons of Tyrannosaurus (left) and Apatosaurus (right) at the American Museum of
Natural History
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
(unranked): Ornithodira
(unranked): Dinosauromorpha
(unranked): Dinosauriformes
Dinosauria
Superorder:
Owen, 1842
Orders and suborders
• †Ornithischia
• †Cerapoda
• †Thyreophora
• Saurischia
• †Sauropodomorpha
• Theropoda
Dinosaurs are a diverse group of animals that were the dominant terrestrial vertebrates for over
160 million years, from the late Triassic period (about 230 million years ago) until the end of the Cretaceous
(about 65 million years ago). The extinction of most dinosaur species occurred during the Cretaceous–
Tertiary extinction event. The fossil record indicates that birds evolved within theropod dinosaurs during the
Jurassic period. Some of them survived the Cretaceous–Tertiary extinction event, including the ancestors of
all modern birds. Consequently, in modern classification systems, birds are considered a type of dinosaur—
the only group of which that has survived to the present day.[1][2]
Dinosaurs are a diverse and varied group of animals; birds, at over 9,000 species, are the most
diverse group of vertebrate besides perciform fish.[3] Paleontologists have identified over 500 distinct
genera[4] and more than 1,000 different species of non-avian dinosaurs.[5] Dinosaurs are represented on
every continent by both extant species and fossil remains.[6] Some dinosaurs are or were herbivorous,
others carnivorous. Some have been bipedal, others quadrupedal, and others have been able to shift
between these body postures. Many non-avian species developed elaborate skeletal modifications such as
bony armor, horns or crests. Avian dinosaurs have been the planet's dominant flying vertebrate since the
extinction of the pterosaurs. Although generally known for the large size of some species, most dinosaurs
were human-sized or even smaller. Most groups of dinosaurs are known to have built nests and laid eggs.
The term "dinosaur" was coined in 1842 by the English paleontologist Richard Owen, and derives
from Greek δεινός (deinos) "terrible, powerful, wondrous" + σαῦρος (sauros) "lizard". Through the first half of
the twentieth century, most of the scientific community believed dinosaurs to have been sluggish,
unintelligent cold-blooded animals. Most research conducted since the 1970s, however, has indicated that
dinosaurs were active animals with elevated metabolisms and numerous adaptations for social interaction.
Since the first dinosaur fossils were recognized in the early nineteenth century, mounted dinosaur
skeletons have been major attractions at museums around the world, and dinosaurs have become a part of
world culture. They have been featured in best-selling books and films such as Jurassic Park, and new
discoveries are regularly covered by the media. In informal speech, the word "dinosaur" is used to describe
things that are impractically large, slow-moving, obsolete, or bound for extinction,[7] reflecting the outdated
view that dinosaurs were maladapted monsters of the ancient world.
Contents
[hide]
• 1 Etymology
• 2 Modern definition
• 2.1 General description
• 2.2 Distinguishing anatomical features
• 3 Natural history
• 3.1 Origins and early evolution
• 3.2 Low diversification in the Cretaceous
• 3.3 Classification
• 3.4 Evolution and paleobiogeography
• 4 Paleobiology
• 4.1 Size
• 4.1.1 Largest and smallest
• 4.2 Behavior
• 4.3 Communication and vocalization
• 4.4 Physiology
• 4.5 Soft tissue and DNA
• 5 Feathers and the origin of birds
• 5.1 Feathers
• 5.2 Skeleton
• 5.3 Soft anatomy
Etymology
The taxon Dinosauria was formally named in 1842 by Sir Richard Owen, who used it to refer to the
"distinct tribe or sub-order of Saurian Reptiles" that were then being recognized in England and around the
world.[8]:103 The term is derived from the Greek words δεινός (deinos meaning "terrible", "powerful", or
"wondrous") and σαῦρος (sauros meaning "lizard" or "reptile").[8]:103[9] Though the taxonomic name has
often been interpreted as a reference to dinosaurs' teeth, claws, and other fearsome characteristics, Owen
intended it merely to evoke their size and majesty.[10] In colloquial English "dinosaur" is sometimes used to
describe an obsolete or unsuccessful thing or person,[11] despite the dinosaurs' 160 million year reign and
the global abundance and diversity of their avian descendants: modern-day birds.
Modern definition
Triceratops skeleton at the American Museum of Natural History in New York City
Under phylogenetic taxonomy, dinosaurs are usually defined as the group consisting of " Triceratops,
Neornithes [modern birds], their most recent common ancestor, and all descendants."[12] It has also been
suggested that Dinosauria be defined with respect to the most recent common ancestor of Megalosaurus and
Iguanodon, because these were two of the three genera cited by Richard Owen when he recognized the
Dinosauria.[13] Both definitions result in the same set of animals being defined as dinosaurs, including
theropods (mostly bipedal carnivores), sauropodomorphs (mostly large herbivorous quadrupeds with long
necks and tails), ankylosaurians (armored herbivorous quadrupeds), stegosaurians (plated herbivorous
quadrupeds), ceratopsians (herbivorous quadrupeds with horns and frills), and ornithopods (bipedal or
quadrupedal herbivores including "duck-bills"). These definitions are written to correspond with scientific
conceptions of dinosaurs that predate the modern use of phylogenetics. The continuity of meaning is
intended to prevent confusion about what the term "dinosaur" means.
There is a wide consensus among paleontologists that birds are the descendants of theropod
dinosaurs. Using the strict cladistical definition that all descendants of a single common ancestor must be
included in a group for that group to be natural, birds would thus be dinosaurs and dinosaurs are, therefore,
not extinct. Birds are classified by most paleontologists as belonging to the subgroup Maniraptora, which are
coelurosaurs, which are theropods, which are saurischians, which are dinosaurs.[14]
From the point of view of cladistics, birds are dinosaurs, but in ordinary speech the word "dinosaur"
does not include birds. Additionally, referring to dinosaurs that are not birds as "non-avian dinosaurs" is
cumbersome. For clarity, this article will use "dinosaur" as a synonym for "non-avian dinosaur". The term
"non-avian dinosaur" will be used for emphasis as needed.
General description
Stegosaurus skeleton, Field Museum, Chicago

Using one of the above definitions, dinosaurs (aside from birds) can be generally described as
terrestrial archosaurian reptiles with limbs held erect beneath the body, that existed from the Late Triassic
(first appearing in the Carnian faunal stage) to the Late Cretaceous (going extinct at the end of the
Maastrichtian).[15] Many prehistoric animals are popularly conceived of as dinosaurs, such as ichthyosaurs,
mosasaurs, plesiosaurs, pterosaurs, and Dimetrodon, but are not classified scientifically as dinosaurs.
Marine reptiles like ichthyosaurs, mosasaurs, and plesiosaurs were neither terrestrial nor archosaurs;
pterosaurs were archosaurs but not terrestrial; and Dimetrodon was a Permian animal more closely related to
mammals.[16] Dinosaurs were the dominant terrestrial vertebrates of the Mesozoic, especially the Jurassic
and Cretaceous. Other groups of animals were restricted in size and niches; mammals, for example, rarely
exceeded the size of a cat, and were generally rodent-sized carnivores of small prey.[17] One notable
exception is Repenomamus giganticus, a triconodont weighing between 12 kilograms (26 lb) and
14 kilograms (31 lb) that is known to have eaten small dinosaurs like young Psittacosaurus.[18]
Dinosaurs were an extremely varied group of animals; according to a 2006 study, over 500 dinosaur
genera have been identified with certainty so far, and the total number of genera preserved in the fossil
record has been estimated at around 1850, nearly 75% of which remain to be discovered.[4] An earlier study
predicted that about 3400 dinosaur genera existed, including many which would not have been preserved in
the fossil record.[19] As of September 17, 2008, 1047 different species of dinosaurs have been named.[5]
Some were herbivorous, others carnivorous. Some dinosaurs were bipeds, some were quadrupeds, and
others, such as Ammosaurus and Iguanodon, could walk just as easily on two or four legs. Many had bony
armor, or cranial modifications like horns and crests. Although known for large size, many dinosaurs were
human-sized or smaller. Dinosaur remains have been found on every continent on Earth, including
Antarctica.[6] No dinosaurs are known to have lived in marine or aerial habitats, although it is possible some
feathered theropods were flyers. There is also evidence that some spinosaurids had semi-aquatic habits.[20]
Distinguishing anatomical features

While recent discoveries have made it more difficult to present a universally agreed-upon list of
dinosaurs' distinguishing features, nearly all dinosaurs discovered so far share certain modifications to the
ancestral archosaurian skeleton. Although some later groups of dinosaurs featured further modified versions
of these traits, they are considered typical across Dinosauria; the earliest dinosaurs had them and passed
them on to all their descendants. Such common features across a taxonomic group are called
synapomorphies.
Dinosaur synapomorphies include an elongated crest on the humerus, or upper arm bone, to
accommodate the attachment of deltopectoral muscles; a shelf at the rear of the ilium, or main hip bone; a
tibia, or shin bone, featuring a broad lower edge and a flange pointing out and to the rear; and an ascending
projection on the astragalus, one of the ankle bones, which secures it to the tibia.[12]
Edmontonia was an armored dinosaur of the group Ankylosauria

A variety of other skeletal features were shared by many dinosaurs. However, because they were
either common to other groups of archosaurs or were not present in all early dinosaurs, these features are
not considered to be synapomorphies. For example, as diapsid reptiles, dinosaurs ancestrally had two pairs
of temporal fenestrae (openings in the skull behind the eyes), and as members of the diapsid group
Archosauria, had additional openings in the snout and lower jaw.[21] Additionally, several characteristics
once thought to be synapomorphies are now known to have appeared before dinosaurs, or were absent in
the earliest dinosaurs and independently evolved by different dinosaur groups. These include an elongated
scapula, or shoulder blade; a sacrum composed of three or more fused vertebrae (three are found in some
other archosaurs, but only two are found in Herrerasaurus);[12] and an acetabulum, or hip socket, with a hole
at the center of its inside surface (closed in Saturnalia, for example).[22] Another difficulty of determining
distinctly dinosaurian features is that early dinosaurs and other archosaurs from the Late Triassic are often
poorly known and were similar in many ways; these animals have sometimes been misidentified in the
literature.[23]
Hip joints and hindlimb postures

Dinosaurs stood erect in a manner similar to most modern mammals, but distinct from most other
reptiles, whose limbs sprawl out to either side.[24] Their posture was due to the development of a laterally
facing recess in the pelvis (usually an open socket) and a corresponding inwardly facing distinct head on the
femur.[25] Their erect posture enabled dinosaurs to breathe easily while moving, which likely permitted
stamina and activity levels that surpassed those of "sprawling" reptiles.[26] Erect limbs probably also helped
support the evolution of large size by reducing bending stresses on limbs.[27] Some non-dinosaurian
archosaurs, including rauisuchians, also had erect limbs but achieved this by a "pillar erect" configuration of
the hip joint, where instead of having a projection from the femur insert on a socket on the hip, the upper
pelvic bone was rotated to form an overhanging shelf.[27]
Natural history
Origins and early evolution
Main article: Evolution of dinosaurs
Marasuchus, a dinosaur-like ornithodiran

For a long time many scientists thought dinosaurs were polyphyletic with multiple groups of unrelated
"dinosaurs" evolving due to similar pressures,[28][29][30] but dinosaurs are now known to have formed a
single group.[12][31]
Dinosaurs diverged from their archosaur ancestors approximately 230 million years ago during the
Middle to Late Triassic period, roughly 20 million years after the Permian–Triassic extinction event wiped out
an estimated 95% of all life on Earth.[32][33] Radiometric dating of the rock formation that contained fossils
from the early dinosaur genus Eoraptor establishes its presence in the fossil record at this time.
Paleontologists believe Eoraptor resembles the common ancestor of all dinosaurs;[34] if this is true, its traits
suggest that the first dinosaurs were small, bipedal predators.[35] The discovery of primitive, dinosaur-like
ornithodirans such as Marasuchus and Lagerpeton in Argentinian Middle Triassic strata supports this view;
analysis of recovered fossils suggests that these animals were indeed small, bipedal predators.
When dinosaurs appeared, terrestrial habitats were occupied by various types of basal archosaurs
and therapsids, such as aetosaurs, cynodonts, dicynodonts, ornithosuchids, rauisuchias, and rhynchosaurs.
Most of these other animals became extinct in the Triassic, in one of two events. First, at about the boundary
between the Carnian and Norian faunal stages (about 215 million years ago), dicynodonts and a variety of
basal archosauromorphs, including the prolacertiforms and rhynchosaurs, became extinct. This was followed
by the Triassic–Jurassic extinction event (about 200 million years ago), that saw the end of most of the other
groups of early archosaurs, like aetosaurs, ornithosuchids, phytosaurs, and rauisuchians. These losses left
behind a land fauna of crocodylomorphs, dinosaurs, mammals, pterosaurians, and turtles.[12]
The early forms Herrerasaurus (large), Eoraptor (small) and a Plateosaurus skull
The first few lines of primitive dinosaurs diversified through the Carnian and Norian stages of the
Triassic, most likely by occupying the niches of groups that became extinct. Traditionally, dinosaurs were
thought to have replaced the variety of other Triassic land animals by proving superior through a long period
of competition. This now appears unlikely, for several reasons. Dinosaurs do not show a pattern of steadily
increasing in diversity and numbers, as would be predicted if they were competitively replacing other groups;
instead, they were very rare through the Carnian, making up only 1–2% of individuals present in faunas. In
the Norian, however, after the extinction of several other groups, they became significant components of
faunas, representing 50–90% of individuals. Also, what had been viewed as a key adaptation of dinosaurs,
their erect stance, is now known to have been present in several contemporaneous groups that were not as
successful (aetosaurs, ornithosuchids, rauisuchians, and some groups of crocodylomorphs). Finally, the Late
Triassic itself was a time of great upheaval in life, with shifts in plant life, marine life, and climate.[12]
Crurotarsans, today represented only by crocodilians but in the Late Triassic also encompassing such now-
extinct groups as aetosaurs, phytosaurs, ornithosuchians, and rauisuchians, were actually more diverse in
the Late Triassic than dinosaurs, indicating that the survival of dinosaurs had more to do with luck than
superiority.[36]
Low diversification in the Cretaceous

Statistical analyses based on raw data suggest that dinosaurs diversified, i.e. the number of species
increased, in the Late Cretaceous. However in July 2008 Graeme T. Lloyd et al. argued that this apparent
diversification was an illusion caused by sampling bias, because Late Cretaceous rocks have been very
heavily studied. Instead, they wrote, dinosaurs underwent only two significant diversifications in the Late
Cretaceous, the initial radiations of the euhadrosaurs and ceratopsians. In the Mid Cretaceous, the flowering
angiosperm plants became a major part of terrestrial ecosystems, which had previous been dominated by
gymnosperms such as conifers. Dinosaur coprolites (fossilized dung) indicate that, while some ate
angiosperms, most herbivorous dinosaurs mainly ate gymnosperms. Meanwhile herbivorous insects and
mammals diversified rapidly to take advantage of the new type of plant food, while lizards, snakes,
crocodilians and birds also diversified at the same time. Lloyd et al. suggest that dinosaurs' failure to diversify
as ecosystems were changing doomed them to extinction.[37]
Classification
Main article: Dinosaur classification
Dinosaurs (including birds) are archosaurs, like modern crocodilians. Archosaurs' diapsid skulls have
two holes, called temporal fenestrae, located where the jaw muscles attach, and an additional antorbital
fenestra in front of the eyes. Most reptiles (including birds) are diapsids; mammals, with only one temporal
fenestra, are called synapsids; and turtles, with no temporal fenestra, are anapsids. Anatomically, dinosaurs
share many other archosaur characteristics, including teeth that grow from sockets rather than as direct
extensions of the jawbones. Within the archosaur group, dinosaurs are differentiated most noticeably by their
gait. Dinosaur legs extend directly beneath the body, whereas the legs of lizards and crocodilians sprawl out
to either side.
Collectively, dinosaurs are usually regarded as a superorder or an unranked clade. They are divided
into two orders, Saurischia and Ornithischia, depending upon pelvic structure. Saurischia includes those taxa
sharing a more recent common ancestor with birds than with Ornithischia, while Ornithischia includes all taxa
sharing a more recent common ancestor with Triceratops than with Saurischia. Saurischians ("lizard-hipped",
from the Greek sauros (σαυρος) meaning "lizard" and ischion (ισχιον) meaning "hip joint") retained the hip
structure of their ancestors, with a pubis bone directed cranially, or forward.[25] This basic form was modified
by rotating the pubis backward to varying degrees in several groups ( Herrerasaurus,[38] therizinosauroids,
[39] dromaeosaurids,[40] and birds[14]). Saurischia includes the theropods (bipedal and mostly carnivores,
except for birds) and sauropodomorphs (long-necked quadrupedal herbivores).
By contrast, ornithischians ("bird-hipped", from the Greek ornitheios (ορνιθειος) meaning "of a bird"
and ischion (ισχιον) meaning "hip joint") had a pelvis that superficially resembled a bird's pelvis: the pubis
bone was oriented caudally (rear-pointing) Unlike birds, the ornithischian pubis also usually had an additional
forward-pointing process. Ornithischia includes a variety of herbivores. (NB: the terms "lizard hip" and "bird
hip" are misnomers – birds evolved from dinosaurs with "lizard hips".)
Edmontosaurus
Tyrannosaurus Ornithischian pelvis pelvis (showing ornithischian
Saurischian pelvis pelvis (showing structure (left side) structure – left side)
structure (left side) saurischian structure –
left side)
The following is a simplified classification of dinosaur families. A more detailed version can be found
at List of dinosaur classifications.
Several macronarian Sauropods: from left to right Camarasaurus, Brachiosaurus, Giraffatitan, and
Euhelopus
Various ornithopod dinosaurs and one heterodontosaurid. Far left: Camptosaurus, left: Iguanodon,
center background: Shantungosaurus, center foreground: Dryosaurus, right: Corythosaurus, far right (small):
Heterodontosaurus, far right (large) Tenontosaurus.
• Dinosauria
• Saurischia (theropods and sauropods)
• †Herrerasaurians (early bipedal predators)
• Theropods (all bipedal; most were carnivores)
• †Coelophysoids (Coelophysis and close relatives)
• †Ceratosaurians (Ceratosaurus and abelisaurids – the latter were
important Late Cretaceous predators in southern continents)
• †Spinosauroids (long bodies; short arms; some with crocodile-like skulls
and bony "sails" on their backs)
• †Carnosaurians (Allosaurus and close relatives, like
Carcharodontosaurus)
• Coelurosaurians (diverse, with a range of body sizes and niches)
• †Tyrannosauroids (small to gigantic, often with reduced forelimbs)
• †Ornithomimosaurians ("ostrich-mimics"; mostly toothless;
carnivores to possible herbivores)
• †Therizinosauroids (bipedal herbivores with large hand claws and
small heads)
• †Oviraptorosaurians (mostly toothless; their diet and lifestyle are
uncertain)
• †Dromaeosaurids (popularly known as "raptors"; bird-like
carnivores)
• †Troodontids (similar to dromaeosaurids, but more lightly built)
• Avialans (flying dinosaurs, including modern birds: the only living
dinosaurs)
• †Sauropodomorphs (quadrupedal herbivores with small heads, long necks and
tails, and elephant-like bodies)
• †"Prosauropods" (early relatives of sauropods; small to quite large; some
possibly omnivorous; bipeds and quadrupeds)
• †Sauropods (very large, usually over 15 meters long [49 ft])
• †Diplodocoids (skulls and tails elongated; teeth typically narrow
and pencil-like)
• †Macronarians (boxy skulls; spoon-shaped or pencil-shaped teeth)
• †Brachiosaurids (very long necks; forelimbs longer than
hindlimbs)
• †Titanosaurians (diverse; stocky, with wide hips; most
common in the Late Cretaceous of southern continents)
• †Ornithischians (diverse bipedal and quadrupedal
herbivores)
• †Heterodontosaurids (meter- or yard-scale herbivores or omnivores with
prominent canine teeth)
• †Thyreophorans (armored dinosaurs; mostly quadrupeds)
• †Ankylosaurians (scutes as primary armor; some had club-like tails)
• †Stegosaurians (spikes and plates as primary armor)
• †Ornithopods (diverse, from meter- or yard-scale bipeds to 12-meter (39 ft)
animals that could move as both bipeds and quadrupeds; evolved a method of chewing
using skull flexibility and large numbers of teeth)
• †Hadrosaurids ("duckbilled dinosaurs")
• †Pachycephalosaurians ("bone-heads"; bipeds with domed or knobby growth on
skulls)
• †Ceratopsians (dinosaurs with horns and frills, although most early forms had
only the beginnings of these features)
Evolution and paleobiogeography

Dinosaur evolution after the Triassic follows changes in vegetation and the location of continents. In
the Late Triassic and Early Jurassic, the continents were connected as the single landmass Pangaea, and
there was a worldwide dinosaur fauna mostly composed of coelophysoid carnivores and prosauropod
herbivores.[41] Gymnosperm plants (particularly conifers), a potential food source, radiated in the Late
Triassic. Prosauropods did not have sophisticated mechanisms for processing food in the mouth, and so
must have employed other means of breaking down food farther along the digestive tract.[42] The general
homogeneity of dinosaurian faunas continued into the Middle and Late Jurassic, where most localities had
predators consisting of ceratosaurians, spinosauroids, and carnosaurians, and herbivores consisting of
stegosaurian ornithischians and large sauropods. Examples of this include the Morrison Formation of North
America and Tendaguru Beds of Tanzania. Dinosaurs in China show some differences, with specialized
sinraptorid theropods and unusual, long-necked sauropods like Mamenchisaurus.[41] Ankylosaurians and
ornithopods were also becoming more common, but prosauropods had become extinct. Conifers and
pteridophytes were the most common plants. Sauropods, like the earlier prosauropods, were not oral
processors, but ornithischians were evolving various means of dealing with food in the mouth, including
potential cheek-like organs to keep food in the mouth, and jaw motions to grind food.[42] Another notable
evolutionary event of the Jurassic was the appearance of true birds, descended from maniraptoran
coelurosaurians.[14]
By the Early Cretaceous and the ongoing breakup of Pangaea, dinosaurs were becoming strongly
differentiated by landmass. The earliest part of this time saw the spread of ankylosaurians, iguanodontians,
and brachiosaurids through Europe, North America, and northern Africa. These were later supplemented or
replaced in Africa by large spinosaurid and carcharodontosaurid theropods, and rebbachisaurid and
titanosaurian sauropods, also found in South America. In Asia, maniraptoran coelurosaurians like
dromaeosaurids, troodontids, and oviraptorosaurians became the common theropods, and ankylosaurids
and early ceratopsians like Psittacosaurus became important herbivores. Meanwhile, Australia was home to
a fauna of basal ankylosaurians, hypsilophodonts, and iguanodontians.[41] The stegosaurians appear to
have gone extinct at some point in the late Early Cretaceous or early Late Cretaceous. A major change in the
Early Cretaceous, which would be amplified in the Late Cretaceous, was the evolution of flowering plants. At
the same time, several groups of dinosaurian herbivores evolved more sophisticated ways to orally process
food. Ceratopsians developed a method of slicing with teeth stacked on each other in batteries, and
iguanodontians refined a method of grinding with tooth batteries, taken to its extreme in hadrosaurids.[42]
Some sauropods also evolved tooth batteries, best exemplified by the rebbachisaurid Nigersaurus.[43]
There were three general dinosaur faunas in the Late Cretaceous. In the northern continents of North
America and Asia, the major theropods were tyrannosaurids and various types of smaller maniraptoran
theropods, with a predominantly ornithischian herbivore assemblage of hadrosaurids, ceratopsians,
ankylosaurids, and pachycephalosaurians. In the southern continents that had made up the now-splitting
Gondwana, abelisaurids were the common theropods, and titanosaurian sauropods the common herbivores.
Finally, in Europe, dromaeosaurids, rhabdodontid iguanodontians, nodosaurid ankylosaurians, and
titanosaurian sauropods were prevalent.[41] Flowering plants were greatly radiating,[42] with the first grasses
appearing by the end of the Cretaceous.[44] Grinding hadrosaurids and shearing ceratopsians became
extremely diverse across North America and Asia. Theropods were also radiating as herbivores or
omnivores, with therizinosaurians and ornithomimosaurians becoming common.[42]
The Cretaceous–Tertiary extinction event, which occurred approximately 65 million years ago at the
end of the Cretaceous period, caused the extinction of all dinosaurs except for the birds. Some other diapsid
groups, such as crocodilians, lizards, snakes, sphenodontians, and choristoderans, also survived the event.
[45]
Paleobiology
Knowledge about dinosaurs is derived from a variety of fossil and non-fossil records, including
fossilized bones, feces, trackways, gastroliths, feathers, impressions of skin, internal organs and soft tissues.
[46][47] Many fields of study contribute to our understanding of dinosaurs, including physics (especially
biomechanics), chemistry, biology, and the earth sciences (of which paleontology is a sub-discipline). Two
topics of particular interest and study have been dinosaur size and behavior.
Size
Main article: Dinosaur size
Scale diagram comparing the largest known dinosaurs in five major clades and a human
While the evidence is incomplete, it is clear that, as a group, dinosaurs were large. Even by dinosaur
standards, the sauropods were gigantic. For much of the dinosaur era, the smallest sauropods were larger
than anything else in their habitat, and the largest were an order of magnitude more massive than anything
else that has since walked the Earth. Giant prehistoric mammals such as the Indricotherium and the
Columbian mammoth were dwarfed by the giant sauropods, and only a handful of modern aquatic animals
approach or surpass them in size – most notably the blue whale, which reaches up to 173000 kg (381000 lb)
and over 30 meters (100 ft) in length.[48] There are several proposed advantages for the large size of
sauropods, including protection from predation, reduction of energy use, and longevity, but it may be that the
most important advantage was dietary. Large animals are more efficient at digestion than small animals,
because food spends more time in their digestive systems. This also permits them to subsist on food with
lower nutritive value than smaller animals. Sauropod remains are mostly found in rock formations interpreted
as dry or seasonally dry, and the ability to eat large quantities of low-nutrient browse would have been
advantageous in such environments.[49]
Most dinosaurs, however, were much smaller than the giant sauropods. Current evidence suggests
that dinosaur average size varied through the Triassic, early Jurassic, late Jurassic and Cretaceous periods.
[34] Theropod dinosaurs, when sorted by estimated weight into categories based on order of magnitude,
most often fall into the 100 to 1000 kilogram (220 to 2200 lb) category, whereas recent predatory carnivorans
peak in the 10 to 100 kilogram (22 to 220 lb) category.[50] The mode of dinosaur body masses is between
one and ten metric tonnes.[51] This contrasts sharply with the size of Cenozoic mammals, estimated by the
National Museum of Natural History as about 2 to 5 kilograms (5 to 10 lb).[52]
Largest and smallest

Only a tiny percentage of animals ever fossilize, and most of these remain buried in the earth. Few of
the specimens that are recovered are complete skeletons, and impressions of skin and other soft tissues are
rare. Rebuilding a complete skeleton by comparing the size and morphology of bones to those of similar,
better-known species is an inexact art, and reconstructing the muscles and other organs of the living animal
is, at best, a process of educated guesswork. As a result, scientists will probably never be certain of the
largest and smallest dinosaurs.
Comparative size of Giraffatitan
The tallest and heaviest dinosaur known from good skeletons is Giraffatitan brancai (previously
classified as a species of Brachiosaurus). Its remains were discovered in Tanzania between 1907–12. Bones
from multiple similar-sized individuals were incorporated into the skeleton now mounted and on display at the
Humboldt Museum of Berlin;[53] this mount is 12 meters (39 ft) tall and 22.5 meters (74 ft) long, and would
have belonged to an animal that weighed between 30000 and 60000 kilograms (70000 and 130000 lb). The
longest complete dinosaur is the 27-meter (89 ft) long Diplodocus, which was discovered in Wyoming in the
United States and displayed in Pittsburgh's Carnegie Natural History Museum in 1907.
Comparative size of Eoraptor

There were larger dinosaurs, but knowledge of them is based entirely on a small number of
fragmentary fossils. Most of the largest herbivorous specimens on record were all discovered in the 1970s or
later, and include the massive Argentinosaurus, which may have weighed 80000 to 100000 kilograms (90 to
110 short tons); some of the longest were the 33.5 meters (110 ft) long Diplodocus hallorum[49] (formerly
Seismosaurus) and the 33 meters (110 ft) long Supersaurus;[54] and the tallest, the 18 meters (59 ft) tall
Sauroposeidon, which could have reached a sixth-floor window. The longest of them all may have been
Amphicoelias fragillimus, known only from a now lost partial vertebral neural arch described in 1878.
Extrapolating from the illustration of this bone, the animal may have been 58 meters (190 ft) long and
weighed over 120000 kg (260000 lb).[49] The largest known carnivorous dinosaur was Spinosaurus,
reaching a length of 16 to 18 meters (50 to 60 ft), and weighing in at 8150 kg (18000 lb).[55] Other large
meat-eaters included Giganotosaurus, Carcharodontosaurus and Tyrannosaurus.[56]
Not including modern birds, the smallest dinosaurs known were about the size of a pigeon.[57] The
theropods Anchiornis and Epidexipteryx both had a total skeletal length of under 35 centimeters (1.1 ft).[57]
[58] Anchiornis is currently the smallest dinosaur described from an adult specimen, with an estimated weight
of 110 grams.[58] The smallest herbivorous dinosaurs included Microceratus and Wannanosaurus, at about
60 cm (2 ft) long each.[59][60]
Behavior
A nesting ground of Maiasaura was discovered in 1978

Interpretations of dinosaur behavior are generally based on the pose of body fossils and their habitat,
computer simulations of their biomechanics, and comparisons with modern animals in similar ecological
niches. As such, the current understanding of dinosaur behavior relies on speculation, and will likely remain
controversial for the foreseeable future. However, there is general agreement that some behaviors which are
common in crocodiles and birds, dinosaurs' closest living relatives, were also common among dinosaurs.
The first potential evidence of herding behavior was the 1878 discovery of 31 Iguanodon dinosaurs
which were then thought to have perished together in Bernissart, Belgium, after they fell into a deep, flooded
sinkhole and drowned.[61] Other mass-death sites have been subsequently discovered. Those, along with
multiple trackways, suggest that gregarious behavior was common in many dinosaur species. Trackways of
hundreds or even thousands of herbivores indicate that duck-bills (hadrosaurids) may have moved in great
herds, like the American Bison or the African Springbok. Sauropod tracks document that these animals
traveled in groups composed of several different species, at least in Oxfordshire, England,[62] although there
is not evidence for specific herd structures.[63] Dinosaurs may have congregated in herds for defense, for
migratory purposes, or to provide protection for their young. There is evidence that many types of dinosaurs,
including various theropods, sauropods, ankylosaurians, ornithopods, and ceratopsians, formed
aggregations of immature individuals. One example is a site in Inner Mongolia that has yielded the remains of
over twenty Sinornithomimus, from one to seven years old. This assemblage is interpreted as a social group
that was trapped in mud.[64] The interpretation of dinosaurs as gregarious has also extended to depicting
carnivorous theropods as pack hunters working together to bring down large prey.[65][66] However, this
lifestyle is uncommon among the modern relatives of dinosaurs (crocodiles and other reptiles, and birds –
Harris's Hawk is a well-documented exception), and the taphonomic evidence suggesting pack hunting in
such theropods as Deinonychus and Allosaurus can also be interpreted as the results of fatal disputes
between feeding animals, as is seen in many modern diapsid predators.[67]
Fossilized egg of the oviraptorid Citipati, American Museum of Natural History
Jack Horner's 1978 discovery of a Maiasaura ("good mother dinosaur") nesting ground in Montana
demonstrated that parental care continued long after birth among the ornithopods.[68] There is also evidence
that other Cretaceous-era dinosaurs, like Patagonian titanosaurian sauropods (1997 discovery), also nested
in large groups.[69] The Mongolian oviraptorid Citipati was discovered in a chicken-like brooding position in
1993, which may mean it was covered with an insulating layer of feathers that kept the eggs warm.[70]
Parental care is also implied by other finds. For example, the fossilized remains of a grouping of
Psittacosaurus has been found, consisting of one adult and 34 juveniles; in this case, the large number of
juveniles may be due to communal nesting.[71] Additionally, a dinosaur embryo (pertaining to the
prosauropod Massospondylus) was found without teeth, indicating that some parental care was required to
feed the young dinosaur.[72] Trackways have also confirmed parental behavior among ornithopods from the
Isle of Skye in northwestern Scotland.[73] Nests and eggs have been found for most major groups of
dinosaurs, and it appears likely that dinosaurs communicated with their young, in a manner similar to modern
birds and crocodiles.
Artist's rendering of two Centrosaurus, herbivorous ceratopsid dinosaurs from the late Cretaceous
fauna of North America
The crests and frills of some dinosaurs, like the marginocephalians, theropods and lambeosaurines,
may have been too fragile to be used for active defense, and so they were likely used for sexual or
aggressive displays, though little is known about dinosaur mating and territorialism. Head wounds from bites
suggest that theropods, at least, engaged in active aggressive confrontations.[74]
From a behavioral standpoint, one of the most valuable dinosaur fossils was discovered in the Gobi
Desert in 1971. It included a Velociraptor attacking a Protoceratops,[75] providing evidence that dinosaurs
did indeed attack each other.[76] Additional evidence for attacking live prey is the partially healed tail of an
Edmontosaurus, a hadrosaurid dinosaur; the tail is damaged in such a way that shows the animal was bitten
by a tyrannosaur but survived.[76] Cannibalism amongst some species of dinosaurs was confirmed by tooth
marks found in Madagascar in 2003, involving the theropod Majungasaurus.[77]
Based on current fossil evidence from dinosaurs such as Oryctodromeus, some herbivorous species
seem to have led a partially fossorial (burrowing) lifestyle,[78] and some bird-like species may have been
arboreal (tree-climbing), most notably primitive dromaeosaurids such as Microraptor[79] and the enigmatic
scansoriopterygids.[80] However, most dinosaurs seem to have relied on land-based locomotion. A good
understanding of how dinosaurs moved on the ground is key to models of dinosaur behavior; the science of
biomechanics, in particular, has provided significant insight in this area. For example, studies of the forces
exerted by muscles and gravity on dinosaurs' skeletal structure have investigated how fast dinosaurs could
run,[81] whether diplodocids could create sonic booms via whip-like tail snapping,[82] and whether
sauropods could float.[83]
Communication and vocalization

The nature of dinosaur communication remains enigmatic, and is an active area of research. In 2008,
paleontologist Phil Senter examined the evidence for vocalization in Mesozoic animal life, including
dinosaurs.[84] Senter found that, contrary to popular depictions of roaring dinosaurs in motion pictures, it is
likely that most dinosaurs were not capable of creating any vocalizations. To draw this conclusion, Senter
studied the distribution of vocal organs in reptiles and birds. He found that vocal chords in the larynx probably
evolved multiple times among reptiles, including crocodilians, which are able to produce guttural roars. Birds,
on the other hand, lack a larynx. Instead, bird calls are produced by the syrinx, a vocal organ found only in
birds, and which is not related to the larynx, meaning it evolved independently from the vocal organs in
reptiles. The syrinx depends on the air sac system in birds to function; specifically, it requires the presence of
a clavicular air sac near the wishbone or collar bone. This air sac leaves distinctive marks or opening on the
bones, including a distinct opening in the upper arm bone ( humerus). While many dinosaurs show evidence
of extensive air sac systems, almost none possess the clavicular air sac necessary to vocalize (one
exception, Aerosteon, probably evolved its clavicular air sac independently of birds for reasons other than
vocalization).[84]
The most primitive animals with evidence of a vocalizing syrinx are the enantironithine birds. Any
bird-line archosaurs more primitive than this probably did not make vocal calls. Rather, several lines of
evidence suggest that dinosaurs used primarily visual communication, in the form of distinctive-looking (and
possibly brightly colored) horns, frills, crests, sails and feathers. This is similar to some modern reptile groups
such as lizards, in which many forms are largely silent (though like dinosaurs they possess well-developed
senses of hearing) but use complex coloration and display behaviors to communicate.[84]
Also, though they may not have been able to vocalize, some dinosaurs may have used other
methods of producing sound for communication. Modern animals, including reptiles and birds, use a wide
variety of non-vocal sound communication, including hissing, jaw grinding or clapping, use of environment
(such as splashing), and wing beating (which would have been possible in winged maniraptoran dinosaurs).
[84]
Some studies have suggested that the hollow crests of the lambeosaurines may have functioned as
resonance chambers used for a wide range of vocalizations.[85][86] However, Senter (2008) noted that such
chambers are also used in modern non-vocal animals to accentuate or deepen non-vocal sounds like
hissing. For example, many snakes, which lack vocal chords, have resonating chambers in the skull.[84]
Physiology
Main article: Physiology of dinosaurs
Tyrannosaurus rex skull and upper vertebral column, Palais de la Découverte, Paris
A vigorous debate on the subject of temperature regulation in dinosaurs has been ongoing since the
1960s. Originally, scientists broadly disagreed as to whether dinosaurs were capable of regulating their body
temperatures at all. More recently, dinosaur endothermy has become the consensus view, and debate has
focused on the mechanisms of temperature regulation.
After dinosaurs were discovered, paleontologists first posited that they were ectothermic creatures:
"terrible lizards" as their name suggests. This supposed cold-bloodedness implied that dinosaurs were
relatively slow, sluggish organisms, comparable to modern reptiles, which need external sources of heat in
order to regulate their body temperature. Dinosaur ectothermy remained a prevalent view until Robert T.
"Bob" Bakker, an early proponent of dinosaur endothermy, published an influential paper on the topic in
1968.
Modern evidence indicates that dinosaurs thrived in cooler temperate climates, and that at least
some dinosaur species must have regulated their body temperature by internal biological means (perhaps
aided by the animals' bulk). Evidence of endothermy in dinosaurs includes the discovery of polar dinosaurs in
Australia and Antarctica (where they would have experienced a cold, dark six-month winter), the discovery of
dinosaurs whose feathers may have provided regulatory insulation, and analysis of blood-vessel structures
within dinosaur bone that are typical of endotherms. Skeletal structures suggest that theropods and other
dinosaurs had active lifestyles better suited to an endothermic cardiovascular system, while sauropods
exhibit fewer endothermic characteristics. It is certainly possible that some dinosaurs were endothermic while
others were not. Scientific debate over the specifics continues.[87]
Eubrontes, a dinosaur footprint in the Lower Jurassic Moenave Formation at the St. George Dinosaur
Discovery Site at Johnson Farm, southwestern Utah
Complicating the debate is the fact that warm-bloodedness can emerge based on more than one
mechanism. Most discussions of dinosaur endothermy tend to compare them with average-sized birds or
mammals, which expend energy to elevate body temperature above that of the environment. Small birds and
mammals also possess insulation, such as fat, fur, or feathers, which slows down heat loss. However, large
mammals, such as elephants, face a different problem because of their relatively small ratio of surface area
to volume (Haldane's principle). This ratio compares the volume of an animal with the area of its skin: as an
animal gets bigger, its surface area increases more slowly than its volume. At a certain point, the amount of
heat radiated away through the skin drops below the amount of heat produced inside the body, forcing
animals to use additional methods to avoid overheating. In the case of elephants, they have little hair as
adults, have large ears which increase their surface area, and have behavioral adaptations as well (such as
using the trunk to spray water on themselves and mud-wallowing). These behaviors increase cooling through
evaporation.
Large dinosaurs would presumably have had to deal with similar issues; their body size suggest they
lost heat relatively slowly to the surrounding air, and so could have been what are called inertial
homeotherms, animals that are warmer than their environments through sheer size rather than through
special adaptations like those of birds or mammals. However, so far this theory fails to account for the
numerous dog- and goat-sized dinosaur species, or the young of larger species.
Modern computerized tomography (CT) scans of a dinosaur's chest cavity (conducted in 2000) found
the apparent remnants of a four-chambered heart, much like those found in today's mammals and birds.[88]
The idea is controversial within the scientific community, coming under fire for bad anatomical science[89] or
simply wishful thinking.[90] The question of how this find reflects on metabolic rate and dinosaur internal
anatomy may be moot, though, regardless of the object's identity: both modern crocodilians and birds, the
closest living relatives of dinosaurs, have four-chambered hearts (albeit modified in crocodilians), and so
dinosaurs probably had them as well.[91]
Soft tissue and DNA
One of the best examples of soft-tissue impressions in a fossil dinosaur was discovered in Petraroia,
Italy. The discovery was reported in 1998, and described the specimen of a small, very young coelurosaur,
Scipionyx samniticus. The fossil includes portions of the intestines, colon, liver, muscles, and windpipe of this
immature dinosaur.[46]
In the March 2005 issue of Science, the paleontologist Mary Higby Schweitzer and her team
announced the discovery of flexible material resembling actual soft tissue inside a 68-million-year-old
Tyrannosaurus rex leg bone from the Hell Creek Formation in Montana. After recovery, the tissue was
rehydrated by the science team.[47]
When the fossilized bone was treated over several weeks to remove mineral content from the
fossilized bone-marrow cavity (a process called demineralization), Schweitzer found evidence of intact
structures such as blood vessels, bone matrix, and connective tissue (bone fibers). Scrutiny under the
microscope further revealed that the putative dinosaur soft tissue had retained fine structures
(microstructures) even at the cellular level. The exact nature and composition of this material, and the
implications of Schweitzer's discovery, are not yet clear; study and interpretation of the material is ongoing.
[47]
Newer research, published in PloS One (30 July 2008), has challenged the claims that the material
found is the soft tissue of Tyrannosaurus. Thomas Kaye of the University of Washington and his co-authors
contend that what was really inside the tyrannosaur bone was slimy biofilm created by bacteria that coated
the voids once occupied by blood vessels and cells.[92] The researchers found that what previously had
been identified as remnants of blood cells, because of the presence of iron, were actually framboids,
microscopic mineral spheres bearing iron. They found similar spheres in a variety of other fossils from
various periods, including an ammonite. In the ammonite they found the spheres in a place where the iron
they contain could not have had any relationship to the presence of blood.[93]
The successful extraction of ancient DNA from dinosaur fossils has been reported on two separate
occasions, but, upon further inspection and peer review, neither of these reports could be confirmed.[94]
However, a functional visual peptide of a theoretical dinosaur has been inferred using analytical phylogenetic
reconstruction methods on gene sequences of related modern species such as reptiles and birds.[95] In
addition, several proteins, including hemoglobin,[96] have putatively been detected in dinosaur fossils.[97]
Feathers and the origin of birds

Main article: Origin of birds
The possibility that dinosaurs were the ancestors of birds was first suggested in 1868 by Thomas
Henry Huxley.[98] After the work of Gerhard Heilmann in the early 20th century, the theory of birds as
dinosaur descendants was abandoned in favor of the idea of their being descendants of generalized
thecodonts, with the key piece of evidence being the supposed lack of clavicles in dinosaurs.[99] However,
as later discoveries showed, clavicles (or a single fused wishbone, which derived from separate clavicles)
were not actually absent;[14] they had been found as early as 1924 in Oviraptor, but misidentified as an
interclavicle.[100] In the 1970s, John Ostrom revived the dinosaur–bird theory,[101] which gained momentum
in the coming decades with the advent of cladistic analysis,[102] and a great increase in the discovery of
small theropods and early birds.[21] Of particular note have been the fossils of the Yixian Formation, where a
variety of theropods and early birds have been found, often with feathers of some type.[14] Birds share over
a hundred distinct anatomical features with theropod dinosaurs, which are now generally accepted to have
been their closest ancient relatives.[103] They are most closely allied with maniraptoran coelurosaurs.[14] A
minority of scientists, most notably Alan Feduccia and Larry Martin, have proposed other evolutionary paths,
including revised versions of Heilmann's basal archosaur proposal,[104] or that maniraptoran theropods are
the ancestors of birds but themselves are not dinosaurs, only convergent with dinosaurs.[105]
Feathers
Main article: Feathered dinosaurs
The famous Berlin Specimen of Archaeopteryx lithographica
Archaeopteryx, the first good example of a "feathered dinosaur", was discovered in 1861. The initial
specimen was found in the Solnhofen limestone in southern Germany, which is a lagerstätte, a rare and
remarkable geological formation known for its superbly detailed fossils. Archaeopteryx is a transitional fossil,
with features clearly intermediate between those of modern reptiles and birds. Brought to light just two years
after Darwin's seminal The Origin of Species, its discovery spurred the nascent debate between proponents
of evolutionary biology and creationism. This early bird is so dinosaur-like that, without a clear impression of
feathers in the surrounding rock, at least one specimen was mistaken for Compsognathus.[106]
Since the 1990s, a number of additional feathered dinosaurs have been found, providing even
stronger evidence of the close relationship between dinosaurs and modern birds. Most of these specimens
were unearthed in the lagerstätte of the Yixian Formation, Liaoning, northeastern China, which was part of an
island continent during the Cretaceous. Though feathers have been found in only a few locations, it is
possible that non-avian dinosaurs elsewhere in the world were also feathered. The lack of widespread fossil
evidence for feathered non-avian dinosaurs may be because delicate features like skin and feathers are not
often preserved by fossilization and thus are absent from the fossil record. To this point, protofeathers (thin,
filament-like structures) are known from dinosaurs at the base of Coelurosauria, such as compsognathids like
Sinosauropteryx and tyrannosauroids (Dilong),[107] but barbed feathers are known only among the
coelurosaur subgroup Maniraptora, which includes oviraptorosaurs, troodontids, dromaeosaurids, and birds.
[14][108] The description of feathered dinosaurs has not been without controversy; perhaps the most vocal
critics have been Alan Feduccia and Theagarten Lingham-Soliar, who have proposed that protofeathers are
the result of the decomposition of collagenous fiber that underlaid the dinosaurs' integument,[109][110][111]
and that maniraptoran dinosaurs with barbed feathers were not actually dinosaurs, but convergent with
dinosaurs.[105][110] However, their views have for the most part not been accepted by other researchers, to
the point that the question of the scientific nature of Feduccia's proposals has been raised.[112]
Skeleton
Because feathers are often associated with birds, feathered dinosaurs are often touted as the
missing link between birds and dinosaurs. However, the multiple skeletal features also shared by the two
groups represent another important line of evidence for paleontologists. Areas of the skeleton with important
similarities include the neck, pubis, wrist (semi-lunate carpal), arm and pectoral girdle, furcula (wishbone),
and breast bone. Comparison of bird and dinosaur skeletons through cladistic analysis strengthens the case
for the link.
Soft anatomy
Pneumatopores on the left ilium of Aerosteon riocoloradensis

Large meat-eating dinosaurs had a complex system of air sacs similar to those found in modern
birds, according to an investigation which was led by Patrick O'Connor of Ohio University. The lungs of
theropod dinosaurs (carnivores that walked on two legs and had bird-like feet) likely pumped air into hollow
sacs in their skeletons, as is the case in birds. "What was once formally considered unique to birds was
present in some form in the ancestors of birds", O'Connor said.[113] In a 2008 paper published in the online
journal PLoS ONE, scientists described Aerosteon riocoloradensis, the skeleton of which supplies the
strongest evidence to date of a dinosaur with a bird-like breathing system. CT-scanning revealed the
evidence of air sacs within the body cavity of the Aerosteon skeleton.[114][115]
Another piece of evidence that birds and dinosaurs are closely related is the use by both of gizzard
stones. These stones are swallowed by animals to aid digestion and break down food and hard fibers once
they enter the stomach. When found in association with fossils, gizzard stones are called gastroliths.[116]
Reproductive biology
A discovery of features in a Tyrannosaurus rex skeleton recently provided more evidence that
dinosaurs and birds evolved from a common ancestor and, for the first time, allowed paleontologists to
establish the sex of a dinosaur. When laying eggs, female birds grow a special type of bone between the
hard outer bone and the marrow of their limbs. This medullary bone, which is rich in calcium, is used to make
eggshells. The presence of endosteally derived bone tissues lining the interior marrow cavities of portions of
the Tyrannosaurus rex specimen's hind limb suggested that T. rex used similar reproductive strategies, and
revealed the specimen to be female.[117] Further research has found medullary bone in the theropod
Allosaurus and the ornithopod Tenontosaurus. Because the line of dinosaurs that includes Allosaurus and
Tyrannosaurus diverged from the line that led to Tenontosaurus very early in the evolution of dinosaurs, this
suggests that dinosaurs in general produced medullary tissue. Medullary bone has been found in specimens
of sub-adult size, which suggests that dinosaurs reached sexual maturity rather quickly for such large
animals.[118]
Behavioral evidence
A recently discovered troodont fossil demonstrates that some dinosaurs slept with their heads tucked
under their arms.[119] This behavior, which may have helped to keep the head warm, is also characteristic of
modern birds.
Extinction
Main articles: Cretaceous–Tertiary extinction event and K–T boundary
Non-avian dinosaurs suddenly became extinct approximately 65 million years ago. Many other
groups of animals also became extinct at this time, including ammonites (nautilus-like mollusks), mosasaurs,
plesiosaurs, pterosaurs, most birds, and many groups of mammals.[6] This mass extinction is known as the
Cretaceous–Tertiary extinction event. The nature of the event that caused this mass extinction has been
extensively studied since the 1970s; at present, several related theories are supported by paleontologists.
Though the consensus is that an impact event was the primary cause of dinosaur extinction, some scientists
cite other possible causes, or support the idea that a confluence of several factors was responsible for the
sudden disappearance of dinosaurs from the fossil record.
At the peak of the Mesozoic, there were no polar ice caps, and sea levels are estimated to have been
from 100 to 250 meters (300 to 800 ft) higher than they are today. The planet's temperature was also much
more uniform, with only 25 °C (45 °F) separating average polar temperatures from those at the equator. On
average, atmospheric temperatures were also much higher; the poles, for example, were 50 °C (90 °F)
warmer than today.[120][121]
The atmosphere's composition during the Mesozoic was vastly different as well. Carbon dioxide
levels were up to 12 times higher than today's levels, and oxygen formed 32 to 35% of the
atmosphere[citation needed], as compared to 21% today. However, by the late Cretaceous, the environment
was changing dramatically. Volcanic activity was decreasing, which led to a cooling trend as levels of
atmospheric carbon dioxide dropped. Oxygen levels in the atmosphere also started to fluctuate and would
ultimately fall considerably. Some scientists hypothesize that climate change, combined with lower oxygen
levels, might have led directly to the demise of many species. If the dinosaurs had respiratory systems similar
to those commonly found in modern birds, it may have been particularly difficult for them to cope with
reduced respiratory efficiency, given the enormous oxygen demands of their very large bodies.[6]
Impact event
Main article: Chicxulub crater
The Chicxulub Crater at the tip of the Yucatán Peninsula; the impactor that formed this crater may
have caused the dinosaur extinction.
The asteroid collision theory, which was brought to wide attention in 1980 by Walter Alvarez and
colleagues, links the extinction event at the end of the Cretaceous period to a bolide impact approximately
65.5 million years ago. Alvarez et al. proposed that a sudden increase in iridium levels, recorded around the
world in the period's rock stratum, was direct evidence of the impact.[122] The bulk of the evidence now
suggests that a bolide 5 to 15 kilometers (3 to 9 mi) wide hit in the vicinity of the Yucatán Peninsula, creating
the approximately 180 km (110 mi) Chicxulub Crater and triggering the mass extinction.[123][124] Scientists
are not certain whether dinosaurs were thriving or declining before the impact event. Some scientists
propose that the meteorite caused a long and unnatural drop in Earth's atmospheric temperature, while
others claim that it would have instead created an unusual heat wave.
Although the speed of extinction cannot be deduced from the fossil record alone, various models
suggest that the extinction was extremely rapid. The consensus among scientists who support this theory is
that the impact caused extinctions both directly (by heat from the meteorite impact) and also indirectly (via a
worldwide cooling brought about when matter ejected from the impact crater reflected thermal radiation from
the sun).
In September 2007, U.S. researchers led by William Bottke of the Southwest Research Institute in
Boulder, Colorado, and Czech scientists used computer simulations to identify the probable source of the
Chicxulub impact. They calculated a 90% probability that a giant asteroid named Baptistina, approximately
160 km (100 mi) in diameter, orbiting in the asteroid belt which lies between Mars and Jupiter, was struck by
a smaller unnamed asteroid about 55 km (35 mi) in diameter about 160 million years ago. The impact
shattered Baptistina, creating a cluster which still exists today as the Baptistina family. Calculations indicate
that some of the fragments were sent hurtling into earth-crossing orbits, one of which was the 10 km (6 mi)
wide meteorite which struck Mexico's Yucatan peninsula 65 million years ago, creating the Chicxulub crater.
[125]
A similar but more controversial explanation proposes that "passages of the [hypothetical] solar
companion star Nemesis through the Oort comet cloud would trigger comet showers."[126] One or more of
these comets then collided with the Earth at approximately the same time, causing the worldwide extinction.
As with the impact of a single asteroid, the end result of this comet bombardment would have been a sudden
drop in global temperatures, followed by a protracted cool period.[126]
Deccan Traps
Main article: Deccan Traps
Before 2000, arguments that the Deccan Traps flood basalts caused the extinction were usually
linked to the view that the extinction was gradual, as the flood basalt events were thought to have started
around 68 million years ago and lasted for over 2 million years. However, there is evidence that two-thirds of
the Deccan Traps were created in only 1 million years about 65.5 million years ago, and so these eruptions
would have caused a fairly rapid extinction, possibly over a period of thousands of years, but still longer than
would be expected from a single impact event.[127][128]
The Deccan Traps could have caused extinction through several mechanisms, including the release
into the air of dust and sulphuric aerosols, which might have blocked sunlight and thereby reduced
photosynthesis in plants. In addition, Deccan Trap volcanism might have resulted in carbon dioxide
emissions, which would have increased the greenhouse effect when the dust and aerosols cleared from the
atmosphere.[128] Before the mass extinction of the dinosaurs, the release of volcanic gases during the
formation of the Deccan Traps "contributed to an apparently massive global warming. Some data point to an
average rise in temperature of 8 °C (14 °F) in the last half million years before the impact [at
Chicxulub]."[127][128]
In the years when the Deccan Traps theory was linked to a slower extinction, Luis Alvarez (who died
in 1988) replied that paleontologists were being misled by sparse data. While his assertion was not initially
well-received, later intensive field studies of fossil beds lent weight to his claim. Eventually, most
paleontologists began to accept the idea that the mass extinctions at the end of the Cretaceous were largely
or at least partly due to a massive Earth impact. However, even Walter Alvarez has acknowledged that there
were other major changes on Earth even before the impact, such as a drop in sea level and massive volcanic
eruptions that produced the Indian Deccan Traps, and these may have contributed to the extinctions.[129]
Failure to adapt to changing conditions
Lloyd et al. (2008) noted that, in the Mid Cretaceous, the flowering, angiosperm plants became a
major part of terrestrial ecosystems, which had previously been dominated by gymnosperms such as
conifers. Dinosaur coprolite–fossilized dung–indicate that, while some ate angiosperms, most herbivorous
dinosaurs ate mainly gymnosperms. Statistical analysis by Lloyd et al. concluded that, contrary to earlier
studies, dinosaurs did not diversify very much in the Late Cretaceous. Lloyd et al. suggested that dinosaurs'
failure to diversify as ecosystems were changing doomed them to extinction.[37]
Possible Paleocene survivors

Main article: Paleocene dinosaurs
Non-avian dinosaur remains are occasionally found above the K–T boundary. In 2001,
paleontologists Zielinski and Budahn reported the discovery of a single hadrosaur leg-bone fossil in the San
Juan Basin, New Mexico, and described it as evidence of Paleocene dinosaurs. The formation in which the
bone was discovered has been dated to the early Paleocene epoch, approximately 64.5 million years ago. If
the bone was not re-deposited into that stratum by weathering action, it would provide evidence that some
dinosaur populations may have survived at least a half million years into the Cenozoic Era.[130] Other
evidence includes the finding of dinosaur remains in the Hell Creek Formation up to 1.3 meters (51 in) above
(40000 years later than) the K–T boundary. Similar reports have come from other parts of the world, including
China.[131] Many scientists, however, dismissed the supposed Paleocene dinosaurs as re-worked, i.e.
washed out of their original locations and then re-buried in much later sediments.[132][133] However, direct
dating of the bones themselves has supported the later date, with U-Pb dating methods resulting in a precise
age of 64.8 ± 0.9 million years ago.[134] If correct, the presence of a handful of dinosaurs in the early
Paleocene would not change the underlying facts of the extinction.[132]
History of discovery
Dinosaur fossils have been known for millennia, although their true nature was not recognized. The
Chinese, whose modern word for dinosaur is konglong (恐龍, or "terrible dragon"), considered them to be
dragon bones and documented them as such. For example, Hua Yang Guo Zhi, a book written by Zhang Qu
during the Western Jin Dynasty, reported the discovery of dragon bones at Wucheng in Sichuan Province.
[135] Villagers in central China have long unearthed fossilized "dragon bones" for use in traditional
medicines, a practice that continues today.[136] In Europe, dinosaur fossils were generally believed to be the
remains of giants and other creatures killed by the Great Flood.
Scholarly descriptions of what would now be recognized as dinosaur bones first appeared in the late
17th century in England. Part of a bone, now known to have been the femur of a Megalosaurus,[137] was
recovered from a limestone quarry at Cornwell near Chipping Norton, Oxfordshire, England, in 1676. The
fragment was sent to Robert Plot, Professor of Chemistry at the University of Oxford and first curator of the
Ashmolean Museum, who published a description in his Natural History of Oxfordshire in 1677. He correctly
identified the bone as the lower extremity of the femur of a large animal, and recognized that it was too large
to belong to any known species. He therefore concluded it to be the thigh bone of a giant human similar to
those mentioned in the Bible. In 1699, Edward Lhuyd, a friend of Sir Isaac Newton, was responsible for the
first published scientific treatment of what would now be recognized as a dinosaur when he described and
named a sauropod tooth, "Rutellum implicatum",[138][139] that had been found in Caswell, near Witney,
Oxfordshire.[140]
William Buckland
Between 1815 and 1824, the Rev William Buckland, a professor of geology at Oxford University,
collected more fossilized bones of Megalosaurus and became the first person to describe a dinosaur in a
scientific journal.[137][141] The second dinosaur genus to be identified, Iguanodon, was discovered in 1822
by Mary Ann Mantell – the wife of English geologist Gideon Mantell. Gideon Mantell recognized similarities
between his fossils and the bones of modern iguanas. He published his findings in 1825.[142][143]
The study of these "great fossil lizards" soon became of great interest to European and American
scientists, and in 1842 the English paleontologist Richard Owen coined the term "dinosaur". He recognized
that the remains that had been found so far, Iguanodon, Megalosaurus and Hylaeosaurus, shared a number
of distinctive features, and so decided to present them as a distinct taxonomic group. With the backing of
Prince Albert of Saxe-Coburg-Gotha, the husband of Queen Victoria, Owen established the Natural History
Museum in South Kensington, London, to display the national collection of dinosaur fossils and other
biological and geological exhibits.
In 1858, the first known American dinosaur was discovered, in marl pits in the small town of
Haddonfield, New Jersey (although fossils had been found before, their nature had not been correctly
discerned). The creature was named Hadrosaurus foulkii. It was an extremely important find: Hadrosaurus
was one of the first nearly complete dinosaur skeletons found (the first was in 1834, in Maidstone, Kent,
England), and it was clearly a bipedal creature. This was a revolutionary discovery as, until that point, most
scientists had believed dinosaurs walked on four feet, like other lizards. Foulke's discoveries sparked a wave
of dinosaur mania in the United States.
Othniel Charles Marsh, 19th century photograph

Edward Drinker Cope, 19th century photograph
Dinosaur mania was exemplified by the fierce rivalry between Edward Drinker Cope and Othniel
Charles Marsh, both of whom raced to be the first to find new dinosaurs in what came to be known as the
Bone Wars. The feud probably originated when Marsh publicly pointed out that Cope's reconstruction of an
Elasmosaurus skeleton was flawed: Cope had inadvertently placed the plesiosaur's head at what should
have been the animal's tail end. The fight between the two scientists lasted for over 30 years, ending in 1897
when Cope died after spending his entire fortune on the dinosaur hunt. Marsh 'won' the contest primarily
because he was better funded through a relationship with the US Geological Survey. Unfortunately, many
valuable dinosaur specimens were damaged or destroyed due to the pair's rough methods: for example, their
diggers often used dynamite to unearth bones (a method modern paleontologists would find appalling).
Despite their unrefined methods, the contributions of Cope and Marsh to paleontology were vast: Marsh
unearthed 86 new species of dinosaur and Cope discovered 56, a total of 142 new species. Cope's collection
is now at the American Museum of Natural History in New York, while Marsh's is on display at the Peabody
Museum of Natural History at Yale University.[144]
After 1897, the search for dinosaur fossils extended to every continent, including Antarctica. The first
Antarctic dinosaur to be discovered, the ankylosaurid Antarctopelta oliveroi, was found on Ross Island in
1986, although it was 1994 before an Antarctic species, the theropod Cryolophosaurus ellioti, was formally
named and described in a scientific journal.
Current dinosaur "hot spots" include southern South America (especially Argentina) and China.
China in particular has produced many exceptional feathered dinosaur specimens due to the unique geology
of its dinosaur beds, as well as an ancient arid climate particularly conducive to fossilization.
The "dinosaur renaissance"

Main article: Dinosaur renaissance
The field of dinosaur research has enjoyed a surge in activity that began in the 1970s and is ongoing.
This was triggered, in part, by John Ostrom's discovery of Deinonychus, an active predator that may have
been warm-blooded, in marked contrast to the then-prevailing image of dinosaurs as sluggish and cold-
blooded. Vertebrate paleontology has become a global science. Major new dinosaur discoveries have been
made by paleontologists working in previously unexploited regions, including India, South America,
Madagascar, Antarctica, and most significantly China (the amazingly well-preserved feathered dinosaurs in
China have further consolidated the link between dinosaurs and their conjectured living descendants,
modern birds). The widespread application of cladistics, which rigorously analyzes the relationships between
biological organisms, has also proved tremendously useful in classifying dinosaurs. Cladistic analysis,
among other modern techniques, helps to compensate for an often incomplete and fragmentary fossil record.
Cultural depictions
Main article: Cultural depictions of dinosaurs
By human standards, dinosaurs were creatures of fantastic appearance and often enormous size. As
such, they have captured the popular imagination and become an enduring part of human culture. Entry of
the word "dinosaur" into the common vernacular reflects the animals' cultural importance: in English,
"dinosaur" is commonly used to describe anything that is impractically large, slow-moving, obsolete, or bound
for extinction.[7]
Public enthusiasm for dinosaurs first developed in Victorian England, where in 1854, three decades
after the first scientific descriptions of dinosaur remains, the famous dinosaur sculptures were unveiled in
London's Crystal Palace Park. The Crystal Palace dinosaurs proved so popular that a strong market in
smaller replicas soon developed. In subsequent decades, dinosaur exhibits opened at parks and museums
around the world, ensuring that successive generations would be introduced to the animals in an immersive
and exciting way.[145] Dinosaurs' enduring popularity, in its turn, has resulted in significant public funding for
dinosaur science, and has frequently spurred new discoveries. In the United States, for example, the
competition between museums for public attention led directly to the Bone Wars of the 1880s and 1890s,
during which a pair of feuding paleontologists made enormous scientific contributions.[146]
The popular preoccupation with dinosaurs has ensured their appearance in literature, film and other
media. Beginning in 1852 with a passing mention in Charles Dickens' Bleak House,[147] dinosaurs have
been featured in large numbers of fictional works. Sir Arthur Conan Doyle's 1912 book The Lost World, the
iconic 1933 film King Kong, 1954's Godzilla and its many sequels, the best-selling 1990 novel Jurassic Park
by Michael Crichton and its 1993 film adaptation are just a few notable examples of dinosaur appearances in
fiction. Authors of general-interest non-fictional works about dinosaurs, including some prominent
paleontologists, have often sought to use the animals as a way to educate readers about science in general.
Dinosaurs are ubiquitous in advertising; numerous companies have referenced dinosaurs in printed or
televised advertisements, either in order to sell their own products or in order to characterize their rivals as
slow-moving, dim-witted or obsolete.[148]
See also
Wikimedia Commons has media related to: Dinosauria
Dinosaurs portal

• List of dinosaurs
• List of dinosaur-bearing rock formations
• Physiology of dinosaurs
• Prehistoric reptile
Notes and references

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126.^ a b edited by Christian Koeberl and Kenneth G. MacLeod. (2002). Catastrophic Events
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127.^ a b Hofman, C, Féraud, G & Courtillot, V (2000). "40Ar/39Ar dating of mineral separates
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X.
128.^ a b c Duncan, RA & Pyle, DG (1988). "Rapid eruption of the Deccan flood basalts at the
Cretaceous/Tertiary boundary". Nature 333: 841–843. doi:10.1038/333841a0.
129.^ Alvarez, W (1997). T. rex and the Crater of Doom. Princeton University Press. pp. 130–
146. ISBN 978-0691016306.
130.^ Fassett, JE, Lucas, SG, Zielinski, RA, and Budahn, JR (2001). "Compelling new evidence
for Paleocene dinosaurs in the Ojo Alamo Sandstone, San Juan Basin, New Mexico and Colorado,
USA" (PDF). Catastrophic events and mass extinctions, Lunar and Planetary Contribution 1053: 45–
46. http://www.lpi.usra.edu/meetings/impact2000/pdf/3139.pdf. Retrieved 2007-05-18.
131.^ Sloan, R. E., Rigby, K,. Van Valen, L. M., Gabriel, Diane (1986). "Gradual dinosaur
extinction and simultaneous ungulate radiation in the Hell Creek Formation". Science 232 (4750):
629–633. doi:10.1126/science.232.4750.629. PMID 17781415.
132.^ a b Fastovsky, David E.; Sheehan, Peter M. (2005). "Reply to comment on "The Extinction
of the dinosaurs in North America"" (PDF). GSA Today 15: 11. doi:10.1130/1052-
5173(2005)015[11b:RTEOTD]2.0.CO;2. http://www.geosociety.org/gsatoday/archive/15/7/pdf/i1052-
5173-15-7-11b.pdf.
133.^ Sullivan, RM (2003). "No Paleocene dinosaurs in the San Juan Basin, New Mexico".
Geological Society of America Abstracts with Programs 35 (5): 15.
http://gsa.confex.com/gsa/2003RM/finalprogram/abstract_47695.htm. Retrieved 2007-07-02.
134.^ Fassett, J.E., Heaman, L.M., and A. Simonetti. (2011). "Direct U-Pb dating of Cretaceous
and Paleocene dinosaur bones, San Juan Basin, New Mexico." Geology, 39:159-162.
doi:10.1130/G31466.1
135.^ Dong Zhiming (1992). Dinosaurian Faunas of China. China Ocean Press, Beijing. ISBN 3-
540-52084-8. OCLC 26522845.
136.^ "Dinosaur bones 'used as medicine'". BBC News. 2007-07-06.
http://news.bbc.co.uk/2/hi/asia-pacific/6276948.stm. Retrieved 2007-07-06.
137.^ a b Sarjeant, William A.S. (1997). "The earliert discoveries". In Farlow, James O.; and
Brett-Surman, Michael K. (eds.). The Complete Dinosaur. Bloomington: Indiana University Press.
pp. 3–11. ISBN 0-253-33349-0.
138.^ Lhuyd, E. (1699). Lithophylacii Britannici Ichnographia, sive lapidium aliorumque fossilium
Britannicorum singulari figura insignium. Gleditsch and Weidmann:London.
139.^ Delair, J.B.; Sarjeant, W.A.S. (2002). "The earliest discoveries of dinosaurs: the records
re-examined". Proceedings of the Geologists' Association 113: 185–197.
140.^ Gunther, R.T. (1945). Early Science in Oxford: Life and Letters of Edward Lhuyd , volume
14. Author:Oxford.
141.^ Buckland, W. (1824). "Notice on the Megalosaurus or great Fossil Lizard of Stonesfield."
Transactions of the Geological Society of London, series 2, vol. 1: 390–396.
142.^ Mantell, Gideon A. (1825). "Notice on the Iguanodon, a newly discovered fossil reptile,
from the sandstone of Tilgate forest, in Sussex.". Philosophical Transactions of the Royal Society
115: 179–186. doi:10.1098/rstl.1825.0010. http://links.jstor.org/sici?sici=0261-
0523(1825)115%3C179%3ANOTIAN%3E2.0.CO%3B2-W. Retrieved 2007-02-21.
143.^ Sues, Hans-Dieter (1997). "European Dinosaur Hunters". In James Orville Farlow and M.
K. Brett-Surman (eds.). The Complete Dinosaur. Bloomington: Indiana University Press. p. 14.
ISBN 0-253-33349-0.
144.^ Holmes T (1996). Fossil Feud: The Bone Wars of Cope and Marsh, Pioneers in Dinosaur
Science. Silver Burdett Press. ISBN 978-0382391477. OCLC 34472600.
145.^ Torrens, H.S. (1993). "The dinosaurs and dinomania over 150 years". Modern Geology 18
(2): 257–286.
146.^ Breithaupt, Brent H. (1997). "First golden period in the USA". In Currie, Philip J. and
Padian, Kevin (eds.). The Encyclopedia of Dinosaurs. San Diego: Academic Press. pp. 347–350.
ISBN 978-0122268106.
147.^ "London. Michaelmas term lately over, and the Lord Chancellor sitting in Lincoln's Inn
Hall. Implacable November weather. As much mud in the streets, as if the waters had but newly
retired from the face of the earth, and it would not be wonderful to meet a Megalosaurus, forty feet
long or so, waddling like an elephantine lizard up Holborne Hill. " From page 1 of Dickens, Charles
J.H. (1852). Bleak House. London: Bradbury & Evans.
148.^ Glut, D.F., and Brett-Surman, M.K. (1997). Farlow, James O. and Brett-Surman, Michael
K. (eds.). ed. The Complete Dinosaur. Indiana University Press. pp. 675–697. ISBN 978-
0253213136.
Further reading
• Bakker, Robert T. (1986). The Dinosaur Heresies: New Theories Unlocking the Mystery of
the Dinosaurs and Their Extinction. New York: Morrow. ISBN 0688042872.
• Holtz, Thomas R. Jr. (2007). Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for
Dinosaur Lovers of All Ages. New York: Random House. ISBN 9780375824197.
• Paul, Gregory S. (2000). The Scientific American Book of Dinosaurs . New York: St. Martin's
Press. ISBN 0312262264.
• Paul, Gregory S. (2002). Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs
and Birds. Baltimore: The Johns Hopkins University Press. ISBN 0801867630.
External links
This audio file was created from a revision of Dinosaur dated 2005-12-30, and does not reflect subsequent edits to the article. ( Audio help)
General
• DinoDatabase.com | Hundreds of dinosaurs and dinosaur related topics.
Images
• The Science and Art of Gregory S. Paul Influential paleontologist's anatomy art and paintings
• Skeletal Drawing Professional restorations of numerous dinosaurs, and discussions of
dinosaur anatomy.
Video
• BBC Nature: Watch dinosaurs brought to life and get experts' interpretations with videos from
BBC programmes including Walking with Dinosaurs.
Popular
• Dinosaurs & other extinct creatures: From the Natural History Museum, a well illustrated
dinosaur directory.
• Dinosaurnews (www.dinosaurnews.org) The dinosaur-related headlines from around the
world. Recent news on dinosaurs, including finds and discoveries, and many links.
• Dinosauria From UC Berkeley Museum of Paleontology Detailed information – scroll down for
menu.
• LiveScience.com All about dinosaurs, with current featured articles.
• Zoom Dinosaurs (www.enchantedlearning.com) From Enchanted Learning. Kids' site, info
pages and stats, theories, history.
• Dinosaur genus list contains data tables on nearly every published dinosaur genus.
• LiveScience.com Giant Dinosaurs Get Downsized by LiveScience, June 21, 2009
Technical
• Palaeontologia Electronica From Coquina Press. Online technical journal.
• Dinobase A searchable dinosaur database, from the University of Bristol, with dinosaur lists,
classification, pictures, and more.
• DinoData (www.dinodata.org) Technical site, essays, classification, anatomy.
• Dinosauria On-Line (www.dinosauria.com) Technical site, essays, pronunciation, dictionary.
• Thescelosaurus! By Justin Tweet. Includes a cladogram and small essays on each relevant
genera and species.
• Dinosauromorpha Cladogram From Palaeos. A detailed amateur site about all things paleo.
[hide]v · d · eArchosauromorphs
Kingdom: Animalia · Phylum: Chordata · Class: Sauropsida · Subclass: Diapsida
Primitive Euparkeriidae • Erythrosuchidae • Proterochampsidae •

Archosauromorphs Proterosuchidae • Choristodera • Prolacertiformes • Rhynchosauria •
Trilophosauria • Vancleavea
Crurotarsi Ornithosuchidae • Aetosauria • Phytosauria • Rauisuchia •

Archosaurs Crocodylomorpha • Crocodilia
Avemetatarsalia and Scleromochlus • Pterosauria • Dinosauromorpha • Dinosauria •

Ornithodira Archosaurs Ornithischia • Saurischia • Aves
Avian Archosaurs Avialae • Archaeopteryx • Confuciusornis • Ichthyornis •

Enantiornithes • Hesperornithes • Neornithes • Palaeognathae • Neognathae
Retrieved from "http://en.wikipedia.org/wiki/Dinosaur"
Categories: Dinosaurs | Paleontology | Paleozoology | Greek loanwords

W000
Earth

This article is about the planet. For other uses, see Earth (disambiguation).
Earth
"The Blue Marble" photograph of Earth,
taken from Apollo 17
Designations
Pronunciation i /ˈɜrθ/
Adjective earthly, tellurian, telluric, terran, terrestrial.
Orbital characteristics
Epoch J2000.0[note 1]
152,098,232 km
Aphelion
1.01671388 AU[note 2]
147,098,290 km
Perihelion
0.98329134 AU[note 2]
149,598,261 km
Semi-major axis
1.00000261 AU[1]
Eccentricity 0.01671123[1]
365.256363004 days[2]
Orbital period
1.000017421 yr
29.78 km/s[3]
Average orbital speed
107,200 km/h
Mean anomaly 357.51716°[3]
7.155° to Sun's equator

Inclination
1.57869°[4] to invariable plane
Longitude of ascending node 348.73936°[3][note 3]
Argument of perihelion 114.20783°[3][note 4]

Satellites 1 (the Moon)
Physical characteristics
Mean radius 6,371.0 km[5]
Equatorial radius 6,378.1 km[6]
Polar radius 6,356.8 km[7]
Flattening 0.0033528[6]
40,075.16 km (equatorial)[8]
Circumference
40,008.00 km (meridional)[8]
510,072,000 km2[9][10][note 5]
Surface area 148,940,000 km2 land (29.2 %)
361,132,000 km2 water (70.8 %)
Volume 1.08321×1012 km3[3]

Mass 5.9736×1024 kg[3]
Mean density 5.515 g/cm3[3]
Equatorial surface gravity 9.780327 m/s2[11]

0.99732 g
Escape velocity 11.186 km/s[3]
Sidereal rotation 0.99726968 d[12]

period 23h 56m4.100s
Equatorial rotation velocity 1,674.4 km/h (465.1 m/s)[13]
Axial tilt 23°26'21".4119[2]
0.367 (geometric)[3]
Albedo
0.306 (Bond)[3]
Surface temp.
Kelvin
Celsius
min mean max
184 K[14 287.2 K[15 331 K[16
] ] ]
-89.2 °C 14 °C 57.8 °C
Atmosphere
Surface pressure 101.325 kPa (MSL)
78.08% nitrogen (N2)[3]

20.95% oxygen (O2)
Composition 0.93% argon
0.038% carbon dioxide
About 1% water vapor (varies with climate)
Earth (or the Earth) is the third planet from the Sun and the densest and fifth-largest of the eight
planets in the Solar System. It is also the largest of the Solar System's four terrestrial planets. It is sometimes
referred to as the World, the Blue Planet, or by its Latin name, Terra.[note 6][note 7]
Home to millions of species including humans, Earth is currently the only astronomical body where
life is known to exist.[17] The planet formed 4.54 billion years ago, and life appeared on its surface within a
billion years.[18] Earth's biosphere has significantly altered the atmosphere and other abiotic conditions on
the planet, enabling the proliferation of aerobic organisms as well as the formation of the ozone layer which,
together with Earth's magnetic field, blocks harmful solar radiation, permitting life on land.[19] The physical
properties of the Earth, as well as its geological history and orbit, have allowed life to persist during this
period. The planet is expected to continue supporting life for at least another 500 million years.[20][21]
Earth's outer surface is divided into several rigid segments, or tectonic plates, that migrate across the
surface over periods of many millions of years. About 71% of the surface is covered with salt water oceans,
the remainder consisting of continents and islands which together have many lakes and other sources of
water contributing to the hydrosphere. Liquid water, necessary for all known life, is not known to exist in
equilibrium on any other planet's surface.[note 8] Earth's poles are mostly covered with solid ice (Antarctic ice
sheet) or sea ice (Arctic ice cap). The planet's interior remains active, with a thick layer of relatively solid
mantle, a liquid outer core that generates a magnetic field, and a solid iron inner core.
Earth interacts with other objects in space, especially the Sun and the Moon. At present, Earth orbits
the Sun once for every roughly 366.26 times it rotates about its axis, which is equal to 365.26 solar days, or
one sidereal year.[note 9] The Earth's axis of rotation is tilted 23.4° away from the perpendicular to its orbital
plane, producing seasonal variations on the planet's surface with a period of one tropical year (365.24 solar
days).[22] Earth's only known natural satellite, the Moon, which began orbiting it about 4.53 billion years ago,
provides ocean tides, stabilizes the axial tilt and gradually slows the planet's rotation. Between approximately
3.8 billion and 4.1 billion years ago, numerous asteroid impacts during the Late Heavy Bombardment caused
significant changes to the greater surface environment.
Both the mineral resources of the planet, as well as the products of the biosphere, contribute
resources that are used to support a global human population. These inhabitants are grouped into about 200
independent sovereign states, which interact through diplomacy, travel, trade, and military action. Human
cultures have developed many views of the planet, including personification as a deity, a belief in a flat Earth
or in the Earth as the center of the universe, and a modern perspective of the world as an integrated
environment that requires stewardship.
Contents
[hide]
• 1 Chronology
• 1.1 Evolution of life
• 1.2 Future
• 2 Composition and structure
• 2.1 Shape
• 2.2 Chemical composition
• 2.3 Internal structure
• 2.4 Heat
• 2.5 Tectonic plates
• 2.6 Surface
• 2.7 Hydrosphere
• 2.8 Atmosphere
• 2.8.1 Weather and climate
• 2.8.2 Upper atmosphere
• 2.9 Magnetic field
• 3 Orbit and rotation
• 3.1 Rotation
• 3.2 Orbit
• 3.3 Axial tilt and seasons
• 4 Moon
Chronology
Main article: History of the Earth
See also: Geological history of Earth
Scientists have been able to reconstruct detailed information about the planet's past. The earliest
dated Solar System material was formed 4.5672 ± 0.0006 billion years ago,[23] and by 4.54 billion years ago
(within an uncertainty of 1%)[18] the Earth and the other planets in the Solar System had formed out of the
solar nebula—a disk-shaped mass of dust and gas left over from the formation of the Sun. This assembly of
the Earth through accretion was thus largely completed within 10–20 million years.[24] Initially molten, the
outer layer of the planet Earth cooled to form a solid crust when water began accumulating in the
atmosphere. The Moon formed shortly thereafter, 4.53 billion years ago.[25]
The current consensus model[26] for the formation of the Moon is the giant impact hypothesis, in
which the Moon was created when a Mars-sized object (sometimes called Theia) with about 10% of the
Earth's mass[27] impacted the Earth in a glancing blow.[28] In this model, some of this object's mass would
have merged with the Earth and a portion would have been ejected into space, but enough material would
have been sent into orbit to coalesce into the Moon.
Outgassing and volcanic activity produced the primordial atmosphere of the Earth. Condensing water
vapor, augmented by ice and liquid water delivered by asteroids and the larger proto-planets, comets, and
trans-Neptunian objects produced the oceans.[29] The newly formed Sun was only 70% of its present
luminosity, yet evidence shows that the early oceans remained liquid—a contradiction dubbed the faint young
Sun paradox. A combination of greenhouse gases and higher levels of solar activity served to raise the
Earth's surface temperature, preventing the oceans from freezing over.[30] By 3.5 billion years ago, the
Earth's magnetic field was established, which helped prevent the atmosphere from being stripped away by
the solar wind.[31]
Two major models have been proposed for the rate of continental growth:[32] steady growth to the
present-day[33] and rapid growth early in Earth history.[34] Current research shows that the second option is
most likely, with rapid initial growth of continental crust[35] followed by a long-term steady continental area.
[36][37][38] On time scales lasting hundreds of millions of years, the surface continually reshaped as
continents formed and broke up. The continents migrated across the surface, occasionally combining to form
a supercontinent. Roughly 750 million years ago (Ma), one of the earliest known supercontinents, Rodinia,
began to break apart. The continents later recombined to form Pannotia, 600–540 Ma, then finally Pangaea,
which broke apart 180 Ma.[39]
Evolution of life
At present, Earth provides the only example of an environment that has given rise to the evolution of
life.[40] Highly energetic chemistry is believed to have produced a self-replicating molecule around 4 billion
years ago and half a billion years later the last common ancestor of all life existed.[41] The development of
photosynthesis allowed the Sun's energy to be harvested directly by life forms; the resultant oxygen
accumulated in the atmosphere and formed a layer of ozone (a form of molecular oxygen [O3]) in the upper
atmosphere. The incorporation of smaller cells within larger ones resulted in the development of complex
cells called eukaryotes.[42] True multicellular organisms formed as cells within colonies became increasingly
specialized. Aided by the absorption of harmful ultraviolet radiation by the ozone layer, life colonized the
surface of Earth.[43]
Since the 1960s, it has been hypothesized that severe glacial action between 750 and 580 Ma,
during the Neoproterozoic, covered much of the planet in a sheet of ice. This hypothesis has been termed
"Snowball Earth", and is of particular interest because it preceded the Cambrian explosion, when
multicellular life forms began to proliferate.[44]
Following the Cambrian explosion, about 535 Ma, there have been five major mass extinctions.[45]
The most recent such event was 65 Ma, when an asteroid impact triggered the extinction of the (non-avian)
dinosaurs and other large reptiles, but spared some small animals such as mammals, which then resembled
shrews. Over the past 65 million years, mammalian life has diversified, and several million years ago an
African ape-like animal such as Orrorin tugenensis gained the ability to stand upright.[46] This enabled tool
use and encouraged communication that provided the nutrition and stimulation needed for a larger brain,
which allowed the evolution of the human race. The development of agriculture, and then civilization, allowed
humans to influence the Earth in a short time span as no other life form had,[47] affecting both the nature and
quantity of other life forms.
The present pattern of ice ages began about 40 Ma and then intensified during the Pleistocene about
3 Ma. High-latitude regions have since undergone repeated cycles of glaciation and thaw, repeating every
40–100,000 years. The last continental glaciation ended 10,000 years ago.[48]
Future
Main article: Future of the Earth
See also: Risks to civilization, humans and planet Earth
The life cycle of the Sun
The future of the planet is closely tied to that of the Sun. As a result of the steady accumulation of
helium at the Sun's core, the star's total luminosity will slowly increase. The luminosity of the Sun will grow by
10% over the next 1.1 Gyr (1.1 billion years) and by 40% over the next 3.5 Gyr.[49] Climate models indicate
that the rise in radiation reaching the Earth is likely to have dire consequences, including the loss of the
planet's oceans.[50]
The Earth's increasing surface temperature will accelerate the inorganic CO2 cycle, reducing its
concentration to levels lethally low for plants (10 ppm for C4 photosynthesis) in approximately 500 million[20]
to 900 million years. The lack of vegetation will result in the loss of oxygen in the atmosphere, so animal life
will become extinct within several million more years.[51] After another billion years all surface water will
have disappeared[21] and the mean global temperature will reach 70 °C[51] (158 °F). The Earth is expected
to be effectively habitable for about another 500 million years from that point,[20] although this may be
extended up to 2.3 billion years if the nitrogen is removed from the atmosphere.[52] Even if the Sun were
eternal and stable, the continued internal cooling of the Earth would result in a loss of much of its CO 2 due to
reduced volcanism,[53] and 35% of the water in the oceans would descend to the mantle due to reduced
steam venting from mid-ocean ridges.[54]
The Sun, as part of its evolution, will become a red giant in about 5 Gyr. Models predict that the Sun
will expand out to about 250 times its present radius, roughly 1 AU (150,000,000 km).[49][55] Earth's fate is
less clear. As a red giant, the Sun will lose roughly 30% of its mass, so, without tidal effects, the Earth will
move to an orbit 1.7 AU (250,000,000 km) from the Sun when the star reaches it maximum radius. The
planet was therefore initially expected to escape envelopment by the expanded Sun's sparse outer
atmosphere, though most, if not all, remaining life would have been destroyed by the Sun's increased
luminosity (peaking at about 5000 times its present level).[49] However, a 2008 simulation indicates that
Earth's orbit will decay due to tidal effects and drag, causing it to enter the red giant Sun's atmosphere and
be vaporized.[55]
Composition and structure

Main article: Earth science
Further information: Earth physical characteristics tables
Earth is a terrestrial planet, meaning that it is a rocky body, rather than a gas giant like Jupiter. It is
the largest of the four solar terrestrial planets in size and mass. Of these four planets, Earth also has the
highest density, the highest surface gravity, the strongest magnetic field, and fastest rotation.[56] It also is the
only terrestrial planet with active plate tectonics.[57]
Shape
Main article: Figure of the Earth
Size comparison of inner planets (left to right): Mercury, Venus, Earth and Mars
The shape of the Earth is very close to that of an oblate spheroid, a sphere flattened along the axis
from pole to pole such that there is a bulge around the equator.[58] This bulge results from the rotation of the
Earth, and causes the diameter at the equator to be 43 km larger than the pole to pole diameter.[59] The
average diameter of the reference spheroid is about 12,742 km, which is approximately 40,000 km/π, as the
meter was originally defined as 1/10,000,000 of the distance from the equator to the North Pole through
Paris, France.[60]
Local topography deviates from this idealized spheroid, though on a global scale, these deviations
are very small: Earth has a tolerance of about one part in about 584, or 0.17%, from the reference spheroid,
which is less than the 0.22% tolerance allowed in billiard balls.[61] The largest local deviations in the rocky
surface of the Earth are Mount Everest (8848 m above local sea level) and the Mariana Trench (10,911 m
below local sea level). Because of the equatorial bulge, the surface locations farthest from the center of the
Earth are the summits of Mount Chimborazo in Ecuador and Huascarán in Peru.[62][63][64]
Chemical composition of the crust[65]
Composition
Compound Formula
Continental Oceanic
silica SiO2 60.2% 48.6%
alumina Al2O3 15.2% 16.5%
lime CaO 5.5% 12.3%
magnesia MgO 3.1% 6.8%
iron(II) oxide FeO 3.8% 6.2%
sodium oxide Na2O 3.0% 2.6%
potassium oxide K2O 2.8% 0.4%
iron(III) oxide Fe2O3 2.5% 2.3%
water H2O 1.4% 1.1%
carbon dioxide CO2 1.2% 1.4%

titanium dioxide TiO2 0.7% 1.4%
phosphorus pentoxide P2O5 0.2% 0.3%
Total 99.6% 99.9%
Chemical composition
See also: Abundance of elements on Earth
The mass of the Earth is approximately 5.98×1024 kg. It is composed mostly of iron (32.1%), oxygen (30.1%),
silicon (15.1%), magnesium (13.9%), sulfur (2.9%), nickel (1.8%), calcium (1.5%), and aluminium (1.4%);
with the remaining 1.2% consisting of trace amounts of other elements. Due to mass segregation, the core
region is believed to be primarily composed of iron (88.8%), with smaller amounts of nickel (5.8%), sulfur
(4.5%), and less than 1% trace elements.[66]
The geochemist F. W. Clarke calculated that a little more than 47% of the Earth's crust consists of
oxygen. The more common rock constituents of the Earth's crust are nearly all oxides; chlorine, sulfur and
fluorine are the only important exceptions to this and their total amount in any rock is usually much less than
1%. The principal oxides are silica, alumina, iron oxides, lime, magnesia, potash and soda. The silica
functions principally as an acid, forming silicates, and all the commonest minerals of igneous rocks are of this
nature. From a computation based on 1,672 analyses of all kinds of rocks, Clarke deduced that 99.22% were
composed of 11 oxides (see the table at right). All the other constituents occur only in very small quantities.
[67]
Internal structure
Main article: Structure of the Earth
The interior of the Earth, like that of the other terrestrial planets, is divided into layers by their
chemical or physical (rheological) properties, but unlike the other terrestrial planets, it has a distinct outer and
inner core. The outer layer of the Earth is a chemically distinct silicate solid crust, which is underlain by a
highly viscous solid mantle. The crust is separated from the mantle by the Mohorovičić discontinuity, and the
thickness of the crust varies: averaging 6 km under the oceans and 30–50 km on the continents. The crust
and the cold, rigid, top of the upper mantle are collectively known as the lithosphere, and it is of the
lithosphere that the tectonic plates are comprised. Beneath the lithosphere is the asthenosphere, a relatively
low-viscosity layer on which the lithosphere rides. Important changes in crystal structure within the mantle
occur at 410 and 660 kilometers below the surface, spanning a transition zone that separates the upper and
lower mantle. Beneath the mantle, an extremely low viscosity liquid outer core lies above a solid inner core.
[68] The inner core may rotate at a slightly higher angular velocity than the remainder of the planet,
advancing by 0.1–0.5° per year.[69]
Geologic layers of the Earth[70]
Depth[ Density
Component
71]
Earth cutaway from core to exosphere. Not to km
Layer g/cm3
scale.
Lithosphere[not
0–60 —
e 10]
0–35 Crust[note 11] 2.2–2.9

35–60 Upper mantle 3.4–4.4
35–
Mantle 3.4–5.6
2890
100–
Asthenosphere —
700
2890–
Outer core 9.9–12.2
5100
5100– 12.8–
Inner core
6378 13.1
Heat
Earth's internal heat comes from a combination of residual heat from planetary accretion (about 20%)
and heat produced through radioactive decay (80%).[72] The major heat-producing isotopes in the Earth are
potassium-40, uranium-238, uranium-235, and thorium-232.[73] At the center of the planet, the temperature
may be up to 7,000 K and the pressure could reach 360 GPa.[74] Because much of the heat is provided by
radioactive decay, scientists believe that early in Earth history, before isotopes with short half-lives had been
depleted, Earth's heat production would have been much higher. This extra heat production, twice present-
day at approximately 3 billion years ago,[72] would have increased temperature gradients within the Earth,
increasing the rates of mantle convection and plate tectonics, and allowing the production of igneous rocks
such as komatiites that are not formed today.[75]
Present-day major heat-producing isotopes[76]
Heat Half-life Mean mantle Heat
Isotope release concentration release
W/kg isotope years kg isotope/kg mantle W/kg mantle
238U 4.47 × 2.91 ×

9.46 × 10−5 30.8 × 10−9
109 10−12
235U 7.04 × 1.25 ×

5.69 × 10−4 0.22 × 10−9
108 10−13
232Th 1.40 × 3.27 ×

2.64 × 10−5 124 × 10−9
1010 10−12
40K 1.25 × 1.08 ×

2.92 × 10−5 36.9 × 10−9
109 10−12
The mean heat loss from the Earth is 87 mW m−2, for a global heat loss of 4.42 × 1013 W.[77] A
portion of the core's thermal energy is transported toward the crust by mantle plumes; a form of convection
consisting of upwellings of higher-temperature rock. These plumes can produce hotspots and flood basalts.
[78] More of the heat in the Earth is lost through plate tectonics, by mantle upwelling associated with mid-
ocean ridges. The final major mode of heat loss is through conduction through the lithosphere, the majority of
which occurs in the oceans because the crust there is much thinner than that of the continents.[79]
Tectonic plates
Earth's main plates[80]
Area
Plate name
106 km2
African Plate[note 12] 78.0
Antarctic Plate 60.9
Indo-Australian Plate 47.2
Eurasian Plate 67.8

North American Plate 75.9
South American Plate 43.6
Pacific Plate 103.3

Main article: Plate tectonics
The mechanically rigid outer layer of the Earth, the lithosphere, is broken into pieces called tectonic
plates. These plates are rigid segments that move in relation to one another at one of three types of plate
boundaries: Convergent boundaries, at which two plates come together, Divergent boundaries, at which two
plates are pulled apart, and Transform boundaries, in which two plates slide past one another laterally.
Earthquakes, volcanic activity, mountain-building, and oceanic trench formation can occur along these plate
boundaries.[81] The tectonic plates ride on top of the asthenosphere, the solid but less-viscous part of the
upper mantle that can flow and move along with the plates,[82] and their motion is strongly coupled with
convection patterns inside the Earth's mantle.
As the tectonic plates migrate across the planet, the ocean floor is subducted under the leading
edges of the plates at convergent boundaries. At the same time, the upwelling of mantle material at divergent
boundaries creates mid-ocean ridges. The combination of these processes continually recycles the oceanic
crust back into the mantle. Because of this recycling, most of the ocean floor is less than 100 million years in
age. The oldest oceanic crust is located in the Western Pacific, and has an estimated age of about 200
million years.[83][84] By comparison, the oldest dated continental crust is 4030 million years old.[85]
Other notable plates include the Indian Plate, the Arabian Plate, the Caribbean Plate, the Nazca
Plate off the west coast of South America and the Scotia Plate in the southern Atlantic Ocean. The Australian
Plate fused with the Indian Plate between 50 and 55 million years ago. The fastest-moving plates are the
oceanic plates, with the Cocos Plate advancing at a rate of 75 mm/yr[86] and the Pacific Plate moving 52–
69 mm/yr. At the other extreme, the slowest-moving plate is the Eurasian Plate, progressing at a typical rate
of about 21 mm/yr.[87]
Surface
Main articles: Landform and Extreme points of Earth
The Earth's terrain varies greatly from place to place. About 70.8%[88] of the surface is covered by
water, with much of the continental shelf below sea level. The submerged surface has mountainous features,
including a globe-spanning mid-ocean ridge system, as well as undersea volcanoes,[59] oceanic trenches,
submarine canyons, oceanic plateaus and abyssal plains. The remaining 29.2% not covered by water
consists of mountains, deserts, plains, plateaus, and other geomorphologies.
The planetary surface undergoes reshaping over geological time periods because of tectonics and
erosion. The surface features built up or deformed through plate tectonics are subject to steady weathering
from precipitation, thermal cycles, and chemical effects. Glaciation, coastal erosion, the build-up of coral
reefs, and large meteorite impacts[89] also act to reshape the landscape.
Present day Earth altimetry and bathymetry. Data from the National Geophysical Data Center's
TerrainBase Digital Terrain Model.
The continental crust consists of lower density material such as the igneous rocks granite and
andesite. Less common is basalt, a denser volcanic rock that is the primary constituent of the ocean floors.
[90] Sedimentary rock is formed from the accumulation of sediment that becomes compacted together.
Nearly 75% of the continental surfaces are covered by sedimentary rocks, although they form only about 5%
of the crust.[91] The third form of rock material found on Earth is metamorphic rock, which is created from the
transformation of pre-existing rock types through high pressures, high temperatures, or both. The most
abundant silicate minerals on the Earth's surface include quartz, the feldspars, amphibole, mica, pyroxene
and olivine.[92] Common carbonate minerals include calcite (found in limestone) and dolomite.[93]
The pedosphere is the outermost layer of the Earth that is composed of soil and subject to soil formation
processes. It exists at the interface of the lithosphere, atmosphere, hydrosphere and biosphere. Currently the
total arable land is 13.31% of the land surface, with only 4.71% supporting permanent crops.[10] Close to
40% of the Earth's land surface is presently used for cropland and pasture, or an estimated 1.3×10 7 km2 of
cropland and 3.4×107 km2 of pastureland.[94]
The elevation of the land surface of the Earth varies from the low point of −418 m at the Dead Sea, to
a 2005-estimated maximum altitude of 8,848 m at the top of Mount Everest. The mean height of land above
sea level is 840 m.[95]
Hydrosphere
Main article: Hydrosphere
Elevation histogram of the surface of the Earth
The abundance of water on Earth's surface is a unique feature that distinguishes the "Blue Planet"
from others in the Solar System. The Earth's hydrosphere consists chiefly of the oceans, but technically
includes all water surfaces in the world, including inland seas, lakes, rivers, and underground waters down to
a depth of 2,000 m. The deepest underwater location is Challenger Deep of the Mariana Trench in the Pacific
Ocean with a depth of −10,911.4 m.[note 13][96]
The mass of the oceans is approximately 1.35×10 18 metric tons, or about 1/4400 of the total mass of the
Earth. The oceans cover an area of 3.618×108 km2 with a mean depth of 3,682 m, resulting in an estimated
volume of 1.332×109 km3.[97] If all the land on Earth were spread evenly, water would rise to an altitude of
more than 2.7 km.[note 14] About 97.5% of the water is saline, while the remaining 2.5% is fresh water. Most
fresh water, about 68.7%, is currently ice.[98]
The average salinity of the Earth's oceans is about 35 grams of salt per kilogram of sea water
(35 ‰).[99] Most of this salt was released from volcanic activity or extracted from cool, igneous rocks.[100]
The oceans are also a reservoir of dissolved atmospheric gases, which are essential for the survival of many
aquatic life forms.[101] Sea water has an important influence on the world's climate, with the oceans acting
as a large heat reservoir.[102] Shifts in the oceanic temperature distribution can cause significant weather
shifts, such as the El Niño-Southern Oscillation.[103]
Atmosphere
Main article: Atmosphere of Earth
The atmospheric pressure on the surface of the Earth averages 101.325 kPa, with a scale height of
about 8.5 km.[3] It is 78% nitrogen and 21% oxygen, with trace amounts of water vapor, carbon dioxide and
other gaseous molecules. The height of the troposphere varies with latitude, ranging between 8 km at the
poles to 17 km at the equator, with some variation resulting from weather and seasonal factors.[104]
Earth's biosphere has significantly altered its atmosphere. Oxygenic photosynthesis evolved 2.7
billion years ago, forming the primarily nitrogen-oxygen atmosphere of today. This change enabled the
proliferation of aerobic organisms as well as the formation of the ozone layer which blocks ultraviolet solar
radiation, permitting life on land. Other atmospheric functions important to life on Earth include transporting
water vapor, providing useful gases, causing small meteors to burn up before they strike the surface, and
moderating temperature.[105] This last phenomenon is known as the greenhouse effect: trace molecules
within the atmosphere serve to capture thermal energy emitted from the ground, thereby raising the average
temperature. Carbon dioxide, water vapor, methane and ozone are the primary greenhouse gases in the
Earth's atmosphere. Without this heat-retention effect, the average surface temperature would be −18 °C and
life would likely not exist.[88]
Weather and climate

Main articles: Weather and Climate
Satellite cloud cover image of Earth using NASA's Moderate-Resolution Imaging Spectroradiometer.
The Earth's atmosphere has no definite boundary, slowly becoming thinner and fading into outer
space. Three-quarters of the atmosphere's mass is contained within the first 11 km of the planet's surface.
This lowest layer is called the troposphere. Energy from the Sun heats this layer, and the surface below,
causing expansion of the air. This lower density air then rises, and is replaced by cooler, higher density air.
The result is atmospheric circulation that drives the weather and climate through redistribution of heat
energy.[106]
The primary atmospheric circulation bands consist of the trade winds in the equatorial region below
30° latitude and the westerlies in the mid-latitudes between 30° and 60°.[107] Ocean currents are also
important factors in determining climate, particularly the thermohaline circulation that distributes heat energy
from the equatorial oceans to the polar regions.[108]
Water vapor generated through surface evaporation is transported by circulatory patterns in the
atmosphere. When atmospheric conditions permit an uplift of warm, humid air, this water condenses and
settles to the surface as precipitation.[106] Most of the water is then transported to lower elevations by river
systems and usually returned to the oceans or deposited into lakes. This water cycle is a vital mechanism for
supporting life on land, and is a primary factor in the erosion of surface features over geological periods.
Precipitation patterns vary widely, ranging from several meters of water per year to less than a millimeter.
Atmospheric circulation, topological features and temperature differences determine the average
precipitation that falls in each region.[109]
The amount of solar energy reaching the Earth's decreases with increasing latitude. At higher
latitudes the sunlight reaches the surface at a lower angles and it must pass through thicker columns of the
atmosphere. As a result, the mean annual air temperature at sea level decreases by about 0.4°C per per
degree of latitude away from the equator.[110] The Earth can be sub-divided into specific latitudinal belts of
approximately homogeneous climate. Ranging from the equator to the polar regions, these are the tropical
(or equatorial), subtropical, temperate and polar climates.[111] Climate can also be classified based on the
temperature and precipitation, with the climate regions characterized by fairly uniform air masses. The
commonly used Köppen climate classification system (as modified by Wladimir Köppen's student Rudolph
Geiger) has five broad groups (humid tropics, arid, humid middle latitudes, continental and cold polar), which
are further divided into more specific subtypes.[107]
Upper atmosphere
This view from orbit shows the full Moon partially obscured and deformed by the Earth's atmosphere.
NASA image.
See also: Outer space
Above the troposphere, the atmosphere is usually divided into the stratosphere, mesosphere, and
thermosphere.[105] Each layer has a different lapse rate, defining the rate of change in temperature with
height. Beyond these, the exosphere thins out into the magnetosphere, where the Earth's magnetic fields
interact with the solar wind.[112] Within the stratosphere is the ozone layer, a component that partially
shields the surface from ultraviolet light and thus is important for life on Earth. The Kármán line, defined as
100 km above the Earth's surface, is a working definition for the boundary between atmosphere and space.
[113]
Thermal energy causes some of the molecules at the outer edge of the Earth's atmosphere have
their velocity increased to the point where they can escape from the planet's gravity. This results in a slow but
steady leakage of the atmosphere into space. Because unfixed hydrogen has a low molecular weight, it can
achieve escape velocity more readily and it leaks into outer space at a greater rate than other gasses.[114]
The leakage of hydrogen into space contributes to the pushing of the Earth from an initially reducing state to
its current oxidizing one. Photosynthesis provided a source of free oxygen, but the loss of reducing agents
such as hydrogen is believed to have been a necessary precondition for the widespread accumulation of
oxygen in the atmosphere.[115] Hence the ability of hydrogen to escape from the Earth's atmosphere may
have influenced the nature of life that developed on the planet.[116] In the current, oxygen-rich atmosphere
most hydrogen is converted into water before it has an opportunity to escape. Instead, most of the hydrogen
loss comes from the destruction of methane in the upper atmosphere.[117]
Magnetic field
Schematic of Earth's magnetosphere. The solar wind flows from left to right
Main article: Earth's magnetic field
The Earth's magnetic field is shaped roughly as a magnetic dipole, with the poles currently located
proximate to the planet's geographic poles. At the equator of the magnetic field, the magnetic field strength at
the planet's surface is 3.05 × 10−5 T, with global magnetic dipole moment of 7.91 × 1015 T m3.[118]
According to dynamo theory, the field is generated within the molten outer core region where heat creates
convection motions of conducting materials, generating electric currents. These in turn produce the Earth's
magnetic field. The convection movements in the core are chaotic; the magnetic poles drift and periodically
change alignment. This results in field reversals at irregular intervals averaging a few times every million
years. The most recent reversal occurred approximately 700,000 years ago.[119][120]
The field forms the magnetosphere, which deflects particles in the solar wind. The sunward edge of
the bow shock is located at about 13 times the radius of the Earth. The collision between the magnetic field
and the solar wind forms the Van Allen radiation belts, a pair of concentric, torus-shaped regions of energetic
charged particles. When the plasma enters the Earth's atmosphere at the magnetic poles, it forms the aurora.
[121]
Orbit and rotation

Rotation
Main article: Earth's rotation
Earth's axial tilt (or obliquity) and its relation to the rotation axis and plane of orbit.
Earth's rotation period relative to the Sun—its mean solar day—is 86,400 seconds of mean solar time
(86,400.0025 SI seconds).[122] As the Earth's solar day is now slightly longer than it was during the 19th
century because of tidal acceleration, each day varies between 0 and 2 SI ms longer.[123][124]
Earth's rotation period relative to the fixed stars, called its stellar day by the International Earth
Rotation and Reference Systems Service (IERS), is 86164.098903691 seconds of mean solar time (UT1), or
23h 56m 4.098903691s. [2][note 15] Earth's rotation period relative to the precessing or moving mean vernal
equinox, misnamed its sidereal day, is 86164.09053083288 seconds of mean solar time (UT1) (23 h 56m
4.09053083288s).[2] Thus the sidereal day is shorter than the stellar day by about 8.4 ms.[125] The length of
the mean solar day in SI seconds is available from the IERS for the periods 1623–2005[126] and 1962–2005.
[127]
Apart from meteors within the atmosphere and low-orbiting satellites, the main apparent motion of
celestial bodies in the Earth's sky is to the west at a rate of 15°/h = 15'/min. For bodies near the celestial
equator, this is equivalent to an apparent diameter of the Sun or Moon every two minutes; from the planet's
surface, the apparent sizes of the Sun and the Moon are approximately the same.[128][129]
Orbit
Main article: Earth's orbit
Earth orbits the Sun at an average distance of about 150 million kilometers every
365.2564 mean solar days, or one sidereal year. From Earth, this gives an apparent movement of the Sun
eastward with respect to the stars at a rate of about 1°/day, or a Sun or Moon diameter every 12 hours.
Because of this motion, on average it takes 24 hours—a solar day—for Earth to complete a full rotation about
its axis so that the Sun returns to the meridian. The orbital speed of the Earth averages about 30 km/s
(108,000 km/h), which is fast enough to cover the planet's diameter (about 12,600 km) in seven minutes, and
the distance to the Moon (384,000 km) in four hours.[3]
The Moon revolves with the Earth around a common barycenter every 27.32 days relative to the
background stars. When combined with the Earth–Moon system's common revolution around the Sun, the
period of the synodic month, from new moon to new moon, is 29.53 days. Viewed from the celestial north
pole, the motion of Earth, the Moon and their axial rotations are all counter-clockwise. Viewed from a vantage
point above the north poles of both the Sun and the Earth, the Earth appears to revolve in a counterclockwise
direction about the Sun. The orbital and axial planes are not precisely aligned: Earth's axis is tilted some
23.5 degrees from the perpendicular to the Earth–Sun plane, and the Earth–Moon plane is tilted about
5 degrees against the Earth-Sun plane. Without this tilt, there would be an eclipse every two weeks,
alternating between lunar eclipses and solar eclipses.[3][130]
The Hill sphere, or gravitational sphere of influence, of the Earth is about 1.5 Gm (or 1,500,000
kilometers) in radius.[131][note 16] This is maximum distance at which the Earth's gravitational influence is
stronger than the more distant Sun and planets. Objects must orbit the Earth within this radius, or they can
become unbound by the gravitational perturbation of the Sun.
Illustration of the Milky Way Galaxy, showing the location of the Sun
Earth, along with the Solar System, is situated in the Milky Way galaxy, orbiting about 28,000 light
years from the center of the galaxy. It is currently about 20 light years above the galaxy's equatorial plane in
the Orion spiral arm.[132]
Axial tilt and seasons

Main article: Axial tilt
Because of the axial tilt of the Earth, the amount of sunlight reaching any given point on the surface
varies over the course of the year. This results in seasonal change in climate, with summer in the northern
hemisphere occurring when the North Pole is pointing toward the Sun, and winter taking place when the pole
is pointed away. During the summer, the day lasts longer and the Sun climbs higher in the sky. In winter, the
climate becomes generally cooler and the days shorter. Above the Arctic Circle, an extreme case is reached
where there is no daylight at all for part of the year—a polar night. In the southern hemisphere the situation is
exactly reversed, with the South Pole oriented opposite the direction of the North Pole.
Earth and Moon from Mars, imaged by Mars Reconnaissance Orbiter. From space, the Earth can be
seen to go through phases similar to the phases of the Moon.
By astronomical convention, the four seasons are determined by the solstices—the point in the orbit of
maximum axial tilt toward or away from the Sun—and the equinoxes, when the direction of the tilt and the
direction to the Sun are perpendicular. In the northern hemisphere, Winter Solstice occurs on about
December 21, Summer Solstice is near June 21, Spring Equinox is around March 20 and Autumnal Equinox
is about September 23. In the Southern hemisphere, the situation is reversed, with the Summer and Winter
Solstices exchanged and the Spring and Autumnal Equinox dates switched.[133]
The angle of the Earth's tilt is relatively stable over long periods of time. However, the tilt does
undergo nutation; a slight, irregular motion with a main period of 18.6 years.[134] The orientation (rather than
the angle) of the Earth's axis also changes over time, precessing around in a complete circle over each
25,800 year cycle; this precession is the reason for the difference between a sidereal year and a tropical
year. Both of these motions are caused by the varying attraction of the Sun and Moon on the Earth's
equatorial bulge. From the perspective of the Earth, the poles also migrate a few meters across the surface.
This polar motion has multiple, cyclical components, which collectively are termed quasiperiodic motion. In
addition to an annual component to this motion, there is a 14-month cycle called the Chandler wobble. The
rotational velocity of the Earth also varies in a phenomenon known as length of day variation.[135]
In modern times, Earth's perihelion occurs around January 3, and the aphelion around July 4.
However, these dates change over time due to precession and other orbital factors, which follow cyclical
patterns known as Milankovitch cycles. The changing Earth-Sun distance results in an increase of about
6.9%[note 17] in solar energy reaching the Earth at perihelion relative to aphelion. Since the southern
hemisphere is tilted toward the Sun at about the same time that the Earth reaches the closest approach to
the Sun, the southern hemisphere receives slightly more energy from the Sun than does the northern over
the course of a year. However, this effect is much less significant than the total energy change due to the
axial tilt, and most of the excess energy is absorbed by the higher proportion of water in the southern
hemisphere.[136]
Moon
Characteristics
Diameter 3,474.8 km
Mass 7.349×1022 kg
Semi-major axis 384,400 km
Orbital period 27 d 7 h 43.7 m

Main article: Moon
The Moon is a relatively large, terrestrial, planet-like satellite, with a diameter about one-quarter of
the Earth's. It is the largest moon in the Solar System relative to the size of its planet, although Charon is
larger relative to the dwarf planet Pluto. The natural satellites orbiting other planets are called "moons" after
Earth's Moon.
The gravitational attraction between the Earth and Moon causes tides on Earth. The same effect on
the Moon has led to its tidal locking: its rotation period is the same as the time it takes to orbit the Earth. As a
result, it always presents the same face to the planet. As the Moon orbits Earth, different parts of its face are
illuminated by the Sun, leading to the lunar phases; the dark part of the face is separated from the light part
by the solar terminator.
Because of their tidal interaction, the Moon recedes from Earth at the rate of approximately 38 mm a
year. Over millions of years, these tiny modifications—and the lengthening of Earth's day by about 23 µs a
year—add up to significant changes.[137] During the Devonian period, for example, (approximately 410
million years ago) there were 400 days in a year, with each day lasting 21.8 hours.[138]
Details of the Earth-Moon system. Besides the radius of each object, the radius to the Earth-Moon
barycenter is shown. Photos from NASA. Data from NASA. The Moon's axis is located by Cassini's third law.
The Moon may have dramatically affected the development of life by moderating the planet's climate.
Paleontological evidence and computer simulations show that Earth's axial tilt is stabilized by tidal
interactions with the Moon.[139] Some theorists believe that without this stabilization against the torques
applied by the Sun and planets to the Earth's equatorial bulge, the rotational axis might be chaotically
unstable, exhibiting chaotic changes over millions of years, as appears to be the case for Mars.[140]
Viewed from Earth, the Moon is just far enough away to have very nearly the same apparent-sized
disk as the Sun. The angular size (or solid angle) of these two bodies match because, although the Sun's
diameter is about 400 times as large as the Moon's, it is also 400 times more distant.[129] This allows total
and annular solar eclipses to occur on Earth.
The most widely accepted theory of the Moon's origin, the giant impact theory, states that it formed
from the collision of a Mars-size protoplanet called Theia with the early Earth. This hypothesis explains
(among other things) the Moon's relative lack of iron and volatile elements, and the fact that its composition is
nearly identical to that of the Earth's crust.[141]
Earth has at least two co-orbital asteroids, 3753 Cruithne and 2002 AA29.[142]
A scale representation of the relative sizes of, and average distance between, Earth and Moon
Habitability
See also: Planetary habitability
A planet that can sustain life is termed habitable, even if life did not originate there. The Earth
provides the (currently understood) requisite conditions of liquid water, an environment where complex
organic molecules can assemble, and sufficient energy to sustain metabolism.[143] The distance of the Earth
from the Sun, as well as its orbital eccentricity, rate of rotation, axial tilt, geological history, sustaining
atmosphere and protective magnetic field all contribute to the conditions believed necessary to originate and
sustain life on this planet.[144]
Biosphere
Main article: Biosphere
The planet's life forms are sometimes said to form a "biosphere". This biosphere is generally believed
to have begun evolving about 3.5 billion years ago. Earth is the only place where life is known to exist. The
biosphere is divided into a number of biomes, inhabited by broadly similar plants and animals. On land,
biomes are separated primarily by differences in latitude, height above sea level and humidity. Terrestrial
biomes lying within the Arctic or Antarctic Circles, at high altitudes or in extremely arid areas are relatively
barren of plant and animal life; species diversity reaches a peak in humid lowlands at equatorial latitudes.
[145]
Natural resources and land use

Main article: Natural resource
The Earth provides resources that are exploitable by humans for useful purposes. Some of these are
non-renewable resources, such as mineral fuels, that are difficult to replenish on a short time scale.
Large deposits of fossil fuels are obtained from the Earth's crust, consisting of coal, petroleum,
natural gas and methane clathrate. These deposits are used by humans both for energy production and as
feedstock for chemical production. Mineral ore bodies have also been formed in Earth's crust through a
process of Ore genesis, resulting from actions of erosion and plate tectonics.[146] These bodies form
concentrated sources for many metals and other useful elements.
The Earth's biosphere produces many useful biological products for humans, including (but far from
limited to) food, wood, pharmaceuticals, oxygen, and the recycling of many organic wastes. The land-based
ecosystem depends upon topsoil and fresh water, and the oceanic ecosystem depends upon dissolved
nutrients washed down from the land.[147] Humans also live on the land by using building materials to
construct shelters. In 1993, human use of land is approximately:
Arable Permanent Permanent Forests Urban
Land use O
land crops pastures and woodland areas
13.13%
Percentage 4.71%[10] 26% 32% 1.5%
[10]
The estimated amount of irrigated land in 1993 was 2,481,250 km2.[10]
Natural and environmental hazards

Large areas of the Earth's surface are subject to extreme weather such as tropical cyclones,
hurricanes, or typhoons that dominate life in those areas. Many places are subject to earthquakes,
landslides, tsunamis, volcanic eruptions, tornadoes, sinkholes, blizzards, floods, droughts, and other
calamities and disasters.
Many localized areas are subject to human-made pollution of the air and water, acid rain and toxic
substances, loss of vegetation (overgrazing, deforestation, desertification), loss of wildlife, species extinction,
soil degradation, soil depletion, erosion, and introduction of invasive species.
According to the United Nations, a scientific consensus exists linking human activities to global
warming due to industrial carbon dioxide emissions. This is predicted to produce changes such as the
melting of glaciers and ice sheets, more extreme temperature ranges, significant changes in weather and a
global rise in average sea levels.[148]
Human geography
Main article: Human geography
See also: World
Cartography, the study and practice of map making, and vicariously geography, have historically
been the disciplines devoted to depicting the Earth. Surveying, the determination of locations and distances,
and to a lesser extent navigation, the determination of position and direction, have developed alongside
cartography and geography, providing and suitably quantifying the requisite information.
Earth has approximately 6,803,000,000 human inhabitants as of December 12, 2009.[149]
Projections indicate that the world's human population will reach seven billion in 2013 and 9.2 billion in 2050.
[150] Most of the growth is expected to take place in developing nations. Human population density varies
widely around the world, but a majority live in Asia. By 2020, 60% of the world's population is expected to be
living in urban, rather than rural, areas.[151]
It is estimated that only one-eighth of the surface of the Earth is suitable for humans to live on—three-
quarters is covered by oceans, and half of the land area is either desert (14%),[152] high mountains (27%),
[153] or other less suitable terrain. The northernmost permanent settlement in the world is Alert, on
Ellesmere Island in Nunavut, Canada.[154] (82°28′N) The southernmost is the Amundsen-Scott South Pole
Station, in Antarctica, almost exactly at the South Pole. (90°S)
The Earth at night, a composite of DMSP/OLS ground illumination data on a simulated night-time
image of the world. This image is not photographic and many features are brighter than they would appear to
a direct observer.
Independent sovereign nations claim the planet's entire land surface, except for some parts of
Antarctica and the odd unclaimed area of Bir Tawil between Egypt and Sudan. As of 2007 there are 201
sovereign states, including the 192 United Nations member states. In addition, there are 59 dependent
territories, and a number of autonomous areas, territories under dispute and other entities.[10] Historically,
Earth has never had a sovereign government with authority over the entire globe, although a number of
nation-states have striven for world domination and failed.[155]
The United Nations is a worldwide intergovernmental organization that was created with the goal of
intervening in the disputes between nations, thereby avoiding armed conflict.[156] It is not, however, a world
government. The U.N. serves primarily as a forum for international diplomacy and international law. When
the consensus of the membership permits, it provides a mechanism for armed intervention.[157]
The first human to orbit the Earth was Yuri Gagarin on April 12, 1961.[158] In total, about 400 people
visited outer space and reached Earth orbit as of 2004, and, of these, twelve have walked on the Moon.[159]
[160][161] Normally the only humans in space are those on the International Space Station. The station's
crew, currently six people, is usually replaced every six months.[162] The furthest humans have travelled
from Earth is 400,171 km, achieved during the 1970 Apollo 13 mission.[163]
Cultural viewpoint
Main article: Earth in culture
The first photograph ever taken by astronauts of an "Earthrise", from Apollo 8
The name "Earth" derives from the Anglo-Saxon word erda, which means ground or soil, and is
related to the German word erde. It became eorthe later, and then erthe in Middle English.[164] The standard
astronomical symbol of the Earth consists of a cross circumscribed by a circle.[165]
Unlike the rest of the planets in the Solar System, humankind did not begin to view the Earth as a
moving object in orbit around the Sun until the 16th century.[166] Earth has often been personified as a deity,
in particular a goddess. In many cultures the mother goddess is also portrayed as a fertility deity. Creation
myths in many religions recall a story involving the creation of the Earth by a supernatural deity or deities. A
variety of religious groups, often associated with fundamentalist branches of Protestantism[167] or Islam,
[168] assert that their interpretations of these creation myths in sacred texts are literal truth and should be
considered alongside or replace conventional scientific accounts of the formation of the Earth and the origin
and development of life.[169] Such assertions are opposed by the scientific community[170][171] and by
other religious groups.[172][173][174] A prominent example is the creation-evolution controversy.
In the past there were varying levels of belief in a flat Earth,[175] but this was displaced by the
concept of a spherical Earth due to observation and circumnavigation.[176] The human perspective
regarding the Earth has changed following the advent of spaceflight, and the biosphere is now widely viewed
from a globally integrated perspective.[177][178] This is reflected in a growing environmental movement that
is concerned about humankind's effects on the planet.[179]
See also
Earth sciences portal
Book:Solar System

• Geodesy
• Geology
Notes
1. ^ All astronomical quantities vary, both secularly and periodically. The quantities given are
the values at the instant J2000.0 of the secular variation, ignoring all periodic variations.
2. ^ a b aphelion = a × (1 + e); perihelion = a × (1 - e), where a is the semi-major axis and e is
the eccentricity.
3. ^ The reference lists the longitude of the ascending node as -11.26064°, which is equivalent
to 348.73936° by the fact that any angle is equal to itself plus 360°.
4. ^ The reference lists the longitude of perihelion, which is the sum of the longitude of the
ascending node and the argument of perihelion. That is, 114.20783° + (-11.26064°) = 102.94719°.
5. ^ Due to natural fluctuations, ambiguities surrounding ice shelves, and mapping conventions
for vertical datums, exact values for land and ocean coverage are not meaningful. Based on data
from the Vector Map and Global Landcover datasets, extreme values for coverage of lakes and
streams are 0.6% and 1.0% of the earth's surface. The ice shields of Antarctica and Greenland are
counted as land, even though much of the rock which supports them lies below sea level.
6. ^ "Blue Planet" is used as the title of two films ( Blue Planet and The Blue Planet), in the Life
issue "The Incredible Year '68", which featured the Earthrise photo with lines from poet James
Dickey ("Behold/The blue planet steeped in its dream/Of reality"), and in the title of the European
Space Agency bulletin report Exploring the water cycle of the 'Blue Planet'.
7. ^ By International Astronomical Union convention, the term terra is used only for naming
extensive land masses on celestial bodies other than the Earth. Cf. Blue, Jennifer (2007-07-05).
"Descriptor Terms (Feature Types)". Gazetteer of Planetary Nomenclature. USGS.
http://planetarynames.wr.usgs.gov/jsp/append5.jsp. Retrieved 2007-07-05.
8. ^ At present, the other planets in the Solar System are either too hot or too cold to support
liquid water on the surface in vapor-liquid equilibrium. As of 2007, water vapor has been detected in
the atmosphere of only one extrasolar planet, and it is a gas giant. See: Tinetti, G.; Vidal-Madjar, A.;
Liang, M.C.; Beaulieu, J. P.; Yung, Y.; Carey, S.; Barber, R. J.; Tennyson, J.; Ribas, I (July 2007).
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http://www.nature.com/nature/journal/v448/n7150/abs/nature06002.html.
9. ^ The number of solar days is one less than the number of sidereal days because the orbital
motion of the Earth about the Sun results in one additional revolution of the planet about its axis.
10.^ Locally varies between 5 and 200 km.
11.^ Locally varies between 5 and 70 km.
12.^ Including the Somali Plate, which is currently in the process of formation out of the African
Plate. See: Chorowicz, Jean (October 2005). "The East African rift system". Journal of African Earth
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13.^ This is the measurement taken by the vessel Kaikō in March 1995 and is believed to be the
most accurate measurement to date. See the Challenger Deep article for more details.
14.^ The total surface area of the Earth is 5.1×108 km2. To first approximation, the average depth would
be the ratio of the two, or 2.7 km.
15.^ Aoki, the ultimate source of these figures, uses the term "seconds of UT1" instead of
"seconds of mean solar time".—Aoki, S. (1982). "The new definition of universal time". Astronomy and
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16.^ For the Earth, the Hill radius is
,
where m is the mass of the Earth, a is an Astronomical Unit, and M is the mass of the Sun. So the
radius in A.U. is about: .

17.^ Aphelion is 103.4% of the distance to perihelion. Due to the inverse square law, the
radiation at perihelion is about 106.9% the energy at aphelion.
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Further reading
• Comins, Neil F. (2001). Discovering the Essential Universe (Second ed.). W. H. Freeman.
ISBN 0-7167-5804-0. http://adsabs.harvard.edu/abs/2003deu..book.....C. Retrieved 2007-03-17.
External links
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• Global Measured Extremes of Temperature and Precipitation National Climatic Data Center
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Categories: Earth | Geography | Geology | Terrestrial planets
W000
Macroevolution

Evolution
Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation
Introduction
Evidence
History
Level of support
Modern synthesis
Social effect
Theory and fact
Cladistics
Ecological genetics
Molecular evolution
Phylogenetics
Population genetics
Systematics
Macroevolution is evolution on a scale of separated gene pools.[1] Macroevolutionary studies focus
on change that occurs at or above the level of species, in contrast with microevolution,[2] which refers to
smaller evolutionary changes (typically described as changes in allele frequencies) within a species or
population.[3]
The process of speciation may fall within the purview of either, depending on the forces thought to
drive it. Paleontology, evolutionary developmental biology, comparative genomics and genomic
phylostratigraphy contribute most of the evidence for the patterns and processes that can be classified as
macroevolution. An example of macroevolution is the appearance of feathers during the evolution of birds
from theropod dinosaurs.
Abrupt transformations from one biologic system to another, for example the passing of life from
water into land or the transition from invertebrates to vertebrates, are rare. Few major biological types have
emerged during the evolutionary history of life and most of them survive till today. When lifeforms take such
giant leaps, they meet little to no competition and are able to exploit a plethora of available niches, following
a pattern of adaptive radiation. This can lead to convergent evolution, where unrelated populations display
similar adaptations.[4]
The evolutionary course of Equidae (wide family including all horses and related animals) is often
viewed as a typical example of macroevolution. The earliest known genus, Hyracotherium (now reclassified
as a palaeothere), was a herbivore animal resembling a dog that lived in the early cenozoic. As its habitat
transformed into an open arid grassland, selective pressure required that the animal become a fast grazer.
Thus elongation of legs and head as well as reduction of toes gradually occurred, producing the only extant
genus of Equidae, Equus.[4]
Contents
[hide]
• 1 Origin of the term
• 2 Macroevolution and the modern evolutionary synthesis
• 3 Research topics
• 4 Misuse
• 5 See also
• 6 References
[edit] Origin of the term
Russian entomologist Yuri Filipchenko first coined the terms "macroevolution" and "microevolution"
in 1927 in his German language work, "Variabilität und Variation". Since the inception of the two terms, their
meanings have been revised several times and the term macroevolution fell into limited disfavour when it
was taken over by such writers as the geneticist Richard Goldschmidt (1940) and the paleontologist Otto
Schindewolf to describe their orthogenetic theories.[5]
A more practical definition of the term describes it as changes occurring on geological time scales, in contrast
to microevolution, which occurs on the timescale of human lifetimes.[6] This definition reflects the spectrum
between micro- and macro-evolution, whilst leaving a clear difference between the terms: because the
geological record rarely has a resolution better than 10,000 years, and humans rarely live longer than 100
years, "meso-evolution" is never observed.[7]
As a result, apart from Dobzhansky, Bernhard Rensch and Ernst Mayr, very few neo-Darwinian writers used
the term, preferring instead to talk of evolution as changes in allele frequencies without mention of the level
of the changes (above species level or below). Those who did were generally working within the continental
European traditions (as Dobzhansky, Ernst Mayr, Bernhard Rensch, Richard Goldschmidt, and Otto
Schindewolf were) and those who didn't were generally working within the Anglo-American tradition (such as
John Maynard Smith and Richard Dawkins). Hence, use of the term "macroevolution" is sometimes wrongly
used as a litmus test of whether the writer is "properly" neo-Darwinian or not.[5]
[edit] Macroevolution and the modern evolutionary synthesis
Within the Modern Synthesis school of thought, macroevolution is thought of as the compounded
effects of microevolution.[8] Thus, the distinction between micro- and macroevolution is not a fundamental
one – the only difference between them is of time and scale. As Ernst W. Mayr observes, "transspecific
evolution is nothing but an extrapolation and magnification of the events that take place within populations
and species...it is misleading to make a distinction between the causes of micro- and macroevolution”.[8]
However, time is not a necessary distinguishing factor – macroevolution can happen without gradual
compounding of small changes; whole-genome duplication can result in speciation occurring over a single
generation - this is especially common in plants.[9]
Changes in the genes regulating development have also been proposed as being important in
producing speciation through large and relatively sudden changes in animals' morphology.[10][11]
[edit] Research topics

Some examples of subjects whose study falls within the realm of macroevolution:
• Adaptive radiations such as The Cambrian Explosion.
• Changes in biodiversity through time.
• Genomic evolution, like horizontal gene transfer, genome fusions in endosymbioses, and
adaptive changes in genome size.
• Mass extinctions.
• Speciation and extinction rates.
• The debate between punctuated equilibrium and gradualism.
• The role of development in shaping evolution, particularly such topics as heterochrony and
phenotypic plasticity.
[edit] Misuse
Main article: Objections to evolution
See also: Speciation
The term "macroevolution" frequently arises within the context of the evolution/creation debate,
usually used by creationists alleging a significant difference between the evolutionary changes observed in
field and laboratory studies and the larger scale macroevolutionary changes that scientists believe to have
taken thousands or millions of years to occur. They may accept that evolutionary change is possible within
species ("microevolution"), but deny that one species can evolve into another ("macroevolution").[1] Contrary
to this belief among the anti-evolution movement proponents, evolution of life forms beyond the species level
("macroevolution", i.e. speciation in a specific case) has indeed been observed multiple times under both
controlled laboratory conditions and in nature.[12] The claim that macroevolution does not occur, or is
impossible, is thus demonstrably false and without support in the scientific community.
Such claims are rejected by the scientific community on the basis of ample evidence that
macroevolution is an active process both presently and in the past.[5][13] The terms macroevolution and
microevolution relate to the same processes operating at different scales, but creationist claims misuse the
terms in a vaguely defined way which does not accurately reflect scientific usage, acknowledging well
observed evolution as "microevolution" and denying that "macroevolution" takes place.[5][14] Evolutionary
theory (including macroevolutionary change) remains the dominant scientific paradigm for explaining the
origins of Earth's biodiversity. Its occurrence is not disputed within the scientific community.[15] While details
of macroevolution are continuously studied by the scientific community, the overall theory behind
macroevolution (i.e. common descent) has been overwhelmingly consistent with empirical data. Predictions
of empirical data from the theory of common descent have been so consistent that biologists often refer to it
as the "fact of evolution".[16][17]
Nicholas Matzke and Paul R. Gross have accused creationists of using "strategically elastic"
definitions of micro- and macroevolution when discussing the topic.[1] The actual definition of macroevolution
accepted by scientists is "any change at the species level or above" (phyla, group, etc.) and microevolution is
"any change below the level of species." Matzke and Gross state that many creationist critics define
macroevolution as something that cannot be attained, as these critics describe any observed evolutionary
change as "just microevolution".[1]
[edit] See also

• List of transitional fossils
• Transitional fossil
• Microevolution
• Speciation
[edit] References
1. ^ a b c d Matzke, Nicholas J. and Paul R. Gross. 2006. Analyzing Critical Analysis: The
Fallback Antievolutionist Strategy. In Eugenie Scott and Glenn Branch, Not in Our Classrooms: Why
Intelligent Design is Wrong for Our Schools, Beacon Press, Boston ISBN 0807032786[page needed]
2. ^ Dobzhansky, Theodosius Grigorievich (1937). Genetics and the origin of species. New
York: Columbia Univ. Press. p. 12. LCCN 37-033383.
3. ^ Reznick DN, Ricklefs RE (February 2009). "Darwin's bridge between microevolution and
macroevolution". Nature 457 (7231): 837–42. doi:10.1038/nature07894. PMID 19212402.
4. ^ a b Introduction to Ecology (1983) - J.C. Emberlin, chapter 8
5. ^ a b c d Macroevolution: Its definition, Philosophy and History
6. ^ Gingerich, P. D. (1987). "Evolution and the fossil record: patterns, rates, and processes".
Canadian Journal of Zoology 65: 1053–1060. doi:10.1139/z87-169. edit
7. ^ Gingerich, P. D. (1987). "Evolution and the fossil record: patterns, rates, and processes".
Canadian Journal of Zoology 65: 1053–1060. doi:10.1139/z87-169. edit
8. ^ a b Kutschera U, Niklas KJ (June 2004). "The modern theory of biological evolution: an
expanded synthesis". Die Naturwissenschaften 91 (6): 255–76. doi:10.1007/s00114-004-0515-y.
PMID 15241603.
9. ^ Rieseberg LH, Willis JH (August 2007). "Plant speciation". Science 317 (5840): 910–4.
doi:10.1126/science.1137729. PMID 17702935.
10.^ Valentine JW, Jablonski D (2003). "Morphological and developmental macroevolution: a
paleontological perspective". The International Journal of Developmental Biology 47 (7-8): 517–22.
PMID 14756327. http://www.intjdevbiol.com/paper.php?doi=14756327.
11.^ Johnson NA, Porter AH (2001). "Toward a new synthesis: population genetics and
evolutionary developmental biology". Genetica 112-113: 45–58. doi:10.1023/A:1013371201773.
PMID 11838782.
12.^ Rice, W.R.; Hostert (1993). "Laboratory experiments on speciation: what have we learned
in 40 years". Evolution 47 (6): 1637–1653. doi:10.2307/2410209. http://links.jstor.org/sici?sici=0014-
3820(199312)47%3A6%3C1637%3ALEOSWH%3E2.0.CO%3B2-T. Retrieved 2008-05-19.
*Jiggins CD, Bridle JR (2004). "Speciation in the apple maggot fly: a blend of vintages?". Trends
Ecol. Evol. (Amst.) 19 (3): 111–4. doi:10.1016/j.tree.2003.12.008. PMID 16701238.
*Boxhorn, J (1995). "Observed Instances of Speciation". TalkOrigins Archive.
*Weinberg JR, Starczak VR, Jorg, D (1992). "Evidence for Rapid Speciation Following a Founder
Event in the Laboratory". Evolution 46 (4): 1214–20. doi:10.2307/2409766.
*Kirkpatrick, Mark; Virginie Ravigné (2002-03). "Speciation by Natural and Sexual Selection: Models
and Experiments". The American Naturalist 159 (3): S22–S35. doi:10.1086/338370. ISSN 00030147.
PMID 18707367. http://www.jstor.org/stable/3078919. Retrieved 2010-03-11.
13.^ CB901: No Macroevolution
14.^ CB902: Microevolution is distinct from macroevolution.
15.^ Myers 2006; NSTA 2007; IAP 2006; AAAS 2006; and Pinholster 2006; Ruling, Kitzmiller v.
Dover page 83
16.^ 29+ Evidences for Macroevolution: The Scientific Case for Common Descent , Douglas L.
Theobald, TalkOrigins Archive, Vers. 2.83, 2004, 12 Jan, 2004.
17.^ Laurence Moran (1993). "Evolution is a Fact and a Theory". TalkOrigins Archive.
http://www.talkorigins.org/faqs/evolution-fact.html. Retrieved 2008-02-07.

• Introduction to macroevolution
• Macroevolution as the common descent of all life
• Macroevolution in the 21st Century Macroevolution as an independent discipline.
• Macroevolution FAQ
Processes of
Population genetic
Evolutionary
developmental
concepts

Modes of

Retrieved from "http://en.wikipedia.org/wiki/Macroevolution"
Categories: Evolutionary biology

W000
Human evolution

Human evolution, or anthropogeny,[1] is the origin and evolution of Homo sapiens as a distinct
species from other hominids, great apes and placental mammals. The study of human evolution uses many
scientific disciplines, including physical anthropology, primatology, archaeology, linguistics and genetics.[2]
The term "human" in the context of human evolution refers to the genus Homo, but studies of human
evolution usually include other hominids, such as the Australopithecines, from which the genus Homo had
diverged by about 2.3 to 2.4 million years ago in Africa.[3][4] Scientists have estimated that humans
branched off from their common ancestor with chimpanzees about 5–7 million years ago. Several species and
subspecies of Homo evolved and are now extinct. These include Homo erectus, which inhabited Asia, and
Homo sapiens neanderthalensis, which inhabited Europe. Archaic Homo sapiens evolved between 400,000
and 250,000 years ago.
The dominant view among scientists concerning the origin of anatomically modern humans is the
"Out of Africa" or recent African origin hypothesis,[5][6][7][8] which argues that Homo sapiens arose in Africa
and migrated out of the continent around 50,000 to 100,000 years ago, replacing populations of Homo
erectus in Asia and Homo neanderthalensis in Europe. Scientists supporting the alternative multiregional
hypothesis argue that Homo sapiens evolved as geographically separate but interbreeding populations
stemming from a worldwide migration of Homo erectus out of Africa nearly 2.5 million years ago. This theory
has been contradicted by recent evidence, although it has been suggested that non Homo sapiens
Neanderthal genomes may have contributed about 4% of non-African heredity, and the recently discovered
Denisova hominin may have contributed 6% of the genome of Melanesians.[9][10]
==History of ideas== The word homo, the name of the biological genus to which humans belong, is
Latin for "human". It was chosen originally by Carolus Linnaeus in his classification system. The word
"human" is from the Latin humanus, the adjectival form of homo. The Latin "homo" derives from the Indo-
European root, dhghem, or "earth".[11]
Carolus Linnaeus and other scientists of his time also considered the great apes to be the closest
relatives of humans due to morphological and anatomical similarities. The possibility of linking humans with
earlier apes by descent only became clear after 1859 with the publication of Charles Darwin's On the Origin
of Species. This argued for the idea of the evolution of new species from earlier ones. Darwin's book did not
address the question of human evolution, saying only that "Light will be thrown on the origin of man and his
history".
Fossil Hominid Evolution Display at The Museum of Osteology, Oklahoma City, USA
The first debates about the nature of human evolution arose between Thomas Huxley and Richard
Owen. Huxley argued for human evolution from apes by illustrating many of the similarities and differences
between humans and apes, and did so particularly in his 1863 book Evidence as to Man's Place in Nature.
However, many of Darwin's early supporters (such as Alfred Russel Wallace and Charles Lyell) did not agree
that the origin of the mental capacities and the moral sensibilities of humans could be explained by natural
selection. Darwin applied the theory of evolution and sexual selection to humans when he published The
Descent of Man in 1871.[12]
A major problem was the lack of fossil intermediaries. It was only in the 1920s that such fossils were
discovered in Africa. In 1925, Raymond Dart described Australopithecus africanus. The type specimen was
the Taung Child, an Australopithecine infant discovered in a cave. The child's remains were a remarkably
well-preserved tiny skull and an endocranial cast of the individual's brain. Although the brain was small
(410 cm³), its shape was rounded, unlike that of chimpanzees and gorillas, and more like a modern human
brain. Also, the specimen showed short canine teeth, and the position of the foramen magnum was evidence
of bipedal locomotion. All of these traits convinced Dart that the Taung baby was a bipedal human ancestor,
a transitional form between apes and humans.
The classification of humans and their relatives has changed considerably over time. The gracile
Australopithecines are now thought to be ancestors of the genus Homo, the group to which modern humans
belong. Both Australopithecines and Homo sapiens are part of the tribe Hominini. Recent data suggests
Australopithecines were a diverse group and that A. africanus may not be a direct ancestor of modern
humans. Reclassification of Australopithecines that originally were split into either gracile or robust varieties
has put the latter into a family of its own, Paranthropus. Taxonomists place humans, Australopithecines and
related species in the same family as other great apes, in the Hominidae.'Italic text'Italic text'
Hominin species distributed through time edit

Note: 1e+06 years = 1 × 106 years = 1 million years ago = 1 Ma''''Italic text'Italic text'Italic text''Italic
text'''
Contents
[hide]
• 1 Before Homo
• 1.1 Evolution of the great apes
• 1.2 Divergence of the human lineage from other
Great Apes
• 2 Genus Homo
• 2.1 H. habilis and H. gautengensis
• 2.2 H. rudolfensis and H. georgicus
• 2.3 H. ergaster and H. erectus
• 2.4 H. cepranensis and H. antecessor
• 2.5 H. heidelbergensis
• 2.6 H. rhodesiensis, and the Gawis cranium
• 2.7 H. neanderthalensis
• 2.8 H. sapiens
• 2.9 H. floresiensis
• 2.10 Denisova hominin
• 2.11 Comparative table of Homo species
• 3 Use of tools
• 3.1 Stone tools
• 3.2 Modern humans and the "Great Leap Forward"
debate
[edit] Before Homo
For evolutionary history before primates, see evolution of mammals and evolutionary history of life.
[edit] Evolution of the great apes
Plesiadapis
The evolutionary history of the primates can be traced back 65 million years, as one of the oldest of
all surviving placental mammal groups. The oldest known primate-like mammal species, the Plesiadapis,
come from North America, but they were widespread in Eurasia and Africa during the tropical conditions of
the Paleocene and Eocene.
Notharctus
The beginning of modern climates was marked by the formation of the first Antarctic ice in the early
Oligocene around 30 million years ago. A primate from this time was Notharctus. Fossil evidence found in
Germany in the 1980s was determined to be about 16.5 million years old, some 1.5 million years older than
similar species from East Africa and challenging the original theory regarding human ancestry originating on
the African continent.
David Begun[13] says that these primates flourished in Eurasia and that the lineage leading to the
African apes and humans— including Dryopithecus—migrated south from Europe or Western Asia into Africa.
The surviving tropical population, which is seen most completely in the upper Eocene and lowermost
Oligocene fossil beds of the Fayum depression southwest of Cairo, gave rise to all living primates—lemurs of
Madagascar, lorises of Southeast Asia, galagos or "bush babies" of Africa, and the anthropoids; platyrrhines
or New World monkeys, and catarrhines or Old World monkeys and the great apes and humans.
The earliest known catarrhine is Kamoyapithecus from uppermost Oligocene at Eragaleit in the
northern Kenya Rift Valley, dated to 24 million years ago. Its ancestry is generally thought to be species
related to Aegyptopithecus, Propliopithecus, and Parapithecus from the Fayum, at around 35 million years
ago.[citation needed] In 2010, Saadanius was described as a close relative of the last common ancestor of
the crown catarrhines, and tentatively dated to 29–28 million years ago, helping to fill an 11-million-year gap
in the fossil record.[14]
Reconstructed tailless Proconsul skeleton

In the early Miocene, about 22 million years ago, the many kinds of arboreally adapted primitive
catarrhines from East Africa suggest a long history of prior diversification. Fossils at 20 million years ago
include fragments attributed to Victoriapithecus, the earliest Old World Monkey. Among the genera thought to
be in the ape lineage leading up to 13 million years ago are Proconsul, Rangwapithecus, Dendropithecus,
Limnopithecus, Nacholapithecus, Equatorius, Nyanzapithecus, Afropithecus, Heliopithecus, and
Kenyapithecus, all from East Africa. The presence of other generalized non-cercopithecids of middle
Miocene age from sites far distant— Otavipithecus from cave deposits in Namibia, and Pierolapithecus and
Dryopithecus from France, Spain and Austria—is evidence of a wide diversity of forms across Africa and the
Mediterranean basin during the relatively warm and equable climatic regimes of the early and middle
Miocene. The youngest of the Miocene hominoids, Oreopithecus, is from 9 million year old coal beds in Italy.
Molecular evidence indicates that the lineage of gibbons (family Hylobatidae) became distinct from
Great Apes between 18 and 12 million years ago, and that of orangutans (subfamily Ponginae) became
distinct from the other Great Apes at about 12 million years; there are no fossils that clearly document the
ancestry of gibbons, which may have originated in a so-far-unknown South East Asian hominoid population,
but fossil proto-orangutans may be represented by Ramapithecus from India and Griphopithecus from
Turkey, dated to around 10 million years ago.
[edit] Divergence of the human lineage from other Great Apes

Species close to the last common ancestor of gorillas, chimpanzees and humans may be
represented by Nakalipithecus fossils found in Kenya and Ouranopithecus found in Greece. Molecular
evidence suggests that between 8 and 4 million years ago, first the gorillas, and then the chimpanzees
(genus Pan) split off from the line leading to the humans; human DNA is approximately 98.4% identical to
that of chimpanzees when comparing single nucleotide polymorphisms (see human evolutionary genetics).
The fossil record of gorillas and chimpanzees is quite limited. Both poor preservation (rain forest soils tend to
be acidic and dissolve bone) and sampling bias probably contribute to this problem.
Other hominines likely adapted to the drier environments outside the equatorial belt, along with
antelopes, hyenas, dogs, pigs, elephants, and horses. The equatorial belt contracted after about 8 million
years ago. Fossils of these hominans - the species in the human lineage following divergence from the
chimpanzees - are relatively well known.
The earliest are Sahelanthropus tchadensis (7 Ma) and Orrorin tugenensis (6 Ma), followed by:
• Ardipithecus (5.5–4.4 Ma), with species Ar. kadabba and Ar. ramidus;
• Australopithecus (4–1.8 Ma), with species Au. anamensis, Au. afarensis, Au. africanus, Au.
bahrelghazali, Au. garhi, and Au. sediba;
• Kenyanthropus (3–2.7 Ma), with species Kenyanthropus platyops;
• Paranthropus (3–1.2 Ma), with species P. aethiopicus, P. boisei, and P. robustus;
• Homo (2 Ma–present), with species Homo habilis, Homo rudolfensis, Homo ergaster, Homo
georgicus, Homo antecessor, Homo cepranensis, Homo erectus, Homo heidelbergensis, Homo
rhodesiensis, Homo neanderthalensis, Homo sapiens idaltu, Archaic Homo sapiens, Homo
floresiensis.
[edit] Genus Homo

Homo sapiens is the only extant species of its genus, Homo. While some other, extinct Homo
species might have been ancestors of Homo sapiens, many were likely our "cousins", having speciated away
from our ancestral line.[15][16] There is not yet a consensus as to which of these groups should count as
separate species and which as subspecies. In some cases this is due to the dearth of fossils, in other cases it
is due to the slight differences used to classify species in the Homo genus.[16] The Sahara pump theory
(describing an occasionally passable "wet" Sahara Desert) provides an explanation of the early variation in
the genus Homo.
Based on archaeological and paleontological evidence, it has been possible to infer, to some extent,
the ancient dietary practices of various Homo species and to study the role of diet in physical and behavioral
evolution within Homo.[17][18][19][20][21]
[edit] H. habilis and H. gautengensis

Homo habilis lived from about 2.4 to 1.4 Ma. Homo habilis evolved in South and East Africa in the
late Pliocene or early Pleistocene, 2.5–2 Ma, when it diverged from the Australopithecines. Homo habilis had
smaller molars and larger brains than the Australopithecines, and made tools from stone and perhaps animal
bones. One of the first known hominids, it was nicknamed 'handy man' by its discoverer, Louis Leakey due to
its association with stone tools. Some scientists have proposed moving this species out of Homo and into
Australopithecus due to the morphology of its skeleton being more adapted to living on trees rather than to
moving on two legs like Homo sapiens.[22]
It was considered to be the first species of the genus Homo until May 2010, when a new species,
Homo gautengensis was discovered in South Africa, that most likely arose earlier than Homo habilis.[23]
[edit] H. rudolfensis and H. georgicus

These are proposed species names for fossils from about 1.9–1.6 Ma, the relation of which with
Homo habilis is not yet clear.
• Homo rudolfensis refers to a single, incomplete skull from Kenya. Scientists have suggested
that this was another Homo habilis, but this has not been confirmed.[24]
• Homo georgicus, from Georgia, may be an intermediate form between Homo habilis and
Homo erectus,[25] or a sub-species of Homo erectus.[26]
[edit] H. ergaster and H. erectus
One current view of the temporal and geographical distribution of hominid populations.[27] Other
interpretations differ mainly in the taxonomy and geographical distribution of hominid species.
The first fossils of Homo erectus were discovered by Dutch physician Eugene Dubois in 1891 on the
Indonesian island of Java. He originally gave the material the name Pithecanthropus erectus based on its
morphology that he considered to be intermediate between that of humans and apes.[28] Homo erectus (H
erectus) lived from about 1.8 Ma to about 70,000 years ago (which would indicate that they were probably
wiped out by the Toba catastrophe; however, Homo erectus soloensis and Homo floresiensis survived it).
Often the early phase, from 1.8 to 1.25 Ma, is considered to be a separate species, Homo ergaster, or it is
seen as a subspecies of Homo erectus, Homo erectus ergaster. In the early Pleistocene, 1.5–1 Ma, in Africa,
Asia, and Europe, some populations of Homo habilis are thought to have evolved larger brains and made
more elaborate stone tools; these differences and others are sufficient for anthropologists to classify them as
a new species, Homo erectus. In addition Homo erectus was the first human ancestor to walk truly upright.
[29] This was made possible by the evolution of locking knees and a different location of the foramen
magnum (the hole in the skull where the spine enters). They may have used fire to cook their meat.
See also: Control of fire by early humans
A famous example of Homo erectus is Peking Man; others were found in Asia (notably in Indonesia),
Africa, and Europe. Many paleoanthropologists now use the term Homo ergaster for the non-Asian forms of
this group, and reserve Homo erectus only for those fossils that are found in Asia and meet certain skeletal
and dental requirements which differ slightly from H. ergaster.
[edit] H. cepranensis and H. antecessor

These are proposed as species that may be intermediate between H. erectus and H.
heidelbergensis.
• H. antecessor is known from fossils from Spain and England that are dated 1.2 Ma–500 ka.
[30][31]
• H. cepranensis refers to a single skull cap from Italy, estimated to be about 800,000 years
old.[32]
[edit] H. heidelbergensis
H. heidelbergensis (Heidelberg Man) lived from about 800,000 to about 300,000 years ago. Also
proposed as Homo sapiens heidelbergensis or Homo sapiens paleohungaricus.[33]
[edit] H. rhodesiensis, and the Gawis cranium

• H. rhodesiensis, estimated to be 300,000–125,000 years old. Most current experts believe
Rhodesian Man to be within the group of Homo heidelbergensis, though other designations such as
Archaic Homo sapiens and Homo sapiens rhodesiensis have also been proposed.
• In February 2006 a fossil, the Gawis cranium, was found which might possibly be a species
intermediate between H. erectus and H. sapiens or one of many evolutionary dead ends. The skull
from Gawis, Ethiopia, is believed to be 500,000–250,000 years old. Only summary details are known,
and no peer reviewed studies have been released by the finding team. Gawis man's facial features
suggest its being either an intermediate species or an example of a "Bodo man" female.[34]
[edit] H. neanderthalensis
Le Ferrassie Neanderthal skull (cast)

H. neanderthalensis lived from 400,000[35] to about 30,000 years ago. Also proposed as Homo
sapiens neanderthalensis.[36] Evidence from sequencing mitochondrial DNA indicated that no significant
gene flow occurred between H. neanderthalensis and H. sapiens, and, therefore, the two were separate
species that shared a common ancestor about 660,000 years ago.[37][38] In 1997, Mark Stoneking stated:
"These results [based on mitochondrial DNA extracted from Neanderthal bone] indicate that Neanderthals
did not contribute mitochondrial DNA to modern humans… Neanderthals are not our ancestors". Subsequent
investigation of a second source of Neanderthal DNA supported these findings.[39]
However, the 2010 sequencing of the Neanderthal genome indicated that Neanderthals did indeed
interbreed with H. sapiens circa 75,000 BC (after H. sapiens moved out from Africa, but before they
separated into Europe, the Middle East, and Asia).[40] Nearly all modern humans have 1% to 4% of their
DNA derived from Neanderthal DNA.[40] This 1–4% bit of DNA is only present in non-African humans.[40]
However, supporters of the multiregional hypothesis point to recent studies indicating non-African nuclear
DNA heritage dating to one Ma,[41] although the reliability of these studies has been questioned.[42]
Competition from Homo sapiens probably contributed to Neanderthal extinction.[43][44] They could have
coexisted in Europe for as long as 10,000 years.[45]
[edit] H. sapiens
Main article: Early Homo sapiens
H. sapiens (the adjective sapiens is Latin for "wise" or "intelligent") have lived from about 250,000
years ago to the present. Between 400,000 years ago and the second interglacial period in the Middle
Pleistocene, around 250,000 years ago, the trend in skull expansion and the elaboration of stone tool
technologies developed, providing evidence for a transition from H. erectus to H. sapiens. The direct
evidence suggests there was a migration of H. erectus out of Africa, then a further speciation of H. sapiens
from H. erectus in Africa. A subsequent migration within and out of Africa eventually replaced the earlier
dispersed H. erectus. This migration and origin theory is usually referred to as the recent single origin or Out
of Africa theory. Current evidence does not preclude some multiregional evolution or some admixture of the
migrant H. sapiens with existing Homo populations. This is a hotly debated area of paleoanthropology.
Current research has established that humans are genetically highly homogenous; that is, the DNA
of individuals is more alike than usual for most species, which may have resulted from their relatively recent
evolution or the possibility of a population bottleneck resulting from cataclysmic natural events such as the
Toba catastrophe.[46][47][48] Distinctive genetic characteristics have arisen, however, primarily as the result
of small groups of people moving into new environmental circumstances. These adapted traits are a very
small component of the Homo sapiens genome, but include various characteristics such as skin color and
nose form, in addition to internal characteristics such as the ability to breathe more efficiently at high
altitudes.
H. sapiens idaltu, from Ethiopia, is an extinct sub-species who lived about 160,000 years ago.
[edit] H. floresiensis
Main article: Homo floresiensis
H. floresiensis, which lived from approximately 100,000 to 12,000 before present, has been
nicknamed hobbit for its small size, possibly a result of insular dwarfism.[49] H. floresiensis is intriguing both
for its size and its age, being a concrete example of a recent species of the genus Homo that exhibits derived
traits not shared with modern humans. In other words, H. floresiensis share a common ancestor with modern
humans, but split from the modern human lineage and followed a distinct evolutionary path. The main find
was a skeleton believed to be a woman of about 30 years of age. Found in 2003 it has been dated to
approximately 18,000 years old. The living woman was estimated to be one meter in height, with a brain
volume of just 380 cm3 (considered small for a chimpanzee and less than a third of the H. sapiens average of
1400 cm3).
However, there is an ongoing debate over whether H. floresiensis is indeed a separate species.[50]
Some scientists presently believe that H. floresiensis was a modern H. sapiens suffering from pathological
dwarfism.[51] This hypothesis is supported in part, because some modern humans who live on Flores, the
island where the skeleton was found, are pygmies. This coupled with pathological dwarfism, it is argued,
could indeed create a hobbit-like human. The other major attack on H. floresiensis is that it was found with
tools only associated with H. sapiens.[51]
The hypothesis of pathological dwarfism, however, fails to explain additional anatomical features that
are unlike those of modern humans (diseased or not) but much like those of ancient members of our genus.
Aside from cranial features, these features include the form of bones in the wrist, forearm, shoulder, knees,
and feet.
[edit] Denisova hominin

Main article: Denisova hominin
In 2008, archaeologists working at the site of Denisova Cave in the Altai Mountains of Siberia
uncovered a small bone fragment from the fifth finger of a juvenile hominin, dubbed the "X-woman" (referring
to the maternal descent of mitochondrial DNA),[52] or the Denisova hominin. Artifacts, including a bracelet,
excavated in the cave at the same level were carbon dated to around 40,000 BP. As DNA had survived in the
fossil fragment due to the cool climate of the Denisova Cave, a team of scientists from the Max Planck
Institute for Evolutionary Anthropology in Leipzig, Germany sequenced mtDNA extracted from the fragment.
[53]
The analysis indicated that modern humans, Neanderthals, and the Denisova hominin last shared a
common ancestor around 1 million years ago.[54] Modern humans are known to have overlapped with
Neanderthals in Europe for more than 10,000 years, and the discovery raises the possibility that
Neanderthals, modern humans and the Denisovan hominin may have co-existed together.
The DNA analysis further indicated that this new hominin species was the result of an early migration
out of Africa, distinct from the later out-of-Africa migrations associated with Neanderthals and modern
humans, but also distinct from the earlier African exodus of Homo erectus.[54] Professor Chris Stringer,
human origins researcher at London's Natural History Museum and one of the leading proponents of the
recent single-origin hypothesis, remarked: "This new DNA work provides an entirely new way of looking at
the still poorly understood evolution of humans in central and eastern Asia." Pääbo noted that the existence
of this distant branch creates a much more complex picture of humankind during the Late Pleistocene.[52]
[edit] Comparative table of Homo species

Comparative table of Homo species view · talk · edit
Lived when Lived Adult Adult Crani
Species
(Ma) where height mass capacity (cm³)
1.75 m 90 kg
H. antecessor 1.2 – 0.8 Spain 1,000
(5.7 ft) (200 lb)
H. cepranensis 0.9 – 0.8? Italy 1,000
Africa,
850
Eurasia (Java, 1.8 m 60 kg
H. erectus 1.5 – 0.2 (early) – 1,100
China, India, (5.9 ft) (130 lb)
(late)
Caucasus)
Eastern
1.9 m
H. ergaster 1.9 – 1.4 and Southern 700–85
(6.2 ft)
Africa
1.0 m 25 kg
H. floresiensis 0.10? – 0.012 Indonesia 400
(3.3 ft) (55 lb)
South 1.0 m
H. gautengensis >2 – 0.6
Africa (3.3 ft)
H. georgicus 1.8 Georgia 600
1.0–1.5 33–55 kg
H. habilis 2.3 – 1.4 Africa 510–66
m (3.3–4.9 ft) (73–120 lb)
Europe, 1.8 m 60 kg 1,100–

H. heidelbergensis 0.6 – 0.35
Africa, China (5.9 ft) (130 lb) 1,400
55–70 kg
Europe, 1.6 m 1,200–
H. neanderthalensis 0.35 – 0.03 (120–150 lb)
Western Asia (5.2 ft) 1,900
(heavily built)
H. rhodesiensis 0.3 – 0.12 Zambia 1,300
H. rudolfensis 1.9 Kenya
H. sapiens idaltu 0.16 – 0.15 Ethiopia 1,450
H. sapiens sapiens 1.4–1.9 50–100 1,000–

0.2 – present Worldwide
(modern humans) m (4.6–6.2 ft) kg (110–220 lb) 1,850
[edit] Use of tools
"A sharp rock", an Oldowan pebble tool, the most basic of human stone tools
Fire, one of the greatest human discoveries
An Acheulean hand axe, the pinnacle of Homo erectus stone working

Venus of Willendorf, an example of Paleolithic art
See also: Hunting hypothesis
Using tools has been interpreted as a sign of intelligence, and it has been theorized that tool use may
have stimulated certain aspects of human evolution—most notably the continued expansion of the human
brain. Paleontology has yet to explain the expansion of this organ over millions of years despite being
extremely demanding in terms of energy consumption. The brain of a modern human consumes about 20
watts (400 kilocalories per day), which is one fifth of the energy consumption of a human body. Increased tool
use would allow hunting for energy-rich meat products, and would enable processing more energy-rich plant
products. Researchers have suggested that early hominids were thus under evolutionary pressure to
increase their capacity to create and use tools.[55]
Precisely when early humans started to use tools is difficult to determine, because the more primitive
these tools are (for example, sharp-edged stones) the more difficult it is to decide whether they are natural
objects or human artifacts. There is some evidence that the australopithecines (4 Ma) may have used broken
bones as tools, but this is debated.
It should be noted that many species make and use tools, but it is the human species that dominates
the areas of making and using more complex tools. The oldest known tools are the "Oldowan stone tools"
from Ethiopia. It was discovered that these tools are from 2.5 to 2.6 million years old, which predates the
earliest known "Homo" species. There is no known evidence that any "Homo" specimens appeared by 2.5
Ma. A Homo fossil was found near some Oldowan tools, and its age was noted at 2.3 million years old,
suggesting that maybe the Homo species did indeed create and use these tools. It is surely possible, but not
solid evidence. Bernard Wood noted that "Paranthropus" coexisted with the early Homo species in the area
of the "Oldowan Industrial Complex" over roughly the same span of time. Although there is no direct evidence
that points to Paranthropus as the tool makers, their anatomy lends to indirect evidence of their capabilities in
this area. Most paleoanthropologists agree that the early "Homo" species were indeed responsible for most
of the Oldowan tools found. They argue that when most of the Oldowan tools were found in association with
human fossils, Homo was always present, but Paranthropus was not.[56]
In 1994, Randall Susman used the anatomy of opposable thumbs as the basis for his argument that
both the Homo and Paranthropus species were toolmakers. He compared bones and muscles of human and
chimpanzee thumbs, finding that humans have 3 muscles that chimps lack. Humans also have thicker
metacarpals with broader heads, making the human hand more successful at precision grasping than the
chimpanzee hand. Susman defended that modern anatomy of the human thumb is an evolutionary response
to the requirements associated with making and handling tools and that both species were indeed
toolmakers.[56]
[edit] Stone tools

Stone tools are first attested around 2.6 Ma, when H. habilis in Eastern Africa used so-called pebble
tools, choppers made out of round pebbles that had been split by simple strikes.[57] This marks the
beginning of the Paleolithic, or Old Stone Age; its end is taken to be the end of the last Ice Age, around
10,000 years ago. The Paleolithic is subdivided into the Lower Paleolithic (Early Stone Age, ending around
350,000–300,000 years ago), the Middle Paleolithic (Middle Stone Age, until 50,000–30,000 years ago), and
the Upper Paleolithic.
The period from 700,000–300,000 years ago is also known as the Acheulean, when H. ergaster (or
erectus) made large stone hand-axes out of flint and quartzite, at first quite rough (Early Acheulian), later
"retouched" by additional, more subtle strikes at the sides of the flakes. After 350,000 BP (Before Present)
the more refined so-called Levallois technique was developed. It consisted of a series of consecutive strikes,
by which scrapers, slicers ("racloirs"), needles, and flattened needles were made.[57] Finally, after about
50,000 BP, ever more refined and specialized flint tools were made by the Neanderthals and the immigrant
Cro-Magnons (knives, blades, skimmers). In this period they also started to make tools out of bone.
[edit] Modern humans and the "Great Leap Forward" debate
See also: Behavioral modernity
Until about 50,000–40,000 years ago the use of stone tools seems to have progressed stepwise.
Each phase (H. habilis, H. ergaster, H. neanderthalensis) started at a higher level than the previous one, but
once that phase started further development was slow. These Homo species were culturally conservative,
but after 50,000 BC modern human culture started to change at a much greater speed. Jared Diamond,
author of The Third Chimpanzee, and some anthropologists characterize this as a "Great Leap Forward".
Modern humans started burying their dead, making clothing out of hides, developing sophisticated
hunting techniques (such as using trapping pits or driving animals off cliffs), and engaging in cave painting.
[58] As human culture advanced, different populations of humans introduced novelty to existing technologies:
artifacts such as fish hooks, buttons and bone needles show signs of variation among different populations of
humans, something that had not been seen in human cultures prior to 50,000 BP. Typically, H.
neanderthalensis populations do not vary in their technologies.
Among concrete examples of Modern human behavior, anthropologists include specialization of
tools, use of jewellery and images (such as cave drawings), organization of living space, rituals (for example,
burials with grave gifts), specialized hunting techniques, exploration of less hospitable geographical areas,
and barter trade networks. Debate continues as to whether a "revolution" led to modern humans ("the big
bang of human consciousness"), or whether the evolution was more gradual.[59]
[edit] Models of human evolution

Today, all humans belong to one population of Homo sapiens sapiens, undivided by species barrier.
However, according to the "Out of Africa" model this is not the first species of hominids: the first species of
genus Homo, Homo habilis, evolved in East Africa at least 2 Ma, and members of this species populated
different parts of Africa in a relatively short time. Homo erectus evolved more than 1.8 Ma, and by 1.5 Ma had
spread throughout the Old World.
Anthropologists have been divided as to whether current human population evolved as one
interconnected population (as postulated by the Multiregional Evolution hypothesis), or evolved only in East
Africa, speciated, then migrated out of Africa and replaced human populations in Eurasia (called the "Out of
Africa" Model or the "Complete Replacement" Model).
[edit] Multiregional model

Main article: Multiregional hypothesis
Multiregional evolution, a model to account for the pattern of human evolution , was proposed by
Milford H. Wolpoff[60] in 1988.[61] Multiregional evolution holds that human evolution from the beginning of
the Pleistocene 2.5 million years BP to the present day has been within a single, continuous human species,
evolving worldwide to modern Homo sapiens.
According to the multiregional hypothesis, fossil and genomic data are evidence for worldwide
human evolution and contradict the recent speciation postulated by the Recent African origin hypothesis. The
fossil evidence was insufficient for Richard Leakey to resolve this debate.[62] Studies of haplogroups in Y-
chromosomal DNA and mitochondrial DNA have largely supported a recent African origin.[63] Evidence from
autosomal DNA also supports the Recent African origin. However the presence of archaic admixture in
modern humans remains a possibility and has been suggested by some studies.[64]
[edit] Out of Africa
See also: Recent single origin hypothesis and Early human migrations
According to the Out of Africa model, developed by Chris Stringer and Peter Andrews, modern H.
sapiens evolved in Africa 200,000 years ago. Homo sapiens began migrating from Africa between 70,000 –
50,000 years ago and eventually replaced existing hominid species in Europe and Asia.[65][66] Out of Africa
has gained support from research using mitochondrial DNA (mtDNA). After analysing genealogy trees
constructed using 133 types of mtDNA, researchers concluded that all were descended from a woman from
Africa, dubbed Mitochondrial Eve. Out of Africa is also supported by the fact that mitochondrial genetic
diversity is highest among African populations.[67]
There are differing theories on whether there was a single exodus or several. A multiple dispersal
model involves the Southern Dispersal theory,[68] which has gained support in recent years from genetic,
linguistic and archaeological evidence. In this theory, there was a coastal dispersal of modern humans from
the Horn of Africa around 70,000 years ago. This group helped to populate Southeast Asia and Oceania,
explaining the discovery of early human sites in these areas much earlier than those in the Levant. A second
wave of humans dispersed across the Sinai peninsula into Asia, resulting in the bulk of human population for
Eurasia. This second group possessed a more sophisticated tool technology and was less dependent on
coastal food sources than the original group. Much of the evidence for the first group's expansion would have
been destroyed by the rising sea levels at the end of the Holocene era.[68] The multiple dispersal model is
contradicted by studies indicating that the populations of Eurasia and the populations of Southeast Asia and
Oceania are all descended from the same mitochondrial DNA lineages, which support a single migration out
of Africa that gave rise to all non-African populations.[69]
The broad study of African genetic diversity headed by Dr. Sarah Tishkoff found the San people to
express the greatest genetic diversity among the 113 distinct populations sampled, making them one of 14
"ancestral population clusters". The research also located the origin of modern human migration in south-
western Africa, near the coastal border of Namibia and Angola.[70]
According to the Toba catastrophe theory to which some anthropologists and archeologists
subscribe, the supereruption of Lake Toba on Sumatra island in Indonesia roughly 70,000 years ago had
global consequences,[71] killing most humans then alive and creating a population bottleneck that affected
the genetic inheritance of all humans today.[72]
[edit] Recent and current human evolution

See also: Transhuman
Natural selection is being observed in contemporary human populations, with recent findings
demonstrating the population which is at risk of the severe debilitating disease kuru has significant over-
representation of an immune variant of the prion protein gene G127V versus non-immune alleles. Scientists
postulate one of the reasons for the rapid selection of this genetic variant is the lethality of the disease in non-
immune persons.[73][74] Other reported evolutionary trends in other populations include a lengthening of the
reproductive period, reduction in cholesterol levels, blood glucose and blood pressure.[75]
In their 2009 book The 10,000 Year Explosion, Gregory Cochran and Henry Harpending argue that
human evolution has accelerated since and as a result of the development of agriculture and civilization
since some 50,000 years ago, and that there are consequently substantial genetic differences between
different current human populations.
[edit] Genetics
Main articles: Human evolutionary genetics and Human genetic variation
Human evolutionary genetics studies how one human genome differs from the other, the evolutionary
past that gave rise to it, and its current effects. Differences between genomes have anthropological, medical
and forensic implications and applications. Genetic data can provide important insight into human evolution.
[edit] Notable human evolution researchers

See also: Category:Human evolution theorists
• Robert Broom, a Scottish physician and palaeontologist whose work on South Africa led to
the discovery and description of the Paranthropus genus of hominins, and of "Mrs. Ples"
• Raymond Dart, an Australian anatomist and palaeoanthropologist, whose work at Taung, in
South Africa, led to the discovery of Australopithecus africanus
• Charles Darwin, a British naturalist who documented considerable evidence that species
originate through evolutionary change
• Henry McHenry, an American anthropologist who specializes in studies of human evolution,
the origins of bipedality, and paleoanthropology
• Donald Johanson, credited with the discovery of Australopithecus afarensis
• Jeffrey Laitman, an American anatomist and physical anthropologist whose work has
explored the evolution of the vocal tract and speech
• Louis Leakey, an African archaeologist and naturalist whose work was important in
establishing human evolutionary development in Africa
• Mary Leakey, a British archaeologist and anthropologist whose discoveries in Africa include
the Laetoli footprints
• Richard Leakey, an African paleontologist and archaeologist, son of Louis and Mary Leakey
• Svante Pääbo, a Swedish biologist specializing in evolutionary genetics
• David Pilbeam, a paleoanthropologist, researcher and writer on a range of topics involving
human and primate evolution.
• Jeffrey H. Schwartz, an American physical anthropologist and professor of biological
anthropology
• Chris Stringer, anthropologist, leading proponent of the recent single origin hypothesis
• Alan Templeton, geneticist and statistician, proponent of the multiregional hypothesis
• Philip V. Tobias, a South African palaeoanthropologist is one of the world's leading
authorities on the evolution of humankind
• Erik Trinkaus, a prominent American paleoanthropologist and expert on Neanderthal biology
and human evolution
• Milford H. Wolpoff, an American paleoanthropologist who is the leading proponent of the
multiregional evolution hypothesis.
[edit] Species list

This list is in chronological order across the page by genus.
• Sahelanthropus • Australopithecus • Homo
• Sahela • Australo • Homo
nthropus pithecus gautengensis
tchadensis anamensis • Homo
• Orrorin • Australo habilis
• Orrorin pithecus • Homo
tugenensis afarensis rudolfensis
• Ardipithecus • Australo • Homo
• Ardipith pithecus ergaster
ecus kadabba bahrelghazali • Homo
• Ardipith • Australo georgicus
ecus ramidus pithecus • Homo
africanus erectus
• Australo • Homo
pithecus garhi cepranensis
• Australo • Homo
pithecus sediba antecessor
• Paranthropus • Homo
• Paranthr heidelbergensis
opus • Homo
aethiopicus rhodesiensis
• Paranthr • Homo
opus boisei neanderthalensis
• Paranthr • Homo
opus robustus sapiens idaltu
• Kenyanthropus • Homo
• Kenyant sapiens (Cro-
hropus platyops Magnon)
• Homo
sapiens sapiens
• Homo
floresiensis
[edit] See also

• List of human evolution
fossils
• Timeline of human
evolution • Evolutionary psychology
• Archaeogenetics • History of Earth
• Dual inheritance theory • Hominid intelligence
• Dysgenics • Human behavioral ecology
• Evolutionary anthropology • Human skeletal changes due to
• Evolutionary medicine bipedalism
• Evolutionary neuroscience • Human vestigiality
• Evolution of human • Ida (fossil)
intelligence • Pan prior
• Evolution of morality • Physical anthropology
• Evolutionary origin of • Sahara pump theory
religions • Sexual selection in human evolution
• Sociocultural evolution
[edit] References
[edit] Notes
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Main article: Bibliography of evolution and human behavior
• Hill, Andrew; Ward, Steven (1988). "Origin of the hominidae: The record of african large
hominoid evolution between 14 my and 4 my". Yearbook of Physical Anthropology 31 (59): 49–83.
doi:10.1002/ajpa.1330310505
• Alexander, R. D. (1990). "How Did Humans Evolve? Reflections on the Uniquely Unique
Species". University of Michigan Museum of Zoology Special Publication (University of Michigan
Museum of Zoology) (1): 1–38. http://insects.ummz.lsa.umich.edu/pdfs/Alexander1990.pdf.
• Flinn, M. V., Geary, D. C., & Ward, C. V. (2005). Ecological dominance, social competition,
and coalitionary arms races: Why humans evolved extraordinary intelligence. Evolution and Human
Behavior, 26, 10-46. Full text.PDF (345 KB)
• edited by Steve Jones, Robert Martin, and David Pilbeam ; foreword by Richard Dawkins.
(1994). Jones, S., Martin, R., & Pilbeam, D.. ed. The Cambridge Encyclopedia of Human Evolution.
Cambridge: Cambridge University Press. ISBN 978-0-521-32370-3. Also ISBN 978-0-521-46786-5
• Wolfgang Enard et al. (2002-08-22). "Molecular evolution of FOXP2, a gene involved in
speech and language". Nature 418: 870.
• DNA Shows Neandertals Were Not Our Ancestors
• J. W. IJdo, A. Baldini, D. C. Ward, S. T. Reeders, R. A. Wells (October 1991). "Origin of
human chromosome 2: An ancestral telomere-telomere fusion" (PDF). Genetics 88: 9051–9055.
http://www.pnas.org/cgi/reprint/88/20/9051.pdf. —two ancestral ape chromosomes fused to give rise
to human chromosome 2.
• Ovchinnikov, et al.; Götherström, Anders; Romanova, Galina P.; Kharitonov, Vitaliy M.;
Lidén, Kerstin; Goodwin, William (2000). "Molecular analysis of Neanderthal DNA from the Northern
Caucasus". Nature 404 (6777): 490. doi:10.1038/35006625. PMID 10761915.
• Heizmann, Elmar P J, Begun, David R (2001). "The oldest Eurasian hominoid". Journal of
Human Evolution 41 (5): 463. doi:10.1006/jhev.2001.0495. PMID 11681862.
• BBC: Finds test human origins theory. 2007-08-08 Homo habilis and Homo erectus are sister
species that overlapped in time.

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Anthropology
W000
Amniote

Amniote
Amniotes
Temporal range: Mississippian–Recent, 340–0 Ma
PreЄ
g
A baby tortoise emerges from an amniotic egg
Kingdom: Animalia
Phylum: Chordata
Superclass: Tetrapoda
(unranked): Reptiliomorpha
Amniota
(unranked):
Haeckel, 1866
Clades
• Synapsida
• Sauropsida
The amniotes are a group of tetrapod vertebrates (four-footed animals with backbones or spinal
columns) that have a terrestrially adapted egg. They include mammals (synapsids) and sauropsids (reptiles
and birds), as well as their fossil ancestors. Amniote embryos, whether laid as eggs or carried by the female,
are protected and aided by several extensive membranes. In eutherian mammals (such as humans), these
membranes include the amniotic sac that surrounds the fetus. These embryonic membranes, and the lack of
a larval stage, distinguish amniotes from tetrapod amphibians.[1]
The first amniotes (referred to as "basal amniotes" or "stem amniotes"), such as Casineria,
resembled small lizards and had evolved within reptiliomorphs which were amphibian about 340 million years
ago, in the Carboniferous geologic period. Their eggs could survive out of the water, allowing amniotes to
branch out into drier environments. The eggs could also "breathe" and cope with waste, allowing the eggs
and the amniotes themselves to evolve into larger forms. The amniotes spread across the globe and became
the dominant land vertebrates.
Very early in the evolutionary history of amniotes, basal amniotes evolved into two main lines of
amniotes, the synapsids and the sauropsids, both of which persist into the modern era. The oldest known
fossil synapsid is Protoclepsydrops from about 320 million years ago, while the oldest known sauropsid is
probably Paleothyris, in the order Captorhinida, from the Middle Pennsylvanian epoch (ca. 306-312 million
years ago).
Contents
[hide]
• 1 Description
• 2 Definition and classification
• 2.1 Traditional classification
• 2.2 Phylogenetic classification
• 2.3 Cladogram of Amniotes
• 3 References
[edit] Description
Amniotes can be characterized in part by embryonic development that includes the formation of
several extensive membranes, the amnion, chorion, and allantois. Amniotes develop directly into a (typically)
terrestrial form with limbs and a thick stratified epithelium, rather than first entering a feeding larval tadpole
stage followed by metamorphosis as in amphibians. In amniotes the transition from a two-layered periderm to
cornified epithelium is triggered by thyroid hormone during embryonic development, rather than
metamorphosis.[2] The unique embryonic features of amniotes may reflect specializations of eggs to survive
drier environments, or the massive size and yolk content of eggs evolved for direct development to a larger
size.
Anatomy of an amniotic egg

1. Eggshell
2. Outer membrane
3. Inner membrane
4. Chalaza
5. Exterior albumen (outer thin albumen)
6. Middle albumen (inner thick albumen)
7. Vitelline membrane
8. Nucleus of Pander
9. Germinal disk (blastoderm)
10. Yellow yolk
11. White yolk
12. Internal albumen
13. Chalaza
14. Air cell
15. Cuticula
Features of amniotes evolved for survival on land include a sturdy but porous leathery or hard
eggshell and an allantois evolved to facilitate respiration while providing a reservoir for disposal of wastes.
Their kidneys and large intestines are also well-suited to water retention. Most mammals do not lay eggs, but
corresponding structures may be found inside the placenta.
The first amniotes, such as Casineria kiddi, which lived about 340 million years ago, evolved from
amphibian reptiliomorphs and resembled small lizards. Their eggs were small and covered with a membrane,
not a hard shell like most modern amniote eggs. Although some modern amphibians lay eggs on land, with
or without significant protection, they all lack advanced traits like an amnion. This kind of egg only became
possible with internal fertilization. The outer membrane, a soft shell, evolved as a protection against the
harsher environments on land, as species evolved to lay their eggs on land where they were safer than in the
water. One can assume the ancestors of the amniotes laid their eggs in moist places, as such modest-sized
animals would not have difficulty finding depressions under fallen logs or other suitable places in the ancient
forests, and dry conditions were probably not the main reason why the soft shell emerged.[3]
In fish and amphibians there is only one inner membrane, also called an embryonic membrane. In
amniotes the inner anatomy of the egg has evolved further and new structures have developed to take care
of the gas exchanges between the embryo and the atmosphere, as well as dealing with the waste problems.
To grow a thicker and tougher shell required new ways to supply the embryo with oxygen, as diffusion alone
was not enough. After the egg developed these structures, further sophistication allowed amniotes to lay
much bigger eggs in much drier habitats. Bigger eggs allowed for bigger offspring, and bigger adults could
produce bigger eggs, so amniotes grew bigger than their ancestors. Real growth was not possible, however,
until they stopped relying on small invertebrates as their main food source and started to eat plants or other
vertebrates, or returned to the water. New habits and heavier bodies meant further evolution for the
amniotes, both in behavior and anatomy.
There are three main lines of amniotes, which may be distinguished by the structure of the skull and
in particular the number of temporal fenestrae (openings) behind the eye. In anapsids (turtles) there are
none, in synapsids (mammals and their extinct relatives) there is one, and in most diapsids (non-anapsid
reptiles, including dinosaurs and birds) there are two.[4]
The skeletal remains of amniotes can be identified from their Labyrinthodont ancestors by their
having at least two pairs of sacral ribs, a sternum in the pectoral girdle (some amniotes have lost it) and an
astragalus bone in the ankle.[5]
[edit] Definition and classification

Amniota was first formally described by embryologist Ernst Haeckel in 1866 on the presence of the
amnion, hence the name. A problem with this definition is that the trait (apomorphy) in question does not
fossilize, and the status of fossil forms has to be inferred from other traits. Thus Jacques Gauthier and
colleagues forwarded a definition of Amniota in 1988 as "the most recent common ancestor of extant
mammals and reptiles, and all its descendants".[5] Gauthiers definition being a node-based crown group, his
definition of the group has a slightly different content than the group defined as biological amniotes
(apomorphy-based clade).[6]
[edit] Traditional classification

Classifications of the amniotes have traditionally recognised three classes based on major traits and
physiology:
• Class Reptilia (reptiles)
• Subclass Anapsida ("proto-reptiles", possibly including turtles)
• Subclass Diapsida (majority of reptiles, progenitors of birds)
• Subclass Synapsida (mammal-like reptiles, progenitors of mammals)
• Class Aves (birds)
• Subclass Neornithes (all modern birds, several extinct subclasses recognised)
• Class Mammalia (mammals)
• Subclass Monotremata (egg-laying mammals)
• Subclass Theria (marsupials and placental mammals)
This rather orderly scheme is the one most commonly found in popular and basic scientific works. It
has come under critique from cladistics, as the class Reptilia is paraphyletic, that is, it has given rise to two
other classes not included in Reptilia.
[edit] Phylogenetic classification
With the advent of cladistics, some researchers have attempted to establish new classes, based on
phylogeny, but disregarding the physiological and anatomical unity of the groups. One such classification, by
Michael Benton, is presented in simplified form below.[7]
• Clade Amniota ("reptiles")
• Class Synapsida - includes mammal-like reptiles
• *Order Pelycosauria †
• Order Therapsida
• Class Mammalia - Mammals
• Class Sauropsida
• Subclass Anapsida
• Order Testudines - Turtles
• Subclass Diapsida
• Order Araeoscelidia †
• Order Younginiformes †
• Infraclass Ichthyosauria †
• Infraclass Lepidosauromorpha
• Superorder Sauropterygia †
• Order Placodontia †
• Order Nothosauroidea †
• Order Plesiosauria †
• Superorder Lepidosauria
• Order Sphenodontida - Tuatara
• Order Squamata - Lizards & snakes
• Infraclass Archosauromorpha
• Order Prolacertiformes †
• Division Archosauria
• Subdivision Crurotarsi
• Order Crocodylia -
Crocodilians
• Subdivision Avemetatarsalia
• Order Pterosauria †
• Superorder Dinosauria
• Order Ornithischia †
• Order Saurischia
• Class Aves - Birds
[edit] Cladogram of Amniotes

The cladogram presented here illustrates the phylogeny (family tree) of amniotes, and follows a
simplified version of the relationships found by Laurin & Reisz (1995).[8] The cladogram covers the group as
defines under Gauthier's definition.
Amniota
Synaps
Sauropsida
Mesosauridae
Reptilia
Parareptilia
Millerettidae
unnamed
Pareiasauria
unnamed
Procolophonoidea
Testudines
(turtles, tortoises, and
terrapins)
Eureptilia
Captorhinidae
Romeriida
Protorothyrididae *
Diapsida (lizards, snakes, crocodiles,

birds, etc.)
[edit] References
1. ^ Benton, Michael J. (1997). Vertebrate Palaeontology. London: Chapman & Hall. pp. 105–
109. ISBN 0-412-73810-4.
2. ^ www.pnas.org
3. ^ Stewart J. R. (1997): Morphology and evolution of the egg of oviparous amniotes. In: S.
Sumida and K. Martin (ed.) Amniote Origins-Completing the Transition to Land (1): 291-326. London:
Academic Press.
4. ^ Romer, A.S. & Parsons, T.S. (1985): The Vertebrate Body. (6th ed.) Saunders,
Philadelphia.
5. ^ a b Gauthier, J., Kluge, A.G. and Rowe, T. (1988). "The early evolution of the Amniota." Pp.
103-155 in Benton, M.J. (ed.), The phylogeny and classification of the tetrapods, Volume 1:
amphibians, reptiles, birds. Oxford: Clarendon Press.
6. ^ Lee, M.S.Y. & Spencer, P.S. (1997): Crown clades, key characters and taxonomic stability:
when is an amniote not an amniote? In: Sumida S.S. & Martin K.L.M. (eds.) Amniote Origins:
completing the transition to land. Academic Press, pp 61-84. Google books
7. ^ Benton, M.J. (2004). Vertebrate Paleontology. Blackwell Publishers. xii-452. ISBN 0-632-
05614-2.
8. ^ Laurin, M. and Reisz, R.R. (1995). "A reevaluation of early amniote phylogeny." Zoological
Journal of the Linnean Society, 113: 165-223.
challenged and removed. (January 2009)

• Out of the Swamps
Retrieved from "http://en.wikipedia.org/wiki/Amniote"
Categories: Tetrapods
W000
Cretaceous

Cretaceous Period
145.5 – 65.5 million years ago
PreЄ
g
Mean atmospheric O2 content over period duration ca. 30 Vol %[1]

(150 % of modern level)
Mean atmospheric CO2 content over period duration ca. 1700 ppm[2]
(6 times pre-industrial level)
ca. 18 °C[3]
Mean surface temperature over period duration (4 °C above modern level)
The Cretaceous (pronounced /krɪˈteɪʃəs/), Latin for "chalky", usually abbreviated K for its German
translation Kreide (chalk), is a geologic period and system from circa 145.5 ± 4 to 65.5 ± 0.3 million years
(Ma) ago. In the geologic timescale, the Cretaceous follows on the Jurassic Period and is followed by the
Paleogene Period of the Cenozoic Era. It is the youngest period of the Mesozoic Era, and at 80 million years
long, the longest period of the Phanerozoic Eon. The end of the Cretaceous defines the boundary between
the Mesozoic and Cenozoic eras. In many languages this period is known as "chalk period".
The Cretaceous was a period with a relatively warm climate and high eustatic sea level. The oceans
and seas were populated with now extinct marine reptiles, ammonites and rudists; and the land by dinosaurs.
At the same time, new groups of mammals and birds as well as flowering plants appeared. The Cretaceous
ended with one of the largest mass extinctions in Earth history, the K-T extinction, when many species,
including non-avian dinosaurs, pterosaurs, and large marine reptiles, disappeared.
A plate with Nematonotus sp. , Pseudostacus sp., and a partial Dercetis triqueter from Cretaceous
found in Hakel, Lebanon
Contents
[hide]
• 1 The Cretaceous world
• 1.1 Paleogeography
• 1.2 Climate
• 2 Geology
• 2.1 Research history
• 2.2 Stratigraphic subdivisions
• 2.3 Rock formations
• 3 Life
• 3.1 Plants
• 3.2 Terrestrial fauna
• 3.3 Marine fauna
• 3.4 Extinction
• 4 See also
• 5 Notes
• 6 References
[edit] The Cretaceous world
[edit] Paleogeography
During the Cretaceous, the late-Paleozoic-to-early-Mesozoic supercontinent of Pangaea completed
its tectonic breakup into present day continents, although their positions were substantially different at the
time. As the Atlantic Ocean widened, the convergent-margin orogenies that had begun during the Jurassic
continued in the North American Cordillera, as the Nevadan orogeny was followed by the Sevier and
Laramide orogenies.
Geography of the US in the Late Cretaceous Period

Though Gondwana was still intact in the beginning of the Cretaceous, it broke up as South America,
Antarctica and Australia rifted away from Africa (though India and Madagascar remained attached to each
other); thus, the South Atlantic and Indian Oceans were newly formed. Such active rifting lifted great
undersea mountain chains along the welts, raising eustatic sea levels worldwide. To the north of Africa the
Tethys Sea continued to narrow. Broad shallow seas advanced across central North America (the Western
Interior Seaway) and Europe, then receded late in the period, leaving thick marine deposits sandwiched
between coal beds. At the peak of the Cretaceous transgression, one-third of Earth's present land area was
submerged.[4]
The Cretaceous is justly famous for its chalk; indeed, more chalk formed in the Cretaceous than in
any other period in the Phanerozoic.[5] Mid-ocean ridge activity—or rather, the circulation of seawater through
the enlarged ridges—enriched the oceans in calcium; this made the oceans more saturated, as well as
increased the bioavailability of the element for calcareous nanoplankton.[6] These widespread carbonates
and other sedimentary deposits make the Cretaceous rock record especially fine. Famous formations from
North America include the rich marine fossils of Kansas's Smoky Hill Chalk Member and the terrestrial fauna
of the late Cretaceous Hell Creek Formation. Other important Cretaceous exposures occur in Europe (e.g.,
the Weald) and China (the Yixian Formation). In the area that is now India, massive lava beds called the
Deccan Traps were erupted in the very late Cretaceous and early Paleocene.
[edit] Climate
The Berriasian epoch showed a cooling trend that had been seen in the last epoch of the Jurassic.
There is evidence that snowfalls were common in the higher latitudes and the tropics became wetter than
during the Triassic and Jurassic.[7] Glaciation was however restricted to alpine glaciers on some high-
latitude mountains, though seasonal snow may have existed farther south. Rafting by ice of stones into
marine environments occurred during much of the Cretaceous but evidence of deposition directly from
glaciers is limited to the Early Cretaceous of the Eromanga Basin in southern Australia.[8][9]
After the end of the Berriasian, however, temperatures increased again, and these conditions were
almost constant until the end of the period.[7] This trend was due to intense volcanic activity which produced
large quantities of carbon dioxide. The development of a number of mantle plumes across the widening mid-
ocean ridges further pushed sea levels up, so that large areas of the continental crust were covered with
shallow seas. The Tethys Sea connecting the tropical oceans east to west also helped in warming the global
climate. Warm-adapted plant fossils are known from localities as far north as Alaska and Greenland, while
dinosaur fossils have been found within 15 degrees of the Cretaceous south pole.[10]
A very gentle temperature gradient from the equator to the poles meant weaker global winds,
contributing to less upwelling and more stagnant oceans than today. This is evidenced by widespread black
shale deposition and frequent anoxic events.[11] Sediment cores show that tropical sea surface
temperatures may have briefly been as warm as 42 °C (107 °F), 17 °C (31 °F) warmer than at present, and
that they averaged around 37 °C (99 °F). Meanwhile deep ocean temperatures were as much as 15 to 20 °C
(27 to 36 °F) higher than today's.[12][13]
Further information: Cool tropics paradox
[edit] Geology
Key events in the Cretaceous

-140 —
–
-130 —
–
-120 —
–
-110 —
–
-100 —
–
-90 —
–
-80 —
–
-70 —
–
Maastrichtian
Campanian
Santonian
Coniacian
Turonian
Cenomanian
Albian
Aptian
Barremian
Hauterivian
Valanginian
Berriasian
Jurassic
Paleogene
C
r
e
t
a
c
e
o
u
s
Mesozoic
Cenozoic
An approximate timescale of key Cretaceous events.

[edit] Research history

The Cretaceous as a separate period was first defined by a Belgian geologist Jean d'Omalius
d'Halloy in 1822, using strata in the Paris Basin[14] and named for the extensive beds of chalk (calcium
carbonate deposited by the shells of marine invertebrates, principally coccoliths), found in the upper
Cretaceous of western Europe. The name Cretaceous was derived from Latin creta, meaning chalk.[15] The
name of the island Crete has the same origin.
[edit] Stratigraphic subdivisions

The Cretaceous is divided into Early and Late Cretaceous epochs or Lower and Upper Cretaceous
series. In older literature the Cretaceous is sometimes divided into three series: Neocomian (lower/early),
Gallic (middle) and Senonian (upper/late). A subdivision in eleven stages, all origining from European
stratigraphy, is now used worldwide. In many parts of the world, alternative local subdivisions are still in use.
As with other older geologic periods, the rock beds of the Cretaceous are well identified but the exact
ages of the system's top and base are uncertain by a few million years. No great extinction or burst of
diversity separates the Cretaceous from the Jurassic. However, the top of the system is sharply defined,
being placed at an iridium-rich layer found worldwide that is believed to be associated with the Chicxulub
impact crater in Yucatan and the Gulf of Mexico. This layer has been tightly dated at 65.5 Ma.[16]
[edit] Rock formations
Drawing of fossil jaws of Mosasaurus hoffmanni, from the Maastrichtian of Dutch Limburg, by Dutch
geologist Pieter Harting (1866).
The high eustatic sea level and warm climate of the Cretaceous meant a large area of the continents
was covered by warm shallow seas. The Cretaceous was named for the extensive chalk deposits of this age
in Europe, but in many parts of the world, the Cretaceous system consists for a major part of marine
limestone, a rock type that is formed under warm, shallow marine circumstances. Due to the high sea level
there was extensive accommodation space for sedimentation so that thick deposits could form. Because of
the relatively young age and great thickness of the system, Cretaceous rocks crop out in many areas
worldwide.
Chalk is a rock type characteristic for (but not restricted to) the Cretaceous. It consists of coccoliths,
microscopically small calcite skeletons of coccolithophores, a type of algae that prospered in the Cretaceous
seas.
In northwestern Europe, chalk deposits from the Upper Cretaceous are characteristic for the Chalk
Group, which forms the white cliffs of Dover on the south coast of England and similar cliffs on the French
Normandian coast. The group is found in England, northern France, the low countries, northern Germany,
Denmark and in the subsurface of the southern part of the North Sea. Chalk is not easily consolidated and
the Chalk Group still consists of loose sediments in many places. The group also has other limestones and
arenites. Among the fossils it contains are sea urchins, belemnites, ammonites and sea reptiles such as
Mosasaurus.
In southern Europe, the Cretaceous is usually a marine system consisting of competent limestone
beds or incompetent marls. Because the Alpine mountain chains did not yet exist in the Cretaceous, these
deposits formed on the southern edge of the European continental shelf, at the margin of the Tethys Ocean.
Stagnation of deep sea currents in middle Cretaceous times caused anoxic circumstances in the sea
water. In many places around the world, dark anoxic shales were formed during this interval.[17] These
shales are an important source rock for oil and gas, for example in the subsurface of the North Sea.
[edit] Life
[edit] Plants
Flowering plants (angiosperms) spread during this period, although they did not become
predominant until the Campanian stage near the end of the epoch. Their evolution was aided by the
appearance of bees; in fact angiosperms and insects are a good example of coevolution. The first
representatives of many leafy trees, including figs, planes and magnolias, appeared in the Cretaceous. At the
same time, some earlier Mesozoic gymnosperms like Conifers continued to thrive; pehuéns (Monkey Puzzle
trees, Araucaria) and other conifers being notably plentiful and widespread. Some fern orders such as
Gleicheniales[18] appeared as early in the fossil record as the Cretaceous, and achieved an early broad
distribution. Gymnosperm taxa like Bennettitales died out before the end of the period.[citation needed]
[edit] Terrestrial fauna
Tyrannosaurus rex, one of the largest land predators of all time, lived during the late Cretaceous.
A pterosaur, Anhanguera piscator
On land, mammals were a small and still relatively minor component of the fauna. Early marsupial
mammals evolved in the Early Cretaceous, with true placentals emerging in the Late Cretaceous period. The
fauna was dominated by archosaurian reptiles, especially dinosaurs, which were at their most diverse stage.
Pterosaurs were common in the early and middle Cretaceous, but as the Cretaceous proceeded they faced
growing competition from the adaptive radiation of birds, and by the end of the period only two highly
specialized families remained.
The Liaoning lagerstätte (Chaomidianzi formation) in China provides a glimpse of life in the Early
Cretaceous, where preserved remains of numerous types of small dinosaurs, birds, and mammals have been
found. The coelurosaur dinosaurs found there represent types of the group Maniraptora, which is transitional
between dinosaurs and birds, and are notable for the presence of hair-like feathers.
During the Cretaceous, insects began to diversify, and the oldest known ants, termites and some
lepidopterans, akin to butterflies and moths, appeared. Aphids, grasshoppers, and gall wasps appeared.[19]
[edit] Marine fauna
Discoscaphites iris, Owl Creek Formation (Upper Cretaceous), Ripley, Mississippi.

In the seas, rays, modern sharks and teleosts became common.[20] Marine reptiles included
ichthyosaurs in the early and middle of the Cretaceous, becoming extinct during the late Cretaceous,
plesiosaurs throughout the entire period, and mosasaurs appearing in the Late Cretaceous.
Baculites, an ammonite genus with a straight shell, flourished in the seas along with reef-building
rudist clams. The Hesperornithiformes were flightless, marine diving birds that swam like grebes.
Globotruncanid Foraminifera and echinoderms such as sea urchins and starfish (sea stars) thrived. The first
radiation of the diatoms (generally siliceous, rather than calcareous) in the oceans occurred during the
Cretaceous; freshwater diatoms did not appear until the Miocene.[19] The Cretaceous was also an important
interval in the evolution of bioerosion, the production of borings and scrapings in rocks, hardgrounds and
shells (Taylor and Wilson, 2003).
[edit] Extinction
Main article: Cretaceous–Tertiary extinction event
There was a progressive decline in biodiversity during the Maastrichtian stage of the Cretaceous
Period prior to the suggested ecological crisis induced by events at the K-T boundary. Furthermore,
biodiversity required a substantial amount of time to recover from the K-T event, despite the probable
existence of an abundance of vacant ecological niches.[21]
Despite the severity of this boundary event, there was significant variability in the rate of extinction
between and within different clades. Species which depended on photosynthesis declined or became extinct
because of the reduction in solar energy reaching the Earth's surface due to atmospheric particles blocking
the sunlight. As is the case today, photosynthesizing organisms, such as phytoplankton and land plants,
formed the primary part of the food chain in the late Cretaceous. Evidence suggests that herbivorous
animals, which depended on plants and plankton as their food, died out as their food sources became
scarce; consequently, top predators such as Tyrannosaurus rex also perished.[22]
Coccolithophorids and molluscs, including ammonites, rudists, freshwater snails and mussels, as
well as organisms whose food chain included these shell builders, became extinct or suffered heavy losses.
For example, it is thought that ammonites were the principal food of mosasaurs, a group of giant marine
reptiles that became extinct at the boundary.[23]
Omnivores, insectivores and carrion-eaters survived the extinction event, perhaps because of the
increased availability of their food sources. At the end of the Cretaceous there seem to have been no purely
herbivorous or carnivorous mammals. Mammals and birds which survived the extinction fed on insects,
larvae, worms, and snails, which in turn fed on dead plant and animal matter. Scientists theorise that these
organisms survived the collapse of plant-based food chains because they fed on detritus.[24][21][25]
In stream communities, few groups of animals became extinct. Stream communities rely less on food
from living plants and more on detritus that washes in from land. This particular ecological niche buffered
them from extinction.[26] Similar, but more complex patterns have been found in the oceans. Extinction was
more severe among animals living in the water column, than among animals living on or in the sea floor.
Animals in the water column are almost entirely dependent on primary production from living phytoplankton,
while animals living on or in the ocean floor feed on detritus or can switch to detritus feeding.[21]
The largest air-breathing survivors of the event, crocodilians and champsosaurs, were semi-aquatic
and had access to detritus. Modern crocodilians can live as scavengers and can survive for months without
food, and their young are small, grow slowly, and feed largely on invertebrates and dead organisms or
fragments of organisms for their first few years. These characteristics have been linked to crocodilian survival
at the end of the Cretaceous.[24]
Numerous borings in Cretaceous Rudist bivalves from
a Cretaceous cobble, hardground from Texas with the Cretaceous of the Omani
Faringdon, England; these encrusting oysters and Mountains, United Arab Inoceramus from the
are excellent examples of borings. The scale bar is Emirates. Scale bar is Cretaceous of South Dakota.
fossil bioerosion. 10 mm. 10 mm.
[edit] See also

Geology portal
Paleontology portal
Time portal
• Chalk Formation
• South Polar dinosaurs
• Western Interior Seaway
[edit] Notes
1. ^ Image:Sauerstoffgehalt-1000mj.svg
2. ^ Image:Phanerozoic Carbon Dioxide.png
3. ^ Image:All palaeotemps.png
4. ^ Dougal Dixon et al., Atlas of Life on Earth, (New York: Barnes & Noble Books, 2001), p.
215.
5. ^ Stanley, Steven M. Earth System History. New York: W.H. Freeman and Company, 1999.
ISBN 0-7167-2882-6 p. 280
6. ^ Stanley, pp. 279–81
7. ^ a b The Berriasian Age
8. ^ Alley, N.F. and Frakes, L.A. 2003. "First known Cretaceous glaciation: Livingston Tillite,
South Australia". Australian Journal of Earth Science 50:134–150.
9. ^ Frakes, L.A. and Francis, J. E. 1988. "A guide to Phanerozoic cold climates from high
latitude ice rafting in the Cretaceous". Nature 333:547–549.
10.^ Stanley, pp. 480–2
11.^ Stanley, pp. 481–2
12.^ "Warmer than a Hot Tub: Atlantic Ocean Temperatures Much Higher in the Past"
PhysOrg.com. Retrieved 12/3/06.
13.^ Skinner, Brian J., and Stephen C. Porter. The Dynamic Earth: An Introduction to Physical
Geology. 3rd ed. New York: John Wiley & Sons, Inc., 1995. ISBN 0-471-59549-7. p. 557
14.^ (in Russian) Great Soviet Encyclopedia (3rd ed.). Moscow: Sovetskaya Enciklopediya.
1974. vol. 16, p. 50.
15.^ Glossary of Geology (3rd ed.). Washington, D.C.: American Geological Institute. 1972.
pp. 165.
16.^ The official geologic timescale of the ICS (in 2008) gives 65.5 Ma as upper boundary of the
Cretaceous, new callibrations by Kuiper et al. (2008) yield 65.9 Ma
17.^ See Stanley (1999), pp. 481–482
18.^ C.Michael Hogan. 2010. Fern. Encyclopedia of Earth. National council for Science and the
Environment. Washington, DC
19.^ a b http://www.ucmp.berkeley.edu/mesozoic/cretaceous/cretlife.html
20.^ http://www.talkorigins.org/origins/geo_timeline.html
21.^ a b c MacLeod, N, Rawson, PF, Forey, PL, Banner, FT, Boudagher-Fadel, MK, Bown, PR,
Burnett, JA, Chambers, P, Culver, S, Evans, SE, Jeffery, C, Kaminski, MA, Lord, AR, Milner, AC,
Milner, AR, Morris, N, Owen, E, Rosen, BR, Smith, AB, Taylor, PD, Urquhart, E & Young, JR (1997).
"The Cretaceous–Tertiary biotic transition". Journal of the Geological Society 154 (2): 265–292.
doi:10.1144/gsjgs.154.2.0265.
22.^ Wilf, P & Johnson KR (2004). "Land plant extinction at the end of the Cretaceous: a
quantitative analysis of the North Dakota megafloral record". Paleobiology 30 (3): 347–368.
doi:10.1666/0094-8373(2004)030<0347:LPEATE>2.0.CO;2.
23.^ Kauffman, E (2004). "Mosasaur Predation on Upper Cretaceous Nautiloids and Ammonites
from the United States Pacific Coast". PALAIOS (Society for Sedimentary Geology) 19 (1): 96–100.
doi:10.1669/0883-1351(2004)019<0096:MPOUCN>2.0.CO;2.
http://palaios.geoscienceworld.org/cgi/reprint/19/1/96. Retrieved 2007-06-17.
24.^ a b Shehan, P & Hansen, TA (1986). "Detritus feeding as a buffer to extinction at the end of
the Cretaceous". Geology 14 (10): 868–870. doi:10.1130/0091-
7613(1986)14<868:DFAABT>2.0.CO;2.
25.^ Aberhan, M, Weidemeyer, S, Kieesling, W, Scasso, RA, & Medina, FA (2007). "Faunal
evidence for reduced productivity and uncoordinated recovery in Southern Hemisphere Cretaceous-
Paleogene boundary sections". Geology 35 (3): 227–230. doi:10.1130/G23197A.1.
26.^ Sheehan, PM & Fastovsky, DE (1992). "Major extinctions of land-dwelling vertebrates at
the Cretaceous–Tertiary boundary, eastern Montana". Geology 20 (6): 556–560. doi:10.1130/0091-
7613(1992)020<0556:MEOLDV>2.3.CO;2. http://www.geoscienceworld.org/cgi/georef/1992034409.
[edit] References
• Kashiyama, Yuichiro; Nanako O. Ogawa, Junichiro Kuroda, Motoo Shiro, Shinya Nomoto,
Ryuji Tada, Hiroshi Kitazato, Naohiko Ohkouchi (2008-05). "Diazotrophic cyanobacteria as the major
photoautotrophs during mid-Cretaceous oceanic anoxic events: Nitrogen and carbon isotopic
evidence from sedimentary porphyrin". Organic Geochemistry 39 (5): 532–549.
doi:10.1016/j.orggeochem.2007.11.010. http://www.sciencedirect.com/science?
_ob=ArticleURL&_udi=B6V7P-4R98K6R-
1&_user=1080547&_rdoc=1&_fmt=&_orig=search&_sort=d&view=c&_acct=C000051389&_version=
1&_urlVersion=0&_userid=1080547&md5=49204479929f0c87061bf3c69d7b1949. Retrieved 2008-
05-10.
• Neal L Larson, Steven D Jorgensen, Robert A Farrar and Peter L Larson. Ammonites and
the other Cephalopods of the Pierre Seaway. Geoscience Press, 1997.
• Ogg, Jim; June, 2004, Overview of Global Boundary Stratotype Sections and Points
(GSSP's) http://www.stratigraphy.org/gssp.htm Accessed April 30, 2006.
• Ovechkina, M.N. and Alekseev, A.S. 2005. Quantitative changes of calcareous nannoflora in
the Saratov region (Russian Platform) during the late Maastrichtian warming event. Journal of Iberian
Geology 31 (1): 149–165. PDF
• Rasnitsyn, A.P. and Quicke, D.L.J. (2002). History of Insects. Kluwer Academic Publishers.
ISBN 1-4020-0026-X. —detailed coverage of various aspects of the evolutionary history of the insects.
• Skinner, Brian J., and Stephen C. Porter. The Dynamic Earth: An Introduction to Physical
Geology. 3rd ed. New York: John Wiley & Sons, Inc., 1995. ISBN 0-471-60618-9}
• Stanley, Steven M. Earth System History. New York: W.H. Freeman and Company, 1999.
ISBN 0-7167-2882-6
• Taylor, P.D. and Wilson, M.A., 2003. Palaeoecology and evolution of marine hard substrate
communities. Earth-Science Reviews 62: 1–103.[1]

Look up cretaceous in Wiktionary, the free dictionary.
Wikimedia Commons has media related to: Cretaceous
• UCMP Berkeley Cretaceous page

• Bioerosion website at The College of Wooster
• Cretaceous Microfossils: 180+ images of Foraminifera
Cretaceous Period
Lower/Early Cretaceous Upper/Late Cretaceous
Berriasian | Valanginian | Hauterivian Cenomanian | Turonian | Coniacian

Barremian | Aptian | Albian Santonian | Campanian | Maastrichtian
Preceded
542 Ma - Paleozoic Era - 251 Ma
by Proterozoic Eon
Cambrian Ordovician Silurian Devonian Carboniferous
[hide]v · d · eGeologic history of Earth
[show] Precambrian (4.57 Gya – 542 Mya)
In left column are eons; right column: bold are eras; not bold are periods:
Hadean
(4.57 – 4 Gya) (informal)
Eoarchean (4 – 3.6 Gya)
Archean Paleoarchean (3.6 – 3.2 Gya)
(4 – 2.5 Gya) Mesoarchean (3.2 – 2.8 Gya)
Neoarchean (2.8 – 2.5 Gya)
Paleoproterozoic (2.5 – 1.6 Gya): Siderian (2.5 – 2.3 Gya) · Rhyacian (2.3 –
2.05 Gya) · Orosirian (2.05 – 1.8 Gya) · Statherian (1.8 – 1.6 Gya)
Proterozoic Mesoproterozoic (1.6 – 1 Gya): Calymmian (1.6 – 1.4 Gya) · Ectasian (1.4 – 1.2
(2.5 Gya – 542 Mya) Gya) · Stenian (1.2 – 1 Gya)
Neoproterozoic (1 Gya – 542 Mya): Tonian (1 Gya – 850 Mya) · Cryogenian (850 –
635 Mya) · Ediacaran (635 – 542 Mya)
Mya = millions years ago. Gya = billions years ago.
[hide] Phanerozoic (542 – 0 Mya)
In horizontal bars are eras; in left column are periods; right column: bold are epochs; not bold not
italic are ages; italic are chrons:
[show] Paleozoic (542 – 251 Mya)
Terreneuvian (542 – 521 Mya): Fortunian (542 – 528 Mya) · Age 2* (528 –
521 Mya)
Epoch 2* (521 – 510 Mya): Age 3* (521 – 515 Mya) · Age 4* (515 – 510 Mya)
Cambrian
Epoch 3* (510 – 499 Mya): Age 5* (510 – 506.5 Mya) · Drumian (506.5 – 503
(542 – 488.3 Mya)
Mya) · Guzhangian (503 – 499 Mya)
Furongian (499 – 488.3 Mya): Paibian (499 – 496 Mya) · Age 9* (496 – 492 Mya) ·
Age 10* (492 – 488.3 Mya)
Early Ordovician (488.3 – 471.8 Mya): Tremadocian (488.3 – 478.6 Mya) ·

Floian (478.6 – 471.8 Mya)
Ordovician Middle Ordovician (471.8 – 460.9 Mya): Dapingian (471.8 – 468.1 Mya) ·
(488.3 – 443.7 Mya) Darriwilian (468.1 – 460.9 Mya)
Late Ordovician (460.9 – 443.7 Mya): Sandbian (460.9 – 455.8 Mya) · Katian
(455.8 – 445.6 Mya) · Hirnantian (445.6 – 443.7 Mya)
Llandovery (443.7 – 428.2 Mya): Rhuddanian (443.7 – 439 Mya) ·

Aeronian (439 – 436 Mya) · Telychian (436 – 428.2 Mya)
Silurian
Wenlock (428.2 – 422.9 Mya): Sheinwoodian (428.2 – 426.2 Mya) · Homerian
(443.7 – 416 Mya)
(426.2 – 422.9 Mya)
Ludlow (422.9 – 418.7 Mya): Gorstian (422.9 – 421.3 Mya) · Ludfordian (421.3 –
418.7 Mya)
Pridoli (418.7 – 416 Mya)
Early Devonian (416 – 397.5 Mya): Lochkovian (416 – 411.2 Mya) ·

Pragian (411.2 – 407 Mya) · Emsian (407 – 397.5 Mya)
Devonian Middle Devonian (397.5 – 385.3 Mya): Eifelian (397.5 – 391.8 Mya) · Givetian
(416 – 359.2 Mya) (391.8 – 385.3 Mya)
Late Devonian (385.3 – 359.2 Mya): Frasnian (385.3 – 374.5 Mya) · Famennian
(374.5 – 359.2 Mya)
Mississippian (359.2 – 318.1 Mya): Tournaisian / Early Mississippian

(359.2 – 345.3 Mya) · Viséan / Middle Mississippian (345.3 – 328.3 Mya) ·
Serpukhovian / Late Mississippian (328.3 – 318.1 Mya)
Carboniferous
Pennsylvanian (318.1 – 299 Mya): Bashkirian / Early Pennsylvanian (318.1 –
(359.2 – 299 Mya)
311.7 Mya) · Moscovian / Middle Pennsylvanian (311.7 – 307.2 Mya) · Late
Pennsylvanian (307.2 – 299 Mya): Kasimovian (307.2 – 303.4 Mya) · Gzhelian
(303.4 – 299 Mya)
Permian Cisuralian (299 – 270.6 Mya): Asselian (299 – 294.6 Mya) · Sakmarian
(299 – 251 Mya) (294.6 – 284.4 Mya) · Artinskian (284.4 – 275.6 Mya) · Kungurian (275.6 – 270.6
Mya)
Guadalupian (270.6 – 260.4 Mya): Roadian (270.6 – 268 Mya) · Wordian (268 –
265.8 Mya) · Capitanian (265.8 – 260.4 Mya)
Lopingian (260.4 – 251 Mya): Wuchiapingian (260.4 – 253.8 Mya) ·
Changhsingian (253.8 – 251 Mya)
[hide] Mesozoic (251 – 65.5 Mya)
Early Triassic (251 – 245.9 Mya): Induan (251 – 249.5 Mya) · Olenekian
(249.5 – 245.9 Mya)
Triassic Middle Triassic (245.9 – 228.7 Mya): Anisian (245.9 – 237 Mya) · Ladinian (237 –
(251 – 199.6 Mya) 228.7 Mya)
Late Triassic (228.7 – 199.6 Mya): Carnian (228.7 – 216.5 Mya) · Norian (216.5 –
203.6 Mya) · Rhaetian (203.6 – 199.6 Mya)
Early Jurassic (199.6 – 175.6 Mya): Hettangian (199.6 – 196.5 Mya) ·

Sinemurian (196.5 – 189.6 Mya) · Pliensbachian (189.6 – 183 Mya) · Toarcian (183
– 175.6 Mya)
Jurassic Middle Jurassic (175.6 – 161.2 Mya): Aalenian (175.6 – 171.6 Mya) · Bajocian
(199.6 – 145.5 Mya) (171.6 – 167.7 Mya) · Bathonian (167.7 – 164.7 Mya) · Callovian (164.7 – 161.2
Mya)
Late Jurassic (161.2 – 145.5 Mya): Oxfordian (161.2 – 155.6 Mya) · Kimmeridgian
(155.6 – 150.8 Mya) · Tithonian (150.8 – 145.5 Mya)
Early Cretaceous (145.5 – 99.6 Mya): Berriasian (145.5 – 140.2 Mya) ·
Valanginian (140.2 – 133.9 Mya) · Hauterivian (133.9 – 130 Mya) · Barremian (130
Cretaceous – 125 Mya) · Aptian (125 – 112 Mya) · Albian (112 – 99.6 Mya)
(145.5 – 65.5 Mya) Late Cretaceous (99.6 – 65.5 Mya): Cenomanian (99.6 – 93.6 Mya) · Turonian
(93.6 – 88.6 Mya) · Coniacian (88.6 – 85.8 Mya) · Santonian (85.8 – 83.5 Mya) ·
Campanian (83.5 – 70.6 Mya) · Maastrichtian (70.6 – 65.5 Mya)
[hide] Cenozoic (65.5 – 0 Mya)
Paleogene, Neogene and early Pleistocene comprise former Tertiary* (65.5 – 1.8 Mya) period.
Gelasian and Calabrian comprise Early Pleistocene (2.588 Mya – 781 kya) subepoch.
Paleocene (65.5 – 55.8 Mya): Danian (65.5 – 61.1 Mya) · Selandian (61.1 –
58.7 Mya) · Thanetian (58.7 – 55.8 Mya)
Paleogene Eocene (55.8 – 33.9 Mya): Ypresian (55.8 – 48.6 Mya) · Lutetian (48.6 – 40.4 Mya) ·
(65.5 – 23.03 Mya) Bartonian (40.4 – 37.2 Mya) · Priabonian (37.2 – 33.9 Mya)
Oligocene (33.9 – 23.03 Mya): Rupelian (33.9 – 28.4 Mya) · Chattian (28.4 – 23.03
Mya)
Neogene Miocene (23.03 – 5.332 Mya): Aquitanian (23.03 – 20.43 Mya) · Burdigalian
(23.03 – 2.588 Mya) (20.43 – 15.97 Mya) · Langhian (15.97 – 13.82 Mya) · Serravallian (13.82 – 11.608
Mya) · Tortonian (11.608 – 7.246 Mya) · Messinian (7.246 – 5.332 Mya)
Pliocene (5.332 – 2.588 Mya): Piacenzian (5.332 – 3.6 Mya) · Zanclean (3.6 – 2.588
Mya)
Pleistocene (2.588 Mya – 11.4 kya): Gelasian (2.588 – 1.806 Mya) ·

Calabrian (1.806 Mya – 781 kya) · Middle Pleistocene / Ionian (781 – 126 kya) · Late
Pleistocene / Tarantian (126 – 11.4 kya): Oldest Dryas* (18 – 14.67 kya) · Bølling*
Quaternary
(14.67 – 14 kya) · Older Dryas* (14 – 13.7 kya) · Allerød* (13.7 – 12.8 kya) · Younger
(2.588 – 0 Mya)
Dryas* (12.8 – 11.4 kya)
Holocene (11.4 – 0 kya): Preboreal* (11.4 – 9 kya) · Boreal* (9 – 8 kya) · Atlantic* (8 –
5 kya) · Subboreal* (5 – 2.5 kya) · Subatlantic* (2.5 – 0 kya)
kya = thousands years ago. Mya = millions years ago. * Not officially recognized by the I.C.S.
Source: International Stratigraphic Chart. International Commission on Stratigraphy. Retrieved 8

February 2008.
Retrieved from "http://en.wikipedia.org/wiki/Cretaceous"

Categories: Cretaceous | Mesozoic
W000
W000
Eugene S. Gaffney

Eugene S. Gaffney is an American paleontologist and leading authority on the evolutionary history of
turtles.
Contents
[hide]
• 1 Biography
• 2 Accolades
• 3 Selected publications
• 4 Notes
[edit] Biography
He graduated from Rutgers State University in 1965; and received his PhD in 1969 with a thesis on
"The North American Baenoidea and the Cryptodire-Pleurodire Dichotomy" from Columbia University, where
he also taught for several years. He then moved to the Department of Vertebrate Paleontology, American
Museum of Natural History, becoming full Curator in 1980, a position he still holds. He has authored
numerous publications on turtle systematics and phylogeny.[1]
Dr. Gaffney pioneered the use of cladistics in turtle research.[2] He has done fieldwork in Canada
and the United States, central Europe, southern Africa, China, Argentina, Brazil, and especially Australia,
where he has studied the evolution of the Meiolania, the giant horned tortoise.[3]
[edit] Accolades
A symposium on fossil turtles was held in his honor at the Royal Tyrrell Museum in Drumheller,
Alberta, Canada, in 2009.[4][5]
Several species of fossil turtles, including the protostegid sea turtle, Santanachelys gaffneyi , and the
Macrobaenid Aurorachelys gaffneyi[6] have also been named in his honor.
[edit] Selected publications

• Gaffney, E. S. (1975). A phylogeny and classification of the higher categories of turtles.
Bulletin of the American Museum of Natural History 155(5): 389-436. on-line
• Gaffney, E. S. (1979). The Jurassic turtles of North America. Bulletin of the American
Museum of Natural History 162 (3): 91-136.
• Gaffney, E. S. (1979). An introduction to the logic of phylogeny reconstruction. In J. Cracraft
and N. Eldredge (editors), Phylogenetic analysis and paleontology: 79-111. New York: Columbia
University Press.
• Gaffney, E. S. (1980). Phylogenetic relationships of the major groups of amniotes. In A. L.
Panchen (editor), The terrestrial environment and the origin of land vertebrates : 593-610.
LonLondon, New York: Academic Press.
• Gaffney, E. S., & P. A. Meylan. (1988). A phylogeny of turtles. In M. J. Benton, (editor), The
phylogeny and classification of tetrapods: 157-219. Oxford: Clarendon Press.
• Gaffney, E. S. (1990). Dinosaurs A Golden Guide. Western Publishing Company, Inc. 160
pp.
• Norell, M. A., Gaffney, E. S., & Dingus, L. (1995). Discovering Dinosaurs in the American
Museum of Natural History. Alfred A. Knopf, 204 pp.
• Gaffney, E. S. (1996) The postcranial morphology of Meiolania platyceps and a review of the
Meiolaniidae. Bulletin of the American Museum of Natural History ; no. 229 on-line
• Gaffney, E. S., Tong, H., & Meylan, P. A. (2006) Evolution of the side-necked turtles : the
families Bothremydidae, Euraxemydidae, and Araripemydidae. Bulletin of the American Museum of
Natural History, no. 300 on-line
[edit] Notes
1. ^ Q.V. and full list of publications
2. ^ Eugene S. Gaffney, (1975). A phylogeny and classification of the higher categories of
turtles. Bulletin of the American Museum of Natural History 155(5): 389-436.
3. ^ Eugene S. Gaffney - American Museum of Natural History
4. ^ Royal Tyrrell Museum: Gaffney Turtle Symposium. Retrieved September 14, 2010
5. ^ Gaffney Turtle Symposium, Abstracts and Program. Compiled by D.R. Braman. Royal
Tyrrell Museum of Palaeontology, Drumheller, Alberta, October 17–18, 2009
6. ^ Deborah Vandermark, John A. Tarduno, Donald B. Brinkman, Rory D. Cottrell and
Stephanie Mason, (2009). New Late Cretaceous macrobaenid turtle with Asian affinities from the
High Canadian Arctic: Dispersal via ice-free polar routes. Geology February 2009 v. 37 no. 2 p. 183-
186 abstract

• Phylogeny of turtles, American Museum of Natural History, the official site of Eugene Gaffney
Persondata
Name Gaffney, Eugene S.
Alternative names
Short description
Date of birth
Place of birth
Date of death
Place of death
Retrieved from "http://en.wikipedia.org/wiki/Eugene_S._Gaffney"
Categories: American paleontologists | Rutgers University alumni | Living people

W000
On the Origin of Species


"Origin of Species" redirects here. For other uses, see Origin of Species (disambiguation).
Author Charles Darwin
Country United Kingdom
Language English
Natural selection
Subject(s)
Evolutionary biology
Genre(s) science, biology
Publisher John Murray
Publication date 24 November 1859[1]
Media type Print (Hardback)
Pages 502
ISBN N/A
On the Tendency of Species to form Varieties; and on the Perpetuation of

Preceded by
Varieties and Species by Natural Means of Selection
Followed by Fertilisation of Orchids
Charles Darwin's On the Origin of Species, published on 24 November 1859, is a work of scientific
literature which is considered to be the foundation of evolutionary biology. Its full title was On the Origin of
Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life . For
the sixth edition of 1872, the short title was changed to The Origin of Species. Darwin's book introduced the
scientific theory that populations evolve over the course of generations through a process of natural
selection. It presented a body of evidence that the diversity of life arose by common descent through a
branching pattern of evolution. Darwin included evidence that he had gathered on the Beagle expedition in
the 1830s and his subsequent findings from research, correspondence, and experimentation.
Various evolutionary ideas had already been proposed to explain new findings in biology. There was
growing support for such ideas among dissident anatomists and the general public, but during the first half of
the 19th century the English scientific establishment was closely tied to the Church of England, while science
was part of natural theology. Ideas about the transmutation of species were controversial as they conflicted
with the beliefs that species were unchanging parts of a designed hierarchy and that humans were unique,
unrelated to animals. The political and theological implications were intensely debated, but transmutation
was not accepted by the scientific mainstream.
The book was written for non-specialist readers and attracted widespread interest upon its
publication. As Darwin was an eminent scientist, his findings were taken seriously and the evidence he
presented generated scientific, philosophical, and religious discussion. The debate over the book contributed
to the campaign by T.H. Huxley and his fellow members of the X Club to secularise science by promoting
scientific naturalism. Within two decades there was widespread scientific agreement that evolution, with a
branching pattern of common descent, had occurred, but scientists were slow to give natural selection the
significance that Darwin thought appropriate. During the "eclipse of Darwinism" from the 1880s to the 1930s,
various other mechanisms of evolution were given more credit. With the development of the modern
evolutionary synthesis in the 1930s and 1940s, Darwin's concept of evolutionary adaptation through natural
selection became central to modern evolutionary theory, now the unifying concept of the life sciences.
Contents
[hide]
• 1 Summary of Darwin's theory
• 2 Background
• 2.1 Developments before Darwin's theory
• 2.2 Inception of Darwin's theory
• 2.3 Further development
• 3 Publication
• 3.1 Events leading to publication
• 3.2 Time taken to publish
• 3.3 Publication and subsequent editions
• 3.4 Publication outside Great Britain
• 4 Content
• 4.1 Title pages and introduction
• 4.2 Variation under domestication and under nature
• 4.3 Struggle for existence, natural selection, and
divergence
• 4.4 Variation and heredity
• 4.5 Difficulties for the theory
• 4.6 Geologic record
• 4.7 Geographic distribution
• 4.8 Classification, morphology, embryology,
[edit] Summary of Darwin's theory
Darwin pictured shortly before publication

Darwin's theory of evolution is based on key facts and the inferences drawn from them, which
biologist Ernst Mayr summarised as follows:[3]
• Every species is fertile enough that if all offspring survived to reproduce the population would
grow (fact).
• Despite periodic fluctuations, populations remain roughly the same size (fact).
• Resources such as food are limited and are relatively stable over time (fact).
• A struggle for survival ensues (inference).
• Individuals in a population vary significantly from one another (fact).
• Much of this variation is inheritable (fact).
• Individuals less suited to the environment are less likely to survive and less likely to
reproduce; individuals more suited to the environment are more likely to survive and more likely to
reproduce and leave their inheritable traits to future generations, which produces the process of
natural selection (inference).
• This slowly effected process results in populations changing to adapt to their environments,
and ultimately, these variations accumulate over time to form new species (inference).
[edit] Background
See also: History of evolutionary thought and History of biology
[edit] Developments before Darwin's theory

In later editions of the book, Darwin traced evolutionary ideas as far back as Aristotle;[4] the text he
cites is a summary by Aristotle of the ideas of the earlier Greek philosopher Empedocles.[5] Early Christian
Church Fathers and Medieval European scholars interpreted the Genesis creation myth allegorically rather
than as a literal historical account;[6] organisms were described by their mythological and heraldic
significance as well as by their physical form. Nature was widely believed to be unstable and capricious, with
monstrous births from union between species, and spontaneous generation of life.[7]
Cuvier's 1799 paper on living and fossil elephants helped establish the reality of extinction.
The Protestant Reformation inspired a literal interpretation of the Bible, with concepts of creation that
conflicted with the findings of an emerging science seeking explanations congruent with the mechanical
philosophy of René Descartes and the empiricism of the Baconian method. After the turmoil of the English
Civil War, the Royal Society wanted to show that science did not threaten religious and political stability.
John Ray developed an influential natural theology of rational order; in his taxonomy, species were static and
fixed, their adaptation and complexity designed by God, and varieties showed minor differences caused by
local conditions. In God's benevolent design, carnivores caused mercifully swift death, but the suffering
caused by parasitism was a puzzling problem. The biological classification introduced by Carolus Linnaeus in
1735 also viewed species as fixed according to the divine plan. In 1766, Georges Buffon suggested that
some similar species, such as horses and asses, or lions, tigers, and leopards, might be varieties descended
from a common ancestor. The Ussher chronology of the 1650s had calculated creation at 4004 BC, but by
the 1780s geologists assumed a much older world. Wernerians thought strata were deposits from shrinking
seas, but James Hutton proposed a self-maintaining infinite cycle, anticipating uniformitarianism.[8]
Charles Darwin's grandfather Erasmus Darwin outlined a hypothesis of transmutation of species in
the 1790s, and Jean-Baptiste Lamarck published a more developed theory in 1809. Both envisaged that
spontaneous generation produced simple forms of life that progressively developed greater complexity,
adapting to the environment by inheriting changes in adults caused by use or disuse. This process was later
called Lamarckism. Lamarck thought there was an inherent progressive tendency driving organisms
continuously towards greater complexity, in parallel but separate lineages with no extinction.[9] Geoffroy
contended that embryonic development recapitulated transformations of organisms in past eras when the
environment acted on embryos, and that animal structures were determined by a constant plan as
demonstrated by homologies. Georges Cuvier strongly disputed such ideas, holding that unrelated, fixed
species showed similarities that reflected a design for functional needs.[10] His paleontological work in the
1790s had established the reality of extinction, which he explained by local catastrophes, followed by
repopulation of the affected areas by other species.[11]
In Britain, William Paley's Natural Theology saw adaptation as evidence of beneficial "design" by the
Creator acting through natural laws. All naturalists in English universities were Church of England clergymen,
and science became a search for these laws.[12] Geologists adapted catastrophism to show repeated
worldwide annihilation and creation of new fixed species adapted to a changed environment, initially
identifying the most recent catastrophe as the biblical flood.[13] Some anatomists such as Robert Grant were
influenced by Lamarck and Geoffroy, but most naturalists regarded their ideas of transmutation as a threat to
divinely appointed social order.[14]
[edit] Inception of Darwin's theory

See also: Charles Darwin's education and Inception of Darwin's theory
Darwin went to Edinburgh University in 1825 to study medicine. In his second year he neglected his
medical studies for natural history and spent four months assisting Robert Grant's research into marine
invertebrates. Grant revealed his enthusiasm for the transmutation of species, but Darwin rejected it.[15] At
Cambridge University starting in 1827, Darwin learnt science as natural theology from botanist John Stevens
Henslow, and read Paley, John Herschel and Alexander von Humboldt. Filled with zeal for science, he
studied catastrophist geology with Adam Sedgwick.[16][17]
In mid-July 1837 Darwin started his "B" notebook on Transmutation of Species, and on page 36
wrote "I think" above his first evolutionary tree.
In December 1831, he joined the Beagle expedition as a geologist and naturalist. He read Charles
Lyell's Principles of Geology and from the first stop ashore, at St. Jago, found Lyell's uniformitarianism a key
to the geological history of landscapes. Darwin discovered fossils resembling huge armadillos, and noted the
geographical distribution of modern species in hope of finding their "centre of creation".[18] The three
Fuegian missionaries the expedition returned to Tierra del Fuego were friendly and civilised, yet to Darwin
their relatives on the island seemed "miserable, degraded savages",[19] and he no longer saw an
unbridgeable gap between humans and animals.[20] As the Beagle neared England in 1836, he noted that
species might not be fixed.[21]
Richard Owen showed that fossils of extinct species Darwin found in South America were allied to
living species on the same continent. In March 1837, ornithologist John Gould announced that Darwin's Rhea
was a separate species from the previously described rhea (though their territories overlapped), that
mockingbirds collected on the Galápagos Islands represented three separate species each unique to a
particular island, and that several distinct birds from those islands were all classified as finches.[22] Darwin
began speculating, in a series of notebooks, on the possibility that "one species does change into another" to
explain these findings, and around July sketched a genealogical branching of a single evolutionary tree,
discarding Lamarck's independent lineages progressing to higher forms.[23] Unconventionally, Darwin asked
questions of fancy pigeon and animal breeders as well as established scientists. At the zoo he had his first
sight of an ape, and was profoundly impressed by how human the orangutan seemed.[24]
In late September 1838, he started reading Thomas Malthus's An Essay on the Principle of
Population with its statistical proof that human populations breed beyond their means and struggle to survive.
Darwin related this to the struggle for existence among wildlife and botanist de Candolle's "warring of the
species" in plants; he immediately envisioned "a force like a hundred thousand wedges" pushing well-
adapted variations into "gaps in the economy of nature", so that the survivors would pass on their form and
abilities, and unfavourable variations would be destroyed.[25][26] By December 1838, he had noted a
similarity between the act of breeders selecting traits and a Malthusian Nature selecting among variants
thrown up by "chance" so that "every part of newly acquired structure is fully practical and perfected".[27]
Darwin now had the framework of his theory of natural selection "by which to work",[28] but he was
fully occupied with his career as a geologist and held off writing a sketch of his theory until his book on The
Structure and Distribution of Coral Reefs was completed in May 1842.[29][30]
[edit] Further development

See also: Development of Darwin's theory
Darwin continued to research and extensively revise his theory while focusing on his main work of
publishing the scientific results of the Beagle voyage.[29] He tentatively wrote of his ideas to Lyell in January
1842;[31] then in June he roughed out a 35-page "Pencil Sketch" of his theory.[32] Darwin began
correspondence about his theorising with the botanist Joseph Dalton Hooker in January 1844, and by July
had rounded out his "sketch" into a 230-page "Essay", to be expanded with his research results and
published if he died prematurely.[33]
Darwin researched how the skulls of different pigeon breeds varied, as shown in his Variation of
Plants and Animals Under Domestication of 1868.
In November 1844, the anonymously published popular science book Vestiges of the Natural History
of Creation, written by Scottish journalist Robert Chambers, widened public interest in the concept of
transmutation of species. Vestiges used evidence from the fossil record and embryology to support the claim
that living things had progressed from the simple to the more complex over time. But it proposed a linear
progression rather than the branching common descent theory behind Darwin's work in progress, and it
ignored adaptation. Darwin read it soon after publication, and scorned its amateurish geology and zoology,
[34] but he carefully reviewed his own arguments after leading scientists, including Adam Sedgwick, attacked
its morality and scientific errors.[35] Vestiges had significant influence on public opinion, and the intense
debate helped to pave the way for the acceptance of the more scientifically sophisticated Origin by moving
evolutionary speculation into the mainstream. While few naturalists were willing to consider transmutation,
Herbert Spencer became an active proponent of Lamarckism and progressive development in the 1850s.[36]
Hooker was persuaded to take away a copy of the "Essay" in January 1847, and eventually sent a
page of notes giving Darwin much needed feedback. Reminded of his lack of expertise in taxonomy, Darwin
began an eight year study of barnacles, becoming the leading expert on their classification. Using his theory,
he discovered homologies showing that slightly changed body parts served different functions to meet new
conditions, and he found an intermediate stage in the evolution of distinct sexes.[37]
Darwin's barnacle studies convinced him that variation arose constantly and not just in response to
changed circumstances. In 1854, he completed the last part of his Beagle-related writing and began working
full-time on evolution. His thinking changed from the view that species formed in isolated populations only, as
on islands, to an emphasis on speciation without isolation; that is, he saw increasing specialisation within
large stable populations as continuously exploiting new ecological niches. He conducted empirical research
focusing on difficulties with his theory. He studied the developmental and anatomical differences between
different breeds of many domestic animals, became actively involved in fancy pigeon breeding, and
experimented (with the help of his son Francis) on ways that plant seeds and animals might disperse across
oceans to colonise distant islands. By 1856, his theory was much more sophisticated, with a mass of
supporting evidence.[37][38]
[edit] Publication
Main article: Publication of Darwin's theory
[edit] Events leading to publication
A photograph of Alfred Russel Wallace (1823–1913) taken in Singapore in 1862

An 1855 paper on the "introduction" of species, written by Alfred Russel Wallace, claimed that
patterns in the geographical distribution of species and fossils could be explained if every new species
always came into existence near an already existing, closely related species.[39] Charles Lyell recognised
the implications of Wallace's paper and its possible connection to Darwin's work, although Darwin did not,
and in the spring of 1856 Lyell urged Darwin to publish his theory to establish priority. Darwin was torn
between the desire to set out a full and convincing account and the pressure to quickly produce a short
paper. He decided he did not want to expose his ideas to review by an editor as would have been required to
publish in an academic journal. On 14 May 1856, he began a "sketch" account, and by July had decided to
produce a full technical treatise on species.[40]
Darwin was hard at work on his "big book" on Natural Selection, when on 18 June 1858 he received
a parcel from Wallace, who was working in Borneo. It enclosed twenty pages describing an evolutionary
mechanism, a response to Darwin's recent encouragement, with a request to send it on to Lyell if Darwin
thought it worthwhile. The mechanism was similar to Darwin's own theory.[40] Darwin wrote to Lyell that
"your words have come true with a vengeance, ... forestalled" and he would "of course, at once write and
offer to send [it] to any journal" that Wallace chose, adding that "all my originality, whatever it may amount to,
will be smashed".[41] Lyell and Hooker agreed that a joint paper should be presented at the Linnean Society,
and on 1 July 1858, the papers entitled On the Tendency of Species to form Varieties; and on the
Perpetuation of Varieties and Species by Natural Means of Selection , by Wallace and Darwin respectively,
were read out but drew little reaction. While Darwin considered Wallace's idea to be identical to his concept
of natural selection, historians have pointed out differences. Darwin described natural selection as being
analogous to the artificial selection practised by animal breeders, and emphasised competition between
individuals; Wallace drew no comparison to selective breeding, and focused on ecological pressures that
kept different varieties adapted to local conditions.[42][43][44]
After the meeting, Darwin decided to write "an abstract of my whole work".[45] He started work on 20
July 1858, while on holiday at Sandown,[46] and wrote parts of it from memory.[47] Lyell discussed
arrangements with publisher John Murray III, of the publishing house John Murray,[48] who responded
immediately to Darwin's letter of 31 March 1859[49] with an agreement to publish the book without even
seeing the manuscript, and an offer to Darwin of 2⁄3 of the profits.[50] (eventually Murray paid £180 to Darwin
for the 1st edition and by Darwin's death in 1882 the book was in its 6th edition, earning Darwin nearly
£3000.[51]) Darwin had initially decided to call it An abstract of an Essay on the Origin of Species and
Varieties Through natural selection , but with Murray's persuasion it was eventually changed to the snappier
title: On the Origin of Species, with the title page adding by Means of Natural Selection, or the Preservation
of Favoured Races in the Struggle for Life.[1] Here the term "races" is used as an alternative for "varieties"
and does not carry the modern connotation of human races—the first use in the book refers to "the several
races, for instance, of the cabbage" and proceeds to a discussion of "the hereditary varieties or races of our
domestic animals and plants".[52]
[edit] Time taken to publish

Darwin had his basic theory of natural selection "by which to work" by December 1838, yet almost
twenty years later, when Wallace's letter arrived on 18 June 1858, Darwin was still not ready to publish his
theory. It was long thought that Darwin avoided or delayed making his ideas public for personal reasons.
Reasons suggested have included fear of religious persecution or social disgrace if his views were revealed,
and concern about upsetting his clergymen naturalist friends or his pious wife Emma. Charles Darwin's
illness caused repeated delays. His paper on Glen Roy had proved embarrassingly wrong, and he may have
wanted to be sure he was correct. David Quammen has suggested all these factors may have contributed,
and notes Darwin's large output of books and busy family life during that time.[53]
A more recent study by science historian John van Wyhe has determined that the idea that Darwin
delayed publication only dates back to the 1940s, and Darwin's contemporaries thought the time he took was
reasonable. Darwin always finished one book before starting another. While he was researching, he told
many people about his interest in transmutation without causing outrage. He firmly intended to publish, but it
was not until September 1854 that he could work on it full time. His estimate that writing his "big book" would
take five years was optimistic.[54]
[edit] Publication and subsequent editions

On the Origin of Species was first published on Thursday 24 November 1859, priced at fifteen
shillings. The book had been offered to booksellers at Murray's autumn sale on Tuesday 22 November, and
all available copies had been taken up immediately. In total, 1,250 copies were printed but after deducting
presentation and review copies, and five for Stationers' Hall copyright, around 1,170 copies were available
for sale.[1] The second edition of 3,000 copies was quickly brought out on 7 January 1860,[55] and
incorporated numerous corrections as well as a response to religious objections by the addition of a new
epigraph on page ii, a quotation from Charles Kingsley, and the phrase "by the Creator" amended to the
closing sentence.[56] During Darwin's lifetime the book went through six editions, with cumulative changes
and revisions to deal with counter-arguments raised. The third edition came out in 1861, with a number of
sentences rewritten or added and an introductory appendix, An Historical Sketch of the Recent Progress of
Opinion on the Origin of Species,[57] while the fourth in 1866 had further revisions. The fifth edition,
published on 10 February 1869, incorporated more changes and for the first time included the phrase
"survival of the fittest", which had been coined by the philosopher Herbert Spencer in his Principles of
Biology (1864).[58]
In January 1871, George Jackson Mivart's On the Genesis of Species listed detailed arguments
against natural selection, and claimed it included false metaphysics.[59] Darwin made extensive revisions to
the sixth edition of the Origin (this was the first edition in which he used the word "evolution"[60]), and added
a new chapter VII, Miscellaneous objections, to address Mivart's arguments.[61] The sixth edition was
published by Murray on 19 February 1872 with "On" dropped from the title. Darwin had told Murray of
working men in Lancashire clubbing together to buy the 5th edition at fifteen shillings and wanted it made
more widely available; the price was halved to 7s 6d by printing in a smaller font. It includes a glossary
compiled by W.S. Dallas. Book sales increased from 60 to 250 per month.[61]
[edit] Publication outside Great Britain

American botanist Asa Gray (1810–1888)
In the United States, Asa Gray negotiated with a Boston publisher for publication of an authorised
American version, but learnt that two New York publishing firms were already planning to exploit the absence
of international copyright to print Origin.[62] Darwin was delighted by the popularity of the book, and asked
Gray to keep any profits.[63] Gray managed to negotiate a 5% royalty with Appleton's of New York,[64] who
got their edition out in mid January 1860, and the other two withdrew. In a May letter, Darwin mentioned a
print run of 2,500 copies, but it is not clear if this referred to the first printing only as there were four that year.
[1][65]
The book was widely translated in Darwin's life time, but problems arose with translating concepts
and metaphors, and some translations were biased by the translator's own agenda.[66] Darwin distributed
presentation copies in France and Germany, hoping that suitable applicants would come forward, as
translators were expected to make their own arrangements with a local publisher. He welcomed the
distinguished elderly naturalist and geologist Heinrich Georg Bronn, but the German translation published in
1860 imposed Bronn's own ideas, adding controversial themes that Darwin had deliberately omitted. Bronn
translated "favoured races" as "perfected races", and added essays on issues including the origin of life, as
well as a final chapter on religious implications partly inspired by Bronn's adherence to Naturphilosophie.[67]
In 1862, Bronn produced a second edition based on the third English edition and Darwin's suggested
additions, but then died of a heart attack.[68] Darwin corresponded closely with Julius Victor Carus, who
published an improved translation in 1867.[69] Darwin's attempts to find a translator in France fell through,
and the translation by Clémence Royer published in 1862 added an introduction praising Darwin's ideas as
an alternative to religious revelation and promoting ideas anticipating social Darwinism and eugenics, as well
as numerous explanatory notes giving her own answers to doubts that Darwin expressed. Darwin
corresponded with Royer about a second edition published in 1866 and a third in 1870, but he had difficulty
getting her to remove her notes and was troubled by these editions.[68][70] He remained unsatisfied until a
translation by Edmond Barbier was published in 1876.[1] A Dutch translation was published in 1860.[71] By
1864, additional translations had appeared in Italian and Russian.[66] In Darwin's lifetime, Origin was
published in Swedish in 1869, Danish in 1872, Polish in 1873, Hungarian in 1873–1874, Spanish in 1877 and
Serbian in 1878. By 1977, it had appeared in an additional 18 languages.[72]
[edit] Content
[edit] Title pages and introduction
John Gould's illustration of Darwin's Rhea was published in 1841. The existence of two rhea species
with overlapping ranges influenced Darwin.
Page ii contains quotations by William Whewell and Francis Bacon on the theology of natural laws,
[73] harmonising science and religion in accordance with Isaac Newton's belief in a rational God who
established a law-abiding cosmos.[74] In the second edition, Darwin added an epigraph from Joseph Butler
affirming that God could work through scientific laws as much as through miracles, in a nod to the religious
concerns of his oldest friends.[56] The Introduction establishes Darwin's credentials as a naturalist and
author,[75] then refers to John Herschel's letter suggesting that the origin of species "would be found to be a
natural in contradistinction to a miraculous process":[76]
WHEN on board H.M.S. 'Beagle,' as naturalist, I was much struck with certain facts in the
distribution of the inhabitants of South America, and in the geological relations of the present to
the past inhabitants of that continent. These facts seemed to me to throw some light on the
origin of species—that mystery of mysteries, as it has been called by one of our greatest
philosophers.[77]
Darwin refers specifically to the distribution of the species rheas, and to that of the Galápagos
tortoises and mockingbirds. He mentions his years of work on his theory, and the arrival of Wallace at the
same conclusion, which led him to "publish this Abstract" of his incomplete work. He outlines his ideas, and
sets out the essence of his theory:
As many more individuals of each species are born than can possibly survive; and as,
consequently, there is a frequently recurring struggle for existence, it follows that any being, if it
vary however slightly in any manner profitable to itself, under the complex and sometimes
varying conditions of life, will have a better chance of surviving, and thus be naturally selected.
From the strong principle of inheritance, any selected variety will tend to propagate its new and
modified form.[78]
Starting with the third edition, Darwin prefaced the introduction with a historical sketch that traced the
development of evolutionary ideas.[79] In that sketch he acknowledged that Patrick Matthew had, unknown
to Wallace or himself, anticipated the concept of natural selection in an appendix to a book published in
1831;[80] in the fourth edition he mentioned that William Charles Wells had done so as early as 1813.[81]
[edit] Variation under domestication and under nature

Chapter I covers animal husbandry and plant breeding, going back to ancient Egypt. Darwin
discusses contemporary opinions on the origins of different breeds under cultivation to argue that many have
been produced from common ancestors by selective breeding.[82] As an illustration of artificial selection, he
describes fancy pigeon breeding,[83] noting that "[t]he diversity of the breeds is something astonishing", yet
all were descended from one species of rock pigeon.[84] Darwin saw two distinct kinds of variation: (1) rare
abrupt changes he called "sports" or "monstrosities" (example: ancon sheep with short legs), and (2)
ubiquitous small differences (example: slightly shorter or longer bill of pigeons).[85] Both types of hereditary
changes can be used by breeders. However, for Darwin the small changes were most important in evolution.
In Chapter II, Darwin specifies that the distinction between species and varieties is arbitrary, with
experts disagreeing and changing their decisions when new forms were found. He concludes that "a well-
marked variety may be justly called an incipient species" and that "species are only strongly marked and
permanent varieties".[86] He argues for the ubiquity of variation in nature.[87] Historians have noted that
naturalists had long been aware that the individuals of a species differed from one another, but had generally
considered such variations to be limited and unimportant deviations from the archetype of each species, that
archetype being a fixed ideal in the mind of God. Darwin and Wallace made variation among individuals of
the same species central to understanding the natural world.[83]
[edit] Struggle for existence, natural selection, and divergence

In Chapter III, Darwin asks how varieties "which I have called incipient species" become distinct
species, and in answer introduces the key concept he calls "natural selection";[88] in the fifth edition he adds,
"But the expression often used by Mr. Herbert Spencer, of the Survival of the Fittest, is more accurate, and is
sometimes equally convenient."[89]
Owing to this struggle for life, any variation, however slight and from whatever cause
proceeding, if it be in any degree profitable to an individual of any species, in its infinitely
complex relations to other organic beings and to external nature, will tend to the preservation of
that individual, and will generally be inherited by its offspring ... I have called this principle, by
which each slight variation, if useful, is preserved, by the term of Natural Selection, in order to
mark its relation to man's power of selection.[88]
He notes that both A. P. de Candolle and Charles Lyell had stated that all organisms are exposed to
severe competition. Darwin emphasises that he used the phrase "struggle for existence" in "a large and
metaphorical sense, including dependence of one being on another"; he gives examples ranging from plants
struggling against drought to plants competing for birds to eat their fruit and disseminate their seeds. He
describes the struggle resulting from population growth: "It is the doctrine of Malthus applied with manifold
force to the whole animal and vegetable kingdoms." He discusses checks to such increase including complex
ecological interdependencies, and notes that competition is most severe between closely related forms
"which fill nearly the same place in the economy of nature".[90]
Chapter IV details natural selection under the "infinitely complex and close-fitting ... mutual relations
of all organic beings to each other and to their physical conditions of life".[91] Darwin takes as an example a
country where a change in conditions led to extinction of some species, immigration of others and, where
suitable variations occurred, descendants of some species became adapted to new conditions. He remarks
that the artificial selection practised by animal breeders frequently produced sharp divergence in character
between breeds, and suggests that natural selection might do the same, saying:
But how, it may be asked, can any analogous principle apply in nature? I believe it can and does
apply most efficiently, from the simple circumstance that the more diversified the descendants
from any one species become in structure, constitution, and habits, by so much will they be
better enabled to seize on many and widely diversified places in the polity of nature, and so be
enabled to increase in numbers.[92]
Historians have remarked that here Darwin anticipated the modern concept of an ecological niche.
[93] He did not suggest that every favourable variation must be selected, nor that the favoured animals were
better or higher, but merely more adapted to their surroundings.
This tree diagram, used to show the divergence of species, is the only illustration in the Origin of
Species.
Darwin proposes sexual selection, driven by competition between males for mates, to explain
sexually dimorphic features such as lion manes, deer antlers, peacock tails, bird songs, and the bright
plumage of some male birds.[94] He analysed sexual selection more fully in The Descent of Man, and
Selection in Relation to Sex (1871). Natural selection was expected to work very slowly in forming new
species, but given the effectiveness of artificial selection, he could "see no limit to the amount of change, to
the beauty and infinite complexity of the coadaptations between all organic beings, one with another and with
their physical conditions of life, which may be effected in the long course of time by nature's power of
selection". Using a tree diagram and calculations, he indicates the "divergence of character" from original
species into new species and genera. He describes branches falling off as extinction occurred, while new
branches formed in "the great Tree of life ... with its ever branching and beautiful ramifications".[95]
[edit] Variation and heredity

In Darwin's time there was no agreed-upon model of heredity;[96] in Chapter I Darwin admitted, "The
laws governing inheritance are quite unknown."[97] He accepted a version of the inheritance of acquired
characteristics (which after Darwin's death came to be called Lamarckism), and Chapter V discusses what he
called the effects of use and disuse; he wrote that he thought "there can be little doubt that use in our
domestic animals strengthens and enlarges certain parts, and disuse diminishes them; and that such
modifications are inherited", and that this also applied in nature.[98] Darwin stated that some changes that
were commonly attributed to use and disuse, such as the loss of functional wings in some island dwelling
insects, might be produced by natural selection. In later additions of Origin, Darwin expanded the role
attributed to the inheritance of acquired characteristics. Darwin also admitted ignorance of the source of
inheritable variations, but speculated they might be produced by environmental factors.[99][100] However,
one thing was clear: whatever the exact nature and causes of new variations, Darwin knew from observation
and experiment that breeders were able to select such variations and produce huge differences in many
generations of selection.[85] The observation that selection works in domestic animals is not destroyed by
lack of understanding of the underlying hereditary mechanism.
Breeding of animals and plants showed related varieties varying in similar ways, or tending to revert
to an ancestral form, and similar patterns of variation in distinct species were explained by Darwin as
demonstrating common descent. He recounted how Lord Morton's mare apparently demonstrated telegony,
offspring inheriting characteristics of a previous mate of the female parent, and accepted this process as
increasing the variation available for natural selection.[101]
More detail was given in Darwin's 1868 book on The Variation of Animals and Plants under
Domestication, which tried to explain heredity through his hypothesis of pangenesis. Although Darwin had
privately questioned blending inheritance, he struggled with the theoretical difficulty that novel individual
variations would tend to blend into a populationa. However, inherited variation could be seen,[102] and
Darwin's concept of selection working on a population with a range of small variations was workable.[103] It
was not until the modern evolutionary synthesis in the 1930s and 1940s that a model of heredity became
completely integrated with a model of variation.[104]
[edit] Difficulties for the theory

Chapter VI begins by saying the next three chapters will address possible objections to the theory,
the first being that often no intermediate forms between closely related species are found, though the theory
implies such forms must have existed. Darwin attributed this to the competition between different forms,
combined with the small number of individuals of intermediate forms, often leading to extinction of such
forms.[105] The rest of the chapter deals with whether natural selection could produce complex specialised
structures, and the behaviours to use them, when it would be difficult to imagine how intermediate forms
could be functional. Darwin said:
Secondly, is it possible that an animal having, for instance, the structure and habits of a bat,
could have been formed by the modification of some animal with wholly different habits? Can we
believe that natural selection could produce, on the one hand, organs of trifling importance, such
as the tail of a giraffe, which serves as a fly-flapper, and, on the other hand, organs of such
wonderful structure, as the eye, of which we hardly as yet fully understand the inimitable
perfection?[106]
His answer was that in many cases animals exist with intermediate structures that are functional. He
presented flying squirrels, and flying lemurs as examples of how bats might have evolved from non-flying
ancestors.[107] He discussed various simple eyes found in invertebrates, starting with nothing more than an
optic nerve coated with pigment, as examples of how the vertebrate eye could have evolved. Darwin
concludes: "If it could be demonstrated that any complex organ existed, which could not possibly have been
formed by numerous, successive, slight modifications, my theory would absolutely break down. But I can find
out no such case."[108]
Chapter VII (of the first edition) addresses the evolution of instincts. His examples included two he
had investigated experimentally: slave-making ants and the construction of hexagonal cells by honey bees.
Darwin noted that some species of slave-making ants were more dependent on slaves than others, and he
observed that many ant species will collect and store the pupae of other species as food. He thought it
reasonable that species with an extreme dependency on slave workers had evolved in incremental steps. He
suggested that bees that make hexagonal cells evolved in steps from bees that made round cells, under
pressure from natural selection to economise wax. Darwin concluded:
Finally, it may not be a logical deduction, but to my imagination it is far more satisfactory to look
at such instincts as the young cuckoo ejecting its foster-brothers, —ants making slaves, —the
larvæ of ichneumonidæ feeding within the live bodies of caterpillars, —not as specially endowed
or created instincts, but as small consequences of one general law, leading to the advancement
of all organic beings, namely, multiply, vary, let the strongest live and the weakest die.[109]
Chapter VIII addresses the idea that species had special characteristics that prevented hybrids from
being fertile in order to preserve separately created species. Darwin said that, far from being constant, the
difficulty in producing hybrids of related species, and the viability and fertility of the hybrids, varied greatly,
especially among plants. Sometimes what were widely considered to be separate species produced fertile
hybrid offspring freely, and in other cases what were considered to be mere varieties of the same species
could only be crossed with difficulty. Darwin concluded: "Finally, then, the facts briefly given in this chapter do
not seem to me opposed to, but even rather to support the view, that there is no fundamental distinction
between species and varieties."[110]
In the sixth edition Darwin inserted a new chapter VII (renumbering the subsequent chapters) to
respond to criticisms of earlier editions, including the objection that many features of organisms were not
adaptive and could not have been produced by natural selection. He said some such features could have
been by-products of adaptive changes to other features, and that often features seemed non-adaptive
because their function was unknown, as shown by his book on Fertilisation of Orchids that explained how
their elaborate structures facilitated pollination by insects. Much of the chapter responds to George Jackson
Mivart's criticisms, including his claim that features such as baleen filters in whales, flatfish with both eyes on
one side and the camouflage of stick insects could not have evolved through natural selection because
intermediate stages would not have been adaptive. Darwin proposed scenarios for the incremental evolution
of each feature.[111]
[edit] Geologic record

Chapter IX deals with the fact that the geologic record appears to show forms of life suddenly arising,
without the innumerable transitional fossils expected from gradual changes. Darwin borrowed Charles Lyell's
argument in Principles of Geology that the record is extremely imperfect as fossilisation is a very rare
occurrence, spread over vast periods of time; since few areas had been geologically explored, there could
only be fragmentary knowledge of geological formations, and fossil collections were very poor. Evolved local
varieties which migrated into a wider area would seem to be the sudden appearance of a new species.
Darwin did not expect to be able to reconstruct evolutionary history, but continuing discoveries gave him well
founded hope that new finds would occasionally reveal transitional forms.[112][113] To show that there had
been enough time for natural selection to work slowly, he again cited Principles of Geology and other
observations based on sedimentation and erosion, including an estimate that erosion of The Weald had
taken 300 million years.[114] The initial appearance of entire groups of well developed organisms in the
oldest fossil-bearing layers, now known as the Cambrian explosion, posed a problem. Darwin had no doubt
that earlier seas had swarmed with living creatures, but stated that he had no satisfactory explanation for the
lack of fossils.[115] Fossil evidence of pre-Cambrian life has since been found, extending the history of life
back for billions of years.[116]
Chapter X examines whether patterns in the fossil record are better explained by common descent
and branching evolution through natural selection than by the individual creation of fixed species. Darwin
expected species to change slowly, but not at the same rate – some organisms such as Lingula were
unchanged since the earliest fossils. The pace of natural selection would depend on variability and change in
the environment.[117] This distanced his theory from Lamarckian laws of inevitable progress.[112] It has
been argued that this anticipated the punctuated equilibrium hypothesis,[113][118] but other scholars have
preferred to emphasise Darwin's commitment to gradualism.[119] He cited Richard Owen's findings that the
earliest members of a class were a few simple and generalised species with characteristics intermediate
between modern forms, and were followed by increasingly diverse and specialised forms, matching the
branching of common descent from an ancestor.[112] Patterns of extinction matched his theory, with related
groups of species having a continued existence until extinction, then not reappearing. Recently extinct
species were more similar to living species than those from earlier eras, and as he had seen in South
America, and William Clift had shown in Australia, fossils from recent geological periods resembled species
still living in the same area.[117]
[edit] Geographic distribution
Chapter XI deals with evidence from biogeography, starting with the observation that differences in
flora and fauna from separate regions cannot be explained by environmental differences alone; South
America, Africa, and Australia all have regions with similar climates at similar latitudes, but those regions
have very different plants and animals. The species found in one area of a continent are more closely allied
with species found in other regions of that same continent than to species found on other continents. Darwin
noted that barriers to migration played an important role in the differences between the species of different
regions. The coastal sea life of the Atlantic and Pacific sides of Central America had almost no species in
common even though the Isthmus of Panama was only a few miles wide. His explanation was a combination
of migration and descent with modification. He went on to say: "On this principle of inheritance with
modification, we can understand how it is that sections of genera, whole genera, and even families are
confined to the same areas, as is so commonly and notoriously the case."[120] Darwin explained how a
volcanic island formed a few hundred miles from a continent might be colonised by a few species from that
continent. These species would become modified over time, but would still be related to species found on the
continent, and Darwin observed that this was a common pattern. Darwin discussed ways that species could
be dispersed across oceans to colonise islands, many of which he had investigated experimentally.[121]
Chapter XII continues the discussion of biogeography. After a brief discussion of freshwater species,
it returns to oceanic islands and their peculiarities; for example on some islands roles played by mammals on
continents were played by other animals such as flightless birds or reptiles. The summary of both chapters
says:
... I think all the grand leading facts of geographical distribution are explicable on the theory of
migration (generally of the more dominant forms of life), together with subsequent modification
and the multiplication of new forms. We can thus understand the high importance of barriers,
whether of land or water, which separate our several zoological and botanical provinces. We
can thus understand the localisation of sub-genera, genera, and families; and how it is that
under different latitudes, for instance in South America, the inhabitants of the plains and
mountains, of the forests, marshes, and deserts, are in so mysterious a manner linked together
by affinity, and are likewise linked to the extinct beings which formerly inhabited the same
continent ... On these same principles, we can understand, as I have endeavoured to show, why
oceanic islands should have few inhabitants, but of these a great number should be endemic or
peculiar; ...[122]
[edit] Classification, morphology, embryology, rudimentary organs

Chapter XIII starts by observing that classification depends on species being grouped together in a
multilevel system of groups and sub groups based on varying degrees of resemblance. After discussing
classification issues, Darwin concludes:
All the foregoing rules and aids and difficulties in classification are explained, if I do not greatly
deceive myself, on the view that the natural system is founded on descent with modification; that
the characters which naturalists consider as showing true affinity between any two or more
species, are those which have been inherited from a common parent, and, in so far, all true
classification is genealogical; that community of descent is the hidden bond which naturalists
have been unconsciously seeking, ...[123]
Darwin discusses morphology, including the importance of homologous structures. He says, "What
can be more curious than that the hand of a man, formed for grasping, that of a mole for digging, the leg of
the horse, the paddle of the porpoise, and the wing of the bat, should all be constructed on the same pattern,
and should include the same bones, in the same relative positions?"[124] He notes that animals of the same
class often have extremely similar embryos. Darwin discusses rudimentary organs, such as the wings of
flightless birds and the rudiments of pelvis and leg bones found in some snakes. He remarks that some
rudimentary organs, such as teeth in baleen whales, are found only in embryonic stages.
[edit] Concluding remarks

The final chapter reviews points from earlier chapters, and Darwin concludes by hoping that his
theory might produce revolutionary changes in many fields of natural history. Although he avoids the
controversial topic of human origins in the rest of the book so as not to prejudice readers against his theory,
here he ventures a cautious hint that psychology would be put on a new foundation and that "Light will be
thrown on the origin of man".[125] Darwin ends with a passage that became well known and much quoted:
It is interesting to contemplate an entangled bank, clothed with many plants of many kinds, with
birds singing on the bushes, with various insects flitting about, and with worms crawling through
the damp earth, and to reflect that these elaborately constructed forms, so different from each
other, and dependent on each other in so complex a manner, have all been produced by laws
acting around us ... Thus, from the war of nature, from famine and death, the most exalted
object which we are capable of conceiving, namely, the production of the higher animals,
directly follows. There is grandeur in this view of life, with its several powers, having been
originally breathed into a few forms or into one; and that, whilst this planet has gone cycling on
according to the fixed law of gravity, from so simple a beginning endless forms most beautiful
and most wonderful have been, and are being, evolved.[126]
[edit] Structure and style
[edit] Nature and structure of Darwin's argument
Darwin's aims were twofold: to show that species had not been separately created, and to show that
natural selection had been the chief agent of change.[127] He knew that his readers were already familiar
with the concept of transmutation of species from Vestiges, and his introduction ridicules that work as failing
to provide a viable mechanism.[128] Therefore the first four chapters lay out his case that selection in nature,
caused by the struggle for existence, is analogous to the selection of variations under domestication, and that
the accumulation of adaptive variations provides a scientifically testable mechanism for evolutionary
speciation.[129][130]
Later chapters provide evidence that evolution has occurred, supporting the idea of branching,
adaptive evolution without directly proving that selection is the mechanism. Darwin presents supporting facts
drawn from many disciplines, showing that his theory could explain a myriad of observations from many fields
of natural history that were inexplicable under the alternate concept that species had been individually
created.[130][131][132] The structure of Darwin's argument showed the influence of John Herschel, whose
philosophy of science maintained that a mechanism could be called a vera causa (true cause) if three things
could be demonstrated: its existence in nature, its ability to produce the effects of interest, and its ability to
explain a wide range of observations.[133]
[edit] Literary style

The Examiner review of 3 December 1859 commented, "Much of Mr. Darwin's volume is what
ordinary readers would call 'tough reading;' that is, writing which to comprehend requires concentrated
attention and some preparation for the task. All, however, is by no means of this description, and many parts
of the book abound in information, easy to comprehend and both instructive and entertaining."[128][134]
While the book was readable enough to sell, its dryness ensured that it was seen as aimed at
specialist scientists and could not be dismissed as mere journalism or imaginative fiction. Unlike the still-
popular Vestiges, it avoided the narrative style of the historical novel and cosmological speculation, though
the closing sentence clearly hinted at cosmic progression. Darwin had long been immersed in the literary
forms and practices of specialist science, and made effective use of his skills in structuring arguments.[128]
David Quammen has described the book as written in everyday language for a wide audience, but noted that
Darwin's literary style was uneven: in some places he used convoluted sentences that are difficult to read; in
other places his writing was beautiful. Quammen advised that later editions were weakened by Darwin
making concessions and adding details to address his critics, and recommended the first edition.[135] James
T. Costa said that because the book was an abstract produced in haste in response to Wallace's essay, it
was more approachable than the big book on natural selection Darwin had been working on, which would
have been encumbered by scholarly footnotes and much more technical detail. He added that parts of Origin
are dense, but parts are almost lyrical, and the case studies and observations are presented in a narrative
style unusual in serious scientific books, which broadened its audience.[136]
[edit] Reception
See also: Reaction to Darwin's theory
British cartoonists presented Darwin's theory in an unthreatening way. In the 1870s iconic caricatures
of Darwin with an ape or monkey body emphasised his significance in transforming ideas, and contributed to
widespread identification of evolutionism with Darwinism.[137]
The book aroused international interest[138] and a widespread debate, with no sharp line between
scientific issues and ideological, social and religious implications.[139] Much of the initial reaction was
hostile, but Darwin had to be taken seriously as a prominent and respected name in science. There was
much less controversy than had greeted the 1844 publication Vestiges of Creation, which had been rejected
by scientists,[138] but had influenced a wide public readership into believing that nature and human society
were governed by natural laws.[25] The Origin of Species as a book of wide general interest became
associated with ideas of social reform. Its proponents made full use of a surge in the publication of review
journals, and it was given more popular attention than almost any other scientific work, though it failed to
match the continuing sales of Vestiges.[140] Darwin's book legitimised scientific discussion of evolutionary
mechanisms, and the newly coined term Darwinism was used to cover the whole range of evolutionism, not
just his own ideas. By the mid 1870s, evolutionism was triumphant.[139]
With the exception of a brief hint in the final chapter, Darwin had avoided the subject of human
evolution. Despite this, the first review claimed it made a creed of the "men from monkeys" idea from
Vestiges.[141] Human evolution became central to the debate and was strongly argued by Huxley who
featured it in his popular "working-men's lectures". Darwin did not publish his own views on this until 1871.
[142] The naturalism of natural selection conflicted with presumptions of purpose in nature and while this
could be reconciled by theistic evolution, other mechanisms implying more progress or purpose were more
acceptable. Herbert Spencer had already incorporated Lamarckism into his popular philosophy of
progressive free market human society. He popularised the terms evolution and survival of the fittest, and
many thought Spencer was central to evolutionary thinking.[143]
[edit] Impact on the scientific community

See also: History of evolutionary thought
Scientific readers were already aware of arguments that species changed through processes that
were subject to laws of nature, but the transmutational ideas of Lamarck and the vague "law of development"
of Vestiges had not found scientific favour. Darwin presented natural selection as a scientifically testable
mechanism while accepting that other mechanisms such as inheritance of acquired characters were
possible. His strategy established that evolution through natural laws was worthy of scientific study, and by
1875, most scientists accepted that evolution occurred but few thought natural selection was significant.
Darwin's scientific method was also disputed, with his proponents favouring the empiricism of John Stuart
Mill's A System of Logic, while opponents held to the idealist school of William Whewell's Philosophy of the
Inductive Sciences, in which investigation could begin with the intuitive truth that species were fixed objects
created by design.[144] Early support for Darwin's ideas came from the findings of field naturalists studying
biogeography and ecology, including Joseph Dalton Hooker in 1860, and Asa Gray in 1862. Henry Walter
Bates presented research in 1861 that explained insect mimicry using natural selection. Alfred Russel
Wallace discussed evidence from his Malay archipelago research, including an 1864 paper with an
evolutionary explanation for the Wallace line.[145]
Huxley used illustrations to show that humans and apes had the same basic skeletal structure.[146]
Evolution had less obvious applications to anatomy and morphology, and at first had little impact on
the research of the anatomist Thomas Henry Huxley.[147] Despite this, Huxley strongly supported Darwin on
evolution; though he called for experiments to show whether natural selection could form new species, and
questioned if Darwin's gradualism was sufficient without sudden leaps to cause speciation. Huxley wanted
science to be secular, without religious interference, and his article in the April 1860 Westminster Review
promoted scientific naturalism over natural theology,[148][149] praising Darwin for "extending the domination
of Science over regions of thought into which she has, as yet, hardly penetrated" and coining the term
"Darwinism" as part of his efforts to secularise and professionalise science.[150] Huxley gained influence,
and initiated the X Club, which used the journal Nature to promote evolution and naturalism, shaping much of
late Victorian science. Later, the German morphologist Ernst Haeckel would convince Huxley that
comparative anatomy and palaeontology could be used to reconstruct evolutionary genealogies.[147][151]
The leading naturalist in Britain was the anatomist Richard Owen, an idealist who had shifted to the
view in the 1850s that the history of life was the gradual unfolding of a divine plan.[152] Owen's review of the
Origin in the April 1860 Edinburgh Review bitterly attacked Huxley, Hooker and Darwin, but also signalled
acceptance of a kind of evolution as a teleological plan in a continuous "ordained becoming", with new
species appearing by natural birth. Others that rejected natural selection, but supported "creation by birth",
included the Duke of Argyll who explained beauty in plumage by design.[153] Since 1858, Huxley had
emphasised anatomical similarities between apes and humans, contesting Owen's view that humans were a
separate sub-class. Their disagreement over human origins came to the fore at the British Association for the
Advancement of Science meeting featuring the legendary 1860 Oxford evolution debate.[154] In two years of
acrimonious public dispute that Charles Kingsley satirised as the "Great Hippocampus Question" and
parodied in The Water-Babies as the "great hippopotamus test", Huxley showed that Owen was incorrect in
asserting that ape brains lacked a structure present in human brains.[155] Others, including Charles Lyell
and Alfred Russel Wallace, thought that humans shared a common ancestor with apes, but higher mental
faculties could not have evolved through a purely material process. Darwin published his own explanation in
the Descent of Man (1871).[156]
[edit] Impact outside Great Britain
Haeckel showed a main trunk leading to mankind with minor branches to various animals, unlike
Darwin's branching evolutionary tree.[157]
Evolutionary ideas, although not natural selection, were accepted by German biologists accustomed
to ideas of homology in morphology from Goethe's Metamorphosis of Plants and from their long tradition of
comparative anatomy. Bronn's alterations in his German translation added to the misgivings of
conservatives, but enthused political radicals. Ernst Haeckel was particularly ardent, aiming to synthesise
Darwin's ideas with those of Lamarck and Goethe while still reflecting the spirit of Naturphilosophie.[67][158]
Their ambitious programme to reconstruct the evolutionary history of life was joined by Huxley and supported
by discoveries in palaeontology. Haeckel used embryology extensively in his recapitulation theory, which
embodied a progressive, almost linear model of evolution. Darwin was cautious about such histories, and
had already noted that von Baer's laws of embryology supported his idea of complex branching.[157]
Asa Gray promoted and defended Origin against those American naturalists with an idealist
approach, notably Louis Agassiz who viewed every species as a distinct fixed unit in the mind of the Creator,
classifying as species what others considered merely varieties. Edward Drinker Cope and Alpheus Hyatt
reconciled this view with evolutionism in a form of neo-Lamarckism involving recapitulation theory.[158]
French speaking naturalists in several countries showed appreciation of the much modified French
translation by Clémence Royer, but Darwin's ideas had little impact in France, where any scientists
supporting evolutionary ideas opted for a form of Lamarckism.[70] The intelligentsia in Russia had accepted
the general phenomenon of evolution for several years before Darwin had published his theory, and
scientists were quick to take it into account, although the Malthusian aspects were felt to be relatively
unimportant. The political economy of struggle was criticised as a British stereotype by Karl Marx and by Leo
Tolstoy, who had the character Levin in his novel Anna Karenina voice sharp criticism of the morality of
Darwin's views.[66]
[edit] Challenges to natural selection
There were serious scientific objections to the process of natural selection as the key mechanism of
evolution, including Karl von Nägeli's insistence that a trivial characteristic with no adaptive advantage could
not be developed by selection. Darwin conceded that these could be linked to adaptive characteristics. His
estimate that the age of the Earth allowed gradual evolution was disputed by William Thomson (later
awarded the title Lord Kelvin), who calculated that it had cooled in less than 100 million years. Darwin
accepted blending inheritance, but Fleeming Jenkin calculated that as it mixed traits, natural selection could
not accumulate useful traits. Darwin tried to meet these objections in the 5th edition. Mivart supported
directed evolution, and compiled scientific and religious objections to natural selection. In response, Darwin
made considerable changes to the sixth edition. The problems of the age of the Earth and heredity were only
resolved in the 20th century.[59][159]
By the mid 1870s, most scientists accepted evolution, but relegated natural selection to a minor role
as they believed evolution was purposeful and progressive. The range of evolutionary theories during "the
eclipse of Darwinism" included forms of "saltationism" in which new species were thought to arise through
"jumps" rather than gradual adaptation, forms of orthogenesis claiming that species had an inherent
tendency to change in a particular direction, and forms of neo-Lamarckism in which inheritance of acquired
characteristics led to progress. The minority view of August Weismann, that natural selection was the only
mechanism, was called neo-Darwinism. It was thought that the rediscovery of Mendelian inheritance
invalidated Darwin's views.[160][161]
[edit] Religious attitudes

The book produced a wide range of religious responses at a time of changing ideas and increasing
secularisation. The issues raised were complex and there was a large middle ground. Developments in
geology meant that there was little opposition based on a literal reading of Genesis,[162] but defence of the
argument from design and natural theology was central to debates over the book in the English speaking
world.[163][164]
The liberal theologian Baden Powell defended evolutionary ideas by arguing that the introduction of
new species should be considered a natural rather than a miraculous process.[165]
Natural theology was not a unified doctrine, and while some such as Louis Agassiz were strongly
opposed to the ideas in the book, others sought a reconciliation in which evolution was seen as purposeful.
[162] In the Church of England, some liberal clergymen interpreted natural selection as an instrument of
God's design, with the cleric Charles Kingsley seeing it as "just as noble a conception of Deity".[166][167] In
the second edition of January 1860, Darwin quoted Kingsley as "a celebrated cleric", and added the phrase
"by the Creator" to the closing sentence, which from then on read "life, with its several powers, having been
originally breathed by the Creator into a few forms or into one".[168] While some commentators have taken
this as a concession to religion that Darwin later regretted,[56] Darwin's view at the time was of God creating
life through the laws of nature,[169][170] and even in the first edition there are several references to
"creation".[171]
Baden Powell praised "Mr Darwin's masterly volume [supporting] the grand principle of the self-
evolving powers of nature".[172] In America, Asa Gray argued that evolution is the secondary effect, or
modus operandi, of the first cause, design,[173] and published a pamphlet defending the book in terms of
theistic evolution, Natural Selection is not inconsistent with Natural Theology .[166][174][175] Theistic
evolution became a popular compromise, and St. George Jackson Mivart was among those accepting
evolution but attacking Darwin's naturalistic mechanism. Eventually it was realised that supernatural
intervention could not be a scientific explanation, and naturalistic mechanisms such as neo-Lamarckism were
favoured over natural selection as being more compatible with purpose.[162]
Even though the book had barely hinted at human evolution, it quickly became central to the debate
as mental and moral qualities were seen as spiritual aspects of the immaterial soul, and it was believed that
animals did not have spiritual qualities. This conflict could be reconciled by supposing there was some
supernatural intervention on the path leading to humans, or viewing evolution as a purposeful and
progressive ascent to mankind's position at the head of nature.[162] While many conservative theologians
accepted evolution, Charles Hodge argued in his 1874 critique "What is Darwinism?" that "Darwinism",
defined narrowly as including rejection of design, was atheism though he accepted that Asa Gray did not
reject design.[176][177] Asa Gray responded that this charge misrepresented Darwin's text.[178] By the early
20th century, four noted authors of The Fundamentals were explicitly open to the possibility that God created
through evolution,[179] but fundamentalism inspired the American creation–evolution controversy that began
in the 1920s. Some conservative Roman Catholic writers and influential Jesuits opposed evolution in the late
19th and early 20th century, but other Catholic writers, starting with Mivart, pointed out that early Church
Fathers had not interpreted Genesis literally in this area.[180] The Vatican stated its official position in a 1950
papal encyclical, which stated that evolution was not inconsistent with Catholic teaching.[181][182]
[edit] Modern influence

A modern phylogenetic tree based on genome analysis shows the three-domain system.
Various alternative evolutionary mechanisms favoured during "the eclipse of Darwinism" became
untenable as more was learned about inheritance and mutation. The full significance of natural selection was
at last accepted in the 1930s and 1940s as part of the modern evolutionary synthesis. During that synthesis
biologists and statisticians, including R. A. Fisher, Sewall Wright and J.B.S. Haldane, merged Darwinian
selection with a statistical understanding of Mendelian genetics.[161]
Modern evolutionary theory continues to develop. Darwin's theory of evolution by natural selection,
with its tree-like model of branching common descent, has become the unifying theory of the life sciences.
The theory explains the diversity of living organisms and their adaptation to the environment. It makes sense
of the geologic record, biogeography, parallels in embryonic development, biological homologies, vestigiality,
cladistics, phylogenetics and other fields, with unrivalled explanatory power; it has also become essential to
applied sciences such as medicine and agriculture.[183][184] Despite the scientific consensus, a religion-
based political controversy has developed over how evolution is taught in schools, especially in the United
States.[185]
Interest in Darwin's writings continues, and scholars have generated an extensive literature, the
Darwin Industry, about his life and work. The text of Origin itself has been subject to much analysis including
a variorum, detailing the changes made in every edition, first published in 1959,[186] and a concordance, an
exhaustive external index published in 1981.[187] Worldwide commemorations of the 150th anniversary of
the publication of On the Origin of Species and the bicentenary of Darwin's birth were scheduled for 2009.
[188] They celebrate the ideas which "over the last 150 years have revolutionised our understanding of
nature and our place within it".[189]
[edit] See also
Book:Evolution

• Charles Darwin bibliography
• The Complete Works of Charles Darwin Online
• The Descent of Man, and Selection in Relation to Sex , published in 1871 and his second
great book on evolutionary theory.
• Transmutation of species
• Modern evolutionary synthesis
[edit] Notes
1. ^ a b c d e Freeman 1977
2. ^ Darwin 1859, p. iii
3. ^ Mayr 1982, pp. 479–480
4. ^ Darwin 1872, p. xiii
5. ^ Aristotle, Physics, translated by Hardie, R. P. and Gayle, R. K. and hosted by MIT's Internet
Classics Archive, http://classics.mit.edu/Aristotle/physics.2.ii.html, retrieved 2009-04-23
6. ^ Forster & Marston 1999, pp. 26–27
7. ^ Bowler 2003, pp. 27, 43, 45
8. ^ Bowler 2003, pp. 27–36, 39–42, 57–62, 67, 70, 77–80
9. ^ Bowler 2003, pp. 84–90
10.^ Desmond 1989, pp. 47–54
11.^ Bowler 2003, pp. 111–114
12.^ Browne 1995, pp. 91, 129
13.^ Bowler 2003, pp. 115–117
14.^ Desmond & Moore 1991, pp. 34–35
15.^ Browne 1995, pp. 80–88
16.^ Bowler 2003, pp. 148–149
17.^ Browne 1995, pp. 133–140
18.^ Larson 2004, pp. 59–62
19.^ Darwin 1845, pp. 205–208
20.^ Browne 1995, pp. 244–250
21.^ Keynes 2000, pp. xix–xx; Eldredge 2006
22.^ Quammen 2006, pp. 24–25
23.^ Herbert 1980, pp. 7–10. van Wyhe 2008, p. 44; Darwin's Notebook B: Transmutation of
species. pp. 1–13, 26, 36, 74, http://darwin-online.org.uk/content/frameset?
viewtype=side&itemID=CUL-DAR121.-&pageseq=1, retrieved 2009-03-16
24.^ Desmond & Moore 1991, pp. 240–244
25.^ a b van Wyhe 2009
26.^ Larson 2004, pp. 66–70; Darwin's Notebook D: Transmutation of species. pp. 134–135 ,
http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=CUL-DAR123.-&pageseq=112,
retrieved 2009-04-08
27.^ Darwin's Notebook E: Transmutation of species. p. 75, http://darwin-
online.org.uk/content/frameset?viewtype=text&itemID=CUL-DAR124.-&pageseq=63, retrieved 2009-
03-14
28.^ Darwin 1958, p. 120
29.^ a b van Wyhe 2007, pp. 186–187
30.^ Browne 1995, p. 436
31.^ Desmond & Moore 1991, p. 292
32.^ Browne 1995, pp. 436–437
33.^ van Wyhe 2007, p. 188
34.^ Darwin Correspondence Project – Letter 814—Darwin, C. R. to Hooker, J. D., (7 Jan 1845) ,
http://www.darwinproject.ac.uk/darwinletters/calendar/entry-814.html#back-mark-814.f5, retrieved
2008-11-24
35.^ Browne 1995, pp. 461–465
36.^ Bowler 2003, pp. 135–140
37.^ a b Bowler 2003, pp. 169–173
Darwin 1958, pp. 117–121
38.^ Quammen 2006, pp. 138–142
39.^ Wallace, Alfred R. (1855), "On the law which has regulated the introduction of new
species", Annals and Magazine of Natural History, including Zoology, Botany, and Geology 16: 184–
196, http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=A11&pageseq=1
40.^ a b Quammen 2006, pp. 135–158
41.^ Darwin Correspondence Project – Letter 2285—Darwin to Lyell (June 1858),
http://www.darwinproject.ac.uk/darwinletters/calendar/entry-2285.html, retrieved 2008-03-15
42.^ Larson 2004, pp. 74–75
43.^ Quammen 2006, pp. 162–163
44.^ Bowler 2003, pp. 175–176
45.^ "Letter 2303 — Darwin, C. R. to Hooker, J. D., 5 July (1858)". Darwin Correspondence
Project. http://www.darwinproject.ac.uk/entry-2303. Retrieved 2010-09-07.
46.^ Darwin 2006, p. 36 verso
47.^ "Letter 2432 — Darwin, C. R. to Hooker, J. D., 15 Mar (1859)". Darwin Correspondence
48.^ "Letter 2437 — Darwin, C. R. to Lyell, Charles, 28 Mar (1859)". Darwin Correspondence
49.^ "Letter 2441 — Darwin, C. R. to Murray, John (b), 31 Mar (1859)". Darwin Correspondence
50.^ "Letter 2443 — Murray, John (b) to Darwin, C. R., 1 Apr 1859". Darwin Correspondence
51.^ "Charles Darwin and his publisher". Darwin Correspondence Project. 2010.
http://www.darwinproject.ac.uk/charles-darwin-and-john-murray. Retrieved 2010-09-07.
52.^ Darwin 1859, p. 15
53.^ Quammen 2006, pp. 84–92
54.^ van Wyhe 2007
55.^ Darwin 1958, p. 122
56.^ a b c Browne 2002, pp. 95–96
57.^ Darwin 1861, p. xiii
58.^ "This survival of the fittest, which I have here sought to express in mechanical terms, is that
which Mr. Darwin has called 'natural selection', or the preservation of favoured races in the struggle
for life." Spencer 1864, pp. 444–445
59.^ a b Mivart 1871
60.^ Used in the transformist sense by Charles Lyell in 1832, Principles of Geology vol 2, p. 11;
and by Darwin in The Descent of Man in 1871, p. 2 onwards.
61.^ a b Desmond & Moore 1991, pp. 577, 582, 590, 592–593; Freeman 1977
62.^ Darwin Correspondence Project – Letter 2592—Darwin, C. R. to Gray, Asa, 21 Dec (1859) ,
http://www.darwinproject.ac.uk/darwinletters/calendar/entry-2592.html#mark-2592.f3, retrieved 2008-
12-06
63.^ Darwin Correspondence Project – Letter 2665—Darwin, C. R. to Gray, Asa, 28 Jan (1860) ,
http://www.darwinproject.ac.uk/darwinletters/calendar/entry-2665.html#back-mark-2665.f9, retrieved
2008-12-06
64.^ Darwin Correspondence Project – Letter 2706—Gray, Asa to Darwin, C. R., 20 Feb 1860 ,
http://www.darwinproject.ac.uk/darwinletters/calendar/entry-2706.html, retrieved 2008-12-06
65.^ Desmond & Moore 1991, p. 492
66.^ a b c Browne 2002, pp. 256–259
67.^ a b Browne 2002, pp. 140–142
68.^ a b Darwin Correspondence Project – The correspondence of Charles Darwin, volume 10:
1862, http://www.darwinproject.ac.uk/content/view/28/38/, retrieved 2009-03-06
69.^ Darwin Correspondence Project – The correspondence of Charles Darwin, volume 14:
1866, http://www.darwinproject.ac.uk/content/view/32/38/, retrieved 2009-03-06
70.^ a b Browne 2002, pp. 142–144
71.^ Ch. Darwin, Het ontstaan der soorten van dieren en planten door middel van de natuurkeus
of het bewaard blijven van bevoorregte rassen in de strijd des levens , transl. by T.C. Winkler
(Haarlem 1860) Source: Teyler, Winkler, Darwin Lecture by Marijn van Hoorn MA at the Congress of
the European Botanical and Horticultural Libraries Group, Prague, 23 April 2009
72.^ Freeman 1977, pp. 101–111
73.^ Darwin 1859, p. ii.
74.^ Phipps 1983
75.^ Secord 2000, p. 510
76.^ van Wyhe 2007, p. 197
77.^ Darwin 1859, p. 1.
78.^ Darwin 1859, p. 5.
79.^ Darwin & Costa 2009, p. 1
80.^ Darwin 1861, p. xiii.
81.^ Darwin 1866, p. xiv-xv.
82.^ Darwin & Costa 2009, p. 7
83.^ a b Quammen 2006, pp. 184–186
84.^ Darwin 1859, pp. 20–28
85.^ a b David Reznick (2009) The Origin Then and Now, Princeton University Press, p.49.
86.^ Darwin 1859, pp. 44–59 Chap. II.
87.^ Darwin & Costa 2009, p. 44
88.^ a b Darwin 1859, pp. 60–61 Chap. III
89.^ Darwin 1869, pp. 72
90.^ Darwin 1859, pp. 62–76
91.^ Darwin 1859, p. 80
92.^ Darwin 1859, p. 112
93.^ Quammen 2006, p. 189
94.^ Darwin 1859, pp. 87–101
95.^ Darwin 1859, pp. 117–130.
96.^ Larson 2004, p. 85
97.^ Darwin 1859, p. 13
98.^ Darwin 1859, p. 134.
99.^ Larson 2004, pp. 86–87
100.^ Darwin 1859, pp. 131–150
101.^ Quammen 2006, pp. 159–167
Darwin 1859, pp. 159–167
102.^ Richard Dawkins (8 February 2003). "An early flowering of genetics | Books". The
Guardian. http://www.guardian.co.uk/books/2003/feb/08/peopleinscience.evolution. Retrieved 2010-
10-24.
103.^ Bowler 2003, pp. 200–201
104.^ Bowler 1989
105.^ Darwin 1859, pp. 171–178
106.^ Darwin 1859, pp. 171–172
107.^ Darwin 1859, pp. 180–181
108.^ Darwin 1859, pp. 187–190
109.^ Darwin 1859, pp. 243–244
110.^ Darwin 1859, pp. 245–278
111.^ Darwin 1872, pp. 168–205
112.^ a b c Bowler 2003, p. 182
113.^ a b Wesley R. Elsberry (1996), Punctuated Equilibria, http://www.talkorigins.org/faqs/punc-
eq.html#pe-vs-pg, retrieved 2009-04-30
114.^ Darwin 1859, pp. 282–287
115.^ Darwin 1859, pp. 306–308
116.^ Schopf 2000
117.^ a b Darwin 1859, pp. 312–345
118.^ Rhodes 1987
119.^ Darwin & Costa 2009, p. 108
120.^ Darwin 1859, pp. 350–351
121.^ Darwin 1859, pp. 346–382
122.^ Darwin 1859, pp. 408–409
123.^ Darwin 1859, p. 420
124.^ Darwin 1859, p. 434
125.^ Browne 2002, pp. 60–61
Larson 2004, p. 92
Darwin & Costa 2009, p. 488
Quammen 2006, p. 196
Darwin 1859, p. 488
126.^ Darwin 1859, pp. 489–490
127.^ Darwin 1871, p. 152
128.^ a b c Secord 2000, pp. 508–511
129.^ Quammen 2006, pp. 183–188
130.^ a b Bowler 2003, pp. 180–181
131.^ Quammen 2006, pp. 190, 200–201
132.^ Larson 2004, pp. 88–89
133.^ Darwin & Costa 2009, p. xvii
134.^ Crawford 1859
135.^ Quammen 2006, pp. 176–181
136.^ Darwin & Costa 2009, p. ix
137.^ Browne 2002, pp. 376–379
138.^ a b van Wyhe 2008, pp. 48–49
139.^ a b Bowler 2003, pp. 177–180
140.^ Browne 2002, pp. 102–103
141.^ Browne 2002, p. 87
Leifchild 1859
142.^ Bowler 2003, pp. 207–209
Huxley 1863
143.^ Bowler 2003, pp. 203–207, 220–222
144.^ Bowler 2003, pp. 179–180, 197–198
145.^ Bowler 2003, pp. 183–184, 189
146.^ Bowler 2003, p. 208
147.^ a b Bowler 2003, pp. 184–185
148.^ Browne 2002, pp. 105–106
149.^ Huxley 1860
150.^ Bowler 2003, p. 184
151.^ Larson 2004, p. 108
152.^ Bowler 2003, pp. 124–126
153.^ Desmond & Moore 1991, pp. 490–491, 545–547
Secord 2000, p. 512
154.^ Lucas 1979
Desmond & Moore 1991, pp. 464–465, 493–499
155.^ Browne 2002, pp. 160–161
156.^ Bowler 2003, pp. 208–211, 214–216
157.^ a b Bowler 2003, pp. 169–170, 190–192
158.^ a b Bowler 2003, pp. 186–187, 237, 241
159.^ Bowler 2003, pp. 198–200, 234–236
160.^ Bowler 2003, p. 225
161.^ a b Quammen 2006, pp. 205–234
162.^ a b c d Bowler 2003, pp. 202–208
163.^ Dewey 1994, p. 26
164.^ Larson 2004, pp. 89–92
165.^ Bowler 2003, p. 139
166.^ a b Darwin and design: historical essay, Darwin Correspondence Project, 2007,
http://www.darwinproject.ac.uk/content/view/110/104/, retrieved 2008-09-17
167.^ Darwin Correspondence Project – Letter 2534—Kingsley, Charles to Darwin, C. R., 18 Nov
1859, http://www.darwinproject.ac.uk/darwinletters/calendar/entry-2534.html, retrieved 2009-04-11
168.^ Darwin 1860, p. 490.
169.^ Quammen 2006, p. 119
170.^ Moore 2006
171.^ Barlow 1963, p. 207.
172.^ Desmond & Moore 1991, pp. 487–488, 500
173.^ Dewey 1994, p. 27
174.^ Miles 2001
175.^ Gray, Asa (1860), "Natural Selection is not inconsistent with Natural Theology", Atlantic
Monthly (Darwin Correspondence Project – Essay: Natural selection & natural theology),
http://www.darwinproject.ac.uk/content/view/84/69/, retrieved 2009-04-11
176.^ Forster & Marston 1999, pp. 37–40
177.^ Hodge 1874, p. 177
178.^ Gray, Asa (28 May 1874), "What is Darwinism?", The Nation (Darwin Correspondence
Project), http://www.darwinproject.ac.uk/content/view/86/71/, retrieved 2009-03-03
179.^ Forster & Marston 1999, pp. 41–43
180.^ Bolwer 2003, pp. 323–324
181.^ Pius XII (1950), Humani Generis, Vatican,
http://www.vatican.va/holy_father/pius_xii/encyclicals/documents/hf_p-xii_enc_12081950_humani-
generis_en.html, retrieved 2009-06-08
182.^ Kreeft 2001, p. 49
183.^ Biography, The Complete Works of Charles Darwin Online, 21 January 2009,
http://darwin-online.org.uk/biography.html, retrieved 2009-04-23
184.^ Larson 2004, pp. 287–288, 295
185.^ Darwin & Costa 2009, p. x
186.^ Peckham 1959, recently reprinted.
187.^ Quammen 2006, pp. 179, 271–283
188.^ The ISTC of On the Origin of Species is A02-2009-00000001-4. As a tribute to its
influence, this work has been the first one to be registered by The International ISTC Agency.
189.^ Darwin 200: Celebrating Charles Darwin's bicentenary – What is Darwin200? , The Natural
History Museum, http://www.darwin200.org/what-is.html, retrieved 2009-04-23
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[edit] Contemporary reviews

• Carpenter, William Benjamin (1859), "Darwin on the Origin of Species", National Review 10
(December 1859): 188–214, http://darwin-online.org.uk/content/frameset?
• Gray, Asa (1860), "(Review of) The Origin of Species", Athenaeum (1710: 4 August 1860):
161, http://darwin-online.org.uk/content/frameset?itemID=A213&viewtype=text&pageseq=1 . Extract
from Proceedings of the American Academy of Arts and Sciences 4 (1860): 411–415.
• Huxley, Thomas Henry (1859), "Time and Life: Mr Darwin's "Origin of Species"", Macmillan's
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• Huxley, Thomas Henry (1859), "Darwin on the Origin of Species", The Times (26 December
1859): 8–9, http://darwin-online.org.uk/content/frameset?itemID=A166&viewtype=text&pageseq=1 .
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• Jenkin, Fleeming (1867), "(Review of) The Origin of Species", North British Review 46 (June
1867): 277–318, http://darwin-online.org.uk/content/frameset?
• Murray, Andrew (1860), "On Mr Darwin's Theory of the Origin of Species", Proceedings of
the Royal Society of Edinburgh 4: 274–291, http://darwin-online.org.uk/content/frameset?
itemID=A14&viewtype=text&pageseq=1 .
• Owen, Richard (1860), "Review of Darwin's Origin of Species", Edinburgh Review 3 (April
1860): 487–532, http://darwin-online.org.uk/content/frameset?
• Wilberforce, Samuel (1860), "(Review of) On the Origin of Species, by means of Natural
Selection; or the Preservation of Favoured Races in the Struggle for Life", Quarterly Review 108
(215: July 1860): 225–264, http://darwin-online.org.uk/content/frameset?
• For further reviews, see Darwin Online: Reviews & Responses to Darwin, Darwin Online, 10
March 2009, http://darwin-online.org.uk/reviews.html, retrieved 2009-06-18

Wikimedia Commons has media related to: The Origin of Species
Wikisource has original text related to this article:

• The Complete Works of Charles Darwin Online:
• Table of contents, bibliography of On the Origin of Species – links to text and images
of all six British editions of The Origin of Species, the 6th edition with additions and
corrections (final text), the first American edition, and translations into Danish, Dutch, French,
German, Polish, Russian and Spanish.
• Online Variorum, showing every change between the six British editions.
• The Origin of Species: An Outline , A short, accessible outline of the book.
• Origin of Species, full text with embedded audio.
• Victorian Science Texts
• Darwin Correspondence Project Home Page , University Library, Cambridge.
• PDF scans at Archive.org
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W000
Ordovician
The Ordovician /ɔrdəˈvɪʃən/ is a geologic period and system,
Ordovician Period the second of six of the Paleozoic Era,
488.3 – 443.7 million years ago
PreЄ
g
Mean atmospheric O2 content over ca. 13.5 Vol %[1]

period duration (68 % of modern level)
Mean atmospheric CO2 content ca. 4200 ppm[2]

over period duration (15 times pre-industrial level)
Mean surface temperature over ca. 16 °C[3]

period duration (2 °C above modern level)
180m; rising to 220m in Caradoc and falling sharply to

Sea level (above present day)
140m in end-Ordovician glaciations[4]
While recognition of the distinct Ordovician Period was slow in the United Kingdom, other areas of
the world accepted it quickly. It received international sanction in 1906, when it was adopted as an official
period of the Paleozoic Era by the International Geological Congress.
Contents
[hide]
• 1 Dating
• 2 Subdivisions
• 3 Paleogeography
• 4 Geochemistry
• 5 Climate and sea level
• 6 Life
• 6.1 Fauna
• 6.2 Flora
• 7 End of the period
• 8 References
[edit] Dating
The Ordovician Period started at a major extinction event called the Cambrian-Ordovician extinction
events some time about 488.3 ± 1.7 Mya (million years ago), and lasted for about 44.6 million years. It ended
with the Ordovician–Silurian extinction event, about 443.7 ± 1.5 Mya (ICS, 2004) that wiped out 60% of
marine genera.
The dates given are recent radiometric dates and vary slightly from those used in other sources. This
second period of the Paleozoic era created abundant fossils and in some regions, major petroleum and gas
reservoirs.
The boundary chosen for the beginning both of the Ordovician Period and the Tremadocian stage is
highly useful. Since it correlates well with the occurrence of widespread graptolite, conodont, and trilobite
species, the base of the Tremadocian allows scientists not only to relate these species to each other, but to
species that occur with them in other areas as well. This makes it easier to place many more species in time
relative to the beginning of the Ordovician Period.
[edit] Subdivisions
Key events in the Ordovician

-490 —
–
-485 —
–
-480 —
–
-475 —
–
-470 —
–
-465 —
–
-460 —
–
-455 —
–
-450 —
–
-445 —
–
Cambrian
Tremadocian
Floian
Dapingian
Darriwilian
Sandbian
Katian
Hirnantian
Hirnantian
Early Ordovician
Llanvirn
Llandeilo
Caradoc
Ashgill
Silurian
←
First land plant spores[6]
O
r
d
o
v
i
c
i
a
n
Palæozoic
Key events of the Ordovician Period.

Left: ICS approved stages.
Right: "General" stages.
A number of regional terms have been used to refer to subdivisions of the Ordovician Period. In
2008, the ICS erected a formal international system of subdivisions, illustrated to the right.[7]
The Ordovician Period in Britain was traditionally broken into Early (Tremadocian and Arenig), Middle
(Llanvirn [subdivided into Abereiddian and Llandeilian] and Llandeilo) and Late (Caradoc and Ashgill)
epochs. The corresponding rocks of the Ordovician System are referred to as coming from the Lower,
Middle, or Upper part of the column. The faunal stages (subdivisions of epochs) from youngest to oldest are:
• Hirnantian/Gamach (Late Ordovician: Ashgill)
• Rawtheyan/Richmond (Late Ordovician: Ashgill)
• Cautleyan/Richmond (Late Ordovician: Ashgill)
• Pusgillian/Maysville/Richmond (Late Ordovician: Ashgill)
• Trenton (Middle Ordovician: Caradoc)
• Onnian/Maysville/Eden (Middle Ordovician: Caradoc)
• Actonian/Eden (Middle Ordovician: Caradoc)
• Marshbrookian/Sherman (Middle Ordovician: Caradoc)
• Longvillian/Sherman (Middle Ordovician: Caradoc)
• Soundleyan/Kirkfield (Middle Ordovician: Caradoc)
• Harnagian/Rockland (Middle Ordovician: Caradoc)
• Costonian/Black River (Middle Ordovician: Caradoc)
• Chazy (Middle Ordovician: Llandeilo)
• Llandeilo (Middle Ordovician: Llandeilo)
• Whiterock (Middle Ordovician: Llanvirn)
• Llanvirn (Middle Ordovician: Llanvirn)
• Cassinian (Early Ordovician: Arenig)
• Arenig/Jefferson/Castleman (Early Ordovician: Arenig)
• Tremadoc/Deming/Gaconadian (Early Ordovician: Tremadoc)
[edit] Paleogeography
Sea levels were high during the Ordovician; in fact during the Tremadocian, marine transgressions
worldwide were the greatest for which evidence is preserved in the rocks.
During the Ordovician, the southern continents were collected into a single continent called
Gondwana. Gondwana started the period in equatorial latitudes and, as the period progressed, drifted toward
the South Pole. Early in the Ordovician, the continents Laurentia (present-day North America), Siberia, and
Baltica (present-day northern Europe) were still independent continents (since the break-up of the
supercontinent Pannotia earlier), but Baltica began to move towards Laurentia later in the period, causing the
Iapetus Ocean to shrink between them. The small continent Avalonia separated from Gondwana and began
to head north towards Baltica and Laurentia. The Rheic Ocean between Gondwana and Avalonia was
formed as a result.
A major mountain-building episode was the Taconic orogeny that was well under way in Cambrian
times. In the beginning of the Late Ordovician, from 460 to 450 Ma, volcanoes along the margin of the
Iapetus Ocean spewed massive amounts of carbon dioxide into the atmosphere, turning the planet into a
hothouse. These volcanic island arcs eventually collided with proto North America to form the Appalachian
mountains. By the end of the Late Ordovician these volcanic emissions had stopped. Gondwana had by that
time neared or approached the pole and was largely glaciated.
[edit] Geochemistry
The Ordovician was a time of calcite sea geochemistry in which low-magnesium calcite was the
primary inorganic marine precipitate of calcium carbonate. Carbonate hardgrounds were thus very common,
along with calcitic ooids, calcitic cements, and invertebrate faunas with dominantly calcitic skeletons.[8][9]
Unlike Cambrian times, when calcite production was dominated by microbial and non-biological
processes, animals (and macroalgae) became a dominant source of calcareous material in Ordovician
deposits.[10]
[edit] Climate and sea level

The Ordovician saw the highest sea levels of the Palaeozoic, and the low relief of the continents led
to many shelf deposits being formed under hundreds of metres of water.[10] Sea level rose more or less
continuously throughout the Early Ordovician, levelling off somewhat during the middle of the period.[10]
Locally, some regressions occurred, but sea level rise continued in the beginning of the Late Ordovician. A
change was soon on the cards, however, and sea levels fell steadily in accord with the cooling temperatures
for the ~30 million years leading up to the Hirnantian glaciation. Within this icy stage, sea level seems to
have risen and dropped somewhat, but despite much study the details remain unresolved.[10]
At the beginning of the period, around 480 million years ago, the climate was very hot due to high
levels of CO2, which gave a strong greenhouse effect. The marine waters are assumed to have been around
45°C, which restricted the diversification of complex multi-cellular organisms. But over time, the climate
become cooler, and around 460 million years ago, the ocean temperatures became comparable to those of
present day equatorial waters.[11]
As with North America and Europe, Gondwana was largely covered with shallow seas during the
Ordovician. Shallow clear waters over continental shelves encouraged the growth of organisms that deposit
calcium carbonates in their shells and hard parts. The Panthalassic Ocean covered much of the northern
hemisphere, and other minor oceans included Proto-Tethys, Paleo-Tethys, Khanty Ocean, which was closed
off by the Late Ordovician, Iapetus Ocean, and the new Rheic Ocean.
As the Ordovician progressed, we see evidence of glaciers on the land we now know as Africa and
South America. At the time these land masses were sitting at the South Pole, and covered by ice caps.
[edit] Life
Nautiloids like Orthoceras were among the largest predators in the Ordovician.
A diorama depicting Ordovician flora and fauna.

For most of the Late Ordovician, life continued to flourish, but at and near the end of the period there
were mass-extinction events that seriously affected planktonic forms like conodonts, graptolites, and some
groups of trilobites (Agnostida and Ptychopariida, which completely died out, and the Asaphida, which were
much reduced). Brachiopods, bryozoans and echinoderms were also heavily affected, and the endocerid
cephalopods died out completely, except for possible rare Silurian forms. The Ordovician-Silurian Extinction
Events may have been caused by an ice age that occurred at the end of the Ordovician period as the end of
the Late Ordovician was one of the coldest times in the last 600 million years of earth history.
[edit] Fauna
On the whole, the fauna that emerged in the Ordovician set the template for the remainder of the
Palaeozoic.[10] The fauna was dominated by tiered communities of suspension feeders, mainly with short
food chains; this said, the ecological system reached a new grade of complexity far beyond that of the
Cambrian fauna,[10] which has persisted until the present day.[10]
Though less famous than the Cambrian explosion, the Ordovician featured an adaptive radiation, the
Ordovician radiation, that was no less remarkable; marine faunal genera increased fourfold, resulting in 12%
of all known Phanerozoic marine fauna.[12] Another change in the fauna was the strong increase in filter
feeding organisms.[13] The trilobite, inarticulate brachiopod, archaeocyathid, and eocrinoid faunas of the
Cambrian were succeeded by those that dominated the rest of the Paleozoic, such as articulate brachiopods,
cephalopods, and crinoids. Articulate brachiopods, in particular, largely replaced trilobites in shelf
communities.[14] Their success epitomizes the greatly increased diversity of carbonate shell-secreting
organisms in the Ordovician compared to the Cambrian.[14]
In North America and Europe, the Ordovician was a time of shallow continental seas rich in life.
Trilobites and brachiopods in particular were rich and diverse. Although solitary corals date back to at least
the Cambrian, reef-forming corals appeared in the early Ordovician, corresponding to an increase in the
stability of carbonate and thus a new abundance of calcifying animals.[10]
Molluscs, which appeared during the Cambrian or even the Ediacaran, became common and varied,
especially bivalves, gastropods, and nautiloid cephalopods.
Now-extinct marine animals called graptolites thrived in the oceans. Some new cystoids and crinoids
appeared.
It was long thought that the first true vertebrates (fish — Ostracoderms) appeared in the Ordovician,
but recent discoveries in China reveal that they probably originated in the Early Cambrian. The very first
gnathostome (jawed fish) appeared in the Late Ordovician epoch.
During the Middle Ordovician there was a large increase in the intensity and diversity of bioeroding
organisms. This is known as the Ordovician Bioerosion Revolution.[15] It is marked by a sudden abundance
of hard substrate trace fossils such as Trypanites, Palaeosabella and Petroxestes.
In the Early Ordovician, trilobites were joined by many new types of organisms, including tabulate
corals, strophomenid, rhynchonellid, and many new orthid brachiopods, bryozoans, planktonic graptolites
and conodonts, and many types of molluscs and echinoderms, including the ophiuroids ("brittle stars") and
the first sea stars. Nevertheless the trilobites remained abundant, with all the Late Cambrian orders
continuing, and being joined by the new group Phacopida. The first evidence of land plants also appeared;
see Evolutionary history of life.
In the Middle Ordovician, the trilobite-dominated Early Ordovician communities were replaced by
generally more mixed ecosystems, in which brachiopods, bryozoans, molluscs, cornulitids, tentaculitids and
echinoderms all flourished, tabulate corals diversified and the first rugose corals appeared; trilobites were no
longer predominant. The planktonic graptolites remained diverse, with the Diplograptina making their
appearance. Bioerosion became an important process, particularly in the thick calcitic skeletons of corals,
bryozoans and brachiopods, and on the extensive carbonate hardgrounds that appear in abundance at this
time. One of the earliest known armoured agnathan ("ostracoderm") vertebrate, Arandaspis, dates from the
Middle Ordovician.
Trilobites in the Ordovician were very different than their predecessors in the Cambrian. Many
trilobites developed bizarre spines and nodules to defend against predators such as primitive sharks and
nautiloids while other trilobites such as Aeglina prisca evolved to become swimming forms. Some trilobites
even developed shovel-like snouts for ploughing through muddy sea bottoms. Another unusual clade of
trilobites known as the trinucleids developed a broad pitted margin around their head shields.[16] Some
trilobites such as Asaphus kowalewski evolved long eyestalks to assist in detecting predators whereas other
trilobite eyes in contrast disappeared completely.[17]
Outcrop of Upper
The Upper Fossil Mountain, Ordovician rubbly limestone
Ordovician edrioasteroid west-central Utah; Middle and shale, southern Indiana;
Cystaster stellatus on a Ordovician fossiliferous College of Wooster Outcrop of Upper
cobble from the Kope shales and limestones in the students. Ordovician limestone and
Formation in northern lower half. minor shale, central
Kentucky. In the background Tennessee; College of
is the cyclostome bryozoan
Wooster students.
Corynotrypa.
Trypanites borings
in an Ordovician Outcrop of Bryozoan fossils in
Petroxestes borings
hardground, southeastern Ordovician kukersite oil Ordovician kukersite oil
in an Ordovician
Indiana.[18] shale, northern Estonia. shale, northern Estonia.
hardground, southern Ohio.
[15]
Prasopora, a
Brachiopods and Platystrophia trepostome bryozoan from
bryozoans in an Ordovician ponderosa, Maysvillian the Ordovician of Iowa.
The Ordovician
limestone, southern
cystoid Echinosphaerites
(Upper Ordovician) near (an extinct echinoderm) from
Minnesota. Madison, Indiana. Scale bar northeastern Estonia;
is 5.0 mm. approximately 5 cm in
diameter.
The heliolitid coral

An Ordovician Zygospira modesta,
Protaraea richmondensis Graptolites
strophomenid brachiopod spiriferid brachiopods,
encrusting a gastropod; ( Amplexograptus ) from the
with encrusting inarticulate preserved in their original
Cincinnatian (Upper Ordovician near Caney
brachiopods and a positions on a trepostome
Ordovician) of southeastern Springs, Tennessee.
bryozoan. bryozoan; Cincinnatian
Indiana.
(Upper Ordovician) of
southeastern Indiana.
[edit] Flora
Green algae were common in the Late Cambrian (perhaps earlier) and in the Ordovician. Terrestrial
plants probably evolved from green algae, first appearing in the form of tiny non-vascular mosses resembling
liverworts. Fossil spores from land plants have been identified in uppermost Ordovician sediments.
Among the first land fungi may have been arbuscular mycorrhiza fungi (Glomerales), playing a
crucial role in facilitating the colonization of land by plants through mycorrhizal symbiosis, which makes
mineral nutrients available to plant cells; such fossilized fungal hyphae and spores from the Ordovician of
Wisconsin have been found with an age of about 460 million years ago, a time when the land flora most likely
only consisted of plants similar to non-vascular bryophytes.[19]
[edit] End of the period

Main article: Ordovician-Silurian extinction events
The Ordovician came to a close in a series of extinction events that, taken together, comprise the
second largest of the five major extinction events in Earth's history in terms of percentage of genera that went
extinct. The only larger one was the Permian-Triassic extinction event.
The extinctions occurred approximately 447–444 million years ago and mark the boundary between
the Ordovician and the following Silurian Period. At that time all complex multicellular organisms lived in the
sea, and about 49% of genera of fauna disappeared forever; brachiopods and bryozoans were greatly
reduced, along with many trilobite, conodont and graptolite families.
The most commonly accepted theory is that these events were triggered by the onset of most cold
conditions in the late Katian, followed by an ice age, in the Hirnantian faunal stage, that ended the long,
stable greenhouse conditions typical of the Ordovician.
The ice age was possibly not long-lasting, study of oxygen isotopes in fossil brachiopods showing
that its duration could have been only 0.5 to 1.5 million years.[9] Other researchers (Page et al.) estimate
more temperate conditions did not return until the late Silurian.
The late Ordovician glaciation event was preceded by a fall in atmospheric carbon dioxide (from
7000 ppm to 4400 ppm[citation needed]), which selectively affected the shallow seas where most organisms
lived. As the southern supercontinent Gondwana drifted over the South Pole, ice caps formed on it, which
have been detected in Upper Ordovician rock strata of North Africa and then-adjacent northeastern South
America, which were south-polar locations at the time.
Glaciation locks up water from the world-ocean, and the interglacials free it, causing sea levels
repeatedly to drop and rise; the vast shallow intra-continental Ordovician seas withdrew, which eliminated
many ecological niches, then returned carrying diminished founder populations lacking many whole families
of organisms, then withdrew again with the next pulse of glaciation, eliminating biological diversity at each
change.[20] Species limited to a single epicontinental sea on a given landmass were severely affected.[9]
Tropical lifeforms were hit particularly hard in the first wave of extinction, while cool-water species were hit
worst in the second pulse.[9]
Surviving species were those that coped with the changed conditions and filled the ecological niches
left by the extinctions.
At the end of the second event, melting glaciers caused the sea level to rise and stabilise once more.
The rebound of life's diversity with the permanent re-flooding of continental shelves at the onset of the
Silurian saw increased biodiversity within the surviving Orders.
Melott et al. (2006) suggested a ten-second gamma ray burst could have destroyed the ozone layer
and exposed terrestrial and marine surface-dwelling life to deadly radiation and initiated global cooling.[21]
[edit] References
4. ^ Haq, B. U.; Schutter, SR (2008). "A Chronology of Paleozoic Sea-Level Changes". Science
322 (5898): 64–68. doi:10.1126/science.1161648. PMID 18832639.
5. ^ Gradstein, Felix M.; Ogg, J. G.; Smith, A. G. (2004). A Geologic Time Scale 2004.
Cambridge: Cambridge University Press. ISBN 0521786738.
6. ^ Wellman, C.H., Gray, J. (2000). "The microfossil record of early land plants". Phil. Trans. R.
Soc. B 355 (1398): 717–732. doi:10.1098/rstb.2000.0612.
7. ^ Details on the Dapingian are available at Wang, X.; Stouge, S.; Chen, X.; Li, Z.; Wang, C.
(2009). "Dapingian Stage: standard name for the lowermost global stage of the Middle Ordovician
Series". Lethaia 42: 377–380. doi:10.1111/j.1502-3931.2009.00169.x. edit
8. ^ Template:Cite doi 10.1016/S0031-0182(98)00109-6
9. ^ a b c d Stanley, S. M.; Hardie, L. A. (1999). "Hypercalcification; paleontology links plate
tectonics and geochemistry to sedimentology". GSA Today 9: 1–7.
10.^ a b c d e f g h Munnecke, A.; Calner, M.; Harper, D. A. T.; Servais, T. (2010). "Ordovician and
Silurian sea-water chemistry, sea level, and climate: A synopsis". Palaeogeography,
Palaeoclimatology, Palaeoecology 296: 389. doi:10.1016/j.palaeo.2010.08.001. edit
11.^ Explosion in marine biodiversity explained by climate change
12.^ Dixon, Dougal; et al. (2001). Atlas of Life on Earth. New York: Barnes & Noble Books.
pp. 87. ISBN 0760719578.
13.^ Palaeos Paleozoic : Ordovician : The Ordovician Period
14.^ a b Cooper, John D.; Miller, Richard H.; Patterson, Jacqueline (1986). A Trip Through Time:
Principles of Historical Geology. Columbus: Merrill Publishing Company. pp. 247, 255–259.
ISBN 0675201403.
15.^ a b Wilson, M. A.; Palmer, T. J. (2006). "Patterns and processes in the Ordovician
Bioerosion Revolution" (PDF). Ichnos 13: 109–112. doi:10.1080/10420940600850505.
http://www3.wooster.edu/geology/WilsonPalmer06.pdf.
16.^ "Palaeos Paleozoic : Ordovician : The Ordovician Period". April 11, 2002.
http://www.palaeos.com/Paleozoic/Ordovician/Ordovician.htm#Life.
17.^ A Guide to the Orders of Trilobites
18.^ Wilson, M. A.; Palmer, T. J. (2001). "Domiciles, not predatory borings: a simpler
explanation of the holes in Ordovician shells analyzed by Kaplan and Baumiller, 2000". PALAIOS 16:
524–525. doi:10.1669/0883-1351(2001)016<0524:DNPBAS>2.0.CO;2.
19.^ Redecker, D.; Kodner, R. ; Graham, L. E. (2000). "Glomalean fungi from the Ordovician".
20.^ Emiliani (1992), 491
21.^ Melott, Adrian; et al. (2004). "Did a gamma-ray burst initiate the late Ordovician mass
extinction?". International Journal of Astrobiology 3: 55–61. doi:10.1017/S1473550404001910.

Wikimedia Commons has media related to: Ordovician
• Ogg, Jim (June 2004). "Overview of Global Boundary Stratotype Sections and Points
(GSSP's)". Archived from the original on 2006-04-23.
http://web.archive.org/web/20060423084018/http://www.stratigraphy.org/gssp.htm. Retrieved 2006-
04-30.
• Mehrtens, Charlotte. "Chazy Reef at Isle La Motte".
http://www.anr.state.vt.us/dec/geo/chazytxt.htm. An Ordovician reef in Vermont.
• Examples of Ordovician Fossils
• Ordovician fossils of the famous Cincinnatian Group
• The Dry Dredgers, an active group of amateur paleontologists in the Cincinnati area
Preceded
by Proterozoic Eon
Hadean

[hide] Paleozoic (542 – 251 Mya)
521 Mya)
Cambrian
(542 – 488.3 Mya)
Age 10* (492 – 488.3 Mya)

Floian (478.6 – 471.8 Mya)

Silurian Aeronian (439 – 436 Mya) · Telychian (436 – 428.2 Mya)
(443.7 – 416 Mya) Wenlock (428.2 – 422.9 Mya): Sheinwoodian (428.2 – 426.2 Mya) · Homerian
(426.2 – 422.9 Mya)
418.7 Mya)

(416 – 359.2 Mya) (391.8 – 385.3 Mya)
(374.5 – 359.2 Mya)

Carboniferous
(359.2 – 299 Mya)
(303.4 – 299 Mya)
Permian Cisuralian (299 – 270.6 Mya): Asselian (299 – 294.6 Mya) · Sakmarian
(299 – 251 Mya) (294.6 – 284.4 Mya) · Artinskian (284.4 – 275.6 Mya) · Kungurian (275.6 – 270.6
Mya)
(249.5 – 245.9 Mya)
(251 – 199.6 Mya) 228.7 Mya)
203.6 Mya) · Rhaetian (203.6 – 199.6 Mya)

– 175.6 Mya)
Mya)
Valanginian (140.2 – 133.9 Mya) · Hauterivian (133.9 – 130 Mya) · Barremian (130 –
Cretaceous 125 Mya) · Aptian (125 – 112 Mya) · Albian (112 – 99.6 Mya)
Mya)
Neogene Miocene (23.03 – 5.332 Mya): Aquitanian (23.03 – 20.43 Mya) · Burdigalian
(20.43 – 15.97 Mya) · Langhian (15.97 – 13.82 Mya) · Serravallian (13.82 – 11.608
(23.03 – 2.588 Mya)
Mya)

Quaternary
(2.588 – 0 Mya)
Dryas* (12.8 – 11.4 kya)

February 2008.
Retrieved from "http://en.wikipedia.org/wiki/Ordovician"
Categories: Ordovician
W000
Organism

"Life on Earth" redirects here. For the BBC series, see Life on Earth (TV series).
challenged and removed. (December 2010)
See also: Creature and Life form
Life on Earth
Temporal range: Archaean - Recent
These Escherichia coli cells provide an example of a prokaryotic microorganism
(unranked): Life on Earth (Gaeabionta)
Domains and Kingdoms
• Cellular life
• Bacteria
• Archaea
• Eukarya
• Bikonta
• Rhizaria
• Excavata
• Heterokonta
• Alveolata
• Plantae
• Unikonta
• Amoebozoa
• Fungi
• Animalia
• Non-cellular life (viruses) **
A polypore mushroom has parasitic relationship with its host

An ericoid mycorrhizal fungus
In biology, an organism is any contiguous living system (such as animal, plant, fungus, or micro-
organism). In at least some form, all organisms are capable of response to stimuli, reproduction, growth and
development, and maintenance of homoeostasis as a stable whole. An organism may either be unicellular
(single-celled) or be composed of, as in humans, many trillions of cells grouped into specialized tissues and
organs. The term multicellular (many-celled) describes any organism made up of more than one cell.
The term "organism" (Greek ὀργανισμός – organismos, from Ancient Greek ὄργανον – organon
"organ, instrument, tool") first appeared in the English language in 1701 and took on its current definition by
1834 (Oxford English Dictionary).
Scientific classification in biology considers organisms synonymous with "life on Earth". Based on
cell type, organisms may be divided into the prokaryotic and eukaryotic groups. The prokaryotes represent
two separate domains, the Bacteria and Archaea. Eukaryotic organisms, with a membrane-bounded cell
nucleus, also contain organelles, namely mitochondria and (in plants) plastids, generally considered to be
derived from endosymbiotic bacteria.[1] Fungi, animals and plants are examples of species that are
eukaryotes.
More recently a clade, Neomura, has been proposed, which groups together the Archaea and
Eukarya. Neomura is thought to have evolved from Bacteria, more specifically from Actinobacteria.[2]
Contents
[hide]
• 1 Semantics
• 1.1 Viruses
• 2 Organizational terminology
• 3 Chemistry
• 3.1 Macromolecules
• 4 Structure
• 4.1 The cell
• 5 Life span
• 6 Evolution
• 6.1 History of life
• 6.2 Horizontal gene transfer, and the history of life
• 6.3 Future of life (cloning and synthetic organisms)
• 7 Notes
[edit] Semantics
The word organism may broadly be defined as an assembly of molecules functioning as a more or
less stable whole which exhibits the properties of life. However, many sources propose definitions that
exclude viruses and theoretically possible man-made non-organic life forms.[3] Viruses are dependent on the
biochemical machinery of a host cell for reproduction.
Chambers Online Reference provides a broad definition: "any living structure, such as a plant,
animal, fungus or bacterium, capable of growth and reproduction".[4]
In multicellular terms, "organism" usually describes the whole hierarchical assemblage of systems
(for example circulatory, digestive, or reproductive) themselves collections of organs; these are, in turn,
collections of tissues, which are themselves made of cells. In some plants and the nematode Caenorhabditis
elegans, individual cells are totipotent.
A superorganism is an organism consisting of many individuals working together as a single
functional or social unit.
[edit] Viruses
Viruses are not typically considered to be organisms because they are incapable of "independent" or
autonomous reproduction or metabolism. This controversy is problematic because some cellular organisms
are also incapable of independent survival (but not of independent metabolism and procreation) and live as
obligatory intracellular parasites. Although viruses have a few enzymes and molecules characteristic of living
organisms, they have no metabolism of their own and cannot synthesize and organize the organic
compounds that form them. Naturally, this rules out autonomous reproduction and they can only be passively
replicated by the machinery of the host cell. In this sense they are similar to inanimate matter. While viruses
sustain no independent metabolism, and thus are usually not accounted organisms, they do have their own
genes and they do evolve by similar mechanisms by which organisms evolve.
[edit] Organizational terminology
definition by genus and differentia. Intermediate minor rankings are not shown.
All organisms are classified by the science of alpha taxonomy into either taxa or clades.
Taxa are ranked groups of organisms, which run from the general (domain) to the specific (species).
A broad scheme of ranks in hierarchical order is:
1. Domain
2. Kingdom
3. Phylum
4. Class
5. Order
6. Family
7. Genus
8. Species
To give an example, Homo sapiens is the Latin binomial equating to modern humans. All members of
the species sapiens are, at least in theory, genetically able to interbreed. Several species may belong to a
genus, but the members of different species within a genus are unable to interbreed to produce fertile
offspring. Homo, however, only has one surviving species (sapiens), Homo erectus, Homo neanderthalensis,
etc. having become extinct thousands of years ago. Several genera belong to the same family and so on up
the hierarchy. Eventually, the relevant kingdom (Animalia, in the case of humans) is placed into one of the
three domains depending upon certain genetic and structural characteristics.
All living organisms known to science are given classification by this system such that the species
within a particular family are more closely related and genetically similar than the species within a particular
phylum.
[edit] Chemistry
Organisms are complex chemical systems, organized in ways that promote reproduction and some
measure of sustainability or survival. The molecular phenomena of chemistry are fundamental in
understanding organisms, but it is a philosophical error (reductionism) to reduce organismal biology to mere
chemistry. It is generally the phenomena of entire organisms that determine their fitness to an environment
and therefore the survivability of their DNA-based genes.
Organisms clearly owe their origin, metabolism, and many other internal functions to chemical
phenomena, especially the chemistry of large organic molecules. Organisms are complex systems of
chemical compounds that, through interaction and environment, play a wide variety of roles.
Organisms are semi-closed chemical systems. Although they are individual units of life (as the
definition requires) they are not closed to the environment around them. To operate they constantly take in
and release energy. Autotrophs produce usable energy (in the form of organic compounds) using light from
the sun or inorganic compounds while heterotrophs take in organic compounds from the environment.
The primary chemical element in these compounds is carbon. The physical properties of this element
such as its great affinity for bonding with other small atoms, including other carbon atoms, and its small size
making it capable of forming multiple bonds, make it ideal as the basis of organic life. It is able to form small
three-atom compounds (such as carbon dioxide), as well as large chains of many thousands of atoms that
can store data (nucleic acids), hold cells together, and transmit information (protein).
[edit] Macromolecules
Compounds that make up organisms may be divided into macromolecules and other, smaller
molecules. The four groups of macromolecule are nucleic acids, proteins, carbohydrates and lipids. Nucleic
acids (specifically deoxyribonucleic acid, or DNA) store genetic data as a sequence of nucleotides. The
particular sequence of the four different types of nucleotides (adenine, cytosine, guanine, and thymine)
dictate the many characteristics that constitute the organism. The sequence is divided up into codons, each
of which is a particular sequence of three nucleotides and corresponds to a particular amino acid. Thus a
sequence of DNA codes for a particular protein that, due to the chemical properties of the amino acids it is
made from, folds in a particular manner and so performs a particular function.
These protein functions have been recognized:
1. Enzymes, which catalyze all of the reactions of metabolism
2. Structural proteins, such as tubulin, or collagen
3. Regulatory proteins, such as transcription factors or cyclins that regulate the cell cycle
4. Signaling molecules or their receptors such as some hormones and their receptors
5. Defensive proteins, which can include everything from antibodies of the immune system, to
toxins (e.g., dendrotoxins of snakes), to proteins that include unusual amino acids like canavanine
Lipids make up the membrane of cells that constitutes a barrier, containing everything within the cell
and preventing compounds from freely passing into, and out of, the cell. In some multicellular organisms they
serve to store energy and mediate communication between cells. Carbohydrates also store and transport
energy in some organisms, but are more easily broken down than lipids.
[edit] Structure
All organisms consist of monomeric units called cells; some contain a single cell (unicellular) and
others contain many units (multicellular). Multicellular organisms are able to specialize cells to perform
specific functions. A group of such cells is a tissue, the four basic types of which are epithelium, nervous
tissue, muscle tissue, and connective tissue. Several types of tissue work together in the form of an organ to
produce a particular function (such as the pumping of the blood by the heart, or as a barrier to the
environment as the skin). This pattern continues to a higher level with several organs functioning as an organ
system to allow for reproduction, digestion, etc. Many multicelled organisms consist of several organ
systems, which coordinate to allow for life.
[edit] The cell

The cell theory, first developed in 1839 by Schleiden and Schwann, states that all organisms are
composed of one or more cells; all cells come from preexisting cells; all vital functions of an organism occur
within cells, and cells contain the hereditary information necessary for regulating cell functions and for
transmitting information to the next generation of cells.
There are two types of cells, eukaryotic and prokaryotic. Prokaryotic cells are usually singletons,
while eukaryotic cells are usually found in multicellular organisms. Prokaryotic cells lack a nuclear membrane
so DNA is unbound within the cell, eukaryotic cells have nuclear membranes.
All cells, whether prokaryotic or eukaryotic, have a membrane, which envelops the cell, separates its
interior from its environment, regulates what moves in and out, and maintains the electric potential of the cell.
Inside the membrane, a salty cytoplasm takes up most of the cell volume. All cells possess DNA, the
hereditary material of genes, and RNA, containing the information necessary to build various proteins such
as enzymes, the cell's primary machinery. There are also other kinds of biomolecules in cells.
All cells share several abilities:[5]
• Reproduction by cell division (binary fission, mitosis or meiosis).
• Use of enzymes and other proteins coded for by DNA genes and made via messenger RNA
intermediates and ribosomes.
• Metabolism, including taking in raw materials, building cell components, converting energy,
molecules and releasing by-products. The functioning of a cell depends upon its ability to extract and
use chemical energy stored in organic molecules. This energy is derived from metabolic pathways.
• Response to external and internal stimuli such as changes in temperature, pH or nutrient
levels.
• Cell contents are contained within a cell surface membrane that contains proteins and a lipid
bilayer.
[edit] Life span

One of the basic parameters of an organism is its life span. Some organisms live as short as one
day, while some plants can live thousands of years. Aging is important when determining the life span of
most organisms, bacterium, a virus or even a prion.[citation needed]
[edit] Evolution
See also: Common descent and Origin of life
In biology, the theory of universal common descent proposes that all organisms on Earth are
descended from a common ancestor or ancestral gene pool. Evidence for common descent may be found in
traits shared between all living organisms. In Darwin's day, the evidence of shared traits was based solely on
visible observation of morphologic similarities, such as the fact that all birds have wings, even those that do
not fly.
Today, there is debate over whether or not all organisms descended from a common ancestor, or a
"last universal ancestor" (LUA), also called the "last universal common ancestor" (LUCA). The universality of
genetic coding suggests common ancestry. For example, every living cell makes use of nucleic acids as its
genetic material, and uses the same twenty amino acids as the building blocks for proteins, although
exceptions to the basic twenty amino acids have been found. However, throughout history groupings based
on appearance or function of species have sometimes been polyphyletic due to convergent evolution.
A hypothetical phylogenetic tree of all extant organisms, based on 16S rRNA gene sequence data,
showing the evolutionary history of the three domains of life, bacteria, archaea, and eukaryotes. Originally
proposed by Carl Woese.
The "last universal ancestor" (LUA), or "last universal common ancestor" (LUCA), is the name given to the
hypothetical unicellular organism or single cell that gave rise to all life on Earth 3.5 to 3.8 billion years ago;[6]
however, this hypothesis has since been refuted on many grounds. For example, it was once thought that the
genetic code was universal (see: Universal genetic code), but many variations have been discovered[7]
including various alternative mitochondrial codes.[8] Back in the early 1970s, evolutionary biologists thought
that a given piece of DNA specified the same protein subunit in every living thing, and that the genetic code
was thus universal. This was interpreted as evidence that every organism had inherited its genetic code from
a single common ancestor, aka, an LUCA. In 1979, however, exceptions to the code were found in
mitochondria, the tiny energy factories inside cells. Researchers studying human mitochondrial genes
discovered that they used an alternative code, and many slight variants have been discovered since,[7]
including various alternative mitochondrial codes,[8] as well as small variants such as Mycoplasma
translating the codon UGA as tryptophan. Biologists subsequently found exceptions in bacteria and in the
nuclei of algae and single-celled animals. For example, certain proteins may use alternative initiation (start)
codons not normally used by that species.[9] In certain proteins, non-standard amino acids are substituted for
standard stop codons, depending upon associated signal sequences in the messenger RNA: UGA can code
for selenocysteine and UAG can code for pyrrolysine. Selenocysteine is now viewed as the 21st amino acid,
and pyrrolysine is viewed as the 22nd. A detailed description of variations in the genetic code can be found at
the NCBI web site.
Information about the early development of life includes input from many different fields, including
geology and planetary science. These sciences provide information about the history of the Earth and the
changes produced by life. However, a great deal of information about the early Earth has been destroyed by
geological processes over the course of time.
[edit] History of life
Main article: Timeline of evolution
The chemical evolution from self-catalytic chemical reactions to life (see Origin of life) is not a part of
biological evolution, but it is unclear at which point such increasingly complex sets of reactions became what
we would consider, today, to be living organisms.
Precambrian stromatolites in the Siyeh Formation, Glacier National Park. In 2002, William Schopf of
UCLA published a controversial paper in the journal Nature arguing that formations such as this possess 3.5
billion year old fossilized algae microbes. If true, they would be the earliest known life on earth.
Not much is known about the earliest developments in life. However, all existing organisms share
certain traits, including cellular structure and genetic code. Most scientists interpret this to mean all existing
organisms share a common ancestor, which had already developed the most fundamental cellular
processes, but there is no scientific consensus on the relationship of the three domains of life (Archaea,
Bacteria, Eukaryota) or the origin of life. Attempts to shed light on the earliest history of life generally focus on
the behavior of macromolecules, particularly RNA, and the behavior of complex systems.
The emergence of oxygenic photosynthesis (around 3 billion years ago) and the subsequent
emergence of an oxygen-rich, non-reducing atmosphere can be traced through the formation of banded iron
deposits, and later red beds of iron oxides. This was a necessary prerequisite for the development of aerobic
cellular respiration, believed to have emerged around 2 billion years ago.
In the last billion years, simple multicellular plants and animals began to appear in the oceans. Soon
after the emergence of the first animals, the Cambrian explosion (a period of unrivaled and remarkable, but
brief, organismal diversity documented in the fossils found at the Burgess Shale) saw the creation of all the
major body plans, or phyla, of modern animals. This event is now believed to have been triggered by the
development of the Hox genes. About 500 million years ago, plants and fungi colonized the land, and were
soon followed by arthropods and other animals, leading to the development of today's land ecosystems.
The evolutionary process may be exceedingly slow. Fossil evidence indicates that the diversity and
complexity of modern life has developed over much of the history of the earth. Geological evidence indicates
that the Earth is approximately 4.6 billion years old. Studies on guppies by David Reznick at the University of
California, Riverside, however, have shown that the rate of evolution through natural selection can proceed
10 thousand to 10 million times faster than what is indicated in the fossil record.[10] Such comparative
studies however are invariably biased by disparities in the time scales over which evolutionary change is
measured in the laboratory, field experiments, and the fossil record.
[edit] Horizontal gene transfer, and the history of life
The ancestry of living organisms has traditionally been reconstructed from morphology, but is
increasingly supplemented with phylogenetics—the reconstruction of phylogenies by the comparison of
genetic (DNA) sequence.
Sequence comparisons suggest recent horizontal transfer of many genes among diverse
species including across the boundaries of phylogenetic "domains". Thus determining the
phylogenetic history of a species can not be done conclusively by determining evolutionary trees
for single genes.[11]
Biologist Gogarten suggests "the original metaphor of a tree no longer fits the data from recent
genome research", therefore "biologists [should] use the metaphor of a mosaic to describe the different
histories combined in individual genomes and use [the] metaphor of a net to visualize the rich exchange and
cooperative effects of HGT among microbes."[12]
[edit] Future of life (cloning and synthetic organisms)

In modern terms, the category of organism cloning refers to the procedure of creating a new
multicellular organism, genetically identical to another. However, cloning also has the potential of creating
entirely new species of organisms. Organism cloning is the subject of much ethical debate. (see Bioethics,
Ethics of cloning, and Designer baby articles)
The J. Craig Venter Institute has recently assembled a synthetic bacterial genome, Mycoplasma
genitalium, by using recombination in yeast of 25 overlapping DNA fragments in a single step. "The use of
yeast recombination greatly simplifies the assembly of large DNA molecules from both synthetic and natural
fragments."[13] Other companies, such as Synthetic Genomics, have already been formed to take advantage
of the many commercial uses of custom designed genomes.
[edit] Notes
1. ^ T.Cavalier-Smith (1987) The origin of eukaryote and archaebacterial cells, Annals of the
New York Academy of Sciences 503, 17–54
2. ^ T. Cavalier-Smith (2002) The neomuran origin of archaebacteria, the negibacterial root of
the universal tree and bacterial megaclassification. International Journal of Systematic and
Evolutionary Microbiology 52, 7–76
3. ^ "organism". Oxford English Dictionary (online ed.). 2004.
4. ^ "organism". Chambers 21st Century Dictionary (online ed.). 1999.
5. ^ The Universal Features of Cells on Earth in Chapter 1 of Molecular Biology of the Cell
fourth edition, edited by Bruce Alberts (2002) published by Garland Science.
6. ^ Doolittle, W. Ford (February, 2000). Uprooting the tree of life. Scientific American 282 (6):
90–95.
7. ^ a b NCBI: "The Genetic Codes", Compiled by Andrzej (Anjay) Elzanowski and Jim Ostell
8. ^ a b Jukes TH, Osawa S, The genetic code in mitochondria and chloroplasts., Experientia.
1990 Dec 1;46(11-12):1117-26.
9. ^ Genetic Code page in the NCBI Taxonomy section (Downloaded 27 April 2007.)
10.^ Evaluation of the Rate of Evolution in Natural Populations of Guppies (Poecilia reticulata)
"[1]"
11.^ Oklahoma State – Horizontal Gene Transfer
12.^ esalenctr.org
13.^ Gibsona, Daniel G.; Benders, Gwynedd A.; Axelroda, Kevin C.; et al. (2008). "One-step
assembly in yeast of 25 overlapping DNA fragments to form a complete synthetic Mycoplasma
genitalium genome". PNAS 105 (51): 20404–20409. doi:10.1073/pnas.0811011106. PMID 19073939.
PMC 2600582. http://www.pnas.org/content/105/51/20404.full.pdf.

Book:Organism

• BBCNews: 27 September, 2000, When slime is not so thick Citat: "It means that some of the
lowliest creatures in the plant and animal kingdoms, such as slime and amoeba, may not be as
primitive as once thought"
• SpaceRef.com, July 29, 1997: Scientists Discover Methane Ice Worms On Gulf Of
Mexico Sea Floor
• The Eberly College of Science: Methane Ice Worms discovered on Gulf of
Mexico Sea Floor download Publication quality photos
• Artikel, 2000: Methane Ice Worms: Hesiocaeca methanicola. Colonizing Fossil Fuel
Reserves
• SpaceRef.com, May 04, 2001: Redefining "Life as We Know it" Hesiocaeca
methanicola In 1997, Charles Fisher, professor of biology at Penn State, discovered this
remarkable creature living on mounds of methane ice under half a mile of ocean on the floor
of the Gulf of Mexico.
• BBCNews, 18 December, 2002, 'Space bugs' grown in lab Citat: "Bacillus simplex and
Staphylococcus pasteuri...Engyodontium album The strains cultured by Dr Wainwright seemed to be
resistant to the effects of UV - one quality required for survival in space"
• BBCNews, 19 June, 2003, Ancient organism challenges cell evolution Citat: "It appears that
this organelle has been conserved in evolution from prokaryotes to eukaryotes, since it is present in
both"
• Interactive Syllabus for General Biology - BI 04, Saint Anselm College, Summer 2003
• Jacob Feldman: Stramenopila
• NCBI Taxonomy entry: root (rich)
• Saint Anselm College: Survey of representatives of the major Kingdoms Citat: "Number of
kingdoms has not been resolved...Bacteria present a problem with their diversity...Protista present a
problem with their diversity...",
• Species 2000 Indexing the world's known species. Species 2000 has the objective of
enumerating all known species of plants, animals, fungi and microbes on Earth as the baseline
dataset for studies of global biodiversity. It will also provide a simple access point enabling users to
link from here to other data systems for all groups of organisms, using direct species-links.
• The largest organism in the world may be a fungus carpeting nearly 10 square kilometers of
an Oregon forest, and may be as old as 10500 years.
• The Tree of Life.
• Frequent questions from kids about life and their answers


Category · Portal
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Retrieved from "http://en.wikipedia.org/wiki/Organism"
Categories: Life | Organisms | Greek loanwords

W000
Photosynthesis

Composite image showing the global distribution of photosynthesis, including both oceanic
phytoplankton and vegetation
Overall equation for the type of photosynthesis that occurs in plants

Photosynthesis ( /foʊtoʊˈsɪnθəsɪs/; from the Greek φώτο- [photo-], "light," and σύνθεσις [synthesis],
"putting together", "composition") is a process that converts carbon dioxide into organic compounds,
especially sugars, using the energy from sunlight.[1] Photosynthesis occurs in plants, algae, and many
species of bacteria, but not in archaea. Photosynthetic organisms are called photoautotrophs, since they can
create their own food. In plants, algae, and cyanobacteria, photosynthesis uses carbon dioxide and water,
releasing oxygen as a waste product. Photosynthesis is vital for all aerobic life on Earth. As well as
maintaining the normal level of oxygen in the atmosphere, nearly all life either depends on it directly as a
source of energy, or indirectly as the ultimate source of the energy in their food[2] (the exceptions are
chemoautotrophs that live in rocks or around deep sea hydrothermal vents). The rate of energy capture by
photosynthesis is immense, approximately 100 terawatts,[3] which is about six times larger than the power
consumption of human civilization.[4] As well as energy, photosynthesis is also the source of the carbon in all
the organic compounds within organisms' bodies. In all, photosynthetic organisms convert around 100–115
teragrams of carbon into biomass per year.[5][6]
Although photosynthesis can happen in different ways in different species, some features are always
the same. For example, the process always begins when energy from light is absorbed by proteins called
photosynthetic reaction centers that contain chlorophylls. In plants, these proteins are held inside organelles
called chloroplasts, while in bacteria they are embedded in the plasma membrane. Some of the light energy
gathered by chlorophylls is stored in the form of adenosine triphosphate (ATP). The rest of the energy is used
to remove electrons from a substance such as water. These electrons are then used in the reactions that turn
carbon dioxide into organic compounds. In plants, algae and cyanobacteria, this is done by a sequence of
reactions called the Calvin cycle, but different sets of reactions are found in some bacteria, such as the
reverse Krebs cycle in Chlorobium. Many photosynthetic organisms have adaptations that concentrate or
store carbon dioxide. This helps reduce a wasteful process called photorespiration that can consume part of
the sugar produced during photosynthesis.
Overview of cycle between autotrophs and heterotrophs. Photosynthesis is the main means by which
plants, algae and many bacteria produce organic compounds and oxygen from carbon dioxide and water
(green arrow).
The first photosynthetic organisms probably evolved about 3,500 million years ago, early in the
evolutionary history of life, when all forms of life on Earth were microorganisms and the atmosphere had
much more carbon dioxide. They most likely used hydrogen or hydrogen sulfide as sources of electrons,
rather than water.[7] Cyanobacteria appeared later, around 3,000 million years ago, and drastically changed
the Earth when they began to oxygenate the atmosphere, beginning about 2,400 million years ago.[8] This
new atmosphere allowed the evolution of complex life such as protists. Eventually, no later than a billion
years ago, one of these protists formed a symbiotic relationship with a cyanobacterium, producing the
ancestor of many plants and algae.[9] The chloroplasts in modern plants are the descendants of these
ancient symbiotic cyanobacteria.[10]
Contents
[hide]
• 1 Overview
• 2 Photosynthetic membranes and organelles
• 3 Light reactions
• 3.1 Z scheme
• 3.2 Water photolysis
• 4 Light-independent reactions
• 4.1 The Calvin Cycle
• 4.2 Carbon concentrating mechanisms
• 4.2.1 On land
• 4.2.2 In water
• 5 Order and kinetics
• 6 Efficiency
• 7 Evolution
• 7.1 Symbiosis and the origin of chloroplasts
• 7.2 Cyanobacteria and the evolution of
photosynthesis
• 8 Discovery
• 9 Factors
• 9.1 Light intensity (irradiance), wavelength and
temperature
Overview
Photosynthesis changes the energy from the sun into chemical energy, splits water to liberate O 2,
and fixes CO2 into sugar.
Photosynthetic organisms are photoautotrophs, which means that they are able to synthesize food
directly from carbon dioxide using energy from light. However, not all organisms that use light as a source of
energy carry out photosynthesis, since photoheterotrophs use organic compounds, rather than carbon
dioxide, as a source of carbon.[2] In plants, algae and cyanobacteria, photosynthesis releases oxygen. This
is called oxygenic photosynthesis. Although there are some differences between oxygenic photosynthesis in
plants, algae and cyanobacteria, the overall process is quite similar in these organisms. However, there are
some types of bacteria that carry out anoxygenic photosynthesis, which consumes carbon dioxide but does
not release oxygen.
Carbon dioxide is converted into sugars in a process called carbon fixation. Carbon fixation is a
redox reaction, so photosynthesis needs to supply both a source of energy to drive this process, and the
electrons needed to convert carbon dioxide into carbohydrate, which is a reduction reaction. In general
outline, photosynthesis is the opposite of cellular respiration, where glucose and other compounds are
oxidized to produce carbon dioxide, water, and release chemical energy. However, the two processes take
place through a different sequence of chemical reactions and in different cellular compartments.
The general equation for photosynthesis is therefore:
2n CO2 + 2n H2O + photons → 2(CH2O)n + n O2 + 2n A
Carbon dioxide + electron donor + light energy → carbohydrate + oxygen + oxidized electron donor
Since water is used as the electron donor in oxygenic photosynthesis, the equation for this process
is:
2n CO2 + 2n H2O + photons → 2(CH2O)n + 2n O2
carbon dioxide + water + light energy → carbohydrate + oxygen

Other processes substitute other compounds (such as arsenite) for water in the electron-supply role;
the microbes use sunlight to oxidize arsenite to arsenate:[11] The equation for this reaction is:
(AsO33–) + CO2 + photons → CO + (AsO43–)[12]
carbon dioxide + arsenite + light energy → arsenate + carbon monoxide (used to build other
compounds in subsequent reactions)
Photosynthesis occurs in two stages. In the first stage, light-dependent reactions or light reactions
capture the energy of light and use it to make the energy-storage molecules ATP and NADPH. During the
second stage, the light-independent reactions use these products to capture and reduce carbon dioxide.
Most organisms that utilize photosynthesis to produce oxygen use visible light to do so, although at
least three use infrared radiation.[13]
Photosynthetic membranes and organelles

Chloroplast ultrastructure:
1. outer membrane
2. intermembrane space
3. inner membrane (1+2+3: envelope)
4. stroma (aqueous fluid)
5. thylakoid lumen (inside of thylakoid)
6. thylakoid membrane
7. granum (stack of thylakoids)
8. thylakoid (lamella)
9. starch
10. ribosome
11. plastidial DNA
12. plastoglobule (drop of lipids)
Main articles: Chloroplast and Thylakoid
The proteins that gather light for photosynthesis are embedded within cell membranes. The simplest
way these are arranged is in photosynthetic bacteria, where these proteins are held within the plasma
membrane.[14] However, this membrane may be tightly folded into cylindrical sheets called thylakoids,[15] or
bunched up into round vesicles called intracytoplasmic membranes.[16] These structures can fill most of the
interior of a cell, giving the membrane a very large surface area and therefore increasing the amount of light
that the bacteria can absorb.[15]
In plants and algae, photosynthesis takes place in organelles called chloroplasts. A typical plant cell
contains about 10 to 100 chloroplasts. The chloroplast is enclosed by a membrane. This membrane is
composed of a phospholipid inner membrane, a phospholipid outer membrane, and an intermembrane space
between them. Within the membrane is an aqueous fluid called the stroma. The stroma contains stacks
(grana) of thylakoids, which are the site of photosynthesis. The thylakoids are flattened disks, bounded by a
membrane with a lumen or thylakoid space within it. The site of photosynthesis is the thylakoid membrane,
which contains integral and peripheral membrane protein complexes, including the pigments that absorb light
energy, which form the photosystems.
Plants absorb light primarily using the pigment chlorophyll, which is the reason that most plants have
a green color. Besides chlorophyll, plants also use pigments such as carotenes and xanthophylls.[17] Algae
also use chlorophyll, but various other pigments are present as phycocyanin, carotenes, and xanthophylls in
green algae, phycoerythrin in red algae (rhodophytes) and fucoxanthin in brown algae and diatoms resulting
in a wide variety of colors.
These pigments are embedded in plants and algae in special antenna-proteins. In such proteins all
the pigments are ordered to work well together. Such a protein is also called a light-harvesting complex.
Although all cells in the green parts of a plant have chloroplasts, most of the energy is captured in the
leaves. The cells in the interior tissues of a leaf, called the mesophyll, can contain between 450,000 and
800,000 chloroplasts for every square millimeter of leaf. The surface of the leaf is uniformly coated with a
water-resistant waxy cuticle that protects the leaf from excessive evaporation of water and decreases the
absorption of ultraviolet or blue light to reduce heating. The transparent epidermis layer allows light to pass
through to the palisade mesophyll cells where most of the photosynthesis takes place.
Light reactions
Light-dependent reactions of photosynthesis at the thylakoid membrane

Main article: Light-dependent reactions
In the light reactions, one molecule of the pigment chlorophyll absorbs one photon and loses one
electron. This electron is passed to a modified form of chlorophyll called pheophytin, which passes the
electron to a quinone molecule, allowing the start of a flow of electrons down an electron transport chain that
leads to the ultimate reduction of NADP to NADPH. In addition, this creates a proton gradient across the
chloroplast membrane; its dissipation is used by ATP synthase for the concomitant synthesis of ATP. The
chlorophyll molecule regains the lost electron from a water molecule through a process called photolysis,
which releases a dioxygen (O2) molecule. The overall equation for the light-dependent reactions under the
conditions of non-cyclic electron flow in green plants is:[18]
2 H2O + 2 NADP+ + 3 ADP + 3 Pi + light → 2 NADPH + 2 H+ + 3 ATP + O2
Not all wavelengths of light can support photosynthesis. The photosynthetic action spectrum
depends on the type of accessory pigments present. For example, in green plants, the action spectrum
resembles the absorption spectrum for chlorophylls and carotenoids with peaks for violet-blue and red light.
In red algae, the action spectrum overlaps with the absorption spectrum of phycobilins for blue-green light,
which allows these algae to grow in deeper waters that filter out the longer wavelengths used by green
plants. The non-absorbed part of the light spectrum is what gives photosynthetic organisms their color (e.g.,
green plants, red algae, purple bacteria) and is the least effective for photosynthesis in the respective
organisms.
Z scheme
The "Z scheme"

In plants, light-dependent reactions occur in the thylakoid membranes of the chloroplasts and use
light energy to synthesize ATP and NADPH. The light-dependent reaction has two forms: cyclic and non-
cyclic. In the non-cyclic reaction, the photons are captured in the light-harvesting antenna complexes of
photosystem II by chlorophyll and other accessory pigments (see diagram at right). When a chlorophyll
molecule at the core of the photosystem II reaction center obtains sufficient excitation energy from the
adjacent antenna pigments, an electron is transferred to the primary electron-acceptor molecule, pheophytin,
through a process called photoinduced charge separation. These electrons are shuttled through an electron
transport chain, the so called Z-scheme shown in the diagram, that initially functions to generate a
chemiosmotic potential across the membrane. An ATP synthase enzyme uses the chemiosmotic potential to
make ATP during photophosphorylation, whereas NADPH is a product of the terminal redox reaction in the
Z-scheme. The electron enters a chlorophyll molecule in Photosystem I. The electron is excited due to the
light absorbed by the photosystem. A second electron carrier accepts the electron, which again is passed
down lowering energies of electron acceptors. The energy created by the electron acceptors is used to move
hydrogen ions across the thylakoid membrane into the lumen. The electron is used to reduce the co-enzyme
NADP, which has functions in the light-independent reaction. The cyclic reaction is similar to that of the non-
cyclic, but differs in the form that it generates only ATP, and no reduced NADP (NADPH) is created. The
cyclic reaction takes place only at photosystem I. Once the electron is displaced from the photosystem, the
electron is passed down the electron acceptor molecules and returns back to photosystem I, from where it
was emitted, hence the name cyclic reaction.
Water photolysis
Main articles: Photodissociation and Oxygen evolution
The NADPH is the main reducing agent in chloroplasts, providing a source of energetic electrons to
other reactions. Its production leaves chlorophyll with a deficit of electrons (oxidized), which must be
obtained from some other reducing agent. The excited electrons lost from chlorophyll in photosystem I are
replaced from the electron transport chain by plastocyanin. However, since photosystem II includes the first
steps of the Z-scheme, an external source of electrons is required to reduce its oxidized chlorophyll a
molecules. The source of electrons in green-plant and cyanobacterial photosynthesis is water. Two water
molecules are oxidized by four successive charge-separation reactions by photosystem II to yield a molecule
of diatomic oxygen and four hydrogen ions; the electron yielded in each step is transferred to a redox-active
tyrosine residue that then reduces the photoxidized paired-chlorophyll a species called P680 that serves as
the primary (light-driven) electron donor in the photosystem II reaction center. The oxidation of water is
catalyzed in photosystem II by a redox-active structure that contains four manganese ions and a calcium ion;
this oxygen-evolving complex binds two water molecules and stores the four oxidizing equivalents that are
required to drive the water-oxidizing reaction. Photosystem II is the only known biological enzyme that carries
out this oxidation of water. The hydrogen ions contribute to the transmembrane chemiosmotic potential that
leads to ATP synthesis. Oxygen is a waste product of light-dependent reactions, but the majority of
organisms on Earth use oxygen for cellular respiration, including photosynthetic organisms.[19][20]
Light-independent reactions
The Calvin Cycle
Main articles: Calvin cycle, Carbon fixation, and Light-independent reaction
In the Light-independent or dark reactions the enzyme RuBisCO captures CO2 from the atmosphere
and in a process that requires the newly formed NADPH, called the Calvin-Benson Cycle, releases three-
carbon sugars, which are later combined to form sucrose and starch. The overall equation for the light-
independent reactions in green plants is:[18]
3 CO2 + 9 ATP + 6 NADPH + 6 H+ → C3H6O3-phosphate + 9 ADP + 8 Pi + 6 NADP+ + 3 H2O
Overview of the Calvin cycle and carbon fixation
To be more specific, carbon fixation produces an intermediate product, which is then converted to the
final carbohydrate products. The carbon skeletons produced by photosynthesis are then variously used to
form other organic compounds, such as the building material cellulose, as precursors for lipid and amino acid
biosynthesis, or as a fuel in cellular respiration. The latter occurs not only in plants but also in animals when
the energy from plants gets passed through a food chain.
The fixation or reduction of carbon dioxide is a process in which carbon dioxide combines with a five-
carbon sugar, ribulose 1,5-bisphosphate (RuBP), to yield two molecules of a three-carbon compound,
glycerate 3-phosphate (GP), also known as 3-phosphoglycerate (PGA). GP, in the presence of ATP and
NADPH from the light-dependent stages, is reduced to glyceraldehyde 3-phosphate (G3P). This product is
also referred to as 3-phosphoglyceraldehyde (PGAL) or even as triose phosphate. Triose is a 3-carbon sugar
(see carbohydrates). Most (5 out of 6 molecules) of the G3P produced is used to regenerate RuBP so the
process can continue (see Calvin-Benson cycle). The 1 out of 6 molecules of the triose phosphates not
"recycled" often condense to form hexose phosphates, which ultimately yield sucrose, starch and cellulose.
The sugars produced during carbon metabolism yield carbon skeletons that can be used for other metabolic
reactions like the production of amino acids and lipids.
Carbon concentrating mechanisms
On land
Overview of C4 carbon fixation
In hot and dry conditions, plants close their stomata to prevent the loss of water. Under these
conditions, CO2 will decrease, and oxygen gas, produced by the light reactions of photosynthesis, will
decrease in the stem, not leaves, causing an increase of photorespiration by the oxygenase activity of
ribulose-1,5-bisphosphate carboxylase/oxygenase and decrease in carbon fixation. Some plants have
evolved mechanisms to increase the CO2 concentration in the leaves under these conditions.
Main article: C4 carbon fixation

C4 plants chemically fix carbon dioxide in the cells of the mesophyll by adding it to the three-carbon
molecule phosphoenolpyruvate (PEP), a reaction catalyzed by an enzyme called PEP carboxylase and
which creates the four-carbon organic acid, oxaloacetic acid. Oxaloacetic acid or malate synthesized by this
process is then translocated to specialized bundle sheath cells where the enzyme, rubisco, and other Calvin
cycle enzymes are located, and where CO 2 released by decarboxylation of the four-carbon acids is then
fixed by rubisco activity to the three-carbon sugar 3-phosphoglyceric acids. The physical separation of
rubisco from the oxygen-generating light reactions reduces photorespiration and increases CO 2 fixation and
thus photosynthetic capacity of the leaf.[21] C4 plants can produce more sugar than C3 plants in conditions of
high light and temperature. Many important crop plants are C 4 plants, including maize, sorghum, sugarcane,
and millet. Plants that do not use PEP-carboxylase in carbon fixation are called C3 plants because the
primary carboxylation reaction, catalyzed by rubisco, produces the three-carbon sugar 3-phosphoglyceric
acids directly in the Calvin-Benson cycle. Over 90% of plants use C 3 carbon fixation, compared to 3% that
use C4 carbon fixation.[22]
Main article: CAM photosynthesis
Xerophytes, such as cacti and most succulents, also use PEP carboxylase to capture carbon dioxide
in a process called Crassulacean acid metabolism (CAM). In contrast to C4 metabolism, which physically
separates the CO2 fixation to PEP from the Calvin cycle, CAM temporally separates these two processes.
CAM plants have a different leaf anatomy from C 3 plants, and fix the CO2 at night, when their stomata are
open. CAM plants store the CO2 mostly in the form of malic acid via carboxylation of phosphoenolpyruvate to
oxaloacetate, which is then reduced to malate. Decarboxylation of malate during the day releases CO 2 inside
the leaves, thus allowing carbon fixation to 3-phosphoglycerate by rubisco. 16,000 species of plants use
CAM.[23]
In water
Cyanobacteria possess carboxysomes which increase the concentration of CO2 around rubisco to
increase the rate of photosynthesis. This operates by carbonic anhydrase producing hydrocarbonate ions
(HCO3-) which are then pumped into the carboxysome, before being processed by a different carbonic
anhydrase to produce CO2.[24] Pyrenoids in algae and hornworts also act to concentrate CO2 around
rubisco.[25]
Order and kinetics

The overall process of photosynthesis takes place in four stages. The first, energy transfer in
antenna chlorophyll takes place in the femtosecond (1 femtosecond (fs) = 10 −15 s) to picosecond (1
picosecond (ps) = 10−12 s) time scale. The next phase, the transfer of electrons in photochemical reactions,
takes place in the picosecond to nanosecond time scale (1 nanosecond (ns) = 10 −9 s). The third phase, the
electron transport chain and ATP synthesis, takes place on the microsecond (1 microsecond (μs) = 10 −6 s) to
millisecond (1 millisecond (ms) = 10−3 s) time scale. The final phase is carbon fixation and export of stable
products and takes place in the millisecond to second time scale. The first three stages occur in the thylakoid
membranes.
Efficiency
Main article: Photosynthetic efficiency
Plants usually convert light into chemical energy with a photosynthetic efficiency of 3–6%.[26] Actual
plants' photosynthetic efficiency varies with the frequency of the light being converted, light intensity,
temperature and proportion of carbon dioxide in the atmosphere, and can vary from 0.1% to 8%.[27] By
comparison, solar panels convert light into electric energy at an efficiency of approximately 6–20% for mass-
produced panels, and up to 41% in a research laboratory.[28]
Evolution
Plant cells with visible chloroplasts (from a moss, Plagiomnium affine)

Early photosynthetic systems, such as those from green and purple sulfur and green and purple
nonsulfur bacteria, are thought to have been anoxygenic, using various molecules as electron donors. Green
and purple sulfur bacteria are thought to have used hydrogen and sulfur as an electron donor. Green
nonsulfur bacteria used various amino and other organic acids. Purple nonsulfur bacteria used a variety of
nonspecific organic molecules. The use of these molecules is consistent with the geological evidence that
the atmosphere was highly reduced at that time.[citation needed]
Fossils of what are thought to be filamentous photosynthetic organisms have been dated at 3.4
billion years old.[29][30]
The main source of oxygen in the atmosphere is oxygenic photosynthesis, and its first appearance is
sometimes referred to as the oxygen catastrophe. Geological evidence suggests that oxygenic
photosynthesis, such as that in cyanobacteria, became important during the Paleoproterozoic era around 2
billion years ago. Modern photosynthesis in plants and most photosynthetic prokaryotes is oxygenic.
Oxygenic photosynthesis uses water as an electron donor, which is oxidized to molecular oxygen (O2) in the
photosynthetic reaction center.
Symbiosis and the origin of chloroplasts

Several groups of animals have formed symbiotic relationships with photosynthetic algae. These are
most common in corals, sponges and sea anemones, possibly due to these animals having particularly
simple body plans and large surface areas compared to their volumes.[31] In addition, a few marine mollusks
Elysia viridis and Elysia chlorotica also maintain a symbiotic relationship with chloroplasts they capture from
the algae in their diet and then store in their bodies. This allows the molluscs to survive solely by
photosynthesis for several months at a time.[32][33] Some of the genes from the plant cell nucleus have even
been transferred to the slugs, so that the chloroplasts can be supplied with proteins that they need to survive.
[34]
An even closer form of symbiosis may explain the origin of chloroplasts. Chloroplasts have many
similarities with photosynthetic bacteria, including a circular chromosome, prokaryotic-type ribosomes, and
similar proteins in the photosynthetic reaction center.[35][36] The endosymbiotic theory suggests that
photosynthetic bacteria were acquired (by endocytosis) by early eukaryotic cells to form the first plant cells.
Therefore, chloroplasts may be photosynthetic bacteria that adapted to life inside plant cells. Like
mitochondria, chloroplasts still possess their own DNA, separate from the nuclear DNA of their plant host
cells and the genes in this chloroplast DNA resemble those in cyanobacteria.[37] DNA in chloroplasts codes
for redox proteins such as photosynthetic reaction centers. The CoRR Hypothesis proposes that this Co-
location is required for Redox Regulation.
Cyanobacteria and the evolution of photosynthesis

The biochemical capacity to use water as the source for electrons in photosynthesis evolved once, in
a common ancestor of extant cyanobacteria. The geological record indicates that this transforming event took
place early in Earth's history, at least 2450–2320 million years ago (Ma), and possibly much earlier.[38]
Available evidence from geobiological studies of Archean (>2500 Ma) sedimentary rocks indicates that life
existed 3500 Ma, but the question of when oxygenic photosynthesis evolved is still unanswered. A clear
paleontological window on cyanobacterial evolution opened about 2000 Ma, revealing an already-diverse
biota of blue-greens. Cyanobacteria remained principal primary producers throughout the Proterozoic Eon
(2500–543 Ma), in part because the redox structure of the oceans favored photoautotrophs capable of
nitrogen fixation.[citation needed] Green algae joined blue-greens as major primary producers on continental
shelves near the end of the Proterozoic, but only with the Mesozoic (251–65 Ma) radiations of dinoflagellates,
coccolithophorids, and diatoms did primary production in marine shelf waters take modern form.
Cyanobacteria remain critical to marine ecosystems as primary producers in oceanic gyres, as agents of
biological nitrogen fixation, and, in modified form, as the plastids of marine algae.[39]
A 2010 study by researchers at Tel Aviv University discovered that the Oriental hornet (Vespa
orientalis) converts sunlight into electric power using a pigment called xanthopterin. This is the first scientific
evidence of a member of the animal kingdom engaging in photosynthesis.[40]
Discovery
Although some of the steps in photosynthesis are still not completely understood, the overall
photosynthetic equation has been known since the 19th century.
Jan van Helmont began the research of the process in the mid-17th century when he carefully
measured the mass of the soil used by a plant and the mass of the plant as it grew. After noticing that the soil
mass changed very little, he hypothesized that the mass of the growing plant must come from the water, the
only substance he added to the potted plant. His hypothesis was partially accurate—much of the gained mass
also comes from carbon dioxide as well as water. However, this was a signaling point to the idea that the bulk
of a plant's biomass comes from the inputs of photosynthesis, not the soil itself.
Joseph Priestley, a chemist and minister, discovered that when he isolated a volume of air under an
inverted jar, and burned a candle in it, the candle would burn out very quickly, much before it ran out of wax.
He further discovered that a mouse could similarly "injure" air. He then showed that the air that had been
"injured" by the candle and the mouse could be restored by a plant.
In 1778, Jan Ingenhousz, court physician to the Austrian Empress, repeated Priestley's experiments.
He discovered that it was the influence of sunlight on the plant that could cause it to revive a mouse in a
matter of hours.
In 1796, Jean Senebier, a Swiss pastor, botanist, and naturalist, demonstrated that green plants
consume carbon dioxide and release oxygen under the influence of light. Soon afterwards, Nicolas-Théodore
de Saussure showed that the increase in mass of the plant as it grows could not be due only to uptake of
CO2, but also to the incorporation of water. Thus the basic reaction by which photosynthesis is used to
produce food (such as glucose) was outlined.
Cornelis Van Niel made key discoveries explaining the chemistry of photosynthesis. By studying
purple sulfur bacteria and green bacteria he was the first scientist to demonstrate that photosynthesis is a
light-dependent redox reaction, in which hydrogen reduces carbon dioxide.
Robert Emerson discovered two light reactions by testing plant productivity using different
wavelengths of light. With the red alone, the light reactions were suppressed. When blue and red were
combined, the output was much more substantial. Thus, there were two photosystems, one aborbing up to
600 nm wavelengths, the other up to 700. The former is known as PSII, the latter is PSI. PSI contains only
chlorophyll a, PSII contains primarily chlorophyll a with most of the available chlorophyll b, among other
pigments.[41]
Further experiments to prove that the oxygen developed during the photosynthesis of green plants
came from water, were performed by Robert Hill in 1937 and 1939. He showed that isolated chloroplasts give
off oxygen in the presence of unnatural reducing agents like iron oxalate, ferricyanide or benzoquinone after
exposure to light. The Hill reaction is as follows:
2 H2O + 2 A + (light, chloroplasts) → 2 AH2 + O2
where A is the electron acceptor. Therefore, in light the electron acceptor is reduced and oxygen is
evolved. Cyt b6, now known as a plastoquinone, is one electron acceptor.
Samuel Ruben and Martin Kamen used radioactive isotopes to determine that the oxygen liberated in
photosynthesis came from the water.
Melvin Calvin and Andrew Benson, along with James Bassham, elucidated the path of carbon
assimilation (the photosynthetic carbon reduction cycle) in plants. The carbon reduction cycle is known as the
Calvin cycle, which inappropriately ignores the contribution of Bassham and Benson. Many scientists refer to
the cycle as the Calvin-Benson Cycle, Benson-Calvin, and some even call it the Calvin-Benson-Bassham (or
CBB) Cycle.
A Nobel Prize winning scientist, Rudolph A. Marcus, was able to discover the function and
significance of the electron transport chain.
Otto Heinrich Warburg and Dean Burk discovered the I-quantum photosynthesis reaction that splits
the CO2, activated by the respiration.[42]
Factors
The leaf is the primary site of photosynthesis in plants.

There are three main factors affecting photosynthesis and several corollary factors. The three main
are:
• Light irradiance and wavelength
• Carbon dioxide concentration
• Temperature.
Light intensity (irradiance), wavelength and temperature

In the early 20th century Frederick Frost Blackman along with Albert Einstein investigated the effects
of light intensity (irradiance) and temperature on the rate of carbon assimilation.
• At constant temperature, the rate of carbon assimilation varies with irradiance, initially
increasing as the irradiance increases. However at higher irradiance this relationship no longer holds
and the rate of carbon assimilation reaches a plateau.
• At constant irradiance, the rate of carbon assimilation increases as the temperature is
increased over a limited range. This effect is only seen at high irradiance levels. At low irradiance,
increasing the temperature has little influence on the rate of carbon assimilation.
Carbon assimilation at a constant temperature.
These two experiments illustrate vital points: firstly, from research it is known that photochemical
reactions are not generally affected by temperature. However, these experiments clearly show that
temperature affects the rate of carbon assimilation, so there must be two sets of reactions in the full process
of carbon assimilation. These are of course the light-dependent 'photochemical' stage and the light-
independent, temperature-dependent stage. Second, Blackman's experiments illustrate the concept of
limiting factors. Another limiting factor is the wavelength of light. Cyanobacteria, which reside several meters
underwater, cannot receive the correct wavelengths required to cause photoinduced charge separation in
conventional photosynthetic pigments. To combat this problem, a series of proteins with different pigments
surround the reaction center.This unit is called a phycobilisome.
Carbon dioxide levels and photorespiration
As carbon dioxide concentrations rise, the rate at which sugars are made by the light-independent
reactions increases until limited by other factors. RuBisCO, the enzyme that captures carbon dioxide in the
light-independent reactions, has a binding affinity for both carbon dioxide and oxygen. When the
concentration of carbon dioxide is high, RuBisCO will fix carbon dioxide. However, if the carbon dioxide
concentration is low, RuBisCO will bind oxygen instead of carbon dioxide. This process, called
photorespiration, uses energy, but does not produce sugars.
RuBisCO oxygenase activity is disadvantageous to plants for several reasons:
1. One product of oxygenase activity is phosphoglycolate (2 carbon) instead of 3-
phosphoglycerate (3 carbon). Phosphoglycolate cannot be metabolized by the Calvin-Benson cycle
and represents carbon lost from the cycle. A high oxygenase activity, therefore, drains the sugars
that are required to recycle ribulose 5-bisphosphate and for the continuation of the Calvin-Benson
cycle.
2. Phosphoglycolate is quickly metabolized to glycolate that is toxic to a plant at a high
concentration; it inhibits photosynthesis.
3. Salvaging glycolate is an energetically expensive process that uses the glycolate pathway
and only 75% of the carbon is returned to the Calvin-Benson cycle as 3-phosphoglycerate. The
reactions also produce ammonia (NH3) which is able to diffuse out of the plant leading to a loss of
nitrogen.
A highly simplified summary is:
2 glycolate + ATP → 3-phosphoglycerate + carbon dioxide + ADP +NH 3
The salvaging pathway for the products of RuBisCO oxygenase activity is more commonly known as
photorespiration, since it is characterized by light-dependent oxygen consumption and the release of carbon
dioxide.
See also
Environment portal
Ecology portal
• Jan Anderson (scientist)

• Artificial photosynthesis
• Calvin-Benson cycle
• Carbon fixation
• Cellular respiration
• Chemosynthesis
• Light-dependent reaction
• Photobiology
• Photoinhibition
• Photosynthetic reaction center
• Photosynthetically active radiation
• Quantum biology
• Red edge
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38.^ "Cyanobacteria: Fossil Record". Ucmp.berkeley.edu.
http://www.ucmp.berkeley.edu/bacteria/cyanofr.html. Retrieved 2010-08-26.
39.^ Herrero A and Flores E (editor). (2008). The Cyanobacteria: Molecular Biology, Genomics
and Evolution (1st ed.). Caister Academic Press. ISBN 978-1-904455-15-8.
40.^ Plotkin, M.; Hod, I.; Zaban, A.; Boden, S. A.; Bagnall, D. M.; Galushko, D.; Bergman, D. J.
(2010). "Solar energy harvesting in the epicuticle of the oriental hornet (Vespa orientalis)".
Naturwissenschaften 97 (12): 1067. doi:10.1007/s00114-010-0728-1. PMID 21052618. edit
41.^ ed. and technical staff: Mark D. Licker ... (2007). McGraw Hill Encyclopedia of Science &
Technology (Mcgraw Hill Encyclopedia of Science and Technology), vol. 13 . McGraw-Hill
Professional. p. 470. ISBN 0-07-144143-3.
42.^ http://nobelprize.org/nobel_prizes/medicine/laureates/1931/warburg.html
Further reading
• Asimov, Isaac (1968). Photosynthesis. New York, London: Basic Books, Inc.. ISBN 0-465-
05703-9.
• Bidlack JE; Stern KR, Jansky S (2003). Introductory plant biology. New York: McGraw-Hill.
ISBN 0-07-290941-2.
• Blankenship RE (2008). Molecular Mechanisms of Photosynthesis (2nd ed.). John Wiley &
Sons Inc. ISBN 0-470-71451-4.
• Govindjee (1975). Bioenergetics of photosynthesis. Boston: Academic Press. ISBN 0-12-
294350-3.
• Govindjee Beatty JT,Gest H, Allen JF (2006). Discoveries in Photosynthesis. Advances in
Photosynthesis and Respiration. 20. Berlin: Springer. ISBN 1-4020-3323-0.
• Gregory RL (1971). Biochemistry of photosynthesis. New York: Wiley-Interscience. ISBN 0-
471-32675-5.
• Rabinowitch E, Govindjee (1969). Photosynthesis. London: J. Wiley. ISBN 0-471-70424-5.
• Reece, J, Campbell, N (2005). Biology. San Francisco: Pearson, Benjamin Cummings.
ISBN 0-8053-7146-X.
External links
Wikimedia Commons has media related to: Photosynthesis
• A collection of photosynthesis pages for all levels from a renowned expert (Govindjee)
• In depth, advanced treatment of photosynthesis, also from Govindjee
• Science Aid: Photosynthesis Article appropriate for high school science
• Liverpool John Moores University, Dr.David Wilkinson
• Metabolism, Cellular Respiration and Photosynthesis – The Virtual Library of Biochemistry
and Cell Biology
• Overall examination of Photosynthesis at an intermediate level
• Overall Energetics of Photosynthesis
• Photosynthesis Discovery Milestones – experiments and background
• The source of oxygen produced by photosynthesis Interactive animation, a textbook tutorial

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W000
Ape

For other uses, see Ape (disambiguation).
Apes[1]
Temporal range: Late Oligocene - Recent
Lar Gibbon (Hylobates lar)
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Subclass: Theria
Infraclass: Eutheria
Order: Primates
Suborder: Haplorrhini
Infraorder: Simiiformes
Parvorder: Catarrhini
Hominoidea
Superfamily:
Gray, 1825
Families
Hylobatidae
Hominidae
†Proconsulidae
†Dryopithecidae
†Oreopithecidae
†Pliopithecidae
Under the current classification system there are two families of hominoids:
• the family Hylobatidae consists of 4 genera and 14 species of gibbon, including the Lar
Gibbon and the Siamang, collectively known as the lesser apes.
• the family Hominidae consisting of chimpanzees, gorillas, humans and orangutans[1][2]
collectively known as the great apes.
A few other primates, such as the Barbary Ape, have the word ape in their common names (usually
to indicate lack of a tail), but they are not regarded as true apes.
Except for gorillas and humans, all true apes are agile climbers of trees. Their diet is best described
as omnivorous, their diet consisting of fruit, including grass seeds, and in most cases other animals, either
hunted or scavenged, along with anything else available and easily digested. They are native to Africa and
Asia, although humans have spread to all parts of the world.
Most nonhuman ape species are rare or endangered. The chief threat to most of the endangered
species is loss of tropical rainforest habitat, though some populations are further imperiled by hunting for
bushmeat.
Contents
[hide]
• 1 Historical and modern terminology
• 2 Biology
• 3 History of hominoid taxonomy
• 3.1 Changes in taxonomy
• 4 Classification and evolution
• 5 Behaviour and cognition
• 6 Cultural aspects of non-human apes
• 7 See also
• 8 References
[edit] Historical and modern terminology

"Ape", from Old English apa, is possibly an onomatopoetic imitation of animal chatter. The term has a
history of rather imprecise usage. Its earliest meaning was a tailless (and therefore exceptionally human-like)
non-human primate, but as zoological knowledge developed, it became clear that taillessness occurred in a
number of different and otherwise unrelated species.
The original usage of "ape" in English might have referred to the baboon, an African monkey. Two
tailless species of macaque are commonly named as apes, the Barbary ape of North Africa (introduced into
Gibraltar), Macaca sylvanus, and the Sulawesi black ape or Celebes crested macaque, M. nigra.
Until a few decades ago, humans were thought to be distinctly set apart from the other apes (even
from the other great apes), so much so that most people still do not think of the term "apes" to include
humans at all. However, it is not considered accurate by many biologists to think of apes in a biological sense
without considering humans to be included.[citation needed] The terms "non-human apes" or "non-human
great apes" is used with increasing frequency to show the monophyletic relationship of humans to the other
apes while yet talking only about the non-human species.[ citation needed]
A group of apes may be referred to as a troop or a shrewdness.[3]
[edit] Biology
The gibbon family, Hylobatidae, is composed of fifteen medium-sized species. Their major distinction
is their long arms, which they use to brachiate through the trees. As an evolutionary adaptation to this
arboreal lifestyle, their wrists are ball and socket joints. The largest of the gibbons, the Siamang, weighs up
to 14 kg (31 lb). In comparison, the smallest great ape is the Common Chimpanzee at a modest 40 to 65 kg
(88 to 143 lb).
The great ape family was previously referred to as Pongidae, and humans (and fossil hominids) were
omitted from it, but there is no biological case for doing this. This definition is still used by many
anthropologists and by lay people; however, it makes Pongidae paraphyletic, whereas most taxonomists
nowadays encourage monophyletic groups. Chimpanzees, gorillas, humans and orangutans are all more
closely related to one another than any of these four genera are to the gibbons. However, the term "hominid"
is still used with the specific meaning of extinct animals more closely related to humans than the other great
apes (for example, australopithecines), even though "hominin" is now correct in that usage. It is now usual to
use even finer divisions, such as subfamilies and tribes to distinguish which hominoids are being discussed.
Current evidence implies that humans share a common, extinct, ancestor with the chimpanzee line, from
which we separated more recently than the gorilla line.
Both great apes and lesser apes fall within Catarrhini, which also includes the Old World monkeys of
Africa and Eurasia. Within this group, both families of apes can be distinguished from these monkeys by the
number of cusps on their molars (apes have five—the "Y-5" molar pattern, Old World monkeys have only four
in a bilophodont pattern). Apes have more mobile shoulder joints and arms due to the dorsal position of the
scapula, broad ribcages that are flatter front-to-back, and a shorter, less mobile spine compared to Old World
monkeys (with caudal vertebrae greatly reduced, resulting in tail loss in some species). These are all
anatomical adaptations to vertical hanging and swinging locomotion (brachiation) in the apes, as well as
better balance in a bipedal pose. All living members of the Hylobatidae and Hominidae are tailless, and
humans can therefore accurately be referred to as bipedal apes. However, there are also primates in other
families that lack tails, and at least one (the Pig-Tailed Langur) that has been known to walk significant
distances bipedally. The front skull is characterised by its sinuses, fusion of the frontal bone and post-orbital
constriction.
Although the hominoid fossil record is far from complete, and the evidence is often fragmentary,
there is enough to give a good outline of the evolutionary history of humans. The time of the split between
humans and living apes used to be thought to have occurred 15 to 20 million years ago, or even up to 30 or
40 million years ago. Some apes occurring within that time period, such as Ramapithecus, used to be
considered as hominins, and possible ancestors of humans. Later fossil finds indicated that Ramapithecus
was more closely related to the orangutan, and new biochemical evidence indicated that the last common
ancestor of humans and other hominins occurred between 5 and 10 million years ago, and probably in the
lower end of that range.
[edit] History of hominoid taxonomy
The history of hominoid taxonomy is somewhat confusing and complex. The names of subgroups
have changed their meaning over time as new evidence, from fossil discoveries and comparisons of anatomy
and DNA sequences, has changed understanding of the relationships between hominoids. The story of the
hominoid taxonomy is one of gradual demotion of humans from a special position in the taxonomy to being
one branch among many. It also illustrates the growing influence of cladistics (the science of classifying living
things by strict descent) on taxonomy.
As of 2006, there are eight extant genera of hominoids. They are the four great ape genera ( Homo
(humans), Pan (chimpanzees and bonobos), Gorilla, and Pongo (orangutans)), and the four genera of
gibbons (Hylobates, Hoolock, Nomascus, and Symphalangus).[1] (The genus for the hoolock gibbons was
recently changed from Bunopithecus to Hoolock.[4])
In 1758, Carolus Linnaeus, relying on second- or third-hand accounts, placed a second species in
Homo along with H. sapiens: Homo troglodytes ("cave-dwelling man"). It is not clear to which animal this
name refers, as Linnaeus had no specimen to refer to, hence no precise description. Linnaeus named the
orangutan Simia satyrus ("satyr monkey"). He placed the three genera Homo, Simia and Lemur in the order
of Primates.
The troglodytes name was used for the chimpanzee by Blumenbach in 1775 but moved to the genus
Simia. The orangutan was moved to the genus Pongo in 1799 by Lacépède.
Linnaeus's inclusion of humans in the primates with monkeys and apes was troubling for people who
denied a close relationship between humans and the rest of the animal kingdom. Linnaeus's Lutheran
Archbishop had accused him of "impiety." In a letter to Johann Georg Gmelin dated 25 February 1747,
Linnaeus wrote:
It is not pleasing to me that I must place humans among the primates, but man is intimately
familiar with himself. Let's not quibble over words. It will be the same to me whatever name is applied.
But I desperately seek from you and from the whole world a general difference between men and
simians from the principles of Natural History. I certainly know of none. If only someone might tell me
one! If I called man a simian or vice versa I would bring together all the theologians against me.
Perhaps I ought to, in accordance with the law of Natural History.[5]
Accordingly, Johann Friedrich Blumenbach in the first edition of his Manual of Natural History (1779),
proposed that the primates be divided into the Quadrumana (four-handed, i.e. apes and monkeys) and
Bimana (two-handed, i.e. humans). This distinction was taken up by other naturalists, most notably Georges
Cuvier. Some elevated the distinction to the level of order.
However, the many affinities between humans and other primates — and especially the great apes —
made it clear that the distinction made no scientific sense. Charles Darwin wrote, in The Descent of Man:
The greater number of naturalists who have taken into consideration the whole structure of
man, including his mental faculties, have followed Blumenbach and Cuvier, and have placed man in a
separate Order, under the title of the Bimana, and therefore on an equality with the orders of the
Quadrumana, Carnivora, etc. Recently many of our best naturalists have recurred to the view first
propounded by Linnaeus, so remarkable for his sagacity, and have placed man in the same Order with
the Quadrumana, under the title of the Primates. The justice of this conclusion will be admitted: for in
the first place, we must bear in mind the comparative insignificance for classification of the great
development of the brain in man, and that the strongly marked differences between the skulls of man
and the Quadrumana (lately insisted upon by Bischoff, Aeby, and others) apparently follow from their
differently developed brains. In the second place, we must remember that nearly all the other and
more important differences between man and the Quadrumana are manifestly adaptive in their nature,
and relate chiefly to the erect position of man; such as the structure of his hand, foot, and pelvis, the
curvature of his spine, and the position of his head.[6]
[edit] Changes in taxonomy

Until about 1960, the hominoids were
usually divided into two families: humans and
their extinct relatives in Hominidae, the other
apes in Pongidae.[7]
The 1960s saw the application of

techniques from molecular biology to primate
taxonomy. Goodman used his 1964
immunological study of serum proteins to
propose a division of the hominoids into three
families, with the non-human great apes in
Pongidae and the lesser apes (gibbons) in
Hylobatidae.[8] The trichotomy of hominoid
families, however, prompted scientists to ask
which family speciated first from the common
hominoid ancestor.
Within the superfamily Hominoidea,
gibbons are the outgroup: this means that the
rest of the hominoids are more closely related
to each other than any of them are to gibbons.
This led to the placing of the other great apes
into the family Hominidae along with humans,
by demoting the Pongidae to a subfamily; the
Hominidae family now contained the
subfamilies Homininae and Ponginae. Again,
the three-way split in Ponginae led scientists to
ask which of the three genera is least related to
the others.
Investigation showed orangutans to be

the outgroup, but comparing humans to all
three other hominid genera showed that African
apes (chimpanzees and gorillas) and humans
are more closely related to each other than any
of them are to orangutans. This led to the
placing of the African apes in the subfamily
Homininae, forming another three-way split.
This classification was first proposed by M.
Goodman in 1974.[9]
To try to resolve the hominine
trichotomy, some authors proposed the division
of the subfamily Homininae into the tribes
Gorillini (African apes) and Hominini (humans).
However, DNA comparisons provide

convincing evidence that within the subfamily
Homininae, gorillas are the outgroup. This
suggests that chimpanzees should be in
Hominini along with humans. This classification
was first proposed (though one rank lower) by
M. Goodman et al. in 1990.[2] See Human
evolutionary genetics for more information on
the speciation of humans and great apes.
Later DNA comparisons split the

gibbon genus Hylobates into four genera:
Hylobates, Hoolock, Nomascus, and
Symphalangus.[1][4]
[edit] Classification and evolution

As discussed above, hominoid taxonomy has undergone several changes. Genetic analysis shows
that apes diverged from the Old World monkeys between 29 million and 34.5 million years ago.[10] The
lesser and greater apes split about 18 mya, and the hominid splits happened 14 mya ( Pongo), 7 mya
(Gorilla), and 3-5 mya (Homo & Pan).[citation needed]
Listed are the families and genera of apes; also listed are the extant species.
• Superfamily Hominoidea[1]
• Family Hylobatidae: gibbons
• Genus Hylobates
• Lar Gibbon or White-handed Gibbon, H. lar
• Agile Gibbon or Black-handed Gibbon, H. agilis
• Müller's Bornean Gibbon, H. muelleri
• Silvery Gibbon, H. moloch
• Pileated Gibbon or Capped Gibbon, H. pileatus
• Kloss's Gibbon or Mentawai Gibbon or Bilou, H. klossii
• Genus Hoolock
• Western Hoolock Gibbon, H. hoolock
• Eastern Hoolock Gibbon, H. leuconedys
• Genus Symphalangus
• Siamang, S. syndactylus
• Genus Nomascus
• Black Crested Gibbon, N. concolor
• Eastern Black Crested Gibbon, N. nasutus
• Hainan Gibbon, N. hainanus
• Southern White-cheeked Gibbon N. siki
• White-cheeked Crested Gibbon, N. leucogenys
• Yellow-cheeked Gibbon, N. gabriellae
• Family Hominidae: great apes
• Genus Pongo: orangutans
• Bornean Orangutan, P. pygmaeus
• Sumatran Orangutan, P. abelii
• Genus Gorilla: gorillas
• Western Gorilla, G. gorilla
• Eastern Gorilla, G. beringei
• Genus Homo: humans
• Human, H. sapiens
• Genus Pan: chimpanzees
• Common Chimpanzee, P. troglodytes
• Bonobo, P. paniscus
• Bili Ape, newly discovered, no taxonomy
[edit] Behaviour and cognition

Although there had been earlier studies, the scientific investigation of behaviour and cognition in non-
human apes expanded enormously during the latter half of the twentieth century. Major studies of behaviour
in the field were completed on the three better-known great apes, for example by Jane Goodall, Dian Fossey
and Birute Galdikas (field work on gibbons and the Bonobo is still relatively underdeveloped). These studies
have shown that in their natural environments, the different apes show sharply varying social structure:
gibbons are monogamous, territorial pair-bonders, orangutans are solitary, gorillas live in small troops with a
single adult male leader, while chimpanzees live in larger troops with Bonobos exhibiting promiscuous sexual
behaviour. Their diets also vary; gorillas are foliovores while the others are all primarily frugivores, although
the Common Chimpanzee does some hunting for meat. Foraging behaviour is correspondingly variable.
All the apes are generally thought of as highly intelligent, and scientific study has broadly confirmed
that they perform outstandingly well on a wide range of cognitive tests - though again there is relatively little
data on gibbon cognition. The early studies by Wolfgang Köhler demonstrated exceptional problem-solving
abilities in chimpanzees, which Köhler attributed to insight. The use of tools has been repeatedly
demonstrated; more recently, the manufacture of tools has been documented, both in the wild and in
laboratory tests. Imitation is much more easily demonstrated in great apes than in other primate species.
Almost all the studies in animal language acquisition have been completed with great apes, and though there
is continuing dispute as to whether they demonstrate real language abilities, there is no doubt that they
involve significant feats of learning. Chimpanzees in different parts of Africa have developed tools that are
used in food acquisition, demonstrating a form of animal culture.[11]
[edit] Cultural aspects of non-human apes

Often, non-human apes are said to be the result of a curse—a Jewish folktale claims that one of the
races who built the Tower of Babel became apes as punishment, while Muslim lore says that the Jews of
Eilat became non-human apes as punishment for fishing on the Sabbath. Some sects of Christianity have
folklore that claims that these apes are a symbol of lust and were created by Satan in response to God's
creation of humans.[citation needed] It is uncertain whether any of these references are to any specific apes.
All of these concepts date from a period when neither the distinction between apes and monkeys, nor the fact
that humans are apes, was widely understood, if understood at all.
[edit] See also
Mammals portal
• Chimp Haven
• Declaration on Great Apes from the Great Ape Project
• Human
• List of human evolution fossils
• List of apes (for notable non-fictional apes)
• List of fictional apes
• Planet of the Apes
[edit] References
1. ^ a b c d e Groves, C. (2005). Wilson, D. E., & Reeder, D. M, eds. ed. Mammal Species of the
World (3rd ed.). Baltimore: Johns Hopkins University Press. pp. 178–184. ISBN 0-801-88221-4.
OCLC 62265494. http://www.bucknell.edu/msw3.
2. ^ a b M. Goodman, D. A. Tagle, D. H. Fitch, W. Bailey, J. Czelusniak, B. F. Koop, P. Benson,
J. L. Slightom (1990). "Primate evolution at the DNA level and a classification of hominoids". Journal
of Molecular Evolution 30 (3): 260–266. doi:10.1007/BF02099995. PMID 2109087.
3. ^ "?". http://www.hintsandthings.co.uk/kennel/collectives.htm.
4. ^ a b Mootnick, A.; Groves, C. P. (2005). "A new generic name for the hoolock gibbon
(Hylobatidae)". International Journal of Primatology 26 (26): 971–976. doi:10.1007/s10764-005-5332-
4.
5. ^ "Letter, Carl Linnaeus to Johann Georg Gmelin. Uppsala, Sweden, 25 February 1747".
Swedish Linnaean Society. http://linnaeus.c18.net/Letters/display_txt.php?id_letter=L0783.
6. ^ Charles Darwin (1871). The Descent of Man.
7. ^ G. G. Simpson (1945). "The principles of classification and a classification of mammals".
Bull. Am. Mus. Nat. Hist. 85: 1–350.
8. ^ M. Goodman (1964). "Man’s place in the phylogeny of the primates as reflected in serum
proteins". In S. L. Washburn. Classification and human evolution. Aldine, Chicago. pp. 204–234.
9. ^ M. Goodman (1974). "Biochemical Evidence on Hominid Phylogeny". Annual Review of
Anthropology 3: 203–228. doi:10.1146/annurev.an.03.100174.001223.
10.^ "Apes monkeys split earlier than_fossils had indicated/".
http://www.redorbit.com/news/science/117352/apes_monkeys_split_earlier_than_fossils_had_indicat
ed/.
11.^ William McGrew (1992). Chimpanzee material culture: implications for human evolution .

Find more about Ape on Wikipedia's sister projects:

Wikispecies has information related to: Ape

Hominoidea
• Hominoid systematics: The soft evidence by David Pilbeam Entrez Pubmed 10995486.
Agreement between cladograms based on molecular and anatomical data.
• Dirtpage: Apes




Hominid
Retrieved from "http://en.wikipedia.org/wiki/Ape"
Categories: Apes
W000
Primate

This article is about the type of animal. For other uses, see Primate (disambiguation).
Primates[1]
Temporal range: Late Paleocene–recent
PreЄ
g
Olive Baboon, Papio anubis
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Infraclass: Eutheria
Superorder: Euarchontoglires
Primates
Order:
Linnaeus, 1758
Families
• 16, See classification
Range of the non-human primates (green)

A primate (pronounced /ˈpraɪmeɪt/, US dict: prī′·māt) is a member of the biological order Primates
(/praɪˈmeɪtiːz/ prī·mā′·tēz; Latin: "prime, first rank"[2]), the group that contains prosimians (including lemurs,
lorises, galagos and tarsiers) and simians (monkeys and apes).[3] With the exception of humans, who inhabit
every continent on Earth,[a] most primates live in tropical or subtropical regions of the Americas, Africa and
Asia.[4] Primates range in size from the Madame Berthe's Mouse Lemur, which weighs only 30 grams (1.1
oz) to the Mountain Gorilla weighing 200 kilograms (440 lb). According to fossil evidence, the primitive
ancestors of primates may have existed in the late Cretaceous period around 65 million years ago, and the
oldest known primate is the Late Paleocene Plesiadapis, c. 55–58 million years ago.[5] Molecular clock
studies suggest that the primate branch may be even older, originating in the mid-Cretaceous period around
85 mya.[5]
The Primates order has traditionally been divided into two main groupings: prosimians and simians.
Prosimians have characteristics most like those of the earliest primates, and included the lemurs of
Madagascar, lorisiforms and tarsiers. Simians included the monkeys and apes. More recently, taxonomists
have created the suborder Strepsirrhini, or curly-nosed primates, to include non-tarsier prosimians and the
suborder Haplorrhini, or dry-nosed primates, to include tarsiers and the simians. Simians are divided into two
groups: the platyrrhines ("flat nosed") or New World monkeys of South and Central America and the
catarrhine (narrow nosed) monkeys of Africa and southeastern Asia. The New World monkeys include the
capuchin, howler and squirrel monkeys, and the catarrhines include the Old World monkeys (such as
baboons and macaques) and the apes. Humans are the only catarrhines that have spread successfully
outside of Africa, South Asia, and East Asia, although fossil evidence shows many species once existed in
Europe as well.
Considered generalist mammals, primates exhibit a wide range of characteristics. Some primates
(including some great apes and baboons) do not live primarily in trees, but all species possess adaptations
for climbing trees. Locomotion techniques used include leaping from tree to tree, walking on two or four
limbs, knuckle-walking, and swinging between branches of trees (known as brachiation). Primates are
characterized by their large brains, relative to other mammals, as well as an increased reliance on
stereoscopic vision at the expense of smell, the dominant sensory system in most mammals. These features
are most significant in monkeys and apes, and noticeably less so in lorises and lemurs. Three-color vision
has developed in some primates. Most also have opposable thumbs and some have prehensile tails. Many
species are sexually dimorphic, which means males and females have different physical traits, including
body mass, canine tooth size, and coloration. Primates have slower rates of development than other similarly
sized mammals, and reach maturity later but have longer lifespans. Some species live in solitude, others live
in male–female pairs, and others live in groups of up to hundreds of members.
Contents
[hide]
• 1.1 Evolution
• 1.2 Classification of living primates
• 1.3 Hybrids
• 2 Distinguishing features
• 3 Anatomy, physiology, and morphology
• 3.1 Sexual dimorphism
• 3.2 Locomotion
• 4 Behavior
• 4.1 Social systems
• 4.2 Interspecific associations
• 4.3 Cognition and communication
• 4.4 Life history
• 4.5 Diet and feeding
• 5 Habitat and distribution
• 6 Interactions with humans
• 6.1 Legal and social status
• 6.2 Role in scientific research
• 6.3 Conservation
• 7 See also
Euarchontoglires
Glires
Rodentia (rodents)
Lagomorpha (rabbits, hares,

pikas)
Euarchonta
Scandentia
(treeshrews)
Primatomorpha
Dermoptera
(colugos)
†Plesiadapiforme
s
Primates
The Primates order are a part of the clade Euarchontoglires which is nested within the Eutheria clade
of the class Mammalia. Recent molecular genetic research on primates, colugos, and treeshrews has shown
that the two species of colugos are more closely related to the primates than the treeshrews,[6] even though
the treeshrews were at one time considered primates.[7] These three orders make up the Euarchonta clade.
This clade combines with the Glires clade (composed of the Rodentia and Lagomorpha) to form the
Euarchontoglires clade. Variously, both Euarchonta and Euarchontoglires are ranked as superorders. Some
scientists consider Dermoptera a suborder of Primates and call the "true" primates the suborder Euprimates.
[8]
[edit] Evolution
Further information: List of fossil primates and Lemur evolutionary history
The primate lineage is thought to go back at least 65 mya,[9] even though the oldest known primate
from the fossil record is Plesiadapis (c. 55–58 mya) from the Late Paleocene.[10][11] Other studies, including
molecular clock studies, have estimated the origin of the primate branch to have been in the mid-Cretaceous
period, around 85 mya.[12][13][14]
In modern cladistic reckonings, the Primates order is monophyletic. The suborder Strepsirrhini, the
curly-nosed or "wet-nosed" primates, is generally thought to have split off from the primitive primate line
about 63 mya (million years ago),[15] although earlier dates are also supported.[16] The seven strepsirhine
families are the five related lemur families and the two remaining families that include the lorisids and the
galagos.[1][17] Older classification schemes wrap the Lepilemuridae into the Lemuridae and the Galagidae
into the Lorisidae, yielding a three-two family split instead of the five-two split as presented here.[1] During
the Eocene, most of the northern continents were dominated by two groups, the adapiforms and the
omomyids.[18][19] The former is considered a member of Strepsirrhini, but it does not have a toothcomb like
modern lemurs; recent analysis has suggested Darwinius masillae fits into this grouping.[20] The latter was
related closely to tarsiers, monkeys, and apes. It is unclear exactly how these two groups relate to extant
primates. Omomyids perished about 30 mya,[19] while Adapids survived until about 10 mya.[21]
Ring-tailed Lemur, a strepsirrhine primate
According to genetic studies, the lemurs of Madagascar diverged from the lorisiforms approximately
75 mya.[16] These studies, as well as chromosomal and molecular evidence, also show that lemurs are more
closely related to each other than to other strepsirrhine primates.[16][22] However, Madagascar split from
Africa at 160 mya and from India at 90 mya.[23] For lemurs to be more closely related to each other than
other strepsirrhine primates, it is thought that a very small ancestral population came to Madagascar via a
single rafting event between 50 and 80 million years ago.[16][22][23] Other colonization options have been
examined, such as multiple colonizations from Africa and India, but none are supported by the genetic and
molecular evidence.[18]
Until recently the Aye-aye has been difficult to place within Strepsirrhini.[1] Theories had been
proposed that its family, Daubentoniidae, was either a lemuriform primate (meaning its ancestors split from
lemur line more recently than the lemurs and lorises split) or a sister group to all the other strepsirrhines. In
2008, the Aye-aye family (Daubentoniidae) was confirmed to be mostly closely related to the Malagasy
lemurs, likely having descended from the same ancestral population that colonized the island.[16]
The suborder Haplorrhini, the simple-nosed or "dry-nosed" primates, is composed of two sister
clades.[1] The prosimian tarsiers in family Tarsiidae (monotypic in its own infraorder Tarsiiformes), represent
the most primitive division at about 58 mya.[24][25] The Simiiformes infraorder emerged about 40 mya,[19]
and contains the two clades: the parvorder Platyrrhini that developed in South America and contains New
World monkeys, and the parvorder Catarrhini that developed in Africa and contains the Old World monkeys,
humans and the other apes.[1] A third clade, which included the eosimiids, developed in Asia but went extinct
millions of years ago.[26]
Like the lemurs, the New World monkeys have unclear origins. Molecular sequence studies based on
concatenated sequences have yielded wide variations in the estimated date of the divergence between
platyrrhines and catarrhines, ranging from 33 to 70 mya, while studies based on mitochondrial sequences
suggest a more consistent date of 35 to 43 mya.[5][27] It has been postulated that there is a single origin for
the anthropoids in Africa some migrated and subsequently speciation occurred.[18] It is possible that the
anthropoid primates traversed the Atlantic ocean during the Eocene, facilitated by Atlantic Ocean ridges and
a lowered sea level, then island-hopped to South America.[18] Once again, a rafting event may explain this
transoceanic colonization. Due to continental drift, the young Atlantic Ocean was not nearly as wide as it is
today,[18] and research suggests that a small 1 kg (2.2 lb) primate could have survived 13 days on a raft of
vegetation.[28] Given estimated current and wind speeds, this would have provided enough time to make the
voyage between the continents.
Emperor Tamarin, a New World monkey

Apes and monkeys spread from Africa into Europe and Asia starting in the Miocene.[29] Soon after,
the lorises and tarsiers made the same journey. The first hominid fossils were discovered in Northern Africa
and date back 5–8 mya.[19] Old World monkeys disappeared from Europe about 1.8 mya.[30] Some
molecular and fossil studies generally show that modern humans originated in Africa 100–200 tya (thousand
years ago).[31]
Although primates are well studied in comparison to other animal groups, several new species have
been recently discovered; genetic tests on some populations have revealed previously unrecognised
species. Primate Taxonomy listed about 350 species of primates in 2001,[32] the author, Colin Groves,
increased that number to 376 for his contribution to the third edition of Mammal Species of the World
(MSW3).[1] However, MSW3 falls short of current understanding as its collection was completed in 2003; a
number of publications have since pushed the number to 424 species, or 658 including subspecies.[33]
[edit] Classification of living primates
A 1927 drawing of chimpanzees, a gibbon (top right) and two orangutans (center and bottom center).
The chimp in the upper left is brachiating; the orang at the bottom center is knuckle-walking.
Homo sapiens, a member of the order Primates
The following is the listing of the various families of primates:[1][17][33]
• Order Primates
• Suborder Strepsirrhini: non-tarsier prosimians
• Infraorder Lemuriformes
• Family Cheirogaleidae: dwarf lemurs and mouse-lemurs (32 species)
• Family Daubentoniidae: Aye-aye (1 species)
• Family Lemuridae: lemurs (22 species)
• Family Lepilemuridae: sportive lemurs (26 species)
• Family Indriidae: woolly lemurs and allies (19 species)
• Infraorder Lorisiformes
• Family Lorisidae: lorises, pottos and allies (9 species)
• Family Galagidae: galagos (19 species)
• Suborder Haplorrhini: tarsiers, monkeys and apes
• Infraorder Tarsiiformes
• Family Tarsiidae: tarsiers (9 species)
• Infraorder Simiiformes
• Parvorder Platyrrhini: New World monkeys
• Family Callitrichidae: marmosets and tamarins (42 species)
• Family Cebidae: capuchins and squirrel monkeys (17
species)
• Family Aotidae: night or owl monkeys (douroucoulis) (10
species)
• Family Pitheciidae: titis, sakis and uakaris (42 species)
• Family Atelidae: howler, spider and woolly monkeys (28
species)
• Parvorder Catarrhini
• Superfamily Cercopithecoidea
• Family Cercopithecidae: Old World monkeys (135
species)
• Superfamily Hominoidea
• Family Hylobatidae: gibbons or "lesser apes" (13
species)
• Family Hominidae: great apes, including humans (7
species)
Philippine Tarsier, once considered a prosimian, now predominantly considered a haplorrhine

The order Primates was established by Carl Linnaeus in 1758, in the tenth edition of his book
Systema Naturae,[34] for the genera Homo (humans), Simia (other apes and monkeys), Lemur (prosimians)
and Vespertilio (bats). In the first edition of the same book (1735), he had used the name Anthropomorpha
for Homo, Simia and Bradypus (sloths).[35] In 1839, Henri Marie Ducrotay de Blainville, following Linnaeus
and imitating his nomenclature, established the orders Secundates (including the suborders Chiroptera,
Insectivora and Carnivora), Tertiates (or Glires) and Quaternates (including Gravigrada, Pachydermata and
Ruminantia),[36] but these new taxa were not accepted.
Before Anderson and Jones introduced the classification of Strepsirhini and Haplorhini in 1984,[37]
(followed by McKenna and Bell's 1997 work Classification of Mammals: Above the species level),[38] the
Primates were divided into two superfamilies: Prosimii and Anthropoidea.[39] The Prosimii included all of the
prosimians: all of Strepsirrhini plus the tarsiers. The Anthropoidea contained all of the simians.
[edit] Hybrids
Primate hybrids usually arise in captivity,[40] but there have also been examples in the wild.[41][42]
Hybridization occurs where two species' range overlap to form hybrid zones; hybrids may be created by
humans when animals are placed in zoos or due to environmental pressures such as predation.[41]
Intergeneric hybridizations, hybrids of different genera, have also been found in the wild. Although they
belong to genera that have been distinct for several million years, interbreeding still occurs between the
Gelada and the Hamadryas Baboon.[43]

Primates have diversified in arboreal habitats (trees and bushes) and retain many characteristics that
are adaptations to this environment.[44] They are distinguished by:
• retention of the collar bone in the pectoral girdle;[44]
• shoulder joints which allow high degrees of movement in all directions;[44]
• five digits on the fore and hind limbs with opposable thumbs and big toes;[44]
• nails on the fingers and toes (in most species);[45]
• a flat nail on the hallux (in all extant species);[45]
• sensitive tactile pads on the ends of the digits;[44]
• orbits encircled in bone;[46]
• a trend towards a reduced snout and flattened face, attributed to a reliance on vision at the
expense of olfaction (most notably in haplorrhines, and less so in strepsirrhines);[46]
• a complex visual system with stereoscopic vision, high visual acuity and color vision;[44]
• a brain having a well developed cerebellum with posterior lobe and a Calcarine fissure;[46]
• a large brain in comparison to body size, especially in simians;[44]
• differentiation of an enlarged cerebral cortex;[44]
• reduced number of teeth compared to primitive mammals;[44]
• three kinds of teeth;[46]
• a well-developed cecum;[46]
• two pectoral mammary glands;[44]
• typically one young per pregnancy;[44]
• a pendulous penis and scrotal testes;[46]
• a long gestation and developmental period;[44] and
• a trend towards holding the torso upright leading to bipedalism.[44]
Not all primates exhibit these anatomical traits, nor is every trait unique to primates. For example,
other mammals have collar bones, three kinds of teeth and a pendulous penis, while spider monkeys have
greatly reduced thumbs, ruffed lemurs have six mammary glands and strepsirrhines generally have longer
snouts and a strong sense of smell.[46]
In regard to behavior, primates are frequently highly social, with flexible dominance hierarchies.[47]
New World species form monogamous pair bonds, and show substantial paternal care of their young, unlike
most Old World monkeys.[48]
[edit] Anatomy, physiology, and morphology

Primates have forward-facing eyes on the front of the skull; binocular vision allows accurate distance
perception, useful for the brachiating ancestors of all great apes.[44] There is a bony ridge above the eye
sockets; this ridge reinforces weaker bones in the face which are put under strain during chewing.
Strepsirrhines have a postorbital bar, a bone which runs around the eye socket, to protect their eyes; in
contrast, the higher primates, haplorrhines, have evolved fully enclosed sockets.[49]
Primate crania. The labels indicate mass of brain.
The primate skull has a large domed cranium which is particularly prominent in anthropoids. The
cranium protects the large brain, a distinguishing characteristic of this group.[44] The endocranial volume
(the volume within the skull) is three times greater in humans than in the greatest non-human primate,
reflecting a larger brain size.[50] The mean endocranial volume is 1201 cubic centimeters in humans,
469 cm3 in gorillas, 400 cm3 in chimpanzees and 397 cm3 in orangutans.[50] The primary evolutionary trend
of primates has been the elaboration of the brain, in particular the neocortex (a part of the cerebral cortex),
which is involved with sensory perception, generation of motor commands, spatial reasoning, conscious
thought and, in humans, language.[4] While other mammals rely heavily on their sense of smell, the arboreal
life of primates has led to a tactile, visually dominant sensory system,[4] a reduction in the olfactory region of
the brain and increasingly complex social behavior.[51]
An 1893 drawing of the hands and feet of primates
Primates generally have five digits on each limb (pentadactyly), with keratin nails on the end of each
finger. The bottom sides of the hands and feet have sensitive pads on the fingertips. Most have opposable
thumbs, a characteristic primate feature; however, opposing thumbs are not limited to this order (opossums,
for example, also have them).[44] Thumbs allow some species to use tools. In primates, the combination of
opposing thumbs, short fingernails (rather than claws) and long, inward-closing fingers is a relic of the
ancestral practice of gripping branches, and has, in part, allowed some species to develop brachiation
(swinging by the arms from tree limb to tree limb) as a significant means of transportation. Prosimians have
clawlike nails on the second toe of each foot, called toilet-claws, which they use for grooming.[44]
The primate collar bone is retained as prominent element of the pectoral girdle; this allows the
shoulder joint broad mobility.[47] Apes have more mobile shoulder joints and arms due to the dorsal position
of the scapula, broad ribcages that are flatter front-to-back, and a shorter, less mobile spine compared to Old
World monkeys (with lower vertebrae greatly reduced, resulting in tail loss in some species). Old World
monkeys are unlike apes in that most have tails. The only primate family with prehensile tails are the New
World Atelids, including the howler, spider and woolly monkeys.
Primates show an evolutionary trend towards a reduced snout.[47] Technically, Old World monkeys
are distinguished from New World monkeys by the structure of the nose, and from apes by the arrangement
of their teeth.[51] In New World monkeys the nostrils face sideways; in Old World monkeys, they face
downwards.[51] There is a considerably varied dental pattern in primates and although some have lost most
of their incisors, all retain at least one lower incisor.[51] In most strepsirhines, the lower incisors and canines
form a toothcomb, which is used in grooming and sometimes foraging,[46][51] and the first lower premolar is
shaped like a canine.[46] Old World monkeys have eight premolars, compared with twelve in New World
monkeys.[51] The Old World species are divided into apes and monkeys depending on the number of cusps
on their molars; apes have five, Old World monkeys have four,[51] although humans may have 4 or 5.[52]
The main hominid molar cusp (hypocone) evolved in early primate history, while the cusp of the
corresponding primitive lower molar (paraconid) was lost. Prosimians are distinguished by their immobilized
upper lips, the moist tip of their nose and forward-facing lower front teeth.
The evolution of color vision in primates is unique among most eutherian mammals. While the
remote vertebrate ancestors of the primates possessed three color vision (trichromaticism), the nocturnal,
warm-blooded, mammalian ancestors lost one of three cones in the retina during the Mesozoic period. Fish,
reptiles and birds are therefore trichromatic or tetrachromatic while all mammals, with the exception of some
primates and marsupials,[53] are dichromats or monochromats (totally color blind).[46] Nocturnal primates,
such as the night monkeys and bush babies, are often monochromatic. Catarrhines are routinely trichromatic
due to a gene duplication of the red-green opsin gene at the base of their lineage, 30 to 40 million years ago.
[46][54] Platyrrhines, on the other hand, are trichromatic in a few cases only.[55] Specifically, individual
females must be heterozygous for two alleles of the opsin gene (red and green) located on the same locus of
the X chromosome.[46] Males, therefore, can only be dichromatic, while females can be either dichromatic or
trichromatic. Color vision in strepsirrhines is not as well understood; however, research indicates a range of
color vision similar to that found in platyrrhines.[46]
Like catarrhines, Howler monkeys (a family of platyrrhines) show routine trichromatism that has been
traced to an evolutionarily recent gene duplication.[56] Howler monkeys are one of the most specialized leaf-
eaters of the New World monkeys; fruits are not a major part of their diet,[57] and the type of leaves they
prefer to consume (young, nutritive, and digestible) are detectable only by a red-green signal. Field work
exploring the dietary preferences of howler monkeys suggests that routine trichromaticism was
environmentally selected for.[55]
[edit] Sexual dimorphism

Main article: Sexual dimorphism in non-human primates
Distinct sexual size dimorphism can be seen between the male Hamadryas Baboons (grey) and the
female (brown).
Sexual dimorphism, the variation between individuals of different sex in the same species, is often
exhibited in simians, though to a greater degree in Old World species (apes and some monkeys) than New
World species. Recent studies involve comparing DNA to examine both the variation in the expression of the
dimorphism among primates and the fundamental causes of sexual dimorphism. Primates usually have
dimorphism in body mass[58][59] and canine tooth size[60][61] along with pelage and skin color.[62] The
dimorphism can be attributed to and affected by different factors, including mating system,[63] size,[63]
habitat and diet.[64]
Comparative analyses have generated a more complete understanding of the relationship between
sexual selection, natural selection, and mating systems in primates. Studies have shown that dimorphism is
the product of changes in both male and female traits.[65] Ontogenetic scaling, where relative extension of a
common growth trajectory occurs, may give some insight into the relationship between sexual dimorphism
and growth patterns.[66] Some evidence from the fossil record suggests that there was convergent evolution
of dimorphism, and some extinct hominids probably had greater dimorphism than any living primate.[65]
[edit] Locomotion
Diademed Sifaka, a vertical clinger and leaper, ready to leap to another tree
Primate species move by brachiation, bipedalism, leaping, arboreal and terrestrial quadrupedalism,
climbing, knuckle-walking or by a combination of these methods. Several prosimians are primarily vertical
clinger and leapers. These include many bushbabies, all indriids (i.e., sifakas, avahis and indris), sportive
lemurs, and all tarsiers.[67] Other prosimians are arboreal quadrupeds and climbers. Some are also
terrestrial quadrupeds, while some are leapers. Most monkeys are both arboreal and terrestrial quadrupeds
and climbers. Gibbons, muriquis and spider monkeys all use brachiation extensively.[30] Woolly monkeys
also sometimes brachiate.[57] Orangutans use a similar form of locomotion called quadramanous climbing,
in which they use their arms and legs to carry their heavy bodies through the trees.[30] Chimpanzees and
gorillas knuckle walk,[30] and can move bipedally for short distances. Although numerous species, such as
the Australopithecines and early hominids, have exhibited fully bipedal locomotion, humans are the only
extant species with this trait.
[edit] Behavior
[edit] Social systems
Richard Wrangham stated that social systems of non-human primates are best classified by the
amount of movement by females occurring between groups.[68] He proposed four categories:
• Female transfer systems – females move away from the group in which they were born.
Females of a group will not be closely related whereas males will have remained with their natal
groups, and this close association may be influential in social behavior. The groups formed are
generally quite small. This organization can be seen in chimpanzees, where the males, who are
typically related, will cooperate in defense of the group's territory. Among New World Monkeys,
spider monkeys and muriquis use this system.[69]
Japanese Macaques bathe together in Jigokudani Hot Spring
• Male transfer systems – while the females remain in their natal groups, the males will
emigrate as adolescents. Polygynous and multi-male societies are classed in this category. Group
sizes are usually larger. This system is common among the Ring-tailed Lemur, capuchin monkeys
and cercopithecine monkeys.[30]
• Monogamous species – a male–female bond, sometimes accompanied by a juvenile
offspring. There is shared responsibility of parental care and territorial defense. The offspring leaves
the parents' territory during adolescence. Gibbons essentially use this system, although "monogamy"
in this context does not necessarily mean absolute sexual fidelity.[70]
• Solitary species – often males who defend territories that include the home ranges of several
females. This type of organization is found in the prosimians. Orangutans do not defend their territory
but effectively have this organization.[71]
Other systems are known to occur as well. For example, with howler monkeys both the males and
females typically transfer from their natal group on reaching sexual maturity, resulting in groups in which
neither the males nor females are typically related.[57] Some prosimians, colobine monkeys and callitrichid
monkeys use this system.[30]
Chimpanzees are social animals.

Primatologist Jane Goodall, who studied in the Gombe Stream National Park, noted fission-fusion
societies in chimpanzees.[72] There is fission where the main group splits up to forage during the day, then
fusion when the group returns at night to sleep as a group. This social structure can also be observed in the
Hamadryas Baboon,[73] spider monkeys[57] and the Bonobo.[73] The Gelada has a similar social structure
in which many smaller groups come together to form temporary herds of up to 600 monkeys.[73]
These social systems are affected by three main ecological factors: distribution of resources, group
size and predation.[48] Within a social group there is a balance between cooperation and competition.
Cooperative behaviors include social grooming (removing skin parasites and cleaning wounds), food sharing,
and collective defense against predators or of a territory. Aggressive behaviors often signal competition for
availability of food, sleeping sites or mates. Aggression is also used in establishing dominance hierarchies.
[48][74]
[edit] Interspecific associations

Several species of primates are known to associate in the wild. Some of these associations have
been extensively studied. In the Tai Forest of Africa several species coordinate anti-predator behavior. These
include the Diana Monkey, Campbell's Mona Monkey, Lesser Spot-nosed Monkey, Western Red Colobus,
King Colobus and Sooty Mangabey, which coordinate anti-predator alarm calls.[75] Among the predators of
these monkeys is the Common Chimpanzee.[76]
The Red-tailed Monkey associates with several species, including the Western Red Colobus, Blue
Monkey, Wolf's Mona Monkey, Mantled Guereza, Black Crested Mangabey and Allen's Swamp Monkey.[73]
Several of these species are predated on by the Common Chimpanzee.[77]
In South America, squirrel monkeys associate with capuchin monkeys.[78] This may have more to do
with foraging benefits to the squirrel monkeys rather than anti-predation benefits.[78]
[edit] Cognition and communication

Main article: Primate cognition
Further information: Great ape language
Primates have advanced cognitive abilities: some make tools and use them to acquire food and for
social displays;[79][80] some have sophisticated hunting strategies requiring cooperation, influence and rank;
[81] they are status conscious, manipulative and capable of deception;[82] they can recognise kin and
conspecifics;[83][84] and they can learn to use symbols and understand aspects of human language
including some relational syntax and concepts of number and numerical sequence.[85][86][87] Research in
primate cognition explores problem solving, memory, social interaction, a theory of mind, and numerical,
spatial, and abstract concepts.[88]
Lemurs, lorises, tarsiers, and New World monkeys rely on olfactory signals for many aspects of
social and reproductive behavior.[4] Specialized glands are used to mark territories with pheromones, which
are detected by the vomeronasal organ; this process forms a large part of the communication behavior of
these primates.[4] In Old World monkeys and apes this ability is mostly vestigial, having regressed as
trichromatic eyes evolved to become the main sensory organ.[89] Primates also use vocalizations, gestures,
and facial expressions to convey psychological state.[90]
[edit] Life history

Primates have slower rates of development than other mammals.[30] All non-human primate infants
are breastfed by their mothers and rely on them for grooming and transportation.[30] In some species, infants
are protected and transported by males in the group, particularly males who may be their fathers.[30] Other
relatives of the infant, such as siblings and aunts, may participate in its care as well.[30] Most primate
mothers cease ovulation while breastfeeding an infant; once the infant is weaned the mother can reproduce
again.[30] This often leads to weaning conflict with infants who attempt to continue breastfeeding.[30]
Primates have a longer juvenile period between weaning and sexual maturity than other mammals of
similar size.[30] During the juvenile period, primates are more susceptible than adults to predation and
starvation; they gain experience in feeding and avoiding predators during this time [30] They learn social and
fighting skills, often through playing.[30]
Primates, especially females, have longer lifespans than other similarly sized mammals.[30]
[edit] Diet and feeding
Leaf eating Mantled Guereza, a species of black-and-white colobus

Crab-eating Macaques forage and temporarily store food in their cheeks pouches
Primates exploit a variety of food sources. It has been said that many characteristics of modern
primates, including humans, derive from an early ancestor's practice of taking most of its food from the
tropical canopy.[91] Most primates include fruit in their diets to obtain easily digested carbohydrates and
lipids for energy.[30] However, they require other foods, such as leaves or insects, for amino acids, vitamins
and minerals. Primates in the main suborder Strepsirrhini (non-tarsier prosimians) are able to synthesize
vitamin C, while primates of the suborder of Haplorrhini (tarsiers, monkeys and apes) have lost the ability to
synthesize vitamin C, and require it in the diet.[92]
Many primates have anatomical specializations that enable them to exploit particular foods, such as
fruit, leaves, gum or insects.[30] For example, leaf eaters such as howler monkeys, black-and-white
colobuses and sportive lemurs have extended digestive tracts which enable them to absorb nutrients from
leaves that can be difficult to digest.[30] Marmosets, which are gum eaters, have strong incisor teeth,
enabling them to open tree bark to get to the gum, and claws rather than nails, enabling them to cling to trees
while feeding.[30] The Aye-aye combines rodent-like teeth with a long, thin middle finger to fill the same
ecological niche as a woodpecker. It taps on trees to find insect larvae, then gnaws holes in the wood and
inserts its elongated middle finger to pull the larvae out.[93] Some species have additional specializations.
For example, the Grey-cheeked Mangabey has thick enamel on its teeth, enabling it to open hard fruits and
seeds that other monkeys cannot.[30]
The Gelada is the only primate species that feeds primarily on grass.[94] Tarsiers are the only extant
obligate carnivorous primates, exclusively eating insects, crustaceans, small vertebrates and snakes
(including venomous species).[95] Capuchin monkeys, on the other hand, can exploit many different types of
food, including fruit, leaves, flowers, buds, nectar, seeds, insects and other invertebrates, bird eggs, and
small vertebrates such as birds, lizards, squirrels and bats.[57] The Common Chimpanzee has a varied diet
that includes predation on other primate species, such as the Western Red Colobus monkey.[76][77]
[edit] Habitat and distribution
Rhesus Macaque at Agra Fort, India

Primates evolved from arboreal animals, and many species live most of their lives in trees. Most
primate species live in tropical rain forests. The number of primate species within tropical areas has been
shown to be positively correlated to the amount of rainfall and the amount of rain forest area.[96] Accounting
for 25% to 40% of the fruit-eating animals (by weight) within tropical rainforests, primates play an important
ecological role by dispersing seeds of many tree species.[97]
Some species are partially terrestrial, such as baboons and Patas Monkeys, and a few species are
fully terrestrial, such as Geladas and Humans. Non-human primates live in a diverse number of forested
habitats in the tropical latitudes of Africa, India, Southeast Asia, and South America, including rainforests,
mangrove forests, and montane forests. There are some examples of non-human primates that live outside
of the tropics; the mountain-dwelling Japanese Macaque lives in the north of Honshū where there is snow-
cover eight months of the year; the Barbary Macaque lives in the Atlas Mountains of Algeria and Morocco.
Primate habitats span a range of altitudes: the Black Snub-nosed Monkey has been found living in the
Hengduan Mountains at altitudes of 4,700 meters (15,400 ft),[98] the Mountain Gorilla can be found at 4,200
meters (13,200 ft) crossing the Virunga Mountains,[99] and the Gelada has been found at elevations of up to
5,000 meters (16,400 ft) in the Ethiopian Highlands. Although most species are generally shy of water, a few
are good swimmers and are comfortable in swamps and watery areas, including the Proboscis Monkey, De
Brazza's Monkey and Allen's Swamp Monkey, which has developed small webbing between its fingers.
Some primates, such as the Rhesus Macaque and gray langurs, can exploit human-modified environments
and even live in cities.[73][100]
[edit] Interactions with humans

Close interactions between humans and non-human primates (NHPs) can create pathways for the
transmission of zoonotic diseases. Viruses such as Herpesviridae (most notably Herpes B Virus), Poxviridae,
measles, ebola, rabies, the Marburg virus and viral hepatitis can be transmitted to humans; in some cases
the viruses produce potentially fatal diseases in both humans and non-human primates.[101]
[edit] Legal and social status
Only humans are recognized as persons and protected in law by the United Nations Universal
Declaration of Human Rights.[b] The legal status of NHPs, on the other hand, is the subject of much debate,
with organizations such as the Great Ape Project (GAP) campaigning to award at least some of them legal
rights.[102] In June 2008, Spain became the first country in the world to recognize the rights of some NHPs
when its parliament's cross-party environmental committee urged the country to comply with GAP's
recommendations, which are that chimpanzees, bonobos, orangutans, and gorillas not be used for animal
experiments.[103][104]
Capuchin monkeys' manual dexterity is one reason they can assist quadriplegic humans.
Many species of NHP are kept as pets by humans. GAP estimates that around 3,000 NHPs live as
exotic pets in the United States, while the Humane Society of the United States puts the figure much higher,
at around 15,000.[105] The expanding Chinese middle class has increased demand for NHPs as exotic pets
in recent years.[106] Although NHP import for the pet trade was banned in the U.S. in 1975, smuggling still
occurs along the United States – Mexico border, with prices ranging from US$3000 for monkeys to $30,000
for apes.[107]
Primates are used as model organisms in laboratories and have been used in space missions.[108]
They serve as service animals for disabled humans. Capuchin monkeys can be trained to assist quadriplegic
humans; their intelligence, memory, and manual dexterity make them ideal helpers.[109]
NHPs are kept in zoos around the globe. Historically, zoos were primarily a form of entertainment,
but more recently have shifted their focus to conservation, education and research. Many zoos now feature
naturalistic exhibits and educational material for the public; in the United States many participate in the
Species Survival Plan (SSP), developed by the Association of Zoos and Aquariums (AZA), to maximize
genetic diversity through captive breeding. Zoos and other animal welfare supporters generally oppose
animal rights initiatives and the GAP's insistence that all NHPs be released from captivity for two primary
reasons. First, captive-born primates lack the knowledge and experience to survive in the wild if released.
Second, zoos provide living space for primates and other animals threatened with extinction in the wild.
[edit] Role in scientific research

Further information: Animal testing on non-human primates and International trade in primates
Thousands of non-human primates are used around the world in research because of their
psychological and physiological similarity to humans.[110][111] In particular, the brains and eyes of NHPs
more closely parallel human anatomy than those of any other animals. NHPs are commonly used in
preclinical trials, neuroscience, ophthalmology studies, and toxicity studies. Rhesus Macaques are often
used, as are other Macaques, African green monkeys, chimpanzees, baboons, squirrel monkeys, and
marmosets, both wild-caught and purpose-bred.[110][112] In 2005, GAP reported that 1,280 of the 3,100
NHPs living in captivity in the United States were used for experiments.[102] In 2004, the European Union
used around 10,000 NHPs in such experiments; in 2005 in Great Britain, 4,652 experiments were conducted
on 3,115 NHPs.[113] Governments of many nations have strict care requirements of NHPs kept in captivity.
In the US, federal guidelines extensively regulate aspects of NHP housing, feeding, enrichment, and
breeding.[114] European groups such as the European Coalition to End Animal Experiments are seeking a
ban on all NHP use in experiments as part of the European Union's review of animal testing legislation.[115]
[edit] Conservation
The International Union for Conservation of Nature (IUCN) lists more than a third of primates as
critically endangered or vulnerable. Common threats to primate species include deforestation, forest
fragmentation, monkey drives (resulting from primate crop raiding),[116] and primate hunting for use in
medicines, as pets, and for food. Large-scale tropical forest clearing is widely regarded as the process that
most threatens primates.[117][118][119] More than 90% of primate species occur in tropical forests.[118]
[120] The main cause of forest loss is clearing for agriculture, although commercial logging, subsistence
harvesting of timber, mining, and dam construction contribute to tropical forest depletion too.[120] In
Indonesia large areas of lowland forest have been cleared to increase palm oil production, and one analysis
of satellite imagery concluded that during 1998 and 1999 there was a loss of 1,000 Sumatran Orangutans per
year in the Leuser Ecosystem alone.[121]
Critically endangered Sumatran Orangutan
Primates with a large body size (over 5 kg) have an increased extinction risk due to their increased
profitability to poachers compared to smaller primates.[120] They reach sexual maturity later than other
animals and have a longer period between births. Populations therefore have a slower recovery time after the
loss of members to poaching or the pet trade.[122] Data for some African cities show that half of all protein
consumed in urban areas comes from the bushmeat trade.[123] Endangered primates such as guenons and
the Drill are hunted at levels that far exceed sustainable levels.[123] This is due to their large body size, ease
of transport and profitability per animal.[123] As farming encroaches on forest habitats, primates feed on the
crops, causing the farmers large economic losses.[124] Primate crop raiding gives locals a negative
impression of primates, hindering conservation efforts.[125]
Madagascar, home to five endemic primate families, has experienced the greatest extinction of the
recent past; since human settlement 1,500 years ago, at least eight classes and fifteen species have become
extinct due to hunting and habitat destruction.[4] Among the primates wiped out were Archaeoindris (a lemur
larger than a silverback gorilla) and the families Palaeopropithecidae and Archaeolemuridae.[4]
In Asia, Hinduism, Buddhism, and Islam prohibit eating primate meat; however, primates are still
hunted for food.[120] Some smaller traditional religions allow the consumption of primate meat.[126][127]
The pet trade and traditional medicine also increase demand for illegal hunting.[106][128][129] The Rhesus
Macaque, a model organism, was protected after overtrapping threatened its numbers in the 1960s; the
program was so effective that the macaques are now seen as a pest throughout their range.[119]
The critically endangered Cross River Gorilla

In Central and South America forest fragmentation and hunting are the two main problems for
primates. Large tracts of forest are now rare in Central America.[117][130] This increases the amount of
forest vulnerable to edge effects such as farmland encroachment, lower levels of humidity and a change in
plant life.[131][132] Movement restriction results in a greater amount of inbreeding, which can cause
deleterious effects leading to a population bottleneck, whereby a significant percentage of the population is
lost.[133][134]
There are 21 critically endangered primates, 7 of which have remained on the IUCN's "The World's
25 Most Endangered Primates" list since the year 2000: the Silky Sifaka, Delacour's Langur, the White-
headed Langur, the Gray-shanked Douc, the Tonkin Snub-nosed Langur, the Cross River Gorilla and the
Sumatran Orangutan.[135] Miss Waldron's Red Colobus was recently declared extinct when no trace of the
subspecies could be found from 1993 to 1999.[136] A few hunters have found and killed individuals since
then, and the species' prospects remain bleak.[137]
[edit] See also

Primates portal
• List of primates
• List of fossil primates
• Human evolution
• Arboreal theory
• Primatology
• Simian shelf
[edit] Footnotes
• a Humans inhabit every continent if one includes the scientific and meteorological stations in
Antarctica.
• b Article 6: Everyone has the right to recognition everywhere as a person before the law.[138]

• David J. Chivers, Bernard A. Wood & Alan Bilsborough, ed (1984). Food Acquisition and
Processing in Primates. New York & London: Plenum Press. ISBN 0-306-41701-4
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hominoids and the effective population size of the common ancestor of humans and chimpanzees".
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112.^ Conlee, K. M., Hoffeld, E. H. & Stephens, M. L. (2004). "A Demographic Analysis of
Primate Research in the United States" (pdf). ATLA (Alternatives to Laboratory Animals) 32 (Sup 1):
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113.^ presented to Parliament by the Secretary of State for the Home Department by Command
of Her Majesty, July 2006. (July 2006) (pdf). Statistics of scientific procedures on living animals:
Great Britain 2005. The Stationery Office. ISBN 0101687729. http://www.official-
documents.gov.uk/document/cm68/6877/6877.pdf. Retrieved 2008-06-16.
114.^ "Nonhuman Primates: Research Animals". Animal Welfare Information Center. United
States Department of Agriculture. http://awic.nal.usda.gov/nal_display/index.php?
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Leone, and the Gambia. St. Ives: Trendrine. pp. 214–219. ISBN 0951256246.
117.^ a b Chapman, C. A. & Peres, C. A. (2001). "Primate conservation in the new millennium:
the role of scientists". Evolutionary Anthropology 10: 16–33. doi:10.1002/1520-
6505(2001)10:1<16::AID-EVAN1010>3.0.CO;2-O.
118.^ a b Mittermeier, R. A. & Cheney, D. L. (1987). "Conservation of primates and their
habitats". In Smuts, B. B., Cheney, D. L., Seyfarth, R. M., Wrangham, R. W. & Struhsaker, T. T..
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original on October 12, 2003.
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University of Chicago Press. ISBN 9780226116372.
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orangutan numbers in the Leuser Ecosystem, northern Sumatra". Oryx 35 (1): 14–25.
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risk in declining species". Proceedings of the Royal Society of London 267 (1456): 1947–1952.
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126.^ Choudhury, A. (2001). "Primates in Northeast India: an overview of their distribution and
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Wikimedia Commons has media related to: Primates
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• High-Resolution Cytoarchitectural Primate Brain Atlases
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• PrimateImages: Natural History Collection
• Tree of Life web project
[hide]v · d · eExtant mammal orders by infraclass
Kingdom Animalia · Phylum Chordata · Subphylum Vertebrata · (unranked) Amniota

Australosphenida Monotremata (Platypus and echidnas)
Paucituberculata (Shrew opossums) ·

Ameridelphia
Didelphimorphia (Opossums)
Metatheria
(Marsupial inclusive) Microbiotheria (Monito del Monte) ·
Notoryctemorphia (Marsupial moles) · Dasyuromorphia
Australidelphia
(Quolls and dunnarts) · Peramelemorphia (Bilbies and
bandicoots) · Diprotodontia (Kangaroos and relatives)
Cingulata (Armadillos) · Pilosa (Anteaters and

Xenarthra
sloths)
Afrosoricida (Tenrecs and golden moles) ·

Eutheria Macroscelidea (Elephant shrews) · Tubulidentata
Afrotheria
(Placental inclusive) (Aardvark) · Hyracoidea (Hyraxes) · Proboscidea
(Elephants) · Sirenia (Dugongs and manatees)
Soricomorpha (Shrews and moles) ·

Laurasiatheria Erinaceomorpha (Hedgehogs and relatives) ·
Chiroptera (Bats) · Pholidota (Pangolins) · Carnivora ·
Perissodactyla (Odd-toed ungulates) · Artiodactyla
(Even-toed ungulates) · Cetacea (Whales and
dolphins)
Rodentia (Rodents) · Lagomorpha (Rabbits

Euarchontoglires and relatives) · Scandentia (Treeshrews) ·
Dermoptera (Colugos) · Primates
[hide]v · d · eExtant primate families by suborder
Kingdom Animalia · Phylum Chordata · Class Mammalia · Infraclass Eutheria · Superorder

Euarchontoglires
Strepsirrhini Cheirogaleidae · Lemuridae · Lepilemuridae · Indriidae · Daubentoniidae ·

Lorisidae · Galagidae
Haplorrhini Tarsiidae · Cebidae · Callitrichidae · Aotidae · Pitheciidae · Atelidae ·

Cercopithecidae · Hylobatidae · Hominidae
Retrieved from "http://en.wikipedia.org/wiki/Primate"

Categories: Primates
W000
Biology

For other uses, see Biology (disambiguation).
Biology deals with the study of the many varieties of living organisms. Clockwise from top left:
Salmonella typhimurium, Phascolarctos cinereus, Athyrium filix-femina, Amanita muscaria, Agalychnis
callidryas, and Brachypelma smithi
Biology is a natural science concerned with the study of life and living organisms, including their
structure, function, growth, origin, evolution, distribution, and taxonomy.[1] Biology is a vast subject
containing many subdivisions, topics, and disciplines. Among the most important topics are five unifying
principles that can be said to be the fundamental axioms of modern biology:[2]
1. Cells are the basic unit of life
2. New species and inherited traits are the product of evolution
3. Genes are the basic unit of heredity
4. An organism regulates its internal environment to maintain a stable and constant condition
5. Living organisms consume and transform energy .
Subdisciplines of biology are recognized on the basis of the scale at which organisms are studied
and the methods used to study them: biochemistry examines the rudimentary chemistry of life; molecular
biology studies the complex interactions of systems of biological molecules; cellular biology examines the
basic building block of all life, the cell; physiology examines the physical and chemical functions of the
tissues, organs, and organ systems of an organism; and ecology examines how various organisms interact
and associate with their environment.[3]
Contents
[hide]
• 1 History
• 2 Foundations of modern biology
• 2.1 Cell theory
• 2.2 Evolution
• 2.3 Genetics
• 2.4 Homeostasis
• 2.5 Energy
• 3 Research
• 3.1 Structural
• 3.2 Physiological
• 3.3 Evolutionary
• 3.4 Systematics
• 3.5 Ecology
• 4 Branches of biology
• 5 See also
• 6 Notes and references
History
Main article: History of biology
Ernst Haeckel's Tree of Life (1879)
The term biology in its modern sense appears to have been introduced independently by Karl
Friedrich Burdach (1800), Gottfried Reinhold Treviranus (Biologie oder Philosophie der lebenden Natur,
1802), and Jean-Baptiste Lamarck (Hydrogéologie, 1802).[4][5] It is a classical compound inspired by the
Greek word βίος, bios, "life" and the suffix -λογία, -logia, "study of."
Although biology in its modern form is a relatively recent development, sciences related to and
included within it have been studied since ancient times. Natural philosophy was studied as early as the
ancient civilizations of Mesopotamia, Egypt, the Indian subcontinent, and China. However, the origins of
modern biology and its approach to the study of nature are most often traced back to ancient Greece.[6]
While the formal study of medicine dates back to Hippocrates (ca. 460 BC – ca. 370 BC), it was Aristotle (384
BC – 322 BC) who contributed most extensively to the development of biology. Especially important are his
History of Animals and other works where he showed naturalist leanings, and later more empirical works that
focused on biological causation and the diversity of life. Aristotle's successor at the Lyceum, Theophrastus,
wrote a series of books on botany that survived as the most important contribution of antiquity to the plant
sciences, even into the Middle Ages. Significant advances in the study and development of biology were
promoted through the efforts of such Muslim physicians as the Afro-Arab scholar al-Jahiz (781–869) in
zoology,[7] the Kurdish biologist Al-Dinawari (828–896) in botany,[8] and the Persian physician Rhazes (865–
925) in anatomy and physiology. These philosophers elaborated on, expanded, and improved the Greek
biological theories and systematics. Medicine was especially well studied by Islamic scholars working in
Greek philosopher traditions, while natural history drew heavily on Aristotelian thought, especially in
upholding a fixed hierarchy of life.
Biology began to quickly develop and grow with Antony van Leeuwenhoek's dramatic improvement
of the microscope. It was then that scholars discovered spermatozoa, bacteria, infusoria and the sheer
strangeness and diversity of microscopic life. Investigations by Jan Swammerdam led to new interest in
entomology and built the basic techniques of microscopic dissection and staining.[9]
Advances in microscopy also had a profound impact on biological thinking itself. In the early 19th
century, a number of biologists pointed to the central importance of the cell. In 1838 and 1839, Schleiden and
Schwann began promoting the ideas that (1) the basic unit of organisms is the cell and (2) that individual
cells have all the characteristics of life, although they opposed the idea that (3) all cells come from the
division of other cells. Thanks to the work of Robert Remak and Rudolf Virchow, however, by the 1860s most
biologists accepted all three tenets of what came to be known as cell theory.[10]
Meanwhile, taxonomy and classification became a focus in the study of natural history. Carolus
Linnaeus published a basic taxonomy for the natural world in 1735 (variations of which have been in use ever
since), and in the 1750s introduced scientific names for all his species.[11] Georges-Louis Leclerc, Comte de
Buffon, treated species as artificial categories and living forms as malleable—even suggesting the possibility
of common descent. Though he was opposed to evolution, Buffon is a key figure in the history of evolutionary
thought; his work influenced the evolutionary theories of both Lamarck and Darwin.[12]
Serious evolutionary thinking originated with the works of Jean-Baptiste Lamarck. However, it was
the British naturalist Charles Darwin, combining the biogeographical approach of Humboldt, the
uniformitarian geology of Lyell, Thomas Malthus's writings on population growth, and his own morphological
expertise, that created a more successful evolutionary theory based on natural selection; similar reasoning
and evidence led Alfred Russel Wallace to independently reach the same conclusions.[13]
The discovery of the physical representation of heredity came along with evolutionary principles and
population genetics. In the 1940s and early 1950s, experiments pointed to DNA as the component of
chromosomes that held genes. A focus on new model organisms such as viruses and bacteria, along with
the discovery of the double helical structure of DNA in 1953, marked the transition to the era of molecular
genetics. From the 1950s to present times, biology has been vastly extended in the molecular domain. The
genetic code was cracked by Har Gobind Khorana, Robert W. Holley and Marshall Warren Nirenberg after
DNA was understood to contain codons. Finally, the Human Genome Project was launched in 1990 with the
goal of mapping the general human genome. This project was essentially completed in 2003,[14] with further
analysis still being published. The Human Genome Project was the first step in a globalized effort to
incorporate accumulated knowledge of biology into a functional, molecular definition of the human body and
the bodies of other organisms.
Foundations of modern biology

Much of modern biology can be encompassed within five unifying principles: cell theory, evolution,
genetics, homeostasis, and energy.[2]
Cell theory
Main article: Cell theory
Cells in culture, stained for keratin (red) and DNA (green)
Cell theory states that the cell is the fundamental unit of life, and that all living things are composed
of one or more cells or the secreted products of those cells (e.g. shells). All cells arise from other cells
through cell division. In multicellular organisms, every cell in the organism's body derives ultimately from a
single cell in a fertilized egg. The cell is also considered to be the basic unit in many pathological processes.
[15] Additionally, the phenomenon of energy flow occurs in cells in processes that are part of the function
known as metabolism. Finally, cells contain hereditary information (DNA) which is passed from cell to cell
during cell division.
Evolution
Natural selection of a population for dark coloration.

Main article: Evolution
A central organizing concept in biology is that life changes and develops through evolution, and that
all life-forms known have a common origin. Introduced into the scientific lexicon by Jean-Baptiste de Lamarck
in 1809,[16] evolution was established by Charles Darwin fifty years later as a viable theory when he
articulated its driving force: natural selection.[17][18] (Alfred Russel Wallace is recognized as the co-
discoverer of this concept as he helped research and experiment with the concept of evolution.)[19] Evolution
is now used to explain the great variations of life found on Earth.
Darwin theorized that species and breeds developed through the processes of natural selection and
artificial selection or selective breeding.[20] Genetic drift was embraced as an additional mechanism of
evolutionary development in the modern synthesis of the theory.[21]
The evolutionary history of the species—which describes the characteristics of the various species
from which it descended—together with its genealogical relationship to every other species is known as its
phylogeny. Widely varied approaches to biology generate information about phylogeny. These include the
comparisons of DNA sequences conducted within molecular biology or genomics, and comparisons of fossils
or other records of ancient organisms in paleontology.[22] Biologists organize and analyze evolutionary
relationships through various methods, including phylogenetics, phenetics, and cladistics. (For a summary of
major events in the evolution of life as currently understood by biologists, see evolutionary timeline.)
The theory of evolution postulates that all organisms on the Earth, both living and extinct, have
descended from a common ancestor or an ancestral gene pool. This last universal common ancestor of all
organisms is believed to have appeared about 3.5 billion years ago.[23] Biologists generally regard the
universality and ubiquity of the genetic code as definitive evidence in favor of the theory of universal common
descent for all bacteria, archaea, and eukaryotes (see: origin of life).[24]
Genetics
Main article: Genetics
A Punnett square depicting a cross between two pea plants heterozygous for purple (B) and white (b)
blossoms
Genes are the primary units of inheritance in all organisms. A gene is a unit of heredity and
corresponds to a region of DNA that influences the form or function of an organism in specific ways. All
organisms, from bacteria to animals, share the same basic machinery that copies and translates DNA into
proteins. Cells transcribe a DNA gene into an RNA version of the gene, and a ribosome then translates the
RNA into a protein, a sequence of amino acids. The translation code from RNA codon to amino acid is the
same for most organisms, but slightly different for some. For example, a sequence of DNA that codes for
insulin in humans also codes for insulin when inserted into other organisms, such as plants.[25][26]
DNA usually occurs as linear chromosomes in eukaryotes, and circular chromosomes in prokaryotes.
A chromosome is an organized structure consisting of DNA and histones. The set of chromosomes in a cell
and any other hereditary information found in the mitochondria, chloroplasts, or other locations is collectively
known as its genome. In eukaryotes, genomic DNA is located in the cell nucleus, along with small amounts in
mitochondria and chloroplasts. In prokaryotes, the DNA is held within an irregularly shaped body in the
cytoplasm called the nucleoid.[27] The genetic information in a genome is held within genes, and the
complete assemblage of this information in an organism is called its genotype.[28]
Homeostasis
Main article: Homeostasis
The hypothalamus secretes CRH, which directs the pituitary gland to secrete ACTH. In turn, ACTH
directs the adrenal cortex to secrete glucocorticoids, such as cortisol. The GCs then reduce the rate of
secretion by the hypothalamus and the pituitary gland once a sufficient amount of GCs has been released.
[29]
Homeostasis is the ability of an open system to regulate its internal environment to maintain stable
conditions by means of multiple dynamic equilibrium adjustments controlled by interrelated regulation
mechanisms. All living organisms, whether unicellular or multicellular, exhibit homeostasis.[30]
To maintain dynamic equilibrium and effectively carry out certain functions, a system must detect and
respond to perturbations. After the detection of a perturbation, a biological system normally respond through
negative feedback. This means stabilizing conditions by either reducing or increasing the activity of an organ
or system. One example is the release of glucagon when sugar levels are too low.
Energy
Basic overview of energy and human life.

The survival of a living organism depends on the continuous input of energy. Chemical reactions that
are responsible for its structure and function are tuned to extract energy from substances that act as its food
and transform them to help form new cells and sustain them. In this process, molecules of chemical
substances that constitute food play two roles; first, they contain energy that can be transformed for
biological chemical reactions; second, they develop new molecular structures made up of biomolecules.
The organisms responsible for the introduction of energy into an ecosystem are known as producers
or autotrophs. Nearly all of these organisms originally draw energy from the sun.[31] Plants and other
phototrophs use solar energy via a process known as photosynthesis to convert raw materials into organic
molecules, such as ATP, whose bonds can be broken to release energy.[32] A few ecosystems, however,
depend entirely on energy extracted by chemotrophs from methane, sulfides, or other non-luminal energy
sources.[33]
Some of the captured energy is used to produce biomass to sustain life and provide energy for
growth and development. The majority of the rest of this energy is lost as heat and waste molecules. The
most important processes for converting the energy trapped in chemical substances into energy useful to
sustain life are metabolism[34] and cellular respiration.[35]
Research
Structural
Schematic of typical animal cell depicting the various organelles and structures.
Main articles: Molecular biology, Cell biology, Genetics, and Developmental biology
Molecular biology is the study of biology at a molecular level.[36] This field overlaps with other areas
of biology, particularly with genetics and biochemistry. Molecular biology chiefly concerns itself with
understanding the interactions between the various systems of a cell, including the interrelationship of DNA,
RNA, and protein synthesis and learning how these interactions are regulated.
Cell biology studies the structural and physiological properties of cells, including their behaviors,
interactions, and environment. This is done on both the microscopic and molecular levels, for single-celled
organisms such as bacteria as well as the specialized cells in multicellular organisms such as humans.
Understanding the structure and function of cells is fundamental to all of the biological sciences. The
similarities and differences between cell types are particularly relevant to molecular biology.
Anatomy considers the forms of macroscopic structures such as organs and organ systems.[37]
Genetics is the science of genes, heredity, and the variation of organisms.[38][39] Genes encode the
information necessary for synthesizing proteins, which in turn play a large role in influencing (though, in many
instances, not completely determining) the final phenotype of the organism. In modern research, genetics
provides important tools in the investigation of the function of a particular gene, or the analysis of genetic
interactions. Within organisms, genetic information generally is carried in chromosomes, where it is
represented in the chemical structure of particular DNA molecules.
Developmental biology studies the process by which organisms grow and develop. Originating in
embryology, modern developmental biology studies the genetic control of cell growth, differentiation, and
"morphogenesis," which is the process that progressively gives rise to tissues, organs, and anatomy. Model
organisms for developmental biology include the round worm Caenorhabditis elegans,[40] the fruit fly
Drosophila melanogaster,[41] the zebrafish Danio rerio,[42] the mouse Mus musculus,[43], and the weed
Arabidopsis thaliana.[44][45] (A model organism is a species that is extensively studied to understand
particular biological phenomena, with the expectation that discoveries made in that organism provide insight
into the workings of other organisms.)[46]
Physiological
Main article: Physiology
Physiology studies the mechanical, physical, and biochemical processes of living organisms by
attempting to understand how all of the structures function as a whole. The theme of "structure to function" is
central to biology. Physiological studies have traditionally been divided into plant physiology and animal
physiology, but some principles of physiology are universal, no matter what particular organism is being
studied. For example, what is learned about the physiology of yeast cells can also apply to human cells. The
field of animal physiology extends the tools and methods of human physiology to non-human species. Plant
physiology borrows techniques from both research fields.
Physiology studies how for example nervous, immune, endocrine, respiratory, and circulatory
systems, function and interact. The study of these systems is shared with medically oriented disciplines such
as neurology and immunology.
Evolutionary
Evolutionary research is concerned with the origin and descent of species, as well as their change
over time, and includes scientists from many taxonomically oriented disciplines. For example, it generally
involves scientists who have special training in particular organisms such as mammalogy, ornithology,
botany, or herpetology, but use those organisms as systems to answer general questions about evolution.
Evolutionary biology is partly based on paleontology, which uses the fossil record to answer
questions about the mode and tempo of evolution,[47] and partly on the developments in areas such as
population genetics[48] and evolutionary theory. In the 1980s, developmental biology re-entered evolutionary
biology from its initial exclusion from the modern synthesis through the study of evolutionary developmental
biology.[49] Related fields often considered part of evolutionary biology are phylogenetics, systematics, and
taxonomy.
Systematics
A phylogenetic tree of all living things, based on rRNA gene data, showing the separation of the three
domains bacteria, archaea, and eukaryotes as described initially by Carl Woese. Trees constructed with
other genes are generally similar, although they may place some early-branching groups very differently,
presumably owing to rapid rRNA evolution. The exact relationships of the three domains are still being
debated.
Main article: Systematics
Multiple speciation events create a tree structured system of relationships between species. The role
of systematics is to study these relationships and thus the differences and similarities between species and
groups of species.[50] However, systematics was an active field of research long before evolutionary thinking
was common.[51] The classification, taxonomy, and nomenclature of biological organisms is administered by
the International Code of Zoological Nomenclature, International Code of Botanical Nomenclature, and
International Code of Nomenclature of Bacteria for animals, plants, and bacteria, respectively. The
classification of viruses, viroids, prions, and all other sub-viral agents that demonstrate biological
characteristics is conducted by the International Code of Virus classification and nomenclature.[52][53][54]
[55] However, several other viral classification systems do exist.
Traditionally, living things have been divided into five kingdoms: Monera; Protista; Fungi; Plantae;
Animalia.[56]
However, many scientists now consider this five-kingdom system outdated. Modern alternative
classification systems generally begin with the three-domain system: Archaea (originally Archaebacteria);
Bacteria (originally Eubacteria); Eukaryota (including protists, fungi, plants, and animals)[57] These domains
reflect whether the cells have nuclei or not, as well as differences in the chemical composition of the cell
exteriors.[57]
Further, each kingdom is broken down recursively until each species is separately classified. The
order is: Domain; Kingdom; Phylum; Class; Order; Family; Genus; Species.
There is also a series of intracellular parasites that are "on the edge of life"[58] in terms of metabolic
activity, meaning that many scientists do not actually classify these structures as alive, due to their lack of at
least one or more of the fundamental functions that define life. They are classified as viruses, viroids, prions,
or satellites.
The scientific name of an organism is generated from its genus and species. For example, humans
are listed as Homo sapiens. Homo is the genus, and sapiens the species. When writing the scientific name of
an organism, it is proper to capitalize the first letter in the genus and put all of the species in lowercase.
Additionally, the entire term may be italicized or underlined.[59][60]
The dominant classification system is called the Linnaean taxonomy. It includes ranks and binomial
nomenclature. How organisms are named is governed by international agreements such as the International
Code of Botanical Nomenclature (ICBN), the International Code of Zoological Nomenclature (ICZN), and the
International Code of Nomenclature of Bacteria (ICNB).
A merging draft, BioCode, was published in 1997 in an attempt to standardize nomenclature in these
three areas, but has yet to be formally adopted.[61] The BioCode draft has received little attention since
1997; its originally planned implementation date of January 1, 2000, has passed unnoticed. However, a 2004
paper concerning the cyanobacteria does advocate a future adoption of a BioCode and interim steps
consisting of reducing the differences between the codes.[62] The International Code of Virus Classification
and Nomenclature (ICVCN) remains outside the BioCode.
Ecology
Mutual symbiosis between clownfish of the genus Amphiprion that dwell among the tentacles of
tropical sea anemones. The territorial fish protects the anemone from anemone-eating fish, and in turn the
stinging tentacles of the anemone protects the clown fish from its predators
Main articles: Ecology, Ethology, Behavior, and Biogeography
Ecology studies the distribution and abundance of living organisms, and the interactions between
organisms and their environment.[63] The habitat of an organism can be described as the local abiotic
factors such as climate and ecology, in addition to the other organisms and biotic factors that share its
environment.[64] One reason that biological systems can be difficult to study is that so many different
interactions with other organisms and the environment are possible, even on the smallest of scales. A
microscopic bacterium responding to a local sugar gradient is responding to its environment as much as a
lion is responding to its environment when it searches for food in the African savanna. For any given species,
behaviors can be co-operative, aggressive, parasitic, or symbiotic. Matters become more complex when two
or more different species interact in an ecosystem. Studies of this type are within the province of ecology.
Ecological systems are studied at several different levels, from individuals and populations to
ecosystems and the biosphere. The term population biology is often used interchangeably with population
ecology, although population biology is more frequently used when studying diseases, viruses, and
microbes, while population ecology is more commonly when studying plants and animals. As can be
surmised, ecology is a science that draws on several disciplines.
Ethology studies animal behavior (particularly that of social animals such as primates and canids),
and is sometimes considered a branch of zoology. Ethologists have been particularly concerned with the
evolution of behavior and the understanding of behavior in terms of the theory of natural selection. In one
sense, the first modern ethologist was Charles Darwin, whose book, The Expression of the Emotions in Man
and Animals, influenced many ethologists to come.[65]
Biogeography studies the spatial distribution of organisms on the Earth,[66] focusing on topics like
plate tectonics, climate change, dispersal and migration, and cladistics.
Branches of biology
Main article: List of biology disciplines
These are the main branches of biology:[67][68]
• Aerobiology — the study of airborne organic particles
• Agriculture — the study of producing crops from the land, with an emphasis on practical
applications
• Anatomy — the study of form and function, in plants, animals, and other organisms, or
specifically in humans
• Astrobiology- the study of evolution, distribution, and future of life in the universe—also known
as exobiology, exopaleontology, and bioastronomy
• Biochemistry — the study of the chemical reactions required for life to exist and function,
usually a focus on the cellular level
• Bioengineering — the study of biology through the means of engineering with an emphasis on
applied knowledge and especially related to biotechnology
• Bioinformatics — the use of information technology for the study, collection, and storage of
genomic and other biological data
• Biomathematics or Mathematical Biology — the quantitative or mathematical study of
biological processes, with an emphasis on modeling
• Biomechanics — often considered a branch of medicine, the study of the mechanics of living
beings, with an emphasis on applied use through prosthetics or orthotics
• Biomedical research — the study of the human body in health and disease
• Biophysics — the study of biological processes through physics, by applying the theories and
methods traditionally used in the physical sciences
• Biotechnology — a new and sometimes controversial branch of biology that studies the
manipulation of living matter, including genetic modification and synthetic biology
• Building biology — the study of the indoor living environment
• Botany — the study of plants
• Cell biology — the study of the cell as a complete unit, and the molecular and chemical
interactions that occur within a living cell
• Conservation Biology — the study of the preservation, protection, or restoration of the natural
environment, natural ecosystems, vegetation, and wildlife
• Cryobiology — the study of the effects of lower than normally preferred temperatures on living
beings.
• Developmental biology — the study of the processes through which an organism forms, from
zygote to full structure
• Ecology — the study of the interactions of living organisms with one another and with the non-
living elements of their environment
• Embryology — the study of the development of embryo (from fecundation to birth). See also
topobiology.
• Entomology — the study of insects
• Environmental Biology — the study of the natural world, as a whole or in a particular area,
especially as affected by human activity
• Epidemiology — a major component of public health research, studying factors affecting the
health of populations
• Ethology — the study of animal behavior
• Evolutionary Biology — the study of the origin and descent of species over time
• Genetics — the study of genes and heredity
• Herpetology — the study of reptiles and amphibians
• Histology — the study of cells and tissues, a microscopic branch of anatomy
• Ichthyology — the study of fish
• Integrative biology — the study of whole organisms
• Limnology — the study of inland waters
• Mammalogy — the study of mammals
• Marine Biology — the study of ocean ecosystems, plants, animals, and other living beings
• Microbiology — the study of microscopic organisms (microorganisms) and their interactions
with other living things
• Molecular Biology — the study of biology and biological functions at the molecular level, some
cross over with biochemistry
• Mycology — the study of fungi
• Neurobiology — the study of the nervous system, including anatomy, physiology and
pathology
• Oceanography — the study of the ocean, including ocean life, environment, geography,
weather, and other aspects influencing the ocean
• Oncology — the study of cancer processes, including virus or mutation oncogenesis,
angiogenesis and tissues remoldings
• Ornithology — the study of birds
• Population biology — the study of groups of conspecific organisms, including
• Population ecology — the study of how population dynamics and extinction
• Population genetics — the study of changes in gene frequencies in populations of
organisms
• Paleontology — the study of fossils and sometimes geographic evidence of prehistoric life
• Pathobiology or pathology — the study of diseases, and the causes, processes, nature, and
development of disease
• Parasitology — the study of parasites and parasitism
• Pharmacology — the study and practical application of preparation, use, and effects of drugs
and synthetic medicines
• Physiology — the study of the functioning of living organisms and the organs and parts of
living organisms
• Phytopathology — the study of plant diseases (also called Plant Pathology)
• Psychobiology — the study of the biological bases of psychology
• Sociobiology — the study of the biological bases of sociology
• Structural biology — a branch of molecular biology, biochemistry, and biophysics concerned
with the molecular structure of biological macromolecules
• Virology — the study of viruses and some other virus-like agents
• Zoology — the study of animals, including classification, physiology, development, and
behavior (See also Entomology, Ethology, Herpetology, Ichthyology, Mammalogy, and Ornithology)
See also
Biology portal
Environment portal
Ecology portal
Earth_sciences portal
Main article: Outline of biology
• Conservation biology
• The Journal of Life Sciences
• List of biological websites
• List of biologists
• List of Russian biologists
• List of biology topics
• Lists of biology journals and magazines
• Periodic Table of Life Sciences in Tinbergen's four questions
[show]v · d · eTopics related to biology
People and Biologist · Notable biologists · History of biology · Nobel Prize in

history Physiology or Medicine · Timeline of biology and organic chemistry · List of
geneticists and biochemists
Institutions,
publications Bachelor of Science · Publications
Terms and Omne vivum ex ovo · In vivo · In vitro · In utero · In silico

phrases
Related Medicine (Physician) · Physical anthropology · Environmental science ·

disciplines Life Sciences · Biotechnology
Other List of conservation topics · Altricial and Precocial development strategies
Notes and references

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56.^ Margulis, L; Schwartz, KV (1997). Five Kingdoms: An Illustrated Guide to the Phyla of Life
on Earth (3rd ed.). WH Freeman & Co. ISBN 978-0716731832. OCLC 237138975 223623098
237138975.
57.^ a b Woese C, Kandler O, Wheelis M (1990). "Towards a natural system of organisms:
proposal for the domains Archaea, Bacteria, and Eukarya". Proc Natl Acad Sci USA 87 (12): 4576–9.
doi:10.1073/pnas.87.12.4576. PMID 2112744. PMC 54159.
58.^ Rybicki EP (1990) "The classification of organisms at the edge of life, or problems with
virus systematics." S Aft J Sci 86:182–186
59.^ Heather Silyn-Roberts (2000). Writing for Science and Engineering: Papers, Presentation .
Oxford: Butterworth-Heinemann. pp. 198. ISBN 0750646365. http://books.google.com/?
id=hVUU7Gq8QskC&lpg=PA198&dq=species%20epithet
%20capitalize&pg=PA198#v=onepage&q=species%20epithet%20capitalize.
60.^ "Recommendation 60F". International Code of Botanical Nomenclature, Vienna Code.
2006. pp. 60F.1. http://ibot.sav.sk/icbn/frameset/0065Ch7OaGoNSec1a60.htm#recF.
61.^ John McNeill (1996-11-04). "The BioCode: Integrated biological nomenclature for the 21st
century?". Proceedings of a Mini-Symposium on Biological Nomenclature in the 21st Century .
62.^ Ahoren Oren (2004). "A proposal for further integration of the cyanobacteria under the
Bacteriological Code". Int. J. Syst. Evol. Microbiol. 54 (Pt 5): 1895–1902. doi:10.1099/ijs.0.03008-0.
PMID 15388760.
63.^ Begon, M.; Townsend, C. R., Harper, J. L. (2006). Ecology: From individuals to
ecosystems. (4th ed.). Blackwell. ISBN 1405111178.
64.^ Habitats of the world. New York: Marshall Cavendish. 2004. pp. 238. ISBN 978-0-7614-
7523-1. http://books.google.com/?id=U-_mlcy8rGgC&pg=PA238.
65.^ Black, J (Jun 2002). "Darwin in the world of emotions" (Free full text). Journal of the Royal
Society of Medicine 95 (6): 311–3. doi:10.1258/jrsm.95.6.311. ISSN 0141-0768. PMID 12042386.
PMC 1279921. http://www.jrsm.org/cgi/pmidlookup?view=long&pmid=12042386.
66.^ Wiley, 1981
67.^ Branches of Biology on biology-online.org
68.^ Biology on bellaonline.com
Further reading
• Alberts, Bruce; Johnson, A, Lewis, J, Raff, M, Roberts, K & Walter, P (2002). Molecular
Biology of the Cell (4th ed.). Garland. ISBN 978-0815332183. OCLC 48122761 57023651 69932405
145080076 48122761 57023651 69932405.
• Begon, Michael; Townsend, CR & Harper, JL (2005). Ecology: From Individuals to
Ecosystems (4th ed.). Blackwell Publishing Limited. ISBN 978-1405111171. OCLC 57675855
62131207 57639896 57675855 62131207.
• Campbell, Neil (2004). Biology (7th ed.). Benjamin-Cummings Publishing Company. ISBN 0-
8053-7146-X. OCLC 71890442.
• Colinvaux, Paul (1979). Why Big Fierce Animals are Rare: An Ecologist's Perspective
(reissue ed.). Princeton University Press. ISBN 0691023646. OCLC 24132192 10081738 24132192.
• Hoagland, Mahlon (2001). The Way Life Works (reprint ed.). Jones and Bartlett Publishers
inc. ISBN 076371688X. OCLC 45487537 223090105 45487537.
• Janovy, John Jr. (2004). On Becoming a Biologist (2nd ed.). Bison Books.
ISBN 0803276206. OCLC 56964280 55138571 56964280.
• Johnson, George B. (2005). Biology, Visualizing Life. Holt, Rinehart, and Winston. ISBN 0-
03-016723-X. OCLC 36306648.
• Tobin, Allan; Dusheck, Jennie (2005). Asking About Life (3rd ed.). Belmont, CA: Wadsworth.
ISBN 0-534-40653X.
External links
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Cell (biology)

Drawing of the structure of cork as it appeared under the microscope to Robert Hooke from
Micrographia which is the origin of the word "cell" being used to describe the smallest unit of a living
organism
Cells in culture, stained for keratin (red) and DNA (green)
The cell is the functional basic unit of life. It was discovered by Robert Hooke and is the functional
unit of all known living organisms. It is the smallest unit of life that is classified as a living thing, and is often
called the building block of life.[1] Some organisms, such as most bacteria, are unicellular (consist of a single
cell). Other organisms, such as humans, are multicellular. Humans have about 100 trillion or 1014 cells; a
typical cell size is 10 µm and a typical cell mass is 1 nanogram. The largest cells are about 135 µm in the
anterior horn in the spinal cord while granule cells in the cerebellum, the smallest, can be some 4 µm and the
longest cell can reach from the toe to the lower brain stem (Pseudounipolar cells).[2] The largest known cells
are unfertilised ostrich egg cells which weigh 3.3 pounds.[3][4]
In 1835, before the final cell theory was developed, Jan Evangelista Purkyně observed small
"granules" while looking at the plant tissue through a microscope. The cell theory, first developed in 1839 by
Matthias Jakob Schleiden and Theodor Schwann, states that all organisms are composed of one or more
cells, that all cells come from preexisting cells, that vital functions of an organism occur within cells, and that
all cells contain the hereditary information necessary for regulating cell functions and for transmitting
information to the next generation of cells.[5]
The word cell comes from the Latin cellula, meaning, a small room. The descriptive term for the
smallest living biological structure was coined by Robert Hooke in a book he published in 1665 when he
compared the cork cells he saw through his microscope to the small rooms monks lived in.[6]
Contents
[hide]
• 1 Anatomy
• 1.1 Prokaryotic cells
• 1.2 Eukaryotic cells
• 2 Subcellular components
• 2.1 Membrane
• 2.2 Cytoskeleton
• 2.3 Genetic material
• 2.4 Organelles
• 3 Structures outside the cell wall
• 3.1 Capsule
• 3.2 Flagella
• 3.3 Fimbriae (pili)
• 4 Functions
• 4.1 Growth and metabolism
• 4.2 Creation
• 4.3 Protein synthesis
• 5 Movement or motility
• 6 Evolution
• 6.1 Origin of the first cell
• 6.2 Origin of eukaryotic cells
Anatomy
There are two types of cells: eukaryotic and prokaryotic. Prokaryotic cells are usually independent,
while eukaryotic cells are often found in multicellular organisms.
Prokaryotic cells
Main article: Prokaryote
Diagram of a typical prokaryotic cell
The prokaryote cell is simpler, and therefore smaller, than a eukaryote cell, lacking a nucleus and
most of the other organelles of eukaryotes. There are two kinds of prokaryotes: bacteria and archaea; these
share a similar structure.
Nuclear material of prokaryotic cell consist of a single chromosome which is in direct contact with
cytoplasm. Here the undefined nuclear region in the cytoplasm is called nucleoid.
A prokaryotic cell has three architectural regions:
• On the outside, flagella and pili project from the cell's surface. These are structures (not
present in all prokaryotes) made of proteins that facilitate movement and communication between
cells;
• Enclosing the cell is the cell envelope – generally consisting of a cell wall covering a plasma
membrane though some bacteria also have a further covering layer called a capsule. The envelope
gives rigidity to the cell and separates the interior of the cell from its environment, serving as a
protective filter. Though most prokaryotes have a cell wall, there are exceptions such as Mycoplasma
(bacteria) and Thermoplasma (archaea). The cell wall consists of peptidoglycan in bacteria, and acts
as an additional barrier against exterior forces. It also prevents the cell from expanding and finally
bursting (cytolysis) from osmotic pressure against a hypotonic environment. Some eukaryote cells
(plant cells and fungi cells) also have a cell wall;
• Inside the cell is the cytoplasmic region that contains the cell genome (DNA) and ribosomes
and various sorts of inclusions. A prokaryotic chromosome is usually a circular molecule (an
exception is that of the bacterium Borrelia burgdorferi, which causes Lyme disease). Though not
forming a nucleus, the DNA is condensed in a nucleoid. Prokaryotes can carry extrachromosomal
DNA elements called plasmids, which are usually circular. Plasmids enable additional functions, such
as antibiotic resistance.
Eukaryotic cells
Main article: Eukaryote
Diagram of a typical animal (eukaryotic) cell, showing subcellular components.

Organelles:
(1) nucleolus
(2) nucleus
(3) ribosome
(4) vesicle
(5) rough endoplasmic reticulum (ER)
(6) Golgi apparatus
(7) Cytoskeleton
(8) smooth endoplasmic reticulum
(9) mitochondria
(10) vacuole
(11) cytoplasm
(12) lysosome
(13) centrioles within centrosome
Eukaryotic cells are about 15 times wider than a typical prokaryote and can be as much as 1000
times greater in volume. The major difference between prokaryotes and eukaryotes is that eukaryotic cells
contain membrane-bound compartments in which specific metabolic activities take place. Most important
among these is a cell nucleus, a membrane-delineated compartment that houses the eukaryotic cell's DNA.
This nucleus gives the eukaryote its name, which means "true nucleus." Other differences include:
• The plasma membrane resembles that of prokaryotes in function, with minor differences in
the setup. Cell walls may or may not be present.
• The eukaryotic DNA is organized in one or more linear molecules, called chromosomes,
which are associated with histone proteins. All chromosomal DNA is stored in the cell nucleus,
separated from the cytoplasm by a membrane. Some eukaryotic organelles such as mitochondria
also contain some DNA.
• Many eukaryotic cells are ciliated with primary cilia. Primary cilia play important roles in
chemosensation, mechanosensation, and thermosensation. Cilia may thus be "viewed as sensory
cellular antennae that coordinate a large number of cellular signaling pathways, sometimes coupling
the signaling to ciliary motility or alternatively to cell division and differentiation."[7]
• Eukaryotes can move using motile cilia or flagella. The flagella are more complex than those
of prokaryotes.
Table 1: Comparison of features of prokaryotic and eukaryotic cells
Prokaryotes Eukaryotes
Typical organisms bacteria, archaea protists, fungi, plants, animals
~ 10–100 µm (sperm cells, apart from the

Typical size ~ 1–10 µm
tail, are smaller)
nucleoid region; no
Type of nucleus real nucleus with double membrane
real nucleus
linear molecules (chromosomes) with

DNA circular (usually)
histone proteins
RNA-/protein- coupled in RNA-synthesis inside the nucleus

synthesis cytoplasm protein synthesis in cytoplasm
Ribosomes 50S+30S 60S+40S
Cytoplasmatic very few structures highly structured by endomembranes and

structure a cytoskeleton
flagella made of flagella and cilia containing microtubules;

Cell movement
flagellin lamellipodia and filopodia containing actin
one to several thousand (though some

Mitochondria none
lack mitochondria)
Chloroplasts none in algae and plants
single cells, colonies, higher multicellular

Organization usually single cells
organisms with specialized cells
Binary fission Mitosis (fission or budding)

Cell division
(simple division) Meiosis
Table 2: Comparison of structures between animal and plant cells

Typical animal cell Typical plant cell
• Nucleus • Nucleus
• Nucleolus (within • Nucleolus (within
Organelles nucleus) nucleus)
• Rough endoplasmic • Rough ER
reticulum (ER) • Smooth ER
• Smooth ER
• Ribosomes
• Ribosomes
• Cytoskeleton
• Cytoskeleton
• Golgi apparatus
• Golgi apparatus
(dictiosomes)
• Cytoplasm
• Cytoplasm
• Mitochondria
• Mitochondria
• Vesicles
• Plastids and its derivatives
• Lysosomes
• Vacuole(s)
• Centrosome
• Cell wall
• Centrioles
Subcellular components
The cells of eukaryotes (left) and prokaryotes (right)

All cells, whether prokaryotic or eukaryotic, have a membrane that envelops the cell, separates its
interior from its environment, regulates what moves in and out (selectively permeable), and maintains the
electric potential of the cell. Inside the membrane, a salty cytoplasm takes up most of the cell volume. All
cells possess DNA, the hereditary material of genes, and RNA, containing the information necessary to build
various proteins such as enzymes, the cell's primary machinery. There are also other kinds of biomolecules
in cells. This article will list these primary components of the cell, then briefly describe their function.
Membrane
Main article: Cell membrane
The cytoplasm of a cell is surrounded by a cell membrane or plasma membrane. The plasma
membrane in plants and prokaryotes is usually covered by a cell wall. This membrane serves to separate
and protect a cell from its surrounding environment and is made mostly from a double layer of lipids
(hydrophobic fat-like molecules) and hydrophilic phosphorus molecules. Hence, the layer is called a
phospholipid bilayer. It may also be called a fluid mosaic membrane. Embedded within this membrane is a
variety of protein molecules that act as channels and pumps that move different molecules into and out of the
cell. The membrane is said to be 'semi-permeable', in that it can either let a substance (molecule or ion) pass
through freely, pass through to a limited extent or not pass through at all. Cell surface membranes also
contain receptor proteins that allow cells to detect external signaling molecules such as hormones.
Cytoskeleton
Main article: Cytoskeleton
Bovine Pulmonary Artery Endothelial cell: nuclei stained blue, mitochondria stained red, and F-actin,
an important component in microfilaments, stained green. Cell imaged on a fluorescent microscope.
The cytoskeleton acts to organize and maintain the cell's shape; anchors organelles in place; helps
during endocytosis, the uptake of external materials by a cell, and cytokinesis, the separation of daughter
cells after cell division; and moves parts of the cell in processes of growth and mobility. The eukaryotic
cytoskeleton is composed of microfilaments, intermediate filaments and microtubules. There is a great
number of proteins associated with them, each controlling a cell's structure by directing, bundling, and
aligning filaments. The prokaryotic cytoskeleton is less well-studied but is involved in the maintenance of cell
shape, polarity and cytokinesis.[8]
Genetic material
Two different kinds of genetic material exist: deoxyribonucleic acid (DNA) and ribonucleic acid
(RNA). Most organisms use DNA for their long-term information storage, but some viruses (e.g., retroviruses)
have RNA as their genetic material. The biological information contained in an organism is encoded in its
DNA or RNA sequence. RNA is also used for information transport (e.g., mRNA) and enzymatic functions
(e.g., ribosomal RNA) in organisms that use DNA for the genetic code itself. Transfer RNA (tRNA) molecules
are used to add amino acids during protein translation.
Prokaryotic genetic material is organized in a simple circular DNA molecule (the bacterial
chromosome) in the nucleoid region of the cytoplasm. Eukaryotic genetic material is divided into different,
linear molecules called chromosomes inside a discrete nucleus, usually with additional genetic material in
some organelles like mitochondria and chloroplasts (see endosymbiotic theory).
A human cell has genetic material contained in the cell nucleus (the nuclear genome) and in the
mitochondria (the mitochondrial genome). In humans the nuclear genome is divided into 23 pairs of linear
DNA molecules called chromosomes. The mitochondrial genome is a circular DNA molecule distinct from the
nuclear DNA. Although the mitochondrial DNA is very small compared to nuclear chromosomes, it codes for
13 proteins involved in mitochondrial energy production and specific tRNAs.
Foreign genetic material (most commonly DNA) can also be artificially introduced into the cell by a
process called transfection. This can be transient, if the DNA is not inserted into the cell's genome, or stable,
if it is. Certain viruses also insert their genetic material into the genome.
Organelles
Main article: Organelle
The human body contains many different organs, such as the heart, lung, and kidney, with each
organ performing a different function. Cells also have a set of "little organs," called organelles, that are
adapted and/or specialized for carrying out one or more vital functions. Both eukaryotic and prokaryotic cells
have organelles but organelles in eukaryotes are generally more complex and may be membrane bound.
There are several types of organelles in a cell. Some (such as the nucleus and golgi apparatus) are
typically solitary, while others (such as mitochondria, peroxisomes and lysosomes) can be numerous
(hundreds to thousands). The cytosol is the gelatinous fluid that fills the cell and surrounds the organelles.
Cell nucleus – eukaryotes only - a cell's information
center
The cell nucleus is the most conspicuous
organelle found in a eukaryotic cell. It houses the cell's
chromosomes, and is the place where almost all DNA
replication and RNA synthesis (transcription) occur.
The nucleus is spherical and separated from the
cytoplasm by a double membrane called the nuclear
envelope. The nuclear envelope isolates and protects
a cell's DNA from various molecules that could
accidentally damage its structure or interfere with its
processing. During processing, DNA is transcribed, or
copied into a special RNA, called messenger RNA
(mRNA). This mRNA is then transported out of the
nucleus, where it is translated into a specific protein
molecule. The nucleolus is a specialized region within Diagram of a cell nucleus
the nucleus where ribosome subunits are assembled.
In prokaryotes, DNA processing takes place in the
cytoplasm.
Mitochondria and Chloroplasts – eukaryotes only -

the power generators
Mitochondria are self-replicating organelles
that occur in various numbers, shapes, and sizes in
the cytoplasm of all eukaryotic cells. Mitochondria play
a critical role in generating energy in the eukaryotic
cell. Mitochondria generate the cell's energy by
oxidative phosphorylation, using oxygen to release
energy stored in cellular nutrients (typically pertaining
to glucose) to generate ATP. Mitochondria multiply by
splitting in two. Respiration occurs in the cell
mitochondria.
Organelles that are modified chloroplasts are
broadly called plastids, and are involved in energy
storage through photosynthesis, which uses solar
energy to generate carbohydrates and oxygen from
carbon dioxide and water.[citation needed]
Mitochondria and chloroplasts each contain
their own genome, which is separate and distinct from
the nuclear genome of a cell. Both organelles contain
this DNA in circular plasmids, much like prokaryotic
cells, strongly supporting the evolutionary theory of
endosymbiosis; since these organelles contain their
own genomes and have other similarities to
prokaryotes, they are thought to have developed
through a symbiotic relationship after being engulfed
by a primitive cell.[citation needed]
Endoplasmic reticulum – eukaryotes only

The endoplasmic reticulum (ER) is the
transport network for molecules targeted for certain
modifications and specific destinations, as compared
to molecules that will float freely in the cytoplasm. The
ER has two forms: the rough ER, which has ribosomes
on its surface and secretes proteins into the
cytoplasm, and the smooth ER, which lacks them.
Smooth ER plays a role in calcium sequestration and
release.
Golgi apparatus – eukaryotes only
The primary function of the Golgi apparatus is
to process and package the macromolecules such as
proteins and lipids that are synthesized by the cell. It is
particularly important in the processing of proteins for
secretion. The Golgi apparatus forms a part of the
endomembrane system of eukaryotic cells. Vesicles
that enter the Golgi apparatus are processed in a cis
to trans direction, meaning they coalesce on the cis
side of the apparatus and after processing pinch off on
the opposite (trans) side to form a new vesicle in the
animal cell.[citation needed]
Diagram of an endomembrane system

Ribosomes
The ribosome is a large complex of RNA and
protein molecules. They each consist of two subunits,
and act as an assembly line where RNA from the
nucleus is used to synthesise proteins from amino
acids. Ribosomes can be found either floating freely or
bound to a membrane (the rough endoplasmatic
reticulum in eukaryotes, or the cell membrane in
prokaryotes).[9]
Lysosomes and Peroxisomes – eukaryotes only

Lysosomes contain digestive enzymes (acid hydrolases). They digest excess or worn-out
organelles, food particles, and engulfed viruses or bacteria. Peroxisomes have enzymes that rid the
cell of toxic peroxides. The cell could not house these destructive enzymes if they were not contained
in a membrane-bound system. These organelles are often called a "suicide bag" because of their
ability to detonate and destroy the cell.[citation needed]
Centrosome – the cytoskeleton organiser

The centrosome produces the microtubules of a cell – a key component of the cytoskeleton. It
directs the transport through the ER and the Golgi apparatus. Centrosomes are composed of two
centrioles, which separate during cell division and help in the formation of the mitotic spindle. A single
centrosome is present in the animal cells. They are also found in some fungi and algae cells.[ citation
needed]
Vacuoles
Vacuoles store food and waste. Some vacuoles store extra water. They are often described
as liquid filled space and are surrounded by a membrane. Some cells, most notably Amoeba, have
contractile vacuoles, which can pump water out of the cell if there is too much water. The vacuoles of
eukaryotic cells are usually larger in those of plants than animals.
Structures outside the cell wall
Capsule
A gelatinous capsule is present in some bacteria outside the cell wall. The capsule may be
polysaccharide as in pneumococci, meningococci or polypeptide as Bacillus anthracis or hyaluronic acid as
in streptococci.[citation needed] Capsules are not marked by ordinary stain and can be detected by special
stain. The capsule is antigenic. The capsule has antiphagocytic function so it determines the virulence of
many bacteria. It also plays a role in attachment of the organism to mucous membranes.[ citation needed]
Flagella
Flagella are the organelles of cellular mobility. They arise from cytoplasm and extrude through the
cell wall. They are long and thick thread-like appendages, protein in nature. Are most commonly found in
bacteria cells but are found in animal cells as well.
Fimbriae (pili)
They are short and thin hair like filaments, formed of protein called pilin (antigenic). Fimbriae are
responsible for attachment of bacteria to specific receptors of human cell (adherence). There are special
types of pili called (sex pili) involved in conjunction.[ citation needed]
Functions
Growth and metabolism
Main articles: Cell growth and Metabolism
Between successive cell divisions, cells grow through the functioning of cellular metabolism. Cell
metabolism is the process by which individual cells process nutrient molecules. Metabolism has two distinct
divisions: catabolism, in which the cell breaks down complex molecules to produce energy and reducing
power, and anabolism, in which the cell uses energy and reducing power to construct complex molecules
and perform other biological functions. Complex sugars consumed by the organism can be broken down into
a less chemically complex sugar molecule called glucose. Once inside the cell, glucose is broken down to
make adenosine triphosphate (ATP), a form of energy, through two different pathways.
The first pathway, glycolysis, requires no oxygen and is referred to as anaerobic metabolism. Each
reaction is designed to produce some hydrogen ions that can then be used to make energy packets (ATP). In
prokaryotes, glycolysis is the only method used for converting energy.
The second pathway, called the Krebs cycle, or citric acid cycle, occurs inside the mitochondria and
can generate enough ATP to run all the cell functions.
An overview of protein synthesis.
Within the nucleus of the cell (light blue), genes (DNA, dark blue) are transcribed into RNA. This RNA is then
subject to post-transcriptional modification and control, resulting in a mature mRNA (red) that is then
transported out of the nucleus and into the cytoplasm (peach), where it undergoes translation into a protein.
mRNA is translated by ribosomes (purple) that match the three-base codons of the mRNA to the three-base
anti-codons of the appropriate tRNA. Newly synthesized proteins (black) are often further modified, such as
by binding to an effector molecule (orange), to become fully active.
Creation
Main article: Cell division
Cell division involves a single cell (called a mother cell) dividing into two daughter cells. This leads to
growth in multicellular organisms (the growth of tissue) and to procreation (vegetative reproduction) in
unicellular organisms.
Prokaryotic cells divide by binary fission. Eukaryotic cells usually undergo a process of nuclear
division, called mitosis, followed by division of the cell, called cytokinesis. A diploid cell may also undergo
meiosis to produce haploid cells, usually four. Haploid cells serve as gametes in multicellular organisms,
fusing to form new diploid cells.
DNA replication, or the process of duplicating a cell's genome, is required every time a cell divides.
Replication, like all cellular activities, requires specialized proteins for carrying out the job.
Protein synthesis
Main article: Protein biosynthesis
Cells are capable of synthesizing new proteins, which are essential for the modulation and
maintenance of cellular activities. This process involves the formation of new protein molecules from amino
acid building blocks based on information encoded in DNA/RNA. Protein synthesis generally consists of two
major steps: transcription and translation.
Transcription is the process where genetic information in DNA is used to produce a complementary
RNA strand. This RNA strand is then processed to give messenger RNA (mRNA), which is free to migrate
through the cell. mRNA molecules bind to protein-RNA complexes called ribosomes located in the cytosol,
where they are translated into polypeptide sequences. The ribosome mediates the formation of a polypeptide
sequence based on the mRNA sequence. The mRNA sequence directly relates to the polypeptide sequence
by binding to transfer RNA (tRNA) adapter molecules in binding pockets within the ribosome. The new
polypeptide then folds into a functional three-dimensional protein molecule.
Movement or motility
Cells can move during many processes: such as wound healing, the immune response and cancer
metastasis. For wound healing to occur, white blood cells and cells that ingest bacteria move to the wound
site to kill the microorganisms that cause infection.
At the same time fibroblasts (connective tissue cells) move there to remodel damaged structures. In the case
of tumor development, cells from a primary tumor move away and spread to other parts of the body. Cell
motility involves many receptors, crosslinking, bundling, binding, adhesion, motor and other proteins.[10] The
process is divided into three steps – protrusion of the leading edge of the cell, adhesion of the leading edge
and de-adhesion at the cell body and rear, and cytoskeletal contraction to pull the cell forward. Each step is
driven by physical forces generated by unique segments of the cytoskeleton.[11][12]
Evolution
The origin of cells has to do with the origin of life, which began the history of life on Earth.
Origin of the first cell

Further information: Abiogenesis
There are several theories about the origin of small molecules that could lead to life in an early Earth.
One is that they came from meteorites (see Murchison meteorite). Another is that they were created at deep-
sea vents. A third is that they were synthesized by lightning in a reducing atmosphere ( see Miller–Urey
experiment); although it is not clear if Earth had such an atmosphere. There are essentially no experimental
data defining what the first self-replicating forms were. RNA is generally assumed to be the earliest self-
replicating molecule, as it is capable of both storing genetic information and catalyzing chemical reactions
(see RNA world hypothesis). But some other entity with the potential to self-replicate could have preceded
RNA, like clay or peptide nucleic acid.[13]
Cells emerged at least 4.0–4.3 billion years ago. The current belief is that these cells were
heterotrophs. An important characteristic of cells is the cell membrane, composed of a bilayer of lipids. The
early cell membranes were probably more simple and permeable than modern ones, with only a single fatty
acid chain per lipid. Lipids are known to spontaneously form bilayered vesicles in water, and could have
preceded RNA. But the first cell membranes could also have been produced by catalytic RNA, or even have
required structural proteins before they could form.[14]
Origin of eukaryotic cells
The eukaryotic cell seems to have evolved from a symbiotic community of prokaryotic cells. DNA-
bearing organelles like the mitochondria and the chloroplasts are almost certainly what remains of ancient
symbiotic oxygen-breathing proteobacteria and cyanobacteria, respectively, where the rest of the cell seems
to be derived from an ancestral archaean prokaryote cell – a theory termed the endosymbiotic theory.
There is still considerable debate about whether organelles like the hydrogenosome predated the
origin of mitochondria, or viceversa: see the hydrogen hypothesis for the origin of eukaryotic cells.
Sex, as the stereotyped choreography of meiosis and syngamy that persists in nearly all extant
eukaryotes, may have played a role in the transition from prokaryotes to eukaryotes. An 'origin of sex as
vaccination' theory suggests that the eukaryote genome accreted from prokaryan parasite genomes in
numerous rounds of lateral gene transfer. Sex-as-syngamy (fusion sex) arose when infected hosts began
swapping nuclearized genomes containing co-evolved, vertically transmitted symbionts that conveyed
protection against horizontal infection by more virulent symbionts.[15]
History
• 1632–1723: Antonie van Leeuwenhoek teaches himself to grind lenses, builds a microscope
and draws protozoa, such as Vorticella from rain water, and bacteria from his own mouth.
• 1665: Robert Hooke discovers cells in cork, then in living plant tissue using an early
microscope.[6]
• 1839: Theodor Schwann and Matthias Jakob Schleiden elucidate the principle that plants
and animals are made of cells, concluding that cells are a common unit of structure and
development, and thus founding the cell theory.
• The belief that life forms can occur spontaneously ( generatio spontanea) is contradicted by
Louis Pasteur (1822–1895) (although Francesco Redi had performed an experiment in 1668 that
suggested the same conclusion).
• 1855: Rudolf Virchow states that cells always emerge from cell divisions (omnis cellula ex
cellula).
• 1931: Ernst Ruska builds first transmission electron microscope (TEM) at the University of
Berlin. By 1935, he has built an EM with twice the resolution of a light microscope, revealing
previously unresolvable organelles.
• 1953: Watson and Crick made their first announcement on the double-helix structure for DNA
on February 28.
• 1981: Lynn Margulis published Symbiosis in Cell Evolution detailing the endosymbiotic
theory.
See also
Wikimedia Commons has media related to: Cell biology
Main article: Topic outline of cell biology

• Cell biology
• Cell culture
• Cell type
• Cellular component
• Cytorrhysis
• Cytotoxicity
• Plasmolysis
• Stem cell
• Syncytium
References
1. ^ Cell Movements and the Shaping of the Vertebrate Body in Chapter 21 of Molecular
Biology of the Cell fourth edition, edited by Bruce Alberts (2002) published by Garland Science.
The Alberts text discusses how the "cellular building blocks" move to shape developing embryos. It is
also common to describe small molecules such as amino acids as "molecular building blocks".
2. ^ Integrative Biology 131 - Lecture 03: Skeletal System at YouTube first 12 minutes of the
lecture covers cells (by Marian Diamond).
3. ^ Campbell, Neil A.; Brad Williamson; Robin J. Heyden (2006). Biology: Exploring Life.
4. ^ Mitzi Perdue. "Facts about Birds and Eggs". http://www.eggscape.com/birds.htm. Retrieved
2010-04-15.
5. ^ Maton, Anthea; Hopkins, Jean Johnson, Susan LaHart, David Quon Warner, Maryanna
Wright, Jill D (1997). Cells Building Blocks of Life. New Jersey: Prentice Hall. ISBN 0-13-423476-6.
6. ^ a b "... I could exceedingly plainly perceive it to be all perforated and porous, much like a
Honey-comb, but that the pores of it were not regular [..] these pores, or cells, [..] were indeed the
first microscopical pores I ever saw, and perhaps, that were ever seen, for I had not met with any
Writer or Person, that had made any mention of them before this. . ." – Hooke describing his
observations on a thin slice of cork. Robert Hooke
7. ^ Satir, P; Christensen, ST; Søren T. Christensen (2008-03-26). "Structure and function of
mammalian cilia". Histochemistry and Cell Biology (Springer Berlin / Heidelberg) 129 (6): 687–693.
doi:10.1007/s00418-008-0416-9. 1432-119X. PMID 18365235. PMC 2386530.
http://www.springerlink.com/content/x5051hq648t3152q/. Retrieved 2009-09-12.
8. ^ Michie K, Löwe J (2006). "Dynamic filaments of the bacterial cytoskeleton". Annu Rev
Biochem 75: 467–92. doi:10.1146/annurev.biochem.75.103004.142452. PMID 16756499.
9. ^ Ménétret JF, Schaletzky J, Clemons WM, et al., CW; Akey (December 2007). "Ribosome
binding of a single copy of the SecY complex: implications for protein translocation". Mol. Cell 28 (6):
1083–92. doi:10.1016/j.molcel.2007.10.034. PMID 18158904.
10.^ Revathi Ananthakrishnan1 *, Allen Ehrlicher2 ✉. "The Forces Behind Cell Movement".
Biolsci.org. http://www.biolsci.org/v03p0303.htm. Retrieved 2009-04-17.
11.^ Alberts B, Johnson A, Lewis J. et al. Molecular Biology of the Cell, 4e. Garland Science.
2002
12.^ Ananthakrishnan R, Ehrlicher A. The Forces Behind Cell Movement. Int J Biol Sci 2007;
3:303–317. http://www.biolsci.org/v03p0303.htm
13.^ Orgel LE (1998). "The origin of life--a review of facts and speculations". Trends Biochem
Sci 23 (12): 491–5. doi:10.1016/S0968-0004(98)01300-0. PMID 9868373.
14.^ Griffiths G (December 2007). "Cell evolution and the problem of membrane topology".
Nature reviews. Molecular cell biology 8 (12): 1018–24. doi:10.1038/nrm2287. PMID 17971839.
15.^ Sterrer W (2002). "On the origin of sex as vaccination". Journal of Theoretical Biology 216:
387–396. doi:10.1006/jtbi.2002.3008. PMID 12151256.
• This article incorporates public domain material from the NCBI document "Science
Primer".
External links
• Inside the Cell
• Virtual Cell's Educational Animations
• The Inner Life of A Cell, a flash video showing what happens inside of a cell
• The Virtual Cell
• Cells Alive!
• Journal of Cell Biology
• The Biology Project > Cell Biology
• Centre of the Cell online
• The Image & Video Library of The American Society for Cell Biology , a collection of peer-
reviewed still images, video clips and digital books that illustrate the structure, function and biology of
the cell.
Textbooks
• Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). Molecular Biology of the
Cell (4th ed.). Garland. ISBN 0815332181. http://www.ncbi.nlm.nih.gov/books/bv.fcgi?
rid=mboc4.TOC&depth=2.
• Lodish H, Berk A, Matsudaira P, Kaiser CA, Krieger M, Scott MP, Zipurksy SL, Darnell J
(2004). Molecular Cell Biology (5th ed.). WH Freeman: New York, NY. ISBN 978-0716743668.
http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=mcb.TOC.
• Cooper GM (2000). The cell: a molecular approach (2nd ed.). Washington, D.C: ASM Press.
ISBN 0-87893-102-3. http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=cooper.TOC&depth=2.
[hide]v · d · eStructures of the cell / organelles
Cell membrane · Nucleus (and Nucleolus) · Endoplasmic reticulum ·

Golgi apparatus · Parenthesome · Autophagosome
Endomembrane
system Vesicles (Exosome · Lysosome · Endosome · Phagosome · Vacuole)
Cytoplasmic granules: Melanosome · Microbody (Glyoxysome,
Peroxisome) · Weibel-Palade body
Microfilaments · Intermediate filaments · Microtubules · Prokaryotic

cytoskeleton
Cytoskeleton
MTOCs: Centrosome/Centriole · Basal body · Spindle pole body
Myofibril
Endosymbionts Mitochondrion · Plastids (Chloroplast · Chromoplast · Leucoplast)

RNA: Ribosome · Vault
Other internal
Cytoplasm · Proteasome
Undulipodium: Cilium/Flagellum · Axoneme · Radial spoke

External
Cell wall · Acrosome
B strc: edmb (perx), skel (ctrs), epit, cili, mito, nucl (chro)
Retrieved from "http://en.wikipedia.org/wiki/Cell_(biology)"
Categories: Cell biology

W000
Tetrapod

This article is about four-limbed vertebrates. For the structure, see Tetrapod (structure).
This article has multiple issues. Please help improve it or discuss these issues on
the talk page.
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Tagged since July 2009.
Tetrapod
Tetrapods
Temporal range: Middle Devonian - Recent, 395–0 Ma
PreЄ
g
The four classes of extant tetrapods, (clockwise from upper left), Rana (amphibian), Opisthocomus
(bird), Eumeces (reptile) and Mus (mammal)
Kingdom: Animalia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
(unranked): Eugnathostomata
(unranked): Teleostomi
Tetrapoda
Superclass:
Broili, 1913
Subgroups
• Amphibia
• Reptiliomorpha
and see text
Tetrapods (Greek τετραποδη tetrapodē, equivalent to Latin quadruped, "four-footed") are vertebrate
animals having four limbs. Amphibians, reptiles, birds, and mammals are all tetrapods, and even snakes and
other limbless reptiles and amphibians are tetrapods by descent. The earliest tetrapods evolved from the
lobe-finned fishes in the Devonian.[1] They are now a dominant part of the terrestrial fauna, representing all
known larger land animals. Some groups have even returned to an aquatic existence, including the largest
animal known, the blue whale.
Contents
[hide]
• 1 Evolution
• 1.1 Origin of tetrapods
• 1.2 "The Age of Fishes"
• 1.3 Lungs before land
• 1.3.1 Lobe-finned fishes
• 1.3.2 Denizens of the swamp
• 1.4 Devonian tetrapods
• 1.4.1 Excretion in tetrapods
• 1.4.2 Lungs
• 1.4.3 Fossil early tetrapods
• 1.4.4 From water to land
• 1.5 Carboniferous tetrapods
• 1.6 Permian tetrapods
• 1.7 Mesozoic
• 1.8 Extant (Living) tetrapods
• 2.1 Linnaean classification
• 2.2 Phylogenetic classification
• 2.3 Tetrapod groups
• 2.4 Phylogeny
[edit] Evolution
In Late Devonian vertebrate speciation, descendants of pelagic lobe-finned fish — like

Eusthenopteron — exhibited a sequence of adaptations:
•Panderichthys, suited to muddy shallows;
•Tiktaalik with limb-like fins that could take it onto land;
•Early tetrapods in weed-filled swamps, such as;
•Acanthostega, which had feet with eight digits,
•Ichthyostega with limbs.
Descendants also included pelagic lobe-finned fish such as coelacanth species.
[edit] Origin of tetrapods
The evolution of the first tetrapods marked the moment when the two basic forms of vertebrates,
fishes and tetrapods, diverged.[2] This transition, from a body plan for breathing and navigating in water and
a body plan enabling the animal to move on land, involved a series of changes taking place throughout most
of the 57 million years that make up the Devonian period.[3] While it is one of the most profound evolutionary
changes known, it is also one of the best understood, largely thanks to a number of amazing fossil finds in
the late 20th century combined with improved phylogenetic analysis.[4]
[edit] "The Age of Fishes"

The Devonian period is traditionally known as the "Age of Fishes", marking the diversification of
numerous extinct and modern major fish groups.[5] Among them were the early bony fishes, who diversified
and spread in freshwater and brackish environments at the beginning of the period. The early types
resembled their cartilaginous forefathers in many aspects of their anatomy, including a shark-like tailfin, spiral
gut, large pectoral fins stiffened in front by skeletal elements and a largely unossified axial skeleton.[6]
They did however have certain traits separating them from cartilaginous fishes, traits that would
become pivotal in the evolution of terrestrial forms: The gills did not open singly to exterior like in sharks,
rather they were hidden behind a bony operculum, and bound posteriorly by a stout cleithrum bone, also
functioning as anchoring for the pectoral fins. As part of the overall armour of rhomboid cosmin scales, the
skull had a full cover of dermal bone, constituting a skull roof over the otherwise shark-like cartilaginous inner
cranium. Importantly, they also had a swim bladder/lung, a feature lacking in all other fishes.[7]
[edit] Lungs before land
The primary function of the swim bladder/lung is not entirely certain. One consideration is buoyancy.
The heavy scale armour of the early bony fishes would certainly weigh the animals down. In cartilaginous
fishes, lacking a swim bladder, the open sea sharks need to swim constantly to avoid sinking into the depths,
the pectoral fins providing lift.[8] Another factor is oxygen consumption. Ambient oxygen was relatively low in
the early Devonian, possibly about half of modern values.[9] The partial pressure of oxygen is necessarily
much higher in air than in water, and vertebrates are active animals with high energy requirement compared
to invertebrates of similar sizes.[10][11] The Devonian saw increasing oxygen levels, opening up ecological
niches as active, large bodied animals for groups able to exploit aerial oxygen.[9] Both factors may have
been important, modern fishes with open swim bladders do indeed use their bladders both for buoyancy and
for breathing.
[edit] Lobe-finned fishes

The tetrapods have their root in the early Devonian tetrapodomorph fish.[12] Primitive tetrapods
developed from an osteolepid tetrapodomorph lobe-finned fish (sarcopterygian-crossopterygian), with a two-
lobed brain in a flattened skull. The coelacanth group represent marine sarcopterygians that never acquired
these shallow-water adaptations. The sarcopterygians apparently took two different lines of descent and are
accordingly separated into two major groups the Dipnomorpha (including the Dipnoi or lungfishes) and the
Tetrapodomorpha (which include extinct lines of lobe-finned fishes that evolved into the Tetrapoda).
The oldest known tetrapodomorph is Kenichthys from China, dated at around 395 million years old.
Two of the earliest tetrapodomorphs, dating from 380 Ma, were Gogonasus and Panderichthys. They had a
choana and used their fins as paddles in shallow-water habitats choked with plants and detritus.[13] Their
fins could also have been used to attach themselves to plants or similar while they were lying in ambush for
prey. The universal tetrapod characteristics of front limbs that bend backward at the elbow and hind limbs
that bend forward at the knee can plausibly be traced to early tetrapods living in shallow water.
It has been suggested that the evolution of the tetrapod limb from lobe-finned fishes is related to the
loss of the proteins actinodin 1 and actinodin 2, which are involved in fish fin development[14]
[edit] Denizens of the swamp

The first tetrapods are thought to have evolved in coastal and brackish marine environments, and in
shallow and swampy freshwater habitats.[15] Formerly, the timing was thought to be towards the end of the
Devonian. In 2010, this belief was challenged by the discovery of the oldest known tetrapod tracks,
preserved in marine tidal flat sediments of the southern coast of Laurasia, now Świętokrzyskie (Holy Cross)
Mountains of Poland. They were made during the Eifelian stage of the Middle Devonian. The tracks are
dated to about 395 million years ago, 18 million years earlier than the oldest known tetrapod body fossils.[16]
Some tracks show digits, indicating that the animal had the ability to walk on land. Additionally, the tracks
show that the animal was capable of thrusting its arms and legs forward. This type of motion would have
been impossible in tetrapodomorphs such as Tiktaalik. The animal that produced the tracks is estimated to
have been up to 2.5 metres (8.2 ft) long with footpads up to 26 centimetres (10 in) wide, although most tracks
are only 15 centimetres (5.9 in) wide.[17]
[edit] Devonian tetrapods

Research by Jennifer A. Clack and her colleagues showed that the earliest tetrapods, such as
Acanthostega, were wholly aquatic and quite unsuited to life on land. This is in contrast to the earlier view
that fish had first invaded the land — either in search of prey (like modern mudskippers) or to find water when
the pond they lived in dried out — and later evolved legs, lungs, etc.
By the late Devonian, land plants had stabilized freshwater habitats, allowing the first wetland
ecosystems to develop, with increasingly complex food webs that afforded new opportunities. [1] Freshwater
habitats were not the only places to find water filled with organic matter and choked with plants with dense
vegetation near the water's edge. Swampy habitats like shallow wetlands, coastal lagoons and large brackish
river deltas also existed at this time, and there is much to suggest that this is the kind of environment in which
the tetrapods evolved. Early fossil tetrapods have been found in marine sediments, and because fossils of
primitive tetrapods in general are found scattered all around the world, they must have spread by following
the coastal lines — they could not have lived in freshwater only.
[edit] Excretion in tetrapods

The common ancestor of all present gnathostomes lived in freshwater, and later migrated back to the
sea. To deal with the much higher salinity in sea water, they evolved the ability to turn the nitrogen waste
product ammonia into harmless urea, storing it in the body to make the blood as salty as the sea water
without poisoning the organism. This is the system currently found in cartilaginous fishes and the first bony
fishes (acanthodians). Ray-finned fishes (Actinopterygii) later returned to freshwater and lost this ability,
while the fleshy-finned fishes (Sarcopterygii) retained it. Since the blood of ray-finned fishes contains more
salt than freshwater, they could simply get rid of ammonia through their gills. When they finally returned to
the sea again, they did not recover their old trick of turning ammonia to urea, and they had to evolve salt
excreting glands instead. Lungfishes do the same when they are living in water, making ammonia and no
urea, but when the water dries up and they are forced to burrow down in the mud, they switch to urea
production. Like cartilaginous fishes, the coelacanth can store urea in its blood, as can the only known
amphibians that can live for long periods of time in salt water (the toad Bufo marinus and the frog Rana
cancrivora). These are traits they have inherited from their ancestors.
If early tetrapods lived in freshwater, and if they lost the ability to produce urea and used ammonia
only, they would have to evolve it from scratch again later. Not a single species of all the ray-finned fishes
living today has been able to do that, so it is not likely the tetrapods would have done so either. Terrestrial
animals that can only produce ammonia would have to drink constantly, making a life on land impossible (a
few exceptions exist, as some terrestrial woodlice can excrete their nitrogenous waste as ammonia gas).
This probably also was a problem at the start when the tetrapods started to spend time out of water, but
eventually the urea system would dominate completely. Because of this it is not likely they emerged in
freshwater (unless they first migrated into freshwater habitats and then migrated onto land so shortly after
that they still retained the ability to make urea), although some species never left, or returned to, the water
could of course have adapted to freshwater lakes and rivers.
[edit] Lungs
It is now clear that the common ancestor of the bony fishes (Osteichthyes) had a primitive air-
breathing lung -- later evolved into a swim bladder in most actinopterygians (ray-finned fishes). This suggests
that crossopterygians evolved in warm shallow waters, using their simple lung when the oxygen level in the
water became too low.
Fleshy lobe-fins supported on bones rather than ray-stiffened fins seems to have been an ancestral
trait of all bony fishes (Osteichthyes). The lobe-finned ancestors of the tetrapods evolved them further, while
the ancestors of the ray-finned fishes (Actinopterygii) evolved their fins in a different direction. The most
primitive group of actinopterygians, the bichirs, still have fleshy frontal fins.
[edit] Fossil early tetrapods
Nine genera of Devonian tetrapods have been described, several known mainly or entirely from
lower jaw material. All of them were from the European-North American supercontinent, which comprised
Europe, North America and Greenland. The only exception is a single Gondwanan genus, Metaxygnathus,
which has been found in Australia.
The first Devonian tetrapod identified from Asia was recognized from a fossil jawbone reported in
2002. The Chinese tetrapod Sinostega pani was discovered among fossilized tropical plants and lobe-finned
fish in the red sandstone sediments of the Ningxia Hui Autonomous Region of northwest China. This finding
substantially extended the geographical range of these animals and has raised new questions about the
worldwide distribution and great taxonomic diversity they achieved within a relatively short time.
Eusthenopteron
Panderichthys
Tiktaalik
Acanthostega
Ichthyostega
Hynerpeton
Tulerpeton
Crassigyrinus
Diadectes
These earliest tetrapods were not terrestrial. The earliest confirmed terrestrial forms are known from
the early Carboniferous deposits, some 20 million years later. Still, they may have spent very brief periods
out of water and would have used their legs to paw their way through the mud.
Why they went to land in the first place is still debated. One reason could be that the small juveniles
who had completed their metamorphosis had what it took to make use of what land had to offer. Already
adapted to breathe air and move around in shallow waters near land as a protection (just as modern fish (and
amphibians) often spent the first part of their life in the comparative safety of shallow waters like mangrove
forests), two very different niches partially overlapped each other, with the young juveniles in the diffuse line
between. One of them was overcrowded and dangerous while the other was much safer and much less
crowded, offering less competition over resources. The terrestrial niche was also a much more challenging
place for primary aquatic animals, but because of the way evolution and the selection pressure works, those
juveniles who could take advantage of this would be rewarded. Once they gained a small foothold on land,
thanks to their preadaptations and being at the right place at the right time, favourable variations in their
descendants would gradually result in continuing evolution and diversification.
At this time the abundance of invertebrates crawling around on land and near water, in moist soil and
wet litter, offered a food supply. Some were even big enough to eat small tetrapods, but the land was free
from dangers common in the water.
It is plausible that at first adults would be too heavy and slow and have greater needs for large prey.
Small juveniles would be much lighter, faster and could subsist on relatively small invertebrates. Modern
mudskippers are said to be able to snap insects in flight while on land, and the early juvenile tetrapods might
also have shown formidable abilities.[ citation needed]
[edit] From water to land

Initially making only tentative forays onto land, tetrapods adapted to terrestrial environments over
time and spent longer periods away from the water, while also spending a longer part of their juvenile stage
on land before returning to the water for the rest of their life. It is also possible that the adults started to spend
some time on land (as the skeletal modifications in early tetrapods such as Ichthyostega suggests) but only
to bask in the sun close to the water's edge, not to hunt or move around. The first true tetrapods that were
adapted to terrestrial locomotion were small. Only later did they increase in size.
The fully grown kept most of the anatomical adaptations from their juvenile stage, giving them
modified limbs and other traits associated with a terrestrial lifestyle. To be successful adults they first had to
be successful juveniles. The adults of some of the smaller species were in that case probably able to move
on land too when sufficiently evolved.
If some sort of neoteny or dwarfism occurred, making the animals sexually mature and fully grown
while still living on land, they would only need to visit water to drink and reproduce.
[edit] Carboniferous tetrapods

Until the 1990s, there was a 30 million year gap in the fossil record between the late Devonian
tetrapods and the reappearance of tetrapod fossils in recognizable mid-Carboniferous amphibian lineages. It
was referred to as "Romer's Gap", after the palaeontologist who recognized it.
During the "gap", tetrapod backbones developed, as did limbs with digits and other adaptations for
terrestrial life. Ears, skulls and vertebral columns all underwent changes too. The number of digits on hands
and feet became standardized at five, as lineages with more digits died out. The very few tetrapod fossils
found in the "gap" are all the more precious.
The transition from an aquatic lobe-finned fish to an air-breathing amphibian was a momentous
occasion in the evolutionary history of the vertebrates. For an animal to live in a gravity-neutral, aqueous
environment and then invade one that is entirely different required major changes to the overall body plan,
both in form and in function. Eryops is an example of an animal that made such adaptations. It retained and
refined most of the traits found in its fish ancestors. Sturdy limbs supported and transported its body while out
of water. A thicker, stronger backbone prevented its body from sagging under its own weight. Also, by
utilizing vestigial fish jaw bones, a rudimentary ear was developed, allowing Eryops to hear airborne sound.
By the Visean age of mid-Carboniferous times the early tetrapods had radiated into at least three
main branches. Recognizable basal-group tetrapods are representative of the temnospondyls (e.g. Eryops)
lepospondyls (e.g. Diplocaulus) and anthracosaurs, which were the relatives and ancestors of the Amniota.
Depending on whichever authorities one follows, modern amphibians (frogs, salamanders and caecilians)
are derived from either temnospondyls or lepospondyls (or possibly both, although this is now a minority
position). The first amniotes are known from the early part of the Late Carboniferous, and during the Triassic
counted among their number the earliest mammals, turtles, and crocodiles (lizards and birds appeared in the
Jurassic, and snakes in the Cretaceous). As living members of the tetrapod clan — that is of the tetrapod
"crown-group" — these varied tetrapods represent the phylogenetic end-points of these two divergent
lineages. A fourth Carboniferous group, the baphetids, which are thought to be related to temnospondyls, left
no modern survivors.
[edit] Permian tetrapods

In the Permian period, as the separate tetrapod lineages each developed in their own way, the term
"tetrapoda" becomes less useful. In addition to temnospondyl and anthracosaur clades among the early
"amphibia" (labyrinthodonts), there were two important divergent clades of amniotes, the Sauropsida and the
Synapsida, of which the latter were the most important and successful Permian animals. Each of these
lineages, however, remains grouped with the tetrapoda, just as Homo sapiens could be considered a very
highly-specialized kind of lobe-finned fish.
[edit] Mesozoic
The beginning of the Mesozoic saw a major turnover in fauna following the Permian–Triassic
extinction event. Many of the once large and diverse groups died out or were greatly reduced. Life on Earth
seemed to recover quickly after the Permian extinctions, but this was mostly in the form of disaster taxa, such
as the hardy Lystrosaurus. The most recent research indicates that the specialized animals that formed
complex ecosystems, with high biodiversity, complex food webs and a variety of niches, took much longer to
recover. It is thought that this long recovery was due to the successive waves of extinction which inhibited
recovery, as well as to prolonged environmental stress to organisms which continued into the Early Triassic.
Recent research indicates that recovery did not begin until the start of the mid-Triassic, 4M to 6M years after
the extinction;[18] and some writers estimate that the recovery was not complete until 30M years after the P-
Tr extinction, i.e. in the late Triassic.[19]
A small group of reptiles, the diapsids, began to diversify during the Triassic, notably the dinosaurs.
By the late Mesozoic, the large Laborynthodont groups that first appeared during the Paleozoic such as
temnospondyls and reptile-like amphibians had gone extinct. All current major groups of sauropsids evolved
during the Mesozoic, with birds first appearing in the Jurassic as a derived clade of theropod dinosaurs.
Many groups of synapsids such as anomodontians and therocephalians that once comprised the dominant
terrestrial fauna of the Permian also became extinct during this time, but during the Triassic, one group
(Cynodontia) gave rise to the descendant taxon Mammalia, which survived through the Mesozoic to later
diversify into the dominant terrestrial fauna during the Cenozoic.
[edit] Extant (Living) tetrapods

Following the great faunal turnover at the end of the Mesozoic, only three categories of living crown
group tetrapods were left, all of which also include many extinct groups:
• Lissamphibia : Modern frogs and toads, newts and salamanders, and caecilians
• Sauropsida : Turtles, lepidosaurians (tuataras, lizards, amphisbaenians and snakes), birds,
and crocodilians
• Synapsida : Mammals
Tetrapods were originally classified by means of Linnean taxonomy, but currently their taxonomy is
more frequently being evaluated cladistically.
[edit] Linnaean classification

Traditional classification has the tetrapods classed into four classes based on gross anatomical and
physiological traits. [20] Note that snakes and other legless reptiles are considered tetrapods because they
are descended from ancestors who had a full complement of limbs. Similar considerations apply to caecilians
and aquatic mammals:
• Class Amphibia (Amphibians)
• Class Reptilia (Reptiles)
• Class Aves (Birds)
• Class Mammalia (Mammals)
This classification is the one most commonly encountered in school textbooks and popular works.
While orderly and easy to use, has come under critique from cladistics. The earliest tetrapods are grouped
under Class Amphibia, despite several of the groups would have been more closely related to Amniotes than
to modern day amphibians. Reptiles too form a paraphyletic group, as they have given rise to (birds that are
traditionally not considered to be a type of reptile. basal non-mammalian synapsids ("mammal-like reptiles")
traditionally also sort under Class Reptilia as a separate subclass.[20] They are however not among in the
crown group reptiles, and while the early groups may have been reptile-like in biology, the later ones was
decidedly not reptile-like, nor were they true mammals. Thus some authors have argued for a new
classification based purely on phylogeny, disregarding the anatomy and physiology (see below).
[edit] Phylogenetic classification
All early tetrapods and tetrapodomorphs that were not amphibians in the strict phylogenetic sense,
nor amniotes, were once placed together in the paraphyletic group Labyrinthodontia. Labyrinthodonts were
distinguished mainly by their complex dentine infolding tooth structure, a feature shared with crossopterygian
fish. The labyrinthodonts were divided into the Ichthyostegalia (another paraphyletic assemblage of primitive
tetrapods and kin, such as Ichthyostega), the Temnospondyli (possibly members of Amphibia), and the
Anthracosauria (close relatives of amniotes). The main difference between the three groups was based on
their respective vertebral structures. The Anthracosauria had small pleurocentra, which grew and fused,
becoming the true centrum in later vertebrates. In contrast, the Temnospondyli had a conservative vertebral
column in which the pleurocentra remained small in primitive forms, vanishing entirely in the more advanced
ones. The intercentra are large and form a complete ring. Temnospondyli is thought to have been the sister
group of Anthracosauria, which would eventually give rise to amniotes.
[edit] Tetrapod groups
Pederpes finneyae
Lyddekerina huxleyi
Benthosuchus sushkini
A partial taxonomy of the tetrapods:
• Phylum Chordata
• Class Sarcopterygii
• Subclass Tetrapodomorpha
• Eusthenopteron
• Panderichthys
• Tiktaalik
• Ventastega
• Superclass Tetrapoda
• Family Elginerpetontidae
• Family Acanthostegidae
• Family Ichthyostegidae
• Hynerpeton
• Family Tulerpeton
• Family Crassigyrinidae
• Family Loxommatidae
• Family Colosteidae
• Family Whatcheeriidae
• Family Diadectes
• Batrachomorpha (directly above, below, or redundant to Amphibia)
• Class Amphibia — Amphibians
• Subclass Lepospondyli
• Subclass Temnospondyli
• Subclass Lissamphibia — frogs, salamanders
• Superorder Reptiliomorpha contains among others:
• Series Amniota, which contains among others:
• Class Reptilia — Reptiles
• Class Aves — Birds
• Class Synapsida — Mammal-like reptiles
• Class Mammalia — Mammals
[edit] Phylogeny
Cladogram modified after Ruta, Jeffery, & Coates (2003).[12]
Gephyrostegidae
Eoherpetontidae
Embolomeri
Crown group Tetrapoda
Crown Amniota
Diadectomorpha
Crown Amniota
Lepospondyli
Nectridea
Adelospondyli
Aistopoda
Microsauria
Microbrachomorpha
Lysorophidae
Microbrachomorphs
Crown Lissamphibia*
Tuditanomorphs
Temnospondyli
Eryopoidea
Dissorophoidea
Arch
Rhinesuchidae
Rhytidosteidae
Chigutisauridae
Plagiosauridae
Brachyopidae
Mastodonsauroidea
Metoposauroidea
Trematosauroidea
[edit] Anatomical features of early tetrapods
The tetrapod's ancestral fish must have possessed similar traits to those inherited by the early
tetrapods, including internal nostrils (to separate the breathing and feeding passages) and a large fleshy fin
built on bones that could give rise to the tetrapod limb. The rhipidistian crossopterygians fulfill every
requirement for this ancestry. Their palatal and jaw structures were identical to those of early tetrapods, and
their dentition was identical too, with labyrinthine teeth fitting in a pit-and-tooth arrangement on the palate.
The crossopterygian paired fins were smaller than tetrapod limbs, but the skeletal structure was very similar
in that the crossopterygian had a single proximal bone (analogous to the humerus or femur), two bones in the
next segment (forearm or lower leg), and an irregular subdivision of the fin, roughly comparable to the
structure of the carpus / tarsus and phalanges of a hand.
The major difference between crossopterygians and early tetrapods was in relative development of
front and back skull portions; the snout is much less developed than in most early tetrapods and the post-
orbital skull is exceptionally longer than an amphibian's.
A great many kinds of early tetrapods lived during the Carboniferous period. Therefore, their ancestor
would have lived earlier, during the Devonian period. Devonian Ichthyostega were the earliest of true
tetrapods, with a skeleton that is directly comparable to that of rhipidistian ancestors. Early temnospondyls
(Late Devonian to Early Mississippian) still had some ichthyostegid features such as similar skull bone
patterns, labyrinthine tooth structure, the fish skull-hinge, pieces of gill structure between the cheek and
shoulder, and the vertebral column. They had, however, lost several other fish features such as the fin rays in
the tail.
In order to propagate in the terrestrial environment, certain challenges had to be overcome. The
animal's body needed additional support, because buoyancy was no longer a factor. A new method of
respiration was required in order to extract atmospheric oxygen, instead of oxygen dissolved in water. A
means of locomotion would need to be developed to traverse distances between waterholes. Water retention
was now important since it was no longer the living matrix, and it could be lost easily to the environment.
Finally, new sensory input systems were required if the animal was to have any ability to function reasonably
while on land.
[edit] Skull
The most notable characteristics that make a tetrapod's skull different from a fish's are the relative
frontal and rear portion lengths. The fish had a long rear portion while the front was short; the orbital vacuities
were thus located towards the anterior end. In the tetrapod, the front of the skull lengthened, positioning the
orbits farther back on the skull. The lacrimal bone was not in contact with the frontal anymore, having been
separated from it by the prefrontal bone. Also of importance is that the skull was now free to rotate from side
to side, independent of the spine, on the newly forming neck.
A diagnostic character of temnospondyls is that the tabular bones (which formed the posterior
corners of the skull-table) were separated from the respective left and right parietals by a sutural junction
between the postparietals and supratemporals. Also at the rear of the skull, all bones dorsal to the cleithrum
were lost.
The lower jaw of, for example, Eryops resembled its crossopterygian ancestors in that on the outer
surface lay a long dentary that bore teeth. There were also bones below the dentary on the jaw: two
splenials, the angulary and the surangular. On the inside were usually three coronoids that bore teeth and lay
close to the dentary. On the upper jaw was a row of marginal labyrinthine teeth, located on the maxilla and
premaxilla. In Eryops, as in all early amphibians, the teeth were replaced in waves that traveled from the front
of the jaw to the back in such a way that every other tooth was mature, and the ones in between were young.
[edit] Dentition
The "labyrinthodonts" had a peculiar tooth structure from which their name was derived and,
although not exclusive to the group, the labyrinthine dentition is a useful indicator as to proper classification.
The important feature of the tooth is that the enamel and dentine were folded in such a way as to form a
complicated corrugated pattern when viewed in cross section. This infolding resulted in strengthening of the
tooth and increased wear resistance. Such teeth survived for 100 Ma, first among crossopterygian fish, then
stem reptiles. Modern amphibians no longer have this type of dentition but rather pleurodont teeth, in fewer
numbers of the whole group.
[edit] Sensory organs

There is a density difference between air and water that causes smells (certain chemical compounds
detectable by chemoreceptors) to behave differently. An animal first venturing out onto land would have
difficulty in locating such chemical signals if its sensory apparatus was designed for aquatic detection.
Fish have a lateral line system that detects pressure fluctuations in the water. Such pressure is non-
detectable in air, but grooves for the lateral line sense organs were found on the skull of labyrinthodonts,
suggesting a partially aquatic habitat. Modern amphibians, which are semi-aquatic, exhibit this feature
whereas it has been retired by the higher vertebrates. The olfactory epithelium would also have to be
modified in order to detect airborne odors.
In addition to the lateral line organ system, the eye had to change as well. This change came about
because the refractive index of light differs between air and water, so the focal length of the lens was altered
in order to properly function. The eye was now exposed to a relatively dry environment rather than being
bathed by water, so eyelids developed and tear ducts evolved to produce a liquid, moistening the eyeball.
[edit] Hearing
The balancing function of the middle ear was retained from the fish ancestry, but delicate air
vibrations could not set up pulsations through the skull in order for it to function a proper auditory organ.
Typical of most labyrinthodonts, the spiracular gill pouch was retained as the otic notch, closed in by the
tympanum, a thin, tight membrane.
The hyomandibula of fish migrated upwards from its jaw supporting position, and was reduced in size
to form the stapes. Situated between the tympanum and braincase in an air-filled cavity, the stapes was now
capable of transmitting vibrations from the exterior of the head to the interior. Thus the stapes became an
important element in an impedance matching system, coupling airborne sound waves to the receptor system
of the inner ear. This system had evolved independently within several different amphibian lineages.
In order for the impedance matching ear to work, certain conditions had to be met. The stapes must
have been perpendicular to the tympanum, small and light enough to reduce its inertia and suspended in an
air-filled cavity. In modern species that are sensitive to over 1 kHz frequencies, the footplate of the stapes is
1/20th the area of the tympanum. However, in early amphibians the stapes was too large, making the
footplate area oversized, preventing the hearing of high frequencies. So it appears that only high intensity,
low frequency sounds could be detected, with the stapes more probably being used to support the braincase
against the cheek.
[edit] Girdles
The pectoral girdle of early tetrapods such as Eryops was highly developed, with a larger size for
both increased muscle attachment to it and to the limbs. Most notably, the shoulder girdle was disconnected
from the skull, resulting in improved terrestrial locomotion. The crossopterygian cleithrum was retained as the
clavicle, and the interclavicle was well-developed, lying on the underside of the chest. In primitive forms, the
two clavicles and the interclavical could have grown ventrally in such a way as to form a broad chest plate,
although such was not the case in Eryops. The upper portion of the girdle had a flat, scapular blade, with the
glenoid cavity situated below performing as the articulation surface for the humerus, while ventrally there was
a large, flat coracoid plate turning in toward the midline.
The pelvic girdle also was much larger than the simple plate found in fishes, accommodating more
muscles. It extended far dorsally and was joined to the backbone by one or more specialized sacral ribs. The
hind legs were somewhat specialized in that they not only supported weight, but also provided propulsion.
The dorsal extension of the pelvis was the ilium, while the broad ventral plate was composed of the pubis in
front and the ischium in behind. The three bones met at a single point in the center of the pelvic triangle
called the acetabulum, providing a surface of articulation for the femur.
The main strength of the ilio-sacral attachment of Eryops was by ligaments, a condition structurally,
but not phylogenetically, intermediate between that of the most primitive embolomerous amphibians and
early reptiles. The condition that is more usually found in higher vertebrates is that cartilage and fusion of the
sacral ribs to the blade of the ilium are utilized in addition to ligamentous attachments.
[edit] Limbs
The humerus was the largest bone of the arm, its head articulating with the glenoid cavity of the
pectoral girdle, distally with the radius and ulna. The radius resided on the inner side of the forearm and
rested directly under the humerus, supporting much of the weight, while the ulna was located to the outside
of the humerus. The ulna had a head, which muscles pulled on to extend the limb, called the olecranon that
extended above the edge of the humerus.
The radius and the ulna articulated with the carpus, which was a proximal row of three elements: the
radiale underlying the radius, the ulnare underneath the ulna and an intermedium between the two. A large
central element was beneath the last and may have articulated with the radius. There were also three smaller
centralia lying to the radial side. Opposite the head of each toe lay a series of five distal carpals. Each digit
had a first segment, the metacarpal, lying in the palm region.
The pelvic limb bones were essentially the same as in the pectoral limb, but with different names.
The analogue to the humerus was the femur, which was longer and slimmer. The two lower arm bones
corresponded to the tibia and fibula of the hind leg, the former being the innermost and the latter the
outermost bones. The tarsus is the hind version of the carpus and its bones correspond as well.
[edit] Feeding
Early tetrapods had a wide gaping jaw with weak muscles to open and close it. In the jaw were fang-
like palatal teeth that, when coupled with the gape, suggests an inertial feeding habit. This is when the
amphibian would grasp the prey and, lacking any chewing mechanism, toss the head up and backwards,
throwing the prey farther back into the mouth. Such feeding is seen today in the crocodile and alligator.
The tongue of modern adult amphibians is quite fleshy and attached to the front of the lower jaw, so it
is reasonable to speculate that it was fastened in a similar fashion in primitive forms, although it was probably
not specialized like it is in a frog.
It is taken that early tetrapods were not very active, suggesting that they were not predatory. It is
more likely that it fed on fish either in the water or on those that became stranded at the margins of lakes and
swamps. Also abundant at the time was a large supply of terrestrial invertebrates, which may have provided
a fairly adequate food supply.
[edit] Respiration
Modern amphibians breathe by inhaling air into lungs, where oxygen is absorbed. They also breathe
through the moist lining of the mouth and skin, known as cutaneous respiration. Eryops also inhaled, but its
ribs were too closely spaced to suggest that it did this by expanding the rib cage. More likely, it breathed by
buccal pumping in which it opened its mouth and nostrils, depressed the hyoid apparatus to expand the oral
cavity, closed its mouth and nostrils finally and elevated the floor of the mouth to force air back into the lungs
— in other words, it gulped, then swallowed. It probably exhaled by contraction of the elastic tissue in the lung
walls. Other special respiratory methods probably existed.
[edit] Circulation
Early tetrapods most likely had a three-chambered heart, as do modern amphibians and reptiles, in
which oxygenated blood from the lungs and de-oxygenated blood from the respiring tissues enters by
separate atria, and is directed via a spiral valve to the appropriate vessel — aorta for oxygenated blood and
pulmonary vein for deoxygenated blood. The spiral valve is essential to keeping the mixing of the two types
of blood to a minimum, enabling the animal to have higher metabolic rates, and be more active than
otherwise.
[edit] Locomotion
In typical early tetrapod posture the upper arm and upper leg extended nearly straight horizontal from
its body, and the forearm and the lower leg extended downward from the upper segment at a near right
angle. The body weight was not centered over the limbs, but was rather transferred 90 degrees outward and
down through the lower limbs, which touched the ground. Most of the animal's strength was used to just lift its
body off the ground for walking, which was probably slow and difficult. With this sort of posture, it could only
make short broad strides. This has been confirmed by fossilized footprints found in Carboniferous rocks.
Ligamentous attachments within the limbs were present in Eryops, being important because they
were the precursor to bony and cartilaginous variations seen in modern terrestrial animals that use their limbs
for locomotion.
Of all body parts, the spine was the most affected by the move from water to land. It now had to resist
the bending caused by body weight and had to provide mobility where needed. Previously, it could bend
along its entire length. Likewise, the paired appendages had not been formerly connected to the spine, but
the slowly strengthening limbs now transmitted their support to the axis of the body.
[edit] See also

• Geologic timescale
• Body form
• Hexapod
• Octopod
[edit] References
1. ^ Clack, J.A. (2002). Gaining ground: the origin and evolution of tetrapods. Bloomington,
Indiana, USA.: Indiana University Press.
2. ^ Benton 2005, pp. 76–80
3. ^ Zimmer, C. (1999): At the Water's Edge : Fish with Fingers, Whales with Legs, and How
Life Came Ashore but Then Went Back to Sea. 304 pages. Free Press ISBN 0684856239
4. ^ Shubin, N. (2008): Your Inner Fish: A Journey Into the 3.5-Billion-Year History of the
Human Body New York. Pantheon Books. ISBN 978-0375424472
5. ^ Wells, H.G. (1922): A Short History of the World, chapter IV: The Age of Fishes fra
Bartelby.com
6. ^ Colbert, Edwin H. (1969). Evolution of the Vertebrates (2nd ed.). John Wiley & Sons.
pp. 49–53.
7. ^ Benton 2005, pp. 64–9
8. ^ Videler, J.J. (1993): Fish Swimming. New York: Chapman & Hall
9. ^ a b Dahl & al. (2010): Devonian rise in atmospheric oxygen correlated to the radiations of
terrestrial plants and large predatory fish. PNAS vol. 107 no. 42, pp 17911-17915 Article
10.^ Vaquer-Sunyer, R. & Duarte, C. (2008): Thresholds of hypoxia for marine biodiversity.
Proceedings to the Natural Academy of Science, USA, no 105 (40): pp 15452–15457
11.^ Gray J., & Wu, R. & Or, Y. (2002): Effects of hypoxia and organic enrichment on the coastal
marine environment. Marine Ecology Progress Series, no 238: pp 249–279
12.^ a b Ruta, M.; Jeffery, J. E.; and Coates, M. I. (2003). "A supertree of early tetrapods".
Proceedings of the Royal Society B 270 (1532): 2507–16. doi:10.1098/rspb.2003.2524.
PMID 14667343.
13.^ Monash University. "West Australian Fossil Find Rewrites Land Mammal Evolution."
ScienceDaily 19 October 2006. Accessed 11 March 2009
14.^ Zhang, J.; Wagh, P.; Guay, D.; Sanchez-Pulido, L.; Padhi, B. K.; Korzh, V.; Andrade-
Navarro, M. A.; Akimenko, M. A. �E. (2010). "Loss of fish actinotrichia proteins and the fin-to-limb
transition". Nature 466 (7303): 234. doi:10.1038/nature09137. PMID 20574421. edit
15.^ Clack 2002, pp. 86–7
16.^ Grzegorz Niedźwiedzki, Piotr Szrek, Katarzyna Narkiewicz, Marek Narkiewicz, Per E.
Ahlberg (2010). "Tetrapod trackways from the early Middle Devonian period of Poland". Nature 463
(7277): 43–8. doi:10.1038/nature08623. PMID 20054388.
17.^ Rex Dalton (January 6, 2010). "Discovery pushes back date of first four-legged animal".
Nature News. http://www.nature.com/news/2010/100106/full/news.2010.1.html. Retrieved January 8,
2010.
18.^ Lehrmann, D.J., Ramezan, J., Bowring, S.A., et al. (December 2006). "Timing of recovery
from the end-Permian extinction: Geochronologic and biostratigraphic constraints from south China".
Geology 34 (12): 1053–6. doi:10.1130/G22827A.1.
19.^ Sahney, S., Benton, M.J. (2008). "Recovery from the most profound mass extinction of all
time" (PDF). Proceedings of the Royal Society: Biological 275 (1636): 759–65.
doi:10.1098/rspb.2007.1370. PMID 18198148. PMC 2596898.
20.^ a b Romer, A.S. (1949): The Vertebrate Body. W.B. Saunders, Philadelphia. (2nd ed. 1955;
3rd ed. 1962; 4th ed. 1970)
• Long JA, Young GC, Holland T, Senden TJ, Fitzgerald EM (November 2006). "An
exceptional Devonian fish from Australia sheds light on tetrapod origins". Nature 444 (7116): 199–
202. doi:10.1038/nature05243. PMID 17051154.
• Benton, Michael (2005). Vertebrate Palaeontology (3rd ed.). Blackwell Publishing.

• Cladistic analysis of osteolepiform Sarcopterygians
• Tree of Life: Terrestrial Vertebrates
• Tetrapoda: Overview Overview and cladograms of tetrapods at Palaeos.
• UCMP Taxonomy page
• How did fish grow legs?
• New fossils fill gap between water and land animals
• Human Ears Evolved from Ancient Fish Gills
• Scientific American: Getting a Leg Up on Land
[edit] Devonian tetrapods

• Sinostegia discovery reported in National Geographic
• Devonian Times
• Devonian Times — Tetrapod Trackways
• Scientists find 245 million-year-old burrows of land vertebrates in Antarctica
• Walking with tetrapods Video on recently discovered Middle Devonian tetrapod tracks from
Poland by Nature.
• "Map 2010 — here's oldest tetrapods"
[edit] Carboniferous tetrapods

• Clack JA (July 2002). "An early tetrapod from 'Romer's Gap'". Nature 418 (6893): 72–6.
doi:10.1038/nature00824. PMID 12097908.
• Clack JA (July 1998). "A new Early Carboniferous tetrapod with a mélange of crown-group
characters". Nature 394 (6688): 66–9. doi:10.1038/27895.
http://www.nature.com/nature/journal/v394/n6688/full/394066a0.html.
• "Did Our Ancestors Breathe through Their Ears?"
Retrieved from "http://en.wikipedia.org/wiki/Tetrapod"
Categories: Tetrapods | Fossils | Fossil trackways | Teleostomi

W000
History of paleontology

History of science
Background
Theories/sociology
Duria Antiquior - A more Ancient Dorset is a watercolor painted in Historiography
1830 by the geologist Henry De la Beche based on fossils found by Mary Pseudoscience
Anning. The late 18th and early 19th century was a time of rapid and
dramatic changes in ideas about the history of life on earth. By era
The history of paleontology traces the history of the effort to
understand the history of life on Earth by studying the fossil record left In early cultures
behind by living organisms. Since it is concerned with understanding living in Classical Antiquity
organisms of the past paleontology can be considered to be a field of
biology, but its historical development has been closely tied to geology and In the Middle Ages
the effort to understand the history of the Earth itself. In the Renaissance
Scientific Revolution
Romanticism in science
By culture
Islamic
In ancient times Xenophanes (570-480 BC) wrote about fossil sea shells indicating that land was
once under water. During the Middle Ages, fossils were discussed by the Persian naturalist, Ibn Sina (known
as Avicenna in Europe), in The Book of Healing (1027), which proposed a theory of petrifying fluids that
Albert of Saxony would elaborate on in the 14th century. The Chinese naturalist Shen Kuo (1031–1095)
would propose a theory of climate change based on evidence from petrified bamboo.
In early modern Europe, the systematic study of fossils emerged as an integral part of the changes in
natural philosophy that occurred during the Age of Reason. The nature of fossils and their relationship to life
in the past became better understood during the 17th and 18th centuries, and at the end of the 18th century
the work of Georges Cuvier ended a long running debate about the reality of extinction and led to the
emergence of paleontology, in association with comparative anatomy, as a scientific discipline. The
expanding knowledge of the fossil record also played an increasing role in the development of geology,
particularly stratigraphy.
In 1822 the word "paleontology" was invented by the editor of a French scientific journal to refer to
the study of ancient living organisms through fossils, and the first half of the 19th century saw geological and
paleontological activity become increasingly well organized with the growth of geologic societies and
museums and an increasing number of professional geologists and fossil specialists. This contributed to a
rapid increase in knowledge about the past history of life on Earth, and progress towards definition of the
geologic time scale largely based on fossil evidence. As knowledge of life's past history continued to
improve, it became increasingly obvious that there had been some kind of successive order to the
development of life. This would encourage early evolutionary theories on the transmutation of species.[1]
After Charles Darwin published Origin of Species in 1859, much of the focus of paleontology shifted to
understanding evolutionary paths, including human evolution, and evolutionary theory.[1]
The last half of the 19th century saw a tremendous expansion in paleontological activity, especially in
North America. The trend continued in the 20th century with additional regions of the Earth being opened to
systematic fossil collection, as demonstrated by a series of important discoveries in China near the end of the
century. The last few decades of the 20th century saw a renewed interest in mass extinctions and their role in
the evolution of life on Earth.[2] There was also a renewed interest in the Cambrian explosion that saw the
development of the body plans of most animal phyla. The discovery of fossils of the Ediacaran biota and
developments in paleobiology extended knowledge about the history of life back far before the Cambrian.
Contents
[hide]
• 1 Prior to the 17th century
• 2 17th century
• 3 18th century
• 4 19th century before Darwin
• 4.1 The age of reptiles
• 4.2 Paleobotany and the origin of the word
paleontology
• 4.3 Catastrophism, uniformitarianism and the fossil
record
• 4.4 Transmutation of species and the fossil record
• 4.5 Geological time scale and the history of life
• 4.6 Expansion and professionalization of geology
and paleontology
• 5 19th century after Darwin
• 5.1 Evolution
• 5.2 Developments in North America
• 6 Overview of developments in the 20th century
• 6.1 Developments in geology
• 6.2 Geographical expansion of paleontology
• 6.3 Mass extinctions
[edit] Prior to the 17th century
As early as the 6th century BC, the Greek philosopher Xenophanes of Colophon (570-480 BC)
recognized that some fossil shells were remains of shellfish, which he used to argue that what was at the
time dry land was once under the sea.[3] Leonardo da Vinci (1452–1519), in an unpublished notebook, also
concluded that some fossil sea shells were the remains of shellfish. However, in both cases, the fossils were
complete remains of shellfish species that closely resembled living species, and were therefore easy to
classify.[4]
In 1027, the Persian naturalist, Ibn Sina (known as Avicenna in Europe), proposed an explanation of
how the stoniness of fossils was caused in The Book of Healing. He modified an idea of Aristotle's, which
explained it in terms of vaporous exhalations. Ibn Sina modified this into the theory of petrifying fluids
(succus lapidificatus), which was elaborated on by Albert of Saxony in the 14th century and was accepted in
some form by most naturalists by the 16th century.[5]
Shen Kuo (Chinese: 沈括) (1031–1095) of the Song Dynasty used marine fossils found in the Taihang
Mountains to infer the existence of geological processes such as geomorphology and the shifting of
seashores over time.[6] Using his observation of preserved petrified bamboos found underground in Yan'an,
Shanbei region, Shaanxi province, he argued for a theory of gradual climate change, since Shaanxi was part
of a dry climate zone that did not support a habitat for the growth of bamboos.[7]
As a result of a new emphasis on observing, classifying, and cataloging nature, 16th century natural
philosophers in Europe began to establish extensive collections of fossil objects (as well as collections of
plant and animal specimens), which were often stored in specially built cabinets to help organize them.
Conrad Gesner published a 1565 work on fossils that contained one of the first detailed descriptions of such
a cabinet and collection. The collection belonged to a member of the extensive network of correspondents
that Gesner drew on for his works. Such informal correspondence networks among natural philosophers and
collectors became increasingly important during the course of the 16th century and were direct forerunners of
the scientific societies that would begin to form in the 17th century. These cabinet collections and
correspondence networks played an important role in the development of natural philosophy.[8]
However, most 16th century Europeans did not recognize that fossils were the remains of living
organisms. The etymology of the word fossil comes from the Latin for things having been dug up. As this
indicates, the term was applied to wide variety of stone and stone-like objects without regard to whether they
might have an organic origin. 16th century writers such as Gesner and Georg Agricola were more interested
in classifying such objects by their physical and mystical properties than they were in determining the objects'
origins.[9] In addition, the natural philosophy of the period encouraged alternative explanations for the origin
of fossils. Both the Aristotelian and Neoplatonic schools of philosophy provided support for the idea that
stony objects might grow within the earth to resemble living things. Neoplatonic philosophy maintained that
there could be affinities between living and non-living objects that could cause one to resemble the other.
The Aristotelian school maintained that the seeds of living organisms could enter the ground and generate
objects resembling those organisms.[10]
[edit] 17th century
Johann Jakob Scheuchzer tried to explain fossils using Biblical floods in his Herbarium of the Deluge
(1709)
During the Age of Reason, fundamental changes in natural philosophy were reflected in the analysis
of fossils. In 1665 Athanasius Kircher attributed giant bones to extinct races of giant humans in his Mundus
subterraneus. In the same year Robert Hooke published Micrographia, an illustrated collection of his
observations with a microscope. One of these observations was entitled Of Petrify'd wood, and other Petrify'd
bodies, which included a comparison between petrified and ordinary wood. He concluded that petrified wood
was ordinary wood that had been soaked with "water impregnated with stony and earthy particles". He then
suggested that several kinds of fossil sea shells were formed from ordinary shells by a similar process. He
argued against the prevalent view that such objects were "Stones form'd by some extraordinary Plastick
virtue latent in the Earth itself".[11]
This illustration from Steno's 1667 paper shows a shark head and its teeth along with a fossil tooth
for comparison.
In 1667 Nicholas Steno wrote a paper about a shark head he had dissected. He compared the teeth
of the shark with the common fossil objects known as tongue stones. He concluded that the fossils must have
been shark teeth. Steno then took an interest in the question of fossils, and to address some of the objections
to their organic origin he began studying rock strata. The result of this work was published in 1669 as
Forerunner to a Dissertation on a solid naturally enclosed in a solid . In this book, Steno drew a clear
distinction between objects such as rock crystals that really were formed within rocks and those such as
fossil shells and shark teeth that were formed outside of those rocks. Steno realized that certain kinds of rock
had been formed by the successive deposition of horizontal layers of sediment and that fossils were the
remains of living organisms that had become buried in that sediment. Steno who, like almost all 17th century
natural philosophers, believed that the earth was only a few thousand years old, resorted to the Biblical flood
as a possible explanation for fossils of marine organisms that were far from the sea.[12]
Despite the considerable influence of Forerunner, naturalists such as Martin Lister (1638–1712) and
John Ray (1627–1705) continued to question the organic origin of some fossils. They were particularly
concerned about objects such as fossil Ammonites, which Hooke claimed were organic in origin, that did not
resemble any known living species. This raised the possibility of extinction, which they found difficult to
accept for philosophical and theological reasons.[13]
[edit] 18th century
A drawing comparing jaws was added in 1799 when Cuvier's 1796 presentation on living and fossil
elephants was published.
In his 1778 work Epochs of Nature Georges Buffon referred to fossils, in particular the discovery of
fossils of tropical species such as elephants and rhinoceros in northern Europe, as evidence for the theory
that the earth had started out much warmer than it currently was and had been gradually cooling.
In 1796 Georges Cuvier presented a paper on living and fossil elephants comparing skeletal remains
of Indian and African elephants to fossils of mammoths and of an animal he would later name mastodon
utilizing comparative anatomy. He established for the first time that Indian and African elephants were
different species, and that mammoths differed from both and must be extinct. He further concluded that the
mastodon was another extinct species that also differed from Indian or African elephants, more so than
mammoths. Cuvier made another powerful demonstration of the power of comparative anatomy in
paleontology when he presented a second paper in 1796 on a large fossil skeleton from Paraguay, which he
named Megatherium and identified as a giant sloth by comparing its skull to those of two living species of
tree sloth. Cuvier’s ground-breaking work in paleontology and comparative anatomy lead to the widespread
acceptance of extinction.[14] It also lead Cuvier to advocate the geological theory of catastrophism to explain
the succession of organisms revealed by the fossil record. He also pointed out that since mammoths and
wooly rhinoceros were not the same species as the elephants and rhinoceros currently living in the tropics,
their fossils could not be used as evidence for a cooling earth.
This illustration is from William Smith's 1815 work Strata by Organized Fossils.
In a pioneering application of stratigraphy, William Smith, a surveyor and mining engineer, made
extensive use of fossils to help correlate rock strata in different locations. He created the first geological map
of England during the late 1790s and early 19th century. He established the principle of faunal succession,
the idea that each strata of sedimentary rock would contain particular types of fossils, and that these would
succeed one another in a predictable way even in widely separated geologic formations. At the same time,
Cuvier and Alexandre Brongniart, an instructor at the Paris school of mine engineering, used similar methods
in an influential study of the geology of the region around Paris.
[edit] 19th century before Darwin
[edit] The age of reptiles
This illustration of fossil Iguanodon teeth with a modern iguana jaw for comparison is from Mantell's
1825 paper describing Iguanodon.
In 1808, Cuvier identified a fossil found in Maastricht as a giant marine reptile that he named
Mosasaurus. He also identified, from a drawing, another fossil found in Bavaria as a flying reptile and named
it Pterodactylus. He speculated that an age of reptiles had preceded the first mammals.[15] Cuvier's
speculation would be supported by a series of finds that would be made in Great Britain over the course of
the next two decades. Mary Anning, a professional fossil collector since age eleven, collected the fossils of a
number of marine reptiles from the Jurassic marine strata at Lyme Regis. These included the first ichthyosaur
skeleton to be recognized as such, which was collected in 1811, and the first two plesiosaur skeletons ever
found in 1821 and 1823. Many of her discoveries would be described scientifically by the geologists William
Conybeare, Henry De la Beche, and William Buckland.[16] It was Anning who observed that stony objects
known as "bezoar stones" were often found in the abdominal region of ichthyosaur skeletons, and she noted
that if such stones were broken open they often contained fossilized fish bones and scales as well as
sometimes bones from small ichthyosaurs. This lead Buckland to declare they were fossilized feces, which
he named coprolites, and he used them to better understand ancient food chains.[17]
In 1824, Buckland found and described a lower jaw from Jurassic deposits from Stonesfield. He
determined that the bone belonged to a carnivorous land-dwelling reptile he called Megalosaurus. That same
year Gideon Mantell realized that some large teeth he had found in 1822, in Cretaceous rocks from Tilgate,
belonged to a giant herbivorous land-dwelling reptile. He called it Iguanodon, because the teeth resembled
those of an iguana. In 1832 Mantell would find, in Tilgate, a partial skeleton of an armoured reptile he would
call Hylaeosaurus. In 1842 the English anatomist Richard Owen would create a new order of reptiles, which
he called Dinosauria, for Megalosaurus, Iguanodon, and Hylaeosaurus.[18]
This illustration of the fossil jaw of the Stonesfield mammal is from Gideon Mantell's 1848 book
Wonders of Geology.
This evidence that giant reptiles had lived on Earth in the past caused great excitement in scientific
circles,[19] and even among some segments of the general public.[20] Buckland did describe the jaw of a
small primitive mammal, Phascolotherium, that was found in the same strata as Megalosaurus. This
discovery, known as the Stonesfield mammal, was a much discussed anomaly. Cuvier at first thought it was
a marsupial, but Buckland later realized it was a primitive placental mammal. Due to its small size and
primitive nature, Buckland did not believe it invalidated the overall pattern of an "age of reptiles", when the
largest and most conspicuous animals had been reptiles rather than mammals.[21]
[edit] Paleobotany and the origin of the word paleontology
In 1828 Alexandre Brongniart's son, the botanist Adolphe Brongniart, published the introduction to a
longer work on the history of fossil plants. Adolphe Brongniart concluded that the history of plants could
roughly be divided into four parts. The first period was characterized by cryptogams. The second period was
characterized by the appearance of the conifers. The third period brought emergence of the cycads, and the
fourth by the development of the flowering plants (such as the dicotyledons). The transitions between each of
these periods was marked by sharp discontinuities in the fossil record, with more gradual changes within the
periods. Brongniart's work is the foundation of paleobotany and reinforced the theory that life on earth had a
long and complex history, and different groups of plants and animals made their appearances in successive
order.[22]
The increasing attention being paid to fossil plants in the first decades of the 19th century also
caused a significant change in the terminology for the study of past life. The editor of the influential French
scientific journal, Journal de Phisique, a student of Cuvier's named Henri Marie Ducrotay de Blanville, coined
the term "paleozoologie" in 1817 to refer to the work Cuvier and others were doing to reconstruct extinct
animals from fossil bones. However, Blanville began looking for a term that could refer to the study of both
fossil animal and plant remains. After trying some unsuccessful alternatives, he hit on "paleontologie" in
1822. Blanville's term for the study of the living organisms of the past quickly became popular and was
anglicized into "paleontology".[23]
[edit] Catastrophism, uniformitarianism and the fossil record

In Cuvier's landmark 1796 paper on living and fossil elephants, he referred to a single catastrophe
that destroyed life to be replaced by the current forms. As a result of his studies of extinct mammals, he
realized that animals such as Palaeotherium had lived before the time of the mammoths, which lead him to
write in terms of multiple geological catastrophes that had wiped out a series of successive faunas.[24] By
1830, a scientific consensus had formed around his ideas as a result of paleobotany and the dinosaur and
marine reptile discoveries in Britain.[25] In Great Britain, where natural theology was very influential in the
early 19th century, a group of geologists that included Buckland, and Robert Jameson insisted on explicitly
linking the most recent of Cuvier's catastrophes to the biblical flood. Catastrophism had a religious overtone
in Britain that was absent elsewhere.[26]
Partly in response to what he saw as unsound and unscientific speculations by William Buckland and
other practitioners of flood geology, Charles Lyell advocated the geological theory of uniformitarianism in his
influential work Principles of Geology.[27] Lyell amassed evidence, both from his own field research and the
work of others, that most geological features could be explained by the slow action of present day forces,
such as vulcanism, earthquakes, erosion, and sedimentation rather than past catastrophic events.[28] Lyell
also claimed that the apparent evidence for catastrophic changes in the fossil record, and even the
appearance of directional succession in the history of life, were illusions caused by imperfections in that
record. For instance he argued that the absence of birds and mammals from the earliest fossil strata was
merely an imperfection in the fossil record attributable to the fact that marine organisms were more easily
fossilized.[28] Also Lyell pointed to the Stonesfield mammal as evidence that mammals had not necessarily
been preceded by reptiles, and to the fact that certain Pleistocene strata showed a mixture of extinct and still
surviving species, which he said showed that extinction occurred piecemeal rather than as a result of
catastrophic events.[29] Lyell was successful in convincing geologists of the idea that the geological features
of the earth were largely due to the action of the same geologic forces that could be observed in the present
day, acting over an extended period of time. He was not successful in gaining support for his view of the
fossil record, which he believed did not support a theory of directional succession.[30]
[edit] Transmutation of species and the fossil record
See also: Transmutation of species
Jean Baptiste Lamarck used fossils in his arguments for his theory of the transmutation of species in
the early 19th century.[31] Fossil finds, and the emerging evidence that life had changed over time, fueled
speculation on this topic during the next few decades.[32] Robert Chambers used fossil evidence in his 1844
popular science book Vestiges of the Natural History of Creation , which advocated an evolutionary origin for
the cosmos as well as for life on earth. Like Lamarck's theory it maintained that life had progressed from the
simple to the complex.[33] These early evolutionary ideas were widely discussed in scientific circles but were
not accepted into the scientific mainstream.[34] Many of the critics of transmutational ideas used fossil
evidence in their arguments. In the same paper that coined the term dinosaur Richard Owen pointed out that
dinosaurs were at least as sophisticated and complex as modern reptiles, which he claimed contradicted
transmutational theories.[35] Hugh Miller would make a similar argument, pointing out that the fossil fish
found in the Old Red Sandstone formation were fully as complex as any later fish, and not the primitive forms
alleged by Vestiges.[36] While these early evolutionary theories failed to become accepted as mainstream
science, the debates over them would help pave the way for the acceptance of Darwin's theory of evolution
by natural selection a few years later.[37]
This diagram of the geologic time scale from an 1861 book by Richard Owen shows the appearance
of major animal types.
[edit] Geological time scale and the history of life

Geologists such as Adam Sedgwick, and Roderick Murchison continued, in the course of disputes
such as The Great Devonian Controversy, to make advances in stratigraphy. They described new geological
epochs such as the Cambrian, the Silurian, the Devonian, and the Permian. Increasingly, such progress in
stratigraphy depended on the opinions of experts with specialized knowledge of particular types of fossils
such as William Lonsdale (fossil corals), and John Lindley (fossil plants) who both played a role in the
Devonian controversy and its resolution.[38] By the early 1840s much of the geologic time scale had been
developed. In 1841, John Phillips formally divided the geologic column into three major eras, the Paleozoic,
Mesozoic, and Cenozoic, based on sharp breaks in the fossil record.[39] He identified the three periods of
the Mesozoic era and all the periods of the Paleozoic era except the Ordovician. His definition of the
geological time scale is still used today.[40] It remained a relative time scale with no method of assigning any
of the periods' absolute dates. It was understood that not only had there been an "age of reptiles" preceding
the current "age of mammals", but there had a time (during the Cambrian and the Silurian) when life had
been restricted to the sea, and a time (prior to the Devonian) when invertebrates had been the largest and
most complex forms of animal life.
[edit] Expansion and professionalization of geology and paleontology

This rapid progress in geology and paleontology during the 1830s and 1840s was aided by a growing
international network of geologists and fossil specialists whose work was organized and reviewed by an
increasing number of geological societies. Many of these geologists and paleontologists were now paid
professionals working for universities, museums and government geological surveys. The relatively high level
of public support for the earth sciences was due to their cultural impact, and their proven economic value in
helping to exploit mineral resources such as coal.[41]
Another important factor was the development in the late 18th and early 19th centuries of museums
with large natural history collections. These museums received specimens from collectors around the world
and served as centers for the study of comparative anatomy and morphology. These disciplines played key
roles in the development of a more technically sophisticated form of natural history. One of the first and most
important examples was the Museum of Natural History in Paris, which was at the center of many of the
developments in natural history during the first decades of the 19th century. It was founded in 1793 by an act
of the French National Assembly, and was based on an extensive royal collection plus the private collections
of aristocrats confiscated during the French revolution, and expanded by material seized during French
military conquests. The Paris museum was the professional base for Cuvier, and his professional rival
Geoffroy Saint-Hilaire. The English anatomists Robert Grant and Richard Owen both spent time studying
there. Owen would go on to become the leading British morphologist while working at the museum of the
Royal College of Surgeons.[42][43]
[edit] 19th century after Darwin

[edit] Evolution
This photo of the second Archaeopteryx skeleton to be found was taken in 1881 at the Humboldt
Museum in Berlin.
Charles Darwin's publication of the On the Origin of Species in 1859 was a watershed event in all the
life sciences, especially paleontology. Fossils had played a role in the development of Darwin's theory. In
particular he had been impressed by fossils he had collected in South America during the voyage of the
Beagle of giant armadillos, giant sloths, and what at the time he thought were giant llamas that seemed to be
related to species still living on the continent in modern times.[44] The scientific debate that started
immediately after the publication of Origin led to a concerted effort to look for transitional fossils and other
evidence of evolution in the fossil record. There were two areas where early success attracted considerable
public attention, the transition between reptiles and birds, and the evolution of the modern single-toed horse.
[45] In 1861 the first specimen of Archaeopteryx, an animal with both teeth and feathers and a mix of other
reptilian and avian features, was discovered in a limestone quarry in Bavaria and described by Richard
Owen. Another would be found in the late 1870s and put on display at a Museum in Berlin in 1881. Other
primitive toothed birds were found by Othniel Marsh in Kansas in 1872. Marsh also discovered fossils of
several primitive horses in the Western United States that helped trace the evolution of the horse from the
small 5-toed Hyracotherium of the Eocene to the much larger single-toed modern horses of the genus Equus.
Thomas Huxley would make extensive use of both the horse and bird fossils in his advocacy of evolution.
Acceptance of evolution occurred rapidly in scientific circles, but acceptance of Darwin's proposed
mechanism of natural selection as the driving force behind it was much less universal. In particular some
paleontologists such as Edward Drinker Cope and Henry Fairfield Osborn preferred alternatives such as neo-
Lamarckism, the inheritance of characteristics acquired during life, and orthogenesis, an innate drive to
change in a particular direction, to explain what they perceived as linear trends in evolution.[46]
This diagram by O.C. Marsh of the evolution of horse feet and teeth over time was reproduced in T.H
Huxley's 1876 book, Professor Huxley in America.
There was also great interest in human evolution. Neanderthal fossils were discovered in 1856, but
at the time it was not clear that they represented a different species from modern humans. Eugene Dubois
created a sensation with his discovery of Java Man, the first fossil evidence of a species that seemed clearly
intermediate between humans and apes, in 1891.
[edit] Developments in North America

A major development in the second half of the 19th century was a rapid expansion of paleontology in
North America. In 1858 Joseph Leidy described a Hadrosaurus skeleton, which was the first North American
dinosaur to be described from good remains. However, it was the massive westward expansion of railroads,
military bases, and settlements into Kansas and other parts of the Western United States following the
American Civil War that really fueled the expansion of fossil collection.[47] The result was an increased
understanding of the natural history of north America, including the discovery of the Western Interior Sea that
had covered Kansas and much of the rest of the Midwestern United States during parts of the Cretaceous,
the discovery several important fossils of primitive birds and horses, and the discovery of a number of new
dinosaur genera including Allosaurus, Stegosaurus, and Triceratops. Much of this activity was part of a fierce
personal and professional rivalry between two men, Othniel Marsh, and Edward Cope, which has become
known as the Bone Wars.
[edit] Overview of developments in the 20th century

[edit] Developments in geology
Two 20th century developments in geology had a big effect on paleontology. The first was the
development of radiometric dating, which allowed absolute dates to be assigned to the geologic timescale.
The second was the theory of plate tectonics, which helped make sense of the geographical distribution of
ancient life.
[edit] Geographical expansion of paleontology

During the 20th century paleontological exploration intensified everywhere and ceased to be a
largely European and North American activity. In the 135 years between Buckland's first discovery and 1969
a total of 170 dinosaur genera were known. In the 25 years after 1969 that number increased to 315. Much of
this increase was due to the examination of new rock exposures, particularly in previously little-explored
areas in South America and Africa.[48] Near the end of the century the opening of China to systematic
exploration for fossils has yielded a wealth of material on dinosaurs and the origin of birds and mammals.[49]

The 20th century saw a major renewal of interest in mass extinction events and their effect on the
course of the history of life. This was particularly true after 1980 when Luis and Walter Alvarez put forward
the Alvarez hypothesis claiming that an impact event caused the Cretaceous–Tertiary extinction event, which
killed off the non-avian dinosaurs along with many other living things.[50] Also in the early 1980s Jack
Sepkoski and David M. Raup published papers with statistical analysis of the fossil record of marine
invertebrates that revealed a pattern (possibly cyclical) of repeated mass extinctions with significant
implications for the evolutionary history of life.
[edit] Evolutionary paths and theory
Photo shows the fossils of Taung child discovered in South Africa in 1924.
Throughout the 20th century new fossil finds continued to contribute to understanding the paths
taken by evolution. Examples include major taxonomic transitions such as finds in Greenland, starting in the
1930s (with more major finds in the 1980s), of fossils illustrating the evolution of tetrapods from fish, and
fossils in China during the 1990s that shed light on the dinosaur-bird relationship. Other events that have
attracted considerable attention have included the discovery of a series of fossils in Pakistan that have shed
light on whale evolution, and most famously of all a series of finds throughout the 20th century in Africa
(starting with Taung child in 1924[51]) and elsewhere have helped illuminate the course of human evolution.
Increasingly, at the end of the century, the results of paleontology and molecular biology were being brought
together to reveal detailed phylogenetic trees.
The results of paleontology have also contributed to the development of evolutionary theory. In 1944
George Gaylord Simpson published Tempo and Mode in Evolution, which used quantitative analysis to show
that the fossil record was consistent with the branching, non-directional, patterns predicted by the advocates
of evolution driven by natural selection and genetic drift rather than the linear trends predicted by earlier
advocates of neo-Lamarckism and orthogenesis. This integrated paleontology into the modern evolutionary
synthesis.[52] In 1972 Niles Eldredge and Stephen Jay Gould used fossil evidence to advocate the theory of
punctuated equilibrium, which maintains that evolution is characterized by long periods of relative stasis and
much shorter periods of relatively rapid change.[53]
[edit] Cambrian explosion
Photo shows a complete Anomalocaris fossil from the Burgess shale.

One area of paleontology that has seen a lot of activity during the 1980s, 1990s, and beyond is the
study of the Cambrian explosion during which many of the various phyla of animals with their distinctive body
plans first appear. The well-known Burgess Shale Cambrian fossil site was found in 1909 by Charles Doolittle
Walcott, and another important site in Chengjiang China was found in 1912. However, new analysis in the
1980s by Harry B. Whittington, Derek Briggs, Simon Conway Morris and others sparked a renewed interest
and a burst of activity including discovery of an important new fossil site, Sirius Passet, in Greenland, and the
publication of a popular and controversial book, Wonderful Life by Stephen Jay Gould in 1989.[54]
[edit] Pre-Cambrian fossils
Photo shows a Spriggina fossil from the Ediacaran.

Prior to 1950 there was no widely accepted fossil evidence of life before the Cambrian period. When
Charles Darwin wrote The Origin of Species he acknowledged that the lack of any fossil evidence of life prior
to the relatively complex animals of the Cambrian was a potential argument against the theory of evolution,
but expressed the hope that such fossils would be found in the future. In the 1860s there were claims of the
discovery of pre-Cambrian fossils, but these would later be shown not to have an organic origin. In the late
19th century Charles Doolittle Walcott would discover stromatolites and other fossil evidence of pre-
Cambrian life, but at the time the organic origin of those fossils was also disputed. This would start to change
in the 1950s with the discovery of more stromatolites along with microfossils of the bacteria that built them,
and the publication of a series of papers by the Soviet scientist Boris Vasil'evich Timofeev announcing the
discovery of microscopic fossil spores in pre-Cambrian sediments. A key breakthrough would come when
Martin Glaessner would show that fossils of soft bodied animals discovered by Reginald Sprigg during the
late 1940s in the Ediacaran hills of Australia were in fact pre-Cambrian not early Cambrian as Sprigg had
originally believed, making the Ediacaran biota the oldest animals known. By the end of the 20th century
paleobiology had established that the history of life extended back at least 3.5 billion years.[55]
[edit] See also

• List of years in paleontology
• History of geology
• History of science
[edit] Notes
1. ^ a b Buckland W & Gould SJ (1980). Geology and Mineralogy Considered With Reference to
Natural Theology (History of Paleontology). Ayer Company Publishing. ISBN 978-0405127069.
2. ^ Bowler Evolution: The History of an Idea pp. 351-352
3. ^ Desmond p. 692-697.
4. ^ Rudwick The Meaning of Fossils p. 39
5. ^ Rudwick The Meaning of Fossils p. 24
6. ^ Shen Kuo,Mengxi Bitan (梦溪笔谈; Dream Pool Essays) (1088)
7. ^ Needham, Volume 3, p. 614.
8. ^ Rudwick The Meaning of Fossils pp. 9-17
9. ^ Rudwick The Meaning of Fossils pp. 23-33
10.^ Rudwick The Meaning of Fossils pp. 33-36
11.^ Hooke Micrographia observation XVII
12.^ Rudwick The Meaning of Fossils pp 72-73
13.^ Rudwick The Meaning of Fossils pp 61-65
14.^ McGowan the dragon seekers pp. 3-4
15.^ Rudwick Georges Cuvier, Fossil Bones and Geological Catastrophes p. 158
16.^ McGowan pp. 11-27
17.^ Rudwick, Martin Worlds Before Adam: The Reconstruction of Geohistory in the Age of
Reform (2008) pp. 154-155.
18.^ McGowan p. 176
20.^ McGowan p. 109
23.^ Rudwick Worlds before Adam p. 48
28.^ a b McGowan pp. 100-103
30.^ McGowan pp. 100
31.^ Rudwick The Meaning of Fossils p. 119
32.^ McGowan p. 8
33.^ McGowan pp. 188-191
34.^ Larson p. 73
35.^ Larson p. 44
36.^ Ruckwick The Meaning of fossils pp. 206-207
37.^ Larson p. 51
38.^ Rudwick The Great Devonian Controversy p. 94
39.^ Larson pp. 36-37
40.^ Rudwick The Meaning of Fossils p. 213
42.^ Greene and Depew The Philosophy of Biology pp. 128-130
43.^ Bowler and Morus Making Modern Science pp. 168-169
44.^ Bowler Evolution: The History of an Idea p. 150
45.^ Larson Evolution p. 139
46.^ Larson pp. 126-127
47.^ Everhart Oceans of Kansas p. 17
48.^ McGowan p. 105
49.^ Bowler p. 349
50.^ Alvarez, LW, Alvarez, W, Asaro, F, and Michel, HV (1980). "Extraterrestrial cause for the
Cretaceous–Tertiary extinction". Science 208 (4448): 1095–1108.
doi:10.1126/science.208.4448.1095. PMID 17783054.
51.^ Garwin, Laura; Tim Lincoln. "A Century of Nature: Twenty-One Discoveries that Changed
Science and the World". University of Chicago Press. pp. 3–9.
http://www.press.uchicago.edu/Misc/Chicago/284158_brain.html. Retrieved 2009-07-19.
52.^ Bowler p. 337
53.^ Eldredge, Niles and S. J. Gould (1972). "Punctuated equilibria: an alternative to phyletic
gradualism" In T.J.M. Schopf, ed., Models in Paleobiology. San Francisco: Freeman Cooper. pp. 82-
115. Reprinted in N. Eldredge Time frames. Princeton: Princeton Univ. Press, 1985. Available here
[1].
54.^ Briggs, D. E. G.; Fortey, R. A. (2005). "Wonderful strife: systematics, stem groups, and the
phylogenetic signal of the Cambrian radiation". Paleobiology 31 (2 (Supplement)): 94–112.
doi:10.1666/0094-8373(2005)031[0094:WSSSGA]2.0.CO;2.
http://paleobiol.geoscienceworld.org/cgi/reprint/31/2_Suppl/94.pdf.
55.^ Schopf, J. William. "Solution to Darwin's dilemma: Discovery of the missing Precambrian
record of life". Proceedings of the National Academy of Sciences.
[edit] References
• Bowler, Peter J. (2003). Evolution:The History of an Idea. University of California Press.
ISBN 0-52023693-9.
• Bowler, Peter J.; Iwan Rhys Morus (2005). Making Modern Science. The University of
Chicago Press. ISBN 0-226-06861-7.
• Desmond, Adrian (1975). "The Discovery of Marine Transgressions and the Explanation of
Fossils in Antiquity". American Journal of Science, Volume 275.
• Larson, Edward J. (2004). Evolution: the remarkable history of scientific theory . Modern
Library. ISBN 0-679-64288-9.
• McGowan, Christopher (2001). The Dragon Seekers. Persus Publishing. ISBN 0-7382-0282-
7.
• Everhart, Michael J. (2005). Oceans of Kansas: A Natural History of the Western Interior
Sea. Indiana University Press. ISBN 0-253-34547-2.
• Greene, Marjorie; David Depew (2004). The Philosophy of Biology:An Episodic History.
Cambridge University Press. ISBN 0-521-64371-6.
• Needham, Joseph (1986). Science and Civilization in China: Volume 3, Mathematics and the
Sciences of the Heavens and the Earth. Caves Books Ltd. ISBN 0-253-34547-2.
• Robert Hooke (1665) Micrographia The Royal Society
• Palmer, Douglas (2005) Earth Time: Exploring the Deep Past from Victorian England to the
Grand Canyon. Wiley, Chichester. ISBN 9780470022214
• Rudwick, Martin J.S. (1997). Georges Cuvier, Fossil Bones, and Geological Catastrophes.
The University of Chicago Press. ISBN 0-226-73106-5.
• Rudwick, Martin J.S. (1985). The Meaning of Fossils (2nd ed.). The University of Chicago
Press. ISBN 0-226-73103-0.
• Rudwick, Martin J.S. (1985). The Great Devonian Controversy: The Shaping of Scientific
Knowledge among Gentlemanly Specialists. The University of Chicago Press. ISBN 0-226-73102-2.
• Rudwick, Martin J.S. (2008). Worlds Before Adam: The Reconstruction of Geohistory in the
Age of Reform. The University of Chicago Press. ISBN 0-226-73128-6.
• Zittel, Karl Alfred von (1901). History of geology and palaentology to the end of the
Nineteenth Century. Charles Scribner's Sons, London.
http://www.archive.org/details/historyofgeology00zittrich.

• History of palaeoentomology in Russia
Natural history • History of agriculture • History of botany •

History of medicine • History of anatomy • History of zoology
Fields and systematics • History of geology • History of paleontology • History
disciplines of evolutionary thought • History of ecology • History of model
organisms • History of phycology • History of genetics • History of
biochemistry • History of agricultural science • History of
molecular biology • History of RNA biology • History of molecular
evolution • History of immunology • History of biotechnology

Institutions Max Planck Society • Cold Spring Harbor Laboratory • Stazione
Zoologica • Marine Biological Laboratory • Rockefeller University •
Woods Hole Oceanographic Institute • Laboratory of Molecular
Biology
Griffith's experiment • Miller-Urey experiment • Luria–

Experiments Hershey–Chase experiment • Meselson–Stahl experiment • Crick,


Aristotle • Andreas Vesalius • William Harvey • Antonie
van Leeuwenhoek • Carolus Linnaeus • Georges-Louis Leclerc,
Influential Hugo de Vries • E. B. Wilson • Thomas Hunt Morgan • Aleksandr
figures Oparin • Alexander Fleming • J. B. S. Haldane • Sewall Wright • R.
A. Fisher • Konrad Lorenz • Barbara McClintock • Theodosius

Retrieved from "http://en.wikipedia.org/wiki/History_of_paleontology"

Categories: History of paleontology | Paleontologists
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Population genetics
Systematics
The modern evolutionary synthesis is a union of ideas from several biological specialties which
provides a widely accepted account of evolution. It is also referred to as the new synthesis, the modern
synthesis, the evolutionary synthesis, millennium synthesis and the neo-darwinian synthesis.
The synthesis reflects the current overwhelming consensus.[1] The synthesis was produced over a
decade (1936–1947). The previous development of population genetics (1918–1932) was a stimulus, as it
showed that Mendelian genetics was consistent with natural selection and gradual evolution. The synthesis
is still, to a large extent, the current paradigm in evolutionary biology.[2]
The modern synthesis solved difficulties and confusions caused by the specialisation and poor
communication between biologists in the early years of the 20th century. At its heart was the question of
whether Mendelian genetics could be reconciled with gradual evolution by means of natural selection. A
second issue was whether the broad-scale changes ('macroevolution') seen by palaeontologists could be
explained by changes seen in local populations ('microevolution').
The synthesis included evidence from biologists, trained in genetics, who studied populations in the
field and in the laboratory. These studies were crucial to evolutionary theory. The synthesis drew together
ideas from several branches of biology which had become separated, particularly genetics, cytology,
systematics, botany, morphology, ecology and paleontology.
Julian Huxley invented the term, when he produced his book, Evolution: The Modern Synthesis
(1942). Other major figures in the modern synthesis include R. A. Fisher, Theodosius Dobzhansky, J.B.S.
Haldane, Sewall Wright, E.B. Ford, Ernst Mayr, Bernhard Rensch, Sergei Chetverikov, George Gaylord
Simpson, and G. Ledyard Stebbins.
Contents
[hide]
• 1 Summary of the modern synthesis
• 2 Developments leading up to the synthesis
• 2.1 1859–1899
• 2.2 1900–1915
• 2.3 The foundation of population genetics
• 3 The modern synthesis
• 4 Further advances
• 5 After the synthesis
• 5.1 Understanding of Earth history
• 5.1.1 Symbiotic origin of eukaryotic cell
structures
• 5.1.2 Trees of life
• 5.1.3 Evo-devo
• 5.1.4 Fossil discoveries
• 6 See also
• 7 Footnotes
• 8 References
[edit] Summary of the modern synthesis
The modern synthesis bridged the gap between experimental geneticists and naturalists, and
between both and palaeontologists. It states that:[3][4][5]
1. All evolutionary phenomena can be explained in a way consistent with known genetic
mechanisms and the observational evidence of naturalists.
2. Evolution is gradual: small genetic changes, recombination ordered by natural selection.
Discontinuities amongst species (or other taxa) are explained as originating gradually through
geographical separation and extinction (not saltation).
3. Natural selection is by far the main mechanism of change; even slight advantages are
important when continued. The object of selection is the phenotype in its surrounding environment.
4. The role of genetic drift is equivocal. Though strongly supported initially by Dobzhansky, it
was downgraded later as results from ecological genetics were obtained.
5. Thinking in terms of populations, rather than individuals, is primary: the genetic diversity
existing in natural populations is a key factor in evolution. The strength of natural selection in the wild
is greater than previously expected; the effect of ecological factors such as niche occupation and the
significance of barriers to gene flow are all important.
6. In palaeontology, the ability to explain historical observations by extrapolation from
microevolution to macroevolution is proposed. Historical contingency means explanations at different
levels may exist. Gradualism does not mean constant rate of change.
The idea that speciation occurs after populations are reproductively isolated has been much debated.
In plants, polyploidy must be included in any view of speciation. Formulations such as 'evolution consists
primarily of changes in the frequencies of alleles between one generation and another' were proposed rather
later. The traditional view is that developmental biology ('evo-devo') played little part in the synthesis,[6] but
an account of Gavin de Beer's work by Stephen J. Gould suggests he may be an exception.[7]
[edit] Developments leading up to the synthesis

[edit] 1859–1899
Charles Darwin's The Origin of Species was successful in convincing most biologists that evolution
had occurred, but was less successful in convincing them that natural selection was its primary mechanism.
In the 19th and early 20th centuries, variations of Lamarckism, orthogenesis ('progressive' evolution), and
saltationism (evolution by jumps) were discussed as alternatives.[8] Also, Darwin did not offer a precise
explanation of how new species arise. As part of the disagreement about whether natural selection alone
was sufficient to explain speciation, George Romanes coined the term neo-Darwinism to refer to the version
of evolution advocated by Alfred Russel Wallace and August Weismann with its heavy dependence on
natural selection.[9] Weismann and Wallace rejected the Lamarckian idea of inheritance of acquired
characteristics, something that Darwin had not ruled out.[10]
Weismann's idea was that the relationship between the hereditary material, which he called the germ
plasm (de: Keimplasma), and the rest of the body (the soma) was a one-way relationship: the germ-plasm
formed the body, but the body did not influence the germ-plasm, except indirectly in its participation in a
population subject to natural selection. Weismann was translated into English, and though he was influential,
it took many years for the full significance of his work to be appreciated.[11] Later, after the completion of the
modern synthesis, the term neo-Darwinism came to be associated with its core concept: evolution, driven by
natural selection acting on variation produced by genetic mutation, and genetic recombination (chromosomal
crossovers).[9]
[edit] 1900–1915
Gregor Mendel's work was re-discovered by Hugo de Vries and Carl Correns in 1900. News of this
reached William Bateson in England, who reported on the paper during a presentation to the Royal
Horticultural Society in May 1900.[12] It showed that the contributions of each parent retained their integrity
rather than blending with the contribution of the other parent. This reinforced a division of thought, which was
already present in the 1890s.[13] The two schools were:
• Saltationism (large mutations or jumps), favored by early Mendelians who viewed hard
inheritance as incompatible with natural selection[14]
• Biometric school: led by Karl Pearson and Walter Weldon, argued vigorously against it,
saying that empirical evidence indicated that variation was continuous in most organisms, not
discrete as Mendelism predicted.
The relevance of Mendelism to evolution was unclear and hotly debated, especially by Bateson, who
opposed the biometric ideas of his former teacher Weldon. Many scientists believed the two theories
substantially contradicted each other.[15] This debate between the biometricians and the Mendelians
continued for some 20 years and was only solved by the development of population genetics.
T. H. Morgan began his career in genetics as a saltationist, and started out trying to demonstrate that
mutations could produce new species in fruit flies. However, the experimental work at his lab with Drosophila
melanogaster, which helped establish the link between Mendelian genetics and the chromosomal theory of
inheritance, demonstrated that rather than creating new species in a single step, mutations increased the
genetic variation in the population.[16]
[edit] The foundation of population genetics
The first step towards the synthesis was the development of population genetics. R.A. Fisher, J.B.S.
Haldane, and Sewall Wright provided critical contributions. In 1918, Fisher produced the paper "The
Correlation Between Relatives on the Supposition of Mendelian Inheritance",[17] which showed how the
continuous variation measured by the biometricians could be the result of the action of many discrete genetic
loci. In this and subsequent papers culminating in his 1930 book The Genetical Theory of Natural Selection ,
Fisher was able to show how Mendelian genetics was, contrary to the thinking of many early geneticists,
completely consistent with the idea of evolution driven by natural selection.[18] During the 1920s, a series of
papers by J.B.S. Haldane applied mathematical analysis to real world examples of natural selection such as
the evolution of industrial melanism in peppered moths.[18] Haldane established that natural selection could
work in the real world at a faster rate than even Fisher had assumed.[19]
Sewall Wright focused on combinations of genes that interacted as complexes, and the effects of
inbreeding on small relatively isolated populations, which could exhibit genetic drift. In a 1932 paper he
introduced the concept of an adaptive landscape in which phenomena such as cross breeding and genetic
drift in small populations could push them away from adaptive peaks, which would in turn allow natural
selection to push them towards new adaptive peaks.[18] Wright's model would appeal to field naturalists
such as Theodosius Dobzhansky and Ernst Mayr who were becoming aware of the importance of
geographical isolation in real world populations.[19] The work of Fisher, Haldane and Wright founded the
discipline of population genetics. This is the precursor of the modern synthesis, which is an even broader
coalition of ideas.[18][19][20]
[edit] The modern synthesis
Theodosius Dobzhansky, a Ukrainian emigrant, who had been a postdoctoral worker in Morgan's fruit
fly lab, was one of the first to apply genetics to natural populations. He worked mostly with Drosophila
pseudoobscura. He says pointedly: "Russia has a variety of climates from the Arctic to sub-tropical...
Exclusively laboratory workers who neither possess nor wish to have any knowledge of living beings in
nature were and are in a minority".[21] Not surprisingly, there were other Russian geneticists with similar
ideas, though for some time their work was known to only a few in the West. His 1937 work Genetics and the
Origin of Species was a key step in bridging the gap between population geneticists and field naturalists. It
presented the conclusions reached by Fisher, Haldane, and especially Wright in their highly mathematical
papers in a form that was easily accessible to others. It also emphasized that real world populations had far
more genetic variability than the early population geneticists had assumed in their models, and that
genetically distinct sub-populations were important. Dobzhansky argued that natural selection worked to
maintain genetic diversity as well as driving change. Dobzhansky had been influenced by his exposure in the
1920s to the work of a Russian geneticist named Sergei Chetverikov who had looked at the role of recessive
genes in maintaining a reservoir of genetic variability in a population before his work was shut down by the
rise of Lysenkoism in the Soviet Union.[18][19]
Edmund Brisco Ford's work complemented that of Dobzhansky. It was as a result of Ford's work, as
well as his own, that Dobzhansky changed the emphasis in the third edition of his famous text from drift to
selection.[22] Ford was an experimental naturalist who wanted to test natural selection in nature. He virtually
invented the field of research known as ecological genetics. His work on natural selection in wild populations
of butterflies and moths was the first to show that predictions made by R.A. Fisher were correct. He was the
first to describe and define genetic polymorphism, and to predict that human blood group polymorphisms
might be maintained in the population by providing some protection against disease.[23]
Ernst Mayr's key contribution to the synthesis was Systematics and the Origin of Species, published
in 1942. Mayr emphasized the importance of allopatric speciation, where geographically isolated sub-
populations diverge so far that reproductive isolation occurs. He was sceptical of the reality of sympatric
speciation believing that geographical isolation was a prerequisite for building up intrinsic (reproductive)
isolating mechanisms. Mayr also introduced the biological species concept that defined a species as a group
of interbreeding or potentially interbreeding populations that were reproductively isolated from all other
populations.[18][19][24] Before he left Germany for the United States in 1930, Mayr had been influenced by
the work of German biologist Bernhard Rensch. In the 1920s Rensch, who like Mayr did field work in
Indonesia, analyzed the geographic distribution of polytypic species and complexes of closely related
species paying particular attention to how variations between different populations correlated with local
environmental factors such as differences in climate. In 1947, Rensch published Neuere Probleme der
Abstammungslehre: die Transspezifische Evolution (English translation 1959: Evolution above the Species
level). This looked at how the same evolutionary mechanisms involved in speciation might be extended to
explain the origins of the differences between the higher level taxa. His writings contributed to the rapid
acceptance of the synthesis in Germany.[25][26]
George Gaylord Simpson was responsible for showing that the modern synthesis was compatible
with paleontology in his book Tempo and Mode in Evolution published in 1944. Simpson's work was crucial
because so many paleontologists had disagreed, in some cases vigorously, with the idea that natural
selection was the main mechanism of evolution. It showed that the trends of linear progression (in for
example the evolution of the horse) that earlier paleontologists had used as support for neo-Lamarckism and
orthogenesis did not hold up under careful examination. Instead the fossil record was consistent with the
irregular, branching, and non-directional pattern predicted by the modern synthesis.[18][19]
The botanist G. Ledyard Stebbins was another major contributor to the synthesis. His major work,
Variation and Evolution in Plants, was published in 1950. It extended the synthesis to encompass botany
including the important effects of hybridization and polyploidy in some kinds of plants.[18]
[edit] Further advances

The modern evolutionary synthesis continued to be developed and refined after the initial
establishment in the 1930s and 1940s. The work of W. D. Hamilton, George C. Williams, John Maynard
Smith and others led to the development of a gene-centric view of evolution in the 1960s. The synthesis as it
exists now has extended the scope of the Darwinian idea of natural selection to include subsequent scientific
discoveries and concepts unknown to Darwin, such as DNA and genetics, which allow rigorous, in many
cases mathematical, analyses of phenomena such as kin selection, altruism, and speciation.
A particular interpretation most commonly associated with Richard Dawkins, author of The Selfish
Gene, asserts that the gene is the only true unit of selection.[27] Dawkins further extended the Darwinian
idea to include non-biological systems exhibiting the same type of selective behavior of the 'fittest' such as
memes in culture. The synthesis continues to undergo regular review.[28] (See also Current research in
evolutionary biology).
[edit] After the synthesis
The structure of evolutionary biology.

The history and causes of evolution (center) are subject to various subdisciplines of evolutionary biology. The
areas of segments give an impression of the contributions of subdisciplines to the literature of evolutionary
biology.
There are a number of discoveries in earth sciences and biology which have arisen since the
synthesis. Listed here are some of those topics which are relevant to the evolutionary synthesis, and which
seem soundly based.
[edit] Understanding of Earth history

The Earth is the stage on which the evolutionary play is performed. Darwin studied evolution in the
context of Charles Lyell's geology, but our present understanding of Earth history includes some critical
advances made during the last half-century.
• The age of the Earth has been revised upwards. It is now estimated at 4.56 billion years,
about one-third of the age of the universe. It is worth noting that the Phanerozoic only occupies the
last 1/9th of this period of time.[29]
• The triumph of Alfred Wegener's idea of continental drift came around 1960. The key
principle of plate tectonics is that the lithosphere exists as separate and distinct tectonic plates, which
ride on the fluid-like (visco-elastic solid) asthenosphere. This discovery provides a unifying theory for
geology, linking phenomena such as volcanos, earthquakes, orogeny, and providing data for many
paleogeographical questions.[30] One major question is still unclear: when did plate tectonics begin?
[31]
• Our understanding of the evolution of the Earth's atmosphere has progressed. The
substitution of oxygen for carbon dioxide in the atmosphere, which occurred in the Proterozoic,
caused probably by cyanobacteria in the form of stromatolites, caused changes leading to the
evolution of aerobic organisms.[32][33]
• The identification of the first generally accepted fossils of microbial life was made by
geologists. These rocks have been dated as about 3.465 billion years ago.[34] Walcott was the first
geologist to identify pre-Cambrian fossil bacteria from microscopic examination of thin rock slices. He
also thought stromatolites were organic in origin. His ideas were not accepted at the time, but may
now be appreciated as great discoveries.[35]
• Information about paleoclimates is increasingly available, and being used in paleontology.
One example: the discovery of massive ice ages in the Proterozoic, following the great reduction of
CO2 in the atmosphere. These ice ages were immensely long, and led to a crash in microflora.[36]
See also Cryogenian period and Snowball Earth.
• Catastrophism and mass extinctions. A partial reintegration of catastrophism has occurred,
[37] and the importance of mass extinctions in large-scale evolution is now apparent. Extinction
events disturb relationships between many forms of life and may remove dominant forms and release
a flow of adaptive radiation amongst groups that remain. Causes include meteorite strikes (K–T
junction; Upper Devonian); flood basalt provinces (Deccan traps at K/T junction; Siberian traps at P–T
junction); and other less dramatic processes.[38][39]
Conclusion: Our present knowledge of earth history strongly suggests that large-scale geophysical
events influenced macroevolution and megaevolution. These terms refer to evolution above the species
level, including such events as mass extinctions, adaptive radiation, and the major transitions in evolution.
[40][41]
[edit] Symbiotic origin of eukaryotic cell structures

Further information: Endosymbiont and Endosymbiotic theory
Once symbiosis was discovered in lichen and in plant roots (rhizobia in root nodules) in the 19th
century, the idea arose that the process might have occurred more widely, and might be important in
evolution. Anton de Bary invented the concept of symbiosis;[42] several Russian biologists promoted the
idea;[43] Edwin Wilson mentioned it in his epic text The Cell;[44] as did Ivan Emmanuel Wallin in his
Symbionticism and the origin of species;[45] and there was a brief mention by Julian Huxley in 1930;[46] all
in vain because sufficient evidence was lacking. Symbiosis as a major evolutionary force was not discussed
at all in the evolutionary synthesis.[47]
The role of symbiosis in cell evolution was revived partly by Joshua Lederberg,[48] and finally
brought to light by Lynn Margulis in a series of papers and books.[49][50] It turns out that some cell
organelles are of microbial origin: mitochondria and chloroplasts definitely, cilia, flagella and centrioles
possibly, and perhaps the nuclear membrane and much of the chromosome structure as well. What is now
clear is that the evolution of eukaryote cells is either caused by, or at least profoundly influenced by,
symbiosis with bacterial and archaean cells in the Proterozoic.
The origin of the eukaryote cell by symbiosis in several stages was not part of the evolutionary
synthesis. It is, at least on first sight, an example of megaevolution by big jumps. However, what symbiosis
provided was a copious supply of heritable variation from microorganisms, which was fine-tuned over a long
period to produce the cell structure we see today. This part of the process is consistent with evolution by
natural selection.[51]
[edit] Trees of life

Further information: Last universal ancestor and Phylogenetic tree
The ability to analyse sequence in macromolecules (protein, DNA, RNA) provides evidence of
descent, and permits us to work out genealogical trees covering the whole of life, since now there are data on
every major group of living organisms. This project, begun in a tentative way in the 1960s, has become a
search for the universal tree or the universal ancestor, a phrase of Carl Woese.[52][53] The tree that results
has some unusual features, especially in its roots. There are two domains of prokaryotes: bacteria and
archaea, both of which contributed genetic material to the eukaryotes, mainly by means of symbiosis. Also,
since bacteria can pass genetic material to other bacteria, their relationships look more like a web than a
tree. Once eukaryotes were established, their sexual reproduction produced the traditional branching tree-
like pattern, the only diagram Darwin put in the Origin. The last universal ancestor (LUA) would be a
prokaryotic cell before the split between the bacteria and archaea. LUA is defined as most recent organism
from which all organisms now living on Earth descend (some 3.5 to 3.8 billion years ago, in the Archean era).
[54]
This technique may be used to clarify relationships within any group of related organisms. It is now a
standard procedure, and examples are published regularly. April 2009 sees the publication of a tree covering
all the animal phyla, derived from sequences from 150 genes in 77 taxa.[55]
Early attempts to identify relationships between major groups were made in the 19th century by Ernst
Haeckel, and by comparative anatomists such as Thomas Henry Huxley and E. Ray Lankester. Enthusiasm
waned: it was often difficult to find evidence to adjudicate between different opinions. Perhaps for that
reason, the evolutionary synthesis paid surprisingly little attention to this activity. It is certainly a lively field of
research today.
[edit] Evo-devo
Further information: Evolutionary developmental biology
What once was called embryology played a modest role in the evolutionary synthesis,[56] mostly
about evolution by changes in developmental timing (allometry and heterochrony).[57] Man himself was,
according to Bolk, a typical case of evolution by retention of juvenile characteristics (neoteny). He listed many
characters where "Man, in his bodily development, is a primate foetus that has become sexually mature".[58]
Unfortunately, his interpretation of these ideas was non-Darwinian, but his list of characters is both
interesting and convincing.[59]
Modern interest in Evo-devo springs from clear proof that development is closely controlled by
special genetic systems, and the hope that comparison of these systems will tell us much about the
evolutionary history of different groups.[60][61] In a series of experiments with the fruit-fly Drosophila, Edward
B. Lewis was able to identify a complex of genes whose proteins bind to the cis-regulatory regions of target
genes. The latter then activate or repress systems of cellular processes that accomplish the final
development of the organism.[62][63] Furthermore, the sequence of these control genes show co-linearity:
the order of the loci in the chromosome parallels the order in which the loci are expressed along the anterior-
posterior axis of the body. Not only that, but this cluster of master control genes programs the development of
all higher organisms.[64][65] Each of the genes contains a homeobox, a remarkably conserved DNA
sequence. This suggests the complex itself arose by gene duplication.[66][67][68] In his Nobel lecture, Lewis
said "Ultimately, comparisons of the [control complexes] throughout the animal kingdom should provide a
picture of how the organisms, as well as the [control genes] have evolved".
The term deep homology was coined to describe the common origin of genetic regulatory apparatus
used to build morphologically and phylogenetically disparate animal features.[69] It applies when a complex
genetic regulatory system is inherited from a common ancestor, as it is in the evolution of vertebrate and
invertebrate eyes. The phenomenon is implicated in many cases of parallel evolution.[70]
A great deal of evolution may take place by changes in the control of development. This may be
relevant to punctuated equilibrium theory, for in development a few changes to the control system could
make a significant difference to the adult organism. An example is the giant panda, whose place in the
Carnivora was long uncertain.[71] Apparently, the giant panda's evolution required the change of only a few
genetic messages (5 or 6 perhaps), yet the phenotypic and lifestyle change from a standard bear is
considerable.[72][73] The transition could therefore be effected relatively swiftly.
[edit] Fossil discoveries

In the past thirty or so years there have been excavations in parts of the world which had scarcely
been investigated before. Also, there is fresh appreciation of fossils discovered in the 19th century, but then
denied or deprecated: the classic example is the Ediacaran biota from the immediate pre-Cambrian, after the
Cryogenian period. These soft-bodied fossils are the first record of multicellular life. The interpretation of this
fauna is still in flux.
Many outstanding discoveries have been made, and some of these have implications for evolutionary
theory. The discovery of feathered dinosaurs and early birds from the Lower Cretaceous of Liaoning, N.E.
China have convinced most students that birds did evolve from coelurosaurian theropod dinosaurs. Less well
known, but perhaps of equal evolutionary significance, are the studies on early insect flight, on stem
tetrapods from the Upper Devonian,[74][75] and the early stages of whale evolution.[76]
Recent work has shed light on the evolution of flatfish (pleuronectiformes), such as plaice, sole,
turbot and halibut. Flatfish are interesting because they are one of the few vertebrate groups with external
asymmetry. Their young are perfectly symmetrical, but the head is remodelled during a metamorphosis,
which entails the migration of one eye to the other side, close to the other eye. Some species have both eyes
on the left (turbot), some on the right (halibut, sole); all living and fossil flatfish to date show an 'eyed' side
and a 'blind' side.[77] The lack of an intermediate condition in living and fossil flatfish species had led to
debate about the origin of such a striking adaptation. The case was considered by Lamark,[78] who thought
flatfish precursors would have lived in shallow water for a long period, and by Darwin, who predicted a
gradual migration of the eye, mirroring the metamorphosis of the living forms. Darwin's long-time critic St.
George Mivart thought that the intermediate stages could have no selective value,[79] and in the 6th edition
of the Origin, Darwin made a concession to the possibility of acquired traits.[80] Many years later the
geneticist Richard Goldschmidt put the case forward as an example of evolution by saltation, bypassing
intermediate forms.[81][82]
A recent examination of two fossil species from the Eocene has provided the first clear picture of
flatfish evolution. The discovery of stem flatfish with incomplete orbital migration refutes Goldschmidt's ideas,
and demonstrates that "the assembly of the flatfish bodyplan occurred in a gradual, stepwise fashion".[83]
There are no grounds for thinking that incomplete orbital migration was maladaptive, because stem forms
with this condition ranged over two geological stages, and are found in localities which also yield flatfish with
the full cranial asymmetry. The evolution of flatfish falls squarely within the evolutionary synthesis.[77]
[edit] See also

Book:Evolution

• Developmental systems theory
• Evolution
• Gene-centered view of evolution
• Particulate inheritance theory
• Polymorphism (biology)
• Population genetics
• The Origin of Species
[edit] Footnotes
1. ^ "Appendix: Frequently Asked Questions" (php). Science and Creationism: a view from the
National Academy of Sciences (Second ed.). Washington, DC: The National Academy of Sciences.
1999. p. 28. ISBN ISBN-0-309-06406-6. http://www.nap.edu/openbook.php?
record_id=6024&page=27#p200064869970027001. Retrieved September 24, 2009. "The scientific
consensus around evolution is overwhelming." Argumentum ad populum
2. ^ Mayr 2002, p. 270
3. ^ Huxley 2010
4. ^ Mayr & Provine 1998
5. ^ Mayr E. 1982. The growth of biological thought: diversity, evolution & inheritance . Harvard,
Cambs. p567 et seq.
6. ^ Smocovitis, V. Betty. 1996. Unifying Biology: the evolutionary synthesis and evolutionary
biology. Princeton University Press. p192
7. ^ Gould S.J. Ontogeny and phylogeny. Harvard 1977. p221-2
8. ^ Bowler P.J. 2003. Evolution: the history of an idea. pp236–256
9. ^ a b Gould The Structure of Evolutionary Theory p. 216
10.^ Kutschera U. 2003. A comparative analysis of the Darwin-Wallace papers and the
development of the concept of natural selection. Theory in Biosciences 122, 343-359
11.^ Bowler pp. 253–256
12.^ Mike Ambrose. "Mendel's Peas". Genetic Resources Unit, John Innes Centre, Norwich, UK.
http://www.jic.ac.uk/germplas/pisum/zgs4f.htm. Retrieved 2007-09-22.
13.^ Bateson, William 1894. Materials for the study of variation, treated with special regard to
discontinuity in the origin of species. The division of thought was between gradualists of the
Darwinian school, and saltationists such as Bateson. Mutations (as 'sports') and polymorphisms were
well known long before the Mendelian recovery.
14.^ Larson pp. 157–166
15.^ Grafen, Alan; Ridley, Mark (2006). Richard Dawkins: How A Scientist Changed the Way
We Think. New York, New York: Oxford University Press. p. 69. ISBN 0199291160.
16.^ Bowler pp. 271–272
17.^ Transactions of the Royal Society of Edinburgh, 52:399–433
18.^ a b c d e f g h Larson Evolution: The Remarkable History of a Scientific Theory pp. 221–243
19.^ a b c d e f Bowler Evolution: The history of an Idea pp. 325–339
20.^ Gould The Structure of Evolutionary Theory pp. 503–518
21.^ Mayr & Provine 1998 p. 231
22.^ Dobzhansky T. 1951. Genetics and the Origin of Species. 3rd ed, Columbia University
Press N.Y.
23.^ Ford E.B. 1964, 4th edn 1975. Ecological genetics. Chapman and Hall, London.
24.^ Mayr and Provine 1998 pp. 33–34
25.^ Smith, Charles H.. "Rensch, Bernhard (Carl Emmanuel) (Germany 1900–1990)". Western
Kentucky University. http://www.wku.edu/~smithch/chronob/RENS1900.htm. Retrieved 2007-09-22.
26.^ Mayr and Provine 1998 pp. 298–299, 416
27.^ Bowler p.361
28.^ Pigliucci, Massimo 2007. Do we need an extended evolutionary synthesis? Evolution 61
12, 2743–2749.
29.^ Dalrymple, G. Brent 2001. The age of the Earth in the twentieth century: a problem (mostly)
solved. Special Publications, Geological Society of London 190, 205–221.
30.^ Van Andel, Tjeerd 1994. New views on an old planet: a history of global change . 2nd ed.
Cambridge.
31.^ Witz A. 2006. The start of the world as we know it. Nature 442, p128.
32.^ Schopf J.W. and Klein (eds) 1992. The Proterozoic biosphere: a multi-disciplinary study.
Cambridge University Press.
33.^ Lane, Nick 2002. Oxygen: the molecule that made the world. Oxford.
34.^ Schopf J.W. 1999. Cradle of life: the discovery of Earth's earliest fossils. Princeton.
35.^ Yochelson, Ellis L. 1998. Charles Doolittle Walcott: paleontologist. Kent State, Ohio.
36.^ Knoll A.H. and Holland H.D. 1995. Oxygen and Proterozoic evolution: an update. In
National Research Council, Effects of past climates upon life. National Academy, Washington D.C.
37.^ Huggett, Richard J. 1997. Catastrophism. new ed. Verso.
38.^ Hallam A. and Wignall P.B. 1997. Mass extinctions and their aftermath. Columbia, N.Y.
39.^ Elewa A.M.T. (ed) 2008. Mass extinctions. Springer, Berlin.
40.^ The terms (or their equivalents) were used as part of the synthesis by Simpson G.G. 1944.
Tempo and mode in evolution, and Rensch B. 1947. Evolution above the species level. Columbia,
N.Y. They were also used by some non-Darwinian evolutionists such as Yuri Filipchenko and
Richard Goldschmidt. Here we use the terms as part of the evolutionary synthesis: they do not imply
any change in mechanism.
41.^ Maynard Smith J. and Szathmáry E. 1997. The major transitions in evolution. Oxford.
42.^ de Bary, H.A. 1879. Die Erscheinung der Symbiose. Strassburg.
43.^ Khakhina, Liya Nikolaevna 1992. Concepts of symbiogenesis: a historical and critical study
of the research of Russian scientists.
44.^ Wilson E.B. 1925. The cell in development and heredity . Macmillan, N.Y.
45.^ Wallin I.E. 1927. Symbionticism and the origin of species. Williams & Wilkins, Baltimore.
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[edit] References
• Allen, Garland. Thomas Hunt Morgan: The Man and His Science, Princeton University Press,
1978 ISBN 0-691-08200-6
• Bowler, Peter J. (2003). Evolution:The History of an Idea . University of California Press.
ISBN 0-52023693-9.
• Dawkins, Richard. The Blind Watchmaker, W.W. Norton and Company, Reissue Edition
1996 ISBN 0-393-31570-3
• Dobzhansky, T. Genetics and the Origin of Species, Columbia University Press, 1937 ISBN
0-231-05475-0
• Fisher, R. A. The Genetical Theory of Natural Selection, Clarendon Press, 1930 ISBN 0-19-
850440-3
• Futuyma, D.J. Evolutionary Biology, Sinauer Associates, 1986, p. 12 0-87-893189-9
• Gould, Stephen Jay (2002). The Structure of Evolutionary Theory. Belknap Press of Harvard
• Haldane, J. B. S. The Causes of Evolution, Longman, Green and Co., 1932; Princeton
University Press reprint, ISBN 0-691-02442-1
• Huxley, J. S., ed. The New Systematics, Oxford University Press, 1940 ISBN 0-403-01786-6
• Huxley, Julian S. (2010) [1942]. Evolution: the modern synthesis. The MIT Press. pp. 784.
ISBN 0262513668.
• Larson, Edward J. (2004). Evolution:The Remarkable History of a Scientific Theory . Modern
Library. ISBN 0-679-64288-9.
• Margulis, Lynn and Dorion Sagan. "Acquiring Genomes: A Theory of the Origins of Species",
Perseus Books Group, 2002 ISBN 0-465-04391-7
• Mayr, E. Systematics and the Origin of Species, Columbia University Press, 1942; Harvard
University Press reprint ISBN 0-674-86250-3
• Mayr, Ernst (2002). What evolution is. London: Weidenfeld & Nicolson. pp. 336.
ISBN 0753813688.
• Mayr, E. and W. B. Provine, eds. The Evolutionary Synthesis: Perspectives on the
Unification of Biology, Harvard University Press, 1998 ISBN 0-674-27225-0
• Simpson, G. G. Tempo and Mode in Evolution, Columbia University Press, 1944 ISBN 0-
231-05847-0
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Princeton University Press, 1996 ISBN 0-691-27226-9
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• Rose MR, Oakley TH, The new biology: beyond the Modern Synthesis. Biology Direct 2007,
2:30. A review of biology in light of recent innovations since the initiation of modern synthesis.
Processes of
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Evolutionary Canalisation · Inversion · Modularity · Phenotypic plasticity

developmental
biology (Evo-devo)
concepts

Modes of


Retrieved from "http://en.wikipedia.org/wiki/Modern_evolutionary_synthesis"
Categories: History of evolutionary biology | Synthesis | Biology theories

W000
Sperm whale

Sperm Whale[1]
Size compared to an average human
Conservation status
Vulnerable (IUCN 3.1)[2]

Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Cetacea
Suborder: Odontoceti
Family: Physeteridae
Genus: Physeter
Species: P. macrocephalus
Binomial name
Physeter macrocephalus
Linnaeus, 1758
Sperm whale range (in blue)
Synonyms
Physeter catodon Linnaeus, 1758

Physeter australasianus Desmoulins, 1822
The sperm whale, Physeter macrocephalus, is a marine mammal species, order Cetacea, a toothed
whale (odontocete) having the largest brain of any animal. The name comes from the milky-white waxy
substance, spermaceti, found in the animal's head. The sperm whale is the only living member of genus
Physeter. The synonym Physeter catodon refers to the same species. It is one of three extant species in the
sperm whale superfamily, along with the pygmy sperm whale and dwarf sperm whale.
A bull can grow to 20.5 metres (67 ft) long. It is the largest living toothed animal. The head can take
up to one-third of the animal's length. It has a cosmopolitan distribution across the oceans. The species feeds
on squid and fish, diving as deep as 3 kilometres (9,800 ft), which makes it the deepest diving mammal. Its
diet includes giant squid and colossal squid. The sperm whale's clicking vocalization is the loudest sound
produced by any animal, but its functions are uncertain. These whales live in groups called pods. Pods of
females and their young live separately from older males. The females cooperate to protect and nurse their
young. Females give birth every three to six years, and care for the calves for more than a decade.
Historically, the sperm whale was also known as the common cachalot; "cachalot" is derived from an
archaic French word for "tooth". Over most of the period from the early 18th century until the late 20th
century, the sperm whale was hunted to obtain spermaceti and other products, such as sperm oil and
ambergris. Spermaceti found many important uses, such as candles, soap, cosmetics and machine oil. Due
to its size, the sperm whale could sometimes defend itself effectively against whalers. In the most famous
example, a sperm whale attacked and sank the American whaleship Essex in 1820. As a result of whaling,
the sperm whale is currently listed as vulnerable by the IUCN. The sperm whale has few natural predators,
since few are strong enough to successfully attack a healthy adult; orcas attack pods and kill calves. The
sperm whale can live for more than 70 years.
Contents
[hide]
• 1 Name
• 2 Description
• 2.1 Size
• 2.2 Appearance
• 2.3 Jaws and teeth
• 2.4 Respiration and diving
• 2.5 Brain and senses
• 2.6 Spermaceti functions
• 3 Ecology, behaviour, and life history
• 3.1 Distribution
• 3.2 Reproduction
• 3.3 Social behavior
• 3.4 Feeding
• 4 Taxonomy and naming
• 5.1 Fossil record
• 5.2 Phylogeny
• 6 Relationship with humans
• 6.1 Historical hunting
• 6.2 Current conservation status
[edit] Name
The name sperm whale is an apocopation of spermaceti whale. Spermaceti is the semi-liquid, waxy
substance found in the spermaceti organ or case in front of and above the skull bone and also in the junk, the
area below the spermaceti organ and just above the upper jaw.[3] The case consists of a soft white, waxy
substance saturated with spermaceti oil. The junk is composed of cavities filled with the same wax and
spermaceti oil and intervening connective tissue.[3][4][5] The sperm whale is also known as the "cachalot",
which is thought to derive from the archaic French for "tooth" or "big teeth", as preserved for example in
cachau in the Gascon dialect (a word of either Romance[6] or Basque[7] origin). The etymological dictionary
of Corominas says the origin is uncertain, but it suggests that it comes from the vulgar Latin cappula, plural of
cappulum, sword hilt.[8] According to Encarta Dictionary, the word cachalot came to English "via French from
Spanish or Portuguese cachalote, perhaps from [Portuguese] cachola, 'big head'".
[edit] Description
[edit] Size
The Average sizes [9] Length Weight
sperm whale is
the largest 16 metres (52
Bull 41,000 kilograms (40 LT; 45 ST)
toothed whale, ft)
with adult males
measuring up to 11 metres (36
Cow 14,000 kilograms (14 LT; 15 ST)
20.5 metres (67 ft)
ft) long and
Newborn 4 metres (13 ft) 1,000 kilograms (0.98 LT; 1.1 ST)
weighing up to 57,000 kilograms (56 LT; 63 ST).[4][10] By contrast, the second largest toothed whale, Baird's
Beaked Whale measures 12.8 metres (42 ft) and weighs up to 15 short tons (14,000 kg).[11] The Nantucket
Whaling Museum has a 5.5 metres (18 ft)-long jawbone. The museum claims this individual was 80 feet (24
m) long; the whale that sank the Essex (one of the incidents behind Moby-Dick) was claimed to be 85 feet
(26 m).[12][13] Extensive whaling may have decreased their size, as males were highly sought, primarily
after World War II.[13] Today, males do not usually exceed 18.3 metres (60 ft) in length or 51,000 kilograms
(50 LT; 56 ST) in weight.[9]
It is among the most sexually dimorphic of all cetaceans. At birth both sexes are about the same size,
[9] but mature males are typically 30% to 50% longer and three times as massive.[4]
[edit] Appearance
The sperm whale's distinctive shape comes from its very large head, which is typically one-third of
the animal's length. The blowhole is located very close to the front of the head and shifted to the whale's left.
[4] This gives rise to a distinctive bushy, forward-angled spray.
The flukes of a sperm whale as it dives into the Gulf of Mexico (courtesy NMFS)
The sperm whale's flukes are triangular and very thick. The whale lifts its flukes high out of the water
as it begins a dive.[4] It has a series of ridges on the back's caudal third instead of a dorsal fin. The largest
ridge was called the 'hump' by whalers, and can be mistaken for a dorsal fin because of its shape.[9]
In contrast to the smooth skin of most large whales, its back skin is usually knobbly and has been
likened to a prune by whale-watching enthusiasts.[14] Skin is normally a uniform grey in color, though it may
appear brown in sunlight. Albinos have also been reported.[15][16][17]
[edit] Jaws and teeth
The sperm whale has 20 to 26 teeth on each side of its lower jaw.[4] The teeth are cone-shaped and
weigh up to 1 kilogram (2.2 lb).[18] The purpose of the teeth is unknown. Teeth do not appear to be
necessary for capturing or eating squid, and well-fed animals have been found without teeth. One hypothesis
is that the teeth are used in aggression between males.[19] Bulls often show scars which seem to be caused
by the teeth. Rudimentary teeth are also present in the upper jaw, but these rarely emerge into the mouth.
[20]
[edit] Respiration and diving

Sperm whales, along with bottlenose whales and elephant seals, are the deepest-diving mammals.
[4] Sperm whales are believed to be able to reach 3 kilometres (1.9 mi) and remain submerged for
90 minutes.[4][21] More typical dives are around 400 metres (1,300 ft) and 35 minutes in duration.[4] At these
great depths, sperm whales sometimes become entangled in transoceanic telephone cables and drown.[22]
Sperm whale arching back in preparation to dive off Dominica
The sperm whale has adapted to cope with drastic pressure changes when diving. The flexible
ribcage allows lung collapse, reducing nitrogen intake, and metabolism can decrease to conserve oxygen.
[23][24] Myoglobin, which stores oxygen in muscle tissue, is much more abundant than in terrestrial animals.
[25] The blood has a high red blood cell density, which contain oxygen-carrying hemoglobin. The oxygenated
blood can be directed towards the brain and other essential organs only when oxygen levels deplete.[26][27]
[28] The spermaceti organ may also play a role by adjusting buoyancy (see below).[29]
While sperm whales are well adapted to diving, repeated dives to great depths have long term
effects. Bones show pitting that signals decompression sickness in humans. Older skeletons showed the
most extensive pitting, whereas calves showed no damage. This damage may indicate that sperm whales
are susceptible to decompression sickness, and sudden surfacing could be lethal to them.[30]
Between dives, the sperm whale surfaces to breathe for about eight minutes before diving again.[4]
Odontoceti (toothed whales) breathe air at the
surface through a single, S-shaped blowhole.
Sperm whales spout (breathe) 3–5 times per minute
Blowhole
at rest, increasing to 6–7 times per minute after a
dive. The blow is a noisy, single stream that rises Phonic lips
up to 15 metres (49 ft) above the surface and points
forward and left at a 45° angle. On average, Dorsal
females and juveniles blow every 12.5 seconds Cranium
before dives, while large males blow every 17.5
seconds before dives.[31] Melon
Bony nares
[edit] Brain and senses
The brain is the largest known of any Upper mandible
modern or extinct animal, weighing on average Auditory bullae
about 8 kilograms (18 lb),[34][35] though the sperm
whale has a lower encephalization quotient than Lower mandible
many other whale and dolphin species, lower than Outgoing sound
that of non-human anthropoid apes and much lower
than humans'.[35][36] Incoming sound
Like other toothed whales (suborder
odontoceti), sperm whales use echolocation as one
means to find food because their habitat has
favorable acoustic characteristics and light
Echolocation system of a toothed whale[32][33]

absorption by water and suspended material limits visual range. The whale emits a focused wide angle beam
of high-frequency clicks. Passing air generates sounds from the bony nares through the phonic lips (also
known as "monkey lips"), a structure within the head.[32] The skull, melon and various air sacs in the whale's
head all play important roles in forming and focusing the beam of sound. The lower jaw is the primary echo
reception path. A continuous fat-filled canal transmits received sounds to the inner ear.[33]
[edit] Spermaceti functions

Main article: Spermaceti
The spermaceti organs may help adjust the whale's buoyancy. Before diving, cold water enters the
organ and it is likely that the blood vessels constrict, reducing blood flow and hence temperature. The wax
therefore solidifies.[29][37] The increase in specific density generates a down force of about 392 newtons
(860 lb) and allows the whale to dive with less effort. During the hunt, oxygen consumption, together with
blood vessel dilation, produces heat, and melts the spermaceti, increasing its buoyancy, enabling easy
surfacing.[38]
Herman Melville's Moby Dick suggests that the "case" containing the spermaceti evolved as a kind of
battering ram for use in fights between males.[39] This hypothesis is consistent with the well-documented
sinking of the ships Essex and Ann Alexander by attackers estimated to weigh only one-fifth as much as the
ships.[40]
The case may also aid echolocation (see melon).[5] The organ's variable shape narrows or spreads
the sound.[41]
The sperm whale has two nostrils. An external nostril forms the blow hole, and an internal nostril
presses against the bag-like spermaceti container.[5] The male's spermaceti organ is much larger than the
female's. This may be a case of sexual selection, enabling males to compete for females using sound
displays.[42]
[edit] Ecology, behaviour, and life history

[edit] Distribution
The sperm whale is among the most cosmopolitan species. It prefers ice-free waters over
1,000 metres (3,300 ft) deep.[2] Although both sexes range through temperate and tropical oceans and seas,
only adult males populate higher latitudes.[15]
It is relatively abundant from the poles to the equator, and is found in all the oceans. It inhabits the
Mediterranean Sea, but not the Black Sea,[9] while its presence in the Red Sea is uncertain.[2] The shallow
entrances to both the Black Sea and the Red Sea may account for their absence.[43] The Black Sea's lower
layers are also anoxic and contain high concentrations of sulphur compounds such as hydrogen sulphide.[44]
Populations are denser close to continental shelves and canyons.[15] Sperm whales are usually
found in deep off-shore waters, but may be seen closer to shore in areas where the continental shelf is small
and drops quickly to depths of 310–920 metres (1,020–3,020 ft).[9] Coastal areas with significant sperm whale
populations include the Azores and the Caribbean island of Dominica.[45]
[edit] Reproduction
Young sperm whale

Sperm whales can live 70 years or more.[9][15][46] They are a prime example of a species that has
been K-selected, a reproductive strategy associated with stable environmental conditions, a low birth rate,
significant parental aid to offspring, slow maturation and high longevity.[4]
How they choose mates has not been definitively determined. There is evidence that males have
dominance hierarchies and there is also evidence that female choice influences mating.[47] Gestation
requires 14 to 16 months, producing a single calf.[9] Lactation proceeds for 19 to 42 months, but calves may
suckle up to 13 years (although usually less).[9] Calves can suckle from females other than their mothers.[9]
Females generally have birth intervals of three to six years.[9]
Females reach sexual maturity between 7 and 13 years, males follow beginning at 18 years. Upon
reaching sexual maturity, males move to higher latitudes, where the water is colder and feeding is more
productive. Females remain at lower latitudes.[9] Males reach their full size at about age 50.[4]
[edit] Social behavior
Diagram of Marguerite formation

Females stay in groups of about a dozen individuals and their young.[4] Males leave these "nursery
pods" at somewhere between 4 and 21 years of age and join a "bachelor pod" with other males of similar age
and size.[4] As males grow older, they tend to disperse into smaller groups, and the oldest males typically
live solitary lives.[4] Mature males have beached themselves together, suggesting a degree of cooperation
which is not yet fully understood.[4]
The most common non-human attacker of sperm whales is the orca, but pilot whales and the false
killer whale also sometimes attack or harass them.[48][49] Orcas target groups of females with young,
usually trying to extract and kill a calf. Female sperm whales repel these attacks by encircling their calves.
The adults either face inwards to use their tail flukes against the orcas, or outwards, fighting with their teeth.
[4] This marguerite formation, named after the flower, is also used by whales to support an injured pod
member. Early whalers exploited this behaviour, attracting a whole pod by injuring one of its members.[50] If
the orca pod is extremely large, its members may sometimes be able to kill adult female sperm whales.
Large bull sperm whales have no non-human predators, and are believed to be too large and strong to be
threatened by orcas.[51]
[edit] Feeding
A piece of sperm whale skin with giant squid sucker scars

Sperm Whales usually dive between 300 to 800 metres (980 to 2,600 ft), and sometimes 1–2
kilometres (3,300–6,600 ft) to search for food.[52] Such dives can last more than an hour.[52] They feed on
several species, notably the giant squid, the colossal squid, octopuses, and diverse fish like demersal rays,
but the main part of their diet consists of medium-sized squid.[53] Some prey may be taken incidentally while
eating other items.[53] Most of what is known about deep sea squid has been learned from specimens in
captured sperm whale stomachs, although more recent studies analysed fecal matter. One study, carried out
around the Galápagos, found that squid from the genera Histioteuthis (62%), Ancistrocheirus (16%), and
Octopoteuthis (7%) weighing between 12 and 650 grams (0.026 and 1.4 lb) were the most commonly taken.
[54] Battles between sperm whales and colossal squid (which have been measured to weigh nearly
500 kilograms (1,100 lb)) have never been observed by humans; however white scars are believed to be
caused by the large squid. One study published in 2010 collected evidence that suggests that female sperm
whales may collaborate when hunting Humboldt squid.[55]
An older study, examining whales captured by the New Zealand whaling fleet in the Cook Strait
region, found a 1.69:1 ratio of squid to fish by weight.[56] Sperm whales sometimes steal Sablefish and
Toothfish from long lines. Long-line fishing operations in the Gulf of Alaska complain that sperm whales take
advantage of their fishing operations to eat desirable species straight off the line, sparing the whales the
need to hunt.[57] However, the amount of fish taken is very little compared to what the sperm whale needs
per day. New video footage has been captured of a large male sperm whale "bouncing" a long line, to gain
the fish.[58] Sperm whales are believed to prey on the megamouth shark, a rare and large deep-sea species
discovered in the 1970s.[59][60] In one case, three sperm whales were observed attacking or playing with a
megamouth.[61]
The sharp beak of a consumed squid lodged in the whale's intestine may lead to the production of
ambergris, analogous to the production of pearls.[62] The irritation of the intestines caused by squid beaks
stimulates the secretion of this lubricant-like substance. Sperm whales are prodigious feeders and eat around
3% of their body weight per day. The total annual consumption of prey by sperm whales worldwide is
estimated to be about 100,000,000 short tons (91,000,000 t) — a figure greater than the total consumption of
marine animals by humans each year.[63]
It is not well understood why the sperm whale's head is so large in comparison to the lower jaw. One
theory is that the sperm whale's ability to echolocate through its head aids in hunting. However squid, its
main prey, may have acoustic properties too similar to seawater to reflect sounds.[64] The sperm whale's
head contains a structure called monkey lips, through which it blows air. This can create clicks that have a
source level exceeding 230 decibels re 1 micropascal referenced to a distance of 1 metre (3.3 ft) – in other
words it is by far the loudest sound made by any animal, and 10–14 dB louder than a powerful rifle sounds in
air at 1 metre (3.3 ft) away.[65] It has been hypothesised that clicks attempt to stun prey. Experimental
studies attempting to duplicate this effect have been unable to replicate the supposed injuries, casting doubt
on this idea.[66]
[edit] Taxonomy and naming

The sperm whale belongs to the order Cetacea, the order containing all whales and dolphins. It is a
member of the suborder Odontoceti, the suborder containing all the toothed whales and dolphins. It is the
sole extant species of its genus, Physeter, in the family Physeteridae. Two species of the related extant
genus Kogia, the pygmy sperm whale Kogia breviceps and the dwarf sperm whale K. simus, are either
placed in this family, or in the family Kogiidae.[67] In some taxonomic schemes the families Kogiidae and
Physeteridae are combined as the superfamily Physeteroidea (see the separate entry on the sperm whale
family).[68]
The sperm whale is one of the species originally described by Linnaeus in 1758 in his 18th century
work, Systema Naturae. He recognised four species in the genus Physeter.[69] Experts soon realised that
just one such species exists, although there has been debate about whether this should be named P.
catodon or P. macrocephalus, two of the names used by Linnaeus. Both names are still used, although most
recent authors now accept macrocephalus as the valid name, limiting catodon's status to a lesser synonym.
[a]

See also: Sperm whale family
[edit] Fossil record

Although the fossil record is poor,[70] several extinct genera have been assigned to the clade
Physeteroidea, which includes the last common ancestor of the modern sperm whale, pygmy sperm whale
and dwarf sperm whale, plus all of that ancestor's descendants. These fossils include Ferecetotherium,
Idiorophus, Diaphorocetus, Aulophyseter, Orycterocetus, Scaldicetus, Placoziphius, Zygophyseter and
Acrophyseter.[64][68][71] Ferecetotherium, found in Azerbaijan and dated to the late Oligocene (about
28 to 23 million years ago), is the most primitive fossil that has been found which possesses sperm whale-
specific features such as an asymmetric rostrum ("beak" or "snout").[72] Most sperm whale fossils date from
the Miocene period, 23 to 5 million years ago. Diaphorocetus, from Argentina, has been dated to the early
Miocene. Fossil sperm whales from the Middle Miocene include Aulophyseter, Idiorophus and Orycterocetus,
all of which were found on the west coast of the United States, and Scaldicetus, found in Europe and Japan.
[72][73] Orycterocetus fossils have also been found in the North Atlantic Ocean and the Mediterranean Sea,
in addition to the west coast of the United States.[74] Placoziphius, found in Europe, and Acrophyseter, from
Peru, are dated to the late Miocene.[68][72]
Cetacea
Toothed whales
Physeteroidea
Other
Physeteroidea†
Kogiidae
Pygmy sperm
whale
Dwarf sperm whale
Physeteridae
Other
Physeteridae†
Sperm whale
Ganges River
dolphin
Other river
dolphins
Oceanic
dolphins
Porpoises
Arctic whales
Beaked
whales
Baleen
whales
Evolutionary family tree of sperm whales,[75]

including simplified summary of extinct groups (†)[64]
Fossil sperm whales differ from modern sperm whales in tooth count and the shape of the face and
jaws.[72] For example Scaldicetus had a tapered rostrum.[73] Genera from the Oligocene and early and
middle Miocene, with the possible exception of Aulophyseter, had teeth in their upper jaws.[72] Acrophyseter,
from the late Miocene, also had teeth in both the upper and lower jaws as well as a short rostrum and an
upward curving mandible (lower jaw).[68] These anatomical differences suggest that fossil species may not
have necessarily been deep-sea squid eaters like the modern sperm whale, but that some genera mainly ate
fish.[72] Zygophyseter, dated from the middle to late Miocene and found in southern Italy, had teeth in both
jaws and appears to have been adapted to feed on large prey, rather like the modern Orca (Killer Whale).[64]
[edit] Phylogeny
The traditional view has been that Mysticeti (baleen whales) and Odontoceti (toothed whales) arose
from more primitive whales early in the Oligocene period, and that the super-family Physeteroidea, which
contains the sperm whale, dwarf sperm whale and pygmy sperm whale, diverged from other toothed whales
soon after that, over 23 million years ago.[70][72] In 1993–1996 molecular phylogenetics analyses by
Milinkovitch and colleagues, based on comparing the genes of various modern whales, suggested that the
sperm whales are more closely related to the baleen whales than they are to other toothed whales, which
would have meant that Odontoceti were not monophyletic, in other words did not consist of a single ancestral
toothed whale species and all its descendants.[75] However more recent studies, based on various
combinations of comparative anatomy and molecular phylogenetics, criticised Milinkovitch's analysis on
technical grounds and re-affirmed that the Odontoceti are monophyletic.[75][76][77]
These analyses also confirm that there was a rapid evolutionary radiation (diversification) of the
Physeteroidea in the Miocene period.[64] The Kogiidae (dwarf and pygmy sperm whales) diverged from the
Physeteridae (true sperm whales) at least 8 million years ago.[76]
[edit] Relationship with humans
[edit] Historical hunting
See also: Whaling and Sperm whaling
Spermaceti, obtained primarily from the spermaceti organ, and sperm oil, obtained primarily from the
blubber in the body, were much sought after by 18th, 19th and 20th century whalers. These substances
found a variety of commercial applications, such as candles, soap, cosmetics, machine oil, other specialized
lubricants, lamp oil, pencils, crayons, leather waterproofing, rust-proofing materials and many pharmaceutical
compounds.[78][79][80][81] Ambergris, a solid, waxy, flammable substance produced in the digestive system
of sperm whales, was also sought as a fixative in perfumery.
Sperm whaling
Prior to the early 18th century, hunting was mostly by indigenous Indonesians.[82] Legend has it that
sometime in the early 18th century, around 1712, Captain Christopher Hussey, while cruising for Right
Whales near shore, was blown offshore by a northerly wind, where he encountered a Sperm whale pod and
killed one.[83] Although the story may not be true, sperm whales were indeed soon exploited by American
whalers. Judge Paul Dudley, in his Essay upon the Natural History of Whales (1725), states that one Atkins,
ten or twelve years in the trade, was among the first to catch sperm whales sometime around 1720 off the
New England coast.[84]
There were only a few recorded catches during the first few decades (1709-1730s) of offshore sperm
whaling. Instead sloops concentrated on Nantucket Shoals where they would have taken Right Whales or
went to the Davis Strait region to catch Bowhead Whales. By the early 1740s, with the advent of spermaceti
candles (before 1743), American vessels began to focus on sperm whales. The diary of Benjamin Bangs
(1721–1769) shows that, along with the bumpkin sloop he sailed, he found three other sloops flensing sperm
whales off the coast of North Carolina in late May 1743.[85] On returning to Nantucket in the summer 1744
on a subsequent voyage he noted that "45 spermacetes are brought in here this day," another indication that
American sperm whaling was in full swing.[85]
American sperm whaling soon spread from the east coast of the American colonies to the Gulf
Stream, the Grand Banks, West Africa (1763), the Azores (1765) and the South Atlantic (1770s). From 1770
to 1775 Massachusetts, New York, Connecticut, and Rhode Island ports produced 45,000 barrels of sperm
oil annually, compared to 8,500 of whale oil.[86] In the same decade the British began sperm whaling,
employing American ships and personnel.[87] By the following decade the French had entered the trade,
also employing American expertise.[87] Sperm whaling increased until the mid-19th century. Spermaceti oil
was important in public lighting (for example, in lighthouses, where it was used in the United States until
1862, when it was replaced by lard oil, in turn replaced by petroleum) and for lubricating the machines (such
as those used in cotton mills) of the Industrial Revolution. Sperm whaling declined in the second half of the
19th century, as petroleum came in to broader use. In that sense, it may be said to have protected whale
populations from even greater exploitation.[88][89]
Nantucket, in red, is an island off the state of Massachusetts where much sperm whaling originated
Sperm whaling in the 18th century began with small sloops carrying only one or two whaleboats. The
fleet's scope and size increased over time, and larger ships entered the fishery. In the late 18th century and
early 19th century sperm whaling ships sailed to the Pacific, the Indian Ocean, Japan, the coast of Arabia,
Australia and New Zealand.[87][90][91] Hunting could be dangerous to the crew. For example, on November
20, 1820, a sperm whale claimed to be about 25.9 metres (85 ft) long rammed and sank the Nantucket
whaleship Essex. Only 8 out of 21 sailors survived to be rescued by other ships.[92]
Whaling activity declined from the 1880s until 1946, but picked up again after World War II. Modern
whaling was more efficient than open-boat whaling, employing steam-powered ships and exploding
harpoons. Initially, modern whaling activity focused on large baleen whales, but as these populations were
taken, sperm whaling increased. Cosmetics, soap and machine oil were the major buyers. After populations
declined significantly, the International Whaling Commission gave the species full protection in 1985. Hunting
by Japan in the northern Pacific Ocean continued until 1988.[89]
It is estimated that the historic worldwide population numbered 1,100,000 before commercial sperm
whaling began in the early 18th century.[2] By 1880 it had declined an estimated 29 per cent.[2] From that
date until 1946 the population appears to have recovered somewhat as whaling pressure lessened, but after
the Second World War, the population declined even further, to only 33 per cent of the pre-whaling era.[2] It
has been estimated that in the 19th century between 184,000 and 236,000 sperm whales were killed by the
various whaling nations,[93] while in the modern era, at least 770,000 were taken, the majority between 1946
and 1980.[94]
Sperm whales increase the levels of primary production and carbon export by defecating iron rich
faeces into surface waters of the Southern Ocean. The iron rich faeces causes phytoplankton to grow and
take up more carbon from the atmosphere. When the phytoplankton dies, it sinks to the deep ocean and
takes the atmospheric carbon with it. By reducing the abundance of sperm whales in the Southern Ocean,
whaling has resulted in an extra 2 million tonnes of carbon remaining in the atmosphere each year.[95]
Remaining sperm whale populations are large enough that the species' conservation status is rated
as vulnerable rather than endangered.[2] However, the recovery from the whaling years is a slow process,
particularly in the South Pacific, where the toll on breeding-age males was severe.[96]
[edit] Current conservation status

The number of sperm whales throughout the world is unknown, but is thought to be in the hundreds
of thousands.[2] The conservation outlook is brighter than for many other whales. Historically, Japan has
taken ten sperm whales a year, and until 2006 tens of these whales were hunted off Indonesia. They are
protected practically worldwide, and commercial whaling has ceased.[2] Fishermen do not target the
creatures that sperm whales eat.[2] However, long-line fishing operations in the Gulf of Alaska have
complained about sperm whales stealing fish from their lines.[57]
Entanglement in fishing nets and collisions with ships represent the greatest threats to the sperm
whale population currently.[15] Other current threats include ingestion of marine debris, ocean noise, and
chemical pollution.[97] The IUCN regards the sperm whale as being "vulnerable".[2] The species is listed as
endangered on the United States Endangered Species Act.[98]
[edit] Cultural importance

Rope-mounted teeth are important cultural objects throughout the Pacific. In New Zealand, the Māori
know them as "rei puta" and were rare because sperm whales were not actively hunted in traditional Māori
society.[99] Whale ivory and bone were taken from beached whales. In Fiji the teeth are known as tabua and
they were traditionally given as gifts for atonement or esteem (called sevusevu), and were important in
negotiations between rival chiefs.[100] Friedrich Ratzel in The History of Mankind reported in 1896 that, in
Fiji, whales' or cachalots' teeth were the most-demanded article of ornament or value. They occurred often in
necklaces.[101] Today the tabua remains an important item in Fijian life. The teeth were originally rare in Fiji
and Tonga, which exported teeth, but with the Europeans' arrival, teeth flooded the market and this
"currency" collapsed. The oversupply led in turn to the development of the European art of scrimshaw.[102]
Herman Melville's novel Moby-Dick is based on a true story about a sperm whale that attacked the
whaleship Essex.[103][104] Melville associated the sperm whale with the Bible's Leviathan.[104][105] The
fearsome reputation perpetuated by Melville was based on bull whales' ability to fiercely defend themselves
from attacks by early whalers, occasionally resulting in the destruction of the whaling ships.
Jules Verne's Twenty Thousand Leagues Under the Sea, mentions cachalots (perhaps incorrectly)
as preying on fellow whales.
The Sperm Whale was designated as the Connecticut state animal by the CT General Assembly in
1975. It was selected because of its specific contribution to the state's history and because of its present-day
plight as an endangered species.[106]
Female in Dominican Pod, 2005
[edit] Watching sperm whales

See also: Whale watching
Sperm whales are not the easiest of whales to watch, due to their long dive times and ability to travel
long distances underwater. However, due to the distinctive look and large size of the whale, watching is
increasingly popular. Sperm whale watchers often use hydrophones to listen to the clicks of the whales and
locate them before they surface. Popular locations for sperm whale watching include the picturesque
Kaikoura on New Zealand's South Island, where the continental shelf is so narrow that whales can be
observed from the shore, Andenes and Tromsø in Arctic Norway and at the Azores where it can be seen
throughout the year.[45][107] Dominica is believed to be the only Caribbean island with a year-round
residential pod of females and calves.[108]
[edit] See also

Cetaceans portal
Animals portal
• High-finned sperm whale

• Colossal Squid
• Giant squid
• Toothed Whale
• Whale
[edit] Notes
[edit] Footnotes
• a Until 1974 the species was generally known as P. catodon, however in that year Husson &
Holthuis proposed that the correct name should be P. macrocephalus, the second name in the genus
Physeter published by Linnaeus concurrently with P. catodon, on the grounds that the names were
synonyms published simultaneously and therefore the ICZN principle of "First Reviser" should apply,
in this instance leading to the choice of P. macrocephalus over P. catodon, a view re-stated in
Holthuis, 1987. This has been adopted by most subsequent authors, although Schevill (1986 and
1987) argued that macrocephalus was published with an inaccurate description and that therefore
only the species catodon was valid, rendering the principle of "First Reviser" inapplicable. At the
present time, the name P. catodon is used in the Catalogue of Life, however this is expected to be
changed to follow the most recent version of ITIS which has recently altered its usage from P.
catodon to P. macrocephalus following L. B. Holthuis, and recent (2008) discussions with relevant
experts (refer cited ITIS page for additional information).[4][109][110][111][112][113]
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73.^ a b Hirota, K. & Barnes, L. G. (April 5, 2006). "A new species of Middle Miocene sperm
whale of the genus Scaldicetus (Cetacea; Physeteridae) from Shiga-mura, Japan". Island Arc 3 (4):
453–472. doi:10.1111/j.1440-1738.1994.tb00125.x.
74.^ Bianucci, G., Landrini, W. & Varola, W. (September–October 2004). "First discovery of the
Miocene northern Atlantic sperm whale Orycterocetus in the Mediterranean". Geobios 37 (5): 569–
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75.^ a b c Nikaido, M., Matsuno, F., Hamilton, H., Brownwell, R., Cao, Y., Ding, W., Zuoyan, Z.,
Shedlock, A., Fordyce, R. E., Hasegawa, M. & Okada, N. (June 19, 2001). "Retroposon analysis of
major cetacean lineages: The monophyly of toothed whales and the paraphyly of river dolphins".
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76.^ a b Whitehead, H. (2003). "Evolutionary History of the Sperm Whale". Sperm Whales Social
Evolution in the Ocean. University of Chicago Press. pp. 2–3. ISBN 0-226-89518-1.
77.^ Heyning, J. (August 23, 2006). "Sperm Whale Phylogeny Revisited: Analysis of the
Morphological Evidence". Marine Mammal Science 13 (4): 596–613. doi:10.1111/j.1748-
7692.1997.tb00086.x.
78.^ Wilson, D. (1999). The Smithsonian Book of North American Mammals . Vancouver: UBC
Press. p. 300. ISBN 0774807628.
79.^ The Southampton Oceanography Centre & A deFontaubert. "The status of natural
resources on the high seas". IUCN. p. 63. http://cmsdata.iucn.org/downloads/highseas.pdf. Retrieved
2008-10-11.
80.^ Jamieson, A. (1829). A Dictionary of Mechanical Science, Arts, Manufactures, and
Miscellaneous Knowledge. H. Fisher, Son & Co.. p. 566.
81.^ "Aquarium of the Pacific - Sperm Whale".
http://www.aquariumofpacific.org/onlinelearningcenter/print/sperm_whale/. Retrieved 2008-10-11.
82.^ Whitehead, H. (2003). "Sperm Whales and Humans". Sperm Whales Social Evolution in
the Ocean. University of Chicago Press. p. 14. ISBN 0-226-89518-1.
83.^ Simons, B.. "Christopher Hussey Blown Out (Up) to Sea". Nantucket Historical Association.
http://www.nha.org/history/hn/HNsimons-hussey.htm.
84.^ Dudley, P. (1725). "An Essay upon the Natural History of Whales, with a Particular Account
of the Ambergris Found in the Sperma Ceti Whale". Philosophical Transactions (1683-1775), Vol. 33.
The Royal Society. p. 267. http://www.capecodhistory.us/18th/Dudley-whales-1724.htm.
85.^ a b Dolin, E. (2007). Leviathan: The History of Whaling in America . W. W. Norton. pp. 98–
100. ISBN 0393060578.
86.^ Starbuck, A. (1878). History of the American Whale Fishery from its Earliest Inception to
the Year 1876. ISBN 066535343X. http://mysite.du.edu/~ttyler/ploughboy/starbuck.htm#sectiond.
87.^ a b c Bockstoce, J. (December 1984). "From Davis Strait to Bering Strait: The Arrival of the
Commercial Whaling Fleet in North America's West Arctic". Arctic 37 (4): 528–532.
http://pubs.aina.ucalgary.ca/arctic/Arctic37-4-528.pdf.
88.^ Estes, J. (2006). Whales, Whaling, and Ocean Ecosystems. University of California Press.
p. 329. ISBN 0520248848.
89.^ a b Whitehead, H. (2003). "Sperm whales and humans". Sperm Whales Social Evolution in
90.^ Stackpole, E. A. (1972). Whales & Destiny: The Rivalry between America, France, and
Britain for Control of the Southern Whale Fishery, 1785-1825 . The University of Massachusetts
Press. ISBN 0870231049.
91.^ Baldwin, R., Gallagher, M., and van Waerebeek, K.. "A Review of Cetaceans from Waters
off the Arabian Peninsula". p. 6. http://www.whalecoastoman.com/ArabPeninsula.pdf. Retrieved
2008-10-15.
92.^ "The Wreck of the Whaleship Essex". BBC.
http://www.bbc.co.uk/dna/h2g2/classic/A671492. Retrieved 2008-10-11.
93.^ Davis, L, Gallman, R. & Gleiter, K. (1997). In Pursuit of Leviathan: Technology, Institutions,
Productivity, and Profits in American Whaling, 1816-1906 (National Bureau of Economic Research
Series on Long-Term Factors in Economic Dev). University of Chicago Press. p. 135.
ISBN 0226137899.
94.^ Over 680,000 officially reported at "Whaling Statistics". http://luna.pos.to/whale/sta.html.
Retrieved 2008-10-15. . In addition, studies have found that official reports understated USSR
catches by at least 89,000 "Sperm Whale (Physeter macrocephalus) California/Oregon/Washington
Stock". http://www.nmfs.noaa.gov/pr/pdfs/sars/po2007whsp-cow.pdf. Retrieved 2008-10-16. .
Furthermore, other countries, such as Japan have been found to have understated catches "The
RMS - A Question of Confidence: Manipulations and Falsifications in Whaling".
http://www.wdcs.org/submissions_bin/rmsreview.pdf. Retrieved 2008-10-16.
95.^ Lavery, Trish L., Ben Roudnew, Peter Gill, Justin Seymour, Laurent Seuront, Genevieve
Johnson, James G. Mitchell & Victor Smetacek (2010). "Iron defecation by sperm whales stimulates
carbon export in the Southern Ocean". Proceedings of the Royal Society B 277 (1699): 3527–3531.
doi:10.1098/rspb.2010.0863. PMID 20554546.
96.^ Whitehead, H. (2003). "Ghosts of Whaling Past". Sperm Whales Social Evolution in the
Ocean. University of Chicago Press. pp. 360–362. ISBN 0-226-89518-1.
97.^ Whitehead, H. (2003). "New Threats to Sperm Whales". Sperm Whales Social Evolution in
98.^ Species Profile: "Sperm whale (Physeter catodon)" US Fish and Wildlife Service. Accessed
November 16, 2010.
99.^ "Museum of New Zealand Te Papa Tongarewa Collections Online Search - Rei puta".
http://collections.tepapa.govt.nz/search.aspx?term=Rei%20puta. Retrieved 2009-03-15.
100.^ Arno, A. (2005). "Cobo and tabua in Fiji: Two forms of cultural currency in an economy of
sentiment". American Ethnologist 32 (1): 46–62. doi:10.1525/ae.2005.32.1.46. http://cat.inist.fr/?
aModele=afficheN&cpsidt=16581746. Retrieved 2008-10-06.
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Mankind. (London: MacMillan, 1896). Accessed 21 October 2009.
102.^ Constantine, R. (2002). "Folklore and Legends". In Perrin, W., Würsig, B. & Thewissen, J..
Encyclopedia of Marine Mammals. Academic Press. p. 449. ISBN 0-12-551340-2.
103.^ "Chapter 3. Romances of Adventure. Section 2. Herman Melville. Van Doren, Carl. 1921.
The American Novel". Bartleby.com. http://www.bartleby.com/187/5.html. Retrieved 2008-10-19.
104.^ a b Zwart, H. (2000). What is a Whale? Moby Dick, marine science and the sublime .
Tubingen Attempo. 185–214. http://www.filosofie.science.ru.nl/research/hra/whale.pdf.
105.^ Edwards, B.. "The Playful Learnings". Australasian Journal of American Studies: 9.
http://www.anzasa.arts.usyd.edu.au/a.j.a.s/Articles/1_06/EdwardsArticle.pdf.
106.^ "The State Animal", State of Connecticut Sites, Seals and Symbols (Reproduced from the
Connecticut State Register & Manual: State of Connecticut),
http://vvv.state.ct.us/emblems/animal.htm, retrieved December 26 2010
107.^ "Whale and dolphin watching in the Azores". Wildlife Extra.
http://www.wildlifeextra.com/go/whales/azores/. Retrieved 2008-09-26.
108.^ "Whale Watching Dominica". http://www.dominica.dm/site/whalewatching.cfm. Retrieved
2008-09-26.
109.^ Husson, A.M. & Holthuis, L.B. (1974). Physeter macrocephalus Linnaeus, 1758, the valid
name for the sperm whale. Zoologische Mededelingen 48: 205-217.
110.^ Holthuis, L. B. (1987). The scientific name of the sperm whale. Marine Mammal Science
3(1): 87-89.
111.^ Schevill, W.E. (1986). The International Code of Zoological Nomenclature and a
paradigm: the name Physeter catodon Linnaeus 1758. Marine Mammal Science 2(2): 153-157.
112.^ Schevill, W.E. (1987). Reply to L. B. Holthuis "The scientific name of the sperm whale",
Marine Mammal Science 3(1): 89-90.
113.^ Whitehead, H. (2003). Sperm Whales Social Evolution in the Ocean. University of
Chicago Press. p. 3. ISBN 0-226-89518-1.
[edit] References
• Carwardine, Hoyt, Fordyce & Gill (1998). Whales & Dolphins: The Ultimate Guide to Marine
Mammals. London: HarperCollins. ISBN 0-00-220105-4.
• Randall R. Reeves ... et al. (2002). Guide to marine mammals of the world / National
Audubon Society. New York: A.A. Knopf: Distributed by Random House. ISBN 0-375-41141-0.
• William F. Perrin, Bernd Würsig, J.G.M. Thewissen (Eds.) (2002). Encyclopedia of Marine
Mammals. San Diego, Calif.: Academic Press. ISBN 0-12-551340-2.
• Spermaceti in candles July 22, 2007
• Retroposon analysis of major cetacean lineages: The monophyly of toothed whales and the
paraphyly of river dolphins June 19, 2001

Wikimedia Commons has media related to: Physeter macrocephalus
Wikispecies has information related to: Physeter macrocephalus
• US National Marine Fisheries Service Sperm Whale web page

• 70South—information on the sperm whale
• MarineBio: Sperm Whale, Physeter catodon
• "Physty"-stranded sperm whale nursed back to health and released in 1981
• The physiology of the deep diving adaptations of whales
• ARKive—Photographs, video.
• Whale Trackers—An online documentary film exploring the sperm whales in the
Mediterranean Sea.
• Prof Malcolm Clarke discusses anatomy of sperm whales —Clarke has spent a lifetime
studying sperm whales and giant squid.
[show]v · d · eExtant Cetacea species

Laurasiatheria · (unranked) Cetartiodactyla · (unranked) Whippomorpha
[show] Suborder Mysticeti (Baleen whales)
Balaena Bowhead whale (B. mysticetus)
Balaenidae Southern right whale (E. australis) · North Atlantic

Eubalaena
right whale (E. glacialis) · North Pacific right whale (E.
(Right whales)
japonica)
Balaenopteridae Common minke whale (B. acutorostrata) ·

(Rorquals) Balaenoptera Antarctic minke whale (B. bonaerensis) · Sei whale (B.
borealis) · Bryde's whale (B. brydei) · Pygmy Bryde's
whale (B. edeni) · Blue whale (B. musculus) · B. omurai ·
Fin whale (B. physalus)
Megaptera Humpback whale (M. novaeangliae)
Eschrichtiidae Gray whale (E.

Eschrichtius
robustus)
Neobalaenidae Pygmy right whale (C.

Caperea
marginata)
[show] Suborder Odontoceti (Toothed whales) (cont. below)
Commerson's dolphin (C. commersonii) ·

Cephalorhynchus Chilean dolphin (C. eutropia) · Haviside's dolphin
(C. heavisidii) · Hector's dolphin (C. hectori)
Delphinidae
(Oceanic dolphins) Long-beaked common dolphin (D.
Delphinus capensis) · Short-beaked common dolphin (D.
delphis)
Feresa Pygmy killer whale (F. attenuata)
Short-finned pilot whale (G.

Globicephala
macrorhynchus) · Long-finned pilot whale (G.
(Pilot whales)
melas)
Grampus Risso's dolphin (G. griseus)
Lagenodelphis Fraser's dolphin (L. hosei)
Atlantic white-sided dolphin (L. acutus) ·

White-beaked dolphin (L. albirostris) · Peale's
Lagenorhynchus dolphin (L. australis) · Hourglass dolphin (L.
cruciger) · Pacific white-sided dolphin (L.
obliquidens) · Dusky dolphin (L. obscurus)
Lissodelphis Northern right whale dolphin (L. borealis) ·

(Right whale dolphins) Southern right whale dolphin (L. peronii)
Irrawaddy dolphin (O. brevirostris) ·

Orcaella
Australian snubfin dolphin (O. heinsohni)
Orcinus Killer whale (O. orca)
Peponocephala Melon-headed whale (P. electra)
Pseudorca False killer whale (P. crassidens)
Tucuxi (S. fluviatilis) · Costero (S.

Sotalia
guianensis)
Pacific humpback dolphin (S. chinensis) ·

Sousa Indian humpback dolphin (S. plumbea) · Atlantic
humpback dolphin (S. teuszii)
Pantropical spotted dolphin (S. attenuata) ·

Clymene dolphin (S. clymene) · Striped dolphin (S.
Stenella
coeruleoalba) · Atlantic spotted dolphin (S.
frontalis) · Spinner dolphin (S. longirostris)
Steno Rough-toothed dolphin (S. bredanensis)
Indo-Pacific bottlenose dolphin (T.

Tursiops
aduncus) · Common bottlenose dolphin (T.
truncatus)
[hide] Suborder Odontoceti (Toothed whales) (cont. above)
Delphinapterus Beluga (D. leucas)

Monodontidae
Monodon Narwhal (M. monoceros)
Neophocaena Finless porpoise (N. phocaeniodes)
Phocoenidae Spectacled porpoise (P. dioptrica) · Harbor

(Porpoises) Phocoena porpoise (P. phocoena) · Vaquita (P. sinus) · Burmeister's
porpoise (P. spinipinnis)
Phocoenoides Dall's porpoise (P. dalli)
Physeteridae Physeter Sperm whale (P. macrocephalus)
Kogiidae Pygmy sperm whale (K. breviceps) · Dwarf sperm whale

Kogia
(K. simus)
Iniidae Amazon river dolphin (I.
Inia
geoffrensis)
Lipotidae Baiji (L.

Lipotes
vexillifer)
Platanistidae Ganges and Indus River dolphin (P.

Platanista
gangetica)
Pontoporiidae La Plata dolphin (P.

Pontoporia
blainvillei)
Arnoux's beaked shale (B. arnuxii) · Baird's beaked

Berardius
whale (B. bairdii)
Ziphiidae Northern bottlenose whale (H. ampullatus) ·

(Beaked whales) Hyperoodon
Southern bottlenose whale (H. planifrons)
Indopacetus Tropical bottlenose whale (I. pacificus)

Sowerby's beaked whale (M. bidens) · Andrew's
beaked whale (M. bowdoini) · Hubbs' beaked whale (M.
carlhubbsi) · Blainville's beaked whale (M. densirostris) ·
Gervais' beaked whale (M. europaeus) · Ginkgo-toothed
Mesoplodon
beaked whale (M. ginkgodens) · Gray's beaked whale (M.
(Mesoplodont
grayi) · Hector's beaked whale (M. hectori) · Strap-toothed
whales)
whale (M. layardii) · True's beaked whale (M. mirus) ·
Perrin's beaked whale (M. perrini) · Pygmy beaked whale
(M. peruvianus) · Stejneger's beaked whale (M.
stejnegeri) · Spade-toothed whale (M. traversii)
Tasmacetus Shepherd's beaked whale (T. sheperdi)
Ziphius Cuvier's beaked whale (Z. cavirostris)
Retrieved from "http://en.wikipedia.org/wiki/Sperm_whale"
Categories: IUCN Red List vulnerable species | Sperm whales | Cetaceans of Australia | Fauna of the
Atlantic Ocean | Fauna of the Pacific Ocean | Megafauna | Symbols of Connecticut | Monotypic mammal
genera | Animals described in 1758
W000
Plant evolutionary developmental biology


For a more ecological discussion on the evolution of plant morphology, refer to Evolutionary
history of plants
Evolutionary developmental biology (evo-devo) refers to the study of developmental programs and
patterns from an evolutionary perspective.[1] It seeks to understand the various influences shaping the form
and nature of life on the planet. Evo-devo arose as a separate branch of science only in the last decade.[2]
Most of the synthesis in evo-devo has been in the field of animal evolution, one reason being the presence of
elegant model systems like Drosophila melanogaster, C. elegans, zebrafish and Xenopus laevis. However, in
the past couple of decades, a wealth of information on plant morphology, coupled with modern molecular
techniques has helped shed light on the conserved and unique developmental patterns in the plant kingdom
also.[3]
Contents
[hide]
• 1 Historical perspective
• 1.1 Before 1900
• 1.2 1900 to the present
• 2 Organisms, databases and tools
• 3 Evolution of plant morphology
• 3.1 Overview of plant evolution
• 3.2 Evolution of meristems
• 3.2.1 Diversity in meristem architectures
• 3.2.2 Role of the KNOX-family genes
• 3.2.3 Evolution of the meristem architecture
• 3.3 Evolution of leaves
• 3.3.1 Origins of the leaf
• 3.3.2 Factors influencing leaf architectures
• 3.3.3 Genetic evidences for leaf evolution
• 3.4 Evolution of flowers
• 3.4.1 Origins of the flower
• 3.4.2 Evolution of the MADS-box family
• 3.4.3 Factors influencing floral diversity
• 3.4.4 Flowering time
• 3.4.5 Theories of flower evolution
[edit] Historical perspective
[edit] Before 1900
Johann Wolfgang von Goethe

The origin of the term "morphology" is generally attributed to Johann Wolfgang von Goethe (1749–
1832). He was of the opinion that there is an underlying fundamental organisation (Bauplan) in the diversity
of flowering plants. In his book titled The Metamorphosis of Plants, he proposed that the Bauplan enabled us
to predict the forms of plants that had not yet been discovered.[4] Goethe also was the first to make the
perceptive suggestion that flowers consist of modified leaves.
In the middle centuries, several basic foundations of our current understanding of plant morphology
were laid down. Nehemiah Grew, Marcello Malpighi, Robert Hooke, Antonie van Leeuwenhoek, Wilhelm von
Nageli were just some of the people who helped build knowledge on plant morphology at various levels of
organisation. It was the taxonomical classification of Carolus Linnaeus in the eighteenth century though, that
generated a firm base for the knowledge to stand on and expand.[5] The introduction of the concept of
Darwinism in contemporary scientific discourse also had had an effect on the thinking on plant forms and
their evolution.
Wilhelm Hofmeister, one of the most brilliant botanists of his times, was the one to diverge away from
the idealist way of pursuing botany. Over the course of his life, he brought an interdisciplinary outlook into
botanical thinking. He came up with biophysical explanations on phenomena like phototaxis and geotaxis,
and also discovered the alternation of generations in the plant life cycle.[4]
[edit] 1900 to the present
Arabidopsis thaliana. This flowering plant has been a model system for most of plant molecular
studies
The past century witnessed a rapid progress in the study of plant anatomy. The focus shifted from
the population level to more reductionist levels. While the first half of the century saw expansion in
developmental knowledge at the tissue and the organ level, in the latter half, especially since the 1990s,
there has also been a strong impetus on gaining molecular information.
Edward Charles Jeffrey was one of the early evo devo researchers of the 20th century. He performed
a comparative analyses of the vasculatures of living and fossil Gymnosperms and came to the conclusion
that the storage parenchyma has been derived from tracheids.[6] His research[7] focussed primarily on plant
anatomy in the context of phylogeny. This tradition of evolutionary analyses of plant architectures was further
advanced by Katherine Esau, best known for her book The Plant Anatomy. Her work focussed on the origin
and development of various tissues in different plants. Working with Vernon Cheadle[8], she also explained
the evolutionary specialization of the phloem tissue with respect to its function.
In the meantime, by the beginning of the latter half of 1900s, Arabidopsis thaliana had begun to be
used in some developmental studies. The first collection of Arabidopsis thaliana mutants were made around
1945.[9] However it formally became established as a model organism only in 1998.[10]
Wikispecies has information related to: Arabidopsis thaliana
The recent spurt in information on various plant-related processes has largely been a result of the
revolution in molecular biology. Powerful techniques like mutagenesis and complementation were made
possible in Arabidopsis thaliana via generation of T-DNA containing mutant lines, recombinant plasmids,
techniques like Transposon Tagging etc. Availability of complete physical and genetic maps,[11] RNAi
vectors, and rapid transformation[disambiguation needed] protocols are some of the technologies that have
significantly altered the scope of the field.[10] Recently, there has also been a massive increase in the
genome and EST sequences[12] of various non-model species, which, coupled with the Bioinformatics tools
existing today, generate interesting opportunities in the field of plant evo devo research.
Cusset provided a detailed in-depth analysis of the history of plant morphology, including plant
development and evolution, from its beginnings to the end of the 20th century.[13]
[edit] Organisms, databases and tools
The sampling of the Floral Genome Project

The most important model systems in plant development have been arabidopsis and maize. Maize
has traditionally been the favorite of plant geneticists, while extensive resources in almost every area of plant
physiology and development are available for Arabidopsis thaliana. Apart from these, rice, Antirrhinum
majus, Brassica, tomato are also being used in a variety of studies. The genomes of Arabidopsis thaliana
and rice have been completely sequenced, while the others are in process.[14]. It must be emphasized here
that the information from these "model" organisms form the basis of our developmental knowledge. While
Brassica has been used primarily because of its convenient location in the phylogenetic tree in the mustard
family, Antirrhinum majus is a convenient system for studying leaf architecture. Rice has been traditionally
used for studying responses to hormones like abscissic acid and gibberelin as well as responses to stress.
However, recently, not just the domesticated rice strain, but also the wild strains have been studied for their
underlying genetic architectures.[15]
Some people have objected against extending the results of model organisms to the plant world. One
argument is that the effect of gene knockouts in lab conditions wouldn't truly reflect even the same plant's
response in the natural world. Also, these supposedly crucial genes might not be responsible for the
evolutionary origin of that character. For these reasons, a comparative study of plant traits has been
proposed as the way to go now.[16]
Since the past few years, researchers have indeed begun looking at non-model, "non-conventional"
organisms using modern genetic tools. One example of this is the Floral Genome Project, which envisages to
study the evolution of the current patterns in the genetic architecture of the flower through comparative
genetic analyses, with a focus on EST sequences.[17] Like the FGP, there are several such ongoing projects
that aim to find out conserved and diverse patterns in evolution of the plant shape. Expressed sequence tag
(EST) sequences of quite a few non-model plants like Sugarcane, Apple, Lotus, Barley, Cycas, Coffee, to
name a few, are available freely online[18]. The Cycad Genomics Project,[19] for example, aims to
understand the differences in structure and function of genes between gymnosperms and angiosperms
through sampling in the order Cycadales. In the process, it intends to make available information for the
study of evolution of structures like seeds, cones and evolution of life cycle patterns. Presently the most
important sequenced genomes from an evo-devo point of view include those of A.thaliana (a flowering plant),
Poplar (a woody plant), Physcomitrella patens (a bryophyte), Maize (extensive genetic information), and
Chlamydomonas reinhardtii (a green alga). The impact of such a vast amount of information on
understanding common underlying developmental mechanisms can easily be realised.
Apart from EST and genome sequences, several other tools like PCR, Yeast two hybrid system,
microarrays, RNA Interference, SAGE, QTL mapping etc. permit the rapid study of plant developmental
patterns. Recently, cross-species hybridization has begun to be employed on microarray chips, to study the
conservation and divergence in mRNA expression patterns between closely related species.[20] Techniques
for analyzing this kind of data have also progressed over the past decade. We now have better models for
molecular evolution, more refined analysis algorithms and better computing power as a result of advances in
computer sciences.
[edit] Evolution of plant morphology

[edit] Overview of plant evolution
Main article: Evolutionary history of plants
Evidence suggests that an algal scum formed on the land 1,200 million years ago, but it was not until
the Ordovician period, around 500 million years ago, that land plants appeared. These begun to diversify in
the late Silurian period, around 420 million years ago, and the fruits of their diversification are displayed in
remarkable detail in an early Devonian fossil assemblage known as the Rhynie chert. This chert preserved
early plants in cellular detail, petrified in volcanic springs. By the middle of the Devonian period most of the
features recognised in plants today are present, including roots, leaves and seeds. By the late Devonian,
plants had reached a degree of sophistication that allowed them to form forests of tall trees. Evolutionary
innovation continued after the Devonian period. Most plant groups were relatively unscathed by the Permo-
Triassic extinction event, although the structures of communities changed. This may have set the scene for
the evolution of flowering plants in the Triassic (~200 million years ago), which exploded the Cretaceous and
Tertiary. The latest major group of plants to evolve were the grasses, which became important in the mid
Tertiary, from around 40 million years ago. The grasses, as well as many other groups, evolved new
mechanisms of metabolism to survive the low CO2 and warm, dry conditions of the tropics over the last 10
million years.
[edit] Evolution of meristems

The meristematic cells give rise to various organs of the plant, and keep the plant growing. The
Shoot Apical Meristem (SAM) gives rise to organs like the leaves and flowers. The cells of the apical
meristems - SAM and RAM (Root Apical Meristem)- divide rapidly and are considered to be indeterminate, in
that they do not possess any defined end fate. In that sense, the meristematic cells are frequently compared
to the stem cells in animals, that have an analogous behavior and function.
[edit] Diversity in meristem architectures

Is the mechanism of being indeterminate conserved in the SAM's of the plant world? The SAM
contains a population of stem cells that also produce the lateral meristems while the stem elongates. It turns
out that the mechanism of regulation of the stem cell number might indeed be evolutionarily conserved. The
CLAVATA gene CLV2 responsible for maintaining the stem cell population in Arabidopsis thaliana is very
closely related to the Maize gene FASCIATED EAR 2(FEA2) also involved in the same function.[21]
Similarly, in Rice, the FON1-FON2 system seems to bear a close relationship with the CLV signaling system
in Arabidopsis thaliana.[22] These studies suggest that the regulation of stem cell number, identity and
differentiation might be an evolutionarily conserved mechanism in monocots, if not in angiosperms. Rice also
contains another genetic system distinct from FON1-FON2, that is involved in regulating stem cell number.
[22] This example underlines the innovation that goes about in the living world all the time.
[edit] Role of the KNOX-family genes
Note the long spur of the above flower. Spurs attract pollinators and confer pollinator specificity.
(Flower:Linaria dalmatica)
Complex leaves of C. hirsuta are a result of KNOX gene expression
Genetic screens have identified genes belonging to the KNOX family in this function. These genes
essentially maintain the stem cells in an undifferentiated state. The KNOX family has undergone quite a bit of
evolutionary diversification, while keeping the overall mechanism more or less similar. Members of the KNOX
family have been found in plants as diverse as Arabidopsis thaliana, rice, barley and tomato. KNOX-like
genes are also present in some algae, mosses, ferns and gymnosperms. Misexpression of these genes
leads to formation of interesting morphological features. For example, among members of Antirrhinae, only
the species of genus Antirrhinum lack a structure called spur in the floral region. A spur is considered an
evolutionary innovation because it defines pollinator specificity and attraction. Researchers carried out
transposon mutagenesis in Antirrhinum majus, and saw that some insertions led to formation of spurs that
were very similar to the other members of Antirrhinae[23], indicating that the loss of spur in wild Antirrhinum
majus populations could probably be an evolutionary innovation.
The KNOX family has also been implicated in leaf shape evolution (See below for a more detailed
discussion). One study looked at the pattern of KNOX gene expression in A. thaliana, that has simple leaves
and Cardamine hirsuta, a plant having complex leaves. In A. thaliana, the KNOX genes are completely
turned off in leaves, but in C.hirsuta, the expression continued, generating complex leaves.[24] Also, it has
been proposed that the mechanism of KNOX gene action is conserved across all vascular plants, because
there is a tight correlation between KNOX expression and a complex leaf morphology.[25]
[edit] Evolution of the meristem architecture

The meristem architectures do differ between angiosperms, gymnosperms and pteridophytes. The
gymnosperm vegetative meristem lacks organization into distinct tunica and corpus layers. They possess
large cells called Central Mother Cells in the meristem. In angiosperms, the outermost layer of cells divides
anticlinally to generate the new cells, while in gymnosperms, the plane of division in the meristem differs for
different cells. However, the apical cells do contain organelles like large vacuoles and starch grains, like the
angiosperm meristematic cells.
Pteridophytes, like fern, on the other hand, do not possess a multicellular apical meristem. They
possess a tetrahedral apical cell, which goes on to form the plant body. Any somatic mutation in this cell can
lead to hereditary transmission of that mutation.[26] The earliest meristem-like organization is seen in an
algal organism from group Charales that has a single dividing cell at the tip, much like the pteridophytes, yet
simpler. One can thus see a clear pattern in evolution of the meristematic tissue, from pteridophytes to
angiosperms. Pteridophytes, with a single meristematic cell; gymnosperms with a multicellular, but less
defined organization and finally, angiosperms, with the highest degree of organization. The genetic
innovations that contributed to this evolution are yet not clearly known.
[edit] Evolution of leaves

For a discussion on Evolution of Photosynthesis, see Photosynthesis.
[edit] Origins of the leaf

Further information: Evolutionary history of plants#leaves and Megaphyll
Leaf lamina. The leaf architecture probably arose multiple times in the plant lineage
Leaves are the primary photosynthetic organs of a plant. Based on their structure, they are classified
into two types - microphylls, that lack complex venation patterns and megaphylls, that are large and with a
complex venation. It has been proposed that these structures arose independently.[27] Megaphylls,
according to the Telome hypothesis, have evolved from plants that showed a three dimensional branching
architecture, through three transformations—planation, which involved formation of a planar architecture,
webbing, or formation of the outgrowths between the planar branches and fusion, where these webbed
outgrowths fused to form a proper leaf lamina. Studies have revealed that these three steps happened
multiple times in the evolution of today's leaves.[28]
It has been proposed that the before the evolution of leaves, plants had the photosynthetic apparatus
on the stems. Today's megaphyll leaves probably became commonplace some 360mya, about 40my after
the simple leafless plants had colonized the land in the early Devonian period. This spread has been linked
to the fall in the atmospheric carbon dioxide concentrations in the Late Paleozoic era associated with a rise in
density of stomata on leaf surface. This must have allowed for better transpiration rates and gas exchange.
Large leaves with less stomata would have gotten heated up in the sun's heat, but an increased stomatal
density allowed for a better-cooled leaf, thus making its spread feasible[29][30].
[edit] Factors influencing leaf architectures
Spiny leaves of Aciphylla squarrosa. It is thought that these leaves evolved as an adaptation against
the now extinct Moas
Various physical and physiological forces like light intensity, humidity, temperature, wind speeds etc.
are thought to have influenced evolution of leaf shape and size. It is observed that high trees rarely have
large leaves, owing to the obstruction they generate for winds. This obstruction can eventually lead to the
tearing of leaves, if they are large. Similarly, trees that grow in temperate or taiga regions have pointed
leaves, presumably to prevent nucleation of ice onto the leaf surface and reduce water loss due to
transpiration. Herbivory, not only by large mammals, but also small insects has been implicated as a driving
force in leaf evolution, an example being plants of the genus Aciphylla, that are commonly found in New
Zealand. The now extinct Moas fed upon these plants, and its seen that the leaves have spines on their
bodies, which probably functioned to discourage the moas from feeding on them. Other members of
Aciphylla that did not co-exist with the moas, do not have these spines.[31]
[edit] Genetic evidences for leaf evolution

At the genetic level, developmental studies have shown that repression of the KNOX genes is
required for initiation of the leaf primordium. This is brought about by ARP genes, which encode transcription
factors. Genes of this type have been found in many plants studied till now, and the mechanism i.e.
repression of KNOX genes in leaf primordia, seems to be quite conserved. Interestingly, expression of KNOX
genes in leaves produces complex leaves. It is speculated that the ARP function arose quite early in vascular
plant evolution, because members of the primitive group Lycophytes also have a functionally similar gene
[32] Other players that have a conserved role in defining leaf primordia are the phytohormone auxin,
gibberelin and cytokinin.
The diversity of leaves
One interesting feature of a plant is its phyllotaxy. The arrangement of leaves on the plant body is
such that the plant can maximally harvest light under the given constraints, and hence, one might expect the
trait to be genetically robust. However, it may not be so. In maize, a mutation in only one gene called abphyl
(ABNORMAL PHYLLOTAXY) was enough to change the phyllotaxy of the leaves. It implies that sometimes,
mutational tweaking of a single locus on the genome is enough to generate diversity. The abphyl gene was
later on shown to encode a cytokinin response regulator protein.[33]
Once the leaf primordial cells are established from the SAM cells, the new axes for leaf growth are
defined, one important (and more studied) among them being the abaxial-adaxial (lower-upper surface) axes.
The genes involved in defining this, and the other axes seem to be more or less conserved among higher
plants. Proteins of the HD-ZIPIII family have been implicated in defining the adaxial identity. These proteins
deviate some cells in the leaf primordium from the default abaxial state, and make them adaxial. It is believed
that in early plants with leaves, the leaves just had one type of surface - the abaxial one. This is the
underside of today's leaves. The definition of the adaxial identity occurred some 200 million years after the
abaxial identity was established[34]. One can thus imagine the early leaves as an intermediate stage in
evolution of today's leaves, having just arisen from spiny stem-like outgrowths of their leafless ancestors,
covered with stomata all over, and not optimized as much for light harvesting.
How the infinite variety of plant leaves is generated is a subject of intense research. Some common
themes have emerged. One of the most significant is the involvement of KNOX genes in generating
compound leaves, as in tomato (see above). But this again is not universal. For example, pea uses a
different mechanism for doing the same thing[35][36]. Mutations in genes affecting leaf curvature can also
change leaf form, by changing the leaf from flat, to a crinky shape,[37] like the shape of cabbage leaves.
There also exist different morphogen gradients in a developing leaf which define the leaf's axis. Changes in
these morphogen gradients may also affect the leaf form. Another very important class of regulators of leaf
development are the microRNAs, whose role in this process has just begun to be documented. The coming
years should see a rapid development in comparative studies on leaf development, with many EST
sequences involved in the process coming online.
[edit] Evolution of flowers

For a more ecological discussion on the evolution of flowers, go to Flower or Evolutionary history of
plants
The pollen bearing organs of the early flower Crossotheca

A flower is, arguably, one of the most beautiful products of evolution. Flower-like structures first
appear in the fossil records some ~130 mya, in the Cretaceous era[38].
The flowering plants have long been assumed to have evolved from within the gymnosperms;
according to the traditional morphological view, they are closely allied to the gnetales. However, recent
molecular evidence is at odds to this hypothesis,[39][40] and further suggests that gnetales are more closely
related to some gymnosperm groups than angiosperms,[41] and that gymnosperms form a distinct clade to
the angiosperms,[39][40][41]. Molecular clock analysis predicts the divergence of flowering plants
(anthophytes) and gymnosperms to ~300 mya[42]
Phylogeny of anthophytes and gymnosperms, from [43]
Cycads
Ginkgo
Conifers
Anthophytes
Bennettitales
Gnetales
Angiosperms
Angiosperms
Gymnosperms
Cycads
Bennettitales
Ginkgo
Conifers
Gnetales
Traditional view Modern view

The main function of a flower is reproduction, which, before the evolution of the flower and
angiosperms, was the job of microsporophylls and megasporophylls. A flower can be considered a powerful
evolutionary innovation, because its presence allowed the plant world to access new means and
mechanisms for reproduction.
[edit] Origins of the flower
Amborella trichopoda : Amborellaceae is considered the sister family of all flowering plants
(magnified image)
The family Amborellaceae is regarded as the sister family of all living flowering plants. That means
members of this family were most likely the first flowering plants.
It seems that on the level of the organ, the leaf may be the ancestor of the flower, or at least some
floral organs. When we mutate some crucial genes involved in flower development, we end up with a cluster
of leaf-like structures. Thus, sometime in history, the developmental program leading to formation of a leaf
must have been altered to generate a flower. There probably also exists an overall robust framework within
which the floral diversity has been generated. A example of that is a gene called LEAFY (LFY), which is
involved in flower development in Arabidopsis thaliana. The homologs of this gene are found in angiosperms
as diverse as tomato, snapdragon, pea, maize and even gymnosperms. Interestingly, expression of
Arabidopsis thaliana LFY in distant plants like poplar and citrus also results in flower-production in these
plants. The LFY gene regulates the expression of some gene belonging to the MADS-box family. These
genes, in turn, act as direct controllers of flower development.
[edit] Evolution of the MADS-box family

The members of the MADS-box family of transcription factors play a very important and evolutionarily
conserved role in flower development. According to the ABC Model of flower development, three zones - A,B
and C - are generated within the developing flower primordium, by the action of some transcription factors,
that are members of the MADS-box family. Among these, the functions of the B and C domain genes have
been evolutionarily more conserved than the A domain gene. Many of these genes have arisen through gene
duplications of ancestral members of this family. Quite a few of them show redundant functions.
The evolution of the MADS-box family has been extensively studied. These genes are present even
in pteridophytes, but the spread and diversity is many times higher in angiosperms[44]. There appears to be
quite a bit of pattern into how this family has evolved. Consider the evolution of the C-region gene
AGAMOUS (AG). It is expressed in today's flowers in the stamens, and the carpel, which are reproductive
organs. It's ancestor in gymnosperms also has the same expression pattern. Here, it is expressed in the
strobili, an organ that produces pollens or ovules[45]. Similarly, the B-genes' (AP3 and PI) ancestors are
expressed only in the male organs in gymnosperms. Their descendants in the modern angiosperms also are
expressed only in the stamens, the male reproductive organ. Thus, the same, then-existing components
were used by the plants in a novel manner to generate the first flower. This is a recurring pattern in evolution.
[edit] Factors influencing floral diversity
Wikiversity has bloom time data for Linaria vulgaris on the Bloom Clock
The various shapes and colors of flowers

How is the enormous diversity in the shape, color and sizes of flowers established? There is
enormous variation in the developmental program in different plants. For example, monocots possess
structures like lodicules and palea, that were believed to be analogous to the dicot petals and carpels
respectively.It turns out that this is true, and the variation is due to slight changes in the MADS-box genes
and their expression pattern in the monocots. Another example is that of a plant called Linaria vulgaris, which
has two kinds of flower symmetries-radial and bilateral. These symmetries are due to epigenetic changes in
just one gene called CYCLOIDEA.[38]
Large number of petals in roses has probably been a result of human selection
Arabidopsis thaliana has a gene called AGAMOUS that plays an important role in defining how many
petals and sepals and other organs are generated. Mutations in this gene give rise to the floral meristem
obtaining an indeterminate fate, and many floral organs keep on getting produced. We have flowers like
roses, carnations and morning glory, for example, that have very dense floral organs. These flowers have
been selected by horticulturists since long for increased number of petals. Researchers have found that the
morphology of these flowers is because of strong mutations in the AGAMOUS homolog in these plants,
which leads to them making a large number of petals and sepals.[46] Several studies on diverse plants like
petunia, tomato, Impatiens, maize etc. have suggested that the enormous diversity of flowers is a result of
small changes in genes controlling their development[47].
Some of these changes also cause changes in expression patterns of the developmental genes,
resulting in different phenotypes. The Floral Genome Project looked at the EST data from various tissues of
many flowering plants. The researchers confirmed that the ABC Model of flower development is not
conserved across all angiosperms. Sometimes expression domains change, as in the case of many
monocots, and also in some basal angiosperms like Amborella. Different models of flower development like
the The fading boundaries model, or the Overlapping-boundaries model which propose non-rigid domains of
expression, may explain these architectures.[48] There is a possibility that from the basal to the modern
angiosperms, the domains of floral architecture have gotten more and more fixed through evolution.
[edit] Flowering time

Another floral feature that has been a subject of natural selection is flowering time. Some plants
flower early in their life cycle, others require a period of vernalization before flowering. This decision is based
on factors like temperature, light intensity, presence of pollinators and other environmental signals. We know
that genes like CONSTANS (CO), Flowering Locus C (FLC) and FRIGIDA regulate integration of
environmental signals into the pathway for flower development. Variations in these loci have been associated
with flowering time variations between plants. For example, Arabidopsis thaliana ecotypes that grow in the
cold, temperate regions require prolonged vernalization before they flower, while the tropical varieties, and
the most common lab strains, don't. We now know that this variation is due to mutations in the FLC and
FRIGIDA genes, rendering them non-functional.[49]
Quite a few players in this process are conserved across all the plants studied. Sometimes though,
despite genetic conservation, the mechanism of action turns out to be different. For example, rice is a short-
day plant, while Arabidopsis thaliana is a long-day plant. Now, in both plants, the proteins CO and
FLOWERING LOCUS T (FT) are present. But in Arabidopsis thaliana, CO enhances FT production, while in
rice, the CO homolog represses FT production, resulting in completely opposite downstream effects[50].
[edit] Theories of flower evolution
Main article: Evolutionary history of plants#flowers
There are many theories that propose how flowers evolved. Some of them are described below.
The Anthophyte Theory was based upon the observation that a gymnospermic group Gnetales has a
flower-like ovule. It has partially developed vessels as found in the angiosperms, and the megasporangium is
covered by three envelopes, like the ovary structure of angiosperm flowers. However, many other lines of
evidence show that Gnetales is not related to angiosperms.[43]
Further information: anthophyta
The Mostly Male Theory has a more genetic basis. Proponents of this theory point out that the
gymnosperms have two very similar copies of the gene LFY while angiosperms just one. Molecular clock
analysis has shown that the other LFY paralog was lost in angiosperms around the same time as flower
fossils become abundant, suggesting that this event might have led to floral evolution.[51] According to this
theory, loss of one of the LFY paralog led to flowers that were more male, with the ovules being expressed
ectopically. These ovules initially performed the function of attracting pollinators, but sometime later, may
have been integrated into the core flower.
One theory also suggests that humans have been one of the reasons for the diversity of flowers. This
theory suggests that since the early settlers found flowers beautiful, they may have started selecting for them
artificially.[52] The flowers may have evolved to exploit the ecological niche being opened because of
humans finding them attractive. The validity of this theory, however, is debatable, not least because flowers
started diversifying long before they came into contact with humans.
[edit] Evolution of secondary metabolism
Structure of Azadirachtin, a terpenoid produced by the Neem plant, which helps ward off microbes
and insects. Many secondary metabolites have complex structures
Although we know many secondary metabolites produced by plants, the extent of the same is still
unfathomable. Secondary metabolites are essentially low molecular weight compounds, sometimes having
complex structures. They function in processes as diverse as immunity, anti-herbivory, pollinator attraction,
communication between plants, maintaining symbiotic associations with soil flora, enhancing the rate of
fertilization etc., and hence are significant from the evo-devo perspective. The structural and functional
diversity of these secondary metabolites across the plant kingdom is so huge that it is estimated that
hundreds of thousands of enzymes might be involved in this process in the entire of the plant kingdom, with
about 15–25% of the coding genome coding for these enzymes. Despite this, every species has its unique
arsenal of secondary metabolites.[53] Many of these metabolites are of enormous medical significance to
humans.
What is the purpose of having so many secondary metabolites being produced, with a significant
chunk of the metabolome devoted to this activity? It is hypothesized that most of these chemicals help in
generating immunity, and in consequence, the diversity of these metabolites is a result of a constant war
between plants and their parasites. There is evidence that this may be true in many cases. The big question
here is the reproductive cost involved in maintaining such an impressive inventory. Various models have
been suggested that probe into this aspect of the question, but a consensus on the extent of the cost is
lacking.[54] We[who?] still cannot predict whether a plant with more secondary metabolites would be better
off than other plants in its vicinity.
Secondary metabolite production seems to have arisen quite early during evolution. Even bacteria
possess the ability to make these compounds. But they assume more significant roles in life from fungi
onwards to plants. In plants they seem to have spread out using different mechanisms like gene duplications,
evolution of novel genes etc. Furthermore, studies have shown that diversity in some of these compounds
may be positively selected for.
Although the role of novel gene evolution in the evolution of secondary metabolism cannot be denied,
there are several examples where new metabolites have been formed by small changes in the reaction. For
example, cyanogen glycosides have been proposed to have evolved multiple times in different plant
lineages. There are several such instances of convergent evolution. For example, we now know that
enzymes for synthesis of limonene – a terpene – are more similar between angiosperms and gymnosperms
than to their own terpene synthesis enzymes. This suggests independent evolution of the limonene
biosynthetic pathway in these two lineages.[55]
[edit] Mechanisms and players in evolution
The stem-loop secondary structure of a pre-microRNA from Brassica oleracea

While environmental factors are significantly responsible for evolutionary change, they act merely as
agents for natural selection. Change is inherently brought about via phenomena at the genetic level -
mutations, chromosomal rearrangements and epigenetic changes. While the general types of mutations hold
true across the living world, in plants, some other mechanisms have been implicated as highly significant.
Polyploidy is a very common feature in plants. It is believed that at least half (and probably all) plants
are or have been polyploids. Polyploidy leads to genome doubling, thus generating functional redundancy in
most genes. The duplicated genes may attain new function, either by changes in expression pattern or
changes in activity. Polyploidy and gene duplication are believed to be among the most powerful forces in
evolution of plant form. It is not know though, why genome doubling is such a frequent process in plants. One
probable reason is the production of large amounts of secondary metabolites in plant cells. Some of them
might interfere in the normal process of chromosomal segregation, leading to polypoidy.
Extreme left: teosinte, Extreme right: maize, middle: maize-teosinte hybrid

In recent times, plants have been shown to possess significant microRNA families, which are
conserved across many plant lineages. In comparison to animals, while the number of plant miRNA families
are lesser than animals, the size of each family is much larger. The miRNA genes are also much more
spread out in the genome than those in animals, where we find them clustered. It has been proposed that
these miRNA families have expanded by duplications of chromosomal regions.[56] Many miRNA genes
involved in regulation of plant development have been found to be quite conserved between plants studied.
Domestication of plants like maize, rice, barley, wheat etc. has also been a significant driving force in
their evolution. Some studies have tried to look at the origins of the maize plant and it turns out that maize is
a domesticated derivative of a wild plant from Mexico called teosinte. Teosinte belongs to the genus Zea, just
as maize, but bears very small inflorescence, 5-10 hard cobs and a highly branched and spread out stem.
Cauliflower : Brassica oleracea var botrytis

Interestingly, crosses between a particular teosinte variety and maize yields fertile offsprings that are
intermediate in phenotype between maize and teosinte. QTL analysis has also revealed some loci that when
mutated in maize yield a teosinte-like stem or teosinte-like cobs. Molecular clock analysis of these genes
estimates their origins to some 9000 years ago, well in accordance with other records of maize
domestication. It is believed that a small group of farmers must have selected some maize-like natural
mutant of teosinte some 9000 years ago in Mexico, and subjected it to continuous selection to yield the
maize plant as we know today.[57]
Another interesting case is that of cauliflower. The edible cauliflower is a domesticated version of the
wild plant Brassica oleracea, which does not possess the dense undifferentiated inflorescence called the
curd, that cauliflower possesses.
Wikispecies has information related to: Brassicaceae
Cauliflower possesses a single mutation in a gene called CAL, controlling meristem differentiation
into inflorescence. This causes the cells at the floral meristem to gain an undifferentiated identity, and instead
of growing into a flower, they grow into a lump of undifferentiated cells.[58] This mutation has been selected
through domestication at least since the Greek empire.
[edit] See also

• Evolutionary developmental biology
• Plant morphology
• Comparative phylogenetics
• Plant evolution
[edit] Suggested readings

1) The Genetics of plant morphological evolution
2) Plant evolution and development in a post-genomic context
3) Evolution of leaf developmental mechanisms
4) Developmental genetics and plant evolution ISBN 0-415-25791-3
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"[http://www.plantphysiol.org/cgi/content/full/127/4/1533#B18 The Regulation of Compound Leaf
Development"]. Plant Physiol. 127 (4): 1533–1538. PMID 11743098.
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Ecol. And Evol. 15 (4): 144–149.
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phylogeny inferred from all three plant genomes: Monophyly of extant gymnosperms and origin of
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evidence from eight genes" (abstract). American Journal of Botany 89 (10): 1670.
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are conifers". Proceedings of the National Academy of Sciences 97 (8): 4092.
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Box Gene Family Controlling Flower Development in Plants". Mol. Biol. Evol. 20 (9): 1435–1447.
doi:10.1093/molbev/msg152. PMID 12777513.
43.^ a b Crepet, W. L. (2000). "Progress in understanding angiosperm history, success, and
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gene family". Nat Rev Gen 2: 186–195.
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Mind

For other uses, see Mind (disambiguation).
Mind (pronounced /ˈmaɪnd/) is the aspect of intellect and consciousness experienced as
combinations of thought, perception, memory, emotion, will, and imagination, including all unconscious
cognitive processes. The term is often used to refer, by implication, to the thought processes of reason. Mind
manifests itself subjectively as a stream of consciousness.
Theories of mind and its function are numerous. Earliest recorded speculations are from the likes of
Zoroaster, the Buddha, Plato, Aristotle, Adi Shankara and other ancient Greek, Indian and, later, Islamic
philosophers. Pre-scientific theories grounded in theology concentrated on the supposed relationship
between the mind and the soul, a human's supernatural, divine or god-given essence.
Which attributes make up the mind is much debated. Some psychologists argue that only the higher
intellectual functions constitute mind, particularly reason and memory. In this view the emotions—love, hate,
fear, joy—are more primitive or subjective in nature and should be seen as different from the mind as such.
Others argue that various rational and emotional states cannot be so separated, that they are of the same
nature and origin, and should therefore be considered all part of what we call the mind.
In popular usage mind is frequently synonymous with thought: the private conversation with
ourselves that we carry on "inside our heads." Thus we "make up our minds," "change our minds" or are "of
two minds" about something. One of the key attributes of the mind in this sense is that it is a private sphere to
which no one but the owner has access. No one else can "know our mind." They can only interpret what we
consciously or unconsciously communicate.
Contents
[hide]
• 1 Etymology
• 2 Mental faculties
• 3 Brain and mind
• 4 Philosophy of mind
• 5 Science of mind
• 5.1 Psychology
• 5.2 Evolutionary psychology
• 6 Evolutionary history of the human mind
• 7 Mental health
• 8 Animal intelligence
• 9 Artificial intelligence
• 10 Religious perspectives
• 11 Other perspectives
• 11.1 Parapsychology
• 11.2 Memetics
• 12 See also
• 13 References
[edit] Etymology
Further information: Geist
The original meaning of Old English gemynd was the faculty of memory, not of thought in general.
Hence call to mind, come to mind, keep in mind, to have mind of, etc. Old English had other words to express
"mind", such as hyge "mind, spirit".
The generalization of mind to include all mental faculties, thought, volition, feeling and memory,
gradually develops over the 14th and 15th centuries.[1]
The meaning of "memory" is shared with Old Norse, which has munr. The word is originally from a
PIE verbal root *men-, meaning "to think, remember", whence also Latin mens "mind", Sanskrit manas "mind"
and Greek μένος "mind, courage, anger".
[edit] Mental faculties

See also: Reason, Faculty psychology, and Modularity of mind
Thought is a mental process which allows individuals to model the world, and so to deal with it
effectively according to their goals, plans, ends and desires. Words referring to similar concepts and
processes include cognition, idea, and imagination. Thinking involves the cerebral manipulation of
information, as when we form concepts, engage in problem solving, reasoning and making decisions.
Thinking is a higher cognitive function and the analysis of thinking processes is part of cognitive psychology.
Memory is an organism's ability to store, retain, and subsequently recall information. Although
traditional studies of memory began in the realms of philosophy, the late nineteenth and early twentieth
century put memory within the paradigms of cognitive psychology. In recent decades, it has become one of
the principal pillars of a new branch of science called cognitive neuroscience, a marriage between cognitive
psychology and neuroscience.
Imagination is the ability to generate mental representations of objects or experiences, and is based
on internal goals or processes rather than perceptual stimulation by the current environment. The term is
technically used in psychology for the process of reviving in the mind percepts of objects formerly given in
sense perception. Since this use of the term conflicts with that of ordinary language, some psychologists
have preferred to describe this process as "imaging" or "imagery" or to speak of it as "reproductive" as
opposed to "productive" or "constructive" imagination. Imagined images are seen with the "mind's eye".
Among the many useful applications of imagination is the ability to simulate possible futures, and make
decisions that maximize imagined future benefits.
Consciousness in mammals (this includes humans) is an aspect of the mind generally thought to
comprise qualities such as subjectivity, sentience, and the ability to perceive the relationship between oneself
and one's environment. It is a subject of much research in philosophy of mind, psychology, neuroscience,
and cognitive science. Some philosophers divide consciousness into phenomenal consciousness, which is
subjective experience itself, and access consciousness, which refers to the global availability of information
to processing systems in the brain.[2] Phenomenal consciousness has many different experienced qualities,
often referred to as qualia. Phenomenal consciousness is usually consciousness of something or about
something, a property known as intentionality in philosophy of mind.
[edit] Brain and mind

See also: Cognitive science
In animals, the brain, or encephalon (Greek for "in the head"), is the control center of the central
nervous system, responsible for thought. In most animals, the brain is located in the head, protected by the
skull and close to the primary sensory apparatus of vision, hearing, equilibrioception, taste and olfaction.
While all vertebrates have a brain, most invertebrates have either a centralized brain or collections of
individual ganglia. Primitive animals such as sponges do not have a brain at all. Brains can be extremely
complex. For example, the human brain contains more than 100 billion neurons, each linked to as many as
10,000 others[citation needed].
Understanding the relationship between the brain and the mind — mind-body problem is one of the
central issues in the history of philosophy — is a challenging problem both philosophically and scientifically.[3]
There are three major philoosphical schools of thought concerning the answer: dualism, materialism, and
idealism. Dualism holds that the mind exists independently of the brain;[4] materialism holds that mental
phenomena are identical to neuronal phenomena;[5] and idealism holds that only mental phenomena exist.
[5]
The most straightforward scientific evidence that there is a strong relationship between the physical
brain matter and the mind is the impact physical alterations to the brain have on the mind, such as with
traumatic brain injury and psychoactive drug use.[6]
In addition to the philosophical questions, the relationship between mind and brain involves a high
number of scientific questions, including understanding the relationship between mental activity and brain
activity, the exact mechanisms by which drugs influence cognition, and the neural correlates of
consciousness.
Through most of history many philosophers found it inconceivable that cognition could be
implemented by a physical substance such as brain tissue (that is neurons and synapses).[7] Philosophers
such as Patricia Churchland posit that the drug-mind interaction is indicative of an intimate connection
between the brain and the mind, not that the two are the same entity.[8] Descartes, who thought extensively
about mind-brain relationships, found it possible to explain reflexes and other simple behaviors in
mechanistic terms, although he did not believe that complex thought, and language in particular, could be
explained by reference to the physical brain alone.[9]
[edit] Philosophy of mind

See also: Philosophy of mind
Philosophy of mind is the branch of philosophy that studies the nature of the mind, mental events,
mental functions, mental properties, consciousness and their relationship to the physical body. The mind-
body problem, i.e. the relationship of the mind to the body, is commonly seen as the central issue in
philosophy of mind, although there are other issues concerning the nature of the mind that do not involve its
relation to the physical body.[10]
"Aristotelian thought has permeated most Occidental philosophical system until modern times, and the
classification of man's function as vegetative, sensitive, and rational is still useful. In present popular usage,
soul and mind are not clearly differentiated and some people, more or less consciously, still feel that the soul,
and perhaps the mind are not clearly differentiated and some people, more or less consciously, still feel that
the soul, and perhaps the mind, may enter or leave the body as independent entities. "- Jose M.R. Delgado
[11]
Dualism and monism are the two major schools of thought that attempt to resolve the mind-body problem.
Dualism is the position that mind and body are in some way separate from each other. It can be traced back
to Plato,[12] Aristotle[13][14][15] and the Samkhya and Yoga schools of Hindu philosophy,[16] but it was
most precisely formulated by René Descartes in the 17th century.[17] Substance dualists argue that the mind
is an independently existing substance, whereas Property dualists maintain that the mind is a group of
independent properties that emerge from and cannot be reduced to the brain, but that it is not a distinct
substance.[18]
Monism is the position that mind and body are not physiologically and ontologically distinct kinds of
entities. This view was first advocated in Western Philosophy by Parmenides in the 5th Century BC and was
later espoused by the 17th Century rationalist Baruch Spinoza.[19] According to Spinoza's dual-aspect
theory, mind and body are two aspects of an underlying reality which he variously described as "Nature" or
"God". Physicalists argue that only the entities postulated by physical theory exist, and that the mind will
eventually be explained in terms of these entities as physical theory continues to evolve. Idealists maintain
that the mind is all that exists and that the external world is either mental itself, or an illusion created by the
mind. Neutral monists adhere to the position that perceived things in the world can be regarded as either
physical or mental depending on whether one is interested in their relationship to other things in the world or
their relationship to the perceiver. For example, a red spot on a wall is physical in its dependence on the wall
and the pigment of which it is made, but it is mental in so far as its perceived redness depends on the
workings of the visual system. Unlike dual-aspect theory, neutral monism does not posit a more fundamental
substance of which mind and body are aspects. The most common monisms in the 20th and 21st centuries
have all been variations of physicalism; these positions include behaviorism, the type identity theory,
anomalous monism and functionalism.[20]
Many modern philosophers of mind adopt either a reductive or non-reductive physicalist position,
maintaining in their different ways that the mind is not something separate from the body.[20] These
approaches have been particularly influential in the sciences, particularly in the fields of sociobiology,
computer science, evolutionary psychology and the various neurosciences.[21][22][23][24] Other
philosophers, however, adopt a non-physicalist position which challenges the notion that the mind is a purely
physical construct. Reductive physicalists assert that all mental states and properties will eventually be
explained by scientific accounts of physiological processes and states.[25][26][27] Non-reductive physicalists
argue that although the brain is all there is to the mind, the predicates and vocabulary used in mental
descriptions and explanations are indispensable, and cannot be reduced to the language and lower-level
explanations of physical science.[28][29] Continued neuroscientific progress has helped to clarify some of
these issues. However, they are far from having been resolved, and modern philosophers of mind continue to
ask how the subjective qualities and the intentionality (aboutness) of mental states and properties can be
explained in naturalistic terms.[30][31]
[edit] Science of mind

[edit] Psychology
See also: Sigmund Freud, Carl Jung, and Unconscious mind
Psychology is the scientific study of human behaviour, mental functioning, and experience; noology,
the study of thought. As both an academic and applied discipline, Psychology involves the scientific study of
mental processes such as perception, cognition, emotion, personality, as well as environmental influences,
such as social and cultural influences, and interpersonal relationships, in order to devise theories of human
behaviour. Psychology also refers to the application of such knowledge to various spheres of human activity,
including problems of individuals' daily lives and the treatment of mental health problems.
Psychology differs from the other social sciences (e.g., anthropology, economics, political science,
and sociology) due to its focus on experimentation at the scale of the individual, or individuals in small groups
as opposed to large groups, institutions or societies. Historically, psychology differed from biology and
neuroscience in that it was primarily concerned with mind rather than brain. Modern psychological science
incorporates physiological and neurological processes into its conceptions of perception, cognition,
behaviour, and mental disorders.
[edit] Evolutionary psychology
Evolutionary psychology (EP) is an approach within psychology that examines psychological traits —
such as memory, perception, or language — from a Darwinian evolutionary perspective. It seeks to explain
how many human psychological traits are evolved adaptations, that is, the functional products of natural
selection or sexual selection. Adaptationist thinking about physiological mechanisms, such as the heart,
lungs, and immune system, is common in evolutionary biology. Evolutionary psychology applies the same
thinking to psychology.
Evolutionary psychologists argue that much of human behavior is originates as psychological
adaptations that evolved to solve recurrent problems in human ancestral environments.[32]
Further information: Evolutionary neuroscience, Konrad Lorentz, E O Wilson, The Adapted
Mind, and Steven Pinker
[edit] Evolutionary history of the human mind

The evolution of human intelligence refers to a set of theories that attempt to explain how human
intelligence has evolved. The question is closely tied to the evolution of the human brain, and to the
emergence of human language.
The timeline of human evolution spans some 7 million years, from the separation of the Pan genus
until the emergence of behavioral modernity by 50,000 years ago. Of this timeline, the first 3 million years
concern Sahelanthropus, the following 2 million concern Australopithecus, while the final 2 million span the
history of actual human species (the Paleolithic).
Many traits of human intelligence, such as empathy, theory of mind, mourning, ritual, and the use of
symbols and tools, are already apparent in great apes although in lesser sophistication than in humans.
There is a debate between supporters of the idea of a sudden emergence of intelligence, or "Great
leap forward" and those of a gradual or continuum hypothesis.
Theories of the evolution of intelligence include:
• Robin Dunbar's social brain hypothesis[33]
• Geoffrey Miller's sexual selection hypothesis[34]
• The ecological dominance-social competition (EDSC) [35] explained by Mark V. Flinn, David
C. Geary and Carol V. Ward based mainly on work by Richard D. Alexander.
• The idea Intelligence as a signal opf good health and resistance to disease.
• The Group selection theory contends that organism characteristics that provide benefits to a
group (clan, tribe, or larger population) can evolve despite individual disadvantages such as those
cited above.
• The idea that intelligence is connected with nutrition, and thereby with status[36] A higher IQ
could be a signal that an individual comes from and lives in a physical and social environment where
nutrition levels are high, and vice versa.
[edit] Mental health

Main article: Mental health
By analogy with the health of the body, one can speak metaphorically of a state of health of the mind,
or mental health. Merriam-Webster defines mental health as "A state of emotional and psychological well-
being in which an individual is able to use his or her cognitive and emotional capabilities, function in society,
and meet the ordinary demands of everyday life." According to the World Health Organization (WHO), there
is no one "official" definition of mental health. Cultural differences, subjective assessments, and competing
professional theories all affect how "mental health" is defined. In general, most experts agree that "mental
health" and "mental illness" are not opposites. In other words, the absence of a recognized mental disorder is
not necessarily an indicator of mental health.
One way to think about mental health is by looking at how effectively and successfully a person
functions. Feeling capable and competent; being able to handle normal levels of stress, maintaining
satisfying relationships, and leading an independent life; and being able to "bounce back," or recover from
difficult situations, are all signs of mental health.
Psychotherapy is an interpersonal, relational intervention used by trained psychotherapists to aid
clients in problems of living. This usually includes increasing individual sense of well-being and reducing
subjective discomforting experience. Psychotherapists employ a range of techniques based on experiential
relationship building, dialogue, communication and behavior change and that are designed to improve the
mental health of a client or patient, or to improve group relationships (such as in a family). Most forms of
psychotherapy use only spoken conversation, though some also use various other forms of communication
such as the written word, art, drama, narrative story, or therapeutic touch. Psychotherapy occurs within a
structured encounter between a trained therapist and client(s). Purposeful, theoretically based psychotherapy
began in the 19th century with psychoanalysis; since then, scores of other approaches have been developed
and continue to be created.
[edit] Animal intelligence

Animal cognition, or cognitive ethology, is the title given to a modern approach to the mental
capacities of animals. It has developed out of comparative psychology, but has also been strongly influenced
by the approach of ethology, behavioral ecology, and evolutionary psychology. Much of what used to be
considered under the title of animal intelligence is now thought of under this heading. Animal language
acquisition, attempting to discern or understand the degree to which animal cognition can be revealed by
linguistics-related study, has been controversial among cognitive linguists.
[edit] Artificial intelligence

This section needs additional citations for verification.
challenged and removed. (September 2007)
Main article: Philosophy of artificial intelligence
In 1950 Alan M. Turing published "Computing machinery and intelligence" in Mind, in which he
proposed that machines could be tested for intelligence using questions and answers. This process is now
named the Turing Test. The term Artificial Intelligence (AI) was first used by John McCarthy who considers it
to mean "the science and engineering of making intelligent machines".[37] It can also refer to intelligence as
exhibited by an artificial (man-made, non-natural, manufactured) entity. AI is studied in overlapping fields of
computer science, psychology, neuroscience and engineering, dealing with intelligent behavior, learning and
adaptation and usually developed using customized machines or computers.
Research in AI is concerned with producing machines to automate tasks requiring intelligent
behavior. Examples include control, planning and scheduling, the ability to answer diagnostic and consumer
questions, handwriting, natural language, speech and facial recognition. As such, the study of AI has also
become an engineering discipline, focused on providing solutions to real life problems, knowledge mining,
software applications, strategy games like computer chess and other video games. One of the biggest
difficulties with AI is that of comprehension. Many devices have been created that can do amazing things, but
critics of AI claim that no actual comprehension by the AI machine has taken place.
The debate about the nature of the mind is relevant to the development of artificial intelligence. If the
mind is indeed a thing separate from or higher than the functioning of the brain, then hypothetically it would
be much more difficult to recreate within a machine, if it were possible at all. If, on the other hand, the mind is
no more than the aggregated functions of the brain, then it will be possible to create a machine with a
recognisable mind (though possibly only with computers much different from today's), by simple virtue of the
fact that such a machine already exists in the form of the human brain.
[edit] Religious perspectives

Various religious traditions have contributed unique perspectives on the nature of mind. In many
traditions, especially mystical traditions, overcoming the ego is considered a worthy spiritual goal.
Judaism teaches that "moach shalit al halev", the mind rules the heart. Humans can approach the
Divine intellectually, through learning and behaving according to the Divine Will as enclothed in the Torah,
and use that deep logical understanding to elicit and guide emotional arousal during prayer. Christianity has
tended to see the mind as distinct from the soul (Greek nous) and sometimes further distinguished from the
spirit. Western esoteric traditions sometimes refer to a mental body that exists on a plane other than the
physical.
Hinduism's various philosophical schools have debated whether the human soul (Sanskrit atman) is
distinct from, or identical to, Brahman, the divine reality.
Buddhism posits that there is actually no distinct thing as a human being, who merely consists of five
aggregates, or skandhas. According to Buddhist philosopher Dharmakirti, mind is defined as "that which is
clarity and cognizes"—where 'clarity' refers to the formless nature of the mind and 'cognizes' to the function of
mind, namely that every mind must cognize an object.[38] The Indian philosopher-sage Sri Aurobindo
attempted to unite the Eastern and Western psychological traditions with his integral psychology, as have
many philosophers and New religious movements.
Taoism sees the human being as contiguous with natural forces, and the mind as not separate from
the body. Confucianism sees the mind, like the body, as inherently perfectible.
See also: Buddhism and psychology
[edit] Other perspectives

[edit] Parapsychology
Parapsychology is the scientific study of certain types of paranormal phenomena, or of phenomena
which appear to be paranormal.,[39] for instance precognition, telekinesis and telepathy. The term is based
on the Greek para (beside/beyond), psyche (soul/mind), and logos (account/explanation) and was coined by
psychologist Max Dessoir in or before 1889.[40] J. B. Rhine later popularized "parapsychology" as a
replacement for the earlier term "psychical research", during a shift in methodologies which brought
experimental methods to the study of psychic phenomena.[40] Parapsychology is controversial, with many
scientists believing that psychic abilities have not been demonstrated to exist.[41][42][43][44][45] The status
of parapsychology as a science has also been disputed,[46] with many scientists regarding the discipline as
pseudoscience.[47][48][49]
[edit] Memetics
Memetics is a theory of mental content based on an analogy with Darwinian evolution, which was
originated by Richard Dawkins and Douglas Hofstadter in the 1980s. It purports to be an approach to
evolutionary models of cultural information transfer. A meme, analogous to a gene, is an idea, belief, pattern
of behaviour (etc.) which is "hosted" in one or more individual minds, and which can reproduce itself from
mind to mind. Thus what would otherwise be regarded as one individual influencing another to adopt a belief
is seen memetically as a meme reproducing itself. As with genetics, particularly under Dawkins's
interpretation, a meme's success may be due its contribution to the effectiveness of its host (i.e., a the meme
is a useful, beneficial idea), or may be "selfish", in which case it could be considered a "virus of the mind".
[edit] See also

Philosophy portal
Mind and Brain portal
• Cognitive sciences
• Conscience
• Mental state
• Mental energy
• Mind at Large
• Neural Darwinism
• Subjective character of experience
• Theory of mind
• Skandha
[edit] References
1. ^ OED; etymonline.com
2. ^ Ned Block: On a Confusion about a Function of Consciousness" in: The Behavioral and
Brain Sciences, 1995.
3. ^ Churchland, Neurophilosophy
4. ^ Hart, 1996
5. ^ a b Lacey, 1996
6. ^ Boake and Diller, 2005
7. ^ Neurophilosophy, Ch. 6
8. ^ Neurophilosophy, Ch. 8
9. ^ Descartes, Description of the human body
10.^ Kim, J. (1995). Honderich, Ted. ed. Problems in the Philosophy of Mind. Oxford
Companion to Philosophy. Oxford: Oxford University Press.
11.^ James M.R. Delgado (1969). Physical control of the mind; towards a psycho civilized
society. Fitzhenry & Whiteside Limited, Toronto citebook. p. 25.
12.^ Plato (1995). E.A. Duke, W.F. Hicken, W.S.M. Nicoll, D.B. Robinson, J.C.G. Strachan. ed.
Phaedo. Clarendon Press.
13.^ Robinson, H. (1983): ‘Aristotelian dualism’, Oxford Studies in Ancient Philosophy 1, 123–44.
14.^ Nussbaum, M. C. (1984): ‘Aristotelian dualism’, Oxford Studies in Ancient Philosophy, 2,
197–207.
15.^ Nussbaum, M. C. and Rorty, A. O. (1992): Essays on Aristotle's De Anima, Clarendon
Press, Oxford.
16.^ Sri Swami Sivananda. "Sankhya:Hindu philosophy: The Sankhya".
http://www.experiencefestival.com/a/Sankhya/id/23117.
17.^ Descartes, René (1998). Discourse on Method and Meditations on First Philosophy . Hacket
Publishing Company. ISBN 0-87220-421-9.
18.^ Hart, W.D. (1996) "Dualism", in Samuel Guttenplan (org) A Companion to the Philosophy of
Mind, Blackwell, Oxford, 265–7.
19.^ Spinoza, Baruch (1670) Tractatus Theologico-Politicus (A Theologico-Political Treatise).
20.^ a b Kim, J., "Mind-Body Problem", Oxford Companion to Philosophy. Ted Honderich (ed.).
Oxford:Oxford University Press. 1995.
21.^ Pinel, J. Psychobiology, (1990) Prentice Hall, Inc. ISBN 8815071741
22.^ LeDoux, J. (2002) The Synaptic Self: How Our Brains Become Who We Are , New York:
Viking Penguin. ISBN 8870787958
23.^ Russell, Stuart J.; Norvig, Peter (2003), Artificial Intelligence: A Modern Approach (2nd
ed.), Upper Saddle River, New Jersey: Prentice Hall, ISBN 0-13-790395-2,
http://aima.cs.berkeley.edu/
24.^ Dawkins, R. The Selfish Gene (1976) Oxford:Oxford University Press. ISBN
25.^ Churchland, Patricia (1986). Neurophilosophy: Toward a Unified Science of the Mind-
Brain.. MIT Press. ISBN 0-262-03116-7.
26.^ Churchland, Paul (1981). "Eliminative Materialism and the Propositional Attitudes". Journal
of Philosophy (Journal of Philosophy, Inc.) 78 (2): 67–90. doi:10.2307/2025900.
27.^ Smart, J.J.C. (1956). "Sensations and Brain Processes". Philosophical Review.
28.^ Donald Davidson (1980). Essays on Actions and Events. Oxford University Press. ISBN 0-
19-924627-0.
29.^ Putnam, Hilary (1967). "Psychological Predicates", in W. H. Capitan and D. D. Merrill, eds.,
Art, Mind and Religion Pittsburgh: University of Pittsburgh Press.
30.^ Dennett, Daniel (1998). The intentional stance. Cambridge, Mass.: MIT Press. ISBN 0-262-
54053-3.
31.^ Searle, John (2001). Intentionality. A Paper on the Philosophy of Mind . Frankfurt a. M.:
Nachdr. Suhrkamp. ISBN 3-518-28556-4.
32.^ Confer, Easton, Fleischman, Goetz, Lewis, Perilloux & Buss, 2010; Buss, 2005; Durrant &
Ellis, 2003; Pinker, 2002; Tooby & Cosmides, 2005
33.^ The Social Brain Hypothesis
34.^ Miller. The Mating Mind. ISBN 0805857494.
35.^ "Flinn, M. V., Geary, D. C., & Ward, C. V. (2005). Ecological dominance, social competition,
and coalitionary arms races: Why humans evolved extraordinary intelligence".
http://web.missouri.edu/~gearyd/Flinnetal2005.pdf. Retrieved 2007-05-05.
36.^ "'Enhanced nutrition of offspring as a crucial factor for the evolution of intelligence on land'".
http://linkinghub.elsevier.com/retrieve/pii/S0306987704000416.
37.^ What is Artificial Intelligence? by John McCarthy Stanford University
38.^ Gyatso, Geshe Kelsang Gyatso, Understanding the Mind: The Nature and Power of the
Mind, Tharpa Publications (2nd. ed., 1997) ISBN 978-0-948006-78-4
39.^ Parapsychological Association website, Glossary of Key Words Frequently Used in
Parapsychology, Retrieved February 10, 2007
40.^ a b Encyclopedia of Occultism & Parapsychology edited by J. Gordon Melton Gale
Research, ISBN 0-8103-5487-X
41.^ Science Framework for California Public Schools. California State Board of Education.
1990.
42.^ Wheeler, J. A. (1979). "Point of View: Drive the Pseudos Out...". Skeptical Inquirer 3: 12–
13.
43.^ Kurtz, P. (1978). "Is Parapsychology a Science?". Skeptical Inquirer 3: 14–32.
44.^ Druckman, D. and Swets, J. A. eds. (1988). Enhancing Human Performance: Issues,
Theories and Techniques. National Academy Press, Washington, D.C.. p. 22. ISBN 0-309-07465-7.
45.^ Reuters (5 September 2003). "Telepathy gets academic in Sweden". CNN.
http://edition.cnn.com/2003/EDUCATION/09/05/offbeat.telepathy.reut/index.html. Retrieved 9 March
2009. "Despite decades of experimental research ... there is still no proof that gifts such as telepathy
and the ability to see the future exist, mainstream scientists say."
46.^ Flew, Antony (1982). "Parapsychology: Science or Pseudoscience?". In Grim, Patrick.
Philosophy of Science and the Occult.
47.^ Cordón, Luis A. (2005). Popular psychology: an encyclopedia . Westport, Conn: Greenwood
Press. pp. 182. ISBN 0-313-32457-3. "The essential problem is that a large portion of the scientific
community, including most research psychologists, regards parapsychology as a pseudoscience,
due largely to its failure to move beyond null results in the way science usually does. Ordinarily,
when experimental evidence fails repeatedly to support a hypothesis, that hypothesis is abandoned.
Within parapsychology, however, more than a century of experimentation has failed even to
conclusively demonstrate the mere existence of paranormal phenomenon, yet parapsychologists
continue to pursue that elusive goal."
48.^ Bunge, Mario (1991). "A skeptic's beliefs and disbeliefs". New Ideas in Psychology 9 (2):
131–149. doi:10.1016/0732-118X(91)90017-G.
49.^ Blitz, David (1991). "The line of demarcation between science and nonscience: The case of
psychoanalysis and parapsychology". New Ideas in Psychology 9: 163–170. doi:10.1016/0732-
118X(91)90020-M.

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W000
Silurian

For other uses, see Silurian (disambiguation).
The Silurian is a geologic period and system that extends
Silurian Periodfrom the end of the Ordovician Period,
about 443.7 ± 1.5 Mya (million years ago), to the
443.7 beginning
– 416 of theago
million years Devonian Period, about 416.0 ± 2.8 Mya
(ICS, 2004,[5] chart). As with other geologic periods, the rock beds that define the period's start and end are
PreЄ
well identified, but the exact dates are uncertain by several million years. The base of the Silurian is set at a
major extinction eventg when 60% of marine species were wiped out. See Ordovician-Silurian extinction
events.
Mean atmospheric O2 content over period ca. 14 Vol %[1]

duration (70 % of modern level)
Mean atmospheric CO2 content over period ca. 4500 ppm[2]

duration (16 times pre-industrial level)
Mean surface temperature over period ca. 17 °C[3]

duration (3 °C above modern level)
Around 180m, with short-term negative

Sea level (above present day)
excursions[4]
Contents
[hide]
• 1 History
• 2 Subdivisions
• 2.1 Llandovery
• 2.2 Wenlock
• 2.3 Ludlow
• 2.4 Přídolí
• 2.5 Regional stages
• 3 Geography
• 4 Climate and sea level
• 5 Perturbations
• 6 Fauna and flora
• 7 Notes
• 8 References
[edit] History
The Silurian system was first identified by British geologist Sir Roderick Impey Murchison, who was
examining fossil-bearing sedimentary rock strata in south Wales in the early 1830s. He named the
sequences for a Celtic tribe of Wales, the Silures, following the convention his friend Adam Sedgwick had
established for the Cambrian. In 1835 the two men presented a joint paper, under the title On the Silurian
and Cambrian Systems, Exhibiting the Order in which the Older Sedimentary Strata Succeed each other in
England and Wales, which was the germ of the modern geological time scale. As it was first identified, the
"Silurian" series when traced farther afield quickly came to overlap Sedgwick's "Cambrian" sequence,
however, provoking furious disagreements that ended the friendship. Charles Lapworth resolved the conflict
by defining a new Ordovician system including the contested beds. An early alternative name for the Silurian
was "Gotlandian" after the strata of the Baltic island of Gotland.
The French geologist Joachim Barrande, building on Murchison's work, used the term Silurian in a
more comprehensive sense than was justified by subsequent knowledge. He divided the Silurian rocks of
Bohemia into eight stages. His interpretation was questioned in 1854 by Edward Forbes, and the later stages
of Barrande, F, G and H, have since been shown to be Devonian. Despite these modifications in the original
groupings of the strata, it is recognized that Barrande established Bohemia as a classic ground for the study
of the earliest fossils.
[edit] Subdivisions
Key events of the Silurian

-445 —
–
-440 —
–
-435 —
–
-430 —
–
-425 —
–
-420 —
–
-415 —
Ordovician
Llandovery
Wenlock
Ludlow
Pridoli
Devonian
←
Lau event
←
Mulde event[6]
←
Ireviken
event[7]
S
i
l
u
r
i
a
n
Palæozoic
Key events of the Silurian Period.

[edit] Llandovery
The Llandovery epoch lasted from 443.7 ± 1.5 million years ago to 428.2 ± 2.3 million years ago, and
is subdivided into three stages: the Rhuddanian,[8] lasting until 439 million years ago, the Aeronian, lasting to
436 million years ago, and the Telychian. The epoch is named for the town of Llandovery in
Carmarthenshire, Wales.
[edit] Wenlock
The Wenlock, which lasted from 428.2 ± 2.3 million years ago to 422.9 ± 2.5 million years ago, is
subdivided into the Sheinwoodian (to 426.2 million years ago) and Homerian ages. It is named after the
Wenlock Edge in Shropshire, England. During the Wenlock, the oldest known tracheophytes of the genus
Cooksonia, appear. The complexity of slightly younger Gondwana plants like Baragwanathia indicates either
a much longer history for vascular plants, perhaps extending into the early Silurian or even Ordovician. See
Evolutionary history of plants.
[edit] Ludlow
The Ludlow, lasting from 422.9 ± 2.5 million years ago to 418.7 ± 2.7 million years ago, comprises
the Gorstian stage, lasting until 421.3 million years ago, and the Ludfordian stage. It is named for the town of
Ludlow in Shropshire, England.
[edit] Přídolí
The Pridoli, lasting from 418.7 ± 2.7 million years ago to 416 ± 2.8 million years ago, is the final and
shortest epoch of the Silurian. It is named after one locality at natural reserve Homolka a Přídolí near the
Prague suburb Slivenec in the Czech Republic. Přídolí is the old name of a cadastral field area.[9]
[edit] Regional stages

In North America a different suite of regional stages is sometimes used:
• Cayugan (Late Silurian - Ludlow)
• Lockportian (middle Silurian: late Wenlock)
• Tonawandan (middle Silurian: early Wenlock)
• Ontarian (Early Silurian: late Llandovery)
• Alexandrian (earliest Silurian: early Llandovery)
[edit] Geography
Ordovician-Silurian boundary exposed on Hovedøya, Norway, showing the very marked difference
between the light gray Ordovician calcareous sandstone and brown Silurian mudstone. The layers have been
inverted (overturned) by the Caledonian orogeny.
With the supercontinent Gondwana covering the equator and much of the southern hemisphere, a
large ocean occupied most of the northern half of the globe.[10] The high sea levels of the Silurian and the
relatively flat land (with few significant mountain belts) resulted in a number of island chains, and thus a rich
diversity of environmental settings.[10]
During the Silurian, Gondwana continued a slow southward drift to high southern latitudes, but there
is evidence that the Silurian icecaps were less extensive than those of the late Ordovician glaciation. The
southern continents remained united during this period. The melting of icecaps and glaciers contributed to a
rise in sea level, recognizable from the fact that Silurian sediments overlie eroded Ordovician sediments,
forming an unconformity. The continents of Avalonia, Baltica, and Laurentia drifted together near the equator,
starting the formation of a second supercontinent known as Euramerica.
Fossilised Late Silurian shallow sea floor, on display in Bristol City Museum, Bristol, England. From
the Wenlock epoch, in the Wenlock limestone, Dudley, West Midlands, England.
When the proto-Europe collided with North America, the collision folded coastal sediments that had
been accumulating since the Cambrian off the east coast of North America and the west coast of Europe.
This event is the Caledonian orogeny, a spate of mountain building that stretched from New York State
through conjoined Europe and Greenland to Norway. At the end of the Silurian, sea levels dropped again,
leaving telltale basins of evaporites in a basin extending from Michigan to West Virginia, and the new
mountain ranges were rapidly eroded. The Teays River, flowing into the shallow mid-continental sea, eroded
Ordovician strata, leaving traces in the Silurian strata of northern Ohio and Indiana.
The vast ocean of Panthalassa covered most of the northern hemisphere. Other minor oceans
include two phases of the Tethys— the Proto-Tethys and Paleo-Tethys— the Rheic Ocean, a seaway of the
Iapetus Ocean (now in between Avalonia and Laurentia), and the newly formed Ural Ocean.
[edit] Climate and sea level

The Silurian period enjoyed relatively stable and warm temperatures, in contrast the extreme
glaciations of the Ordovician before it, and the extreme heat of the ensuing Devonian.[10] Sea levels rose
from their Hirnantian low throughout the first half of the Silurian; they subsequently fell throughout the rest of
the period, although smaller scale patterns are superimposed on this general trend; fifteen high-stands can
be identified, and the highest Silurian sea level was probably around 140 m higher than the lowest level
reached.[10]
During this period, the Earth entered a long warm greenhouse phase, and warm shallow seas
covered much of the equatorial land masses. Early in the Silurian, glaciers retreated back into the South Pole
until they almost disappeared in the middle of Silurian. The period witnessed a relative stabilization of the
Earth's general climate, ending the previous pattern of erratic climatic fluctuations. Layers of broken shells
(called coquina) provide strong evidence of a climate dominated by violent storms generated then as now by
warm sea surfaces. Later in the Silurian, the climate cooled slightly, but in the Silurian-Devonian boundary,
the climate became warmer.
[edit] Perturbations
End of Silurian extinction. Click on the picture for more information

The climate and carbon cycle appears to be rather unsettled during the Silurian, which has a higher
concentration of isotopic excursions than any other period.[10] The Ireviken event, Mulde event and Lau
event each represent isotopic excursions following a minor mass extinction[11] and associated with rapid
sea-level change, in addition to the larger extinction at the end of the Silurian.[10] Each home leaves a
similar signature in the geological record, both geochemically and biologically; pelagic (free-swimming)
organisms were particularly hard hit, as were brachiopods, corals and trilobites, and extinctions rarely occur
in a rapid series of fast bursts.[10]
[edit] Fauna and flora
Artist's impression of Silurian fish

The first bony fish, the Osteichthyes, appeared, represented by the Acanthodians covered with bony
scales; fish reached considerable diversity and developed movable jaws, adapted from the supports of the
front two or three gill arches. A diverse fauna of Eurypterids (Sea Scorpions) -- some of them several meters
in length—prowled the shallow Silurian seas of North America; many of their fossils have been found in New
York State. Leeches also made their appearance during the Silurian Period. Brachiopods, bryozoa, molluscs,
hederelloids and trilobites were abundant and diverse.
Reef abundance was patchy; sometimes they were everywhere, but at other points they are virtually
absent from the rock record.[10]
Cooksonia, the earliest vascular plant, middle Silurian
The Silurian was the first period to see macrofossils of extensive terrestrial biota, in the form of moss
forests along lakes and streams. However, the land fauna did not have a major impact on the Earth until it
diversified in the Devonian.[10]
The first fossil records of vascular plants, that is, land plants with tissues that carry food, appeared in
the second half of the Silurian period. The earliest known representatives of this group are the Cooksonia
(mostly from the northern hemisphere) and Baragwanathia (from Australia). A primitive Silurian land plant
with xylem and phloem but no differentiation in root, stem or leaf, was much-branched Psilophyton,
reproducing by spores and breathing through stomata on every surface, and probably photosynthesizing in
every tissue exposed to light. Rhyniophyta and primitive lycopods were other land plants that first appear
during this period. Neither mosses nor the earliest vascular plants had deep roots. Silurian rocks often have a
brownish tints, possibly a result of extensive erosion of the early soils.
Some evidence suggests the presence of predatory trigonotarbid arachnoids and myriapods in Late
Silurian facies. Predatory invertebrates would indicate that simple food webs were in place that included non-
predatory prey animals. Extrapolating back from Early Devonian biota, Andrew Jeram et al. in 1990[12]
suggested a food web based on as yet undiscovered detritivores and grazers on microorganisms.[13]
[edit] Notes
4. ^ Haq, B. U.; Schutter, SR (2008). "A Chronology of Paleozoic Sea-Level Changes". Science
322 (5898): 64–68. doi:10.1126/science.1161648. PMID 18832639.
5. ^ Gradstein, Felix M.; Ogg, J. G.; Smith, A. G. (2004). A Geologic Time Scale 2004.
6. ^ Jeppsson, L.; Calner, M. (2007). "The Silurian Mulde Event and a scenario for secundo—
secundo events". Earth and Environmental Science Transactions of the Royal Society of Edinburgh
93 (02): 135-154. doi:10.1017/S0263593300000377.
7. ^ Munnecke, A.; Samtleben, C.; Bickert, T. (2003). "The Ireviken Event in the lower Silurian
of Gotland, Sweden-relation to similar Palaeozoic and Proterozoic events". Palaeogeography,
Palaeoclimatology, Palaeoecology 195 (1): 99-124. doi:10.1016/S0031-0182(03)00304-3.
8. ^ Named for the Cefn-Rhuddan Farm in the Llandovery area; confusingly, Rhuddlan lies on
Silurian strata as well.
9. ^ Štěpán Manda, Jiří Frýda: Silurian-Devonian boundary events and their influence on
cephalopod evolution: evolutionary significance of cephalopod egg size during mass extinctions. In:
Bulletin of Geoscience. Vol. 85 (2010) Heft 3, S. 513-540
10.^ a b c d e f g h i Munnecke, A.; Calner, M.; Harper, D. A. T.; Servais, T. (2010). "Ordovician
and Silurian sea-water chemistry, sea level, and climate: A synopsis". Palaeogeography,
Palaeoclimatology, Palaeoecology 296: 389. doi:10.1016/j.palaeo.2010.08.001. edit
11.^ Samtleben, C.; Munnecke, A.; Bickert, T. (2000). "Development of facies and C/O-isotopes
in transects through the Ludlow of Gotland: Evidence for global and local influences on a shallow-
marine environment". Facies 43: 1. doi:10.1007/BF02536983. edit
12.^ Andrew J. Jeram, Paul A. Selden and Dianne Edwards, "Land Animals in the Silurian:
Arachnids and Myriapods from Shropshire, England", Science 2 November 1990:658-61.
13.^ Anna K. Behrensmeyer, John D. Damuth, et al. Terrestrial Ecosystems Through Time
"Paleozoic Terrestrial Ecosystems" (University of Chicago Press), 1992:209.
[edit] References
• Emiliani, Cesare. (1992). Planet Earth : Cosmology, Geology, & the Evolution of Life & the
Environment. Cambridge University Press. (Paperback Edition ISBN 0-521-40949-7)
• Mikulic, DG, DEG Briggs, and J Kluessendorf. 1985. A new exceptionally preserved biota
from the Lower Silurian of Wisconsin, USA. Philosophical Transactions of the Royal Society of
London, 311B:75-86.
• Moore, RA, DEG Briggs, SJ Braddy, LI Anderson, DG Mikulic, and J Kluessendorf. 2005. A
new synziphosurine (Chelicerata: Xiphosura) from the Late Llandovery (Silurian) Waukesha
Lagerstatte, Wisconsin, USA. Journal of Paleontology:79(2), pp. 242–250.
• Ogg, Jim; June, 2004, Overview of Global Boundary Stratotype Sections and Points
(GSSP's) http://www.stratigraphy.org/gssp.htm Accessed April 30, 2006.

Wikimedia Commons has media related to: Silurian
• Palaeos: Silurian
• UCMP Berkeley: The Silurian
• Paleoportal: Silurian strata in U.S., state by state
• USGS:Silurian and Devonian Rocks (U.S.)
• "International Commission on Stratigraphy (ICS)" . Geologic Time Scale 2004.
http://www.stratigraphy.org/. Retrieved September 19, 2005.
• Examples of Silurian Fossils
• GeoWhen Database for the Silurian
Preceded
by Proterozoic Eon
Hadean

Proterozoic Paleoproterozoic (2.5 – 1.6 Gya): Siderian (2.5 – 2.3 Gya) · Rhyacian (2.3 –
(2.5 Gya – 542 Mya) 2.05 Gya) · Orosirian (2.05 – 1.8 Gya) · Statherian (1.8 – 1.6 Gya)
Mesoproterozoic (1.6 – 1 Gya): Calymmian (1.6 – 1.4 Gya) · Ectasian (1.4 – 1.2
Gya) · Stenian (1.2 – 1 Gya)
[hide] Paleozoic (542 – 251 Mya)
521 Mya)
Cambrian Epoch 2* (521 – 510 Mya): Age 3* (521 – 515 Mya) · Age 4* (515 – 510 Mya)
(542 – 488.3 Mya) Epoch 3* (510 – 499 Mya): Age 5* (510 – 506.5 Mya) · Drumian (506.5 – 503
Age 10* (492 – 488.3 Mya)

Floian (478.6 – 471.8 Mya)

Silurian
(426.2 – 422.9 Mya)
(443.7 – 416 Mya)
418.7 Mya)

(416 – 359.2 Mya) (391.8 – 385.3 Mya)
(374.5 – 359.2 Mya)
Carboniferous
(359.2 – 299 Mya)
(303.4 – 299 Mya)
Cisuralian (299 – 270.6 Mya): Asselian (299 – 294.6 Mya) · Sakmarian

(294.6 – 284.4 Mya) · Artinskian (284.4 – 275.6 Mya) · Kungurian (275.6 – 270.6
Mya)
Permian
(299 – 251 Mya)
Triassic
(249.5 – 245.9 Mya)
(251 – 199.6 Mya)
Middle Triassic (245.9 – 228.7 Mya): Anisian (245.9 – 237 Mya) · Ladinian (237 –
228.7 Mya)
203.6 Mya) · Rhaetian (203.6 – 199.6 Mya)

– 175.6 Mya)
Mya)


Mya)
Miocene (23.03 – 5.332 Mya): Aquitanian (23.03 – 20.43 Mya) · Burdigalian

Neogene
(23.03 – 2.588 Mya)
Mya)
Quaternary Pleistocene (2.588 Mya – 11.4 kya): Gelasian (2.588 – 1.806 Mya) ·
(2.588 – 0 Mya) Calabrian (1.806 Mya – 781 kya) · Middle Pleistocene / Ionian (781 – 126 kya) · Late
Dryas* (12.8 – 11.4 kya)

February 2008.
Retrieved from "http://en.wikipedia.org/wiki/Silurian"
Categories: Silurian
W000
Recent African origin of modern humans


Map of early human migrations according to mitochondrial population genetics
In paleoanthropology, the recent African origin of modern humans is the mainstream model
describing the origin and early dispersal of anatomically modern humans. The theory is called the (Recent)
Out-of-Africa model in the popular press, and academically the recent single-origin hypothesis (RSOH),
Replacement Hypothesis, and Recent African Origin (RAO) model. The hypothesis that humans have a
single origin (monogenesis) was published in Charles Darwin's Descent of Man (1871). The concept was
speculative until the 1980s, when it was corroborated by a study of present-day mitochondrial DNA,
combined with evidence based on physical anthropology of archaic specimens. According to genetic and
fossil evidence, archaic Homo sapiens evolved to anatomically modern humans solely in Africa, between
200,000 and 150,000 years ago.
Members of one branch leaving Africa by between 125,000 and 60,000 years ago and over time
replacing earlier human populations such as Neanderthals and Homo erectus. The date of this original "out
of Africa" migration has mostly been dated to between 70,000 and 60,000 years ago, but a 2011 study
suggested that an early wave of migration may have taken place as early as 125,000 years ago.
The recent single origin of modern humans in East Africa is the near-consensus position held within
the scientific community.[1][2][3][4][5]
However, recent sequencing of the full Neanderthal Genome suggests Neanderthals and some
modern humans share some ancient genetic lineages. The authors of the study suggest that their findings
are consistent with Neanderthal admixture of up to 4% in some populations. But the study also suggests that
there may be other reasons why humans and Neanderthals share ancient genetic lineages.[6][7]
The competing hypothesis is the multiregional origin of modern humans. Some push back the
original "out of Africa" migration—in this case, by Homo erectus, not by Homo sapiens—to two million years
ago.[4][8]
Contents
[hide]
• 1 History of the theory
• 2 Early Homo sapiens
• 3 Genetic reconstruction
• 3.1 Mitochondrial DNA
• 3.2 Y-chromosomal DNA
• 3.3 Genomic analysis
• 4 Exodus from Africa
• 5 Subsequent expansion
• 6 Competing hypotheses
• 7 See also
• 8 References
History of the theory

With the development of anthropology in the early 19th century, scholars disagreed vigorously about
different theories of human development. Those such as Johann Friedrich Blumenbach and James Cowles
Pritchard held that since the creation, the various human races had developed as different varieties sharing
descent from one people (monogenism). Their opponents, such as Louis Agassiz and Josiah C. Nott, argued
for polygenism, or the separate development of human races as separate species, or had developed as
separate species through transmutation of species from apes, with no common ancestor.
Charles Darwin was one of the first to propose common descent of living organisms, and among the
first to suggest that all humans had in common ancestors who lived in Africa. In the Descent of Man, he
speculated that humans had descended from apes which still had small brains but walked upright, freeing
their hands for uses which favoured intelligence. Further, he thought such apes were African:[9]
In each great region of the world the living mammals are closely related to the extinct species of
the same region. It is, therefore, probable that Africa was formerly inhabited by extinct apes
closely allied to the gorilla and chimpanzee; and as these two species are now man's nearest
allies, it is somewhat more probable that our early progenitors lived on the African continent than
elsewhere. But it is useless to speculate on this subject, for an ape nearly as large as a man,
namely the Dryopithecus of Lartet, which was closely allied to the anthropomorphous Hylobates,
existed in Europe during the Upper Miocene period; and since so remote a period the earth has
certainly undergone many great revolutions, and there has been ample time for migration on the
largest scale.
—Charles Darwin, Descent of Man[10]
The prediction was insightful because at the time, in 1871, there were hardly any human fossils of
ancient hominids available. Almost fifty years later, Darwin's speculation was supported when
anthropologists began finding numerous fossils of ancient small-brained hominids in several areas of Africa
(list of hominina fossils).
The debate in anthropology had swung in favour of monogenism by the mid-20th century. Isolated
proponents of polygenism held forth in the mid-20th century, such as Carleton Coon, who hypothesized as
late as 1962 that Homo sapiens arose five separate times from Homo erectus in five separate places.[11]
The "Recent African origin" of modern humans means "single origin" (monogenism) and has been used in
various contexts as an antonym to polygenism.
With the advent of archaeogenetics in the 1990s, scientists were able to date the "out of Africa"
migration with some confidence. In 2000, the mitochondrial DNA (mtDNA) sequence of "Mungo Man" of
ancient Australia was published.[12] This work was later questioned.[13]
and explained by W. James Peacock, leader of the team who sequenced Mungo man's ancient
mtdna.[14][dead link]
The question of whether there was inheritance of other typological (not de facto) Homo subspecies
into the Homo sapiens genetic pool remains under debate.
Early Homo sapiens

Main articles: Anatomically modern humans and Archaic Homo sapiens
Anatomically modern humans originated in Africa about 250,000 years ago. The trend in cranial
expansion and the acheulean elaboration of stone tool technologies which occurred between 400,000 years
ago and the second interglacial period in the Middle Pleistocene (around 250,000 years ago) provide
evidence for a transition from H. erectus to H. sapiens.[citation needed] In the Recent African Origin (RAO)
scenario, migration within and out of Africa eventually replaced the earlier dispersed H. erectus.
Homo sapiens idaltu, found at site Middle Awash in Ethiopia, lived about 160,000 years ago.[15] It is
the oldest known anatomically modern human and classified as an extinct subspecies.[16] Fossils of early
Homo sapiens were found in Qafzeh cave in Israel and have been dated to 80,000 to 100,000 years ago.
However these humans seem to have either become extinct or retreated back to Africa 70,000 to 80,000
years ago, possibly replaced by south bound Neanderthals escaping the colder regions of ice age Europe.
[citation needed] Hua Liu & al. analyzing autosomal microsatellite markers dates to c. 56,000±5,700 years
ago mtDNA evidence. He interprets the paleontological fossil of early modern human from Qafzeh cave as
an isolated early offshoot that retracted back to Africa.[1]
All other fossils of fully modern humans outside Africa have been dated to more recent times. The
oldest well dated fossils found outside Africa are from Lake Mungo, Australia, and have been dated to about
42,000 years ago.[17][18] The Tianyuan cave remains in Liujiang region China have a probable date range
between 38,000 and 42,000 years ago. They are most similar in morphology to Minatogawa Man, modern
humans dated between 17,000 and 19,000 years ago and found on Okinawa Island, Japan.[19][20] However,
others have dated Liujang Man to 111,000 to 139,000 years before the present.[21]
Beginning about 100,000 years ago evidence of more sophisticated technology and artwork begins
to emerge and by 50,000 years ago fully modern behaviour becomes more prominent. Stone tools show
regular patterns that are reproduced or duplicated with more precision while tools made of bone and antler
appear for the first time.[22][23]
Genetic reconstruction
Further information: Archaeogenetics and Early human migrations
Two pieces of the human genome are quite useful in deciphering human history: mitochondrial DNA
and the Y chromosome. These are the only two parts of the genome that are not shuffled about by the
evolutionary mechanisms that generate diversity with each generation: instead, these elements are passed
down intact. According to the hypothesis, all people alive today have inherited the same Mitochondria[24]
from one woman who lived in Africa about 160,000 years ago[25][26] she has been named Mitochondrial
Eve. All men today have inherited their Y chromosomes from a man who lived 60,000 years ago, probably in
Africa. He has been named Y-chromosomal Adam. It is now believed that more men participated in the out of
Africa exodus of early humans than women based on comparing non-sex-specific chromosomes with sex-
specific ones.[27]
Mitochondrial DNA
Further information: Human mitochondrial DNA haplogroup
One model of human migration based on Mitochondrial DNA
The first lineage to branch off from Mitochondrial Eve is L0. This haplogroup is found in high
proportions among the San of Southern Africa, the Sandawe of East Africa. It is also found among the Mbuti
people.[28][29]
These groups branched off early in human history and have remained relatively genetically isolated
since then. Haplogroups L1, L2 and L3 are descendents of L1-6 and are largely confined to Africa. The
macro haplogroups M and N, which are the lineages of the rest of the world outside Africa, descend from L3.
Y-chromosomal DNA
Main articles: Human Y-chromosome DNA haplogroups, Haplogroup CT (Y-DNA), and Haplogroup F
(Y-DNA)
The mutations defining macro-haplogroup CT (all Y haplogroups except A and B) predate the "Out of
Africa" migration, its descendent macro-group DE being confined to Africa. The mutations that distinguish
Haplogroup C from all other descendants of CR have occurred some 60,000 years ago, shortly after the first
Out of Africa migration.
Haplogroup F originated some 45,000 years ago, either in North Africa (in which case it would point
to a second wave of out-of-Africa migration) or in South Asia. More than 90% of males not native to Africa are
descended in direct male line from the first bearer of haplogroup F.
Genomic analysis
Although mitochondrial DNA and Y-chromosomal DNA are particularly useful in deciphering human
history, data on the genomes of dozens of population groups have also been studied. In June 2009, an
analysis of genome-wide SNP data from the International HapMap Project (Phase II) and CEPH Human
Genome Diversity Panel samples was published.[30] Those samples were taken from 1138 unrelated
individuals.[30] Before this analysis, population geneticists expected to find dramatic differences among
ethnic groups, with derived alleles shared among such groups but uncommon or nonexistent in other groups.
[31] Instead the study of 53 populations taken from the HapMap and CEPH data revealed that the population
groups studied fell into just three genetic groups: Africans, Eurasians (which includes natives of Europe and
the Middle East, and Southwest Asians east to present-day Pakistan), and East Asians, which includes
natives of Asia, Japan, Southeast Asia, the Americas, and Oceania.[31] The study determined that most
ethnic group differences can be attributed to genetic drift, with modern African populations having greater
genetic diversity than the other two genetic groups, and modern Eurasians somewhat more than modern
East Asians.[31] The study suggested that natural selection may shape the human genome much more
slowly than previously thought, with factors such as migration within and among continents more heavily
influencing the distribution of genetic variations.[32]
Exodus from Africa
Red Sea crossing

By some 70 millennia ago, a part of the bearers of mitochondrial haplogroup L3 migrated from East
Africa into the Near East. The date of this first wave of "out of Africa" migration has been called into question
in 2011, based on the discovery of stone tools in the United Arab Emirates, indicating the presence of
modern humans between 100,000 and 125,000 years ago.[33]
Some scientists believe that only a few people left Africa in a single migration that went on to
populate the rest of the world,[34] based in the fact that only descents of L3 are found outside Africa. From
that settlement, some others point to the possibility of several waves of expansion. For example, Wells says
that the early travelers followed the southern coastline of Asia, crossed about 250 kilometers [155 miles] of
sea, and colonized Australia by around 50,000 years ago. The Aborigines of Australia, Wells says, are the
descendants of the first wave of migrations.[35]
It has been estimated that from a population of 2,000 to 5,000 individuals in Africa[36], only a small
group of possibly as little as 150 to 1,000 people crossed the Red Sea[37]. Of all the lineages present in
Africa only the female descendants of one lineage, mtDNA haplogroup L3, are found outside Africa. Had
there been several migrations one would expect descendants of more than one lineage to be found outside
Africa. L3's female descendants, the M and N haplogroup lineages, are found in very low frequencies in
Africa (although haplogroup M1 is very ancient and diversified in North and Northeast Africa) and appear to
be recent arrivals. A possible explanation is that these mutations occurred in East Africa shortly before the
exodus and by the founder effect became the dominant haplogroups after the exodus from Africa.
Alternatively, the mutations may have arisen shortly after the exodus from Africa.
Other scientists have proposed a Multiple Dispersal Model, in which there were two migrations out of
Africa, one across the Red Sea travelling along the coastal regions to India (the Coastal Route), which would
be represented by Haplogroup M. Another group of migrants with Haplogroup N followed the Nile from East
Africa, heading northwards and crossing into Asia through the Sinai. This group then branched in several
directions, some moving into Europe and others heading east into Asia. This hypothesis attempts to explain
why Haplogroup N is predominant in Europe and why Haplogroup M is absent in Europe. Evidence of the
coastal migration is hypothesized to have been destroyed by the rise in sea levels during the Holocene
epoch.[38][39] Alternatively, a small European founder population that initially expressed both Haplogroup M
and N could have lost Haplogroup M through random genetic drift resulting from a bottleneck (i.e. a founder
effect).
Today at the Bab-el-Mandeb straits the Red Sea is about 12 miles (20 kilometres) wide, but 50,000
years ago it was much narrower and sea levels were 70 meters lower. Though the straits were never
completely closed, there may have been islands in between which could be reached using simple rafts. Shell
middens 125,000 years old have been found in Eritrea,[40] indicating the diet of early humans included
seafood obtained by beachcombing.
Subsequent expansion
Main article: Early human migrations
Map of early human migrations[41]

1. Homo sapiens
2. Neanderthals
3. Early Hominids
From the Near East, these populations spread east to South Asia by 50,000 years ago, and on to
Australia by 40,000 years ago, Homo sapiens for the first time colonizing territory never reached by Homo
erectus. Europe was reached by Cro-Magnon some 40,000 years ago. East Asia (Korea, Japan) was
reached by 30,000 years ago. It is disputed whether subsequent migration to North America took place
around 30,000 years ago, or only considerably later, around 14,000 years ago.
The group that crossed the Red Sea travelled along the coastal route around the coast of Arabia and
Persia until reaching India, which appears to be the first major settling point. M is found in high frequencies
along the southern coastal regions of Pakistan and India and it has the greatest diversity in India, indicating
that it is here where the mutation may have occurred.[42] Sixty percent of the Indian population belong to
Haplogroup M. The indigenous people of the Andaman Islands also belong to the M lineage. The
Andamanese are thought to be offshoots of some of the earliest inhabitants in Asia because of their long
isolation from mainland Asia. They are evidence of the coastal route of early settlers that extends from India
along the coasts of Thailand and Indonesia all the way to Papua New Guinea. Since M is found in high
frequencies in highlanders from New Guinea as well, and both the Andamanese and New Guineans have
dark skin and Afro-textured hair, some scientists believe they are all part of the same wave of migrants who
departed across the Red Sea ~60,000 years ago in the Great Coastal Migration. Notably, the findings of
Harding et al. (2000, p. 1355) show that, at least with regard to dark skin color, the haplotype background of
Papua New Guineans at MC1R (one of a number of genes involved in melanin production) is identical to that
of Africans (barring a single silent mutation). Thus, although these groups are distinct from Africans at other
loci (due to drift, bottlenecks, etc), it is evident that selection for the dark skin color trait likely continued (at
least at MC1R) following the exodus. This would support the hypothesis that suggests that the original
migrants from Africa resembled pre-exodus Africans (at least in skin color), and that the present day
remnants of this ancient phenotype can be seen among contemporary Africans, Andamanese and New
Guineans. Others suggest that their physical resemblance to Africans could be the result of convergent
evolution.[43][44]
From Arabia to India the proportion of haplogroup M increases eastwards: in eastern India, M
outnumbers N by a ratio of 3:1. However, crossing over into East Asia, Haplogroup N reappears as the
dominant lineage. M is predominant in South East Asia but amongst Indigenous Australians N reemerges as
the more common lineage. This discontinuous distribution of Haplogroup N from Europe to Australia can be
explained by founder effects and population bottlenecks.[45]
Competing hypotheses
Main article: Multiregional origin of modern humans
The multiregional hypothesis, initially proposed by Milford Wolpoff, holds that the evolution of
humans from H. erectus at the beginning of the Pleistocene 1.8 million years BP to the present day has been
within a single, continuous worldwide population. Proponents of multiregional origin reject the assumption of
an infertility barrier between ancient Eurasian and African populations of Homo. Multiregional proponents
point to the fossil record and genetic evidence in chromosomal DNA. One study suggested that at least 5% of
the human modern gene pool can be attributed to ancient admixture, which in Europe would be from the
Neanderthals.[46]
A recently discovered fossilized mandible that is putatively a hybrid between Homo sapiens and an
earlier hominid, that is likely to be 110,000 years old, has been interpreted as a challenge to the recent out-
of-Africa hypothesis. However, some scholars doubt that the fossil represents a Homo sapiens hybrid.[47]
See also
• Archaeogenetics of • Hofmeyr Molecular
the Near East Skull Anthropology portal
• Behavioral • Identical
modernity ancestors point Evolutionary
• Early human • Mitochondri biology portal
migrations al Eve
• Genetic history of • Most recent
the British Isles common ancestor
• Genetic history of • Multiregiona
indigenous peoples of the l hypothesis
Americas • Sahara
• Genetic history of pump theory
Europe • Toba
• Genetic history of catastrophe theory
Italy • Y-
• Genetics and chromosomal Adam
archaeogenetics of South
Asia
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Further reading
• Bowler, Peter J. (2003). Evolution: The History of an Idea (3rd ed.). Berkeley: University of
California Press. ISBN 0-520-23693-9. http://books.google.ca/books?
id=e2b5B0po8fwC&lpg=PP1&dq=Evolution%3A%20The%20History%20of%20an
%20Idea&pg=PP1#v=onepage&q&f=true.
• Darwin, Charles (1871). The Descent of Man, and Selection in Relation to Sex (1st ed.).
London: John Murray. http://darwin-
online.org.uk/EditorialIntroductions/Freeman_TheDescentofMan.html. Retrieved 2009-09-05
• Gibbons A (May 2001). "Human anthropology. Modern men trace ancestry to African
migrants". Science 292 (5519): 1051–2. doi:10.1126/science.292.5519.1051b. PMID 11352048.
• Underhill PA, Passarino G, Lin AA, Shen P, Mirazón Lahr M, Foley RA, Oefner PJ, Cavalli-
Sforza LL (January 2001). "The phylogeography of Y chromosome binary haplotypes and the origins
of modern human populations". Ann. Hum. Genet. 65 (Pt 1): 43–62. PMID 11415522.
http://www.human-evol.cam.ac.uk/Members/Lahr/pubs/AHG-65-01.pdf.
• Neanderthals 'mated with modern humans', BBC News, 21 April 1999
• New analysis shows three human migrations out of Africa - Replacement theory
'demolished', Washington University in St. Louis, 2 February 2006
• Harding RM, Healy E, Ray AJ, Ellis NS, Flanagan N, Todd C, Dixon C, Sajantila A, Jackson
IJ, Birch-Machin MA, Rees JL (April 2000). "Evidence for variable selective pressures at MC1R". Am.
J. Hum. Genet. 66 (4): 1351–61. doi:10.1086/302863. PMID 10733465.
• Long JC, Kittles RA (August 2003). "Human genetic diversity and the nonexistence of
biological races". Hum. Biol. 75 (4): 449–71. PMID 14655871.
http://muse.jhu.edu/journals/human_biology/v075/75.4long.pdf.
• Risch, N., Burchard, E., Ziv, E. and Tang, H. (2002). "Categorization of humans in biomedical
research: genes, race and disease". Genome Biology 3 (7): comment2007.2001 -
comment2007.2012. doi:10.1186/gb-2002-3-7-comment2007. PMID 12184798.
• Tishkoff SA, Kidd KK (November 2004). "Implications of biogeography of human populations
for 'race' and medicine". Nat. Genet. 36 (11 Suppl): S21–7. doi:10.1038/ng1438. PMID 15507999.
• Cavalli-Sforza F, Cavalli-Sforza LL (1995). The great human diasporas: the history of
diversity and evolution. Boston: Addison-Wesley. ISBN 0-201-44231-0.
• Crow TJ, ed (2004). The Speciation of Modern Homo sapiens (Proceedings of the British
Academy). London: British Academy. ISBN 0-19-726311-9.
• Foley R (1995). Humans before humanity: an evolutionary perspective . Oxford: Blackwell.
ISBN 0-631-20528-4.
• Olson S (2003). Mapping human history: genes, race, and our common origins . Boston:
Houghton Mifflin. ISBN 0-618-35210-4.
• Oppenheimer, Stephen (2003). The Real Eve: Modern Man's Journey Out of Africa . New
York, NY: Carroll & Graf. ISBN 0-7867-1192-2.
• McKie R, Stringer C (1997). African exodus: the origins of modern humanity . London:
Pimlico. ISBN 0-7126-7307-5.
• Sykes, Bryan (2004). The Seven Daughters of Eve: The Science That Reveals Our Genetic
Ancestry. Corgi Adult. ISBN 0-552-15218-8.
• Wade N (2006). Before the Dawn : Recovering the Lost History of Our Ancestors . Penguin
Press HC, The. ISBN 1-59420-079-3.
• Wells S (2004). Journey of Man: Genetic Odyssey. Randoum House. ISBN 0-8129-7146-9.
• Wells, Spencer (2006). Deep ancestry: inside the Genographic Project. Washington, D.C:
National Geographic. ISBN 0-7922-6215-8.
• Manica A, Amos W, Balloux F, Hanihara T (July 2007). "The effect of ancient population
bottlenecks on human phenotypic variation". Nature 448 (7151): 346–8. doi:10.1038/nature05951.
PMID 17637668. Lay summary – Science Daily.
• Scholz CA, Johnson TC, Cohen AS, King JW, Peck JA, Overpeck JT, Talbot MR, Brown ET,
Kalindekafe L, Amoako PY, Lyons RP, Shanahan TM, Castañeda IS, Heil CW, Forman SL,
McHargue LR, Beuning KR, Gomez J, Pierson J (October 2007). "East African megadroughts
between 135 and 75 thousand years ago and bearing on early-modern human origins". Proc. Natl.
Acad. Sci. U.S.A. 104 (42): 16416–21. doi:10.1073/pnas.0703874104. PMID 17785420. Lay
summary – Science Daily.
• Cohen AS, Stone JR, Beuning KR, Park LE, Reinthal PN, Dettman D, Scholz CA, Johnson
TC, King JW, Talbot MR, Brown ET, Ivory SJ (October 2007). "Ecological consequences of early
Late Pleistocene megadroughts in tropical Africa". Proc. Natl. Acad. Sci. U.S.A. 104 (42): 16422–7.
doi:10.1073/pnas.0703873104. PMID 17925446. Lay summary – Science Daily.
• Smith L (2007-10-09). "Climate change led mankind out of Africa". Times Online.
http://www.timesonline.co.uk/tol/news/world/africa/article2617296.ece.
• Russell S (2008-02-22). "DNA studies trace human migration from Africa". San Francisco
Chronicle. http://www.sfgate.com/cgi-bin/article.cgi?f=/c/a/2008/02/22/MN5RV6L1C.DTL.
• Serre D, Langaney A, Chech M, Teschler-Nicola M, Paunovic M, Mennecier P, Hofreiter M,
Possnert G, Pääbo S (March 2004). "No evidence of Neandertal mtDNA contribution to early modern
humans". PLoS Biol. 2 (3): E57. doi:10.1371/journal.pbio.0020057. PMID 15024415.
External links
• An mtDNA view of the peopling of the world by Homo sapiens
• National Geographic: Atlas of the Human Journey
• Bradshaw Foundation: The Journey of Mankind
Documentaries
• DNA Mysteries – The Search for Adam - by Spencer Wells - National Geographic, 2008
• The Real Eve: Modern Man's Journey Out of Africa - by Stephen Oppenheimer - Discovery
Channel, 2002
• Journey of Man: A Genetic Odyssey (movie) by Spencer Wells – PBS and National
Geographic Channel, 2003
Hominini
Australopithecines


Retrieved from "http://en.wikipedia.org/wiki/Recent_African_origin_of_modern_humans"
Categories: Recent single origin hypothesis | Human evolution | Hypotheses | Genetic genealogy |
Race | Population genetics | Molecular evolution | Paleolithic | Prehistoric Africa
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Evolutionary ecology

A phylogenetic tree of living things.
Evolutionary ecology lies at the intersection of ecology and evolutionary biology. It approaches the
study of ecology in a way that explicitly considers the evolutionary histories of species and the interactions
between them. Conversely, it can be seen as an approach to the study of evolution that incorporates an
understanding of the interactions between the species under consideration. The main subfields of
evolutionary ecology are life history evolution, sociobiology (the evolution of behavior), the evolution of
interspecific relations (cooperation, predator-prey interactions, parasitism, mutualism) and the evolution of
biodiversity and of communities.
[edit] Evolutionary ecologists
Julia Margaret Cameron’s portrait of Darwin

• Charles Darwin
• George Evelyn Hutchinson
• Robert MacArthur
• Eric Pianka
• Michael Rosenzweig
[edit] References
This article includes a list of references, related reading or external links, but its
sources remain unclear because it lacks inline citations. Please improve this article by
introducing more precise citations where appropriate. (April 2010)
• Fox, C.W., Roff, D.A. and Fairbairn, D.J. 2001. Evolutionary Ecology: Concepts and Case
Studies. Oxford University Press.
• Mayhew, P.J. 2006. Discovering Evolutionary Ecology: Bringing Together Ecology and
Evolution. Oxford University Press.
• Pianka, E.R. 2000. Evolutionary Ecology, 6th ed. Benjamin Cummings.

• Evolutionary Ecology Research - a journal in the field.
• Methods in Ecology and Evolution - a journal the field.
• Main issues of Evolutionary Ecology
[hide]v · d · eEvolutionary ecology
Patterns of
evolution Convergent evolution · Parallel evolution · Divergent evolution
Signals Antipredator adaptation · Aposematism · Mimicry · Crypsis

(Countershading) · Unkenreflex
[hide] Modelling ecosystems - trophic components
Abiotic component · Abiotic stress ·

Behaviour · Biogeochemical cycle · Biomass ·
Biotic component · Biotic stress · Carrying
General capacity · Competition · Ecosystem · Ecosystem
ecology · Ecosystem model · Keystone species ·
List of feeding behaviours · Metabolic theory of
ecology · Productivity
Autotrophs · Chemosynthesis ·
Chemotrophs · Foundation species ·
Mixotrophs · Myco-heterotrophy · Mycotroph ·
Producers Organotrophs · Photoheterotrophs ·
Photosynthesis · Photosynthetic efficiency ·
Phototrophs · Primary nutritional groups ·
Primary production
Consumers Apex predator · Bacterivore ·

Carnivores · Chemoorganotroph · Foraging ·
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Microorganisms Lithotrophy · Microbial food web · Microbial
loop · Microbial metabolism · Phage ecology
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Food webs Intertidal · Kelp forests · Lakes · North Pacific
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exclusion principle · Copiotrophs · Dominance ·
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Energy quality · Energy Systems Language · f-
ratio · Feed conversion ratio · Feeding frenzy ·
Mesotrophic soil · Oligotroph · Paradox of the
plankton · Trophic cascade · Trophic level ·
Trophic mutualism · Trophic state index
Antipredator adaptations · Herbivore

Defense/counte adaptations to plant defense · Plant defense
r against herbivores · Predator avoidance in
schooling fish
[hide] Modelling ecosystems - other components
Abundance · Allee effect · Depensation ·

Ecological yield · Effective population size ·
Intraspecific competition · Logistic function ·
Population Malthusian growth model · Maximum sustainable
ecology yield · Overpopulation in wild animals ·
Overexploitation · Population cycle · Population
dynamics · Population modeling · Population size ·
Predator–prey equations · Recruitment · Resilience ·
Small population size · Stability
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extinction · Endemic species · Flagship species ·
Gradient analysis · Indicator species · Introduced
species · Invasive species · Latitudinal gradients in
Species species diversity · Minimum viable population ·
Occupancy-abundance relationship · Population
viability analysis · Rapoport's rule · Relative
abundance distribution · Relative species
abundance · Species diversity · Species
homogeneity · Species richness · Species
distribution · Species-area curve · Umbrella species
Species Antibiosis · Biological interaction ·

interaction Commensalism · Community ecology · Ecological
facilitation · Interspecific competition · Mutualism
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ecology Ecocline · Ecotone · Ecotype · Disturbance · Edge
effect · Foster's rule · Habitat fragmentation ·
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epidemiology · Landscape limnology ·
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theory · Source–sink dynamics
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of Nature · Biological data visualization · Constructal
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Retrieved from "http://en.wikipedia.org/wiki/Evolutionary_ecology"
Categories: Ecology | Evolutionary biology

W000
Notharctinae

†Notharctinae
Fossil range: Early-Middle Eocene
Notharctus tenebrosus
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Family: †Notharctidae
Subfamily: †Notharctinae
Trouessart, 1879
Genera
†Cantius
†Copelemur
†Hesperolemur
†Notharctus
†Pelycodus
†Smilodectes
Notharctinae is an extinct subfamily of primates that were common in North America during the early
and middle Eocene (55-34 million years ago). The six genera that make up the group ( Cantius, Pelycodus,
Copelemur, Hesperolemur, Notharctus, and Smilodectes) contain species that are among the most primitive
of the adapiform group, which is one of the most primitive groups of primates. The evolutionary history of this
subfamily has been comparatively well documented and has been used to argue for evolutionary gradualism.
Though it is generally accepted that adapiforms gave rise to modern day lemurs and lorises, it is not currently
known which branch of Adapiformes these living species are most closely related to. Notharctines went
extinct in the middle Eocene, most likely because of a combination of factors including climatic change and
competition with other North American primates.
Contents
[hide]
• 1 Taxonomic diversity
• 2 Morphology
• 4 References
[edit] Taxonomic diversity

Notharctinae is one of the two subfamilies, along with Cercamoniinae, of the Notharctidae family,
which is a member of the infraorder Adapiformes along with Adapidae and Sivaladapidae. Compared to other
subfamilies, the notharctines were not terribly diverse with only two or three species occurring
synchronously. However, notharctines are some of the most common species found in early and middle
Eocene deposits. Each genus has between 1 (Hesperolemur) and 11 (Cantius) species for a total of 25
species making up the subfamily. Body size ranges from 1100 grams in early species of Cantius to 6900
grams in late occurring Notharctus with an overall trend of increasing body size.
Overall, each genus was restricted to a small geographic region. Cantius was a northern (Wyoming)
early Eocene genus, while the other two early Eocene genera, Pelycodus and Copelemur, occupied more
southern habitats (New Mexico). Middle Eocene taxa, Notharctus and Smilodectes. were again found in
Wyoming, while Hesperolemur is known only from southern California. This move to the north may be
associated with climatic warming between the early and the middle Eocene.
[edit] Morphology
Overall, the notharctines retain a very primitive primate morphology. They have long broad snouts, a
dental formula of 2.1.4.3, a lacrimal bone within the orbit, and, except for Notharctus, an unfused mandible.
All have small orbits which indicate a diurnal lifestyle and there is some evidence of the reduction of the
sense of smell in favor of the sense of sight. One of the diagnostic features of this subfamily is that later
species acquired a hypocone, or an extra cusp on the upper molars, from the postprotocingulum, instead of
the lingual cingulum as in cercamoniines. Many later species show increasing adaptation for folivory
including increased body size, the loss of the paraconids, and the fusion of the mandible. Hesperolemur is
unique among notharctines in having the tympanic ring partially fused to the auditory bulla.
Postcranial remains of Notharctus have linked notharctine locomotion with that of living lemurs.
There is also evidence of canine sexual dimorphism in Notharctus which may indicate the presence of social
groups.

The earliest widely accepted adapiform was the European cercamoniine Donrussellia, though recent
finds of additional species may soon show otherwise. Donrussellia is closely related to the earlier and more
primitive European notharctine species of Cantius. After surviving the Atlantic crossing, more advanced
species of Cantius gradually got bigger and developed larger mesostyles and hypocones, which go along
with the switch from a primarily frugivorous diet to a folivorous one.
Some have suggested that there were two separate immigrations of Cantius, one which gave rise to
the larger Pelycodus and one which gave rise to the smaller lineages of Copelemur, Smilodectes and
Notharctus. Others have suggested that a single lineage of Cantius split, with one branch leading to
Copelemur, one to Pelycodus, and one gradually acquiring a fused mandible, one of the few diagnostic
features between Cantius and Notharctus. Smilodectes either derived from the lineage that became
Notharctus or from the more southern Copelemur lineage. Hesperolemur, a middle Eocene taxa, has only
recently been described and is currently thought to be an immigrant species.
Though some scientists believe that members of the adapiform radiation gave rise to simians
because of the long list of dental and cranial similarities including a fused mandible, loss of paraconids, and
large, sexually dimorphic canines, normally the European cercamoniines are the specific subfamily cited. It is
more widely agreed upon that adapiforms are closely related to modern lemurs and lorises, although there
much speculation as to which taxa are more closely related.
[edit] References
• Beard, KC. 2004 The Hunt for the Dawn Monkey: Unearthing the Origins of Monkeys Univ. of
California Press, Berkeley.
• Fleagle, J. G. 1999. Primate Adaptation and Evolution. San Diego, Academic Press.
• Mikko's Phylogeny Archive
• Gebo, DL. 2002. Adapiformes: phylogeny and adaptation. The Primate Fossil Record.
Cambridge University Press
• Gingerich, PD and Simons, EL. Systematics, Phylogeny and Evolution of Early Eocene
Adapidae in North America. 1977.
• Godinot, M. A Summary of Adapiform Systematics and Phylogeny. Folia Primatologica, 1998
Retrieved from "http://en.wikipedia.org/wiki/Notharctinae"
Categories: Prehistoric prosimians

W000
Evolution of insects

Evolution has produced astonishing variety in insects. Pictured are some of the possible shapes of
antennae.
Insects are a highly diverse group of organisms with a worldwide distribution. They have conquered
every terrestrial environment and have complex interactions with a wide variety of organisms, including
predatory-prey relationships.[1]
Contents
[hide]
• 1 Taxonomic Affinities
• 2 Early evidence
• 2.1 Odonata
• 3 Evolutionary relationships
• 4 Origin of insect flight
• 4.1 Theories
• 5 Life cycle
• 5.1 Mayflies
• 5.2 Distant ancestors
• 6 See also
• 7 References
[edit] Taxonomic Affinities
The relationships of insects to other animal groups remain unclear. Although more traditionally
grouped with millipedes and centipedes, evidence has emerged favouring closer evolutionary ties with the
crustaceans. In the Pancrustacea theory, insects, together with among others Malacostraca, make up a
monophyletic group (sharing a common ancestor): this is today a well accepted hypothesis[2].
[edit] Early evidence

The oldest definitive insect fossil is the Devonian Rhyniognatha hirsti, estimated at 396-407 million
years old.[3] This species already possessed dicondylic mandibles, a feature associated with winged insects,
suggesting that wings may already have evolved at this time. Thus, the first insects probably appeared
earlier, in the Silurian period.[3]
The subclass Apterygota (wingless insects) is now considered artificial as the silverfish (order
Thysanura) are more closely related to Pterygota (winged insects) than to bristletails (order Archaeognatha).
For instance, just like flying insects, Thysanura have so-called dicondylic mandibles, while Archaeognatha
have monocondylic mandibles. The reason for their resemblance is not due to a particularly close
relationship, but rather because they both have kept a primitive and original anatomy in a much higher
degree than the winged insects. The most primitive order of flying insects, the mayflies (Ephemeroptera), are
also those who are most morphologically and physiologically similar to these wingless insects. Some mayfly
nymphs resemble aquatic thysanurans.
Modern Archaeognatha and Thysanura still have rudimentary appendages on their abdomen called
styli, while more primitive and extinct insects known as Monura had much more developed abdominal
appendages, as seen here. The abdominal and thoracic segments in the earliest terrestrial ancestor of the
insects would have been more similar to each other than they are today, and the head had well developed
compound eyes and long antennae. Their body size is not known yet. As the most primitive group today,
Archaeognatha, is most abundant near the coasts, it could mean that this was the kind of habitat where the
insect ancestors became terrestrial. But this specialization to coastal niches could also have a secondary
origin, just as could their jumping locomotion, as it is the crawling Thysanura who are considered to be most
original (plesiomorphic). By looking at how primitive cheliceratan book gills (still seen in horseshoe crabs)
evolved into book lungs in primitive spiders and finally into tracheae in more advanced spiders (most of them
still have a pair of book lungs intact as well), it is possible the trachea of insects was formed in a similar way,
modifying gills at the base of their appendages.
So far there is nothing that suggests the insects were a particularly successful group of animals
before they got their wings.
[edit] Odonata
The Odonata (dragonflies) are also a good candidate as the oldest living member of the Pterygota.
Mayflies are morphologically and physiologically more primitive, but the derived and advanced
characteristics of dragonflies could have evolved independently in their own direction for a long time. It
seems that orders with aquatic nymphs or larvae become evolutionarily conservative once they had adapted
to water. If mayflies made it to the water first, this could partly explain why they are more primitive than
dragonflies, even if dragonflies have an older origin.
Similarly, stoneflies are the most primitive of the Neoptera, but they were not necessarily the first
order to branch off. This also makes it less likely that an aquatic ancestor would have the evolutionary
potential to give rise to all the different forms and species of insects that we know today.
Dragonfly nymphs have a unique labial "mask" used for catching prey, and the imago has a unique
way of copulating, using a secondary male sex organ on the second abdominal segment. It looks like
abdominal appendages modified for sperm transfer and direct insemination have occurred at least twice in
insect evolution, once in Odonata and once in the other flying insects. If these two different methods are the
original ways of copulating for each group, it is a strong indication that it is the dragonflies who are the oldest,
not the mayflies. There is still not agreement about this. Another scenario is that abdominal appendages
adapted for direct insemination have evolved three times in insects; once Odonata, once in mayflies and
once in the Neoptera, both mayflies and Neoptera choosing the same solution. If so, it is still possible that
mayflies are the oldest order among the flying insects. The power of flight is assumed to have evolved only
once, suggesting sperm transfer in the earliest flying insects still was done indirectly.
One possible scenario on how direct insemination evolved in insects is seen in scorpions. The male
deposits a spermatophore on the ground, locks its claws with the female's claws and then guides her over his
packet of sperm, making sure it comes in contact with her genital opening.
When the early (male) insects laid their spermatophores on the ground, it seems likely that some of
them used the clasping organs at the end of their body to drag the female over the package. The ancestors of
Odonata evolved the habit of grabbing the female behind her head, as they still do today. This action, rather
than not grasping the female at all, would have increased the male's chances of spreading its genes. The
chances would be further increased if they first attached their spermatophore safely on their own abdomen
before they placed their abdominal claspers behind the female's head; the male would then not let the female
go before her abdomen had made direct contact with his sperm storage, allowing the transfer of all sperm.
This also meant increased freedom in searching for a female mate because the males could now
transport the packet of sperm elsewhere if the first female slipped away. This ability would eliminate the need
to either wait for another female at the site of the deposited sperm packet or to produce a new packet,
wasting energy. Other advantages include the possibility of mating in other, safer places than flat ground,
such as in trees or bushes.
If the ancestors of the other flying insects evolved the same habit of clasping the female and
dragging her over their spermathophore, but posterior instead of anterior like the Odonata does, their genitals
would come very close to each others. And from there on, it would be a very short step to modify the vestigial
appendages near the male genital opening to transfer the sperm directly into the female. The same
appendages the male Odonata use to transfer their sperm to their secondary sexual organs at the front of
their abdomen.
All insects with an aquatic nymphal or larval stage seem to have adapted to water secondarily from
terrestrial ancestors. Of the most primitive insects with no wings at all, Archaeognatha and Thysanura, all
members live their entire life cycle in terrestrial environments. As mentioned previously, Archaeognatha were
the first to split off from the branch that led to the winged insects (Pterygota), and then the Thysanura
branched off. This indicates that these three groups (Archaeognatha, Thysanura and Pterygota) have a
common terrestrial ancestor, which probably resembled a primitive model of Apterygota, was an
opportunistic generalist and laid spermatophores on the ground instead of copulating, like Thysanura still do
today. If it had feeding habits similar to the majority of apterygotes of today, it lived mostly as a decomposer.
One should expect that a gill breathing arthropod would modify its gills to breathe air if it were
adapting to terrestrial environments, and not evolve new respiration organs from bottom up next to the
original and still functioning ones.
Then comes the fact that insect (larva and nymph) gills are actually a part of a modified, closed
trachea system specially adapted for water, called tracheal gills. The arthropod trachea can only arise in an
atmosphere and as a consequence of the adaptations of living on land. This too indicates that insects are
descended from a terrestrial ancestor.
And finally when looking at the three most primitive insects with aquatic nymphs (called naiads:
Ephemeroptera, Odonata and Plecoptera), each order has its own kind of tracheal gills that are so different
from one another that they must have separate origins. This would be expected if they evolved from land-
dwelling species.
This means that one of the most interesting parts of insect evolution is what happened between the
Thysanura-Pterygota split and the first flight.
[edit] Evolutionary relationships

Insects have complex evolutionary and ecological relationships with other groups including plants
and other animals. For example, insects are prey for a variety of organisms, including terrestrial vertebrates.
The earliest vertebrates on land existed 400 million years ago and were large amphibious piscivores, through
gradual evolutionary change, insectivory was the next diet type to evolve.[1]
[edit] Origin of insect flight

The origin of insect flight remains obscure, since the earliest winged insects currently known appear
to have been capable fliers. Some extinct insects (e.g. the Palaeodictyoptera) had an additional pair of
winglets attached to the first segment of the thorax, for a total of three pairs.
The wings themselves are thought by many[who?] to be highly modified (tracheal) gills. And there is
no doubt that the tracheal gills of the mayfly nymph in many species look like wings. By comparing a well
developed pair of gill blades in the naiads and a reduced pair of hind wings on the adults, it is not hard to
imagine that the mayfly gills (tergaliae) and insect wings have a common origin, and newer research also
supports this.[citation needed] The tergaliae are not found in any other order of insects, and they have
evolved in different directions with time. In some nymphs/naiads the most anterior pair has become
sclerotized and works as a gill cover for the rest of the gills. Others can form a large sucker, be used for
swimming or modified into other shapes. But it doesn't have to mean that these structures were originally
gills. It could also mean that the tergaliae evolved from the same structures which gave rise to the wings, and
that flying insects evolved from a wingless terrestrial species with pairs of plates on its body segments: three
on the thorax and nine on the abdomen (mayfly nymphs with nine pairs of tergaliae on the abdomen exist,
but so far no living or extinct insects with plates on the last two segments have been found). If these were
primary gills, it would be a mystery why they should have waited so long to be modified when we see the
different modifications in modern mayfly nymphs.
[edit] Theories
When the first forests arose on Earth, new niches for terrestrial animals were created. Spore-feeders
and others who depended on plants and/or the animals living around them would have to adapt too to make
use of them. In a world with no flying animals, it would probably just be a matter of time before some
arthropods who were living in the trees evolved paired structures with muscle attachments from their
exoskeleton and used them for gliding, one pair on each segment. Further evolution in this direction would
give bigger gliding structures on their thorax and gradually smaller ones on their abdomen. Their bodies
would have become stiffer while thysanurans, which didn't evolve flight, kept their flexible abdomen.
Mayfly nymphs must have adapted to water while they still had the "gliders" on their abdomen intact.
So far there is no concrete evidence to support this theory either, but it is one that offers an explanation for
the problems of why presumably aquatic animals evolved in the direction they did.
Leaping and arboreal insects seems like a good explanation for this evolutionary process for several
reasons. Because early winged insects were lacking the sophisticated wing folding mechanism of neopterous
insects, they must have lived in the open and not been able to hide or search for food under leaves, in
cracks, under rocks and other such confined spaced. In these old forests there weren't many open places
where insects with huge structures on their back could have lived without experiencing huge disadvantages.
If insects got their wings on land and not in water, which clearly seems to be the case, the tree canopies
would be the most obvious place where such gliding structures could have emerged, in a time when the air
was a new territory. The question is if the plates used for gliding evolved from "scratch" or by modifying
already existing anatomical details. The thorax in Thysanura and Archaeognatha are known to have some
structures connected to their trachea which share similarities to the wings of primitive insects. This suggests
the origin of both the wings and the spiracles are related.
Gliding requires universal body modifications, as seen in present-day vertebrates such as some
rodents and marsupials, which have grown wide, flat expansions of skin for this purpose. The flying dragons
(genus Draco) of Indonesia has modified its ribs into gliders, and even some snakes can glide through the air
by spreading their ribs. The main difference is that while vertebrates have an inner skeleton, primitive insects
had a flexible and adaptive exoskeleton.
It is clear that there would have been some animals living in the trees, as animals are always taking
advantage of all available niches, both for feeding and protection. At the time, the reproductive organs were
by far the most nutritious part of the plant, and these early plants show signs of arthropod consumption and
adaptations to protect themselves, for example by placing their reproductive organs as high up as possible.
But there will always be some species who will be able to cope with that by following the their food source up
the trees.
Knowing that insects were terrestrial at that time and that some arthropods (like primitive insects)
were living in the tree crowns, it seems less likely that they would have developed their wings down on the
ground or in the water.
In a three dimensional environment such as trees, the ability to glide would increase the insects'
chances to survive a fall, as well as saving energy. This trait has repeated itself in modern wingless species
such as the gliding ants who are living an arboreal life. When the gliding ability first had originated, gliding
and leaping behavior would be a logical next step, which would eventually be reflected in their anatomical
design.
The need to navigate through vegetation and to land safely would mean good muscle control over
the proto-wings, and further improvements would eventually lead to true (but primitive) wings.
While the thorax got the wings, a long abdomen could have served as a stabilizer in flight.
It is also worth remembering that some of the earliest flying insects were large predators. This isn't
surprising since there weren't yet any other predators hunting in the air: it was therefore a totally new
ecological niche. Some of the prey were without a doubt other insects, as insects with proto-wings would
have radiated into other species even before the wings were fully evolved. From this point onwards, the arms
race could continue: the same predator/prey co-evolution which has existed as long as there have been
predators and prey on earth; both the hunters and the hunted were in need of improving and extending their
flight skills even further to keep up with the other.
Insects that had evolved their proto-wings in a world without flying predators could afford to be
exposed openly without risk, but this changed when carnivorous flying insects evolved. It is unknown when
they first evolved, but once these predators had emerged they put a strong selection pressure on their victims
and themselves. Those of the prey who came up with a good solution about how to fold their wings over their
backs in a way that made it possible for them to live in narrow spaces would not only be able to hide from
flying predators (and terrestrial predators if they were on the ground) but also to exploit a wide variety of
niches that were closed to those who couldn't fold their wings in this way. And today the neopterous insects
(those that can fold their wings back over the abdomen) are by far the most dominant group of insects.
The water-skimming theory suggests that skimming on the water surface is the origin of insect flight.
[4] This theory is based on the fact that the first fossil insects, the Devonian Rhyniognatha hirsti, is thought to
have possessed wings, even though the insects closest evolutionary ties are with crustaceans, which are
aquatic.
[edit] Life cycle

[edit] Mayflies
Another primitive trait of the mayflies are the subimago; no other insects have this winged yet
sexually immature stage. A few specialized species have females with no subimago, but retain the subimago
stage for males.
The reasons the subimago still exists in this order could be that there hasn't been enough selection
pressure to get rid of it; it also seems specially adapted to do the transition from water to air.
The male geniatalia are not fully functional at this point. One reason for this could be that the
modification of the abdominal appendages into male copulation organs emerged later than the evolution of
flight. This is indicated by the fact that dragonflies have a different copulation organ than other insects.
As we know, in mayflies the nymphs and the adults are specialized for two different ways of living; in
the water and in the air. The only stage (instar) between these two is the subimago. In more primitive fossil
forms, the preadult individuals had not just one instar but numerous ones (while the modern subimago do not
eat, older and more primitive species with a subimagos were probably feeding in this phase of life too as the
lines between the instars were much more diffuse and gradual than today). Adult form was reached several
moults before maturity. They probably didn't have more instars after becoming fully mature. This way of
maturing is how Apterygota do it, which moult even when mature, but not winged insects.
Modern mayflies have eliminated all the instars between imago and nymph, except the single instar
called subimago, which is still not (at least not in the males) fully sexually mature. The other flying insects
with incomplete metamorphosis (Exopterygota) have gone a little further and completed the trend; here all
the immature structures of the animal from the last nymphal stage are completed at once in a single final
moult. The more advanced insects with larvae and complete metamorphosis (Endopterygota) have gone
even further. An interesting theory here is that the pupal stage is actually a strongly modified and extended
stage of subimago, but so far it is nothing more than a theory. Interestingly enough there are some insects
within the Exopterygota, thrips and whiteflies (Aleyrodidae), who have evolved pupae-like stages too.
[edit] Distant ancestors

The distant ancestor of flying insects, a species with primitive proto-wings, had a more or less
ametabolous life cycle and instars of basically the same type as thysanurans with no defined nymphal,
subimago or adult stages as the individual became older. Individuals developed gradually as they were
growing and moulting, but there were probably no big changes in between instars.
Modern mayfly nymphs do not acquire gills until after their first moult. Before this stage they are so
small that there is no need for gills to extract oxygen from the water. This could be a trait from the common
ancestor all flyers evolved from. An early terrestrial insect would have no need for paired outgrowths from the
body before it started to live in the trees (or in the water, for that matter), so it would not have any.
This would also affect the way their offspring looked like in the early instars, resembling earlier
ametabolous generations even after they had started to adapt to a new way of living, in a habitat where they
actually could have some good use for flaps along their body. Since they matured in the same way as
thysanurans with plenty of moultings as they were growing and very little difference between the adults and
much younger individuals (unlike modern insects, who are hemimetabolous or holometabolous), there
probably wasn't much room for adapting into different niches depending on age and stage. Also, it would
have been difficult for an animal already adapted to a niche to make a switch to a new niche later in life
based on age or size differences alone when these differences were not significant.
So they had to specialize and focus their whole existence on improving a single lifestyle in a
particular niche. The older the species and the single individuals became, the more would they differ from
their original form as they adapted to their new lifestyle better than the generations before. The final body
design was no longer achieved while still inside the egg, but continued to develop for most of the life, causing
a bigger difference between the youngest and oldest individuals. Assuming that mature individuals most
likely mastered their new element better than did the nymphs who had the same lifestyle, it would appear to
be an advantage if the immatures reached adult shape and form as soon as possible. This may explain why
they evolved fewer but more intense instars and a stronger focus on the adult body, and the differences
between the adults and the first instars were greater, instead of just gradually growing bigger as earlier
generations had done. This evolutionary trend explains how they went from ametabolous to hemimetabolous
insects.
Reaching maturity and a fully grown body became only a part of the development process, gradually
also a new anatomy and new abilities only possible in the later stages of life, were included. The anatomy
they were born and grew up with had limitations the adults who had learned to fly didn't have. If they couldn't
live their early life the way adults did, immature individuals had to adapt to the best way of living and
surviving despite their limitations till the moment came when they could leave them behind. This would be a
starting point in the evolution where imago and nymphs started to live in different niches, some more clearly
defined than others. Also, a final anatomy, size and maturity reached at once with a single final nymphal
stage meant less waste of time and energy, and also made a more complex adult body structure. These
strategies obviously became very successful with time.
Late Carboniferous and Early Permian insect orders include both several current very long-lived
groups (mayflies, (Ephemeroptera), dragonflies (Odonata), cockroaches (Blattodea), and Orthoptera
(grasshoppers and their relatives)) and a number of Paleozoic forms. During this era, some giant dragonfly-
like forms – e.g. Meganeura and Meganeuropsis (Order Protodonata) and Mazothairos (Order
Palaeodictyoptera) – reached wingspans of 55 to 70 cm (22 to 28 in), making them far larger than any living
insect. Also their nymphs must have had a very impressive size. This gigantism may have been due to higher
atmospheric oxygen levels (up to 80% above modern levels during the Carboniferous) that allowed increased
respiratory efficiency relative to today. The lack of flying vertebrates could have been another factor.
Most extant orders of insects developed during the Permian period that began around 270 million
years ago. Many of the early groups became extinct during the Permian-Triassic extinction event, the largest
mass extinction in the history of the Earth, around 252 million years ago.
The remarkably successful Hymenopterans appeared in the Cretaceous but achieved their diversity
more recently, in the Cenozoic. A number of highly successful insect groups — especially the Hymenoptera
and Lepidoptera (butterflies), as well as many types of Diptera (flies) and Coleoptera (beetles) — evolved in
conjunction with flowering plants, a powerful illustration of co-evolution.
Many modern insect genera developed during the Cenozoic; insects from this period on are often
found preserved in amber, often in perfect condition. Such specimens are easily compared with modern
species. The study of fossilized insects is called paleoentomology.
[edit] See also

• Butterfly evolution
• Spider evolution
[edit] References
1. ^ a b Sahney, S., Benton, M.J. & Falcon-Lang, H.J. (2010). "Rainforest collapse triggered
Pennsylvanian tetrapod diversification in Euramerica" (PDF). Geology 38: 1079–1082.
2. ^ Rota-Stabelli, O.; Kayal, E.; Gleeson, D.; Daub, J.; Boore, J.; Telford, M.; Pisani, D.;
Blaxter, M. et al. (2010). "Ecdysozoan mitogenomics: evidence for a common origin of the legged
invertebrates, the Panarthropoda". Genome biology and evolution 2: 425–440.
doi:10.1093/gbe/evq030. PMID 20624745. edit
3. ^ a b [1] Michael S. Engel and David A. Grimaldi. (2004). New light shed on the oldest insect.
Nature 427: 627-630 (12 February 2004) | doi:10.1038/nature02291
4. ^ Marden, J.H. and M.G. Kramer. 1994. Surface-skimming stoneflies: a possible intermediate
stage for insect flight evolution. Science 266, 427-430.
• Grimaldi, David and Michael S. Engel (2005-05-16). Evolution of the Insects. Cambridge
University Press. ISBN 0-521-82149-5. — an up to date review of the evolutionary history of the
insects
• Rasnitsyn, A.P. and Quicke, D.L.J. (2002). History of Insects. Kluwer Academic Publishers.
ISBN 1-4020-0026-X. — detail coverage of various aspects of the evolutionary history of the insects
[2]

• What arthropod brains say about arthropod phylogeny
• Ecological history of the terrestrial insects
• Geographical history of the insects
• The Primitive Characters of Extant Mayflies (Ephemeroptera)
• The insect abdomen and terminalia
• Morphology of Ephemeroptera
• International Palaeoentomological Society
Processes of
Population genetic
Evolutionary
developmental
concepts

Modes of

Retrieved from "http://en.wikipedia.org/wiki/Evolution_of_insects"
Categories: Evolution by taxon | Entomology

W000
W000
History of the Earth


For the history of modern humans, see History of the world.
This article's introduction section may not adequately summarize its contents. To
comply with Wikipedia's lead section guidelines, please consider expanding the lead to
provide an accessible overview of the article's key points. (September 2010)
Geological time put in a diagram called a geological clock, showing the relative lengths of the eons of
the Earth's history
The history of the Earth describes the most important events and fundamental stages in the
development of the planet Earth from its formation 4.6 billion years ago to the present day.[1] Nearly all
branches of natural science have contributed to the understanding of the main events of the Earth's past. The
age of Earth is approximately one-third of the age of the universe.[2] Immense geological and biological
changes have occurred during that time span.
Contents
[hide]
• 1 Hadean and Archaean eons
• 1.1 Origin of the solar system
• 1.2 Origin of the Earth's core and first atmosphere
• 1.3 The giant impact hypothesis
• 1.4 Origin of the oceans and atmosphere
• 1.5 The first continents
• 1.6 Origin of life
• 2 Proterozoic eon
• 2.1 The oxygen revolution
• 2.2 Snowball Earth and the origin of the ozone layer
• 2.3 Proterozoic development of life
• 2.4 Supercontinents in the Proterozoic era
• 2.5 Late Proterozoic climate and life
• 3 Phanerozoic eon
• 3.1 Paleozoic era
• 3.1.1 Cambrian explosion
• 3.1.2 Paleozoic tectonics, paleogeography
and climate
• 3.1.3 Colonization of land
• 3.2 Mesozoic era
[edit] Hadean and Archaean eons
Main articles: Hadean and Archaean
Starting with the Earth's formation by accretion from the solar nebula 4.54 billion years ago (4.54
Ga),[1] the first eon in the Earth's history is called the Hadean.[3] It lasted until the Archaean eon, which
began 3.8 Ga. The oldest rocks found on Earth date to about 4.0 Ga, and the oldest detrital zircon crystals in
some rocks have been dated to about 4.4 Ga,[4] close to the formation of the Earth's crust and the Earth
itself. Because not much material from this time is preserved, little is known about Hadean times, but
scientists hypothesize at an estimated 4.53 Ga,[nb 1] shortly after formation of an initial crust, the proto-Earth
was impacted by a smaller protoplanet, which ejected part of the mantle and crust into space and created the
Moon.[6][7][8]
During the Hadean, the Earth's surface was under a continuous bombardment by meteorites, and
volcanism must have been severe due to the large heat flow and geothermal gradient. The detrital zircon
crystals dated to 4.4 Ga show evidence of having undergone contact with liquid water, considered as proof
that the planet already had oceans or seas at that time.[4] From crater counts on other celestial bodies it is
inferred that a period of intense meteorite impacts, called the "Late Heavy Bombardment", began about 4.1
Ga, and concluded around 3.8 Ga, at the end of the Hadean.[9]
By the beginning of the Archaean, the Earth had cooled significantly. It would have been impossible
for most present day life forms to exist due to the composition of the Archaean atmosphere, which lacked
oxygen and an ozone layer. Nevertheless it is believed that primordial life began to evolve by the early
Archaean, with some possible fossil finds dated to around 3.5 Ga.[10] Some researchers, however,
speculate that life could have begun during the early Hadean, as far back as 4.4 Ga, surviving the possible
Late Heavy Bombardment period in hydrothermal vents below the Earth's surface.[11]
[edit] Origin of the solar system
An artist's impression of protoplanetary disk

Main article: Formation and evolution of the Solar System
The Solar System (including the Earth) formed from a large, rotating cloud of interstellar dust and gas
called the solar nebula, orbiting the Milky Way's galactic center. It was composed of hydrogen and helium
created shortly after the Big Bang 13.7 Ga and heavier elements ejected by supernovas.[12] About 4.6 Ga,
the solar nebula began to contract, possibly due to the shock wave of a nearby supernova. Such a shock
wave would have also caused the nebula to rotate and gain angular momentum. As the cloud began to
accelerate its rotation, gravity and inertia flattened it into a protoplanetary disk oriented perpendicularly to its
axis of rotation. Most of the mass concentrated in the middle and began to heat up, but small perturbations
due to collisions and the angular momentum of other large debris created the means by which protoplanets
up to several kilometres in length began to form, orbiting the nebular center.
The infall of material, increase in rotational speed and the crush of gravity created an enormous
amount of kinetic energy at the center. Its inability to transfer that energy away through any other process at
a rate capable of relieving the build-up resulted in the disk's center heating up. Ultimately, nuclear fusion of
hydrogen into helium began, and eventually, after contraction, a T Tauri star ignited to create the Sun.
Meanwhile, as gravity caused matter to condense around the previously perturbed objects outside the
gravitational grasp of the new sun, dust particles and the rest of the protoplanetary disk began separating
into rings. Successively larger fragments collided with one another and became larger objects, ultimately
becoming protoplanets.[13] These included one collection about 150 million kilometers from the center:
Earth. The planet formed about 4.54 billion years ago (within an uncertainty of 1%)[1] and was largely
completed within 10–20 million years.[14] The solar wind of the newly formed T Tauri star cleared out most of
the material in the disk that had not already condensed into larger bodies.
Computer simulations have shown that planets with distances equal to the terrestrial planets in our
solar system can be created from a protoplanetary disk.[15] The now widely accepted nebular hypothesis
suggests that the same process, which gave rise to the solar system's planets, produces accretion disks
around virtually all newly forming stars in the universe, some of which yield planets.[16]
[edit] Origin of the Earth's core and first atmosphere

See also: Planetary differentiation
The Proto-Earth grew by accretion, until the inner part of the protoplanet was hot enough to melt the
heavy, siderophile metals. Such liquid metals, with now higher densities, began to sink to the Earth's center
of mass. This so called iron catastrophe resulted in the separation of a primitive mantle and a (metallic) core
only 10 million years after the Earth began to form, producing the layered structure of Earth and setting up
the formation of Earth's magnetic field.
During the accretion of material to the protoplanet, a cloud of gaseous silica must have surrounded
the Earth, to condense afterwards as solid rocks on the surface. What was left surrounding the planet was an
early atmosphere of light (atmophile) elements from the solar nebula, mostly hydrogen and helium, but the
solar wind and Earth's heat would have driven off this atmosphere.
This changed when Earth accreted to about 40% its present radius, and gravitational attraction
retained an atmosphere which included water.
[edit] The giant impact hypothesis

Main articles: Origin and evolution of the Moon and Giant impact hypothesis
The Earth's relatively large natural satellite, the Moon, is unique.[nb 2] During the Apollo program,
rocks from the Moon's surface were brought to Earth. Radiometric dating of these rocks has shown the Moon
to be 4527 ± 10 million years old,[17] about 30 to 55 million years younger than other bodies in the solar
system.[18] (New evidence suggests the Moon formed even later, 4.48±0.02 Ga, or 70–110 Ma after the start
of the Solar System.[5]) Another notable feature is the relatively low density of the Moon, which must mean it
does not have a large metallic core, like all other terrestrial bodies in the solar system. The Moon has a bulk
composition closely resembling the Earth's mantle and crust together, without the Earth's core. This has led
to the giant impact hypothesis, the idea that the Moon was formed during a giant impact of the proto-Earth
with another protoplanet by accretion of the material blown off the mantles of the proto-Earth and impactor.
[19][8]
The impactor, sometimes named Theia, is thought to have been a little smaller than the current
planet Mars. It could have formed by accretion of matter about 150 million kilometres from the Sun and Earth,
at their fourth or fifth Lagrangian point. Its orbit may have been stable at first, but destabilized as Theia's
mass increased due to the accretion of matter. Theia oscillated in larger and larger orbits around the
Lagrangian point until it finally collided with Earth about 4.533 Ga.[7][nb 1] Models reveal that when an
impactor this size struck the proto-Earth at a low angle and relatively low speed (8–20 km/sec), much material
from the mantles and crusts of the proto-Earth and the impactor was ejected into space, where much of it
stayed in orbit around the Earth. This material would eventually form the Moon. However, the metallic cores
of the impactor would have sunk through the Earth's mantle to fuse with the Earth's core, depleting the Moon
of metallic material.[20] The giant impact hypothesis thus explains the Moon's abnormal composition.[21] The
ejecta in orbit around the Earth could have condensed into a single body within a couple of weeks. Under the
influence of its own gravity, the ejected material became a more spherical body: the Moon.[22]
The radiometric ages show the Earth existed already for at least 10 million years before the impact,
enough time to allow for differentiation of the Earth's primitive mantle and core. Then, when the impact
occurred, only material from the mantle was ejected, leaving the Earth's core of heavy siderophile elements
untouched.
The impact had some important consequences for the young Earth. It released an enormous amount
of energy, causing both the Earth and Moon to be completely molten. Immediately after the impact, the
Earth's mantle was vigorously convecting, the surface was a large magma ocean. The planet's first
atmosphere must have been completely blown away by the impact.[23] The impact is also thought to have
changed Earth’s axis to produce the large 23.5° axial tilt that is responsible for Earth’s seasons (a simple,
ideal model of the planets’ origins would have axial tilts of 0° with no recognizable seasons). It may also have
sped up Earth’s rotation.
[edit] Origin of the oceans and atmosphere

Because the Earth lacked an atmosphere immediately after the giant impact, cooling must have
occurred quickly. Within 150 million years, a solid crust with a basaltic composition must have formed. The
felsic continental crust of today did not yet exist. Within the Earth, further differentiation could only begin
when the mantle had at least partly solidified again. Nevertheless, during the early Archaean (about 3.0 Ga)
the mantle was still much hotter than today, probably around 1600°C. This means the fraction of partially
molten material was still much larger than today.
Steam escaped from the crust, and more gases were released by volcanoes, completing the second
atmosphere. Additional water was imported by bolide collisions, probably from asteroids ejected from the
outer asteroid belt under the influence of Jupiter's gravity.
The large amount of water on Earth can never have been produced by volcanism and degassing
alone. It is assumed the water was derived from impacting comets that contained ice.[24]:130-132 Though
most comets are today in orbits farther away from the Sun than Neptune, computer simulations show they
were originally far more common in the inner parts of the solar system. However, most of the water on Earth
was probably derived from small impacting protoplanets, objects comparable with today's small icy moons of
the outer planets.[25] Impacts of these objects can have enriched the terrestrial planets (Mercury, Venus, the
Earth and Mars) with water, carbon dioxide, methane, ammonia, nitrogen and other volatiles. If all water on
Earth was derived from comets alone, millions of comet impacts would be required to support this theory.
Computer simulations illustrate that this is not an unreasonable number.[24]:131
As the planet cooled, clouds formed. Rain created the oceans. Recent evidence suggests the oceans
may have begun forming by 4.2 Ga,[26] or as early as 4.4 Ga.[4] In any event, by the start of the Archaean
eon the Earth was already covered with oceans. The new atmosphere probably contained water vapor,
carbon dioxide, nitrogen, and smaller amounts of other gases.[27] As the output of the Sun was only 70% of
the current amount, significant amounts of greenhouse gas in the atmosphere most likely prevented the
surface water from freezing.[28] Free oxygen would have been bound by hydrogen or minerals on the
surface. Volcanic activity was intense and, without an ozone layer to hinder its entry, ultraviolet radiation
flooded the surface.
Lithified stromatolites on the shores of Lake Thetis (Western Australia). Stromatolites are formed by
colonies of single celled organisms like cyanobacteria or chlorophyta. These colonies of algae entrap
sedimentary grains, thus forming the draped sedimentary layers of a stromatolite. Archaean stromatolites are
the first direct fossil traces of life on Earth, even though little preserved fossilized cells have been found
inside them. The Archaean and Proterozoic oceans could have been full of algal mats like these.
[edit] The first continents

Mantle convection, the process that drives plate tectonics today, is a result of heat flow from the core
to the Earth's surface. It involves the creation of rigid tectonic plates at mid-oceanic ridges. These plates are
destroyed by subduction into the mantle at subduction zones. The inner Earth was warmer during the
Hadean and Archaean eons, so convection in the mantle must have been faster. When a process similar to
present day plate tectonics did occur, this would have gone faster too. Most geologists believe that during the
Hadean and Archaean, subduction zones were more common, and therefore tectonic plates were smaller.
The initial crust, formed when the Earth's surface first solidified, totally disappeared from a
combination of this fast Hadean plate tectonics and the intense impacts of the Late Heavy Bombardment. It
is, however, assumed that this crust must have been basaltic in composition, like today's oceanic crust,
because little crustal differentiation had yet taken place. The first larger pieces of continental crust, which is a
product of differentiation of lighter elements during partial melting in the lower crust, appeared at the end of
the Hadean, about 4.0 Ga. What is left of these first small continents are called cratons. These pieces of late
Hadean and early Archaean crust form the cores around which today's continents grew.
The oldest rocks on Earth are found in the North American craton of Canada. They are tonalites from
about 4.0 Ga. They show traces of metamorphism by high temperature, but also sedimentary grains that
have been rounded by erosion during transport by water, showing rivers and seas existed then.[24]
Cratons consist primarily of two alternating types of terranes. The first are so called greenstone belts,
consisting of low grade metamorphosed sedimentary rocks. These "greenstones" are similar to the
sediments today found in oceanic trenches, above subduction zones. For this reason, greenstones are
sometimes seen as evidence for subduction during the Archaean. The second type is a complex of felsic
magmatic rocks. These rocks are mostly tonalite, trondhjemite or granodiorite, types of rock similar in
composition to granite (hence such terranes are called TTG-terranes). TTG-complexes are seen as the
relicts of the first continental crust, formed by partial melting in basalt. The alternation between greenstone
belts and TTG-complexes is interpreted as a tectonic situation in which small proto-continents were
separated by a thorough network of subduction zones.
[edit] Origin of life
The replicator in virtually all known life is deoxyribonucleic acid. DNA is far more complex than the
original replicator and its replication systems are highly elaborate.
Main article: Abiogenesis
The details of the origin of life are unknown, but the basic principles have been established. There
are two schools of thought about the origin of life. One suggests that organic components arrived on Earth
from space (see “Panspermia”), while the other argues that they originated on Earth. Nevertheless, both
schools suggest similar mechanisms by which life initially arose.[29]
If life arose on Earth, the timing of this event is highly speculative—perhaps it arose around 4 Ga.[30]
It is possible that, as a result of repeated formation and destruction of oceans during that time period caused
by high energy asteroid bombardment, life may have arisen and been extinguished more than once.[4]
In the energetic chemistry of early Earth, a molecule gained the ability to make copies of itself — a
replicator. (More accurately, it promoted the chemical reactions which produced a copy of itself.) The
replication was not always accurate: some copies were slightly different from their parent.
If the change destroyed the copying ability of the molecule, the molecule did not produce any copies,
and the line “died out”. On the other hand, a few rare changes might have made the molecule replicate faster
or better: those “strains” would become more numerous and “successful”. This is an early example of
evolution on abiotic material. The variations present in matter and molecules combined with the universal
tendency for systems to move towards a lower energy state allowed for an early method of natural selection.
As choice raw materials (“food”) became depleted, strains which could utilize different materials, or perhaps
halt the development of other strains and steal their resources, became more numerous.[31]:563-578
The nature of the first replicator is unknown because its function was long since superseded by life’s
current replicator, DNA. Several models have been proposed explaining how a replicator might have
developed. Different replicators have been posited, including organic chemicals such as modern proteins,
nucleic acids, phospholipids, crystals,[32] or even quantum systems.[33] There is currently no way to
determine whether any of these models closely fits the origin of life on Earth.
One of the older theories, one which has been worked out in some detail, will serve as an example of
how this might occur. The high energy from volcanoes, lightning, and ultraviolet radiation could help drive
chemical reactions producing more complex molecules from simple compounds such as methane and
ammonia.[34]:38 Among these were many of the simpler organic compounds, including nucleobases and
amino acids, which are the building blocks of life. As the amount and concentration of this “organic soup”
increased, different molecules reacted with one another. Sometimes more complex molecules would result—
perhaps clay provided a framework to collect and concentrate organic material.[34]:39
Certain molecules could speed up a chemical reaction. All this continued for a long time, with
reactions occurring at random, until by chance it produced a replicator molecule. In any case, at some point,
the function of the replicator was superseded by DNA; all known life (except some viruses and prions) use
DNA as their replicator, in an almost identical manner (see Genetic code).
A small section of a cell membrane. This modern cell membrane is far more sophisticated than the
original simple phospholipid bilayer (the small blue spheres with two tails). Proteins and carbohydrates serve
various functions in regulating the passage of material through the membrane and in reacting to the
environment.
Modern life has its replicating material packaged inside a cellular membrane. It is easier to
understand the origin of the cell membrane than the origin of the replicator, because a cell membrane is
made of phospholipid molecules, which often form a bilayer spontaneously when placed in water. Under
certain conditions, many such spheres can be formed (see “The bubble theory”).[34]:40
The prevailing theory is that the membrane formed after the replicator, which perhaps by then was
RNA (the RNA world hypothesis), along with its replicating apparatus and other biomolecules. Initial
protocells may have simply burst when they grew too large; the scattered contents may then have
recolonized other “bubbles”. Proteins that stabilized the membrane, or that later assisted in an orderly
division, would have promoted the proliferation of those cell lines.
RNA is a likely candidate for an early replicator, because it can both store genetic information and
catalyze reactions. At some point DNA took over the genetic storage role from RNA, and proteins known as
enzymes took over the catalysis role, leaving RNA to transfer information, synthesize proteins and regulate
the process. There is increasing belief that these early cells evolved in association with undersea volcanic
vents known as black smokers[34]:42 or even hot, deep rocks.[31]:580
It is believed that of this multiplicity of protocells, only one line survived. Current phylogentic evidence
suggests that the last universal common ancestor lived during the early Archean eon, perhaps roughly 3.5
Ga or earlier.[35][36] This “LUCA” cell is the ancestor of all life on Earth today. It was probably a prokaryote,
possessing a cell membrane and probably ribosomes, but lacking a nucleus or membrane-bound organelles
such as mitochondria or chloroplasts.
Like all modern cells, it used DNA as its genetic code, RNA for information transfer and protein
synthesis, and enzymes to catalyze reactions. Some scientists believe that instead of a single organism
being the last universal common ancestor, there were populations of organisms exchanging genes in lateral
gene transfer.[35]
[edit] Proterozoic eon

The Proterozoic is the eon of Earth's history that lasted from 2.5 Ga to 542 Ma. In this time span, the
cratons grew into continents with modern sizes. For the first time plate tectonics took place in a modern
sense. The change to an oxygen-rich atmosphere was a crucial development. Life developed from
prokaryotes into eukaryotes and multicellular forms. The Proterozoic saw a couple of severe ice ages called
snowball Earths. After the end of the last Snowball Earth about 600 Ma, the evolution of life on Earth
accelerated. About 580 Ma, the Ediacara biota formed the prelude for the Cambrian Explosion.
[edit] The oxygen revolution

Main article: Oxygen catastrophe
The harnessing of the sun’s energy led to several major changes in life on Earth.
Graph showing range of estimated partial pressure of atmospheric oxygen through geologic time [37]
A banded iron formation from the 3.15 Ga Moories Group, Barberton Greenstone Belt, South Africa.
Red layers represent the times when oxygen was available, gray layers were formed in anoxic
circumstances.
The first cells were likely heterotrophs, using surrounding organic molecules (including those from
other cells) as raw material and an energy source.[31]:564-566 As the food supply diminished, a new
strategy evolved in some cells. Instead of relying on the diminishing amounts of free-existing organic
molecules, these cells adopted sunlight as an energy source. Estimates vary, but by about 3 Ga, something
similar to modern oxygenic photosynthesis had probably developed, which made the sun’s energy available
not only to autotrophs but also to the heterotrophs that consumed them.[38][39] This type of photosynthesis,
which became by far the most common, used the abundant carbon dioxide and water as raw materials and,
with the energy of sunlight, produced energy-rich organic molecules (carbohydrates).
Moreover, oxygen was released as a waste product of the photosynthesis.[37] At first, it became
bound up with limestone, iron, and other minerals. There is substantial proof of this in iron-oxide rich layers in
geological strata that correspond with this period. The reaction of the minerals with oxygen would have
turned the oceans green. When most of the exposed readily reacting minerals were oxidized, oxygen finally
began to accumulate in the atmosphere. Though each cell only produced a minute amount of oxygen, the
combined metabolism of many cells over a vast time transformed Earth’s atmosphere to its current state.
[34]:50-51 Among the oldest examples of oxygen-producing lifeforms are fossil stromatolites. This was
Earth’s third atmosphere.
Some of the oxygen was stimulated by incoming ultraviolet radiation to form ozone, which collected
in a layer near the upper part of the atmosphere. The ozone layer absorbed, and still absorbs, a significant
amount of the ultraviolet radiation that once had passed through the atmosphere. It allowed cells to colonize
the surface of the ocean and eventually the land:[40] without the ozone layer, ultraviolet radiation bombarding
land and sea would have caused unsustainable levels of mutation in exposed cells.
Photosynthesis had another, major, and world-changing impact. Oxygen was toxic; probably much
life on Earth died out as its levels rose in what is known as the "oxygen catastrophe".[40] Resistant forms
survived and thrived, and some developed the ability to use oxygen to increase their metabolism and obtain
more energy from the same food.
[edit] Snowball Earth and the origin of the ozone layer

An oxygen-rich atmosphere had two principal advantages for life. Organisms not using oxygen for
their metabolism, such as anaerobe bacteria, base their metabolism on fermentation. The abundance of
oxygen makes respiration possible, a much more effective energy source for life than fermentation. The
second advantage of an oxygen-rich atmosphere is that oxygen forms ozone in the higher atmosphere,
causing the emergence of the Earth's ozone layer. The ozone layer protects the Earth's surface from
ultraviolet radiation, which is harmful for life. Without the ozone layer, the development of more complex life
later on would probably have been impossible.[24]:219-220
The natural evolution of the Sun made it progressively more luminous during the Archaean and
Proterozoic eons; the Sun's luminosity increases 6% every billion years.[24]:165 As a result, the Earth began
to receive more heat from the Sun in the Proterozoic eon. However, the Earth did not get warmer. Instead,
the geological record seems to suggest it cooled dramatically during the early Proterozoic. Glacial deposits
found in all cratons show that about 2.3 Ga, the Earth underwent its first big ice age (the Makganyene ice
age).[41] Some scientists suggest this and following Proterozoic ice ages were so severe that the planet was
totally frozen over from the poles to the equator, a hypothesis called Snowball Earth. Not all geologists agree
with this scenario and older, Archaean ice ages have been postulated, but the ice age 2.3 Ga is the first such
event for which the evidence is widely accepted.
The ice age around 2.3 Ga could have been directly caused by the increased oxygen concentration
in the atmosphere, which caused the decrease of methane (CH4) in the atmosphere. Methane is a strong
greenhouse gas, but with oxygen it reacts to form CO 2, a less effective greenhouse gas.[24]:172 When free
oxygen became available in the atmosphere, the concentration of methane could have decreased
dramatically, enough to counter the effect of the increasing heat flow from the Sun.
[edit] Proterozoic development of life

Main article: Endosymbiotic theory
Some of the pathways by which the various endosymbionts might have arisen.
Modern taxonomy classifies life into three domains. The time of the origin of these domains is
uncertain. The Bacteria domain probably first split off from the other forms of life (sometimes called
Neomura), but this supposition is controversial. Soon after this, by 2 Ga,[42] the Neomura split into the
Archaea and the Eukarya. Eukaryotic cells (Eukarya) are larger and more complex than prokaryotic cells
(Bacteria and Archaea), and the origin of that complexity is only now becoming known.
Around this time, the first proto-mitochondrion was formed. A bacterial cell related to today’s
Rickettsia,[43] which had learned how to metabolize oxygen, entered a larger prokaryotic cell, which lacked
that capability. Perhaps the large cell attempted to ingest the smaller one but failed (possibly due to the
evolution of prey defenses). The smaller cell may have tried to parasitize the larger one. In any case, the
smaller cell survived inside the larger cell. Using oxygen, it metabolized the larger cell’s waste products and
derived more energy. Some of this excess energy was returned to the host. The smaller cell replicated inside
the larger one. Soon, a stable symbiosis developed between the large cell and the smaller cells inside it.
Over time, the host cell acquired some of the genes of the smaller cells, and the two kinds became
dependent on each other: the larger cell could not survive without the energy produced by the smaller ones,
and these in turn could not survive without the raw materials provided by the larger cell. The whole cell is
now considered a single organism, and the smaller cells are classified as organelles called mitochondria.
A similar event occurred with photosynthetic cyanobacteria[44] entering large heterotrophic cells and
becoming chloroplasts.[34]:60-61[31]:536-539 Probably as a result of these changes, a line of cells capable
of photosynthesis split off from the other eukaryotes more than 1 billion years ago. There were probably
several such inclusion events, as the figure at right suggests. Besides the well-established endosymbiotic
theory of the cellular origin of mitochondria and chloroplasts, it has been suggested that cells led to
peroxisomes, spirochetes led to cilia and flagella, and that perhaps a DNA virus led to the cell nucleus,[45],
[46] though none of these theories is widely accepted.[47]
Green algae of the genus Volvox are believed to be similar to the first multicellular plants.
Archaeans, bacteria, and eukaryotes continued to diversify and to become more complex and better
adapted to their environments. Each domain repeatedly split into multiple lineages, although little is known
about the history of the archaea and bacteria. Around 1.1 Ga, the supercontinent Rodinia was assembling.
[48] The plant, animal, and fungi lines had all split, though they still existed as solitary cells. Some of these
lived in colonies, and gradually some division of labor began to take place; for instance, cells on the
periphery might have started to assume different roles from those in the interior. Although the division
between a colony with specialized cells and a multicellular organism is not always clear, around 1 billion
years ago[49] the first multicellular plants emerged, probably green algae.[50] Possibly by around 900
Ma[31]:488 true multicellularity had also evolved in animals.
At first it probably resembled today’s sponges, which have totipotent cells that allow a disrupted
organism to reassemble itself.[31]:483-487 As the division of labor was completed in all lines of multicellular
organisms, cells became more specialized and more dependent on each other; isolated cells would die.
[edit] Supercontinents in the Proterozoic era

Wilson cycle timeline from 1 Ga, depicting Rodinia and Pangaea supercontinent formation and
separation
When the theory of Plate tectonics was developed around 1960, geologists began to reconstruct the
movements and positions of the continents in the past. This appeared relatively easy until about 250 million
years ago, when all continents were united in what is called the "supercontinent" Pangaea. Before that time,
reconstructions cannot rely on apparent similarities in coastlines or ages of oceanic crust, but only on
geologic observations and paleomagnetic data.[24]:95
Throughout the history of the Earth, there have been times when the continental mass came together
to form a supercontinent, followed by the break-up of the supercontinent and new continents moving apart
again. This repetition of tectonic events is called a Wilson cycle. The further back in time, the scarcer and
harder to interpret the data get. It is at least clear that, about 1000 to 830 Ma, most continental mass was
united in the supercontinent Rodinia.[51] Rodinia was not the first supercontinent; it formed at ~1.0 Ga by
accretion and collision of fragments produced by breakup of the older supercontinent, called Nuna or
Columbia, which was assembled by global-scale 2.0-1.8 Ga collisional events.[52][53] This means plate
tectonic processes similar to today's must have been active during the Proterozoic.
After the break-up of Rodinia about 800 Ma, it is possible the continents joined again around 550 Ma.
The hypothetical supercontinent is sometimes referred to as Pannotia or Vendia. The evidence for it is a
phase of continental collision known as the Pan-African orogeny, which joined the continental masses of
current-day Africa, South-America, Antarctica and Australia. It is extremely likely, however, that the
aggregation of continental masses was not completed, since a continent called Laurentia (roughly equivalent
to current-day North America) had already started breaking off around 610 Ma. It is at least certain that by the
end of the Proterozoic eon, most of the continental mass lay united in a position around the south pole.[54]
[edit] Late Proterozoic climate and life
A 580 million year old fossil of Spriggina floundensi, an animal from the Ediacaran period. Such life
forms could have been ancestors to the many new forms that origined in the Cambrian Explosion.
The end of the Proterozoic saw at least two Snowball Earths, so severe that the surface of the
oceans may have been completely frozen. This happened about 710 and 640 Ma, in the Cryogenian period.
These severe glaciations are less easy to explain than the early Proterozoic Snowball Earth. Most
paleoclimatologists think the cold episodes had something to do with the formation of the supercontinent
Rodinia. Because Rodinia was centered on the equator, rates of chemical weathering increased and carbon
dioxide (CO2) was taken from the atmosphere. Because CO2 is an important greenhouse gas, climates
cooled globally.
In the same way, during the Snowball Earths most of the continental surface was in permafrost,
which decreased chemical weathering again, leading to the end of the glaciations. An alternative hypothesis
is that enough carbon dioxide escaped through volcanic outgassing that the resulting greenhouse effect
raised global temperatures.[55] Increased volcanic activity resulted from the break-up of Rodinia at about the
same time.
The Cryogenian period was followed by the Ediacaran period, which was characterized by a rapid
development of new multicellular lifeforms. Whether there is a connection between the end of the severe ice
ages and the increase in diversity of life is not clear, but it does not seem coincidental. The new forms of life,
called Ediacara biota, were larger and more diverse than ever. Most scientists think some of them may have
been the precursors of the new life forms of the following Cambrian period. Though the taxonomy of most
Ediacaran life forms is unclear, some are proposed to have been ancestors of groups of modern life.[56]
Important developments were the origin of muscular and neural cells. None of the Ediacaran fossils had hard
body parts like skeletons. These first appear after the boundary between the Proterozoic and Phanerozoic
eons or Ediacaran and Cambrian periods.
[edit] Phanerozoic eon

[edit] Paleozoic era
The Paleozoic era (meaning: era of old life forms) was the first era of the Phanerozoic eon, lasting
from 542 to 251 Ma. During the Paleozoic, many modern groups of life came into existence. Life colonized
the land, first plants, then animals. Life usually evolved slowly. At times, however, there are sudden
radiations of new species or mass extinctions. These bursts of evolution were often caused by unexpected
changes in the environment resulting from natural disasters such as volcanic activity, meteorite impacts or
climate changes.
The continents formed at the break-up of Pannotia and Rodinia at the end of the Proterozoic would
slowly move together again during the Paleozoic. This would eventually result in phases of mountain building
that created the supercontinent Pangaea in the late Paleozoic.
[edit] Cambrian explosion

Main article: Cambrian explosion
Apparently, the rate of the evolution of life accelerated in the Cambrian period (542-488 Ma). The
sudden emergence of many new species, phyla, and forms in this period is called the Cambrian Explosion.
The biological formenting in the Cambrian Explosion was unpreceded before and since that time.[24]:229
Whereas the Ediacaran life forms appear yet primitive and not easy to put in any modern group, at the end of
the Cambrian most modern phyla were already present. The development of hard body parts such as shells,
skeletons or exoskeletons in animals like molluscs, echinoderms, crinoids and arthropods (a well-known
group of arthropods from the lower Paleozoic are the trilobites) made the preservation and fossilisation of
such life forms easier than those of their Proterozoic ancestors.[57] For this reason, much more is known
about life in and after the Cambrian than about that of older periods. The boundary between the Cambrian
and Ordovician (the following period, 488-444 Ma) is characterized by a large mass-extinction, in which some
of the new groups disappeared altogether.[58] Some of these Cambrian groups appear complex but are quite
different from modern life; examples are Anomalocaris and Haikouichthys.
During the Cambrian, the first vertebrate animals, among them the first fishes, had appeared.[59] A
creature that could have been the ancestor of the fishes, or was probably closely related to it, was Pikaia. It
had a primitive notochord, a structure that could have developed into a vertebral column later. The first fishes
with jaws (Gnathostomata) appeared during the Ordovician. The colonisation of new niches resulted in
massive body sizes. In this way, fishes with increasing sizes evolved during the early Paleozoic, such as the
titanic placoderm Dunkleosteus, which could grow 7 meters long.
[edit] Paleozoic tectonics, paleogeography and climate

At the end of the Proterozoic, the supercontinent Pannotia had broken apart in the smaller continents
Laurentia, Baltica, Siberia and Gondwana. During periods when continents move apart, more oceanic crust
is formed by volcanic activity. Because young volcanic crust is relatively hotter and less dense than old
oceanic crust, the ocean floors will rise during such periods. This causes the sea level to rise. Therefore, in
the first half of the Paleozoic, large areas of the continents were below sea level.
Early Paleozoic climates were warmer than today, but the end of the Ordovician saw a short ice age
during which glaciers covered the south pole, where the huge continent Gondwana was situated. Traces of
glaciation from this period are only found on former Gondwana. During the Late Ordovician ice age, a
number of mass extinctions took place, in which many brachiopods, trilobites, Bryozoa and corals
disappeared. These marine species could probably not contend with the decreasing temperature of the sea
water.[60] After the extinctions new species evolved, more diverse and better adapted. They would fill the
niches left by the extinct species.
The continents Laurentia and Baltica collided between 450 and 400 Ma, during the Caledonian
Orogeny, to form Laurussia. Traces of the mountain belt which resulted from this collision can be found in
Scandinavia, Scotland and the northern Appalachians. In the Devonian period (416-359 Ma) Gondwana and
Siberia began to move towards Laurussia. The collision of Siberia with Laurussia caused the Uralian
Orogeny, the collision of Gondwana with Laurussia is called the Variscan or Hercynian Orogeny in Europe or
the Alleghenian Orogeny in North America. The latter phase took place during the Carboniferous period (359-
299 Ma) and resulted in the formation of the last supercontinent, Pangaea.
[edit] Colonization of land
For most of Earth’s history, there were no multicellular organisms on land. Parts of the surface may
have vaguely resembled this view of Mars.[citation needed]
Oxygen accumulation from photosynthesis resulted in the formation of an ozone layer that absorbed
much of Sun’s ultraviolet radiation, meaning unicellular organisms that reached land were less likely to die,
and prokaryotes began to multiply and become better adapted to survival out of the water. Prokaryotes had
probably colonized the land as early as 2.6 Ga[61] even before the origin of the eukaryotes. For a long time,
the land remained barren of multicellular organisms. The supercontinent Pannotia formed around 600 Ma
and then broke apart a short 50 million years later.[62] Fish, the earliest vertebrates, evolved in the oceans
around 530 Ma.[31]:354 A major extinction event occurred near the end of the Cambrian period,[63] which
ended 488 Ma.[64]
Several hundred million years ago, plants (probably resembling algae) and fungi started growing at
the edges of the water, and then out of it.[65]:138-140 The oldest fossils of land fungi and plants date to 480–
460 Ma, though molecular evidence suggests the fungi may have colonized the land as early as 1000 Ma
and the plants 700 Ma.[66] Initially remaining close to the water’s edge, mutations and variations resulted in
further colonization of this new environment. The timing of the first animals to leave the oceans is not
precisely known: the oldest clear evidence is of arthropods on land around 450 Ma,[67] perhaps thriving and
becoming better adapted due to the vast food source provided by the terrestrial plants. There is also some
unconfirmed evidence that arthropods may have appeared on land as early as 530 Ma.[68]
At the end of the Ordovician period, 440 Ma, additional extinction events occurred, perhaps due to a
concurrent ice age.[60] Around 380 to 375 Ma, the first tetrapods evolved from fish.[69] It is thought that
perhaps fins evolved to become limbs which allowed the first tetrapods to lift their heads out of the water to
breathe air. This would allow them to live in oxygen-poor water or pursue small prey in shallow water.[69]
They may have later ventured on land for brief periods. Eventually, some of them became so well adapted to
terrestrial life that they spent their adult lives on land, although they hatched in the water and returned to lay
their eggs. This was the origin of the amphibians. About 365 Ma, another period of extinction occurred,
perhaps as a result of global cooling.[70] Plants evolved seeds, which dramatically accelerated their spread
on land, around this time (by approximately 360 Ma).[71][72]
Pangaea, the most recent supercontinent, existed from 300 to 180 Ma. The outlines of the modern
continents and other land masses are indicated on this map.
Some 20 million years later (340 Ma[31]:293-296), the amniotic egg evolved, which could be laid on
land, giving a survival advantage to tetrapod embryos. This resulted in the divergence of amniotes from
amphibians. Another 30 million years (310 Ma[31]:254-256) saw the divergence of the synapsids (including
mammals) from the sauropsids (including birds and reptiles). Other groups of organisms continued to evolve,
and lines diverged—in fish, insects, bacteria, and so on—but less is known of the details. The most recent
hypothesized supercontinent, called Pangaea, formed 300 Ma.
[edit] Mesozoic era

The most severe extinction event to date took place 250 Ma, at the boundary of the Permian and
Triassic periods; 95% of life on Earth died out and started the Mesozoic era (meaning middle life) that
spanned 187 million years[73]. This extinction event was possibly caused by the Siberian Traps volcanic
event, an asteroid impact, methane hydrate gasification, sea level fluctuations, a major anoxic event, other
events, or some combination of these events. Either the proposed Wilkes Land crater[74] in Antarctica or
Bedout structure off the northwest coast of Australia may indicate an impact connection with the Permian-
Triassic extinction. But it remains uncertain whether either these or other proposed Permian-Triassic
boundary craters are either real impact craters or even contemporaneous with the Permian-Triassic
extinction event. Life persevered, and around 230 Ma,[75] dinosaurs split off from their reptilian ancestors. An
extinction event between the Triassic and Jurassic periods 200 Ma spared many of the dinosaurs,[76] and
they soon became dominant among the vertebrates. Though some of the mammalian lines began to
separate during this period, existing mammals were probably all small animals resembling shrews.[31]:169
By 180 Ma, Pangaea broke up into Laurasia and Gondwana. The boundary between avian and non-
avian dinosaurs is not clear, but Archaeopteryx, traditionally considered one of the first birds, lived around
150 Ma.[77] The earliest evidence for the angiosperms evolving flowers is during the Cretaceous period,
some 20 million years later (132 Ma).[78] Competition with birds drove many pterosaurs to extinction and the
dinosaurs were probably already in decline[79] when, 65 Ma, a 10-kilometre (6.2 mi) meteorite probably
struck Earth just off the Yucatán Peninsula where the Chicxulub crater is today. This ejected vast quantities
of particulate matter and vapor into the air that occluded sunlight, inhibiting photosynthesis. Most large
animals, including the non-avian dinosaurs, became extinct,[80] marking the end of the Cretaceous period
and Mesozoic era. Thereafter, in the Paleocene epoch, mammals rapidly diversified, grew larger, and
became the dominant vertebrates. Perhaps a couple of million years later (around 63 Ma), the last common
ancestor of primates lived.[31]:160 By the late Eocene epoch, 34 Ma, some terrestrial mammals had returned
to the oceans to become animals such as Basilosaurus which eventually led to dolphins and baleen whales.
[81]
[edit] Cenozoic era (Recent life)

Main article: Cenozoic
[edit] Human evolution

Australopithecus africanus, an early hominid.
A small African ape living around six Ma was the last animal whose descendants would include both
modern humans and their closest relatives, the bonobo and chimpanzees.[31]:100-101 Only two branches of
its family tree have surviving descendants. Very soon after the split, for reasons that are still debated, apes in
one branch developed the ability to walk upright.[31]:95-99 Brain size increased rapidly, and by 2 Ma, the first
animals classified in the genus Homo had appeared.[65]:300 Of course, the line between different species or
even genera is somewhat arbitrary as organisms continuously change over generations. Around the same
time, the other branch split into the ancestors of the common chimpanzee and the ancestors of the bonobo
as evolution continued simultaneously in all life forms.[31]:100-101
The ability to control fire probably began in Homo erectus (or Homo ergaster), probably at least
790,000 years ago[82] but perhaps as early as 1.5 Ma.[31]:67 In addition, it has sometimes suggested that
the use and discovery of controlled fire may even predate Homo erectus. Fire was possibly used by the early
Lower Paleolithic (Oldowan) hominid Homo habilis or strong australopithecines such as Paranthropus.[83]
It is more difficult to establish the origin of language; it is unclear whether Homo erectus could speak
or if that capability had not begun until Homo sapiens.[31]:67 As brain size increased, babies were born
earlier, before their heads grew too large to pass through the pelvis. As a result, they exhibited more
plasticity, and thus possessed an increased capacity to learn and required a longer period of dependence.
Social skills became more complex, language became more sophisticated, and tools became more
elaborate. This contributed to further cooperation and intellectual development.[84]:7 Modern humans (Homo
sapiens) are believed to have originated somewhere around 200,000 years ago or earlier in Africa; the oldest
fossils date back to around 160,000 years ago.[85]
The first humans to show signs of spirituality are the Neanderthals (usually classified as a separate
species with no surviving descendants); they buried their dead, often apparently with food or tools.[86]:17
However, evidence of more sophisticated beliefs, such as the early Cro-Magnon cave paintings (probably
with magical or religious significance)[86]:17-19 did not appear until some 32,000 years ago.[87] Cro-
Magnons also left behind stone figurines such as Venus of Willendorf, probably also signifying religious
belief.[86]:17-19 By 11,000 years ago, Homo sapiens had reached the southern tip of South America, the last
of the uninhabited continents (except for Antarctica, which remained undiscovered until 1820 AD).[88] Tool
use and communication continued to improve, and interpersonal relationships became more intricate.
[edit] Civilization
Main article: History of the world
Further information: History of Africa, History of the Americas, History of Antarctica, and History of
Eurasia
Vitruvian Man by Leonardo da Vinci epitomizes the advances in art and science seen during the
Renaissance.
Throughout more than 90% of its history, Homo sapiens lived in small bands as nomadic hunter-
gatherers.[84]:8 As language became more complex, the ability to remember and communicate information
resulted in a new replicator: the meme.[89] Ideas could be exchanged quickly and passed down the
generations.
Cultural evolution quickly outpaced biological evolution, and history proper began. Somewhere
between 8500 and 7000 BC, humans in the Fertile Crescent in Middle East began the systematic husbandry
of plants and animals: agriculture.[90] This spread to neighboring regions, and developed independently
elsewhere, until most Homo sapiens lived sedentary lives in permanent settlements as farmers.
Not all societies abandoned nomadism, especially those in isolated areas of the globe poor in
domesticable plant species, such as Australia.[91] However, among those civilizations that did adopt
agriculture, the relative stability and increased productivity provided by farming allowed the population to
expand.
Agriculture had a major impact; humans began to affect the environment as never before. Surplus
food allowed a priestly or governing class to arise, followed by increasing division of labor. This led to Earth’s
first civilization at Sumer in the Middle East, between 4000 and 3000 BC.[84]:15 Additional civilizations
quickly arose in ancient Egypt, at the Indus River valley and in China.
Starting around 3000 BC, Hinduism, one of the oldest religions still practiced today, began to take
form.[92] Others soon followed. The invention of writing enabled complex societies to arise: record-keeping
and libraries served as a storehouse of knowledge and increased the cultural transmission of information.
Humans no longer had to spend all their time working for survival—curiosity and education drove the pursuit
of knowledge and wisdom.
Various disciplines, including science (in a primitive form), arose. New civilizations sprang up, traded
with one another, and fought for territory and resources. Empires soon began to develop. By around 500 BC,
there were empires in the Middle East, Iran, India, China, and Greece, on nearly equal footing; at times one
empire expanded, only to decline or be driven back later.[84]:3
In the fourteenth century, the Renaissance began in Italy with advances in religion, art, and science.
[84]:317-319 European civilization began to change beginning in 1500, leading to the scientific and industrial
revolutions. That continent began to exert political and cultural dominance over human societies around the
planet.[84]:295-299 From 1914 to 1918 and 1939 to 1945, nations around the world were embroiled in world
wars.
Established following World War I, the League of Nations was a first step in establishing international
institutions to settle disputes peacefully. After failing to prevent World War II, it was replaced by the United
Nations. In 1992, several European nations joined in the European Union. As transportation and
communication improved, the economies and political affairs of nations around the world have become
increasingly intertwined. This globalization has often produced both conflict and collaboration.
[edit] Recent events

Main article: Modern era
See also: Modernity and Future
Four and a half billion years after the planet's formation, Earth’s life broke free of the biosphere. For
the first time in history, Earth was viewed from space.
Change has continued at a rapid pace from the mid-1940s to today. Technological developments
include nuclear weapons, computers, genetic engineering, and nanotechnology. Economic globalization
spurred by advances in communication and transportation technology has influenced everyday life in many
parts of the world. Cultural and institutional forms such as democracy, capitalism, and environmentalism
have increased influence. Major concerns and problems such as disease, war, poverty, violent radicalism,
and recently, human-caused climate change have risen as the world population increases.[93]
In 1957, the Soviet Union launched the first artificial satellite into orbit and, soon afterward, Yuri
Gagarin became the first human in space. Neil Armstrong, an American, was the first to set foot on another
astronomical object, the Moon. Unmanned probes have been sent to all the known planets in the solar
system, with some (such as Voyager) having left the solar system. The Soviet Union and the United States
were the earliest leaders in space exploration in the 20th Century. Five space agencies, representing over
fifteen countries,[94] have worked together to build the International Space Station. Aboard it, there has been
a continuous human presence in space since 2000.[95]
[edit] See also

Astronomy portal
Earth_sciences portal
• Timeline of the Big Bang

• Geologic time scale
• Detailed logarithmic timeline
• Natural history
• History of the world
• End of civilization
• Timetable of the Precambrian
• Geological history of Earth
• Future of the Earth
• Paleoatmosphere
[edit] Notes
1. ^ a b New evidence suggests a later date for the Giant Impact and the Moon's formation of
4.48±0.02 Ga, or 70–110 Ma after the start of the Solar System.[5]
2. ^ The Earth's Moon is larger relative to its planet than any other satellite in the solar system.
Pluto's satellite Charon is relatively larger, but Pluto is considered a dwarf planet.
[edit] References
1. ^ a b c See:
ISBN 0-8047-1569-6.
• Newman, William L. (2007-07-09). "Age of the Earth". Publications Services, USGS.
• Dalrymple, G. Brent (2001). "The age of the Earth in the twentieth century: a problem
doi:10.1144/GSL.SP.2001.190.01.14.
• Stassen, Chris (2005-09-10). "The Age of the Earth". TalkOrigins Archive.
2. ^ "Five-Year Wilkinson Microwave Anisotropy Probe (WMAP) Observations: Data
Processing, Sky Maps, and Basic Results" (PDF). nasa.gov.
http://lambda.gsfc.nasa.gov/product/map/dr3/pub_papers/fiveyear/basic_results/wmap5basic.pdf.
3. ^ "Age of the Earth". U.S. Geological Survey. 1997.
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• WETHERILL, G.W.; 1991: Occurrence of Earth-Like Bodies in Planetary Systems, Science
253(5019), pp. 535–538.
• Young, Edward (July 4, 2005). Executive Summary 2005.
http://nai.arc.nasa.gov/team/index.cfm?page=execsumm&teamid=26&year=7.
• XIAO, S. & LAFLAMME, M.; 2009: On the eve of animal radiation: phylogeny, ecology and
evolution of the Ediacara biota, Trends in Ecology and Evolution 24, pp 31–40.

• Cosmic Evolution — a detailed look at events from the origin of the universe to the present
• Valley, John W. “A Cool Early Earth?” Scientific American. 2005 October 58–65. – discusses
the timing of the formation of the oceans and other major events in Earth’s early history.
• Davies, Paul. “Quantum leap of life”. The Guardian. 2005 December 20. – discusses
speculation into the role of quantum systems in the origin of life
• Evolution timeline (uses Shockwave). Animated story of life since about 13,700,000,000
shows everything from the big bang to the formation of the earth and the development of bacteria
and other organisms to the ascent of man.
• Scientific American Magazine (October 2005 Issue) A Cool Early Earth?
• Artist's Conception of Cold Early Earth
• Taylor, G. Jeffrey (April 26, 2004). "Origin of the Earth and Moon". NASA.
http://solarsystem.nasa.gov/scitech/display.cfm?ST_ID=446. Retrieved 2006-03-27.

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History of RNA biology


Numerous key discoveries in biology have emerged from studies of RNA (ribonucleic acid), including
seminal work in the fields of biochemistry, genetics, microbiology, molecular biology, molecular evolution and
structural biology. As of 2010, 30 scientists have been awarded Nobel Prizes for experimental work that
includes studies of RNA. Specific discoveries of high biological significance are discussed in this article.
For related information, see the articles on History of Molecular Biology and History of Genetics. For
background information, see the articles on RNA and nucleic acid
Contents
[hide]
• 1 1930 - 1950
• 1.1 RNA and DNA have distinct chemical properties
• 2 1951 - 1965
• 2.1 Messenger RNA (mRNA) carries genetic
information that directs protein synthesis
• 2.2 Ribosomes make proteins
• 2.3 Transfer RNA (tRNA) is the physical link
between RNA and protein
• 2.4 The genetic code is solved
• 2.5 RNA polymerase is purified
• 3 1966 - 1975
• 3.1 First complete nucleotide sequence of a
biological nucleic acid molecule
• 3.2 Evolutionary variation of homologous RNA
sequences reveals folding patterns
• 3.3 First complete genomic nucleotide sequence
• 3.4 Reverse transcriptase can copy RNA into DNA
• 3.5 RNA replicons evolve rapidly
• 3.6 Ribosomal RNA (rRNA) sequences provide a
record of the evolutionary history of all life forms
[edit] 1930 - 1950
[edit] RNA and DNA have distinct chemical properties
When first studied in the early 1900s, the chemical and biological differences between RNA and DNA
were not apparent, and they were named after the materials from which they were isolated; RNA was initially
known as "yeast nucleic acid" and DNA was "pancreas nucleic acid". Using diagnostic chemical tests,
carbohydrate chemists showed that the two nucleic acids contained different sugars, whereupon the common
name for RNA became "ribose nucleic acid". Other early biochemical studies showed that RNA was readily
broken down at high pH, while DNA was stable (although denatured) in alkali. Nucleoside composition
analysis showed first that RNA contained similar nucleobases to DNA, with uracil instead of thymine, and that
RNA contained a number of minor nucleobase components, e.g. small amounts of pseudouridine and
dimethylguanine.[1]
[edit] 1951 - 1965

[edit] Messenger RNA (mRNA) carries genetic information that directs protein
synthesis
The concept of messenger RNA emerged during the late 1950s, and is associated with Crick's
description of his "Central Dogma of Molecular Biology", which asserted that DNA led to the formation of
RNA, which in turn led to the synthesis of proteins. During the early 1960s, sophisticated genetic analysis of
mutations in the lac operon of E. coli and in the rII locus of bacteriophage T4 were instrumental in defining
the nature of both messenger RNA and the genetic code. The short-lived nature of bacterial RNAs, together
with the highly complex nature of the cellular mRNA population, made the biochemical isolation of mRNA
very challenging. This problem was overcome in the 1960s by the use of reticulocytes in vertebrates, which
produce large quantities of mRNA that are highly enriched in RNA encoding alpha- and beta-globin (the two
major protein chains of hemoglobin).[2]
[edit] Ribosomes make proteins

In the 1950s, results of labeling experiments in rat liver showed that radioactive amino acids were
found to be associated with "microsomes" (later redefined as ribosomes) very rapidly after administration,
and before they became widely incorporated into cellular proteins. Ribosomes were first visualized using
electron microscopy, and their ribonucleoprotein components were identified by biophysical methods, chiefly
sedimentation analysis within ultracentrifuges capable of generating very high accelerations (equivalent to
hundreds of thousands times gravity). Polysomes (multiple ribosomes moving along a single mRNA
molecule) were identified in the early 1960s, and their study led to an understanding of how ribosomes
proceed to read the mRNA in a 5' to 3' direction, processively generating proteins as they do so.[3]
[edit] Transfer RNA (tRNA) is the physical link between RNA and protein
Biochemical fractionation experiments showed that radioactive amino acids were rapidly
incorporated into small RNA molecules that remained soluble under conditions where larger RNA-containing
particles would precipitate. These molecules were termed soluble (sRNA) and were later renamed transfer
RNA (tRNA). Subsequent studies showed that (i) every cell has multiple species of tRNA, each of which is
associated with a single specific amino acid, (ii) that there are a matching set of enzymes responsible for
linking tRNAs with the correct amino acids, and (iii) that tRNA anticodon sequences form a specific decoding
interaction with mRNA codons.[4]
[edit] The genetic code is solved
The genetic code consists of the translation of particular nucleotide sequences in mRNA to specific
amino acid sequences in proteins (polypeptides). The ability to work out out the genetic code emerged from
the convergence of three different areas of study--(i) new methods to generate synthetic RNA molecules of
defined composition to serve as artificial mRNAs, (ii) development of in vitro translation systems that could
used to translate the synthetic mRNAs into protein, and (iii) experimental and theoretical genetic work which
established that the code was written in three letter "words" (codons). Today, our understanding of the
genetic code permits the prediction of the amino sequence of the protein products of the tens of thousands of
genes whose sequences are being determined in genome studies.[5]
[edit] RNA polymerase is purified

The biochemical purification and characterization of RNA polymerase from the bacterium Escherichia
coli enabled the understanding of the mechanisms through which RNA polymerase initiates and terminates
transcription, and how those processes are regulated to regulate gene expression (i.e. turn genes on and
off). Following the isolation of E. coli RNA polymerase, the three RNA polymerases of the eukaryotic nucleus
were identified, as well as those associated with viruses and organelles. Studies of transcription also led to
the identification of many protein factors that influence transcription, including repressors, activators and
enhancers. The availability of purified preparations of RNA polymerase permitted investigators to develop a
wide range of novel methods for studying RNA in the test tube, and led directly to many of the subsequent
key discoveries in RNA biology.[6]
[edit] 1966 - 1975
[edit] First complete nucleotide sequence of a biological nucleic acid
molecule
Although determining the sequence of proteins was becoming somewhat routine, methods for
sequencing of nucleic acids were not available until the mid-1960s. In this seminal work, a specific tRNA was
purified in substantial quantities, and then sliced into overlapping fragments using a variety of ribonucleases.
Analysis of the detailed nucleotide composition of each fragment provided the information necessary to
deduce the sequence of the tRNA. Today, the sequence analysis of much larger nucleic acid molecules is
highly-automated and enormously faster.[7]
[edit] Evolutionary variation of homologous RNA sequences reveals folding

patterns
Additional tRNA molecules were purified and sequenced. The first comparative sequence analysis
was done and revealed that the sequences varied through evolution in such a way that all of the tRNAs could
fold into very similar secondary structures (two-dimensional structures) and had identical sequences at
numerous positions (e.g. CCA at the 3' end). The radial four-arm structure of tRNA molecules is termed the
'cloverleaf structure', and results from the evolution of sequences with common ancestry and common
biological function. Since the discovery of the tRNA cloverleaf, comparative analysis of numerous other
homologous RNA molecules has led to the identification of common sequences and folding patterns.[8]
[edit] First complete genomic nucleotide sequence
The 3569 nucleotide sequence of all of the genes of the RNA bacteriophage MS2 was determined by
a large team of researchers over several years, and was reported in a series of scientific papers. These
results enabled the analysis of the first complete genome, albeit an extremely tiny one by modern standards.
Several surprising features were identified, including genes that partially overlap one another and the first
clues that different organisms might have slightly different codon usage patterns.[9]
[edit] Reverse transcriptase can copy RNA into DNA

Retroviruses were shown to have a single-stranded RNA genome and to replicate via a DNA
intermediate, the reverse of the usual DNA-to-RNA transcription pathway. They encode a RNA-dependent
DNA polymerase (reverse transcriptase) that is essential for this process. Some retroviruses can cause
diseases, including several that are associated with cancer, and HIV-1 which causes AIDS. Reverse
transcriptase has been widely used as an experimental tool for the analysis of RNA molecules in the
laboratory, in particular the conversion of RNA molecules into DNA prior to molecular cloning and/or
polymerase chain reaction (PCR).[10]
[edit] RNA replicons evolve rapidly

Biochemical and genetic analyses showed that the enzyme systems that replicate viral RNA
molecules (reverse transcriptases and RNA replicases) lack molecular proofreading (3' to 5' exonuclease)
activity, and that RNA sequences do not benefit from extensive repair systems analogous to those that exist
for maintaining and repairing DNA sequences. Consequently, RNA genomes appear to be subject to
significantly higher mutation rates than DNA genomes. For example, mutations in HIV-1 that lead to the
emergence of viral mutants that are insensitive to antiviral drugs are common, and constitute a major clinical
challenge.[11]
[edit] Ribosomal RNA (rRNA) sequences provide a record of the evolutionary

history of all life forms
Analysis of ribosomal RNA sequences from a large number of organisms demonstrated that all
extant forms of life on Earth share common structural and sequence features of the ribosomal RNA,
reflecting a common ancestry. Mapping the similarities and differences among rRNA molecules from different
sources provides clear and quantitative information about the phylogenetic (i.e. evolutionary) relationships
among organisms. Analysis of rRNA molecules led to the identification of a third major kingdom of
organisms, the archaea, in addition to the prokaryotes and eukaryotes.[12]
[edit] Non-encoded nucleotides are added to the ends of RNA molecules

Molecular analysis of mRNA molecules showed that, following transcription, mRNAs have non-DNA-
encoded nucleotides added to both their 5' and 3' ends (guanosine caps and poly-A, respectively). Enzymes
were also identified that add and maintain the universal CCA sequence on the 3' end of tRNA molecules.
These events are among the first discovered examples of RNA processing, a complex series of reactions
that are needed to convert RNA primary transcripts into biologically active RNA molecules.[13]
[edit] 1976 - 1985
[edit] Small RNA molecules are abundant in the eukaryotic nucleus
Small nuclear RNA molecules (snRNAs) were identified in the eukaryotic nucleus using
immunological studies with autoimmune antibodies, which bind to small nuclear ribonucleoprotein complexes
(snRNPs; complexes of the snRNA and protein). Subsequent biochemical, genetic, and phylogenetic studies
established that many of these molecules play key roles in essential RNA processing reactions within the
nucleus and nucleolus, including RNA splicing, polyadenylation, and the maturation of ribosomal RNAs.[14]
[edit] RNA molecules require a specific, complex three-dimensional structure

for activity
The detailed three-dimensional structure of tRNA molecules was determined using X-ray
crystallography, and revealed highly complex, compact three dimensional structures consisting of tertiary
interactions laid upon the basic cloverleaf secondary structure. Key features of tRNA tertiary structure include
the coaxial stacking of adjacent helices and non-Watson-Crick interactions among nucleotides within the
apical loops. Additional crystallographic studies showed that a wide range of RNA molecules (including
ribozymes, riboswitches and ribosomal RNA) also fold into specific structures containing a variety of 3D
structural motifs. The ability of RNA molecules to adopt specific tertiary structures is essential for their
biological activity, and results from the single-stranded nature of RNA. In many ways, RNA folding is more
highly analogous to the folding of proteins rather than to the highly repetitive folded structure of the DNA
double helix.[4]
[edit] Genes are commonly interrupted by introns that must be removed by
RNA splicing
Analysis of mature eukaryotic messenger RNA molecules showed that they are often much smaller
than the DNA sequences that encode them. The genes were shown to be discontinuous, composed of
sequences that are not present in the final mature RNA (introns), located between sequences that are
retained in the mature RNA (exons). Introns were shown to be removed after transcription through a process
termed RNA splicing. Splicing of RNA transcripts requires a highly precise and coordinated sequence of
molecular events, consisting of (a) definition of boundaries between exons and introns, (b) RNA strand
cleavage at exactly those sites, and (c) covalent linking (ligation) of the RNA exons in the correct order. The
discovery of discontinuous genes and RNA splicing was entirely unexpected by the community of RNA
biologists, and stands as one of the most shocking findings in molecular biology research.[15]
[edit] Alternative pre-mRNA splicing generates multiple proteins from a single

gene
The great majority of protein-coding genes encoded within the nucleus of metazoan cells contain
multiple introns. In many cases, these introns were shown to be processed in more than one pattern, thus
generating a family of related mRNAs that differ, for example, by the inclusion or exclusion of particular
exons. The end result of alternative splicing is that a single gene can encode a number of different protein
isoforms that can exhibit a variety of (usually related) biological functions. Indeed, most of the proteins
encoded by the human genome are generated by alternative splicing.[16]
[edit] Discovery of catalytic RNA (ribozymes)
An experimental system was developed in which an intron-containing rRNA precursor from the
nucleus of the ciliated protozoan Tetrahymena could be spliced in vitro. Subsequent biochemical analysis
shows that this group I intron was self-splicing; that is, the precursor RNA is capable of carrying out the
complete splicing reaction in the absence of proteins. In separate work, the RNA component of the bacterial
enzyme ribonuclease P (a ribonucleoprotein complex) was shown to catalyze its tRNA-processing reaction in
the absence of proteins. These experiments represented landmarks in RNA biology, since they revealed that
RNA could play an active role in cellular processes, by catalyzing specific biochemical reactions. Before
these discoveries, it was believed that biological catalysis was solely the realm of protein enzymes.[17] [18]
[edit] RNA was likely critical for prebiotic evolution

The discovery of catalytic RNA (ribozymes) showed that RNA could both encode genetic information
(like DNA) and catalyze specific biochemical reactions (like protein enzymes). This realization led to the RNA
World Hypothesis, a proposal that RNA may have played a critical role in prebiotic evolution at a time before
the molecules with more specialized functions (DNA and proteins) came to dominate biological information
coding and catalysis. Although it is not possible for us to know the course of prebiotic evolution with any
certainty, the presence of functional RNA molecules with common ancestry in all modern-day life forms is a
strong argument that RNA was widely present at the time of the last common ancestor.[19]
[edit] Introns can be mobile genetic elements

Some self-splicing introns can spread through a population of organisms by "homing", inserting
copies of themselves into genes at sites that previously lacked an intron. Because they are self-splicing (that
is, they remove themselves at the RNA level from genes into which they have inserted), these sequences
represent transposons that are genetically silent, i.e. they do not interfere with the expression of the gene into
which they become inserted. These introns can be regarded as examples of selfish DNA. Some mobile
introns encode homing endonucleases, enzymes that initiate the homing process by specifically cleaving
double-stranded DNA at or near the intron-insertion site of alleles lacking an intron. Mobile introns are
frequently members of either the group I or group II families of self-splicing introns.[20]
[edit] Spliceosomes mediate nuclear pre-mRNA splicing

Introns are removed from nuclear pre-mRNAs by splicesomes, large ribonucleoprotein complexes
made up of snRNA and protein molecules whose composition and molecular interactions change during the
course of the RNA splicing reactions. Spliceosomes assemble on and around splice sites (the boundaries
between introns and exons in the unspliced pre-mRNA) in mRNA precursors and use RNA-RNA interactions
to identify critical nucleotide sequences and, probably, to catalyze the splicing reactions. Nuclear pre-mRNA
introns and spliceosome-associated snRNAs show similar structural features to self-splicing group II introns.
In addition, the splicing pathway of nuclear pre-mRNA introns and group II introns shares a similar reaction
pathway. These similarities have led to the hypothesis that these molecules may share a common ancestor.
[21]
[edit] 1986 - 2000

[edit] RNA sequences can be edited within cells
Messenger RNA precursors from a wide range of organisms can be edited before being translated
into protein. In this process, non-encoded nucleotides may be inserted into specific sites in the RNA, and
encoded nucleotides may be removed or replaced. RNA editing was first discovered within the mitochondria
of kinetoplastid protozoans, where it has been shown to be extensive.[22] For example, some protein-coding
genes encode fewer than 50% of the nucleotides found within the mature, translated mRNA. Other RNA
editing events are found in mammals, plants, bacteria and viruses. These latter editing events involve fewer
nucleotide modifications, insertions and deletions than the events within kinetoplast DNA, but still have high
biological significance for gene expression and its regulation.[23]
[edit] Telomerase uses a built-in RNA template to maintain chromosome ends

Telomerase is an enzyme that is present in all eukaryotic nuclei which serves to maintain the ends of
the linear DNA in the linear chromosomes of the eukaryotic nucleus, through the addition of terminal
sequences that are lost in each round of DNA replication. Before telomerase was identified, its activity was
predicted on the basis of a molecular understanding of DNA replication, which indicated that the DNA
polymerases known at that time could not replicate the 3' end of a linear chromosome, due to the absence of
a template strand. Telomerase was shown to be a ribonucleoprotein enzyme that contains an RNA
component that serves as a template strand, and a protein component that has reverse transcriptase activity
and adds nucleotides to the chromosome ends using the internal RNA template.[24]
[edit] Ribosomal RNA catalyzes peptide bond formation

For years, scientists had worked to identify which protein(s) within the ribosome were responsible for
peptidyl transferase function during translation, because the covalent linking of amino acids represents one
of the most central chemical reactions in all of biology. Careful biochemical studies showed that extensively-
deproteinized large ribosomal subunits could still catalyze peptide bond formation, thereby implying that the
sought-after activity might lie within ribosomal RNA rather than ribosomal proteins. Structural biologists,
using X-ray crystallography, localized the peptidyl transferase center of the ribosome to a highly-conserved
region of the large subunit ribosomal RNA (rRNA) that is located at the place within the ribosome where the
amino-acid-bearing ends of tRNA bind, and where no proteins are present. These studies led to the
conclusion that the ribosome is a ribozyme. The rRNA sequences that make up the ribosomal active site
represent some of the most highly conserved sequences in the biological world. Together, these
observations indicate that peptide bond formation catalyzed by RNA was a feature of the last common
ancestor of all known forms of life.[25]
[edit] Combinatorial selection of RNA molecules enables in vitro evolution

Experimental methods were invented that allowed investigators to use large, diverse populations of
RNA molecules to carry out in vitro molecular experiments that utilized powerful selective replication
strategies used by geneticists, and which amount to evolution in the test tube. These experiments have been
described using different names, the most common of which are "combinatorial selection", "in vitro selection",
and SELEX (for Systematic Evolution of Ligands by Exponential Enrichment). These experiments have been
used for isolating RNA molecules with a wide range of properties, from binding to particular proteins, to
catalyzing particular reactions, to binding low molecular weight organic ligands. They have equal applicability
to elucidating interactions and mechanisms that are known properties of naturally-occurring RNA molecules
to isolating RNA molecules with biochemical properties that are not known in nature. In developing in vitro
selection technology for RNA, laboratory systems for synthesizing complex populations of RNA molecules
were established, and used in conjunction with the selection of molecules with user-specified biochemical
activities, and in vitro schemes for RNA replication. These steps can be viewed as (a) mutation, (b) selection,
and (c) replication. Together, then, these three processes enable in vitro molecular evolution.[26]
[edit] 2001 - present
[edit] Many mobile DNA elements use an RNA intermediate
Transposable genetic elements (transposons) are found which can replicate via transcription into an
RNA intermediate which is subsequently converted to DNA by reverse transcriptase. These sequences,
many of which are likely related to retroviruses, constitute much of the DNA of the eukaryotic nucleus,
especially so in plants. Genomic sequencing shows that retrotransposons make up 36% of the human
genome and over half of the genome of major cereal crops (wheat and maize).[27]
[edit] Riboswitches bind cellular metabolites and control gene expression

Segments of RNA, typically embedded within the 5'-untranslated region of a vast number of bacterial
mRNA molecules, have a profound effect on gene expression through a previously-undiscovered mechanism
that does not involve the participation of proteins. In many cases, riboswitches change their folded structure
in response to environmental conditions (e.g. ambient temperature or concentrations of specific metabolites),
and the structural change controls the translation or stability of the mRNA in which the riboswitch is
embedded. In this way, gene expression can be dramatically regulated at the post-transcriptional level.[28]
[edit] Small RNA molecules regulate gene expression by post-transcriptional

gene silencing
Another previously unknown mechanism by which RNA molecules are involved in genetic regulation
was discovered in the 1990s. Small RNA molecules termed microRNA (miRNA) and small interfering RNA
(siRNA) are abundant in eukaryotic cells and exert post-transcriptional control over mRNA expression. They
function by binding to specific sites within the mRNA and inducing cleavage of the mRNA via a specific
silencing-associated RNA degradation pathway.[29]
[edit] Noncoding RNA controls epigenetic phenomena

In addition to their well-established roles in translation and splicing, members of noncoding RNA
(ncRNA) families have recently been found to function in genome defense and chromosome inactivation. For
example, piwi-interacting RNAs (piRNAs) prevent genome instability in germ line cells, while Xist (X-inactive-
specific-transcript) is essential for X-chromosome inactivation in mammals.[30]
[edit] Nobel Laureates in RNA biology

See also: List of RNA biologists
Name Dates Institution Awards
1989 Nobel Prize

Altman, Sidney 1939- Yale University
in Chemistry
California Institute of 1975 Nobel Prize

Baltimore, David 1938-
Technology in Physiology or Medicine
Barré-Sinoussi, 2008 Nobel Prize

1947- Pasteur Institute
Françoise in Physiology or Medicine
Blackburn, University of California, 2009 Nobel Prize
1948-
Elizabeth San Francisco in Physiology or Medicine
2002 Nobel Prize

Brenner, Sydney 1927- Salk Institute
in Physiology or Medicine
University of Colorado, 1989 Nobel Prize

Cech, Thomas 1947-
Boulder in Chemistry
1916- 1962 Nobel Prize

Crick, Francis Salk Institute
2004 in Physiology or Medicine
CNR Institute of 1975 Nobel Prize

Dulbecco, Renato 1914-
Biomedical Technologies (Italy) in Physiology or Medicine
2006 Nobel Prize

Fire, Andrew 1959- Stanford University
1980 Nobel Prize

Gilbert, Walter 1932- Harvard University
in Chemistry
Greider, Carol 1961- Johns Hopkins 2009 Nobel Prize

University in Physiology or Medicine

Holley, Robert Cornell University
1965 Nobel Prize

Jacob, François 1920 Pasteur Institute
Khorana, H. Massachusetts 1968 Nobel Prize

1922-
Gobind Institute of Technology in Physiology or Medicine
Medical Research 1982 Nobel Prize

Klug, Aaron 1926-
Council (UK) in Chemistry
2006 Nobel Prize

Kornberg, Roger 1947- Stanford University
in Chemistry
University of 2006 Nobel Prize

Mello, Craig 1960-
Massachusetts Medical School in Physiology or Medicine

Monod, Jacques Pasteur Institute
2008 Nobel Prize
Montagnier, Luc 1932- Pasteur Institute
Nirenberg, 1927- National Institutes of 1968 Nobel Prize

Marshall 2010 Health (USA) in Physiology or Medicine

Ochoa, Severo New York University
1934- University of 1975 Nobel Prize

Temin, Howard
1994 Wisconsin, Madison in Physiology or Medicine
Ramakrishnan, Medical Research 2009 Nobel Prize

1952-
Venkatraman Council (UK) in Chemistry
1993 Nobel Prize

Roberts, Richard 1943- New England Biolabs
Massachusetts 1993 Nobel Prize

Sharp, Philip 1944-
Institute of Technology in Physiology or Medicine
Steitz, Thomas 1940- Yale University 2009 Nobel Prize

in Chemistry
2009 Nobel Prize

Szostak, Jack 1952- Harvard University
1907- University of 1957 Nobel Prize

Todd, Alexander
1997 Cambridge in Chemistry
Cold Spring Harbor 1962 Nobel Prize

Watson, James 1928-
Laboratory in Physiology or Medicine
Weizmann Institute of 2009 Nobel Prize

Yonath, Ada 1939-
Science in Chemistry
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Transcriptase". Annual Review of Biochemistry 75: 493.
25.^ Noller, HF; Hoffarth, V; Zimniak, L (1992). "Unusual resistance of peptidyl transferase to
protein extraction procedures". Science 256 (5062): 1416–9. PMID 1604315. edit
26.^ Joyce, GF (1994). "In vitro evolution of nucleic acids". Current opinion in structural biology
4: 331–6. PMID 11539574. edit
27.^ Beauregard, A.; Curcio, M. J.; Belfort, M. (2008). "The Take and Give Between
Retrotransposable Elements and their Hosts". Annual Review of Genetics 42: 587.
doi:10.1146/annurev.genet.42.110807.091549. PMID 18680436. edit
28.^ Roth, A.; Breaker, R. R. (2009). "The Structural and Functional Diversity of Metabolite-
Binding Riboswitches". Annual Review of Biochemistry 78: 305.
29.^ Carthew, R. W.; Sontheimer, E. J. (2009). "Origins and Mechanisms of miRNAs and
siRNAs". Cell 136 (4): 642. doi:10.1016/j.cell.2009.01.035. PMID 19239886. edit
30.^ Bonasio, R.; Tu, S.; Reinberg, D. (2010). "Molecular Signals of Epigenetic States". Science
330 (6004): 612. doi:10.1126/science.1191078. PMID 21030644. edit
Retrieved from "http://en.wikipedia.org/wiki/History_of_RNA_biology"
Categories: RNA | Non-coding RNA | History of science

W000
Terrestrial ecozone

The neutrality of this article is disputed. Please see the discussion on the talk page.
Please do not remove this message until the dispute is resolved. (June 2009)
An ecozone, ecorealm, biogeographic realm or biorealm is the largest scale biogeographic division
of the Earth's land surfaces, based on the historic and evolutionary distribution patterns of terrestrial plants
and animals. Ecozones represent large areas of the Earth's surface where plants and animals developed in
relative isolation over long periods of time, and are separated from one another by geologic features, such as
oceans, broad deserts, or high mountain ranges, that formed barriers to plant and animal migration.
Ecozones correspond to the floristic kingdoms of botany or zoogeographic regions of mammal zoology.
Simply they are a definition of the plants and animals in a region further divided by the land form region.
(Example the taiga ecozone in Canada is divided into the taiga plains, and taiga shield.)
Ecozones are characterized by the evolutionary history of the plants and animals they contain. As
such, they are distinct from biomes, also known as major habitat types, which are divisions of the Earth's
surface based on life form, or the adaptation of plants and animals to climatic, soil, and other conditions.
Biomes are characterized by similar climax vegetation. Each ecozone may include a number of different
biomes. A tropical moist broadleaf forest in Central America, for example, may be similar to one in New
Guinea in its vegetation type and structure, climate, soils, etc., but these forests are inhabited by plants and
animals with very different evolutionary histories.
The patterns of plant and animal distribution in the world's ecozones was shaped by the process of
plate tectonics, which has redistributed the world's land masses over geological history.
The term ecozone, as used here, is a fairly recent development, and other terms, including kingdom,
realm, and region, are used by other authorities to denote the same meaning. In some older works, continent
is used, which leads to confusion with the geographic/geologic continents. J. Schultz uses the term
"ecozone" to refer his classification system of biomes.
Contents
[hide]
• 1 WWF Ecozones
• 1.1 Bioregions
• 2 See also
• 3 References
[edit] WWF Ecozones

The ecozones are based largely on the biogeographic realms of Pielou (1979) and Udvardy (1975).
A team of biologists convened by the World Wildlife Fund (WWF) developed a system of eight biogeographic
realms (ecozones) as part of their delineation of the world's over 800 terrestrial ecoregions.
Area
Ecozone million million Notes
square kilometres square miles
including the
Palearctic 54.1 20.9 bulk of Eurasia and
North Africa
including most
Nearctic 22.9 8.8
of North America
including Sub-
Afrotropic 22.1 8.5
Saharan Africa
6 of the 8 ecozones
including Near
Neotropic 19.0 7.3 South America and Palea
the Caribbean Afrotr
Indom
Australasia 7.6 2.9 including
Austra
Australia, New
Guinea, and Neotr
neighbouring islands. Oceania and Antarctic
The northern
boundary of this zone
is known as the
Wallace line.
including the
Indo-Malaya 7.5 2.9 Indian subcontinent
and Southeast Asia
including
Polynesia, Melanesia,
Oceania 1.0 0.39 Micronesia, New
Zealand and some
parts of Australia
including
Antarctic 0.3 0.12
Antarctica.
The WWF scheme is broadly similar to Udvardy's system, the chief difference being the delineation
of the Australasian ecozone relative to the Antarctic, Oceanic, and Indomalayan ecozones. In the WWF
system, The Australasia ecozone includes Australia, Tasmania, the islands of Wallacea, New Guinea, the
East Melanesian islands, New Caledonia, and New Zealand. Udvardy's Australian realm includes only
Australia and Tasmania; he places Wallacea in the Indomalayan Realm, New Guinea, New Caledonia, and
East Melanesia in the Oceanian Realm, and New Zealand in the Antarctic Realm.
[edit] Bioregions
The WWF scheme further subdivides the ecozones into bioregions, defined as "geographic clusters
of ecoregions that may span several habitat types, but have strong biogeographic affinities, particularly at
taxonomic levels higher than the species level (genus, family)." The WWF bioregions are as follows:
• Afrotropic
• Antarctic
• Australasia
• Wallacea
• New Guinea and Melanesia
• Indomalaya
• Indian subcontinent
• Indochina
• Sunda Shelf and Philippines
• Nearctic
• Canadian Shield
• Eastern North America
• Northern Mexico
• Western North America
• Neotropical
• Amazonia
• Caribbean
• Central America
• Central Andes
• Eastern South America
• Northern Andes
• Orinoco
• Southern South America
• Oceania
• Palearctic
[edit] See also

Environment portal
Ecology portal
Geography portal
• Ecological land classification

• Ecoprovince
• Ecoregion
[edit] References
• R. K. Brummitt (2001). "World Geographical Scheme for recording plant distributions" (PDF).
Carnegie Mellon University. http://www.tdwg.org/TDWG_geo2.pdf. Retrieved 2008-03-08.
• Cox, C. Barry; Peter D. Moore (1985). Biogeography: An Ecological and Evolutionary
Approach (Fourth Edition). Blackwell Scientific Publications, Oxford.
• Dinerstein, Eric; David Olson; Douglas J. Graham; et al. (1995). A Conservation Assessment
of the Terrestrial Ecoregions of Latin America and the Caribbean. World Bank, Washington DC.
• Pielou, E.C. (1979), Biogeography. A Wiley-Interscience publication, John Wiley & Sons,
New York, Chichester, Brisbane, Toronto. 351 p. ISBN 0471058459
• Ricketts, Taylor H., Eric Dinerstein, David M. Olson, Colby J. Loucks, et al. (1999).
Terrestrial Ecoregions of North America: a Conservation Assessment . Island Press, Washington DC.
• Schultz, J.: Die Ökozonen der Erde, Ulmer Stuttgart, 3rd ed. 2002 (1st ed. 1988). ISBN 3-
8252-1514-8
• Schultz, J.: Handbuch der Ökozonen, Ulmer Stuttgart 2000. ISBN 3-8252-8200-7
• Schultz, J.: The Ecozones of the World, Springer, Berlin Heidelberg New York, 2n ed. 2005.
ISBN 3540200142
• Udvardy, M.D.F (1975). "A classification of the biogeographical provinces of the world. IUCN
Occasional Paper no. 18" (PDF). Morgues, Switzerland : IUCN.
http://cmsdata.iucn.org/downloads/udvardy.pdf. Retrieved 2009-05-19.
• Wikramanayake, Eric; Eric Dinerstein; Colby J. Loucks; et al. (2002). Terrestrial Ecoregions
of the Indo-Pacific: a Conservation Assessment. Island Press; Washington, DC. ISBN
9781559639231

• Media related to Ecozones at Wikimedia Commons
Retrieved from "http://en.wikipedia.org/wiki/Terrestrial_ecozone"
Categories: Ecozones | Biogeography

W000
Alloenzyme

This article does not cite any references or sources.
may be challenged and removed. (August 2008)
Alloenzymes are common biological enzymes that exhibit high levels of functional evolutionary
conservation throughout specific phylas and kingdoms. They are used by phylogeneticists as molecular
markers to gauge evolutionary histories and relationships between different species of organisms.
These enzymes generally perform very basic functions found commonly throughout all lifeforms,
such as DNA polymerase, the enzyme that repairs and copies DNA. Significant changes in this enzyme
reflect significant events in evolutionary history of organisms. As expected DNA polymerase shows relatively
small differences in its amino acid sequence between phyla and even kingdoms.
The key to choosing which alloenzyme to use in a comparison between multiple species is by
choosing one that is as variable as possible while still being present in all the organisms. By comparing the
amino acid sequence of the enzyme in the species, more amino acid similarities should be seen between
evolutionarily related species and fewer between those that are less well related. The less well conserved the
enzyme is, the more amino acid differences will be present in even closely related species.
[edit] See also

• Comparative genomics
• Phylogenetics
• Molecular phylogeny
• Molecular evolution
• Homology (biology)
Retrieved from "http://en.wikipedia.org/wiki/Alloenzyme"
Categories: Molecular biology | Enzymes | Evolutionary biology | Genomics | Phylogenetics

W000
Diatom

For a molecule of two atoms, see Diatomic molecule.
Diatoms
Marine diatoms.
Domain: Eukaryota
Kingdom: Chromalveolata
Phylum: Heterokontophyta
Bacillariophyceae
Class:
Haeckel 1878
Orders
• Centrales
• Pennales
A diatom. Numbered ticks are 10 microns apart.
Diatoms[1] are a major group of algae, and are one of the most common types of phytoplankton.
Most diatoms are unicellular, although they can exist as colonies in the shape of filaments or ribbons (e.g.
Fragillaria), fans (e.g. Meridion), zigzags (e.g. Tabellaria), or stellate colonies (e.g. Asterionella). Diatoms are
producers within the food chain. A characteristic feature of diatom cells is that they are encased within a
unique cell wall made of silica (hydrated silicon dioxide) called a frustule. These frustules show a wide
diversity in form, but usually consist of two asymmetrical sides with a split between them, hence the group
name. Fossil evidence suggests that they originated during, or before, the early Jurassic Period. Diatom
communities are a popular tool for monitoring environmental conditions, past and present, and are commonly
used in studies of water quality.
Contents
[hide]
• 1 General biology
• 3 Ecology
• 4 Life-Cycle
• 6 Fossil record
• 7 Collection
• 8 EST sequencing
• 9 Genome sequencing
• 10 Nanotechnology research
• 11 See also
• 12 References
[edit] General biology
There are more than 200 genera of living diatoms, and it is estimated that there are approximately
100,000 extant species.[2][3][4] Diatoms are a widespread group and can be found in the oceans, in
freshwater, in soils and on damp surfaces. Most live pelagically in open water, although some live as surface
films at the water-sediment interface (benthic), or even under damp atmospheric conditions. They are
especially important in oceans, where they are estimated to contribute up to 45% of the total oceanic primary
production.[5] Spatial distribution of marine phytoplankton species is restricted both horizontally and
vertically.[6] Diatoms occur in all oceans from the poles to the tropics; polar and subpolar regions contain
relatively few species compared with temperate biota. Although tropical regions exhibit the greatest number
of species, more abundant populations are found in polar to temperate regions.[7] Usually microscopic, some
species of diatoms can reach up to 2 millimetres in length.
Several species of fresh-water diatoms.
Diatoms belong to a large group called the heterokonts, including both autotrophs (e.g. golden algae,
kelp) and heterotrophs (e.g. water moulds). Their yellowish-brown chloroplasts are typical of heterokonts,
with four membranes and containing pigments such as the carotenoid fucoxanthin. Individuals usually lack
flagella, but they are present in gametes and have the usual heterokont structure, except they lack the hairs
(mastigonemes) characteristic in other groups. Most diatoms are non-motile, although some move via
flagellation. As their relatively dense cell walls cause them to readily sink, planktonic forms in open water
usually rely on turbulent mixing of the upper layers by the wind to keep them suspended in sunlit surface
waters. Some species actively regulate their buoyancy with intracellular lipids to counter sinking.
Diatom cells are contained within a unique silicate (silicic acid) cell wall comprising two separate
valves (or shells). The biogenic silica that the cell wall is composed of is synthesised intracellularly by the
polymerisation of silicic acid monomers. This material is then extruded to the cell exterior and added to the
wall. Diatom cell walls are also called frustules or tests, and their two valves typically overlap one over the
other like the two halves of a petri dish. In most species, when a diatom divides to produce two daughter
cells, each cell keeps one of the two halves and grows a smaller half within it. As a result, after each division
cycle the average size of diatom cells in the population gets smaller. Once such cells reach a certain
minimum size, rather than simply divide vegetatively, they reverse this decline by forming an auxospore. This
expands in size to give rise to a much larger cell, which then returns to size-diminishing divisions. Auxospore
production is almost always linked to meiosis and sexual reproduction.
Decomposition and decay of diatoms leads to organic and inorganic (in the form of silicates)
sediment, the inorganic component of which can lead to a method of analyzing past marine environments by
corings of ocean floors or bay muds, since the inorganic matter is embedded in deposition of clays and silts
and forms a permanent geological record of such marine strata.
The study of diatoms is a branch of phycology, and phycologists specializing in diatoms are called
diatomists.
Selections from Ernst Haeckel's 1904 Kunstformen der Natur (Artforms of Nature), showing pennate
(left) and centric (right) frustules.
Diatomaceous earth as viewed under bright field illumination on a light microscope. Diatomaceous
earth is a soft, siliceous, sedimentary rock made up of the cell walls of diatoms and readily crumbles to a fine
powder. This sample consists of a mixture of centric (radially symmetric) and pennate (bilaterally symmetric)
diatoms. This image of diatomaceous earth particles in water is at a scale of 6.236 pixels/μm, the entire
image covers a region of approximately 1.13 by 0.69 mm.
The classification of heterokonts is still unsettled, and they may be treated as a division (or phylum),
kingdom, or something in-between. Accordingly, groups like the diatoms may be ranked anywhere from class
(usually called Diatomophyceae) to division (usually called Bacillariophyta), with corresponding changes in
the ranks of their subgroups.
Diatoms are traditionally divided into two orders:
• centric diatoms (Centrales), which are radially symmetric
• pennate diatoms (Pennales), which are bilaterally symmetric. The former are paraphyletic to
the latter.
A more recent classification[3] divides the diatoms into three classes:
• centric diatoms (Coscinodiscophyceae)
• pennate diatoms
• without a raphe (Fragilariophyceae)
• with a raphe (Bacillariophyceae)
It is probable there will be further revisions as understanding of their relationships increases.[8]
Diatoms generally range in size from ca. 2-200μm,[2] and are composed of a cell wall comprising
silica.[7] This siliceous wall can be highly patterned with a variety of pores, ribs, minute spines, marginal
ridges and elevations; all of which can be utilised to delineate genera and species. The cell itself consists of
two halves, each containing an essentially flat plate, or valve and marginal connecting, or girdle band. One
half, the hypotheca, is slightly smaller than the other half, the epitheca. Diatom morphology varies, typically
though the shape of the cell is circular, although, some cells may be triangular, square, or elliptical.
Cells are solitary or united into colonies of various kinds, which may be linked by siliceous structures;
mucilage pads, or stalks; mucilage tubes; amorphous masses of mucilage and threads of polysaccharide
(chitin), which are secreted through strutted processes. Major pigments of diatoms are chlorophylls a and c,
beta-carotene, fucoxanthin, diatoxanthin and diadinoxanthin.[2] Diatoms are primarily photosynthetic. A few,
however, are obligate heterotrophs, while others can live heterotrophically in the absence of light, provided
an appropriate organic carbon source is available. Storage products are chrysolaminarin and lipids.[7]
Round & Crawford (1990)[3] and Hoek et al. (1995)[9] provide more comprehensive coverage of
diatom taxonomy.
[edit] Ecology
A budget of the ocean's silicon cycle[10]
Planktonic diatoms in freshwater and marine environments typically exhibit a "boom and bust" (or
"bloom and bust") lifestyle. When conditions in the upper mixed layer (nutrients and light) are favourable ( e.g.
at the start of spring) their competitive edge[11] allows them to quickly dominate phytoplankton communities
("boom" or "bloom"). As such they are often classed as opportunistic r-strategists (i.e. those organisms
whose ecology is defined by a high growth rate, r).
When conditions turn unfavourable, usually upon depletion of nutrients, diatom cells typically
increase in sinking rate and exit the upper mixed layer ("bust"). This sinking is induced by either a loss of
buoyancy control, the synthesis of mucilage that sticks diatoms cells together, or the production of heavy
resting spores. Sinking out of the upper mixed layer removes diatoms from conditions unfavourable to
growth, including grazer populations and higher temperatures (which would otherwise increase cell
metabolism). Cells reaching deeper water or the shallow seafloor can then rest until conditions become more
favourable again. In the open ocean, many sinking cells are lost to the deep, but refuge populations can
persist near the thermocline.
Ultimately, diatom cells in these resting populations re-enter the upper mixed layer when vertical
mixing entrains them. In most circumstances, this mixing also replenishes nutrients in the upper mixed layer,
setting the scene for the next round of diatom blooms. In the open ocean (away from areas of continuous
upwelling[12]), this cycle of bloom, bust, then return to pre-bloom conditions typically occurs over an annual
cycle, with diatoms only being prevalent during the spring and early summer. In some locations, however, an
autumn bloom may occur, caused by the breakdown of summer stratification and the entrainment of nutrients
while light levels are still sufficient for growth. Since vertical mixing is increasing, and light levels are falling as
winter approaches, these blooms are smaller and shorter-lived than their spring equivalents.
In the open ocean, the condition that typically causes diatom (spring) blooms to end is a lack of
silicon. Unlike other nutrients, this is only a major requirement of diatoms so it is not regenerated in the
plankton ecosystem as efficiently as, for instance, nitrogen or phosphorus nutrients. This can be seen in
maps of surface nutrient concentrations - as nutrients decline along gradients, silicon is usually the first to be
exhausted (followed normally by nitrogen then phosphorus).
Because of this bloom-and-bust cycle, diatoms are believed to play a disproportionately important
role in the export of carbon from oceanic surface waters[12][13] (see also the biological pump). Significantly,
they also play a key role in the regulation of the biogeochemical cycle of silicon in the modern ocean.[10][14]
Egge & Aksnes (1992)[15] figure.

The use of silicon by diatoms is believed by many researchers to be the key to their ecological
success. In a now classic study, Egge & Aksnes (1992)[15] found that diatom dominance of mesocosm
communities was directly related to the availability of silicic acid — when concentrations were greater than 2
mmol m−3, they found that diatoms typically represented more than 70% of the phytoplankton community.
Raven (1983)[16] noted that, relative to organic cell walls, silica frustules require less energy to synthesize
(approximately 8% of a comparable organic wall), potentially a significant saving on the overall cell energy
budget. Other researchers[17] have suggested that the biogenic silica in diatom cell walls acts as an effective
pH buffering agent, facilitating the conversion of bicarbonate to dissolved CO2 (which is more readily
assimilated). Notwithstanding the possible advantages conferred by silicon, diatoms typically have higher
growth rates than other algae of a corresponding size.[11]
Diatoms occur in virtually every environment that contains water. This includes not only oceans,
seas, lakes and streams, but also soil.
[edit] Life-Cycle
Sexual reproduction of a centric diatom (oogamy)

Sexual reproduction of a pinnate diatom (morphological isogamy, physiological anisogamy)
Diatoms are non-motile; however, sperm found in some species can be flagellated, though motility is
usually limited to a gliding motion.[7] Reproduction among these organisms is primarily asexual by binary
fission, with each daughter cell receiving one of the parent cell's two frustules (or theca). This is used by each
daughter cell as the larger frustule (or epitheca) into which a second, small frustule (or hypotheca) is
constructed.
This form of division results in a size reduction of the offspring and therefore the average cell size of
a diatom population decreases, until the cells are about one-third their maximum size.[2] It has been
observed, however, the ability of certain taxa to divide without causing a reduction in cell size.[18]
Nonetheless, in order to restore the cell size of a diatom population for those that do endure size reduction,
sexual reproduction and auxospore formation must occur.[2] Vegetative cells of diatoms are diploid (2N) and
so meiosis can take place, producing male and female gametes which then fuse to form the zygote. The
zygote sheds its silica theca and grows into a large sphere covered by an organic membrane, the auxospore.
A new diatom cell of maximum size, the initial cell, forms within the auxospore thus beginning a new
generation. Resting spores may also be formed as a response to unfavourable environmental conditions with
germination occurring when conditions improve.[7]
In centric diatoms, the small male gametes have one flagellum while the female gametes are large
and non-motile (oogamous). Conversely, in pinnate diatoms both gametes lack flagella (isoogamous).[2]
Certain araphid species have been documented as anisogamous and are, therefore, considered to represent
a transitional stage between centric and pinnate diatoms.[18]

Heterokont chloroplasts appear to be derived from those of red algae, rather than directly from
prokaryotes as occurred in plants. This suggests they had a more recent origin than many other algae.
However, fossil evidence is scant, and it is really only with the evolution of the diatoms themselves that the
heterokonts make a serious impression on the fossil record.
The earliest known fossil diatoms date from the early Jurassic (~185 Ma),[19] although molecular
clock[19] and sedimentary[20] evidence suggests an earlier origin. It has been suggested that their origin
may be related to the end-Permian mass extinction (~250 Ma), after which many marine niches were
opened.[21] The gap between this event and the time that fossil diatoms first appear may indicate a period
when diatoms were unsilicified and their evolution was cryptic.[22] Since the advent of silicification, diatoms
have made a significant impression on the fossil record, with major deposits of fossil diatoms found as far
back as the early Cretaceous, and some rocks (diatomaceous earth, diatomite, kieselguhr) being composed
almost entirely of them.
Although the diatoms may have existed since the Triassic, the timing of their ascendancy and "take-
over" of the silicon cycle is more recent. Prior to the Phanerozoic (before 544 Ma), it is believed that microbial
or inorganic processes weakly regulated the ocean's silicon cycle.[23][24][25] Subsequently, the cycle
appears dominated (and more strongly regulated) by the radiolarians and siliceous sponges, the former as
zooplankton, the latter as sedentary filter feeders primarily on the continental shelves.[26] Within the last 100
My, it is thought that the silicon cycle has come under even tighter control, and that this derives from the
ecological ascendancy of the diatoms.
However, the precise timing of the "take-over" is unclear, and different authors have conflicting
interpretations of the fossil record. Some evidence, such as the displacement of siliceous sponges from the
shelves,[27] suggests that this takeover began in the Cretaceous (146 Ma to 65 Ma), while evidence from
radiolarians suggests "take-over" did not begin until the Cenozoic (65 Ma to present).[28] The expansion of
grassland biomes and the evolutionary radiation of grasses during the Miocene is believed to have increased
the flux of soluble silicon to the oceans, and it has been argued that this has promoted the diatoms during the
Cenozoic era.[29][30] However, work on the variation of diatom diversity during the Cenozoic suggests
instead that diatom success is decoupled from the evolution of grasses, and that diatoms were most diverse
prior to the diversification of grasses.[31] Nevertheless, regardless of the details of the "take-over" timing, it is
clear that this most recent revolution has installed much tighter biological control over the biogeochemical
cycle of silicon.
The fossil record of diatoms has largely been established through the recovery of their siliceous
frustules in marine and non-marine sediments. Although diatoms have both a marine and non-marine
stratigraphic record, diatom biostratigraphy, which is based on time-constrained evolutionary originations and
extinctions of unique taxa, is only well developed and widely applicable in marine systems. The duration of
diatom species ranges have been documented through the study of ocean cores and rock sequences
exposed on land.[32] Where diatom biozones are well established and calibrated to the geomagnetic polarity
time scale (e.g., Southern Ocean, North Pacific, eastern equatorial Pacific), diatom-based age estimates may
be resolved to within <100,000 years, although typical age resolution for Cenozoic diatom assemblages is
several hundred thousand years.
The Cretaceous record of diatoms is limited, but recent studies reveal a progressive diversification of
diatom types. The Cretaceous-Tertiary extinction event, which in the oceans dramatically affected organisms
with calcareous skeletons, appears to have had relatively little impact on diatom evolution.[33]
Although no mass extinctions of marine diatoms have been observed during the Cenozoic, times of
relatively rapid evolutionary turnover in marine diatom assemblages occurred near the Paleocene–Eocene
boundary[34] and at the Eocene–Oligocene boundary.[35] Further turnover of assemblages took place at
various times between the middle Miocene and late Pliocene,[36] in response to progressive cooling of polar
regions and the development of more endemic diatom assemblages. A global trend toward more delicate
diatom frustules has been noted from the Oligocene to the Quaternary.[32] This coincides with an
increasingly more vigorous circulation of the ocean’s surface and deep waters brought about by increasing
latitudinal thermal gradients at the onset of major ice sheet expansion on Antarctica and progressive cooling
through the Neogene and Quaternary towards a bipolar glaciated world. This drove the diatoms into uptaking
silica more competitively (i.e., to use less silica in formation of their frustules). Increased mixing of the oceans
renews silica and other nutrients necessary for diatom growth in surface waters, especially in regions of
coastal and oceanic upwelling.
[edit] Collection
Living diatoms are often found clinging in great numbers to filamentous algae, or forming gelatinous
masses on various submerged plants. Cladophora is frequently covered with Cocconeis, an elliptically
shaped diatom; Vaucheria is often covered with small forms. Diatoms are frequently present as a brown,
slippery coating on submerged stones and sticks, and may be seen to "stream" with river current.
The surface mud of a pond, ditch, or lagoon will almost always yield some diatoms. They can be
made to emerge by filling a jar with water and mud, wrapping it in black paper and letting direct sunlight fall
on the surface of the water. Within a day, the diatoms will come to the top in a scum and can be isolated.
Since diatoms form an important part of the food of molluscs, tunicates, and fishes, the alimentary
tracts of these animals often yield forms that are not easily secured in other ways. Marine diatoms can be
collected by direct water sampling, though benthic forms can be secured by scraping barnacles, oyster
shells, and other shells.
This section uses text from Methods in Plant Histology.[37]
[edit] EST sequencing

The first insights into the properties of the P. tricornutum gene repertoire was described using 1,000
ESTs.[38] Subsequently, the number of ESTs was extended to 12,000 and the Diatom EST Database was
constructed for functional analyses.[39] These sequences have been used to make a comparative analysis
between P. tricornutum and the putative complete proteomes from the green alga Chlamydomonas
reinhardtii, the red alga Cyanidioschyzon merolae, and T. pseudonana.[40] The diatom EST database now
consists in over 200,000 ESTs from P. tricornutum (16 libraries) and T. pseudonana (7 libraries) cells grown
in a range of different conditions, many of which corresponding to different abiotic stresses (available at
http://www.biologie.ens.fr/diatomics/EST3/).[41]
[edit] Genome sequencing

The entire genomes of the centric diatom, Thalassiosira pseudonana (32.4 Mb),[42] and the pennate
diatom, Phaeodactylum tricornutum (27.4 Mb),[43] have been sequenced. Comparisons of the two fully
sequenced diatom genomes finds that the P. tricornutum genome includes fewer genes (10,402 opposed to
11,776) than T. pseudonana and no major synteny (gene order) could be detected between the two
genomes. T. pseudonana genes show an average of ~1.52 introns per gene as opposed to 0.79 in P.
tricornutum, suggesting recent widespread intron gain in the centric diatom.[43][44] Despite relatively recent
evolutionary divergence (90 million years), the extent of molecular divergence between centrics and
pennates indicates rapid evolutionary rates within the Bacillariophyceae compared to other eukaryotic
groups.[43] Comparative genomics also established that a specific class of transposable elements, the
Diatom Copia-like retrotransposons (or CoDis), has been significantly amplified in the P. tricornutum genome
with respect to T. pseudonana, constituting 5.8 and 1% of the respective genomes.[45]
Importantly, diatom genomics brought much information about the extent and dynamics of the
endosymbiotic gene transfer (EGT) process. Comparison of the T. pseudonana proteins with homologs in
other organisms suggested that hundreds have their closest homologs in the Plantae lineage. EGT towards
diatom genomes can be illustrated by the fact that the T. pseudonana genome encodes six proteins which
are most closely related to genes encoded by the Guillardia theta (cryptomonad) nucleomorph genome. Four
of these genes are also found in red algal plastid genomes, thus demonstrating successive EGT from red
algal plastid to red algal nucleus (nucleomorph) to heterokont host nucleus.[42] More recent phylogenomic
analyses of diatom proteomes provided evidence for a prasinophyte-like endosymbiont in the common
ancestor of chromalveolates as supported by the fact the 70% of diatom genes of Plantae origin are of green
lineage provenance and that such genes are also found in the genome of other stramenopiles. Therefore, it
was proposed that chromalveolates are the product of serial secondary endosymbiosis first with a green
algae, followed by a second one with a red algae that conserved the genomic footprints of the previous but
displaced the green plastid.[46] However, phylogenomic analyses of diatom proteomes and chromalveolate
evolutionary history will likely take advantage of complementary genomic data from under-sequenced
lineages such as red algae.
In addition to EGT, horizontal gene transfer (HGT) can occur independently of an endosymbiotic
event. The publication of the P. tricornutum genome reported that at least 587 P. tricornutum genes appear to
be most closely related to bacterial genes, accounting for more than 5% of the P. tricornutum proteome.
About half of these are also found in the T. pseudonana genome, attesting their ancient incorporation in the
diatom lineage.[43]
[edit] Nanotechnology research

The deposition of silica by diatoms may also prove to be of utility to nanotechnology.[47] Diatom cells
repeatedly and reliably manufacture valves of various shapes and sizes, potentially allowing diatoms to
manufacture micro- or nano-scale structures which may be of use in a range of devices, including: optical
systems; semiconductor nanolithography; and even using diatom valves as vehicles for drug delivery. Using
an appropriate artificial selection procedure, diatoms that produce valves of particular shapes and sizes
could be evolved in the laboratory, and then used in chemostat cultures to mass produce nanoscale
components.[48] It has also been proposed that diatoms could be used as a component of solar cells, by
substituting photosensitive titanium dioxide for the silicon dioxide normally used in the creation of cell walls.
[49]
[edit] See also
• Algae
• Biological pump
• Diatomaceous earth
• Phytoplankton
• Plankton
[edit] References
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24.^ Kidder, D. L. and Erwin, D. H. (2001). Secular distribution of biogenic silica through the
Phanerozoic: Comparison of silica-replaced fossils and bedded cherts at the series level. J. Geol.
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25.^ Grenne, T. and Slack, J. F. (2003). Paleozoic and Mesozoic silica-rich seawater: evidence
from hematitic chert (jasper) deposits. Geology 31, 319-322.
26.^ Racki, G. and Cordey, F. (2000). Radiolarian palaeoecology and radiolarites: is the present
the key to the past? Earth-Science Reviews 52, 83-120.
27.^ Maldonado, M., Carmona, M. C., Uriz, J. M. and Cruzado, A. (1999). Decline in Mesozoic
reef-building sponges explained by silicate limitation. Nature 401, 785-788.
28.^ Harper, H. E. and Knoll, A. H. (1975). Silica, diatoms, and Cenozoic radiolarian evolution.
Geology 3, 175-177.
29.^ Falkowski, P.G.; et al. (2004). "The evolution of modern eukaryotic phytoplankton". Science
305 (5682): 354–60. doi:10.1126/science.1095964. PMID 15256663.
30.^ Kidder, D.L.; Gierlowski-Kordesch, E.H. (2005). "Impact of grassland radiation on the
nonmarine silica cycle and Miocene diatomite". PALAIOS 20: 198–206. doi:10.2110/palo.2003.p03-
108.
31.^ Rabosky, D.L.; Sorhannus, U. (2009). "Diversity dynamics of marine planktonic diatoms
across the Cenozoic". Nature 457 (7226): 183–6. doi:10.1038/nature07435. PMID 19129846.
32.^ a b Scherer, R.P., Gladenkov, A.Yu., and Barron, J.A. (2007). Methods and applications of
Cenozoic marine diatom biostratigraphy. “Paleontological Society Papers” 13, 61-83
33.^ Harwood, D.M., Nikolaev, V.A., and Winter, D.M. (2007). Cretaceous record of diatom
evolution, radiation, and expansion. “Paleontological Society Papers” 13, 33-59
34.^ Strelnikova, N. I. (1990). Evolution of diatoms during the Cretaceous and Paleogene
periods. In: Simola, H. (ed.), “Proceedings of the Tenth International Diatom Symposium”, Koeltz
Scientific Books, Koenigstein, Germany, pp. 195-204
35.^ Baldauf, J.G. (1993). Middle Eocene through early Miocene diatom floral turnover. In:
Prothero D., Berggren, W.H., (eds.), “Eocene-Oligocene climatic and biotic evolution”, Princeton
University Press, Princeton, NJ, USA, pp 310-326
36.^ Barron, J.A. (2003). Appearance and extinction of planktonic diatoms during the past 18
m.y. in the Pacific and Southern oceans. “Diatom Research” 18, 203-224
37.^ Chamberlain, C. J. (1901) Methods in Plant Histology, University of Chicago Press, USA
38.^ Scala, S.; Carels, N., Falciatore, A., Chiusano, M.L. and Bowler, C. (July 2002). "Genome
properties of the diatom Phaeodactylum tricornutum". Plant Physiol. 129 (3): 993–1002.
doi:10.1104/pp.010713. PMID 12114555. PMC 166495.
http://www.plantphysiol.org/cgi/content/full/129/3/993.
39.^ Maheswari, U., Montsant, A., Goll, J., Krishnasamy, S., Rajyashri, K.R., Patell, V.M. and
Bowler, C. (2005). The Diatom EST Database. Nucleic Acids Research 33, 344–347.
40.^ Montsant, A.; Jabbari, K., Maheswari, U. and Bowler, C. (February 2005). "Comparative
genomics of the pennate diatom Phaeodactylum tricornutum". Plant Physiol. 137 (2): 500–13.
doi:10.1104/pp.104.052829. PMID 15665249.
41.^ Maheswari U, Mock T, Armbrust EV, Bowler C: Update of the Diatom EST Database: a new
tool for digital transcriptomics. Nucleic Acids Res 2008.
42.^ a b Armbrust et al. (2004). The genome of the diatom Thalassiosira pseudonana: ecology,
evolution, and metabolism. Science 306, 79-86.
43.^ a b c d Bowler, C.; et al. (2008). "The Phaeodactylum genome reveals the evolutionary
history of diatom genomes". Nature 456 (7219): 239–244. doi:10.1038/nature07410.
PMID 18923393.
44.^ Roy SW, Penny D. A very high fraction of unique intron positions in the intron-rich diatom
Thalassiosira pseudonana indicates widespread intron gain. Mol Biol Evol. 2007 Jul;24(7):1447-57.
Epub 2007 Mar 8. PubMed PMID: 17350938.
45.^ Maumus F, Allen AE, Mhiri C, Hu H, Jabbari K, Vardi A, Grandbastien MA, Bowler C.
Potential impact of stress activated retrotransposons on genome evolution in a marine diatom. BMC
Genomics. 2009 Dec 22;10:624.
46.^ Moustafa A, Beszteri B, Maier UG, Bowler C, Valentin K, Bhattacharya D: Genomic
footprints of a cryptic plastid endosymbiosis in diatoms. Science 2009, 324:1724-1726.
47.^ Bradbury, J. (2004). Nature's Nanotechnologists: Unveiling the Secrets of Diatoms. PLoS
Biology 2, 1512-1515.
48.^ Drum, R.W.; Gordon, R. (2003). "Star Trek replicators and diatom nanotechnology". Trends
Biotechnology 21: 325–328. doi:10.1016/S0167-7799(03)00169-0.
http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6TCW-48XCHCX-
3&_user=126770&_coverDate=08%2F31%2F2003&_rdoc=2&_fmt=high&_orig=browse&_srch=doc-
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49.^ Johnson, R.C. (9 April 2009). "Diatoms could triple solar cell efficiency". EE Times.
http://www.eetimes.com/showArticle.jhtml?articleID=216500176. Retrieved 13 April 2009.

Wikispecies has information related to: Diatoms
Wikimedia Commons has media related to: Diatoms
• Computer simulations of pattern formation in diatoms

• Catalogue of Diatom Names, California Academy of Sciences
• Diatom Genome, Joint Genome Institute
• Diatom EST database, École Normale Supérieure
• Plankton*Net, taxonomic database including images of diatom species
• Life History and Ecology of Diatoms, University of California Museum of Paleontology
• Diatoms: 'Nature's Marbles', Eureka site, University of Bergen
• Diatom life history and ecology, Microfossil Image Recovery and Circulation for Learning and
Education (MIRACLE), University College London
• Diatom page, Royal Botanic Garden Edinburgh
• Geometry and Pattern in Nature 3: The holes in radiolarian and diatom tests
• Art Deco Diatoms, Wim van Egmond
• Diatom QuickFacts, Monterey Bay Aquarium Research Institute
• Diatoms and other algae image database Academy of Natural Sciences in Philadelphia
(ANSP)
• Now, energy from single celled algae Deccan Herald
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W000
Ernst Haeckel

"Haeckel" redirects here. For other uses, see Haeckel (disambiguation).
Ernst Haeckel
Born February 16, 1834(1834-02-16)
Died August 9, 1919(1919-08-09) (aged 85)

Nationality German
Ernst Haeckel.
Ernst Heinrich Philipp August Haeckel (February 16, 1834 – August 9, 1919),[1] also written von
Haeckel, was an eminent German biologist, naturalist, philosopher, physician, professor and artist who
discovered, described and named thousands of new species, mapped a genealogical tree relating all life
forms, and coined many terms in biology, including anthropogeny, ecology, phylum, phylogeny, and the
kingdom Protista. Haeckel promoted and popularized Charles Darwin's work in Germany and developed the
controversial recapitulation theory ("ontogeny recapitulates phylogeny") claiming that an individual
organism's biological development, or ontogeny, parallels and summarizes its species' entire evolutionary
development, or phylogeny.
The published artwork of Haeckel includes over 100 detailed, multi-colour illustrations of animals and
sea creatures (see: Kunstformen der Natur, "Artforms of Nature"). As a philosopher, Ernst Haeckel wrote Die
Welträtsel (1895–1899, in English, The Riddle of the Universe, 1901), the genesis for the term "world riddle"
(Welträtsel); and Freedom in Science and Teaching [2] to support teaching evolution.
In the United States, Mount Haeckel, a 13,418 ft (4,090 m) summit in the Eastern Sierra Nevada,
overlooking the Evolution Basin, is named in his honor, as is another Mount Haeckel, a 2,941 m (9,649 ft)
summit in New Zealand; and the asteroid 12323 Haeckel.
The Ernst Haeckel house ("Villa Medusa") in Jena, Germany contains a historic library.
Contents
[hide]
• 1 Life
• 1.1 Politics
• 1.2 "First World War"
• 2 Research
• 3 Polygenism and racial theory
• 4 Embryology and recapitulation theory
• 4.1 Darwin, Naturphilosophie and Lamarck
• 4.2 Embryological drawings
• 4.3 Controversy
• 5 Awards
• 6 Publications
• 7 See also
• 8 Notes
• 9 References
[edit] Life
Sea anemones from Ernst Haeckel's Kunstformen der Natur (Artforms of Nature) of 1904.
Ernst Haeckel: Christmas of 1860 (age 26)
Ernst Haeckel was born on February 16, 1834, in Potsdam (then part of Prussia). [3] In 1852,
Haeckel completed studies at Cathedral High School ( Domgymnasium) of Merseburg.[3] He then studied
medicine in Berlin and Würzburg, particularly with Albert von Kölliker, Franz Leydig, Rudolf Virchow (with
whom he later worked briefly as assistant), and with anatomist-physiologist Johannes Peter Müller (1801–
1858).[3] Together with Hermann Steudner he attended botany lectures in Würzburg. In 1857, Haeckel
attained a doctorate in medicine (M.D.), and afterwards he received a license to practice medicine. The
occupation of physician appeared less worthwhile to Haeckel, after contact with suffering patients.[3]
Haeckel studied under Karl Gegenbaur at the University of Jena for three years, earning a doctorate
in zoology,[3] before becoming a professor of comparative anatomy at the University of Jena, where he
remained for 47 years, from 1862 to 1909. Between 1859 and 1866, Haeckel worked on many invertebrate
groups, including radiolarians, poriferans (sponges) and annelids (segmented worms).[4] During a trip to the
Mediterranean, Haeckel named nearly 150 new species of radiolarians.[4] Haeckel named thousands of new
species from 1859 to 1887.[5]
From 1866 to 1867, Haeckel made an extended journey to the Canary Islands with Hermann Fol and
during this period, met with Charles Darwin, in 1866 at Down House in Kent, Thomas Huxley and Charles
Lyell.[3] In 1867, he married Agnes Huschke. Their son Walter was born in 1868, their daughters Elizabeth in
1871 and Emma in 1873.[3] In 1869, he traveled as a researcher to Norway, in 1871 to Dalmatia, and in 1873
to Egypt, Turkey, and to Greece.[3] Haeckel retired from teaching in 1909, and in 1910 he withdrew from the
Evangelical church.[3] Haeckel's wife, Agnes, died in 1915, and Haeckel became substantially more frail,
with a broken leg (thigh) and broken arm.[3] He sold the mansion Medusa ("Villa Medusa") in 1918 to the
Carl Zeiss foundation.[3] Haeckel died on August 9, 1919.
[edit] Politics
Haeckel's political beliefs were influenced by his affinity for the German Romantic movement coupled
with his acceptance of a form of Lamarckism. Rather than being a strict Darwinian, Haeckel believed that
racial characteristics were acquired through interactions with the environment and that ontogeny directly
followed phylogeny. He believed the social sciences to be instances of "applied biology". Most of these
arguments have been shown to be over-generalizations at best and flatly incorrect at worst in modern biology
and social studies.[4] In 1905, Haeckel founded a group called the "Monist League" to promote his religious
and political beliefs. This group lasted until 1933 and included such notable members as Wilhelm Ostwald,
Georg von Arco, Helene Stöcker and Walter Arthur Berendsohn.[6]
[edit] "First World War"

Haeckel was the first person known to use the term "First World War". Shortly after the start of the
war Haeckel wrote:
There is no doubt that the course and character of the feared "European War"...will become the
first world war in the full sense of the word.
—Indianapolis Star, September 20, 1914[7]
The "European War" became known as "The Great War", and it was not until 1931, with the
beginning realization that another global war might be possible, that there is any other recorded use of the
term "First World War".[7]
[edit] Research
Haeckel (left) with Nicholai Miklukho-Maklai, his assistant, in the Canaries, 1866.
Haeckel was a zoologist, an accomplished artist and illustrator, and later a professor of comparative
anatomy. Although Haeckel's ideas are important to the history of evolutionary theory, and he was a
competent invertebrate anatomist most famous for his work on radiolaria, many speculative concepts that he
championed are now considered incorrect. For example, Haeckel described and named hypothetical
ancestral microorganisms that have never been found.
He was one of the first to consider psychology as a branch of physiology. He also proposed many
now ubiquitous terms including "anthropogeny", "phylum", "phylogeny", "ecology" ("oekologie"),[5] and
proposed the kingdom Protista[3] in 1866. His chief interests lay in evolution and life development processes
in general, including development of nonrandom form, which culminated in the beautifully illustrated
Kunstformen der Natur (Art forms of nature). Haeckel did not support natural selection, rather believing in a
Lamarckian inheritance of acquired characteristics (Lamarckism). [8]
Haeckel advanced a version of the earlier "recapitulation theory", previously set out by Étienne
Serres in the 1820s and supported by followers of Geoffroy including Robert Edmond Grant,[9] which
proposed a link between ontogeny (development of form) and phylogeny (evolutionary descent), summed up
by Haeckel in the phrase "ontogeny recapitulates phylogeny". His concept of recapitulation has been refuted
in the form he gave it (now called "strong recapitulation"), in favour of the ideas first advanced by Karl Ernst
von Baer. "Strong" recapitulation hypothesis views ontogeny as repeating forms of the ancestors, while
"weak" recapitulation means that what is repeated (and built upon) is the ancestral embryonic development
process.[10] He supported the theory with embryo drawings that have since been shown to be oversimplified
and in part inaccurate, and the theory is now considered an oversimplification of quite complicated
relationships. Haeckel introduced the concept of "heterochrony", which is the change in timing of embryonic
development over the course of evolution.
Haeckel was a flamboyant figure. He sometimes took great (and non-scientific) leaps from available
evidence. For example, at the time that Darwin first published On the Origin of Species by Means of Natural
Selection (1859), no remains of human ancestors had yet been found. Haeckel postulated that evidence of
human evolution would be found in the Dutch East Indies (now Indonesia), and described these theoretical
remains in great detail. He even named the as-of-yet unfound species, Pithecanthropus alalus, and charged
his students to go find it. (Richard and Oskar Hertwig were two of Haeckel's many important students.)
One student did find the remains: a young Dutchman named Eugene Dubois went to the East Indies
and dug up the remains of Java Man, the first human ancestral remains ever found. These remains originally
carried Haeckel's Pithecanthropus label, though they were later reclassified as Homo erectus.
[edit] Polygenism and racial theory

The creationist polygenism of Samuel George Morton and Louis Agassiz, which presented human
races as separately created species, was rejected by Charles Darwin, who argued for the monogenesis of
the human species and the recent African origin of modern humans. In contrast to most of Darwin's
supporters, Haeckel put forward a doctrine of evolutionary polygenism based on the ideas of the linguist
August Schleicher, in which several different language groups had arisen separately from speechless
prehuman Urmenschen, which themselves had evolved from simian ancestors. These separate languages
had completed the transition from animals to man, and, under the influence of each main branch of
languages, humans had evolved — in a kind of Lamarckian use-inheritance — as separate species, which
could be subdivided into races. From this Haeckel drew the implication that languages with the most potential
formed human species with the most potential, led by the Semitic and Indo-Germanic groups, with Berber,
Jewish, Greco-Roman and Germanic varieties to the fore.[11] As Haeckel stated:[12]
We must mention here one of the most important results of the comparative study of
languages, which for the Stammbaum of the species of men is of the highest significance, namely that
human languages probably had a multiple or polyphyletic origin. Human language as such probably
developed only after the species of speechless Urmenschen or Affenmenschen had split into several
species or kinds. With each of these human species, language developed on its own and
independently of the others. At least this is the view of Schleicher, one of the foremost authorities on
this subject.… If one views the origin of the branches of language as the special and principal act of
becoming human, and the species of humankind as distinguished according to their language stem,
then one can say that the different species of men arose independently of one another.
Haeckel's view can be seen as a forerunner of the multi-regional hypothesis, which until the 1990s
remained in contention with developments of Darwin's hypothesis of a recent African origin of modern
humans. The multiregional view then fell from favour, and Darwin's view has more recently been validated by
the decipherment of the human genome.
Haeckel also applied the hypothesis of polygenism to the modern diversity of human groups. He
became a leading proponent of scientific racism, stating for instance:[13]
The Caucasian, or Mediterranean man (Homo Mediterraneus), has from time immemorial
been placed at the head of all the races of men, as the most highly developed and perfect. It is
generally called the Caucasian race, but as, among all the varieties of the species, the Caucasian
branch is the least important, we prefer the much more suitable appellation proposed by Friedrich
Müller, namely, that of Mediterranese. For the most important varieties of this species, which are
moreover the most eminent actors in what is called "Universal History," first rose to a flourishing
condition on the shores of the Mediterranean.… This species alone (with the exception of the
Mongolian) has had an actual history; it alone has attained to that degree of civilization which seems to
raise men above the rest of nature.
[edit] Embryology and recapitulation theory
Illustrations of dog and human embryos, looking almost identical at 4 weeks then differing at 6
weeks, shown above a 6 week turtle embryo and 8 day hen embryo, presented by Haeckel in 1868 as
convincing proof of evolution. The pictures of the earliest embryonic stages are now considered inaccurate.
[14]
When Haeckel was a student in the 1850s he showed great interest in embryology, attending the
rather unpopular lectures twice and in his notes sketched the visual aids: textbooks had few illustrations, and
large format plates were used to show students how to see the tiny forms under a reflecting microscope, with
the translucent tissues seen against a black background. Developmental series were used to show stages
within a species, but inconsistent views and stages made it even more difficult to compare different species.
It was agreed by all European evolutionists that all vertebrates looked very similar at an early stage, in what
was thought of as a common ideal type, but there was a continuing debate from the 1820s between the
Romantic recapitulation theory that human embryos developed through stages of the forms of all the major
groups of adult animals, literally manifesting a sequence of organisms on a linear chain of being, and Karl
Ernst von Baer's opposing view that the early general forms diverged into four major groups of specialised
forms without ever resembling the adult of another species, showing affinity to an archetype but no relation to
other types or any transmutation of species. By the time Haeckel was teaching he was able to use a textbook
with woodcut illustrations written by his own teacher Albert von Kölliker, which purported to explain human
development while also using other mammalian embryos to claim a coherent sequence. Despite the
significance to ideas of transformism, this was not really polite enough for the new popular science writing,
and was a matter for medical institutions and for experts who could make their own comparisons.[15]
[edit] Darwin, Naturphilosophie and Lamarck

Darwin's On the Origin of Species, which made a powerful impression on Haeckel when he read it in
1864, was very cautious about the possibility of ever reconstructing the history of life, but did include a
section reinterpreting von Baer's embryology and revolutionising the field of study, concluding that
"Embryology rises greatly in interest, when we thus look at the embryo as a picture, more or less obscured, of
the common parent-form of each great class of animals." It mentioned von Baer's 1828 anecdote
(misattributing it to Louis Agassiz) that at an early stage embryos were so similar that it could be impossible
to tell whether an unlabelled specimen was of a mammal, a bird, or of a reptile, and Darwin's own research
using embryonic stages of barnacles to show that they are crustaceans, while cautioning against the idea
that one organism or embryonic stage is "higher" or "lower", or more or less evolved.[16] Haeckel
disregarded such caution, and in a year wrote his massive and ambitious Generelle Morphologie, published
in 1866, presenting a revolutionary new synthesis of Darwin's ideas with the German tradition of
Naturphilosophie going back to Goethe and with the progressive evolutionism of Lamarck in what he called
Darwinismus. He used morphology to reconstruct the evolutionary history of life, in the absence of fossil
evidence using embryology as evidence of ancestral relationships. He invented new terms, including
ontogeny and phylogeny, to present his evolutionised recapitulation theory that "ontogeny recapitulated
phylogeny". The two massive volumes sold poorly, and were heavy going: with his limited understanding of
German, Darwin found them impossible to read. Haeckel's publisher turned down a proposal for a “ strictly
scholarly and objective” second edition.[17]
[edit] Embryological drawings

Haeckel's aim was a reformed morphology with evolution as the organizing principle of a cosmic
synthesis unifying science, religion, and art. He was giving successful "popular lectures" on his ideas to
students and townspeople in Jena, in an approach pioneered by his teacher Rudolf Virchow. To meet his
publisher's need for a popular work he used a student's transcript of his lectures as the basis of his
Natürliche Schöpfungsgeschichte of 1868, presenting a comprehensive gospel of evolution. In the Spring of
that year he drew figures for the book, synthesising his views of specimens in Jena and published pictures to
represent types. After publication he told a colleague that the images “are completely exact, partly copied
from nature, partly assembled from all illustrations of these early stages that have hitherto become known.”
There were various styles of embryological drawings at that time, ranging from more schematic
representations to “naturalistic” illustrations of specific specimens. Haeckel believed privately that his figures
were both exact and synthetic, and in public asserted that they were schematic like most figures used in
teaching. The images were reworked to match in size and orientation, and though displaying Haeckel's own
views of essential features, they support von Baer's concept that vertebrate embryos begin similarly and then
diverge. Relating different images on a grid conveyed a powerful evolutionary message. As a book for the
general public, it followed the common practice of not citing sources.[18]
In 1868 Haeckel illustrated von Baer's observation that early embryos of different species could not
be told apart by using the same woodcut three times as dog, chick and turtle embryos: he changed this in the
next edition.
The book sold very well, and while some anatomical experts hostile to Haeckel's evolutionary views
expressed some private concerns that certain figures had been drawn rather freely, the figures showed what
they already knew about similarities in embryos. The first published concerns came from Ludwig Rütimeyer,
a professor of zoology and comparative anatomy at the University of Basel who had placed fossil mammals
in an evolutionary lineage early in the 1860s and had been sent a complimentary copy. At the end of 1868 his
review in the Archiv für Anthropologie wondered about the claim that the work was "popular and scholarly",
doubting whether the second was true, and expressed horror about such public discussion of man's place in
nature with illustrations such as the evolutionary trees being shown to non-experts. Though he made no
suggestion that embryo illustrations should be directly based on specimens, to him the subject demanded the
utmost "scrupulosity and conscientiousness" and an artist must “not arbitrarily model or generalize his
originals for speculative purposes” which he considered proved by comparison with works by other authors.
In particular, "one and the same, moreover incorrectly interpreted woodcut, is presented to the reader three
times in a row and with three different captions as [the] embryo of the dog, the chick, [and] the turtle." He
accused Haeckel of "playing fast and loose with the public and with science", and failing to live up to the
obligation to the truth of every serious researcher. Haeckel responded with angry accusations of bowing to
religious prejudice, but in the second (1870) edition changed the duplicated embryo images to a single image
captioned "embryo of a mammal or bird". Duplication using galvanoplastic stereotypes (clichés) was a
common technique in textbooks, but not on the same page to represent different eggs or embryos. In 1891
Haeckel made the excuse that this "extremely rash foolishness" had occurred in undue haste but was "bona
fide", and since repetition of incidental details was obvious on close inspection, it is unlikely to have been
intentional deception.[19]
The revised 1870 second edition of 1,500 copies attracted more attention, being quickly followed by
further revised editions with larger print runs as the book became a prominent part of the optimistic,
nationalist, anticlerical "culture of progress" in Otto von Bismarck's new German Empire. The similarity of
early vertebrate embryos became common knowledge, and the illustrations were praised by experts such as
Michael Foster of the University of Cambridge. In the introduction to his 1871 The Descent of Man, and
Selection in Relation to Sex, Darwin gave particular praise to Haeckel, writing that if Natürliche
Schöpfungsgeschichte "had appeared before my essay had been written, I should probably never have
completed it." The first chapter included an illustration: "As some of my readers may never have seen a
drawing of an embryo, I have given one of man and another of a dog, at about the same early stage of
development, carefully copied from two works of undoubted accuracy" with a footnote citing the sources and
noting that "Häckel has also given analogous drawings in his Schöpfungsgeschichte." The fifth edition of
Haeckel's book appeared in 1874, with a controversial frontispiece featuring the heads of apes and humans
replaced by a heroic portrait of Haeckel himself.[20]
1874 illustration from Anthropogenie showing "very early", "somewhat later" and "still later" stages of
embryos of fish (F), salamander (A), turtle (T), chick (H), pig (S), cow (R), rabbit (K), and human (M).
[edit] Controversy
Later in 1874, Haeckel's simplified embryology textbook Anthropogenie made the subject into a
battleground over Darwinism aligned with Bismarck's Kulturkampf ("culture struggle") against the Catholic
Church. Haeckel took particular care over the illustrations, changing to the leading zoological publisher
Wilhelm Engelmann of Leipzig and obtaining from them use of illustrations from their other textbooks as well
as preparing his own drawings including a dramatic double page illustration showing "early", "somewhat
later" and "still later" stages of 8 different vertebrates. Though Haeckel's views had attracted continuing
controversy, there had been little dispute about the embryos and he had many expert supporters, but
Wilhelm His now revived the earlier criticisms and introduced new attacks on the 1874 illustrations.[21]
Others joined in, both expert anatomists and Catholic priests and supporters politically opposed to Haeckel's
views.[22]
While it has been widely claimed that Haeckel was charged with fraud by five professors and
convicted by a university court at Jena, there does not appear to be an independently verifiable source for
this claim.[23] Recent analyses (Richardson 1998, Richardson and Keuck 2002) have found that some of the
criticisms of Haeckel's embryo drawings were legitimate, but others were unfounded.[24] [25] There were
multiple versions of the embryo drawings, and Haeckel rejected the claims of fraud. It was later said that
"there is evidence of sleight of hand" on both sides of the feud between Haeckel and Wilhelm His.[26] The
controversy involves several different issues (see more details at: recapitulation theory).
Some creationists have claimed that Darwin relied on Haeckel's embryo drawings as proof of
evolution[27] to support their anti-evolution arguments while both On the Origin of Species (1859), and The
Descent of Man (1871) were published before Haeckel's double-page illustration of eight vertebrate embryos
in 1874.[28]
[edit] Awards
He was awarded the Linnean Society of London's prestigious Darwin-Wallace Medal in 1908.
[edit] Publications
Kunstformen — plate 72: Muscinae

Kunstformen — plate 96: Chaetopoda
Darwin’s 1859 book On the Origin of Species had immense popular influence, but although its sales
exceeded its publisher's hopes it was a technical book rather than a work of popular science: long, difficult
and with few illustrations. One of Haeckel's books did a great deal to explain his version of "Darwinism" to the
world. It was a bestselling, provocatively illustrated book in German, titled Natürliche Schöpfungsgeschichte,
published in Berlin in 1868, and translated into English as The History of Creation in 1876. It was frequently
reprinted until 1926.
Haeckel argued that human evolution consisted of precisely 22 phases, the 21st — the "missing link"
— being a halfway step between apes and humans. He even formally named this missing link
Pithecanthropus alalus, translated as "ape man without speech." (The missing link was what the Dutchman
Eugène Dubois, discoverer of Homo erectus, would later resolve to find.)[citation needed]
Haeckel's entire literary output was extensive, working as a professor at the University of Jena for 47
years, and even at the time of the celebration of his 60th birthday at Jena in 1894, Haeckel had produced 42
works with nearly 13,000 pages, besides numerous scientific memoirs and illustrations. [29]
Haeckel's monographs include:
• Radiolaria (1862)
• Siphonophora (1869)
• Monera (1870)
• Calcareous Sponges (1872)
As well as several Challenger reports:
• Deep-Sea Medusae (1881)
• Siphonophora (1888)
• Deep-Sea Keratosa (1889)
• Radiolaria (1887) — illustrated with 140 plates and enumerating over four thousand (4000)
new species.[29]
Among his many books, Ernst Haeckel wrote:
• Generelle Morphologie der Organismen : allgemeine Grundzüge der organischen Formen-
Wissenschaft, mechanisch begründet durch die von C. Darwin reformirte Decendenz-Theorie. (1866)
Berlin
'Monophyletischer Stambaum der Organismen' from 'Generelle Morphologie der Organismen' (1866)
with the three branches Plantae, Protista, Animalia.
• Natürliche Schöpfungsgeschichte (1868) — in English The History of Creation (1876; 6th ed.:
New York, D. Appleton and Co., 1914, 2 volumes)
• Freie Wissenschaft und freie Lehre (1877), in English, Freedom in Science and Teaching , a
reply to a speech in which Rudolf Virchow objected to the teaching of evolution in schools, on the
grounds that evolution was an unproven hypothesis.[29]
• Die systematische Phylogenie (1894) — "Systematic Phylogeny", which has been considered
as his best book[29]
• Anthropogenie (1874, 5th and enlarged edition 1903) — dealing with the evolution of man
• Die Welträthsel (1895–1899), also spelled Die Welträtsel ("world-riddle") — in English The
Riddle of the Universe, 1901[29]
• Über unsere gegenwärtige Kenntnis vom Ursprung des Menschen (1898) — translated into
English as The Last Link, 1808
• Der Kampf um den Entwickelungsgedanken (1905) — English version, Last Words on
Evolution, 1906
• Die Lebenswunder (1904) — English "Wonder of Life", a supplement to the Riddle of the
Universe
Books of travel:
• Indische Reisebriefe (1882) — "Travel notes of India"
• Aus Insulinde: Malayische Reisebriefe (1901) — "Travel notes of Malaysia", the fruits of
journeys to Ceylon and to Java
• Kunstformen der Natur (1904) — Artforms of Nature, with plates representing detailed marine
animal forms
• Wanderbilder (1905) — "Travel Images", with reproductions of his oil-paintings and water-
color landscapes.[29]
• A visit to Ceylon
The standard author abbreviation Haeckel is used to indicate this individual as the author when citing
a botanical name.[30]
[edit] See also

• Alternative taxonomical classification
• Dysteleology
• Embryology
• Francis Galton
• List of wildlife artists
• Proteus (2004 film), an animated documentary by David Lebrun, largely focussing on Ernst
Haeckel
• Haeckel's Tale, a horror film by John McNaughton, featuring a fictionalized version of Ernst
Haeckel
[edit] Notes
1. ^ "Ernst Haeckel — Britannica Concise" (biography), Encyclopædia Britannica Concise, 2006,
Concise. Britannica.com webpage: CBritannica-Haeckel.
2. ^ Freedom in Science and Teaching. German 1877, English 1879, ISBN 1410211754.
3. ^ a b c d e f g h i j k l "Ernst Haeckel" (article), German Wikipedia, October 26, 2006, webpage:
DE-Wiki-Ernst-Haeckel: last paragraph of "Leben" (Life) section.
4. ^ a b c "Ernst Haeckel" (biography), UC Berkeley, 2004, webpage: BerkeleyEdu-Haeckel.
5. ^ a b "Rudolf Steiner and Ernst Haeckel" (colleagues), Daniel Hindes, 2005,
DefendingSteiner.com webpage: Steiner-Haeckel.
6. ^ Health, Race and German Politics Between National Unification and Nazism by Paul
Weindling, Cambridge University Press, 1993.,pgs. 46, 250
7. ^ a b Fred R. Shapiro, ed (2006). The Yale Book of Quotations. Yale University Press.
pp. 329. ISBN 9780300107982. http://books.google.com/?id=w5-GR-
qtgXsC&printsec=frontcover&dq=The+Yale+book+of+quotations#PPA329,M1. Retrieved 2008-10-
08.
8. ^ Ruse, M. 1979. The Darwinian Revolution. Chicago: University of Chicago Press.
9. ^ Desmond 1989, pp. 53–53, 86–88, 337–340
10.^ Richardson and Keuck, (Biol. Review (2002), 77, pp. 495–528) show that it is a
simplification to suppose that Haeckel held the recapitulation theory in its strong form. They quote
Haeckel as saying "If [recapitulation] was always complete, it would be a very easy task to construct
whole phylogeny on the basis of ontogeny. ... There is certainly, even now, a number of lower
vertebrate animals (e.g. some Anthozoa and Vermes) where we are authorised to interpret each
embryological form directly as the historical representation or portrait-like silhouette of an extinct
ancestral form. But in a great majority of animals, including man, this is not possible because the
infinitely varied conditions of existence have led the embryonic forms themselves to be changed and
to partly lose their original condition (Haeckel, 1903: pp. 435–436)"
11.^ Richards, Robert W. (2008). The Tragic Sense of Life: Ernst Haeckel and the Struggle over
Evolutionary Thought. Chicago: University of Chicago Press. pp. 259–260. ISBN 0-226-71214-1.
12.^ Natürliche Schöpfungsgeschichte (1868), p. 511; quoted after Robert J. Richards, "The
linguistic creation of man: Charles Darwin, August Schleicher, Ernst Haeckel, and the Missing Link in
Nineteenth-Century Evolutionary Theory".[1]
13.^ The History of Creation, 6th edition (1914), volume 2, page 429.
14.^ Richardson MK, Hanken J, Selwood L, Wright GM, Richards RJ, Pieau C, Raynaud A
(1998). "Letters". Science 280 (5366): 983, 985–6. PMID 9616084.
15.^ Hopwood 2006, pp. 264–267
Darwin & Costa 2009, p. 450
16.^ Darwin 1859, pp. 439–450
Darwin & Costa 2009, pp. 439–450
17.^ Hopwood 2006, pp. 2269–270
18.^ Hopwood 2006, pp. 270–274
19.^ Hopwood 2006, pp. 275–276, 282–286
20.^ Hopwood 2006, pp. 285–288
Darwin 1871, pp. 4, 14–17
21.^ Wilhelm His: Unsere Körperform und das physiologische Problem ihrer Entstehung . F.C.W.
Vogel, Leipzig 1875.
22.^ Hopwood 2006, pp. 288–296
23.^ "Ernst Haeckel and the Struggles over Evolution and Religion" Robert J. Richards Annals
of the History and Philosophy of Biology, Vol. 10 (2005): 89-115
24.^ Michael K. Richardson. 1998. "Haeckel's embryos continued." Science 281:1289, quoted in
NaturalScience.com webpage Re: Ontogeny and phylogeny: A Letter from Richard Bassetti; Editor's
note.
25.^ "While some criticisms of the drawings are legitimate, others are more tenditious",
Richardson and Keuck "Haeckel's ABC of evolution and development", Biol. Rev. (2002), 77, pp.
495–528. Quoted from p. 495.
26.^ Richardson & Keuck 2001. See for example, their Fig. 7, showing His's drawing of the
forelimb of a deer embryo developing a clef, compared with a similar drawing (Sakurai, 1906)
showing the forelimb initially developing as a digital plate with rays. Richardson & Keuck say
"Unfortunately His's embryos are mostly at later stages than the nearly identical early stage embryos
illustrated by Haeckel [top row of Haeckel's drawing]. Thus they do not inform the debate and may
themselves be disingenuous.", p. 518.
27.^ "Darwin relied on the work of German biologist Ernst Haeckel ... Darwin based his
inference of common ancestry on the belief that the earliest stages of embryo development are the
most similar. Haeckel's drawings, however, entirely omit the earliest stages ...", Jonathan Wells,
Survival of the Fakest, The American Spectator, December 2000–January 2001. Note however,
Darwin (1871) credits Huxley with the idea of comparing the embryos and quoted a statement by T.
Huxley, that it is "quite in the later stages of development that the young human being presents
marked differences from the young ape ..." (from Huxley's Man’s Place in Nature, 1863, p. 67). Note
the subtle difference between Huxley’s claim — the final stages are most different — and what has
been said Darwin relied on via Haeckel – that the earliest stages are the most similar.
28.^ Kurt M. Pickett; John W. Wenzel and Steven W. Rissing (May 2005). "Iconoclasts of
Evolution: Haeckel, Behe, Wells and the Ontogeny of a Fraud" (PDF). The American Biology
Teacher.
http://www.socialwasps.com/Pickett_Lab_of_Vespid_Taxonomy/Publications_files/Pickett_et_al_200
5a.pdf.
29.^ a b c d e f "Biography of Ernst Heinrich Haeckel, 1834–1919" (article), Missouri Association
for Creation, Inc., based on 1911 Britannica, webpage: Gennet-Haeckel: life, career & beliefs.
30.^ "Author Query". International Plant Names Index.
http://www.ipni.org/ipni/authorsearchpage.do.
[edit] References
• Charles Darwin (1859). On the Origin of Species (by Means of Natural Selection). London:
John Murray. ISBN 8420656070.
• Charles Darwin (2003 edition). The Origin of Species (with introduction by Julian Huxley).
Signet Classics. ISBN 0-451-52906-5.
• Desmond, Adrian J. (1989). The politics of evolution: morphology, medicine, and reform in
radical London. Chicago: University of Chicago Press. ISBN 0-226-14374-0.
• Ernst Haeckel, Freedom in Science and Teaching (1879), reprint edition, University Press of
the Pacific, February 2004, paperback, 156 pages, ISBN 1-4102-1175-4.
• Ernst Haeckel, The History of Creation (1868), translated by E. Ray Lankester, Kegan Paul,
Trench & Co., London, 1883, 3rd edition, Volume 1.
• Ernst Haeckel, Kunstformen der Natur ("Artforms of Nature"), 1904, (from series published
1899–1904): over 100 detailed, multi-color illustrations of animals and sea creatures.
• Ernst Haeckel, Lebenswunder, Stuttgart, 1904.
• Ernst Haeckel, The Riddle of the Universe (Die Weltraetsel, 1895–1899), Publisher:
Prometheus Books, Buffalo, NY, 1992, reprint edition, paperback, 405 pages, illustrated, ISBN 0-
87975-746-9.
• Richard Milner, The Encyclopedia of Evolution: Humanity's Search for Its Origins , Henry Holt,
1993.
• Robert J. Richards, The Tragic Sense of Life: Ernst Haeckel and the Struggle over
Evolutionary Thought, University of Chicago Press, 2008.
• Richardson Michael K.. "Haeckel's embryos continued" (article)". Science 281 (1289): 1998.
• Richardson, M. K. & Keuck, G. (2001) "A question of intent: when is a 'schematic' illustration
a fraud?," Nature 410:144 (vol. 410, no. 6825, page 144), March 8, 2001.
• Richardson, M. K. & Keuck, G. (2002) Haeckel's ABC of evolution and development
Biological Reviews (2002), 77: 495–528
• M. Ruse, The Darwinian Revolution. Chicago: University of Chicago Press, 1979.
• Newman, H.H., 1932, 3rd edition, Evolution, Genetics, and Eugenics , Chicago: University of
Chicago Press, p. 30
• G.G. Simpson and W. Beck, An Introduction to Biology (New York: Harcourt Brace and
World, 1965), p. 241
• New Scientist, 9/6/97, p. 23
• W. Bock, Book Review Science, May 1969, pp. 684–685

• Di Gregorio, Mario A. From here to eternity: Ernst Haeckel and Scientific Faith , Vandenhoeck
& Ruprecht, 2005, ISBN 3525569726
• Haeckel, Ernst. (1900). The Riddle of the Universe at the Close of the Nineteenth Century .
Harper (reissued by Cambridge University Press, 2009; ISBN 9781108000895)
• Haeckel, Ernst, Art Forms from the Ocean: The Radiolarian Atlas of 1862 , Prestel Verlag,
2005, ISBN 3-7913-3327-5.
• Works by Ernst Haeckel at Project Gutenberg.
• Richardson, Michael K., "Haeckel, embryos, and evolution," Science Vol. 280, no. 5366 (May
15, 1998) p. 983, 985–986.
• Spiro, Jonathan P. (2009). Defending the Master Race: Conservation, Eugenics, and the
Legacy of Madison Grant. Univ. of Vermont Press. ISBN 978-1-58465-715-6. Lay summary (29
September 2010).

Wikimedia Commons has media related to: Ernst Haeckel
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Author:Ernst Haeckel
• Woods Hole Marine Biological Laboratory Library — An exhibition of material on Haeckel,
including background on many Kunstformen der Natur plates
• University of California, Berkeley — Ernst Haeckel biography
• Ernst Haeckel – Evolution's controversial artist. A slide-show essay about Ernst Haeckel.
• Kunstformen der Natur, Wikimedia Commons: over 100 detailed animal drawings.
• Kunstformen der Natur, scanned (from biolib.de Stuebers Online Library)
• PNG alpha-transparencies of Haeckel's "Kustformen der natur"
• Proteus — An animated documentary film on the life and work of Ernst Haeckel
• Ernst Haeckel Haus and Ernst Haeckel Museum in Jena
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Name Haeckel, Ernst Heinrich Philipp August
Alternative von Haeckel, Ernst Heinrich Philipp August; Haeckel, Ernst Heinrich
names Philipp August von
Short description German biologist and philosopher
Date of birth February 16, 1834(1834-02-16)
Place of birth Potsdam, Prussia
Date of death August 9, 1919(1919-08-09)
Place of death
Retrieved from "http://en.wikipedia.org/wiki/Ernst_Haeckel"
Categories: Botanists with author abbreviations | 1834 births | 1919 deaths | People from Potsdam |
German religious humanists | German biologists | German-language philosophers | German zoologists |
Evolutionary biologists | Protistologists | Racism | University of Jena faculty | People from the Province of
Brandenburg
W000
Natural history

For other uses, see Natural history (disambiguation).
"Naturalist" redirects here. For other uses, see Naturalist (disambiguation).
Tables of natural history, from the 1728 Cyclopaedia
Natural history is the scientific research of plants or animals, leaning more towards observational
rather than experimental methods of study, and encompasses more research published in magazines than in
academic journals.[1] Grouped among the natural sciences, natural history is the systematic study of any
category of natural objects or organisms. That is a very broad designation in a world filled with many narrowly
focused disciplines. So while modern natural history dates historically from studies in the ancient Greco-
Roman world and the medieval Arabic world through to the scattered European Renaissance scientists
working in near isolation, today's field is more of a cross discipline umbrella of many specialty sciences. For
example, geobiology has a strong multi-disciplinary nature combining scientists and scientific knowledge of
many specialty sciences.
A person who studies natural history is known as a naturalist or "natural historian". Natural history is
categorized among the natural sciences. As a published topic, it originated from studies in the ancient Greco-
Roman world. The modern topic comprises many specialty sciences such as geobiology.
Contents
[hide]
• 1 Definitions
• 1.1 Historical
• 1.2 Modern
• 2 History
• 3 Museums
• 4 Societies
• 5 See also
• 6 References
[edit] Definitions
challenged and removed. (November 2009)
[edit] Historical
The English term 'natural history' is a translation of the Latin naturalis historia. Its meaning has
narrowed considerably over time (see also History below). In antiquity, it covered more-or-less anything
which is connected with nature or which uses materials drawn from nature; see for example the contents of
Pliny's encyclopedia of this title, published circa AD 77-79.
Until well into the nineteenth century, knowledge was considered by Europeans to have two main
divisions: the humanities (including theology), and studies of nature. Studies of nature could in turn be
divided, with natural history being the descriptive counterpart to natural philosophy which was the analytical
study of nature. In modern terms, natural philosophy roughly corresponded to modern physics and chemistry,
while natural history included the biological and geological sciences. The two were strongly associated.
During the heyday of the gentleman scientists, many figures contributed to both fields, and early papers in
both were commonly read at professional science society meetings such as the Royal Society and the
French Academy of Sciences – both founded during the seventeenth century.
[edit] Modern
The growth of many separate scientific disciplines in the twentieth century altered the way in which
the term 'natural history' was used. Since it encompasses research that is now normally published within
distinct disciplines, it may be considered an archaic or popular term.
Although terminology was and remains somewhat vague, a number of increasingly restricted uses
can be distinguished. The less restricted uses are 'umbrella terms' for distinct modern scientific disciplines.
Modern uses exclude chemistry and almost all of physics (astronomy is sometimes included).
• As used in the titles of institutions such as the American Museum of Natural History or the
British Natural History Museum, the term covers most of modern biology and geology.
• A more restricted use excludes those areas of geology not concerned with living organisms.
In this sense, natural history includes all of biology (the study of living organisms such as plants,
animals, fungi, bacteria, etc. and their relationships in natural systems) and paleobiology (the study
of extinct life), but only some life-related areas of geology, such as stratigraphy and petrology.
• Applied only within biology, it is used for the study of particular organisms. Thus the 'natural
history of primates' involves describing the relevant structures, operations and circumstances of
primates, such as their diet, reproduction, social grouping, and interactions with other species.[2]
The term may be used to denote the less strictly organized study, description, and classification of
natural objects, such as animals, plants, minerals, which emphasise fieldwork as opposed to more
systematic scientific investigation such as experimental or laboratory work.[3]
Modern definitions of the term include:
• Natural history is "the scientific study of plants or animals (more observational than
experimental) usually published in popular magazines rather than in academic journals".[1]
• "Natural history is the scientific research of plants and animals in their natural environments.
It is concerned with degrees of organization from individual organisms to an entire ecosystem, and
emphasizes identification, life history, distribution, abundance, and inter-relationships. It may include
an aesthetic component."[4]
[edit] History
Natural history begins with Aristotle and other ancient philosophers who analyzed the diversity of the
natural world. Natural history, as a discipline, had existed since classical times, and fifteenth-century
Europeans were very familiar with Pliny the Elder's Historia Naturalis. From the ancient Greeks until the work
of Carolus Linnaeus (also known as Carl Linnaeus, or Carl von Linné) and other 18th century naturalists, the
main concept of natural history was the scala naturae or Great Chain of Being, a conceptual arrangement of
minerals, vegetables, more primitive forms of animals, and more complex life forms on a linear scale of
increasing "perfection", culminating in our species.
Dioscorides' De Materia Medica is often said to be the oldest and most valuable work in the history of
botany.[5] A Greek manuscript of Aristotle's Biological Works, written in Constantinople in the mid-9th
century, and preserved at Corpus Christi College, Oxford, is probably the oldest surviving manuscript of texts
that founded the science of biology.[6]
While natural history was basically static in medieval Europe, it continued to be developed by Arabic
scholars during the Arab Agricultural Revolution. Al-Jahiz described early natural history ideas such as the
"struggle for existence" (Malthus' phrase),[7] and the idea of a food chain.[8][verification needed] He was an
early adherent of environmental determinism.[9][verification needed] Al-Dinawari is considered the founder of
Arabic botany for his Book of Plants, in which he described at least 637 plants and discussed plant
development from germination (sprouting) to death, describing the phases of plant growth and the production
of flowers and fruit.[10] Abu al-Abbas al-Nabati developed an early scientific method for botany, introducing
empirical and experimental techniques in the testing, description and identification of numerous materia
medica, and separating unverified reports from those supported by actual tests and observations.[11] His
student Ibn al-Baitar wrote a pharmaceutical encyclopedia describing 1,400 plants, foods, and drugs, 300 of
which were his own original discoveries. A Latin translation of his work was useful to European biologists and
pharmacists in the 18th and 19th centuries.[12] Earth sciences such as geology were also studied
extensively by Arabic geologists, but by Avicenna's time, around 1000, the Arab Empire was in decline and
scientists were not free to publish their ideas.[13]
Georges Buffon is best remembered for his Histoire naturelle, a 44 volume encyclopedia describing
everything known about the natural world.
From the 13th century, the work of Aristotle was adapted rather rigidly into Christian philosophy,
particularly by Thomas Aquinas, forming the basis for natural theology. During the Renaissance, scholars
(herbalists and humanists, particularly) returned to direct observation of plants and animals for natural
history, and many began to accumulate large collections of exotic specimens and unusual monsters. Andrea
Cesalpino was the creator of one of the first herbaria and the inventor of botanical systematics. Leonhart
Fuchs was one of the three founding fathers of botany, along with Otto Brunfels and Hieronymus Bock.
Important contributors to the field were also Valerius Cordus, Konrad Gesner (Historiae animalium), Frederik
Ruysch, or Gaspard Bauhin.[6] The rapid increase in the number of known organisms prompted many
attempts at classifying and organizing species into taxonomic groups, culminating in the system of the
Swedish naturalist Carl Linnaeus.[6]
In modern Europe, professional disciplines such as physiology, botany, zoology, geology, and
palaeontology were formed. Natural history, formerly the main subject taught by college science professors,
was increasingly scorned by scientists of a more specialized manner and relegated to an "amateur" activity,
rather than a part of science proper. In Victorian Scotland it was believed that the study of natural history
contributed to good mental health.[14] Particularly in Britain and the United States, this grew into specialist
hobbies such as the study of birds, butterflies, seashells (malacology/conchology), beetles and wildflowers;
meanwhile, scientists tried to define a unified discipline of biology (though with only partial success, at least
until the modern evolutionary synthesis). Still, the traditions of natural history continue to play a part in the
study of biology, especially ecology (the study of natural systems involving living organisms and the inorganic
components of the Earth's biosphere that support them), ethology (the scientific study of animal behavior),
and evolutionary biology (the study of the relationships between life-forms over very long periods of time),
and re-emerges today as integrative organismal biology.
Amateur collectors and natural history entrepreneurs played an important role in building the large
natural history collections of the nineteenth and early twentieth centuries, such as the Smithsonian
Institution's National Museum of Natural History.
[edit] Museums
Further information: List of natural history museums
Natural history museums, which evolved from cabinets of curiosities, played an important role in the
emergence of professional biological disciplines and research programs. Particularly in the 19th century,
scientists began to use their natural history collections as teaching tools for advanced students and the basis
for their own morphological research.
[edit] Societies
The term "natural history" alone, or sometimes together with archeology, forms the name of many
national, regional and local natural history societies that maintain records for birds (ornithology), mammals
(mammalogy), insects (entomology), fungi (mycology) and plants (botany). They may also have
microscopical and geological sections.
Examples of these societies in Britain include the Natural History Society of Northumbria founded in
1829, British Entomological and Natural History Society founded in 1872, Birmingham Natural History
Society, Glasgow Natural History Society, London Natural History Society founded in 1858, Manchester
Microscopical and Natural History Society established in 1880, Scarborough Field Naturalists' Society and
the Sorby Natural History Society, Sheffield, founded in 1918. The growth of natural history societies was
also spurred due to the growth of British colonies in tropical regions with numerous new species to be
discovered. Many civil servants took an interest in their new surroundings, sending specimens back to
museums in Britain. (See also Indian natural history)
[edit] See also

Environment portal
Ecology portal
• Big History
• Natural environment
• Natural landscape
• Natural philosophy
• Natural science
• Naturalism (philosophy)
• Nature documentary
• Nature writing
• Nature
• Nature study
• Terra: The Nature of Our World (video podcast)
-----
• Parson-naturalist
• Russian naturalists
[edit] References
Citations and notes
1. ^ a b Natural History WordNet Search, princeton.edu.
2. ^ Primate Glossary - National Zoo| FONZ
3. ^ nature glossary
4. ^ Herman, Stephen G. (2002), "Wildlife biology and natural history: time for a reunion",
Journal of Wildlife Management 66 (4): 933–946, doi:10.2307/3802927,
http://jstor.org/stable/3802927
5. ^ Gulsel M. Kavalali (2003). "Urtica: therapeutic and nutritional aspects of stinging nettles ".
CRC Press. p.15. ISBN 041530833X
6. ^ a b c "Natural History Timeline". HistoryofScience.com.
7. ^ Conway Zirkle (1941), Natural Selection before the "Origin of Species", Proceedings of the
American Philosophical Society 84 (1): 71-123.
8. ^ Frank N. Egerton, "A History of the Ecological Sciences, Part 6: Arabic Language Science -
Origins and Zoological", Bulletin of the Ecological Society of America , April 2002: 142-146 [143]
9. ^ Lawrence I. Conrad (1982), "Taun and Waba: Conceptions of Plague and Pestilence in
Early Islam", Journal of the Economic and Social History of the Orient 25 (3), pp. 268-307 [278].
10.^ Fahd, Toufic, "Botany and agriculture", pp. 815 , in Morelon, Régis; Rashed, Roshdi (1996),
Encyclopedia of the History of Arabic Science, 3, Routledge, ISBN 0415124107
11.^ Huff, Toby (2003), The Rise of Early Modern Science: Islam, China, and the West ,
Cambridge University Press, p. 218, ISBN 0521529948
12.^ Diane Boulanger (2002), "The Islamic Contribution to Science, Mathematics and
Technology", OISE Papers, in STSE Education, Vol. 3.
13.^ Richard Myers (2003). "The Basics of Chemistry". Greenwood Publishing Group. p.13.
ISBN 0313316643
14.^ Diarmid A. Finnegan (2008), "‘An aid to mental health’: natural history, alienists and
therapeutics in Victorian Scotland", Studies in History and Philosophy of Biological and Biomedical
Sciences 39 (3): 326–337, doi:10.1016/j.shpsc.2008.06.006, PMID 18761284
General information
• Allen, David Elliston (1994), The Naturalist in Britain: a social history, New Jersey: Princeton
University Press, pp. 270, ISBN 0-691-03632-2
• Atran, Scott (1990), Cognitive Foundations of Natural History: Towards an Anthropology of
Science, Cambridge, UK: Cambridge University Press, pp. 376, ISBN 978-0521438711
• Kohler, Robert E. Landscapes and Labscapes: Exploring the Lab-Field Border in Biology.
University of Chicago Press: Chicago, 2002.
• Mayr, Ernst. The Growth of Biological Thought: Diversity, Evolution, and Inheritance . The
Belknap Press of Harvard University Press: Cambridge, Massachusetts, 1982.
• Rainger, Ronald; Keith R. Benson; and Jane Maienschein, editors. The American
Development of Biology. University of Pennsylvania Press: Philadelphia, 1988.
Wikimedia Commons has media related to: Natural history
• A History of the Ecological Sciences by Frank N. Egerton

Institutions Rothamsted Experimental Station • Pasteur Institute •

Zoologica • Marine Biological Laboratory • Rockefeller University •
Biology
Griffith's experiment • Miller-Urey experiment • Luria–
Experiments Hershey–Chase experiment • Meselson–Stahl experiment • Crick,



Oparin • Alexander Fleming • J. B. S. Haldane • Sewall Wright • R.

Retrieved from "http://en.wikipedia.org/wiki/Natural_history"
Categories: Natural history | History of science | History of earth science | History of biology
W000
Honeyeater

For the group of Southern Asian birds formerly known as honeyeaters, see Sunbird or specific
members such as the Purple sunbird.
Honeyeaters
Female Crescent Honeyeater, Phylidonyris pyrrhopterus
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Passeriformes
Suborder: Passeri
Superfamily: Meliphagoidea
Meliphagidae
Family:
Vigors, 1825
Genera[3]
• Acanthagenys
• Acanthorhynchus
• Anthochaera
• Anthornis
• Ashbyia
• Certhionyx[1]
• Conopophila
• Entomyzon
• Epthianura
• Foulehaio
• Glycichaera
• Grantiella
• Guadalcanaria
• Gymnomyza
• Lichenostomus
• Lichmera
• Macgregoria
• Manorina
• Meliarchus
• Melidectes
• Melilestes
• Meliphaga
• Melipotes
• Melithreptus
• Melitograis
• Myza
• Myzomela
• Oreornis
• Philemon
• Phylidonyris[1]
• Plectorhyncha
• Prosthemadera
• Ptiloprora
• Pycnopygius
• Ramsayornis
• Stresemannia
• Timeliopsis
• Trichodere
• Xanthomyza[2]
• Xanthotis
The honeyeaters are a large and diverse family of small to medium sized birds most common in
Australia and New Guinea, but also found in New Zealand, the Pacific islands as far east as Samoa and
Tonga, and the islands to the north and west of New Guinea known as Wallacea. Bali, on the other side of
the Wallace Line, has a single species.
Honeyeaters and the Australian chats make up the family Meliphagidae. In total there are 182
species in 42 genera, roughly half of them native to Australia, many of the remainder occupying New Guinea.
With their closest relatives, the Maluridae (Australian fairy-wrens), Pardalotidae (pardalotes), and
Acanthizidae (thornbills, Australian warblers, scrubwrens, etc.) they comprise the superfamily Meliphagoidea
and originated early in the evolutionary history of the oscine passerine radiation.[4]
Although honeyeaters look and behave very much like other nectar-feeding passerines around the
world (such as the sunbirds and flowerpeckers), they are unrelated, and the similarities are the consequence
of convergent evolution.
The extent of the evolutionary partnership between honeyeaters and Australasian flowering plants is
unknown, but probably substantial. A great many Australian plants are fertilised by honeyeaters, particularly
the Proteaceae, Myrtaceae, and Epacridaceae. It is known that the honeyeaters are important in New
Zealand as well, and assumed that the same applies in other areas.
Contents
[hide]
• 1 Characteristics
• 2 Changes in classification
• 3 See also
• 4 References
[edit] Characteristics
A female Eastern Spinebill feeding. Honeyeaters typically hang from branches while feeding on
nectar.
Unlike the hummingbirds of America, honeyeaters do not have extensive adaptations for hovering
flight, though smaller members of the family do hover hummingbird-style to collect nectar from time to time.
In general, honeyeaters prefer to flit quickly from perch to perch in the outer foliage, stretching up or
sideways or hanging upside down at need. Many genera have a highly developed brush-tipped tongue,
longer in some species than others, frayed and fringed with bristles which soak up liquids readily. The tongue
is flicked rapidly and repeatedly into a flower, the upper mandible then compressing any liquid out when the
bill is closed. These birds are one of only a few birds in the world that can fly backwards. This is because of
their special wings.
In addition to nectar, all or nearly all honeyeaters take insects and other small creatures, usually by
hawking, sometimes by gleaning. A few of the larger species, notably the White-eared Honeyeater, and the
Strong-billed Honeyeater of Tasmania, probe under bark for insects and other morsels. Many species
supplement their diets with a little fruit, and a small number eat considerable amounts of fruit,[5] particularly
in tropical rainforests and, oddly, in semi-arid scrubland. The Painted Honeyeater is a mistletoe specialist.
Most, however, exist on a diet of nectar supplemented by varying quantities of insects. In general, the
honeyeaters with long, fine bills are more nectarivorous, the shorter-billed species less so, but even
specialised nectar eaters like the spinebills take extra insects to add protein to their diet when they are
breeding.
The movements of honeyeaters are poorly understood. Most are at least partially mobile but many
movements seem to be local, possibly between favourite haunts as the conditions change. Fluctuations in
local abundance are common, but the small number of definitely migratory honeyeater species aside, the
reasons are yet to be discovered. Many follow the flowering of favourite food plants. Arid zone species
appear to travel further and less predictably than those of the more fertile areas. It seems probable that no
single explanation will emerge: the general rule for honeyeater movements is that there is no general rule.
[edit] Changes in classification

The genera Cleptornis and Apalopteron (Bonin Honeyeater), formerly treated in the Meliphagidae,
have recently been transferred to the Zosteropidae on genetic evidence. The genus Notiomystis (New
Zealand Stitchbird), formerly classified in the Meliphagidae, has recently been removed to the newly-erected
Notiomystidae of which it is the only member.[6] The "Macgregor's bird-of-paradise," historically considered a
bird of paradise (Paradisaeidae), was recently found to be a honeyeater.[7] It is now known as "Macgregor's
Giant Honeyeater" and is classified in the Meliphagidae.
The Wattled Smoky Honeyeater (Melipotes carolae), described in 2007, has been discovered in
December 2005 in the Foja Mountains of Papua, Indonesia.[1]
In 2008, a study that included molecular phylogenetic analysis of museum specimens in the genera
Moho and Chaetoptila, both extinct genera endemic to the Hawaiian islands, argued that these five species
were not members of the Meliphagidae and instead belong to their own distinct family, the Mohoidae.[8]
[edit] See also

• List of honeyeaters
[edit] References
1. ^ a b Recent molecular phylogenetic work by Driskell & Christidis 2004, indicates this genus
is non-monophyletic and will undergo taxonomic revision in the near future.
2. ^ Recent molecular phylogenetic work by Driskell & Christidis 2004, indicates Xanthomyza
phrygia is contained within the genus Anthochaera and this species will be listed as Anthochaera
phrygia in the next species list of Australian birds (L. Christids, pers. comm.)
3. ^ as listed in Sibley & Monroe 1990
4. ^ Barker et al. 2004
5. ^ Lindsey, Terence (1991). Forshaw, Joseph. ed. Encyclopaedia of Animals: Birds. London:
Merehurst Press. pp. 208. ISBN 1-85391-186-0.
6. ^ Driskell et al. 2007
7. ^ Cracraft & Feinstein 2000
8. ^ Fleischer RC, James HF, and Olson SL. (2008). Convergent evolution of Hawaiian and
Australo-Pacific Honeyeaters from distant songbird ancestors. Current Biology, 18(24): 1927-1931.
• Barker, F.K., Cibois, A., Schikler, P., Feinstein, J., and Cracraft, J. (2004). Phylogeny and
diversification of the largest avian radiation. Proceedings Natl. Acad. Sci., USA 101 11040-11045.
• Christidis, L. and Boles, W.E. (1994). The Taxonomy and Species of Birds of Australia and
its Territories. Royal Australasian Ornithologists Union Monograph 2. Melbourne: RAOU. ISBN
1875122060.
• Cracraft, J. and Feinstein, J. (2000). What is not a bird of paradise? Molecular and
morphological evidence places Macgregoria in the Meliphagidae and the Cnemophilinae near the
base of the corvoid tree. Proc. Roy. Soc. London, B 267 233-241.
• Del Hoyo, J., Elliot, A. and Christie D. (editors). (2006). Handbook of the Birds of the World.
Volume 12: Picathartes to Tits and Chickadees. Lynx Edicions. ISBN 9788496553422 (Epthianura
and Ashbyia only)
• Driskell, A.C. and Christidis, L. (2004). Phylogeny and evolution of the Australo-Papuan
honeyeaters (Passeriformes, Meliphagidae). Molecular Phylogenetics and Evolution 31 943–960.
• Driskell, A.C., Christidis, L., Gill, B., Boles, W.E., Barker, F.K., and Longmore, N.W. (2007). A
new endemic family of New Zealand passerine birds: adding heat to a biodiversity hotspot.
Australian Journal of Zoology 55 1-6.
• Sibley, C.G. and Monroe, B.L. Jr. (1990). Distribution and Taxonomy of Birds of the World.
New Haven: Yale University Press. ISBN 0300049692.
• Honeyeater videos on the Internet Bird Collection
• Meliphagoidea - Highlighting relationships of Maluridae on Tree Of Life Web Project
Retrieved from "http://en.wikipedia.org/wiki/Honeyeater"
Categories: Meliphagidae
W000
Flowering plant

Flowering plants
Temporal range: Early Cretaceous — Recent
PreЄ
g
Magnolia virginiana
Sweet Bay
Kingdom: Plantae
Angiospermae
Division:
Lindley[1] [P.D. Cantino & M.J. Donoghue][2]
Clades
Amborellaceae
Nymphaeales
Austrobaileyales
Mesangiospermae
• Ceratophyllaceae
• Chloranthaceae
• Eudicotyledoneae (eudicots)
• Magnoliidae
• Monocotyledoneae (monocots)
Synonyms
Anthophyta
Magnoliophyta Cronquist, Takht. & W.Zimm., 1966
This article's introduction section may not adequately summarize its contents. To
comply with Wikipedia's lead section guidelines, please consider expanding the lead to
provide an accessible overview of the article's key points. (September 2009)
The flowering plants (angiosperms), also known as Angiospermae or Magnoliophyta, are the most
diverse group of land plants. Angiosperms are seed-producing plants like the gymnosperms and can be
distinguished from the gymnosperms by a series of synapomorphies (derived characteristics). These
characteristics include flowers, endosperm within the seeds, and the production of fruits that contain the
seeds.
The ancestors of flowering plants diverged from gymnosperms around 245–202 million years ago,
and the first flowering plants known to exist are from 140 million years ago. They diversified enormously
during the Lower Cretaceous and became widespread around 100 million years ago, but replaced conifers as
the dominant trees only around 60-100 million years ago.
Contents
[hide]
• 1 Angiosperm derived characteristics
• 2 Evolution
• 3.1 History of classification
• 3.2 Flowering plant diversity
• 4 Vascular anatomy
• 5 The flower, fruit, and seed
• 5.1 Flowers
• 5.2 Fertilization and embryogenesis
• 5.3 Fruit and seed
• 6 Economic importance
• 7 See also
• 8 References
[edit] Angiosperm derived characteristics

• Flowers
The flowers, which are the reproductive organs of flowering plants, are the most remarkable feature
distinguishing them from other seed plants. Flowers aid angiosperms by enabling a wider range of
adaptability and broadening the ecological niches open to them. This has allowed flowering plants to largely
dominate terrestrial ecosystems.
• Stamens with two pairs of pollen sacs
Stamens are much lighter than the corresponding organs of gymnosperms and have contributed to
the diversification of angiosperms through time with adaptations to specialized pollination syndromes, such
as particular pollinators. Stamens have also become modified through time to prevent self-fertilization, which
has permitted further diversification, allowing angiosperms eventually to fill more niches.
• Reduced male parts, three cells
The male gametophyte in angiosperms is significantly reduced in size compared to those of
gymnosperm seed plants. The smaller pollen decreases the time from pollination — the pollen grain reaching
the female plant — to fertilization of the ovary; in gymnosperms fertilization can occur up to a year after
pollination, while in angiosperms the fertilization begins very soon after pollination. The shorter time leads to
angiosperm plants setting seeds sooner and faster than gymnosperms, which is a distinct evolutionary
advantage.
• Closed carpel enclosing the ovules (carpel or carpels and accessory parts may become the
fruit)
The closed carpel of angiosperms also allows adaptations to specialized pollination syndromes and
controls. This helps to prevent self-fertilization, thereby maintaining increased diversity. Once the ovary is
fertilized, the carpel and some surrounding tissues develop into a fruit. This fruit often serves as an attractant
to seed-dispersing animals. The resulting cooperative relationship presents another advantage to
angiosperms in the process of dispersal.
• Reduced female gametophyte, seven cells with eight nuclei
The reduced female gametophyte, like the reduced male gametophyte, may be an adaptation
allowing for more rapid seed set, eventually leading to such flowering plant adaptations as annual
herbaceous life cycles, allowing the flowering plants to fill even more niches.
• Endosperm
Endosperm formation generally begins after fertilization and before the first division of the zygote.
Endosperm is a highly nutritive tissue that can provide food for the developing embryo, the cotyledons, and
sometimes for the seedling when it first appears.
These distinguishing characteristics taken together have made the angiosperms the most diverse
and numerous land plants and the most commercially important group to humans. The major exception to the
dominance of terrestrial ecosystems by flowering plants is the coniferous forest.
[edit] Evolution
Further information: Evolutionary history of plants#Flowers
Flowers of Malus sylvestris (crab apple)
Land plants have existed for about 425 million years.[3] Early land plants reproduced sexually with
flagellated, swimming sperm, like the green algae from which they evolved. An adaptation to terrestrialization
was the development of upright meiosporangia for dispersal by spores to new habitats. This feature is lacking
in the descendants of their nearest algal relatives, the Charophycean green algae. A later terrestrial
adaptation took place with retention of the delicate, avascular sexual stage, the gametophyte, within the
tissues of the vascular sporophyte. This occurred by spore germination within sporangia rather than spore
release, as in non-seed plants. A current example of how this might have happened can be seen in the
precocious spore germination in Sellaginella, the spike-moss. The result for the ancestors of angiosperms
was enclosing them in a case, the seed. The first seed bearing plants, like the ginkgo, and conifers (such as
pines and firs), did not produce flowers. Interestingly, the pollen grains (males) of Ginkgo and cycads
produce a pair of flagellated, mobile sperm cells that "swim" down the developing pollen tube to the female
and her eggs.
The apparently sudden appearance of relatively modern flowers in the fossil record initially posed
such a problem for the theory of evolution that it was called an "abominable mystery" by Charles Darwin.[4]
However, the fossil record has considerably grown since the time of Darwin, and recently discovered
angiosperm fossils such as Archaefructus, along with further discoveries of fossil gymnosperms, suggest
how angiosperm characteristics may have been acquired in a series of steps. Several groups of extinct
gymnosperms, particularly seed ferns, have been proposed as the ancestors of flowering plants but there is
no continuous fossil evidence showing exactly how flowers evolved. Some older fossils, such as the upper
Triassic Sanmiguelia, have been suggested. Based on current evidence, some propose that the ancestors of
the angiosperms diverged from an unknown group of gymnosperms during the late Triassic (245–202 million
years ago). A close relationship between angiosperms and gnetophytes, proposed on the basis of
morphological evidence, has more recently been disputed on the basis of molecular evidence that suggest
gnetophytes are instead more closely related to other gymnosperms.
The earliest known macrofossil confidently identified as an angiosperm, Archaefructus liaoningensis,
is dated to about 125 million years BP (the Cretaceous period),[5] while pollen considered to be of
angiosperm origin takes the fossil record back to about 130 million years BP. However, one study has
suggested that the early-middle Jurassic plant Schmeissneria, traditionally considered a type of ginkgo, may
be the earliest known angiosperm, or at least a close relative.[6] Additionally, circumstantial chemical
evidence has been found for the existence of angiosperms as early as 250 million years ago. Oleanane, a
secondary metabolite produced by many flowering plants, has been found in Permian deposits of that age
together with fossils of gigantopterids.[7][8] Gigantopterids are a group of extinct seed plants that share many
morphological traits with flowering plants, although they are not known to have been flowering plants
themselves.
Recent DNA analysis based on molecular systematics [9][10] showed that Amborella trichopoda,
found on the Pacific island of New Caledonia, belongs to a sister group of the other flowering plants, and
morphological studies [11] suggest that it has features that may have been characteristic of the earliest
flowering plants.
The great angiosperm radiation, when a great diversity of angiosperms appears in the fossil record,
occurred in the mid-Cretaceous (approximately 100 million years ago). However, a study in 2007 estimated
that the division of the five most recent (the genus Ceratophyllum, the family Chloranthaceae, the eudicots,
the magnoliids, and the monocots) of the eight main groups occurred around 140 million years ago.[12] By
the late Cretaceous, angiosperms appear to have dominated environments formerly occupied by ferns and
cycadophytes, but large canopy-forming trees replaced conifers as the dominant trees only close to the end
of the Cretaceous 65 millions years ago or even later, at the beginning of the Tertiary.[13] The radiation of
herbaceous angiosperm occurred much later.[14] Yet, many fossil plants recognizable as belonging to
modern families (including beech, oak, maple, and magnolia) appeared already at late Cretaceous.
Two bees on a flower head of Creeping Thistle, Cirsium arvense

It is generally assumed that the function of flowers, from the start, was to involve mobile animals in
their reproduction processes. That is, pollen can be scattered even if the flower is not brightly colored or
oddly shaped in a way that attracts animals; however, by expending the energy required to create such traits,
angiosperms can enlist the aid of animals and thus reproduce more efficiently.
Island genetics provides one proposed explanation for the sudden, fully developed appearance of
flowering plants. Island genetics is believed to be a common source of speciation in general, especially when
it comes to radical adaptations that seem to have required inferior transitional forms. Flowering plants may
have evolved in an isolated setting like an island or island chain, where the plants bearing them were able to
develop a highly specialized relationship with some specific animal (a wasp, for example). Such a
relationship, with a hypothetical wasp carrying pollen from one plant to another much the way fig wasps do
today, could result in both the plant(s) and their partners developing a high degree of specialization. Note that
the wasp example is not incidental; bees, which apparently evolved specifically due to mutualistic plant
relationships, are descended from wasps.
Animals are also involved in the distribution of seeds. Fruit, which is formed by the enlargement of
flower parts, is frequently a seed-dispersal tool that attracts animals to eat or otherwise disturb it, incidentally
scattering the seeds it contains (see frugivory). While many such mutualistic relationships remain too fragile
to survive competition and to spread widely, flowering proved to be an unusually effective means of
reproduction, spreading (whatever its origin) to become the dominant form of land plant life.
Flower ontogeny uses a combination of genes normally responsible for forming new shoots.[15] The
most primitive flowers are thought to have had a variable number of flower parts, often separate from (but in
contact with) each other. The flowers would have tended to grow in a spiral pattern, to be bisexual (in plants,
this means both male and female parts on the same flower), and to be dominated by the ovary (female part).
As flowers grew more advanced, some variations developed parts fused together, with a much more specific
number and design, and with either specific sexes per flower or plant, or at least "ovary inferior".
Flower evolution continues to the present day; modern flowers have been so profoundly influenced
by humans that some of them cannot be pollinated in nature. Many modern, domesticated flowers used to be
simple weeds, which only sprouted when the ground was disturbed. Some of them tended to grow with
human crops, perhaps already having symbiotic companion plant relationships with them, and the prettiest
did not get plucked because of their beauty, developing a dependence upon and special adaptation to human
affection.[16]
Angiospermae
Amborella
Nymphaeales
Austrobaileyales
Mesangiospermae
magnoliids
Chloranthales
monocots
Ceratophyllu
m
eudicots
There are eight groups of living angiosperms:
• Amborella — a single species of shrub from New Caledonia
• Nymphaeales — about 80 species[19] — water lilies and Hydatellaceae
• Austrobaileyales — about 100 species[19] of woody plants from various parts of the world
• Chloranthales — several dozen species of aromatic plants with toothed leaves
• Magnoliidae — about 9,000 species,[19] characterized by trimerous flowers, pollen with one
pore, and usually branching-veined leaves — for example magnolias, bay laurel, and black pepper
• Monocotyledonae — about 70,000 species,[19] characterized by trimerous flowers, a single
cotyledon, pollen with one pore, and usually parallel-veined leaves — for example grasses, orchids,
and palms
• Ceratophyllum — about 6 species[19] of aquatic plants, perhaps most familiar as aquarium
plants
• Eudicotyledonae — about 175,000 species,[19] characterized by 4- or 5- merous flowers,
pollen with three pores, and usually branching-veined leaves — for example sunflowers, petunia,
buttercup, apples and oaks
The exact relationship between these eight groups is not yet clear, although there is agreement that
the first three groups to diverge from the ancestral angiosperm were Amborellales, Nymphaeales, and
Austrobaileyales.[20] The term basal angiosperms refers to these three groups. The five other groups form
the clade Mesangiospermae. The relationship between the three largest of these groups (magnoliids,
monocots and eudicots) remains unclear. Some analyses make the magnoliids the first to diverge, others the
monocots.[18]. Ceratophyllum seems to group with the eudicots rather than with the monocots.
[edit] History of classification
From 1736, an illustration of Linnaean classification

The botanical term "Angiosperm", from the Ancient Greek αγγείον, angeíon (receptacle, vessel) and
σπέρμα, (seed), was coined in the form Angiospermae by Paul Hermann in 1690, as the name of that one of
his primary divisions of the plant kingdom. This included flowering plants possessing seeds enclosed in
capsules, distinguished from his Gymnospermae, or flowering plants with achenial or schizo-carpic fruits, the
whole fruit or each of its pieces being here regarded as a seed and naked. The term and its antonym were
maintained by Carolus Linnaeus with the same sense, but with restricted application, in the names of the
orders of his class Didynamia. Its use with any approach to its modern scope only became possible after
1827, when Robert Brown established the existence of truly naked ovules in the Cycadeae and Coniferae,
and applied to them the name Gymnosperms. From that time onwards, so long as these Gymnosperms
were, as was usual, reckoned as dicotyledonous flowering plants, the term Angiosperm was used
antithetically by botanical writers, with varying scope, as a group-name for other dicotyledonous plants.
Auxanometer: Device for measuring increase or rate of growth in plants
In 1851, Hofmeister discovered the changes occurring in the embryo-sac of flowering plants, and
determined the correct relationships of these to the Cryptogamia. This fixed the position of Gymnosperms as
a class distinct from Dicotyledons, and the term Angiosperm then gradually came to be accepted as the
suitable designation for the whole of the flowering plants other than Gymnosperms, including the classes of
Dicotyledons and Monocotyledons. This is the sense in which the term is used today.
In most taxonomies, the flowering plants are treated as a coherent group. The most popular
descriptive name has been Angiospermae (Angiosperms), with Anthophyta ("flowering plants") a second
choice. These names are not linked to any rank. The Wettstein system and the Engler system use the name
Angiospermae, at the assigned rank of subdivision. The Reveal system treated flowering plants as
subdivision Magnoliophytina (Frohne & U. Jensen ex Reveal, Phytologia 79: 70 1996), but later split it to
Magnoliopsida, Liliopsida and Rosopsida. The Takhtajan system and Cronquist system treat this group at the
rank of division, leading to the name Magnoliophyta (from the family name Magnoliaceae). The Dahlgren
system and Thorne system (1992) treat this group at the rank of class, leading to the name Magnoliopsida.
The APG system of 1998, and the later 2003[21] and 2009[17] revisions, treat the flowering plants as a clade
called angiosperms without a formal botanical name. However, a formal classification was published
alongside the 2009 revision in which the flowering plants form the Subclass Magnoliidae.[22]
The internal classification of this group has undergone considerable revision. The Cronquist system,
proposed by Arthur Cronquist in 1968 and published in its full form in 1981, is still widely used but is no
longer believed to accurately reflect phylogeny. A consensus about how the flowering plants should be
arranged has recently begun to emerge through the work of the Angiosperm Phylogeny Group (APG), which
published an influential reclassification of the angiosperms in 1998. Updates incorporating more recent
research were published as APG II in 2003[21]and as APG III in 2009.[17][23]
Monocot (left) and dicot seedlings
Traditionally, the flowering plants are divided into two groups, which in the Cronquist system are
called Magnoliopsida (at the rank of class, formed from the family name Magnoliacae) and Liliopsida (at the
rank of class, formed from the family name Liliaceae). Other descriptive names allowed by Article 16 of the
ICBN include Dicotyledones or Dicotyledoneae, and Monocotyledones or Monocotyledoneae, which have a
long history of use. In English a member of either group may be called a dicotyledon (plural dicotyledons)
and monocotyledon (plural monocotyledons), or abbreviated, as dicot (plural dicots) and monocot (plural
monocots). These names derive from the observation that the dicots most often have two cotyledons, or
embryonic leaves, within each seed. The monocots usually have only one, but the rule is not absolute either
way. From a diagnostic point of view the number of cotyledons is neither a particularly handy nor reliable
character.
Recent studies, as by the APG, show that the monocots form a monophyletic group (clade) but that
the dicots do not (they are paraphyletic). Nevertheless, the majority of dicot species do form a monophyletic
group, called the eudicots or tricolpates. Of the remaining dicot species, most belong to a third major clade
known as the Magnoliidae, containing about 9,000 species. The rest include a paraphyletic grouping of
primitive species known collectively as the basal angiosperms, plus the families Ceratophyllaceae and
Chloranthaceae.
[edit] Flowering plant diversity

The number of species of flowering plants is estimated to be in the range of 250,000 to 400,000. [24]
[25] [26] The number of families in APG (1998) was 462. In APG II[21] (2003) it is not settled; at maximum it
is 457, but within this number there are 55 optional segregates, so that the minimum number of families in
this system is 402. In APG III (2009) there are 415 families.[17]
The diversity of flowering plants is not evenly distributed. Nearly all species belong to the eudicot
(75%), monocot (23%) and magnoliid (2%) clades. The remaining 5 clades contain a little over 250 species in
total, i.e., less than 0.1% of flowering plant diversity, divided among 9 families.
The most diverse families of flowering plants, in their APG circumscriptions, in order of number of
species, are:
1. Asteraceae or Compositae (daisy family): 23,600 species[27]
2. Orchidaceae (orchid family): 22,075 species[27]
3. Fabaceae or Leguminosae (pea family): 19,400[27]
4. Rubiaceae (madder family): 13,150[28]
5. Poaceae or Gramineae (grass family): 10,035[27]
6. Lamiaceae or Labiatae (mint family): 7,173[27]
7. Euphorbiaceae (spurge family): 5,735[27]
8. Melastomataceae (melastome family): 5,005[27]
9. Myrtaceae (myrtle family): 4,620[27]
10.Apocynaceae (dogbane family): 4,555[27]
In the list above (showing only the 10 largest families), the Orchidaceae and Poaceae are monocot
families; the others are eudicot families.
[edit] Vascular anatomy
Cross-section of a stem of the angiosperm flax:

1. Pith,
2. Protoxylem,
3. Xylem I,
4. Phloem I,
5. Sclerenchyma (bast fibre),
6. Cortex,
7. Epidermis
The amount and complexity of tissue-formation in flowering plants exceeds that of gymnosperms.
The vascular bundles of the stem are arranged such that the xylem and phloem form concentric rings.
In the dicotyledons, the bundles in the very young stem are arranged in an open ring, separating a
central pith from an outer cortex. In each bundle, separating the xylem and phloem, is a layer of meristem or
active formative tissue known as cambium. By the formation of a layer of cambium between the bundles
(interfascicular cambium) a complete ring is formed, and a regular periodical increase in thickness results
from the development of xylem on the inside and phloem on the outside. The soft phloem becomes crushed,
but the hard wood persists and forms the bulk of the stem and branches of the woody perennial. Owing to
differences in the character of the elements produced at the beginning and end of the season, the wood is
marked out in transverse section into concentric rings, one for each season of growth, called annual rings.
Among the monocotyledons, the bundles are more numerous in the young stem and are scattered
through the ground tissue. They contain no cambium and once formed the stem increases in diameter only in
exceptional cases.
[edit] The flower, fruit, and seed

[edit] Flowers
Main articles: Flower and Plant sexuality
A collection of flowers forming an inflorescence
The characteristic feature of angiosperms is the flower. Flowers show remarkable variation in form
and elaboration, and provide the most trustworthy external characteristics for establishing relationships
among angiosperm species. The function of the flower is to ensure fertilization of the ovule and development
of fruit containing seeds. The floral apparatus may arise terminally on a shoot or from the axil of a leaf (where
the petiole attaches to the stem). Occasionally, as in violets, a flower arises singly in the axil of an ordinary
foliage-leaf. More typically, the flower-bearing portion of the plant is sharply distinguished from the foliage-
bearing or vegetative portion, and forms a more or less elaborate branch-system called an inflorescence.
The reproductive cells produced by flowers are of two kinds. Microspores, which will divide to
become pollen grains, are the "male" cells and are borne in the stamens (or microsporophylls). The "female"
cells called megaspores, which will divide to become the egg-cell (megagametogenesis), are contained in
the ovule and enclosed in the carpel (or megasporophyll).
The flower may consist only of these parts, as in willow, where each flower comprises only a few
stamens or two carpels. Usually other structures are present and serve to protect the sporophylls and to form
an envelope attractive to pollinators. The individual members of these surrounding structures are known as
sepals and petals (or tepals in flowers such as Magnolia where sepals and petals are not distinguishable
from each other). The outer series (calyx of sepals) is usually green and leaf-like, and functions to protect the
rest of the flower, especially the bud. The inner series (corolla of petals) is generally white or brightly colored,
and is more delicate in structure. It functions to attract insect or bird pollinators. Attraction is effected by color,
scent, and nectar, which may be secreted in some part of the flower. The characteristics that attract
pollinators account for the popularity of flowers and flowering plants among humans.
While the majority of flowers are perfect or hermaphrodite (having both male and female parts in the
same flower structure), flowering plants have developed numerous morphological and physiological
mechanisms to reduce or prevent self-fertilization. Heteromorphic flowers have short carpels and long
stamens, or vice versa, so animal pollinators cannot easily transfer pollen to the pistil (receptive part of the
carpel). Homomorphic flowers may employ a biochemical (physiological) mechanism called self-
incompatibility to discriminate between self- and non-self pollen grains. In other species, the male and female
parts are morphologically separated, developing on different flowers.
[edit] Fertilization and embryogenesis

Main articles: Fertilization and Plant embryogenesis
Angiosperm life cycle
Double fertilization refers to a process in which two sperm cells fertilize cells in the ovary. This
process begins when a pollen grain adheres to the stigma of the pistil (female reproductive structure),
germinates, and grows a long pollen tube. While this pollen tube is growing, a haploid generative cell travels
down the tube behind the tube nucleus. The generative cell divides by mitosis to produce two haploid ( n)
sperm cells. As the pollen tube grows, it makes its way from the stigma, down the style and into the ovary.
Here the pollen tube reaches the micropyle of the ovule and digests its way into one of the synergids,
releasing its contents (which include the sperm cells). The synergid that the cells were released into
degenerates and one sperm makes its way to fertilize the egg cell, producing a diploid (2 n) zygote. The
second sperm cell fuses with both central cell nuclei, producing a triploid (3 n) cell. As the zygote develops
into an embryo, the triploid cell develops into the endosperm, which serves as the embryo's food supply. The
ovary now will develop into fruit and the ovule will develop into seed.
[edit] Fruit and seed

Main articles: Seed and Fruit
The fruit of the Aesculus or Horse Chestnut tree

As the development of embryo and endosperm proceeds within the embryo-sac, the sac wall
enlarges and combines with the nucellus (which is likewise enlarging) and the integument to form the seed-
coat. The ovary wall develops to form the fruit or pericarp, whose form is closely associated with the manner
of distribution of the seed.
Frequently the influence of fertilization is felt beyond the ovary, and other parts of the flower take part
in the formation of the fruit, e.g. the floral receptacle in the apple, strawberry and others.
The character of the seed-coat bears a definite relation to that of the fruit. They protect the embryo
and aid in dissemination; they may also directly promote germination. Among plants with indehiscent fruits,
the fruit generally provides protection for the embryo and secures dissemination. In this case, the seed-coat
is only slightly developed. If the fruit is dehiscent and the seed is exposed, the seed-coat is generally well
developed, and must discharge the functions otherwise executed by the fruit.
[edit] Economic importance

Agriculture is almost entirely dependent on angiosperms, either directly or indirectly through livestock
feed. Of all the families plants, the Poaceae, or grass family, is by far the most important, providing the bulk
of all feedstocks (rice, corn — maize, wheat, barley, rye, oats, pearl millet, sugar cane, sorghum). The
Fabaceae, or legume family, comes in second place. Also of high importance are the Solanaceae, or
nightshade family (potatoes, tomatoes, and peppers, among others), the Cucurbitaceae, or gourd family (also
including pumpkins and melons), the Brassicaceae, or mustard plant family (including rapeseed and the
innumerable varieties of the cabbage species Brassica oleracea), and the Apiaceae, or parsley family. Many
of our fruits come from the Rutaceae, or rue family, and the Rosaceae, or rose family (including apples,
pears, cherries, apricots, plums, etc.).
In some parts of the world, certain single species assume paramount importance because of their
variety of uses, for example the coconut ( Cocos nucifera) on Pacific atolls, and the olive (Olea europaea) in
the Mediterranean region.
Flowering plants also provide economic resources in the form of wood, paper, fiber (cotton, flax, and
hemp, among others), medicines (digitalis, camphor), decorative and landscaping plants, and many other
uses. The main area in which they are surpassed by other plants is timber production.
[edit] See also

• List of garden plants
[edit] References
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Orme, Brown, and Green. xxxvi.
2. ^ Cantino, Philip D.; James A. Doyle, Sean W. Graham, Walter S. Judd, Richard G.
Olmstead, Douglas E. Soltis, Pamela S. Soltis, & Michael J. Donoghue (2007). "Towards a
phylogenetic nomenclature of Tracheophyta". Taxon 56 (3): E1–E44.
3. ^ Lindley, D (2000). "The role of mid-palaeozoic mesofossils in the detection of early
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Wikimedia Commons has media related to: Magnoliophyta
Wikispecies has information related to: Magnoliophyta

Magnoliophyta
• Angiosperm Phylogeny Poster - Flowering Plant Systamatics Cole, T.C.H. & Hilger, H.H.
• Anatomy of a Flowering Land Plant
• Cronquist, Arthur. (1981) An Integrated System of Classification of Flowering Plants .
Columbia Univ. Press, New York.
• Dilcher, D. 2000. Toward a new synthesis: Major evolutionary trends in the angiosperm fossil
record. PNAS [Proceedings of the National Academy of Sciences of the United States of America]
97: 7030-7036 (available online here)
• Heywood, V. H., Brummitt, R. K., Culham, A. & Seberg, O. (2007). Flowering Plant Families
of the World. Richmond Hill, Ontario, Canada: Firefly Books. ISBN 1-55407-206-9.
• Oldest Known Flowering Plants Identified By Genes , William J. Cromie, Harvard Gazette,
December 16, 1999.
• L. Watson and M.J. Dallwitz (1992 onwards). The families of flowering plants: descriptions,
illustrations, identification, information retrieval.
• Simpson, M.G. Plant Systematics, 2nd Edition. Elsevier/Academic Press. 2010.
• Raven, P.H., R.F. Evert, S.E. Eichhorn. Biology of Plants, 7th Edition. W.H. Freeman. 2004.
Plant physiology



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Rhodophyceae Bangiophyceae · Florideophyceae (Hildenbrandiales, Acrochaetiales, Nemaliales, Batracho
Gelidiales, Gracilariales, Ceramiales)
Glaucocystophyceae Glaucocystis · Cyanophora · Gloeochaete
Viridiplantae/ Chlorophyta/ Prasinophyceae

Plantae GA
sensu stricto UTC clade: Ulvophyceae · Trebouxiophyceae · Chlorophycea
Streptophyta Charophyta/G Charales · Coleochaetales · Desmidia

A Mesostigmatales · Zygnematales
Embryophyta/ Bryophytes Marchantiophy

Plantae (non-vascular) Bryophyta "Moss" · Ho
sensu strictissimo
Lycopodiophy
Tracheophyta
Euphyllophyta
See also: list of plant orders
This article incorporates text from a publication now in the public domain: Chisholm, Hugh, ed
(1911). Encyclopædia Britannica (Eleventh ed.). Cambridge University Press.
Retrieved from "http://en.wikipedia.org/wiki/Flowering_plant"
Categories: Angiosperms | Plant taxonomy | Plants | Pollination | Plant sexuality

W000
Algae

For other uses, see Algae (disambiguation) and Alga (disambiguation).
Algae
Laurencia, a marine genus of Red Algae from Hawaii.
Domain: Eukaryota
Included groups
• Archaeplastida
• Chlorophyta (Green algae)
• Rhodophyta (Red algae)
• Glaucophyta
• Rhizaria, Excavata
• Chlorarachniophytes
• Euglenids
• Chromista, Alveolata
• Heterokonts
• Bacillariophyceae (Diatoms)
• Axodine
• Bolidomonas
• Eustigmatophyceae
• Phaeophyceae (Brown algae)
• Chrysophyceae (Golden
algae)
• Raphidophyceae
• Synurophyceae
• Xanthophyceae (Yellow-green
algae)
• Cryptophyta
• Dinoflagellates
• Haptophyta
Excluded groups
• Cyanobacteria
• Plantae
The lineage of algae according to Thomas Cavalier-Smith. The exact number and placement of
endosymbiotic events is not yet clear, so this diagram can be taken only as a general guide[1][2] It represents
the most parsimonious way of explaining the three types of endosymbiotic origins of plastids. These types
include the endosymbiotic events of cyanobacteria, red algae and green algae, leading to the hypothesis of
the supergroups Archaeplastida, Chromalveolata and Cabozoa respectively. However, the monophyly of
Cabozoa has been refuted and the monophylies of Archaeplastida and Chromalveolata are currently strongly
challenged. Endosymbiotic events are noted by dotted lines.
Algae (pronounced /ˈældʒiː/ or /ˈælɡiː/; singular alga /ˈælɡə/, Latin for "seaweed") are a large and
diverse group of simple, typically autotrophic organisms, ranging from unicellular to multicellular forms, such
as the giant kelps that grow to 65 meters in length. The US Algal Collection is represented by almost 300,000
accessioned and inventoried herbarium specimens.[3] The largest and most complex marine forms are
called seaweeds. They are photosynthetic like plants, and "simple" because their tissues are not organized
into the many distinct organs found in land plants.
Though the prokaryotic cyanobacteria (commonly referred to as blue-green algae) were traditionally
included as "algae" in older textbooks, many modern sources regard this as outdated[4] as they are now
considered to be bacteria.[5] The term algae is now restricted to eukaryotic organisms.[6] All true algae
therefore have a nucleus enclosed within a membrane and plastids bound in one or more membranes.[4][7]
Algae constitute a paraphyletic and polyphyletic group,[4] as they do not include all the descendants of the
last universal ancestor nor do they all descend from a common algal ancestor, although their plastids seem
to have a single origin.[1] Diatoms are also examples of algae.
Algae are found in the fossil record dating back to approximately 3 billion years in the Precambrian.
They exhibit a wide range of reproductive strategies, from simple, asexual cell division to complex forms of
sexual reproduction.[8]
Algae lack the various structures that characterize land plants, such as phyllids (leaves) and rhizoids
in nonvascular plants, or leaves, roots, and other organs that are found in tracheophytes (vascular plants).
Many are photoautotrophic, although some groups contain members that are mixotrophic, deriving energy
both from photosynthesis and uptake of organic carbon either by osmotrophy, myzotrophy, or phagotrophy.
Some unicellular species rely entirely on external energy sources and have limited or no photosynthetic
apparatus.
Nearly all algae have photosynthetic machinery ultimately derived from the Cyanobacteria, and so
produce oxygen as a by-product of photosynthesis, unlike other photosynthetic bacteria such as purple and
green sulfur bacteria. Fossilized filamentous algae from the Vindhya basin have been dated back to 1.6 to
1.7 billion years ago.[9]
Contents
[hide]
• 1 Etymology and study
• 3 Relationship to higher plants
• 4 Morphology
• 5 Symbiotic algae
• 5.1 Lichens
• 5.2 Coral reefs
• 5.3 Sea sponges
• 6 Life-cycle
• 7 Numbers
• 8 Distribution
• 9 Locations
• 10 Uses
• 10.1 Agar
• 10.2 Alginates
• 10.3 Energy source
• 10.4 Fertilizer
• 10.5 Nutrition
• 10.6 Pollution control
• 10.7 Pigments
[edit] Etymology and study
Title page of Samuel Gottlieb Gmelin, Historia Fucorum, dated 1768.

The singular alga is the Latin word for a particular seaweed and retains that meaning in English.[10]
The etymology is obscure. Although some speculate that it is related to Latin algēre, "be cold",[11] there is no
known reason to associate seaweed with temperature. A more likely source is alliga, "binding, entwining."[12]
Since Algae has become a biological classification, alga can also mean one classification under Algae,
parallel to a fungus being a species of fungi, a plant being a species of plant, and so on.
The ancient Greek word for seaweed was φῦκος (fūkos or phykos), which could mean either the
seaweed, probably Red Algae, or a red dye derived from it. The Latinization, fūcus, meant primarily the
cosmetic rouge. The etymology is uncertain, but a strong candidate has long been some word related to the
Biblical ‫( פוך‬pūk), "paint" (if not that word itself), a cosmetic eye-shadow used by the ancient Egyptians and
other inhabitants of the eastern Mediterranean. It could be any color: black, red, green, blue.[13]
Accordingly the modern study of marine and freshwater algae is called either phycology or algology.
The name Fucus appears in a number of taxa.
False-colour Scanning electron micrograph of the unicellular coccolithophore, Gephyrocapsa

oceanica.
While Cyanobacteria have been traditionally included among the Algae, recent works usually exclude
them due to large differences such as the lack of membrane-bound organelles, the presence of a single
circular chromosome, the presence of peptidoglycan in the cell walls, and ribosomes different in size and
content from those of the Eukaryotes.[14][15] Rather than in chloroplasts, they conduct photosynthesis on
specialized infolded cytoplasmic membranes called thylakoid membranes. Therefore, they differ significantly
from the Algae despite occupying similar ecological niches.
By modern definitions Algae are Eukaryotes and conduct photosynthesis within membrane-bound
organelles called chloroplasts. Chloroplasts contain circular DNA and are similar in structure to
Cyanobacteria, presumably representing reduced cyanobacterial endosymbionts. The exact nature of the
chloroplasts is different among the different lines of Algae, reflecting different endosymbiotic events. The
table below describes the composition of the three major groups of Algae. Their lineage relationships are
shown in the figure in the upper right. Many of these groups contain some members that are no longer
photosynthetic. Some retain plastids, but not chloroplasts, while others have lost plastids entirely.
Phylogeny based on plastid.[16] not nucleocytoplasmic genealogy:
Rhodoplasts
Rhodophytes
Heterokonts
Cryptophytes
Haptophytes
Chloroplasts
Euglenophytes
Chlorophytes
Charophytes
Higher plants
(Embryophyta)
Chlorarachniophytes
Supergroup
Members Endosymbiont Summary
affiliation
These Algae have

primary chloroplasts, i.e. the
chloroplasts are surrounded
by two membranes and
probably developed through a
single endosymbiotic event.
The chloroplasts of Red Algae
have chlorophylls a and c
• Chlorophyta
Primoplantae/ (often), and phycobilins, while
• Rhodophyta Cyanobacteria
Archaeplastida those of Green Algae have
• Glaucophyta chloroplasts with chlorophyll a
and b. Higher plants are
pigmented similarly to Green
Algae and probably developed
from them, and thus
Chlorophyta is a sister taxon
to the plants; sometimes they
are grouped as Viridiplantae.
Excavata and • Chlorarachni Green Algae These groups have

Rhizaria ophytes green chloroplasts containing
chlorophylls a and b.[14] Their
chloroplasts are surrounded
by four and three membranes,
respectively, and were
probably retained from
ingested Green Algae.
Chlorarachniophytes,
which belong to the phylum
Cercozoa, contain a small
nucleomorph, which is a relict
of the algae's nucleus.
• Euglenids
Euglenids, which
belong to the phylum
Euglenozoa, live primarily in
freshwater and have
chloroplasts with only three
membranes. It has been
suggested that the
endosymbiotic Green Algae
were acquired through
myzocytosis rather than
phagocytosis.
Chromista and • Heterokonts Red Algae These groups have

Alveolata • Haptophyta chloroplasts containing
• Cryptomona chlorophylls a and c, and
ds phycobilins.The shape varies
• Dinoflagellat from plant to plant. they may
es be of discoid, plate-like,
reticulate, cup-shaped, spiral
or ribbon shaped. They have
one or more pyrenoids to
preserve protein and starch.
The latter chlorophyll type is
not known from any
prokaryotes or primary
chloroplasts, but genetic
similarities with the Red Algae
suggest a relationship
there[citation needed].
In the first three of
these groups (Chromista), the
chloroplast has four
membranes, retaining a
nucleomorph in
Cryptomonads, and they likely
share a common pigmented
ancestor, although other
evidence casts doubt on
whether the Heterokonts,
Haptophyta, and
Cryptomonads are in fact
more closely related to each
other than to other groups.[2]
[17]
The typical
dinoflagellate chloroplast has
three membranes, but there is
considerable diversity in
chloroplasts within the group,
and it appears there were a
number of endosymbiotic
events.[1] The Apicomplexa, a
group of closely related
parasites, also have plastids
called apicoplasts.
Apicoplasts are not
photosynthetic but appear to
have a common origin with
Dinoflagellate chloroplasts.[1]
W.H.Harvey (1811—1866) was the first to divide the Algae into four divisions based on their
pigmentation. This is the first use of a biochemical criterion in plant systematics. Harvey's four divisions are:
Red Algae (Rhodophyta), Brown Algae (Heteromontophyta), Green Algae (Chlorophyta) and Diatomaceae.
[18]
[edit] Relationship to higher plants

The first plants on earth evolved from shallow freshwater algae much like Chara some 400 million
years ago. These probably had an isomorphic alternation of generations and were probably filamentous.
Fossils of isolated land plant spores suggest land plants may have been around as long as 475 million years
ago.[19][20]
[edit] Morphology
The kelp forest exhibit at the Monterey Bay Aquarium. A three-dimensional, multicellular thallus.
A range of algal morphologies are exhibited, and convergence of features in unrelated groups is
common. The only groups to exhibit three dimensional multicellular thalli are the reds and browns, and some
chlorophytes.[21] Apical growth is constrained to subsets of these groups: the florideophyte reds, various
browns, and the charophytes.[21] The form of charophytes is quite different to those of reds and browns,
because have distinct nodes, separated by internode 'stems'; whorls of branches reminiscent of the
horsetails occur at the nodes.[21] Conceptacles are another polyphyletic trait; they appear in the coralline
algae and the Hildenbrandiales, as well as the browns.[21]
Most of the simpler algae are unicellular flagellates or amoeboids, but colonial and non-motile forms
have developed independently among several of the groups. Some of the more common organizational
levels, more than one of which may occur in the life cycle of a species, are
• Colonial: small, regular groups of motile cells
• Capsoid: individual non-motile cells embedded in mucilage
• Coccoid: individual non-motile cells with cell walls
• Palmelloid: non-motile cells embedded in mucilage
• Filamentous: a string of non-motile cells connected together, sometimes branching
• Parenchymatous: cells forming a thallus with partial differentiation of tissues
In three lines even higher levels of organization have been reached, with full tissue differentiation.
These are the brown algae,[22]—some of which may reach 50 m in length (kelps)[23]—the red algae,[24] and
the green algae.[25] The most complex forms are found among the green algae (see Charales and
Charophyta), in a lineage that eventually led to the higher land plants. The point where these non-algal plants
begin and algae stop is usually taken to be the presence of reproductive organs with protective cell layers, a
characteristic not found in the other alga groups.
[edit] Symbiotic algae

Some species of algae form symbiotic relationships with other organisms. In these symbioses, the
algae supply photosynthates (organic substances) to the host organism providing protection to the algal cells.
The host organism derives some or all of its energy requirements from the algae. Examples are as follows.
[edit] Lichens
Main article: Lichens
Rock lichens in Ireland.

Lichens are defined by the International Association for Lichenology to be "an association of a fungus
and a photosynthetic symbiont resulting in a stable vegetative body having a specific structure."[26] The
fungi, or mycobionts, are from the Ascomycota with a few from the Basidiomycota. They are not found alone
in nature but when they began to associate is not known.[27] One mycobiont associates with the same
phycobiont species, rarely two, from the Green Algae, except that alternatively the mycobiont may associate
with the same species of Cyanobacteria (hence "photobiont" is the more accurate term). A photobiont may be
associated with many specific mycobionts or live independently; accordingly, lichens are named and
classified as fungal species.[28] The association is termed a morphogenesis because the lichen has a form
and capabilities not possessed by the symbiont species alone (they can be experimentally isolated). It is
possible that the photobiont triggers otherwise latent genes in the mycobiont.[29]
[edit] Coral reefs

Main articles: Coral, Coral reef, and Zooxanthella
Floridian coral reef

Coral reefs are accumulated from the calcareous exoskeletons of marine invertebrates of the
Scleractinia order; i.e., the Stony Corals. As animals they metabolize sugar and oxygen to obtain energy for
their cell-building processes, including secretion of the exoskeleton, with water and carbon dioxide as
byproducts. As the reef is the result of a favorable equilibrium between construction by the corals and
destruction by marine erosion, the rate at which metabolism can proceed determines the growth or
deterioration of the reef.
Algae of the Dinoflagellate phylum are often endosymbionts in the cells of marine invertebrates,
where they accelerate host-cell metabolism by generating immediately available sugar and oxygen through
photosynthesis using incident light and the carbon dioxide produced in the host. Endosymbiont algae in the
Stony Corals are described by the term zooxanthellae, with the host Stony Corals called on that account
hermatypic corals, which although not a taxon are not in healthy condition without their endosymbionts.
Zooxanthellae belong almost entirely to the genus Symbiodinium.[30] The loss of Symbiodinium from the
host is known as coral bleaching, a condition which unless corrected leads to the deterioration and loss of the
reef.
[edit] Sea sponges

Main article: Sea sponge
Green Algae live close to the surface of some sponges, for example, breadcrumb sponge
(Halichondria panicea). The alga is thus protected from predators; the sponge is provided with oxygen and
sugars which can account for 50 to 80% of sponge growth in some species.[31]
[edit] Life-cycle
Rhodophyta, Chlorophyta and Heterokontophyta, the three main algal Phyla, have life-cycles which
show tremendous variation with considerable complexity. In general there is an asexual phase where the
seaweed's cells are diploid, a sexual phase where the cells are haploid followed by fusion of the male and
female gametes. Asexual reproduction is advantageous in that it permits efficient population increases, but
less variation is possible. Sexual reproduction allows more variation, but is more costly. Often there is no
strict alternation between the sporophyte and also because there is often an asexual phase, which could
include the fragmentation of the thallus.[23][32][33]
For more details on this topic, see Conceptacle.
[edit] Numbers
Algae on coastal rocks at Shihtiping in Taiwan

The Algal Collection of the U.S. National Herbarium (located in the National Museum of Natural
History) consists of approximately 320500 dried specimens, which, although not exhaustive (no exhaustive
collection exists), gives an idea of the order of magnitude of the number of algal species (that number
remains unknown).[34] Estimates vary widely. For example, according to one standard textbook,[35] in the
British Isles the UK Biodiversity Steering Group Report estimated there to be 20000 algal species in the UK.
Another checklist reports only about 5000 species. Regarding the difference of about 15000 species, the text
concludes: "It will require many detailed field surveys before it is possible to provide a reliable estimate of the
total number of species ...."
Regional and group estimates have been made as well: 5000—5500 species of Red Algae worldwide,
"some 1300 in Australian Seas,"[36] 400 seaweed species for the western coastline of South Africa,[37] 669
marine species from California (U.S.A.),[38] 642 in the check-list of Britain and Ireland,[39] and so on, but
lacking any scientific basis or reliable sources, these numbers have no more credibility than the British ones
mentioned above. Most estimates also omit the microscopic Algae, such as the phytoplankta, entirely.
[edit] Distribution
The topic of distribution of algal species has been fairly well studied since the founding of
phytogeography in the mid-19th century AD.[40] Algae spread mainly by the dispersal of spores analogously
to the dispersal of Plantae by seeds and spores. Spores are everywhere in all parts of the Earth: the waters
fresh and marine, the atmosphere, free-floating and in precipitation or mixed with dust, the humus and in
other organisms, such as humans. Whether a spore is to grow into an organism depends on the combination
of the species and the environmental conditions of where the spore lands.
The spores of fresh-water Algae are dispersed mainly by running water and wind, as well as by living
carriers.[41] The bodies of water into which they are transported are chemically selective. Marine spores are
spread by currents. Ocean water is temperature selective, resulting in phytogeographic zones, regions and
provinces.[42]
To some degree the distribution of Algae is subject to floristic discontinuities caused by geographical
features, such as Antarctica, long distances of ocean or general land masses. It is therefore possible to
identify species occurring by locality, such as "Pacific Algae" or "North Sea Algae". When they occur out of
their localities, it is usually possible to hypothesize a transport mechanism, such as the hulls of ships. For
example, Ulva reticulata and Ulva fasciata travelled from the mainland to Hawaii in this manner.
Mapping is possible for select species only: "there are many valid examples of confined distribution
patterns."[43] For example, Clathromorphum is an arctic genus and is not mapped far south of there.[44] On
the other hand, scientists regard the overall data as insufficient due to the "difficulties of undertaking such
studies."[45]
[edit] Locations
Phytoplankton, Lake Chuzenji

Algae are prominent in bodies of water, common in terrestrial environments and are found in unusual
environments, such as on snow and on ice. Seaweeds grow mostly in shallow marine waters, under
100 metres (330 ft); however some have been recorded to a depth of 360 metres (1,180 ft)[46]
The various sorts of algae play significant roles in aquatic ecology. Microscopic forms that live
suspended in the water column (phytoplankton) provide the food base for most marine food chains. In very
high densities (algal blooms) these algae may discolor the water and outcompete, poison, or asphyxiate
other life forms.
Algae are variously sensitive to different factors, which has made them useful as biological indicators
in the Ballantine Scale and its modification.
[edit] Uses
Harvesting Algae
[edit] Agar
Agar, a gelatinous substance derived from red algae, has a number of commercial uses.[47]
[edit] Alginates
Between 100,000 and 170,000 wet tons of Macrocystis are harvested annually in California for
alginate extraction and abalone feed.[48][49]
[edit] Energy source
Main articles: Algae fuel, Biological hydrogen production, Biohydrogen, Biodiesel, Ethanol fuel,
Butanol fuel, and Vegetable fats and oils
To be competitive and independent from fluctuating support from (local) policy on the long run,
biofuels should equal or beat the cost level of fossil fuels. Here, algae based fuels hold great promise,
directly related to the potential to produce more biomass per unit area in a year than any other form of
biomass. The break-even point for algae-based biofuels should be within reach in about ten to fifteen years.
[50]
[edit] Fertilizer
Seaweed is used as a fertilizer.
For more details on this topic, see Seaweed fertiliser.
For centuries seaweed has been used as a fertilizer; George Owen of Henllys writing in the 16th
century referring to drift weed in South Wales:[51]
This kind of ore they often gather and lay on great heapes, where it heteth and rotteth, and will
have a strong and loathsome smell; when being so rotten they cast on the land, as they do their
muck, and thereof springeth good corn, especially barley ... After spring-tydes or great rigs of
the sea, they fetch it in sacks on horse backes, and carie the same three, four, or five miles, and
cast it on the lande, which doth very much better the ground for corn and grass.
Today Algae are used by humans in many ways; for example, as fertilizers, soil conditioners and
livestock feed.[52] Aquatic and microscopic species are cultured in clear tanks or ponds and are either
harvested or used to treat effluents pumped through the ponds. Algaculture on a large scale is an important
type of aquaculture in some places. Maerl is commonly used as a soil conditioner.
[edit] Nutrition
Seaweed gardens on Inisheer.

See also: Edible seaweed
Naturally growing seaweeds are an important source of food, especially in Asia. They provide many
vitamins including: A, B1, B2, B6, niacin and C, and are rich in iodine, potassium, iron, magnesium and
calcium.[53] In addition commercially cultivated microalgae, including both Algae and Cyanobacteria, are
marketed as nutritional supplements, such as Spirulina,[54] Chlorella and the Vitamin-C supplement,
Dunaliella, high in beta-carotene.
Algae are national foods of many nations: China consumes more than 70 species, including fat choy,
a cyanobacterium considered a vegetable; Japan, over 20 species;[55] Ireland, dulse; Chile, cochayuyo.[56]
Laver is used to make "laver bread" in Wales where it is known as bara lawr; in Korea, gim; in Japan, nori
and aonori. It is also used along the west coast of North America from California to British Columbia, in
Hawaii and by the Māori of New Zealand. Sea lettuce and badderlocks are a salad ingredient in Scotland,
Ireland, Greenland and Iceland.
Dulse, a food.
The oils from some Algae have high levels of unsaturated fatty acids. For example, Parietochloris
incisa is very high in arachidonic acid, where it reaches up to 47% of the triglyceride pool.[57] Some varieties
of Algae favored by vegetarianism and veganism contain the long-chain, essential omega-3 fatty acids,
Docosahexaenoic acid (DHA) and Eicosapentaenoic acid (EPA), in addition to vitamin B12.[citation needed]
The vitamin B12 in algae is not biologically active. Fish oil contains the omega-3 fatty acids, but the original
source is algae (microalgae in particular), which are eaten by marine life such as copepods and are passed
up the food chain.[58] Algae has emerged in recent years as a popular source of omega-3 fatty acids for
vegetarians who cannot get long-chain EPA and DHA from other vegetarian sources such as flaxseed oil,
which only contains the short-chain Alpha-Linolenic acid (ALA).
[edit] Pollution control

• Sewage can be treated with algae, reducing the need for greater amounts of toxic chemicals
than are already used.
• Algae can be used to capture fertilizers in runoff from farms. When subsequently harvested,
the enriched algae itself can be used as fertilizer.
Agricultural Research Service scientists found that 60-90% of nitrogen runoff and 70-100% of
phosphorus runoff can be captured from manure effluents using an algal turf scrubber (ATS). Scientists
developed the ATS, which are shallow, 100-foot raceways of nylon netting where algae colonies can form,
and studied its efficacy for three years. They found that algae can readily be used to reduce the nutrient
runoff from agricultural fields and increase the quality of water flowing into rivers, streams, and oceans. The
enriched algae itself also can be used as a fertilizer. Researchers collected and dried the nutrient-rich algae
from the ATS and studied its potential as an organic fertilizer. They found that cucumber and corn seedlings
grew just as well using ATS organic fertilizer as they did with commercial fertilizers.[59]
[edit] Pigments
The natural pigments produced by algae can be used as an alternative to chemical dyes and coloring
agents.[60]
[edit] Stabilizing substances
Carrageenan, from the red alga Chondrus crispus, is used as a stabiliser in milk products.
Main articles: Carrageenan and Chondrus crispus
[edit] See also

• Algaculture
• AlgaeBase
• AlgaePARC
• Anatoxin
• Eutrophication
• Iron fertilization
• Microalgae
• Microbiofuels
• Microphyte
• Nutrition
• Photobioreactor
• Phytoplankton
• Plant
[edit] Notes
1. ^ a b c d Patrick J. Keeling (2004). "Diversity and evolutionary history of plastids and their
hosts". American Journal of Botany 91: 1481–1493. doi:10.3732/ajb.91.10.1481.
2. ^ a b Laura Wegener Parfrey, Erika Barbero, Elyse Lasser, Micah Dunthorn, Debashish
Bhattacharya, David J Patterson, and Laura A Katz (December 2006). "Evaluating Support for the
Current Classification of Eukaryotic Diversity". PLoS Genet. 2 (12): e220.
doi:10.1371/journal.pgen.0020220. PMID 17194223. PMC 1713255.
http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1713255.
3. ^ Smithsonian National Museum of Natural History; Department of Botany.
http://botany.si.edu/projects/algae/
4. ^ a b c Nabors, Murray W. (2004). Introduction to Botany. San Francisco, CA: Pearson
Education, Inc. ISBN 0-8053-4416-0.
5. ^ Ed. Guiry, M.D., John, D.M., Rindi, F and McCarthy, T.K. 2007. New Survey of Clare Island
Volume 6: The Freshwater and Terrestrial Algae. Royal Irish Academy. isbn 13: 978-1-904890-31-7
6. ^ Allaby, M ed. (1992). "Algae". The Concise Dictionary of Botany. Oxford: Oxford University
Press.
7. ^ Round (1981)
8. ^ Smithsonian National Musem of Natural History; Department of Botany.
http://botany.si.edu/projects/algae/introduction.htm
9. ^ Bengtson, S.; Belivanova, V.; Rasmussen, B.; Whitehouse, M. (May 2009). "The
controversial "Cambrian" fossils of the Vindhyan are real but more than a billion years older".
7734. doi:10.1073/pnas.0812460106. ISSN 0027-8424. PMID 19416859. edit
10.^ "alga, algae". Webster's Third New International Dictionary of the English Language
Unabridged with Seven Language Dictionary. 1. Encyclopedia Britannica, Inc. 1986.
11.^ Partridge, Eric (1983). "algae". Origins.
12.^ Lewis, Charlton T.; Charles Short (1879). "alga". alga. Oxford: Clarendon Press. ISBN 0-
19-864201-6. http://www.perseus.tufts.edu/cgi-bin/ptext?layout.reflang=la;layout.refdoc=Perseus
%3Atext%3A1999.04.0059;layout.reflookup=Alga;layout.refcit=;doc=Perseus%3Atext
%3A1999.04.0059%3Aentry%3D%231812.
13.^ Cheyne, Thomas Kelly; John Sutherland Black (1899–1903). "Paint". Encyclopædia Biblica–
A Dictionary of the Bible. 3. New York: Macmillan Co.. pp. 3524–3525. Downloadable Google Books.
14.^ a b Losos, Jonathan B.; Mason, Kenneth A.; Singer, Susan R. (2007). Biology (8 ed.).
McGraw-Hill. ISBN 0-07-304110-6.
15.^ Jochem, Frank J. "Botany 4404 Lecture Notes". Florida International University (FIU).
http://www.jochemnet.de/fiu/bot4404/BOT4404_12.html. Retrieved 2008-12-20.
16.^ Bhattacharya, D.; Medlin, L. (1998). "Algal Phylogeny and the Origin of Land Plants". Plant
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38.^ Abbott and Hollenberg (1976), page 2.
39.^ Hardy and Guiry (2006).
40.^ Round (1981), Chapter 8, Dispersal, continuity and phytogeography.
41.^ Round (1981), page 360.
42.^ Round (1981), page 362.
43.^ Round (1981), Page 357.
44.^ Round (1981), page 371.
45.^ Round (1981), page 366.
46.^ Round (1981), page 176.
47.^ Lewis, J G; Stanley, N F; Guist, G G (1988). "9 Commercial production of algal
hydrocolloides". In Lembi, C.A.; Waaland, J.R.. Algae and Human Affairs. Cambridge University
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48.^ "Macrocystis C. Agardh 1820: 46". AlgaeBase.
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50.^ Barbosa & Wijffels
51.^ Read, Clare Sewell (1849). "On the Farming of South Wales: Prize Report". Journal of the
Royal Agricultural Society of England (John Murray) 10: 142–143. Downloadable Google Books.
52.^ McHugh, Dennis J. (2003). "9, Other Uses of Seaweeds". A Guide to the Seaweed
Industry: FAO Fisheries Technical Paper 441. Rome: Fisheries and Aquaculture Department, Food
and Agriculture Organization (FAO) of the United Nations. ISBN 9251049580, ISBN 0499345.
http://www.fao.org/DOCREP/006/Y4765E/y4765e0c.htm#TopOfPage.
53.^ Simoons, Frederick J (1991). "6, Seaweeds and Other Algae". Food in China: A Cultural
and Historical Inquiry. CRC Press. pp. 179–190. ISBN 0-936923-29-6.
54.^ Morton, Steve L. "Modern Uses of Cultivated Algae". Ethnobotanical Leaflets. Southern
Illinois University Carbondale. http://www.siu.edu/~ebl/leaflets/algae.htm. Retrieved 2008-12-26.
55.^ Mondragon, J; Mondragon, J (2003). Seaweeds of the Pacific Coast. Monterey, California:
Sea Challengers Publications. ISBN 0-930118-29-4.
56.^ "Durvillaea antarctica (Chamisso) Hariot". AlgaeBase.
http://www.algaebase.org/speciesdetail.lasso?species_id=11752&sk=0&from=results&-
session=abv3:51909EC30802716127sVj3EDC9C7.
57.^ Bigogno, C; I Khozin-Goldberg; S Boussiba; A Vonshak; Z Cohen (2002). "Lipid and fatty
acid composition of the green oleaginous alga Parietochloris incisa, the richest plant source of
arachidonic acid.". Phytochemistry 60 (5): 497–503. doi:10.1016/S0031-9422(02)00100-0.
PMID 12052516.
58.^ Allison Aubrey (Morning Edition, November 1, 2007). "Getting Brain Food Straight from the
Source". National Public Radio. http://www.npr.org/templates/story/story.php?storyId=15823852.
59.^ "Algae: A Mean, Green Cleaning Machine". USDA Agricultural Research Service. May 7,
2010. http://www.ars.usda.gov/is/AR/archive/may10/algae0510.htm.
60.^ Arad, Shoshana; Spharim, Ishai (1998). "Production of Valuable Products from Microalgae:
An Emerging Agroindustry". In Altman, Arie. Agricultural Biotechnology. Books in Soils, Plants, and
the Environment. 61. CRC Press. p. 638. ISBN 978-0-8247-9439-2.
[edit] Bibliography
[edit] General
• Chapman, V.J. (1950). Seaweeds and their Uses. London: Methuen & Co. Ltd. ISBN 0-412-
15740-3.
• Lembi, C.A.; Waaland, J.R. (1988). Algae and Human Affairs. Cambridge: Cambridge
• Round, F E (1981). The Ecology of Algae. London: Cambridge University Press. ISBN 0-521-
22583-3.
• Mumford, T F; Miura, A (1988). "Porphyra as food: cultivation and economic". In Lembi, C A;
Waaland, J R. Algae and Human Affairs. Cambridge University Press. pp. 87–117. ISBN 0-521-
32115-8. .
[edit] Regional
Britain and Ireland
• Brodie, Juliet; Burrows, Elsie M; Chamberlain, Yvonne M.; Christensen, Tyge; Dixon, Peter
Stanley; Fletcher, R.L.; Hommersand, Max H; Irvine, Linda M et al. (1977–2003). Seaweeds of the
British Isles: A Collaborative Project of the British Phycological Society and the British Museum
(Natural History). London, Andover: British Museum (Natural History), HMSO, Intercept.
ISBN 9780565007812. 9780113100453, 9780113100163, 9781898298878, 9780565009809,
0565009818, 0565009923, 0565008714
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• Cullinane, John P (1973). Phycology of the South Coast of Ireland . Cork: Cork University
Press.
• Hardy, F G; Aspinall, R J (1988). An Atlas of the Seaweeds of Northumberland and Durham.
The Hancock Museum, University Newcastle upon Tyne: Northumberland Biological Records Centre.
ISBN 978-0-9509680-5-6.
• Hardy, F G; Guiry, Michael D; Arnold, Henry R (2006). A Check-list and Atlas of the
Seaweeds of Britain and Ireland (Revised ed.). London: British Phycological Society. ISBN 3-
906166-35-X 9783906166353.
• John, D M; Whitton, B A; Brook, J A (2002). The Freshwater Algal Flora of the British Isles.
Cambridge, UK; New York: Cambridge University Press. ISBN 0-521-77051-3.
• Knight, Margery; Parke, Mary W (1931). Manx Algae: An Algal Survey of the South End of
the Isle of Man. Liverpool Marine Biology Committee (LMBC) Memoirs on Typical British Marine
Plants & Animals. XXX. Liverpool: University Press.
• Morton, Osborne (1994). Marine Algae of Northern Ireland . Belfast: Ulster Museum.
ISBN 0900761288 9780900761287.
• Morton, Osborne (1 December 2003). "The Marine Macroalgae of County Donegal, Ireland".
Bulletin of the Irish Biogeographical Society 27: 3–164.
Australia
• Huisman, J M (2000). Marine Plants of Australia. University of Western Australian (UWA)
Press. ISBN 1-876268-33-6.
New Zealand
• Chapman, Valentine Jackson; Lindauer, VW; Aiken, M; Dromgoole, FI (1900, 1956, 1961,
1969, 1970). The Marine algae of New Zealand. London; Lehre, Germany: Linnaean Society of
London; Cramer.
Europe
• Cabioc'h, Jacqueline; Floc'h, Jean-Yves; Le Toquin, Alain; Boudouresque, Charles-François;
Meinesz, Alexandre; Verlaque, Marc (1992) (in French). Guide des algues des mers d'Europe:
Manche/Atlantique-Méditerranée. Lausanne, Suisse: Delachaux et Niestlé. ISBN 2-603-00848-X,
9782603008485.
• Gayral, Paulette (1966) (in French). Les Algues de côtes françaises (manche et atlantique),
notions fondamentales sur l'écologie, la biologie et la systématique des algues marines . Paris: Doin,
Deren et Cie.
• Guiry, M.D.; Blunden, G. (1991). Seaweed Resources in Europe: Uses and Potential. John
Wiley & Sons. ISBN 0-471-92947-6.
• Míguez Rodríguez, Luís (1998) (in Galician). Algas mariñas de Galicia: bioloxía,
gastronomía, industria. Vigo: Edicións Xerais de Galicia. ISBN 84-8302-263-X.
• Otero, J. (2002) (in Galician). Guía das macroalgas de Galicia . A Coruña: Baía Edicións.
ISBN 84-89803-22-6.
• Bárbara, I.; Cremades, J. (1993) (in Spanish). Guía de las algas del litoral gallego. A Coruña:
Concello da Coruña - Casa das Ciencias.
Arctic
• Kjellman, Frans Reinhold (1883). The algae of the Arctic Sea: a survey of the species,
together with an exposition of the general characters and the development of the flora . 20.
Stockholm: Kungl. Svenska vetenskapsakademiens handlingar. pp. 1–350.
Greenland
• Lund, Søren Jensen (1959). The Marine Algae of East Greenland. Kövenhavn: C.A. Reitzel.
ISBN 9584734.
Faroe Islands
• Børgesen, Frederik (1903, 1970 reprint). "Marine Algae". In Warming, Eugene. Botany of the
Faröes Based Upon Danish Investigations. Part II. Det nordiske Forlag. pp. 339–532. .
Canary Islands
• Børgesen, Frederik (1925, 1926, 1927, 1929, 1930, 1936). Marine Algae from the Canary
Islands. København: Bianco Lunos.
Morocco
• Gayral, Paulette (1958) (in French). Algues de la côte atlantique marocaine. Casablanca:
Rabat [Société des sciences naturelles et physiques du Maroc].
South Africa
• Stegenga, H.; Bolton, J.J.; Anderson, R.J. (1997). Seaweeds of the South African West
Coast. Bolus Herbarium, University of Cape Town. ISBN 0-7992-1793-X.
North America
• Abbott, I.A.; Hollenberg, G.J. (1976). Marine Algae of California. California: Stanford
• Greeson, Phillip E. (1982). An annotated key to the identification of commonly occurring and
dominant genera of Algae observed in the Phytoplankton of the United States . Washington, D.C.:
U.S. Department of the Interior, Geological Survey.
http://www.archive.org/details/annotatedkeytoid00gree. Retrieved 2008-12-19.
• Taylor, William Randolph (1937, 1957, 1962, 1969). Marine Algae of the Northeastern Coast
of North America. Ann Arbor: University of Michigan Press. ISBN 0-472-04904-6.
• Wehr, J D; Sheath, R G (2003). Freshwater Algae of North America: Ecology and
Classification. USA: Academic Press. ISBN 0-12-741550-5.

Wikimedia Commons has media related to: Algae
• Guiry, Michael and Wendy. "AlgaeBase". http://www.algaebase.org. - a database of all algal
names including images, nomenclature, taxonomy, distribution, bibliography, uses, extracts
• Algae - Cell Centered Database
• "Algae Research". National Museum of Natural History, Department of Botany. 2008.
http://botany.si.edu/projects/algae/. Retrieved 2008-12-19.
• Anderson, Don; Bruce Keafer; Judy Kleindinst; Katie Shaughnessy; Katherine Joyce;
Danielle Fino; Adam Shepherd (2007). "Harmful Algae". U.S. National Office for Harmful Algal
Blooms. http://www.whoi.edu/redtide/page.do?pid=14779. Retrieved 2008-12-19.
• "Australian Freshwater Algae (AFA)". Department of Environment and Climate Change NSW
Botanic Gardens Trust.
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• Silva, Paul (1997–2004). "Index Nominum Algarum (INA)". Berkeley: University Herbarium,
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• Algae: Protists with Chloroplasts
• "Research on microalgae". Wageningen UR. 2009. http://www.algae.wur.nl/uk/. Retrieved
2009-05-18.
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W000
Potato

"Irish potato" redirects here. For the confectionary, see Irish potato candy.
For other uses, see Potato (disambiguation).
Potato
Kingdom: Plantae
(unranked): Angiosperms
(unranked): Eudicots
(unranked): Asterids
Order: Solanales
Family: Solanaceae
Genus: Solanum
Species: S. tuberosum
Binomial name
Solanum tuberosum
L.
The potato is a starchy, tuberous crop from the perennial Solanum tuberosum of the Solanaceae
family (also known as the nightshades). The word potato may refer to the plant itself as well as the edible
tuber. In the region of the Andes, there are some other closely related cultivated potato species. First
introduced outside the Andes region four centuries ago, today potatoes have become an integral part of
much of the world's cuisine and are the world's fourth-largest food crop, following rice, wheat, and maize.[1]
Long-term storage of potatoes requires specialised care in cold warehouses.[2]
Wild potato species occur throughout the Americas, from the United States to Uruguay.[3] The potato
was originally believed to have been domesticated independently in multiple locations,[4] but later genetic
testing of the wide variety of cultivars and wild species proved a single origin for potatoes in the area of
present-day southern Peru (from a species in the Solanum brevicaule complex), where they were
domesticated 7,000–10,000 years ago.[5][6][7] Following centuries of selective breeding, over a thousand
different types of potatoes currently exist in the world.[6] Of these subspecies, a variety that at one point grew
in the Chiloé Archipelago (the potato's south-central Chilean sub-center of origin) left its germplasm on over
99% of the cultivated potatoes worldwide.[8][9]
Following the Spanish conquest of the Inca Empire, the Spanish introduced the potato to Europe in
the second half of the 16th century. The staple was subsequently conveyed by European mariners to
territories and ports throughout the world. The potato was slow to be adopted by distrustful European
farmers, but soon enough it became an important food staple and field crop that played a major role in the
European 19th century population boom.[7] However, lack of genetic diversity, due to the very limited number
of varieties initially introduced, left the crop vulnerable to disease. In 1845, a plant disease known as late
blight, caused by the fungus-like oomycete Phytophthora infestans, spread rapidly through the poorer
communities of western Ireland, resulting in the crop failures that led to the Great Irish Famine. Nonetheless,
thousands of varieties persist in the Andes, where over 100 cultivars might be found in a single valley, and a
dozen or more might be maintained by a single agricultural household.[10]
The annual diet of an average global citizen in the first decade of the 21st century included about
33 kg (73 lb) of potato. However, the local importance of potato is extremely variable and rapidly changing. It
remains an essential crop in Europe (especially eastern and central Europe), where per capita production is
still the highest in the world, but the most rapid expansion over the past few decades has occurred in
southern and eastern Asia. China is now the world's largest potato-producing country, and nearly a third of
the world's potatoes are harvested in China and India.[11]
Contents
[hide]
• 1 Etymology
• 2 Characteristics
• 3 Genetics
• 4 History
• 4.1 Peru
• 4.2 Europe
• 4.2.1 19th century Europe
• 4.2.2 Ireland
• 4.3 Canary Islands Study
• 4.4 Africa
• 4.4.1 Uses
• 4.4.2 Industrialization of potatoes
• 4.4.3 Rwandan potatoes
• 4.5 Asia
• 4.6 US and Canada
• 5 Role in world food supply
• 6 Nutrition
• 6.1 Toxicity
• 7 Growth and cultivation
• 7.1 Storage
Etymology
"Spud" redirects here. For other uses, see Spud (disambiguation).
The English word potato comes from Spanish patata (the name used in Spain). The Spanish Royal
Academy says the Spanish word is a compound of the Taino batata (sweet potato) and the Quechua papa
(potato).[12] The name potato originally referred to a type of sweet potato rather than the other way around,
although there is actually no close relationship between the two plants. The English confused the two plants
one for the other. In many of the chronicles detailing agriculture and plants, no distinction is made between
the two.[13] The 16th-century English herbalist John Gerard used the terms "bastard potatoes" and "Virginia
potatoes" for this species, and referred to sweet potatoes as "common potatoes".[14] Potatoes are
occasionally referred to as "Irish potatoes" or "white potatoes" in the United States, to distinguish them from
sweet potatoes.[14]
The name spud for a small potato comes from the digging of soil (or a hole) prior to the planting of
potatoes. The word is of unknown origin and was originally (c. 1440) used as a term for a short knife or
dagger, probably related to Dutch spyd and/or the Latin "spad-" root meaning "sword"; cf. Spanish "espada",
English "spade" and "spadroon". The word spud traces back to the 16th century. It subsequently transferred
over to a variety of digging tools. Around 1845 it transferred over to the tuber itself.[15] The origin of "spud"
has erroneously been attributed to a 19th century activist group dedicated to keeping the potato out of Britain,
calling itself The Society for the Prevention of an Unwholesome Diet.[15] It was Mario Pei's 1949 The Story
of Language that can be blamed for the false origin. Pei writes, "the potato, for its part, was in disrepute some
centuries ago. Some Englishmen who did not fancy potatoes formed a Society for the Prevention of
Unwholesome Diet. The initials of the main words in this title gave rise to spud." Like most other pre-20th
century acronymic origins, this one is false.[15]
Characteristics
Flowers of a potato plant
Russet potatoes with sprouts

Potato plants are herbaceous perennials that grow about 60 cm (24 in) high, depending on variety,
the culms dying back after flowering. They bear white, pink, red, blue, or purple flowers with yellow stamens.
In general, the tubers of varieties with white flowers have white skins, while those of varieties with colored
flowers tend to have pinkish skins.[16] Potatoes are cross-pollinated mostly by insects, including
bumblebees, which carry pollen from other potato plants, but a substantial amount of self-fertilizing occurs as
well. Tubers form in response to decreasing day length, although this tendency has been minimized in
commercial varieties.[17]
Potato plants
After potato plants flower, some varieties produce small green fruits that resemble green cherry
tomatoes, each containing up to 300 true seeds. Potato fruit contains large amounts of the toxic alkaloid
solanine and is therefore unsuitable for consumption. All new potato varieties are grown from seeds, also
called "true seed" or "botanical seed" to distinguish it from seed tubers. By finely chopping the fruit and
soaking it in water, the seeds separate from the flesh by sinking to the bottom after about a day (the
remnants of the fruit float). Any potato variety can also be propagated vegetatively by planting tubers, pieces
of tubers, cut to include at least one or two eyes, or also by cuttings, a practice used in greenhouses for the
production of healthy seed tubers. Some commercial potato varieties do not produce seeds at all (they bear
imperfect flowers) and are propagated only from tuber pieces. Confusingly, these tubers or tuber pieces are
called "seed potatoes".
Genetics
The major species grown worldwide is Solanum tuberosum (a tetraploid with 48 chromosomes), and
modern varieties of this species are the most widely cultivated. There are also four diploid species (with 24
chromosomes): S. stenotomum, S. phureja, S. goniocalyx, and S. ajanhuiri. There are two triploid species
(with 36 chromosomes): S. chaucha and S. juzepczukii. There is one pentaploid cultivated species (with 60
chromosomes): S. curtilobum.
There are two major subspecies of Solanum tuberosum: andigena, or Andean; and tuberosum, or
Chilean.[18] The Andean potato is adapted to the short-day conditions prevalent in the mountainous
equatorial and tropical regions where it originated. The Chilean potato is adapted to the long-day conditions
prevalent in the higher latitude region of southern Chile, especially on Chiloé Archipelago where it is thought
to have originated.[19] Genetic testing done in 2005 showed that both subspecies derive from a common
ancestor from the area of southern Peru.[20]
There are about five thousand potato varieties worldwide. Three thousand of them are found in the
Andes alone, mainly in Peru, Bolivia, Ecuador, Chile, and Colombia. They belong to eight or nine species,
depending on the taxonomic school. Apart from the five thousand cultivated varieties, there are about 200
wild species and subspecies, many of which can be cross-bred with cultivated varieties, which has been
done repeatedly to transfer resistances to certain pests and diseases from the gene pool of wild species to
the gene pool of cultivated potato species. Genetically modified varieties have met public resistance in the
United States and in the European Union.[21][22]
Most modern potatoes grown in North America arrived through European settlement and not
independently from the South American sources. However, at least one wild potato species, Solanum
fendleri, is found as far north as Texas and used in breeding for resistance to a nematode species that
attacks cultivated potatoes. A secondary center of genetic variability of the potato is Mexico, where important
wild species that have been used extensively in modern breeding are found, such as the hexaploid Solanum
demissum, as a source of resistance to the devastating late blight disease. Another relative native to this
region, Solanum bulbocastanum, has been used to genetically engineer the potato to resist potato blight.[23]
The International Potato Center, based in Lima, Peru, holds an ISO-accredited collection of potato
germplasm.[24]
Potatoes yield abundantly with little effort, and adapt readily to diverse climates as long as the
climate is cool and moist enough for the plants to gather sufficient water from the soil to form the starchy
tubers. Potatoes do not keep very well in storage and are vulnerable to molds that feed on the stored tubers,
quickly turning them rotten. By contrast, grain can be stored for several years without much risk of rotting.[25]
History
Peru
The potato originated in the region of southern Peru.[5] Potatoes were first domesticated in Peru
between 3000 BC and 2000 BC. In the Altiplano, potatoes provided the principal energy source for the Inca
Empire, its predecessors, and its Spanish successor. In Peru above 10,000 feet altitude, tubers exposed to
the cold night air turned into chuño; when kept in permanently frozen underground storehouses, chuño can
be stored for years with no loss of nutritional value. The Spanish fed chuño to the silver miners who produced
vast wealth in the 16th century for the Spanish government.[6]
Europe
Sailors returning from Peru to Spain with silver presumably brought maize and potatoes for their own
food on the trip. Historians speculate that leftover tubers (and maize) were carried ashore and planted.
Basque fishermen from Spain used potatoes as ships stores for their voyages across Atlantic in the 16th
century, and introduced the tuber to western Ireland, where they landed to dry their cod. In 1553, in the book
Crónica del Peru, Pedro Cieza de Leon mentions he saw it in Quito, Popayán and Pasto. Though the English
privateer Francis Drake, returning from his circumnavigation, or Sir Walter Raleigh's employee Thomas
Harriot[26] are commonly credited with introducing potatoes into England, Mary, Queen of Scots already
complained in 1569 of her enforced confinement at Tutbury Castle that its garden was no more than "a
potato patch... fitter to keep pigs in."[27] In 1588, botanist Carolus Clusius made a painting of what he called
"Papas Peruanorum" from a specimen in Belgium; in 1601 he reported that potatoes were in common use in
northern Italy for animal fodder and for human consumption.[28]
The Spanish had an empire across Europe, and brought potatoes for their armies. Peasants along
the way adopted the crop, which was less often pillaged by marauding armies than above-ground stores of
grain. Across most of northern Europe, where open fields prevailed, potatoes were strictly confined to small
garden plots because field agriculture was strictly governed by custom that prescribed seasonal rhythms for
plowing, sowing, harvesting and grazing animals on fallow and stubble. This meant that potatoes were barred
from large-scale cultivation because the rules allowed only grain to be planted in the open fields.[29] In
France and Germany government officials and noble landowners promoted the rapid conversion of fallow
land into potato fields after 1750. The potato thus became an important staple crop in northern Europe.
Famines in the early 1770s contributed to its acceptance, as did government policies in several European
countries and climate change during the Little Ice Age, when traditional crops in this region did not produce
as reliably as before.[30][31] At times when and where most other crops failed, potatoes could still typically
be relied upon to contribute adequately to food supplies during colder years.[32]
The potato was not popular in France before 1800. It took time to be popularly adopted, but had
widely replaced the turnip and rutabaga by the 19th century.[33] Today, the potato forms an important part of
the traditional cuisines of most of Europe. Belarus has the highest consumption of potato per capita, with
each Belorussian consuming 381 kg in 2005—about two pounds per person per day.[34]
19th century Europe

French physician Antoine Parmentier studied the potato intensely and in Examen chymique des
pommes de terres (Paris, 1774) showed their enormous nutritional value. King Louis XVI and his court
eagerly promoted the new crop, with Queen Marie Antoinette even wearing a headdress of potato flowers at
a fancy dress ball. The annual potato crop of France soared to 21 million hectoliters in 1815 and 117 millions
in 1840, allowing a concomitant growth in population while avoiding the Malthusian trap. Although potatoes
had become widely familiar in Russia by 1800, they were confined to garden plots until the grain failure in
1838–1839 persuaded peasants and landlords in central and northern Russia to devote their fallow fields to
raising potatoes. Potatoes yielded from two to four times more calories per acre than grain did, and
eventually came to dominate the food supply in eastern Europe. Boiled or baked potatoes were cheaper than
rye bread, just as nutritious, and did not require a gristmill for grinding. On the other hand cash-oriented
landlords realized that grain was much easier to ship, store and sell, so both grain and potatoes coexisted.
[35]
Antoine Parmentier holding New World plants, François Dumont 1812
Throughout Europe, the most important new food in the 19th century was the potato, which had three
major advantages over other foods for the consumer: its lower rate of spoilage, its bulk (which easily satisfied
hunger), and its cheapness. The crop slowly spread across Europe, such that, for example, by 1845 it
occupied one-third of Irish arable land.[ citation needed] Potatoes comprised about 10% of the caloric intake
of Europeans.[citation needed] Along with several other foods that either originated in the Americas or were
successfully grown or harvested there, potatoes sustained European populations.[36]
In Britain, the potato promoted economic development by underpinning the Industrial Revolution in
the 19th century. It served as a cheap source of calories and nutrients that was easy for urban workers to
cultivate on small backyard plots. Potatoes became popular in the north of England, where coal was readily
available, so a potato-driven population boom provided ample workers for the new factories. Marxist
Friedrich Engels even declared that the potato was the equal of iron for its "historically revolutionary role.[28]
The Dutch potato-starch industry grew rapidly in the 19th century, especially under the leadership of
entrepreneur Willem Albert Scholten (1819–92).[37]
Ireland
In Ireland, the expansion of potato cultivation was due entirely to the landless laborers, renting tiny
plots from landowners, who were interested only in raising cattle or in producing grain for market. A single
acre of potatoes and the milk of a single cow was enough to feed a whole Irish family a monotonous but
nutritionally adequate diet for a healthy, vigorous (and desperately poor) rural population. Often even poor
families grew enough extra potatoes to feed a pig that they could sell for cash.[38]
A lack of genetic diversity from the low number of varieties left the crop vulnerable to disease. In
1845, a plant disease known as late blight, caused by the fungus-like oomycete Phytophthora infestans,
spread rapidly through the poorer communities of western Ireland, resulting in the crop failures that led to the
Great Irish Famine.[39]
The Lumper potato, widely cultivated in western and southern Ireland before and during the great
famine, was bland, wet, and poorly resistant to the potato blight, but yielded large crops and usually provided
adequate calories for peasants and laborers. Heavy dependence on this potato led to disaster when the
potato blight turned a newly harvested potato into a putrid mush in minutes. The Irish Famine in the western
and southern parts of the British-controlled island of Ireland, 1845–49, was a catastrophic failure in the food
supply that led to approximately a million deaths from famine and (especially) diseases that attacked
weakened bodies, and to massive emigration to Britain, the U.S. and Canada.[40]
Canary Islands Study
Shipping records from 1567 make the Canary Islands islands off the shores northwest Africa the first
known home to potatoes outside of Central and South America. In 2007, David Spooner, a horticulturist with
the U.S. Department of Agriculture and a researcher at the University of Wisconsin, Madison, decided to
analyze potatoes on the island.[41] He found that some potatoes on the Canary Islands had genetic markers
of Andean origins and some had markers indicating Chilean roots.[41] Two subspecies of these wild spuds,
one found in Chile, the other in the Andean highlands of Peru, look very similar but differ genetically.[41]
Most scientists have long assumed that European potatoes, the foundation for all modern cultivated
potatoes, come from the Chilean variety, because Chilean lowlands resemble Europe's environment most
closely.[41] According to Spooner, "The idea that it was a single introduction from Chile just doesn't stand
up,". Spooner suggests that different varieties could have been brought from South America at various times.
[41]
Africa
It is generally believed that Potatoes also entered Africa with colonists, who consumed them as a
vegetable rather than as a staple starch.[42] Shipping records from 1567 make the Canary Islands islands off
the shores northwest Africa the first known home to potatoes outside of Central and South America though.
[41] Like in other continents, in spite of its advantages as an antifamine, high-elevation alternative to grain,
the potato was at first resisted by local farmers who thought it was poisonous. The colonialists also promoted
it as a low cost food to them and so it was a symbol of domination.[42] In former European colonies of Africa,
potatoes were initially consumed only occasionally, but increased production made them a staple in certain
areas. In Africa, as in Europe, the popularity of the tubers increased in wartime because they could be stored
in the ground.[42]
In present day Africa it has been a vegetable or co-staple crop.[42]
Uses
In central African regions of relatively high production, potatoes are beaten with grains and legumes
into a stiff porridge, or boiled or roasted and eaten whole.[42]
Industrialization of potatoes
Like in many parts of the world, diet has been globalized and it is currently also used to make
industrialized produce like produced potato fries (chips), chips (crisps).[42]
Rwandan potatoes
In the upper reaches of Rwanda, potatoes took root as a new staple food crop and contributed to
subsistence, surplus, and population expansion.[42] Prior to its most recent civil conflict, Rwanda in some
localities witnessed per capita consumption as high as 153 to 200 kg per year — higher than that in any
Western European country, including Ireland.[42] There, local farmers in recent years have developed the
potato as a cash crop — the result of the introduction of several new varieties brought back by migrant
laborers from Uganda, the diffusion of other varieties from Kenya, and the comparative advantage of raising
potatoes relative to other cash or subsistence crops.[42]
Asia
The potato diffused widely after 1600, becoming a major food resource in Europe and East Asia.
Following its introduction into China toward the end of the Ming dynasty, the potato immediately became a
delicacy of the imperial family. After the middle period of the Qianlong reign (1735–96), population increases
and a subsequent need to increase grain yields coupled with greater peasant geographic mobility led to the
rapid spread of potato cultivation throughout China, and it was acclimated to local natural conditions.
Boomgaard (2003) looks at the adoption of various root and tuber crops in Indonesia throughout the
colonial period and examines the chronology and reasons for progressive adoption of foreign crops: Sweet
potato, Irish potato, bengkuang (yam beans), and Cassava.
The potato was introduced in the Philippines during the late 16th century, and to Java and China
during the 17th century. It was well-established as a crop in India by the late 18th century and in Africa by the
mid-20th century.[31]
US and Canada
Potatoes were planted in Idaho as early as 1838; by 1900 the state's production exceeded a million
bushels (about 27,000 tonnes[44]). Prior to 1910, the crops were stored in barns or root cellars, but, by the
1920s, potato cellars came into use. U.S. potato production has increased steadily; two-thirds of the crop
comes from Idaho, Washington, Oregon, Colorado, and Maine, and potato growers have strengthened their
position in both domestic and foreign markets.
By the 1960s, the Canadian Potato Research Centre in Fredericton, New Brunswick, was one of the
top six potato research institutes in the world. Established in 1912 as a dominion experimental station, the
station began in the 1930s to concentrate on breeding new varieties of disease-resistant potatoes. In the
1950s–60s the growth of the French fry industry in New Brunswick led to a focus on developing varieties for
the industry. By the 1970s, the station's potato research was broader than ever before, but the station and its
research programs had changed, as emphasis was placed on serving industry rather than potato farmers in
general. Scientists at the station even began describing their work using engineering language rather than
scientific prose.[45] Potatoes are Canada's most important vegetable crop; they are grown commercially in
all its provinces, led by PEI.[46]
Role in world food supply
Worldwide potato production

Top Potato Producers
in 2006
(million metric tons)
People's Republic of China 70
Russia 39
India 24
United States 20
Ukraine 19
Germany 10
Poland 9
Belgium 8
Netherlands 7
France 6
World Total 315
Source:
UN Food & Agriculture Organisation
(FAO)[4]
Potato output in 2005
The United Nations FAO reports that the world production of potatoes in 2008 was 314 million
tonnes.[47] The annual diet of an average global citizen in the first decade of the 21st century included about
33 kg (or 73 lb) of potato. However, the local importance of potato is extremely variable and rapidly changing.
It remains an essential crop in Europe (especially eastern and central Europe), where per capita production
is still the highest in the world, but the most rapid expansion over the past few decades has occurred in
southern and eastern Asia. China is now the world's largest potato-producing country, and nearly a third of
the world's potatoes are harvested in China and India.[11] The geographic shift of potato production has
been away from wealthier countries toward lower-income areas of the world, although the degree of this
trend is ambiguous.[48]
In 2008, several international organizations highlighted the potato's role in world food production, in
the face of developing economic problems. They cited its potential derived from its status as a cheap and
plentiful crop that grows in a wide variety of climates and locales.[49] Due to perishability, only about 5% of
the world's potato crop is traded internationally; its minimal presence in world financial markets contributed to
its stable pricing during the 2007–2008 world food price crisis.[50][51] Thus, the United Nations officially
declared the year 2008 as the International Year of the Potato,[52] to raise its profile in developing nations,
calling the crop a "hidden treasure".[53] This followed the International Rice Year in 2004.
Nutrition
Potato, raw, with peel
Nutritional value per 100 g (3.5 oz)
Energy 321 kJ (77 kcal)
Carbohydrates 19 g
Starch 15 g
Dietary fiber 2.2 g
Fat 0.1 g
Protein 2g
Water 75 g
Thiamine (Vit. B1) 0.08 mg (6%)
Riboflavin (Vit. B2) 0.03 mg (2%)
Niacin (Vit. B3) 1.1 mg (7%)
Vitamin B6 0.25 mg (19%)

Vitamin C 20 mg (33%)
Calcium 12 mg (1%)
Iron 1.8 mg (14%)
Magnesium 23 mg (6%)
Phosphorus 57 mg (8%)
Potassium 421 mg (9%)
Sodium 6 mg (0%)
Percentages are relative to US recommendations for adults.

The potato contains vitamins and minerals, as well as an assortment of phytochemicals, such as
carotenoids and polyphenols. Chlorogenic acid constitutes up to 90% of the potato tuber polyphenols. Others
polyphenols found in potatoes are 4-O-caffeoylquinic (crypto-chlorogenic acid), 5-O-caffeoylquinic (neo-
chlorogenic acid), 3,4-dicaffeoylquinic and 3,5-dicaffeoylquinic acids.[54] A medium-size 150 g (5.3 oz)
potato with the skin provides 27 mg of vitamin C (45% of the Daily Value (DV)), 620 mg of potassium (18% of
DV), 0.2 mg vitamin B6 (10% of DV) and trace amounts of thiamin, riboflavin, folate, niacin, magnesium,
phosphorus, iron, and zinc. The fiber content of a potato with skin (2 g) is equivalent to that of many whole
grain breads, pastas, and cereals.
In terms of nutrition, the potato is best known for its carbohydrate content (approximately 26 grams in
a medium potato). The predominant form of this carbohydrate is starch. A small but significant portion of this
starch is resistant to digestion by enzymes in the stomach and small intestine, and so reaches the large
intestine essentially intact. This resistant starch is considered to have similar physiological effects and health
benefits as fiber: It provides bulk, offers protection against colon cancer, improves glucose tolerance and
insulin sensitivity, lowers plasma cholesterol and triglyceride concentrations, increases satiety, and possibly
even reduces fat storage.[55][56][57] The amount of resistant starch in potatoes depends much on
preparation methods. Cooking and then cooling potatoes significantly increased resistant starch. For
example, cooked potato starch contains about 7% resistant starch, which increases to about 13% upon
cooling.[58]
The nutrients of the potato seem to be fairly evenly distributed between the flesh and the skin. For a
medium potato, with and without the skin, nutritiondata.com gives the following:[59][60]
Nutrient Without skin (156 g) (% RDA) With skin (173 g) (% RDA)
Vitamin C 33 28
Thiamin 11 7
Niacin 11 12
Vitamin B6 23 27
Folate 4 12
Pantothenic Acid 9 7
Iron 3 10
Magnesium 10 12
Potassium 17 26
Copper 17 10
Dietary Fiber 9 15
The cooking method used can significantly impact the nutrient availability of the potato.
Potatoes are often broadly classified as high on the glycemic index (GI) and so are often excluded
from the diets of individuals trying to follow a low-GI diet. In fact, the GI of potatoes can vary considerably
depending on type (such as red, russet, white, or Prince Edward), origin (where it was grown), preparation
methods (i.e., cooking method, whether it is eaten hot or cold, whether it is mashed or cubed or consumed
whole, etc.), and with what it is consumed (i.e., the addition of various high-fat or high-protein toppings).[61]
Potatoes are not considered by the NHS as counting towards the five portions of fruit and vegetables
diet.[62]
Toxicity
Early Rose variety seed tuber with sprouts

Potatoes contain toxic compounds known as glycoalkaloids, of which the most prevalent are solanine
and chaconine. Solanine is also found in other plants in the family Solanaceae, which includes such plants as
the deadly nightshade (Atropa belladonna), henbane (Hyoscyamus niger) and tobacco (Nicotiana) as well as
the potato, eggplant, and tomato. This toxin affects the nervous system, causing weakness and confusion.
These compounds, which protect the plant from its predators, are, in general, concentrated in its
leaves, stems, sprouts, and fruits.[63] Exposure to light, physical damage, and age increase glycoalkaloid
content within the tuber;[64] the highest concentrations occur just underneath the skin. Cooking at high
temperatures (over 170 °C or 340 °F) partly destroys these. The concentration of glycoalkaloid in wild
potatoes suffices to produce toxic effects in humans. Glycoalkaloids may cause headaches, diarrhea,
cramps, and in severe cases coma and death; however, poisoning from potatoes occurs very rarely. Light
exposure causes greening from chlorophyll synthesis, thus giving a visual clue as to areas of the tuber that
may have become more toxic; however, this does not provide a definitive guide, as greening and
glycoalkaloid accumulation can occur independently of each other. Some varieties of potato contain greater
glycoalkaloid concentrations than others; breeders developing new varieties test for this, and sometimes
have to discard an otherwise promising cultivar.
The toxic fruits produced by mature potato plants

Breeders try to keep solanine levels below 200 mg/kg (200 ppmw). However, when these
commercial varieties turn green, even they can approach concentrations of solanine of 1000 mg/kg (1000
ppmw). In normal potatoes, analysis has shown solanine levels may be as little as 3.5% of the breeders'
maximum, with 7–187 mg/kg being found.[65]
The U.S. National Toxicology Program suggests that the average American consume at most
12.5 mg/day of solanine from potatoes (the toxic dose is actually several times this, depending on body
weight). Douglas L. Holt, the State Extension Specialist for Food Safety at the University of Missouri, notes
that no reported cases of potato-source solanine poisoning have occurred in the U.S. in the last 50 years,
and most cases involved eating green potatoes or drinking potato-leaf tea.[ citation needed]
Growth and cultivation
Potato planting in Washington

Potato field in Fort Fairfield, Maine
Potatoes grown in a tall bag are common in gardens as they increase potato yield and minimize the
amount of digging required at harvest
Potato growth has been divided into five phases. During the first phase, sprouts emerge and root
growth begins. During the second, photosynthesis begins as the plant develops leaves and branches. New
tubers develop during the third phase, which is often (but not always) associated with flowering. Tuber
formation halts when soil temperatures reach 80 °F (26.7 °C); hence potatoes are considered a cool-season
crop.[66] Tuber bulking occurs during the fourth phase, when the plant begins investing the majority of its
resources in its newly formed tubers. At this stage, several factors are critical to yield: optimal soil moisture
and temperature, soil nutrient availability and balance, and resistance to pest attacks. The final phase is
maturation: The plant canopy dies back, the tuber skins harden, and their sugars convert to starches.[67]
New tubers may arise at the soil surface. Since exposure to light leads to greening of the skins and
the development of solanin, growers are interested in covering such tubers. Commercial growers usually
address this problem by piling additional soil around the base of the plant as it grows ("hilling", or in British
English "earthing up"). An alternative method used by home gardeners and smaller-scale growers involves
covering the growing area with organic mulches such as straw or with plastic sheets.[67]
Correct potato husbandry can be an arduous task in some circumstances. Good ground preparation,
harrowing, plowing, and rolling are always needed, along with a little grace from the weather and a good
source of water. Three successive plowings, with associated harrowing and rolling, are desirable before
planting. Eliminating all root-weeds is desirable in potato cultivation. In general, the potatoes themselves are
grown from the eyes of another potato and not from seed. Home gardeners often plant a piece of potato with
two or three eyes in a hill of mounded soil. Commercial growers plant potatoes as a row crop using seed
tubers, young plants or microtubers and may mound the entire row. Seed potato crops are 'rogued' in some
countries to eliminate diseased plants or those of a different variety from the seed crop.
Potatoes are sensitive to heavy frosts, which damage them in the ground. Even cold weather makes
potatoes more susceptible to bruising and possibly later rotting, which can quickly ruin a large stored crop.
At harvest time, gardeners usually dig up potatoes with a long-handled, three-prong "grape" (or
graip), i.e., a spading fork, or a potato hook, which is similar to the graip but with tines at a 90 degree angle to
the handle. In larger plots, the plow is the fastest implement for unearthing potatoes. Commercial harvesting
is typically done with large potato harvesters, which scoop up the plant and surrounding earth. This is
transported up an apron chain consisting of steel links several feet wide, which separates some of the dirt.
The chain deposits into an area where further separation occurs. Different designs use different systems at
this point. The most complex designs use vine choppers and shakers, along with a blower system or "Flying
Willard" to separate the potatoes from the plant. The result is then usually run past workers who continue to
sort out plant material, stones, and rotten potatoes before the potatoes are continuously delivered to a wagon
or truck. Further inspection and separation occurs when the potatoes are unloaded from the field vehicles
and put into storage.
Immature potatoes may be sold as "New Potatoes" and are particularly valued for taste. These are
often harvested by the home gardener or farmer by "grabbling", i.e. pulling out the young tubers by hand
while leaving the plant in place.
Potatoes are usually cured after harvest to improve skin-set. Skin-set is the process by which the
skin of the potato becomes resistant to skinning damage. Potato tubers may be susceptible to skinning at
harvest and suffer skinning damage during harvest and handling operations. Curing allows the skin to fully
set and any wounds to heal. Wound-healing prevents infection and water-loss from the tubers during storage.
Curing is normally done at relatively warm temperatures 50 °C (122 °F) to 60 °C (140 °F) with high humidity
and good gas-exchange if at all possible.[68]
Storage
Storage facilities need to be carefully designed to keep the potatoes alive and slow the natural
process of decomposition, which involves the breakdown of starch. It is crucial that the storage area is dark,
well ventilated and for long-term storage maintained at temperatures near 4 °C (39 °F). For short-term
storage before cooking, temperatures of about 7 °C (45 °F) to 10 °C (50 °F) are preferred.[2][69]
On the other hand, temperatures below 4 °C (39 °F) convert potatoes' starch into sugar, which alters
their taste and cooking qualities and leads to higher acrylamide levels in the cooked product, especially in
deep-fried dishes — the discovery of Acrylamides in starchy foods in 2002 has led to many international
health concerns as they are believed to be possible carcinogens and their occurrence in cooked foods are
currently under study as possible influences in potential health problems.[70][71]
Under optimum conditions possible in commercial warehouses, potatoes can be stored for up to ten
to twelve months.[2] When stored at homes, the shelf life is usually only for several weeks.[69] If potatoes
develop green areas or start to sprout, these areas should be trimmed before using.[69]
Commercial storage of potatoes involves several phases: drying of surface moisture; a wound
healing phase at 85% to 95% relative humidity and temperatures below 25 °C (77 °F); a staged cooling
phase; a holding phase; and a reconditioning phase, during which the tubers are slowly warmed. Mechanical
ventilation is used at various points during the process to prevent condensation and accumulation of carbon
dioxide.[2]
When stored in the home, mature potatoes are optimally kept at room temperature, where they last 1
to 2 weeks in paper bag, in a dry, cool, dark, well ventilated location.[72] If mature potatoes are refrigerated,
dark spots can occur and conversion of starch into sugar can give rise to unpleasant sweet flavour when
cooked.[72] Only new potatoes can be refrigerated, and should be kept so, where they have a shelflife of
1 week.[72] If kept in too warm temperatures, both mature and new potatoes will sprout and shrivel.[72]
Exposure to light causes them to turn green. Also, potatoes absorb odours produced by pears.[72]
Varieties
Different types of potato

Bamberg potatoes
Organically grown Russet Burbanks

While there are close to 4000 different varieties of potato,[73] it has been bred into many standard or
well-known varieties, each of which has particular agricultural or culinary attributes. In general, varieties are
categorized into a few main groups, such as russets, reds, whites, yellows (also called Yukons) and purples—
based on common characteristics. Around 80 varieties are commercially available in the UK.[74] For culinary
purposes, varieties are often described in terms of their waxiness. Floury, or mealy (baking) potatoes have
more starch (20–22%) than waxy (boiling) potatoes (16–18%). The distinction may also arise from variation in
the comparative ratio of two potato starch compounds: amylose and amylopectin. Amylose, a long-chain
molecule, diffuses out of the starch granule when cooked in water, and lends itself to dishes where the potato
is mashed. Varieties that contain a slightly higher amylopectin content, a highly branched molecule, help the
potato retain its shape when boiled.[75]
The European Cultivated Potato Database (ECPD) is an online collaborative database of potato
variety descriptions, updated and maintained by the Scottish Agricultural Science Agency within the
framework of the European Cooperative Programme for Crop Genetic Resources Networks (ECP/GR)—which
is organised by the International Plant Genetic Resources Institute (IPGRI).[76]
Popular varieties (cultivars) include:
• Adirondac • Clavela • Pink Eye
k Blue Blanca • Pink Fir
• Adirondac • Désirée Apple
k Red • Fianna • Primura
• Agata • Fingerli • Ratte
• Almond ng • Red
• Amandine • Flava Norland
• Anya • Golden • Red
• Arran Wonder Pontiac
Victory • Innovat • Rooster
• Atlantic or • Russet
• Bamberg • Jersey Burbank
• Belle de Royal • Russet
Fontenay • Kerr's Norkotah
• BF-15 Pink • Selma
• Bildtstar • Kestrel • Shepody
• Bintje • King • Sieglinde
• Blue Edward • Sirco
Congo • Kipfler • Spunta
• Bonnotte • Lady • Stobrawa
• Cabritas Balfour • Vivaldi
• Camota • Linda • Vitelotte
• Chelina • Marfona • Yellow
• Chiloé[77] • Maris Finn
• Cielo Piper • Yukon
• Marquis Gold
• Nicola
• Pachac
oña
Blue varieties
Potato variety 'Blue Swede

The blue potato originated in South America. It has purple skin and flesh, which becomes blue once
cooked. It has a slight whitish scab that seems to be present in all samples. The variety, called Cream of the
Crop has been introduced into Ireland and has proved popular.[78]
A mutation in the varieties' P locus causes production of the antioxidant anthocyanin.[79]
Genetically-modified varieties
Genetic research has produced several genetically modified varieties. 'New Leaf', owned by
Monsanto Company, incorporated genes from Bacillus thuringiensis, which conferred resistance to the
Colorado potato beetle; 'New Leaf Plus' and 'New Leaf Y', approved by US regulatory agencies during the
1990s, also included resistance to viruses. McDonald's, Burger King, Frito-Lay, and Procter & Gamble
announced they would not use genetically modified potatoes, and Monsanto published its intent to
discontinue the line in March 2001.[80] The starch content of 'Amflora', waxy potato variety from the German
chemical company BASF, has been modified to contain only amylopectin, making it inedible but more useful
for industrial purposes. In 2010, the European Commission cleared the way for 'Amflora' to be grown in the
European Union. Nevertheless, under EU rules, individual countries have the right to decide whether they will
allow this potato to be grown on their territory. Commercial planting of 'Amflora' was expected in the Czech
Republic and Germany in spring of 2010, and Sweden and the Netherlands in following years.[81] Another
GM potato variety developed by BASF is 'Fortuna'. In 2010, a team of Indian scientists announced they had
developed a genetically modified potato with 35 to 60% more protein than non-modified potatoes. Protein
content was boosted by adding the gene AmA1 from the grain amaranth. They also found 15 to 25% greater
crop yields with these potatoes.[82] The researchers expected that a key market for the GM potato wouuld be
the developing world, where more than a billion people are chronically undernourished.[83]
New non-GM varieties

Other new potato cultivars are conventionally bred. For example, on 22 September 2007, Benguet
State University (BSU) announced that four potato varieties—'Igorota', 'Solibao', 'Ganza' and one not yet
officially named—possess more than 18% dry matter content required by fast-food chains to make crispy and
sturdy French fries.[84] Since 2005, a natural 100% amylopectin waxy potato variety called 'Eliane' is being
cultivated by the starch company AVEBE.
Some horticulturists sell chimeras, made by grafting a tomato plant onto a potato plant, producing
both edible tomatoes and potatoes. This practice is not very widespread.
Pests
Main article: List of potato diseases
A potato ruined by late blight

The historically significant Phytophthora infestans (late blight) remains an ongoing problem in
Europe[85] and the United States.[86] Other potato diseases include Rhizoctonia, Sclerotinia, black leg,
powdery mildew, powdery scab, leafroll virus, and purple top.
Insects that commonly transmit potato diseases or damage the plants include the Colorado potato
beetle, the potato tuber moth, the green peach aphid (Myzus persicae), the potato aphid, beetleafhoppers,
thrips, and mites. The potato root nematode is a microscopic worm that thrives on the roots, thus causing the
potato plants to wilt. Since its eggs can survive in the soil for several years, crop rotation is recommended.
Pesticides
During the crop year 2008, many of the certified organic potatoes produced in the United Kingdom
and certified by the Soil Association as organic were sprayed with a copper pesticide[87] to control potato
blight (Phytophthora infestans).[88] According to the Soil Association, the total copper that can be applied to
organic land is 6 kg/ha/year.[89]
According to an Environmental Working Group analysis[90] of USDA and FDA pesticide residue tests
performed from 2000 through 2008, 84% of the 2,216 tested potato samples contained detectable traces of
at least one pesticide. A total of 36 unique pesticides were detected on potatoes over the 2,216 samples,
though no individual sample contained more than 6 unique pesticide traces, and the average was 1.29
detectable unique pesticide traces per sample. The average quantity of all pesticide traces found in the 2,216
samples was 1.602 ppm. While this is a very low value of pesticide residue, it is the highest amongst the 50
vegetables analyzed.
Uses
• Potatoes are used to brew alcoholic beverages such as vodka, potcheen, or akvavit.
• They are also used as food for domestic animals.
• Potato starch is used in the food industry as, for example, thickeners and binders of soups
and sauces, in the textile industry, as adhesives, and for the manufacturing of papers and boards.[91]
[92]
• Maine companies are exploring the possibilities of using waste potatoes to obtain polylactic
acid for use in plastic products; other research projects seek ways to use the starch as a base for
biodegradable packaging.[92][93]
Culinary uses
Various potato dishes

Potatoes are prepared in many ways: skin-on or peeled, whole or cut up, with seasonings or without.
The only requirement involves cooking to swell the starch granules. Most potato dishes are served hot, but
some are first cooked, then served cold, notably potato salad and potato chips/crisps.
Common dishes are: mashed potatoes, which are first boiled (usually peeled), and then mashed with
milk or yogurt and butter; whole baked potatoes; boiled or steamed potatoes; French-fried potatoes or chips;
cut into cubes and roasted; scalloped, diced, or sliced and fried (home fries); grated into small thin strips and
fried (hash browns); grated and formed into dumplings, Rösti or potato pancakes. Unlike many foods,
potatoes can also be easily cooked in a microwave oven and still retain nearly all of their nutritional value,
provided they are covered in ventilated plastic wrap to prevent moisture from escaping; this method produces
a meal very similar to a steamed potato, while retaining the appearance of a conventionally baked potato.
Potato chunks also commonly appear as a stew ingredient.
Potatoes are boiled between 10 and 25[94] minutes, depending on size and type, to become soft.
Latin America
Papa rellena
Peruvian cuisine naturally contains the potato as a primary ingredient in many dishes, as around
3,000 varieties of this tuber are grown there.[95] Some of the more notable dishes include boiled potato as a
base for several dishies or with ají-based sauces like in papa a la huancaina or ocopa, diced potato for its
use in soups like in cau cau, or in Carapulca with dried potato (papa seca). Smashed condimented potato is
used in causa Limeña and papa rellena. French-fried potatoes are a typical ingredient in Peruvian stir-fries,
including the classic dish lomo saltado.
Chuño is a freeze-dried potato product traditionally made by Quechua and Aymara communities of
Peru and Bolivia,[96] and is known in various countries of South America, including Peru, Bolivia, Argentina,
and Chile. In Chile's Chiloé Archipelago, potatoes are the main ingredient of many dishes, including milcaos,
chapaleles, curanto and chochoca. In Ecuador, the potato, as well as being a staple with most dishes, is
featured in the hearty locro de papas, a thick soup of potato, squash, and cheese.
European cuisine
Fish and chips
In the UK, potatoes form part of the traditional staple fish and chips. Roast potatoes are commonly
served with a Sunday roast, and mashed potatoes form a major component of several other traditional dishes
such as shepherd's pie, bubble and squeak, and bangers and mash. New potatoes are often cooked with
mint and served with a little melted butter.
The Tattie scone is a popular Scottish dish containing potatoes. Colcannon is a traditional Irish food
made with mashed potato, shredded kale or cabbage, and onion; champ is a similar dish. Boxty pancakes
are eaten throughout Ireland, although associated especially with the north, and in Irish diaspora
communities; they are traditionally made with grated potatoes, soaked to loosen the starch and mixed with
flour, buttermilk and baking powder. A variant eaten and sold in Lancashire, especially Liverpool, is made
with cooked and mashed potatoes.
Bryndzové halušky is the Slovakian national dish, made of a batter of flour and finely grated potatoes
that is boiled to form dumplings. These are then mixed with regionally varying ingredients.[97]
In Northern and Eastern Europe, especially in Scandinavian countries, Poland, Russia, Belarus and
Ukraine, newly harvested, early ripening varieties are considered a special delicacy. Boiled whole and served
un-peeled with dill, these "new potatoes" are traditionally consumed with Baltic herring. Puddings made from
grated potatoes (kugel, kugelis, and potato babka) are popular items of Ashkenazi, Lithuanian, and
Belarussian cuisine.[98]
A baked potato served with butter
In Western Europe, especially in Belgium, sliced potatoes are fried to create frieten, the original
French fried potatoes. Stamppot, a traditional Dutch meal, is based on mashed potatoes mixed with
vegetables.
In France, the most notable potato dish is the Hachis Parmentier, named after Antoine-Augustin
Parmentier, a French pharmacist, nutritionist, and agronomist who, in the late 18th century, was instrumental
in the acceptance of the potato as an edible crop in the country. The pâté aux pommes de terre is a regional
potato dish from the central Allier and Limousin regions.
In the north of Italy, in particular, in the Friuli region of the northeast, potatoes serve to make a type of
pasta called gnocchi.[99] Similarly, cooked and mashed potatoes or potato flour can be used in the Knödel or
dumpling eaten with or added to meat dishes all over central and Eastern Europe, but especially in Bavaria
and Luxembourg. Potatoes form one of the main ingredients in many soups such as the vichyssoise and
Albanian potato and cabbage soup. In western Norway, komle is popular.
A traditional Canary Islands dish is Canarian wrinkly potatoes or papas arrugadas. Tortilla de patatas
(potato omelete) and patatas bravas (a dish of fried potatoes in a spicy tomato sauce) are near-universal
constituent of Spanish tapas.
North America
French fries served with a hamburger

Poutine: Fried potatoes, cheese curds, and gravy
In the United States, potatoes have become one of the most widely consumed crops and thus have a
variety of preparation methods and condiments. French fries and often hash browns are commonly found in
typical American fast-food burger joints and cafeterias. One popular favorite involves a baked potato with
cheddar cheese (or sour cream and chives) on top, and in New England "smashed potatoes" (a chunkier
variation on mashed potatoes, retaining the peel) have great popularity. Potato flakes are popular as an
instant variety of mashed potatoes, which reconstitute into mashed potatoes by adding water, with butter or
oil and salt to taste. A regional dish of Central New York, salt potatoes are bite-size new potatoes boiled in
water saturated with salt then served with melted butter. At more formal dinners, a common practice includes
taking small red potatoes, slicing them, and roasting them in an iron skillet. Among American Jews, the
practice of eating latkes (fried potato pancakes) is common during the festival of Hanukkah.
A traditional Acadian dish from New Brunswick is known as poutine râpée. The Acadian poutine is a
ball of grated and mashed potato, salted, sometimes filled with pork in the center, and boiled. The result is a
moist ball about the size of a baseball. It is commonly eaten with salt and pepper or brown sugar. It is
believed to have originated from the German Klöße, prepared by early German settlers who lived among the
Acadians.
Poutine, by contrast, is a hearty serving of French fries, fresh cheese curds and hot gravy. Tracing its
origins to Quebec in the 1950s, it has become a widespread and popular dish throughout Canada.
Indian Subcontinent
Aloo ki sabzi (Potato Curry) and Puri, a typical morning snack, Varanasi, India
In India, the most popular potato dishes are aloo ki sabzi, batata vada, and samosa, which is spicy
mashed potato mixed with a small amount of vegetable stuffed in conical dough, and deep fried. Potatoes
are also a major ingredient as fast food items, such as aloo chaat, where they are deep fried and served with
chutney. In Northern India, alu dum and alu paratha are a favorite part of the diet; the first is a spicy curry of
boiled potato, the second is a type of stuffed chapati.
A dish called masala dosa from South India is very notable all over India. It is a thin pancake of rice
and pulse paste rolled over spicy smashed potato and eaten with sambhar and chutney. Poori in sound India
in particular in Tamil Nadu is almost always taken with smashed potato masal. Other favorite dishes are alu
tikki and pakoda items.
Vada pav is a popular vegetarian fast food dish in Mumbai and other regions in the Maharashtra in
India.
Aloo posto (a curry with potatoes and poppy seeds) is immensely popular in East India, especially
Bengal.
Far East Asia

In the Far East, rice still dominates the potato, especially in China and Japan. However, it is used in
northern China where rice is not easily grown, with a popular dish being 青椒土豆丝(qing jiao tu dou si), made with
green pepper, vinegar and a thin slices of potato. In the winter, roadside sellers in northern China will also
sell roasted potatoes. It is also occasionally seen in Korean and Thai cuisines.[100]
Art
Potato ceramic from the Moche culture. Larco Museum Collection

The potato has been an essential crop in the Andes since the pre-Columbian Era. The Moche culture
from Northern Peru made ceramics from earth, water, and fire. This pottery was a sacred substance, formed
in significant shapes and used to represent important themes. Potatoes are represented anthropomorphically
as well as naturally.[101]
During the late 19th century, numerous images of potato harvesting appeared in European art,
including the works of Willem Witsen and Anton Mauve.[102] Van Gogh's 1885 painting "The Potato Eaters"
portrays a family eating potatoes.[103]
Invented in 1949 and marketed and sold commercially by Hasbro in 1952, Mr. Potato Head is an
American toy that consists of a plastic potato and attachable plastic parts such as ears and eyes to make a
face. It was the first toy ever advertised on television.[104]
See also
• List of potato museums
• New World crops
• Potato battery
Notes
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http://www.cipotato.org/pressroom/press_releases_detail.asp?cod=17&lang=en Lay summary.
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95.^ Hayes, Monte (24 June 2007). "''Peru Celebrates Potato Diversity''". Washingtonpost.com.
http://www.washingtonpost.com/wp-dyn/content/article/2007/06/24/AR2007062400727.html.
96.^ Timothy Johns: With bitter Herbs They Shall Eat it : Chemical ecology and the origins of
human diet and medicine, The University of Arizona Press, Tucson 1990, ISBN 0-8165-1023-7, p.
82-84
97.^ Sinkovec, Magdalena (2004). "Bryndzové Halušky / Potato Dumplings with 'Bryndza'
Sheep Cheese and Bacon". Culinary Cosmic Top Secrets A Nato Cookbook . Lulu. pp. 115–116.
ISBN 9781411608375. http://books.google.com/books?id=9fNgBO3aEK8C&pg=PA115. Retrieved 1
March 2009.
98.^ von Bremzen, Anya; Welchman, John (1990). Please to the Table: The Russian Cookbook .
New York: Workman Publishing. pp. 319–20. ISBN 0-89480-845-1.
99.^ Roden, Claudia (1990). The Food of Italy. London: Arrow Books. p. 72. ISBN 0-09-976220-
X.
100.^ Solomon, Charmaine (1996). Charmaine Solomon's Encyclopedia of Asian Food.
Melbourne: William Heinemann Australia. p. 293. ISBN 0-85561-688-1.
101.^ Berrin, Katherine & Larco Museum. The Spirit of Ancient Peru:Treasures from the Museo
Arqueológico Rafael Larco Herrera. New York:Thames and Hudson, 1997.
102.^ Steven Adams, Anna Gruetzner Robins (2000). Gendering Landscape Art. University of
Manchester. ISBN 0719056284. http://books.google.com/?id=dY7xwrA-
ibQC&pg=PA67&dq=potatoes++witsen+gathering+potatoes.
103.^ van Tilborgh, Louis (2009). "The Potato Eaters by Vincent van Gogh". The Vincent van
Gogh Gallery. http://www.vggallery.com/visitors/004.htm. Retrieved 11 September 2009.
104.^ "Mr Potato Head". Museum of Childhood website. V&A Museum of Childhood.
http://www.vam.ac.uk/moc/collections/toys/construction_toys/mr_potato_head/index.html. Retrieved
11 September 2009.
References
• Economist. "Llamas and mash", The Economist 28 Feb 2008 online
• Economist. "The potato: Spud we like", (leader) The Economist 28 Feb 2008 online
• Boomgaard, Peter. "In the Shadow of Rice: Roots and Tubers in Indonesian History, 1500–
1950." Agricultural History 2003 77(4): 582–610. Issn: 0002-1482 Fulltext: Ebsco
• Hawkes, J.G. (1990). The Potato: Evolution, Biodiversity & Genetic Resources , Smithsonian
Institution Press, Washington, D.C.
• Lang, James (2001). Notes of a Potato Watcher, Texas A&M University, College Station,
Texas.
• Langer, William L. "American Foods and Europe's Population Growth 1750–1850", Journal of
Social History, Vol. 8, No. 2 (Winter, 1975), pp. 51–66 in JSTOR
• McNeill, William H. "How the Potato Changed the World's History." Social Research 1999
66(1): 67–83. Issn: 0037-783x Fulltext: Ebsco, by a leading historian
• McNeill, William H. "The Introduction of the Potato into Ireland", Journal of Modern History 21
(1948): 218–21. in JSTOR
• Ó Gráda, Cormac. Black '47 and Beyond: The Great Irish Famine in History, Economy, and
Memory. (1999). 272 pp.
1. Ó Gráda, Cormac, Richard Paping, and Eric Vanhaute, eds. When the Potato Failed:
Causes and Effects of the Last European Subsistence Crisis, 1845–1850. (2007). 342 pp. ISBN 978-
2-503-51985-2. 15 essays by scholars looking at Ireland and all of Europe
• Reader, John. Propitious Esculent: The Potato in World History (2008), 315pp a standard
scholarly history
• Salaman, Redcliffe N. (1989). The History and Social Influence of the Potato , Cambridge
University Press (originally published in 1949; reprinted 1985 with new introduction and corrections
by J.G. Hawkes).
• Stevenson, W.R., Loria, R., Franc, G.D., and Weingartner, D.P. (2001) Compendium of
Potato Diseases, 2nd ed, Amer. Phytopathological Society, St. Paul, Minnesota.
• Zuckerman, Larry. The Potato: How the Humble Spud Rescued the Western World. (1998).
304 pp. Douglas & McIntyre. ISBN 0-86547-578-4.
Further reading
• "'Humble' Potato Emerging as World's Next Food Source". column (Asahi Shimbun): p. 20.
11 May 2008.
• Spooner, David; et al. (October 2005). "A single domestication for potato based on multilocus
amplified fragment length polymorphism genotyping". Proc. Natl. Acad. Sci. USA 102 (41): 14694–
14699. doi:10.1073/pnas.0507400102. PMID 16203994. PMC 1253605.
• The World Potato Atlas at Cgiar.org, released by the International Potato Center in 2006 and
regularly updated. Includes current chapters of 15 countries:
• South America: (English and Spanish): Bolivia, Colombia, Ecuador, Peru
• Africa: Cameroon, Ethiopia, Kenya
• Eurasia: Armenia, Bangladesh, China, India, Myanmar, Nepal, Pakistan, Tajikistan
• 38 others as brief "archive" chapters
• Further information links at Cgiar.org.
• World Geography of the Potato at UGA.edu, released in 1993.
• Reference for potato history: The Vegetable Ingredients Cookbook by Christine Ingram,
Lorenz Books, 1996 ISBN 1-85967-264-7
• The History and Social Influence of the Potato by Redcliffe N. Salaman ISBN 0-521-31623-5
• Hamilton, Andy & Dave, (2004), Potatoes: Solanum tuberosums retrieved on 4 May 2005
• Gauldie, Enid (1981). The Scottish Miller 1700–1900. Pub. John Donald. ISBN 0-85976-067-
7.
External links
The Wikibook Gardening has a page on the topic of
Potato
Wikibooks Cookbook has a recipe/module on

Potato
Wikimedia Commons has media related to: Solanum tuberosum
Wikimedia Commons has media related to: Potato
Wikimedia Commons has media related to: Potatoes
Look up potato in Wiktionary, the free dictionary.
• GLKS Potato Database

• Centro Internacional de la Papa: CIP (International Potato Center)
• World Potato Congress
• British Potato Council
• Online Potato Pedigree Database for cultivated varieties
• Potato Information & Exchange
• GMO Safety: Genetic engineering on potatoes Biological safety research projects and results
• International Year of the Potato 2008
• Solanum tuberosum (potato, papas): life cycle, tuber anatomy at GeoChemBio
• Potato Genome Sequencing Consortium
• Potato storage and value Preservation: Pawanexh Kohli, CrossTree techno-visors.
[hide]v · d · eLists of countries and territories by agricultural output rankings
Cereals Barley · Buckwheat · Maize · Millet · Oats · Rice · Rye · Sorghum · Triticale ·
Wheat
Fruit Apples · Apricots · Bananas · Citrus (Oranges) · Plums
Vegetables Artichoke · Onion · Potato · Soybean · Tomatoes
Other Cacao · Cassava · Coffee · Fish · Garlic · Milk · Sugar beet · Sugar cane ·
Sunflower seed · Tea · Tobacco · Wine
Related Irrigation · Land use
Lists of countries and territories · Lists by country · List of international rankings · List of top
international rankings by country
[hide]v · d · ePotato cultivars

Adirondack Blue • Adirondack Red • Agata • Almond • Amandine • Anya •
Désirée • Fingerling • Golden Wonder • Jersey Royal • Kerr's Pink • King Edward •
Marfona • Ratte • Red Pontiac • Rooster • Russet Burbank • Stobrawa • Vitelotte •
Vivaldi • Yellow Finn • Yukon Gold
Retrieved from "http://en.wikipedia.org/wiki/Potato"
Categories: Potatoes | Crops originating from the Americas | Crops originating from Bolivia | Crops
originating from Chile | Crops originating from Peru | Indian ingredients | Puerto Rican ingredients | Quechua
loanwords | Root vegetables | Staple foods
W000
Baleen whale
Baleen whales[1]
Temporal range: latest Eocene – Recent
Humpback Whale breaching
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Cetacea
Mysticeti
Suborder:
Cope, 1891
Families
see text
Baleen
The baleen whales, also called whalebone whales or great whales, form the Mysticeti, one of two
suborders of the Cetacea (whales, dolphins, and porpoises). Baleen whales are characterized by having
baleen plates for filtering food from water, rather than having teeth. This distinguishes them from the other
suborder of cetaceans, the toothed whales or Odontoceti. Living Mysticeti species have teeth only during the
embryonal phase. Fossil Mysticeti had teeth before baleen evolved.
The suborder contains four extant families and fifteen species.
Contents
[hide]
• 1 Etymology
• 2 Anatomy
• 3 Ecology and life history
• 3.1 Behavioral ecology
• 3.1.1 Breaching
• 4 Importance to humans
• 6 Taxonomic classification
• 7 References
[edit] Etymology
The taxonomic name Mysticeti apparently derives from a transmission error in early copies of
Aristotle's Historia Animalium in which "ο μυς το κητος" ("the whale known as 'the mouse' or 'Gutter whale' ")
was mistakenly run together as "ο μυστικητος" ("the Mysticetus").[2] An alternate name for the suborder is
Mystacoceti (from Greek μυσταξ "moustache" + κητος "whale").[3]
[edit] Anatomy
Baleen whales are generally larger than toothed whales, and females are bigger than males. This
group includes the largest known animal species, the Blue Whale.
Humpback whale with two blowholes clearly visible

Baleen whales have two blowholes, causing a V-shaped blow.
[edit] Ecology and life history

[edit] Behavioral ecology
Solitary or in small groups called pods.
[edit] Breaching
In spite of their enormous size, baleen whales are able to leap completely out of the water. They can
grow to 190,000 kilograms (420,000 lb) in weight and 33.5 metres (110 ft) in length.[4] Particularly known for
its acrobatics is the Humpback Whale, but other baleen whales also break through the water surface with
their body or beat it loudly with their fins. Some believe that the male baleen whales try to show off in the
presence of females to increase their mating success. Scientists speculate that baleen whales and other
cetaceans may engage in breaching to dislodge parasites, or scratch irritated skin. Breaching, and other
behaviors like lobtailing, are also used to stun or kill nearby fish or krill.
[edit] Importance to humans

From the 11th to the late 20th centuries, baleen whales were hunted commercially for their oil and
baleen. Their oil was used to make margarine and cooking oils, whilst their baleen was used to stiffen
corsets, as parasol ribs and to crease paper.

See also: Evolution of cetaceans
Parietobalaena palmeri (extinct) skull
Early baleen whales first appeared as far back as the early Oligocene, or perhaps the latest Eocene
(39–29 million years ago; e.g., Llanocetus). Early baleen whales possessed teeth inherited from their
ancestors, as opposed to baleen, in modern species. The Oligocene species Aetiocetus cotylalveus is
considered the evolutionary link between toothed and baleen whales. This species was discovered by fossil
collector Douglas Emlong in 1964 near Seal Rock State Recreation Site, Oregon, in a sandstone formation.
[5] In the early 1990s, the species Janjucetus hunderi was discovered in Victoria, Australia by a surfer and
was described in 2006 by E. M. G. Fitzgerald.[6] Janjucetus was a baleen whale with sharp teeth that hunted
fish and squid as well as larger prey, potentially including sharks and dolphin-like cetaceans. These fossils
hint that early baleen whales were predatory and eventually evolved into the gentler, toothless whales known
today. A recent study identified palatal foramina (bony impressions of blood vessels that "feed" the baleen
racks) in the palate of a toothed mysticete, Aetiocetus weltoni. The scientists involved indicated that this
discovery implies that this whale previously possessed both teeth and baleen, and serves as an intermediate
adaptive role between primitive toothed mysticetes and more advanced toothless mysticetes.[7] The first
baleen-bearing, toothless baleen whales (such as Eomysticetus and Micromysticetus) appeared in the late
Oligocene.[8] Early baleen whales probably could not echolocate; no anatomical evidence preserved in the
skulls and ear regions of any fossil baleen whales show any of the adaptations associated with echolocation
as in toothed whales.[6]
[edit] Taxonomic classification

The "†"'s denote extinct families and genera.
Suborder Mysticeti: Baleen whales
• Family †Aetiocetidae
• † Aetiocetus
• † Ashorocetus
• † Chonecetus
• † Morawanocetus
• † Willungacetus
• Family †Aglaocetidae
• † Aglaocetus
• † Isanacetus
• † Pinocetus
• Family Balaenidae: Right whales and Bowhead Whale
• Balaena Bowhead whales
• †Balaenella
• †Balaenotus
• †Balaenula
• Eubalaena
• Eucetites
• †Morenocetus
• Family Balaenopteridae: Rorquals[9]
• †Archaebalaenoptera
• Balaenoptera
• †Cetotheriophanes
• †Diunatans
• †Mauicetus
• Megaptera
• †Notiocetus
• †Parabalaenoptera
• †Plesiobalaenoptera
• †Praemegaptera
• †Protororqualus
• †Family Cetotheriidae
• †Cephalotropis
• †Cetotherium
• †Herpetocetus
• †Hibacetus
• †Joumocetus
• †Metopocetus
• †Mixocetus
• †Nannocetus
• †Palaeobalaena
• †Piscobalaena
• †Plesiocetopsis
• †Titanocetus
• †Family Cetotheriopsidae
• †Cetotheriopsis
• †Micromysticetus
• †Family Diorocetidae
• †Amphicetus
• †Diorocetus
• †Plesiocetus
• †Thinocetus
• †Uranocetus
• †Family Eomysticetidae
• †Eomysticetus
• Family Eschrichtiidae
• †Archaeschrichtius
• †Eschrichtioides
• Eschrichtius Grey whales
• †Gricetoides
• †Megapteropsis
• †Family Llanocetidae
• †Llanocetus
• †Family Mammalodontidae
• †Janjucetus
• †Mammalodon
• Family Neobalaenidae: Pygmy Right Whale
• Caperea, Pygmy Right Whale
• †Family Pelocetidae
• †Cophocetus
• †Halicetus
• †Parietobalaena
• †Pelocetus
• †Eobalaenoptera
• Family incertae sedis
• †Amphitera
• †Burtinopsis
• †Idiocetus
• †Imerocetus
• †Isocetus
• †Mesocetus
• †Mioceta
• †Otradnocetus
• †Peripolocetus
• †Piscocetus
• †Siphonocetus
• †Tiphyocetus
• †Tretulias
• †Ulias
The earliest-known baleen whale is Llanocetus, discovered on Seymour Island, Antarctica, by E.D.
Mitchell in 1989. The species lived during the Latest Eocene/Earliest Miocene, about 35 mya.
[edit] References
1. ^ Mead, James G.; Brownell, Robert L., Jr. (16 November 2005). "Order Cetacea (pp. 723-
id=14300002.
2. ^ OED 'mysticete' (n, 1)
3. ^ OED 'mystacocete'
4. ^ Dewey, T.; Fox, D. (2002). "Balaenoptera musculus (On-line)". Animal Diversity Web.
http://animaldiversity.ummz.umich.edu/site/accounts/information/Balaenoptera_musculus.html.
5. ^ Wallace, D. R. (2007). Neptune's Ark: From Ichthyosaurs to Orcas. Berkeley ; London:
University of California Press. ISBN 0-520-24322-6.
6. ^ a b Fitzgerald, E. M. G. (2006). "A bizarre new toothed mysticete (Cetacea) from Australia
and the early evolution of baleen whales". Proceedings of the Royal Society - 'B': Biological
Sciences, 273 (1604): 2955–2963. doi:10.1098/rspb.2006.3664. PMID 17015308.
7. ^ Deméré, T., McGowen, M., Berta, A., Gatesy, J. (2008). Morphological and Molecular
Evidence for a Stepwise Evolutionary Transition from Teeth to Baleen in Mysticete Whales.
Systematic Biology, 57(1), 15-37.
8. ^ A. E. Sanders and L. G. Barnes. 2002. Paleontology of the Late Oligocene Ashley and
Chandler Bridge Formations of South Carolina, 3: Eomysticetidae, a new family of primitive
mysticetes (Mammalia: Cetacea). Smithsonian Contributions to Paleobiology 93:313-356.
9. ^ Deméré, T.A.; Berta, A.; McGowen, M.R. (2005). "The taxonomic and evolutionary history
of fossil and modern balaenopteroid mysticetes". Journal of Mammalian Evolution 12 (1/2): 99–143.
doi:10.1007/s10914-005-6944-3.
Cetaceans portal
[hide]v · d · eExtant Cetacea species

Laurasiatheria · (unranked) Cetartiodactyla · (unranked) Whippomorpha
[hide] Suborder Mysticeti (Baleen whales)
Balaena Bowhead whale (B. mysticetus)
Balaenidae Southern right whale (E. australis) · North Atlantic

Eubalaena
right whale (E. glacialis) · North Pacific right whale (E.
(Right whales)
japonica)
Common minke whale (B. acutorostrata) ·

Antarctic minke whale (B. bonaerensis) · Sei whale (B.
Balaenopteridae Balaenoptera borealis) · Bryde's whale (B. brydei) · Pygmy Bryde's
(Rorquals) whale (B. edeni) · Blue whale (B. musculus) · B. omurai ·
Fin whale (B. physalus)
Megaptera Humpback whale (M. novaeangliae)

Eschrichtiidae Gray whale (E.
Eschrichtius
robustus)
Neobalaenidae Pygmy right whale (C.

Caperea
marginata)
[hide] Suborder Odontoceti (Toothed whales) (cont. below)
Delphinidae Commerson's dolphin (C. commersonii) ·

(Oceanic dolphins) Cephalorhynchus Chilean dolphin (C. eutropia) · Haviside's dolphin
(C. heavisidii) · Hector's dolphin (C. hectori)
Long-beaked common dolphin (D.

Delphinus capensis) · Short-beaked common dolphin (D.
delphis)
Feresa Pygmy killer whale (F. attenuata)
Globicephala Short-finned pilot whale (G.

(Pilot whales) macrorhynchus) · Long-finned pilot whale (G.
melas)
Grampus Risso's dolphin (G. griseus)
Lagenodelphis Fraser's dolphin (L. hosei)
Atlantic white-sided dolphin (L. acutus) ·

White-beaked dolphin (L. albirostris) · Peale's
Lagenorhynchus dolphin (L. australis) · Hourglass dolphin (L.
cruciger) · Pacific white-sided dolphin (L.
obliquidens) · Dusky dolphin (L. obscurus)
Lissodelphis Northern right whale dolphin (L. borealis) ·

(Right whale dolphins) Southern right whale dolphin (L. peronii)
Irrawaddy dolphin (O. brevirostris) ·

Orcaella
Australian snubfin dolphin (O. heinsohni)
Orcinus Killer whale (O. orca)
Peponocephala Melon-headed whale (P. electra)

Pseudorca False killer whale (P. crassidens)
Tucuxi (S. fluviatilis) · Costero (S.

Sotalia
guianensis)
Pacific humpback dolphin (S. chinensis) ·

Sousa Indian humpback dolphin (S. plumbea) · Atlantic
humpback dolphin (S. teuszii)
Pantropical spotted dolphin (S. attenuata) ·

Clymene dolphin (S. clymene) · Striped dolphin (S.
Stenella
coeruleoalba) · Atlantic spotted dolphin (S.
frontalis) · Spinner dolphin (S. longirostris)
Steno Rough-toothed dolphin (S. bredanensis)
Indo-Pacific bottlenose dolphin (T.

Tursiops aduncus) · Common bottlenose dolphin (T.
truncatus)
[hide] Suborder Odontoceti (Toothed whales) (cont. above)
Delphinapterus Beluga (D. leucas)

Monodontidae
Monodon Narwhal (M. monoceros)
Neophocaena Finless porpoise (N. phocaeniodes)
Phocoenidae Spectacled porpoise (P. dioptrica) · Harbor

(Porpoises) Phocoena porpoise (P. phocoena) · Vaquita (P. sinus) · Burmeister's
porpoise (P. spinipinnis)
Phocoenoides Dall's porpoise (P. dalli)
Physeteridae Sperm whale (P.

Physeter
macrocephalus)
Kogiidae Pygmy sperm whale (K. breviceps) · Dwarf sperm whale

Kogia
(K. simus)
Iniidae Amazon river dolphin (I.
Inia
geoffrensis)
Lipotidae Baiji (L.

Lipotes
vexillifer)
Platanistidae Ganges and Indus River dolphin (P.

Platanista
gangetica)
Pontoporiidae La Plata dolphin (P.

Pontoporia
blainvillei)
Ziphiidae Arnoux's beaked shale (B. arnuxii) · Baird's beaked

(Beaked whales) Berardius
whale (B. bairdii)
Northern bottlenose whale (H. ampullatus) ·

Hyperoodon
Southern bottlenose whale (H. planifrons)
Indopacetus Tropical bottlenose whale (I. pacificus)
Mesoplodon Sowerby's beaked whale (M. bidens) · Andrew's

beaked whale (M. bowdoini) · Hubbs' beaked whale (M.
carlhubbsi) · Blainville's beaked whale (M. densirostris) ·
Gervais' beaked whale (M. europaeus) · Ginkgo-toothed
beaked whale (M. ginkgodens) · Gray's beaked whale (M.
(Mesoplodont
grayi) · Hector's beaked whale (M. hectori) · Strap-toothed
whales)
whale (M. layardii) · True's beaked whale (M. mirus) ·
Perrin's beaked whale (M. perrini) · Pygmy beaked whale
(M. peruvianus) · Stejneger's beaked whale (M.
stejnegeri) · Spade-toothed whale (M. traversii)
Tasmacetus Shepherd's beaked whale (T. sheperdi)
Ziphius Cuvier's beaked whale (Z. cavirostris)
Retrieved from "http://en.wikipedia.org/wiki/Baleen_whale"
Categories: Baleen whales

W000
Evolution of reptiles
A fossil of a Casineria. The earliest evidence of amniotes, although an amphibian, it laid eggs on land
as reptiles do.
Further information: Evolutionary history of life
Reptiles arose about 310-320 million years ago during the Carboniferous period. Reptiles are defined
as animals that have scales, lay land based hard shelled eggs, and possessing cold-blooded metabolisms.
Today reptiles generally rank low in most ecological food chains, containing very few apex predators, but
many examples of apex reptiles have existed in the past. Although reptiles have lower specie variation than
in the past, they have an extremely diverse evolutionary history that has led to biological successes such as
dinosaurs, mammals, and birds.
Contents
[hide]
• 1 First reptiles
• 1.1 Rise from water
• 1.2 Early reptiles
• 1.3 Anapsids, synapsids, diapsids and sauropsids
• 1.4 Mammalian evolution
• 2 Rise of dinosaurs
• 2.1 Permian reptiles
• 2.2 The Mesozoic era, the "Age of Reptiles"
• 2.3 Bird evolution
• 2.4 Demise of the dinosaurs
• 2.5 Role reversal
• 3 The four orders of reptilia
• 3.1 Testudines
• 3.2 Sphenodontia
• 3.3 Squamata
• 3.4 Crocodilia
• 4 References
[edit] First reptiles
[edit] Rise from water
Reptiles (green field) are a paraphyletic group comprising all non-avian and non-mammalian
amniotes.
Reptiles first arose from amphibians in the swamps of the late Carboniferous. Increasing evolutionary
pressure and the vast untouched niches of the land powered the evolutionary changes in amphibians to
gradually become more and more land based. Environmental selection propelled the development of certain
traits, such as a stronger skeletal structure, muscles, and more protective coating (scales) became more
favorable, the basic foundation of reptiles were founded. The evolution of lungs and legs are the main
transitional steps towards reptiles, but the development of hard-shelled external eggs replacing the
amphibious water bound eggs is the defining feature of the class reptilia (with the exception of certain
squamates) and is what allowed amphibians to fully leave water. Another major difference from amphibians
is the increased brain size, more specifically, the enlarged cerebrum and cerebellum. Although there brain
size is small when compared to mammals and birds, these enhancements prove vital in hunting strategies of
reptiles. The increased size of these two regions of the brain allowed for improved motor skills and an
increase in sensory development.
[edit] Early reptiles
An early reptile Hylonomus

Mesozoic scene showing typical reptilian megafauna: the dinosaurs Europasaurus holgeri and
Iguanodon, and the early bird Archaeopteryx perched on the foreground tree stump.
The origin of the reptiles lies about 320–310 million years ago, in the steaming swamps of the late
Carboniferous period, when the first reptiles evolved from advanced reptiliomorph labyrinthodonts.[1] The
oldest known animal that may have been an amniote, i.e. a primitive reptile rather than an advanced
amphibian is Casineria.[2][3] A series of footprints from the fossil strata of Nova Scotia, dated to 315 million
years show typical reptilian toes and imprints of scales.[4] The tracks are attributed to Hylonomus, the oldest
unquestionable reptile known.[5] It was a small, lizard-like animal, about 20 to 30 cm (8–12 in) long, with
numerous sharp teeth indicating an insectivorous diet.[6] Other examples include Westlothiana (for the
moment considered a reptiliomorph amphibian rather than a true amniote)[7] and Paleothyris, both of similar
build and presumably similar habit. One of the best known early reptiles is Mesosaurus, a genus from the
early Permian that had returned to water, feeding on fish. The earliest reptiles were largely overshadowed by
bigger labyrinthodont amphibians such as Cochleosaurus, and remained a small, inconspicuous part of the
fauna until after the small ice age at the end of the Carboniferous.
[edit] Anapsids, synapsids, diapsids and sauropsids
A = Anapsid, B = Synapsid, C = Diapsid
The first reptiles were anapsids, having a solid skull with holes for only nose, eyes, spinal cord, etc.
[8] Very soon after the first reptiles appeared, they split into two branches.[9] One branch, Synapsida
(including both "mammal-like reptiles" and modern, extant mammals), had one opening in the skull roof
behind each eye; the other branch, Diapsida, possessed a hole in their skulls behind each eye, along with a
second hole located higher on the skull. The function of the holes in both groups was to lighten the skull and
give room for the jaw muscles to move, allowing for a more powerful bite.[8] Diapsids and later anapsids are
classed as the "true reptiles", Sauropsida.
Turtles have been traditionally believed to be surviving anapsids, on the basis of their skull structure.
[10] The rationale for this classification was disputed, with some arguing that turtles are diapsids that reverted
to this primitive state in order to improve their armor (see Parareptilia).[1] Later morphological phylogenetic
studies with this in mind placed turtles firmly within Diapsida.[11] All molecular studies have strongly upheld
the placement of turtles within diapsids, most commonly as a sister group to extant archosaurs.[12][13][14]
[15]
[edit] Mammalian evolution

Main article: Evolution of mammals
Tetrapods
Amphibians
Amniotes
Sauropsids
Synapsids
Pelycosaurs
Therapsids
Mammals
A basic cladogram of the origin of modern mammals from their closest reptilian relatives.
The main points to the transition from reptile to mammal was the evolution from scales to hair or fur,
the evolution of warm-bloodedness, the loss of external eggs (except for in monotremes who have retained
this trait), and the evolution mammary glands (the most defining trait in mammals that allow them to produce
milk for offspring). The evolution of mammals was a gradual process that took approximately 70 million
years, beginning in the mid-Permian. By the mid-Triassic, there were many species that looked like modern
mammals, and the first true mammals appeared in the early Jurassic. The earliest known marsupial,
Sinodelphys, appeared 125 million years ago in the early Cretaceous, around the same time as Eomaia, the
first known eutherian (member of placentals' "parent" group); and the earliest known monotreme,
Teinolophos, appeared two million years later.
[edit] Rise of dinosaurs

[edit] Permian reptiles

With the close of the Carboniferous, reptiles became the dominant tetrapod fauna. While the
terrestrial reptiliomorph labyrinthodonts still existed, the synapsids evolved the first truly terrestrial
megafauna (giant animals) in the form of pelycosaurs such as Edaphosaurus and the carnivorous
Dimetrodon. In the mid-Permian period the climate turned dryer, resulting in a change of fauna: The primitive
pelycosaurs were replaced by the more advanced therapsids.[16]
The anapsid reptiles, whose massive skull roofs had no postorbital holes, continued and flourished
throughout the Permian. The pareiasaurs reached giant proportions in the late Permian, eventually
disappearing at the close of the period (the turtles being possible survivors).[16]
Early in the period, the diapsid reptiles split into two main lineages, the archosaurs (forefathers of
crocodiles and dinosaurs) and the lepidosaurs (predecessors of modern snakes, lizards, and tuataras). Both
groups remained lizard-like and relatively small and inconspicuous during the Permian.
[edit] The Mesozoic era, the "Age of Reptiles"

The close of the Permian saw the greatest mass extinction known (see the Permian–Triassic
extinction event). Most of the earlier anapsid/synapsid megafauna disappeared, being replaced by the
archosauromorph diapsids. The archosaurs were characterized by elongated hind legs and an erect pose,
the early forms looking somewhat like long-legged crocodiles. The archosaurs became the dominant group
during the Triassic period, developing into the well-known dinosaurs and pterosaurs, as well as crocodiles
and phytosaurs. Some of the dinosaurs developed into the largest land animals ever to have lived, making
the Mesozoic era popularly known as the "Age of Reptiles". The dinosaurs also developed smaller forms,
including the feather-bearing smaller theropods. In the mid-Jurassic period, these gave rise to the first birds.
[16]
The lepidosauromorph diapsids may have been ancestral to the sea reptiles.[17] These reptiles
developed into the sauropterygians in the early Triassic and the ichthyosaurs during the Middle Triassic. The
mosasaurs also evolved in the Mesozoic era, emerging during the Cretaceous period.
The therapsids came under increasing pressure from the dinosaurs in the early Mesozoic and
developed into increasingly smaller and more nocturnal forms, the first mammals being the only survivors of
the line by the late Jurassic.
[edit] Bird evolution

Main article: Evolution of birds
An Archaeopteryx specimen in Berlin.
The main points to the transition from reptile to bird is the evolution from scales to feathers, the
evolution of the beak (although independently evolved in other organisms), the hallofication of bones,
development of flight, and warm-bloodedness.
The evolution of birds is thought to have begun in the Jurassic Period, with the earliest birds derived
from theropod dinosaurs. Birds are categorized as a biological class, Aves. The earliest known species Aves
is Archaeopteryx lithographica, from the Late Jurassic period, though 'Archaeopteryx is not commonly
considered to have been a true bird, but a transitional species between Aves and Reptiles. Modern
phylogenetics place birds in the dinosaur clade Theropoda. According to the current consensus, Aves and a
sister group of the order Crocodilia, together are the sole living members of an unranked "reptile" clade, the
Archosauria.
Coelurosaurs
Alvarezs
Therizinosauroids
Oviraptosaurs
Archaeopteryx
and Rahonavis
(birds)
Troodontids
Deinonychosaurs
Confuciusornis
(bird)
Microraptor
(dromaeosaur)
Dromaeosaurs
Simplified cladogram from Mayr et al. (2005)
Groups usually regarded as birds are in bold type.
[edit] Demise of the dinosaurs

Main article: Cretaceous–Tertiary extinction event
The close of the Cretaceous period saw the demise of the Mesozoic era reptilian megafauna. Along
with massive amount of volcanic activity at the time, the meteor impact that created the K-T boundary is
accepted as the main cause for this mass extinction event. Of the large marine reptiles, only sea turtles are
left, and, of the dinosaurs, only the small feathered theropods survived in the form of birds. The end of the
“Age of Reptiles” led into the “Age of Mammals”. Despite the change in phrasing, reptile diversification
continued throughout the Cenozoic, with squamates undergoing a greater radiation than they did in the
Mesozoic. Today squamates make up the majority of extant reptiles today (over 90%), filling many of the
ecological niches left by the mammals who would take advantage of the niches left by the dinosaurs.[18]
There are approximately 8,700 extant species of reptiles,[18] compared with 5,400 species of mammals.
[edit] Role reversal

After the Cretaceous-Tertiary extinction wiped out all of the non-avian dinosaurs (birds are generally
regarded as the surviving dinosaurs) and several other mammalian groups, placental and marsupial
mammals diversified into many new forms and ecological niches throughout the Tertiary era. Some reaching
enormous sizes and almost as wide of a variation as the dinosaurs once did. Due to global climate changes
such as a lowering in total oxygen levels and lower average temperatures, mammalian megafauna never
quite reached the sky-scraper heights as did some sauropods. These evolutionary and climatic changes is
what is responsible for all modern orders of mammals that are now on Earth, including humans.
[edit] The four orders of reptilia

[edit] Testudines
Main article: Testudines
Testudines, or turtles, may have evolved from anaspids, but their exact origin is unknown and heavily
debated. Fossils date back to around 220 million years ago and share remarkably similar characteristics.
These first turtles retain the same body plan as do all modern testudines and are mostly herbivorous. With
some feeding exclusively on small marine organisms. The trade-mark shell is believed to have evolved from
extensions from there backbone and widened ribs which fused together. This is supported by the fossil of
Odontochelys semitestacea, which has an incomplete shell originating from the ribs and back bone. This
species also had teeth with its beak, giving more support to it being a transitional fossil, although this claim is
still controversial. This shell evolved to protect itself from predators, but also slows down the land based
species by a great amount. This has caused many species to go extinct in recent times. Because of alien
species out-competing the slow order for food and lack of ability to escape from humans, there are large
amounts of endangered species in this order.
[edit] Sphenodontia
Main article: Sphenodontia
Sphenodontians arose in the mid Triassic and only consist of one genus tuatara, which only includes
two endangered species on New Zealand and some other minor surrounding islands. Their evolutionary
history is filled with many species. Recent paleogenetic discoveries show that tuataras are prone to quick
speciation.
[edit] Squamata
Main article: Squamata
The most recent order of reptiles, squamates are recognized by having a movable quadrate bone
(giving them upper-jaw movement) and possessing horny scales. They originate from the early Jurassic and
are made up of the three suborders Lacertilia, Serpentes, and Amphisbaenia. Although they are the most
recent order, squamates contain more species than any of the other reptilian orders. Squamates are a
monophyletic group that is a sister group to the Sphenodontia or tuataras. The squamates and tuatara
together are a sister group to crocodiles and birds, the extant archosaurs. Although squamate fossils first
appear in the early Jurassic, mitochondrial phylogenetics suggests that they evolved in the late Permian.
Most evolutionary relationships within the squamates are not yet completely worked out, with the relationship
of snakes to other groups being most problematic. From morphological data, Iguanid lizards have been
thought to have diverged from other squamates very early, but recent molecular phylogenies, both from
mitochondrial and nuclear DNA, do not support this early divergence.[19] Because snakes have a faster
molecular clock than other squamates,[19] and there are few early snake and snake ancestor fossils,[20] it is
difficult to resolve the relationship between snakes and other squamate groups.
[edit] Crocodilia
Main article: Crocodilia
The first organisms that showed similar characteristics of Crocodilians would be the Crurotarsi who
appeared during the early Triassic 250 million years ago. This quickly gave rise to the Eusuchia clade 220
million years ago which would eventually lead to the order of Crocodilians, first of which arose about 85
million years ago during the late cretaceous, originating from the Eusuchia clade. The earliest fossil evidence
of eusuchians is of the species Isisfordia. Early species mainly feed on fish and vegetation. They were land-
based, most having long legs (when compared to modern crocodiles) and many were bipeadal. As
diversification increased, many apex predators arose, all of which is now extinct. Modern Crocodilia arose
through specific evolutionary traits. The complete loss of bipedalism in trade for generally low quadrupedal
stance for an easy and less noticeable entrance to bodies of water. The shape of the skull/jaw changed to
allow further grasp along with up-ward pointing nostrils and eyes. Mimicry is evident, as the back of all
crocodilia resemble some type floating log along with their general color scheme of brown and green to
mimic moss or wood. There tail also took on a paddle shape to increase swimming speed. The only
remaining groups to this order is the alligator, crocodile, caiman, and gharial.
[edit] References
1. ^ a b Laurin, M.; Reisz, R. R. (1995). "A reevaluation of early amniote phylogeny". Zoological
Journal of the Linnean Society 113: 165–223. doi:10.1111/j.1096-3642.1995.tb00932.x. (abstract)
2. ^ R. L. Paton, T. R. Smithson and J. A. Clack, "An amniote-like skeleton from the Early
Carboniferous of Scotland", (abstract), [[Nature (journal)|]] 398, 508-513 (8 April 1999)
3. ^ Monastersky, R. (1999): Out of the Swamps, How early vertebrates established a foothold—
with all 10 toes—on land. Science News, Vol. 155, No. 21, p. 328. article
4. ^ Falcon-Lang, H.J., Benton, M.J. & Stimson, M. (2007): Ecology of early reptiles inferred
from Lower Pennsylvanian trackways. Journal of the Geological Society, London, 164; no. 6; pp
1113-1118. article
5. ^ "Earliest Evidence For Reptiles". Sflorg.com. 2007-10-17.
http://www.sflorg.com/sciencenews/scn101707_01.html. Retrieved 2010-03-16.
6. ^ Palmer, D., ed (1999). The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric
Animals. London: Marshall Editions. p. 62. ISBN 1-84028-152-9.
7. ^ Ruta, M.; Coates, M.I. & Quicke, D.L.J. (2003): Early tetrapod relationships revisited.
Biological Review no 78: pp 251-345.PDF
8. ^ a b Romer, A.S. & T.S. Parsons. 1977. The Vertebrate Body. 5th ed. Saunders,
Philadelphia. (6th ed. 1985)
9. ^ van Tuninen, M. & Hadly, E.A. (2004): Error in Estimation of Rate and Time Inferred from
the Early Amniote Fossil Record and Avian Molecular Clocks. Journal of Mulecular Biology, no 59: pp
267-276 PDF
10.^ Benton, M. J. (2000). Vertebrate Paleontology (2nd ed.). London: Blackwell Science Ltd.
ISBN 0632056142. , 3rd ed. 2004 ISBN 0-632-05637-1
11.^ Rieppel O, DeBraga M (1996). "Turtles as diapsid reptiles". Nature 384: 453–5.
doi:10.1038/384453a0.
12.^ Zardoya, R.; Meyer, A. (1998). "Complete mitochondrial genome suggests diapsid affinities
of turtles". Proc Natl Acad Sci U S A 95 (24): 14226–14231. doi:10.1073/pnas.95.24.14226.
ISSN 0027-8424. PMID 9826682. PMC 24355. http://www.pubmedcentral.gov/articlerender.fcgi?
artid=24355.
13.^ Iwabe, N.; Hara, Y.; Kumazawa, Y.; Shibamoto, K.; Saito, Y.; Miyata, T.; Katoh, K. (2004-
12-29). "Sister group relationship of turtles to the bird-crocodilian clade revealed by nuclear DNA-
coded proteins". Molecular Biology and Evolution 22 (4): 810–813. doi:10.1093/molbev/msi075.
PMID 15625185. http://mbe.oxfordjournals.org/content/22/4/810.long. Retrieved 2010-12-31.
14.^ Roos, Jonas; Aggarwal, Ramesh K.; Janke, Axel (Nov. 2007). "Extended mitogenomic
phylogenetic analyses yield new insight into crocodylian evolution and their survival of the
Cretaceous–Tertiary boundary". Molecular Phylogenetics and Evolution 45 (2): 663–673.
doi:10.1016/j.ympev.2007.06.018. PMID 17719245.
15.^ Katsu, Y.; Braun, E. L.; Guillette, L. J. Jr.; Iguchi, T. (2010-03-17). "From reptilian
phylogenomics to reptilian genomes: analyses of c-Jun and DJ-1 proto-oncogenes". Cytogenetic and
Genome Research 127 (2-4): 79–93. doi:10.1159/000297715. PMID 20234127.
16.^ a b c Colbert, E.H. & Morales, M. (2001): Colbert's Evolution of the Vertebrates: A History of
the Backboned Animals Through Time. 4th edition. John Wiley & Sons, Inc, New York — ISBN 978-0-
471-38461-8.
17.^ Gauthier J. A. (1994): The diversification of the amniotes. In: D. R. Prothero and R. M.
Schoch (ed.) Major Features of Vertebrate Evolution: 129-159. Knoxville, Tennessee: The
Paleontological Society.
18.^ a b "The Reptile Database". http://www.reptile-database.org/db-info/SpeciesStat.html.
Retrieved 19 August 2010.
19.^ a b Kumazawa, Yoshinori (2007). "Mitochondrial genomes from major lizard families
suggest their phylogenetic relationships and ancient radiations". Gene 388 (1-2): 19–26.
doi:10.1016/j.gene.2006.09.026. PMID 17118581.
20.^ "Lizards & Snakes Alive!". American Museum of Natural History.
http://www.amnh.org/exhibitions/lizards/snakes/world.php. Retrieved 2007-12-25.
[hide]v · d · eExtant reptile orders by subclass
Kingdom: Animalia · Phylum: Chordata · Subphylum: Vertebrata
Anapsida Chelonia Testudines
Lepidosauromorpha Sphenodontia · Squamata

Diapsida
Archosauromorpha Crocodilia
Processes of
Population genetic
Evolutionary
developmental
concepts

Modes of

Retrieved from "http://en.wikipedia.org/wiki/Evolution_of_reptiles"
Categories: Evolution | Evolution by taxon | Reptiles | Herpetology | Paleontology | Prehistoric

reptiles
W000
Charles Sibley

Charles Gald Sibley (August 7, 1917 – April 12, 1998) was an American ornithologist and molecular
biologist. He had an immense influence on the scientific classification of birds, and the work that Sibley
initiated has substantially altered our understanding of the evolutionary history of modern birds.
Sibley's taxonomy has been a major influence on the sequences adoped by ornithological
organizations, especially the American Ornithologists' Union.
Charles Sibley is of no known family relation to renowned bird artist David Sibley.
Contents
[hide]
• 1 Life and work
• 2 Acid tongue
• 3 Other ornithological Sibleys
• 4 See also
• 5 References
[edit] Life and work

Educated in California (A.B. 1940; Ph.D. 1948 in Zoology, University of California, Berkeley. Minor
fields: Paleontology, Botany), he did his first fieldwork in Mexico in 1939 and 1941, then in Solomon Islands,
Bismarck Archipelago, New Guinea, and the Philippines during World War II while on leave from the U.S.
Navy, in which he was Ensign to Lieutenant in the Communications and Medical Service Corps. He was
based for much of the war at Emirau Island, in what is now New Ireland Province of Papua New Guinea. His
first job after college was from 1948-1949 as Instructor in Zoology and Curator of Birds, University of Kansas,
followed from 1949-1953 as Assistant Professor of Zoology, San Jose State College, California. From 1953-
1965 he was Associate Professor then Professor of Zoology and director of the ornithological laboratory at
Cornell. From 1965–1986 he was Professor of Biology and William Robertson Coe Prof. of Ornithology, Dept
of Biology; and Curator of Birds at the Peabody Museum at Yale. From 1986–1992 he was Dean's Professor
of Science and Professor of Biology at San Francisco State University, and from 1993 until his death he was
Adjunct Professor of Biology at Sonoma State University.
Sibley developed an interest in hybridisation and its implications for evolution and taxonomy and, in
the early 1960s he began to focus on molecular studies: of blood proteins, and then the electrophoresis of
egg-white proteins.
By the early 1970s Sibley was pioneering DNA-DNA hybridisation studies, with the aim of
discovering, once and for all, the true relationships between the modern orders of birds. These were highly
controversial to begin with, and regarded by colleagues as anything from snake-oil salesmanship on the one
hand to Holy Writ on the other (Ahlquist, 1999). With the passage of time and ever-improving laboratory
methods, the balance of scientific opinion has shifted closer to the latter interpretation, though the picture is
by no means clear-cut and simple. Some of Sibley's results – such as the close relationship of galliform birds
and waterfowl and their distinctness from other neognaths – have been verified. Other results such as the
inclusion of diverse groups into the Ciconiiformes have turned out to be very much in error.
Sibley became estranged from his American co-workers for a time and corresponded with overseas
colleagues extensively. But by the mid to late 1980s, Sibley's ongoing work had reversed the trend. His
revised phylogeny of living birds in the light of DNA analysis, published in various forms from 1986–1993 was
both controversial and highly influential.
In 1986 he was elected a Member, National Academy of Sciences, U.S.A. In 1990 Sibley was elected
President of the International Ornithological Congress. His landmark publications, Phylogeny and
Classification of Birds (written with Jon Ahlquist) and Distribution and Taxonomy of Birds of the World (with
Burt Monroe) are among the most-cited of all ornithological works, the former setting out the influential
Sibley-Ahlquist taxonomy.
[edit] Acid tongue
During the 1970s, Sibley was a highly controversial figure in ornithological circles, for both
professional and personal reasons. His friend Richard Schodde, writing Sibley's obituary in Emu, commented
that he was:
... a rebel with a cause. In argument he would bulldoze through, brooking no contradiction.
Critics were baited with an acid tongue, and, in fits of temper, he could be a cruel mimic. In
short, lesser mortals were not tolerated easily and, as has been said by others, collegiate friends
were few. ... I never found him malicious or vindictive, even against those who had tried to bring
him down. Nor was he particularly sophisticated or cultured, just a big, up-front Yank possessed
by 'the big picture' in avian phylogeny and convinced of the righteousness of his cause and
invincibility of his intellect. (Schodde 2000)
Partly due to personality conflicts, Sibley had few long-term collaborations with other scientists, with
the notable exception of Jon Ahlquist. However, he was effective in persuading others to provide him with the
blood, tissue, and egg white samples which were the key to his work.(Ahlquist 1999, p. 856)
[edit] Other ornithological Sibleys

Charles Gald Sibley is of no known family relation to renowned bird artist David Sibley, although the
families knew each other: one of Charles' daughters babysat David, and David's father Fred worked for
Charles at Yale.
There is some family resemblance, and Charles did a fair amount of genealogical research but could
only establish that any relationship was no closer than fourth cousin.
[edit] See also
• Sibley-Ahlquist taxonomy
[edit] References
• Ahlquist, Jon E., 1999: Charles G. Sibley: A commentary on 30 years of collaboration. The
Auk, vol. 116, no. 3 (July 1999). A PDF or DjVu version of this article can be downloaded from the
issue's table of contents page.
• Schodde, Richard (2000): Obituary: Charles G. Sibley, 1911-1998. Emu 100(1): 75-76.
doi:10.1071/MU00903
• National Academy of Sciences bio
Persondata
Name Sibley, Charles
Alternative names
Short description
Date of birth
Place of birth
Date of death
Place of death
Retrieved from "http://en.wikipedia.org/wiki/Charles_Sibley"
Categories: 1917 births | 1998 deaths | American ornithologists | Cornell University faculty | Sonoma
State University faculty
W000
Ring-tailed Lemur

Ring-tailed Lemur[1]
Conservation status
Near Threatened (IUCN 3.1)[2]

CITES Appendix I
Threatened with extinction (CITES)[3]
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Family: Lemuridae
Lemur
Genus:
Linnaeus, 1758
Species: L. catta
Binomial name
Lemur catta
Linnaeus, 1758
Distribution of Lemur catta[4]
Synonyms
Genus:[5][6]
• Prosimia Brisson, 1762[N 1]
• Procebus Storr, 1780
• Catta Link, 1806[N 2]
• Maki Muirhead, 1819[N 3]
• Mococo Trouessart, 1878[N 4]
• Odorlemur Bolwig, 1960
Species:[5][6]
• Maki mococo Muirhead,
1819[N 5]
The Ring-tailed Lemur (Lemur catta) is a large strepsirrhine primate and the most recognized lemur
due to its long, black and white ringed tail. It belongs to Lemuridae, one of four lemur families. It is the only
member of the Lemur genus. Like all lemurs it is endemic to the island of Madagascar. Known locally in
Malagasy as hira or maky (spelled maki in French), it inhabits gallery forests to spiny scrub in the southern
regions of the island. It is omnivorous and the most terrestrial of lemurs. The animal is diurnal, being active
exclusively in daylight hours.
The Ring-tailed Lemur is highly social, living in groups of up to 30 individuals. It is also female
dominant, a trait common among lemurs. To keep warm and reaffirm social bonds, groups will huddle
together forming a lemur ball. The Ring-tailed Lemur will also sunbathe, sitting upright facing its underside,
with its thinner white fur towards the sun. Like other lemurs, this species relies strongly on its sense of smell
and marks its territory with scent glands. The males perform a unique scent marking behavior called spur
marking and will participate in stink fights by impregnating their tail with their scent and wafting it at
opponents.
As one of the most vocal primates, the Ring-tailed Lemur utilizes numerous vocalizations including
group cohesion and alarm calls. Experiments have shown that the Ring-tailed Lemur, despite the lack of a
large brain (relative to simiiform primates), can organize sequences, understand basic arithmetic operations
and preferentially select tools based on functional qualities.
Despite being listed as Near Threatened by the IUCN Red List and suffering from habitat destruction,
the Ring-tailed Lemur reproduces readily in captivity and is the most populous lemur in zoos worldwide,
numbering more than 2000 individuals. It typically lives 16 to 19 years in the wild and 27 years in captivity.
Contents
[hide]
• 1 Etymology
• 2.1 Taxonomic classification
• 3 Anatomy and physiology
• 3.1 Dentition
• 4 Ecology
• 4.1 Geographic range and habitat
• 5 Behavior
• 5.1 Diet
• 5.2 Social systems
• 5.2.1 Olfactory communication
• 5.2.2 Auditory communication
• 5.3 Breeding and reproduction
• 5.4 Cognitive abilities and tool use
• 6 Conservation status
• 7 Cultural references
• 8 Footnotes
• 9 References
[edit] Etymology
The genus name Lemur was created by Carl Linnaeus, the founder of modern binomial
nomenclature, to describe only three species,[5] but the word eventually became the collective name used
for all primates endemic to Madagascar. Linnaeus was familiar with the historical works of Virgil and Ovid
and their references to the festival of Lemuria, during which specters or ghosts—referred to as lemures—were
exorcised. As an analogy to these ghosts from Roman mythology, he created the name "Lemur" to include
these prosimian primates[7][8] due to their nocturnal habits and ghost-like appearance.[9] Their noiseless
movements at night, reflective eyes, and ghostly cries may also have been a factor. It is even possible that
Linnaeus knew that some Malagasy people have held legends that lemurs are the souls of their ancestors.
[10]
The species name, catta, refers to the Ring-tailed Lemur's cat-like appearance. Its purring
vocalization is similar to that of the Domestic Cat.[5]

All mammalian fossils from Madagascar come from recent times.[11] Thus, little is known about the
evolution of the Ring-tailed Lemur, let alone the rest of the lemur clade, which comprises the entire endemic
primate population of the island. However, chromosomal and molecular evidence suggest that lemurs are
more closely related to each other than to other Strepsirrhine primates. For this to have happened, it is
thought that a very small ancestral population came to Madagascar via a single rafting event between 50 and
80 million years ago.[7][11][12] Subsequent evolutionary radiation and speciation has created the diversity of
Malagasy lemurs seen today.[13]
According to analysis of amino acid sequences, the branching of the family Lemuridae has been
dated to 26.1 ±3.3 mya while rRNA sequences of mtDNA place the split at 24.9 ±3.6 mya. The ruffed lemurs
are the first genus to split away (most basal) in the family, a view that is further supported by analysis of DNA
sequences and karyotypes.[14] Additionally, Molecular data suggests a deep genetic divergence and sister
group relationship between the true lemurs (Eulemur) the remaining three genera: Lemur, Hapalemur, and
Prolemur.[15]
The Ring-tailed Lemur is thought to share closer affinities to the bamboo lemurs of the genera
Hapalemur and Prolemur than to the other two genera in its family.[16][17] This has been supported by
comparisons in communication, chromosomes, genetics, and several morphological traits, such as scent
gland similarities. However, other data concerning immunology and other morphological traits fail to support
this close relationship. For example, Hapalemur and Prolemur have short snouts, while the Ring-tailed Lemur
and the rest of Lemuridae have long snouts. (However, differences in the relationship between the orbit (eye
socket) and the muzzle suggest that the Ring-tailed Lemur and the true lemurs evolved their elongated faces
independently.)[15]
The relationship between the Ring-tailed Lemur and bamboo lemurs (both Hapalemur and Prolemur)
is the least understood. Molecular analysis suggests that either the bamboo lemurs diverged from the Ring-
tailed Lemur, making the group monophyletic and supporting the current 3-genera taxonomy, or that the
Ring-tailed Lemur is nested in with the bamboo lemurs.[15]
The karyotype of the Ring-tailed Lemur has 56 chromosomes, of which four are metacentric (arms of
nearly equal length), four are submetacentric (arms of unequal length), and 46 are acrocentric (the short arm
is hardly observable). The X chromosome is metacentric and the Y chromosome is acrocentric.[5]
[edit] Taxonomic classification

Linnaeus first used the genus name Lemur to describe "Lemur tardigradus" (the Red Slender Loris,
now known as Loris tardigradus) in his 1754 catalog of the Museum of King Adolf Frederick. In 1758, his 10th
edition of Systema Naturae listed the genus Lemur with three included species, only one of which is still
considered to be a lemur while another is no longer considered to be a primate. These species include:
Lemur tardigradus, Lemur catta (the Ring-tailed Lemur), and Lemur volans (the Philippine Colugo, now
known as Cynocephalus volans). In 1911, Oldfield Thomas made Lemur catta the type species for the genus,
despite the term initially being used to describe lorises. On January 10, 1929, the International Commission
on Zoological Nomenclature (ICZN) formalized this decision in its publication of Opinion 122.[5][6]
The Ring-tailed Lemur shares many similarities with ruffed lemurs (genus Varecia) and true lemurs
(genus Eulemur), and its skeleton is nearly indistinguishable from that of the true lemurs.[16] Consequently,
the three genera were once grouped together in the genus Lemur and more recently are sometimes referred
to as subfamily Lemurinae (within family Lemuridae). However, ruffed lemurs were reassigned to the genus
Varecia in 1962,[1] and due to similarities between the Ring-tailed Lemur and the bamboo lemurs,
particularly in regards to molecular evidence and scent glands similarities, the true lemurs were moved to the
genus Eulemur by Yves Rumpler and Elwyn Simons (1988) as well as Colin Groves and Robert H. Eaglen
(1988).[1][5][16][17] In 1991, Ian Tattersall and Jeffrey H. Schwartz reviewed the evidence and came to a
different conclusion, instead favoring to return the members of Eulemur and Varecia to the genus Lemur.
However, this view was not widely accepted and the genus Lemur remained monotypic, containing only the
Ring-tailed Lemur.[1][17][18] Because the differences in molecular data are so minute between the Ring-
tailed Lemur and both genera of bamboo lemurs, it has been suggested that all three genera be merged.[15]
Because of the difficulty in discerning the relationships within family Lemuridae, not all authorities
agree on the taxonomy, although the majority of the primatological community favors the current
classification.[18]
Taxonomy of family Lemuridae[1][17][18] Phylogeny of

family Lemuridae[12][14]
[15][19]
• Family Lemuridae
• Genus Lemur: the Ring-tailed Lemur
• Genus Eulemur: brown lemurs
• Genus Varecia: ruffed lemurs
• Genus Hapalemur: lesser gentle or bamboo lemurs
• Genus Prolemur: the Greater Bamboo Lemur
Lemuridae
Varecia
(ruffed lemurs)
Lemur (Ring-
tailed Lemur)
Prolemur
(Greater Bamboo
Lemur)
Hapalemur
(lesser bamboo lemurs)
Eulemur
(true lemurs)
In 1996, researchers Steven Goodman and Olivier Langrand suggested that the Ring-tailed Lemur
may demonstrate regional variations, particularly a high mountain population at Andringitra Massif that has a
thicker coat, lighter coloration, and variations in its tail rings.[5][20] In 2001, primatologist Colin Groves
concluded that this does not represent a locally occurring subspecies. This decision was later supported by
further fieldwork that showed that the differences fell within the normal range of variation for the species. The
thicker coat was considered a local adaptation to extreme low temperatures in the region, and the fading of
the fur was attributed to increased exposure to solar radiation. Additional genetic studies in 2000 further
supported the conclusion that population did not vary significantly from the other Ring-tailed Lemur
populations on the island.[20][21]
[edit] Anatomy and physiology
The Ring-tailed lemur is a relatively large lemur. Its average weight is 2.2 kilograms (4.9 lb).[18] Its
head–body length ranges between 39 and 46 cm (15 and 18 in), its tail length is 56 and 63 cm (22 and 25 in),
and its total length is 95 and 110 cm (37 and 43 in).[5][18] Other measurements include a hind foot length of
102 and 113 mm (4.0 and 4.4 in), ear length of 40 and 48 mm (1.6 and 1.9 in), and cranium length of 78 and
88 mm (3.1 and 3.5 in).[5]
The Ring-tailed Lemur's tail is longer than its body.

The species has a slender frame and narrow face, fox-like muzzle.[5] The Ring-tailed Lemur's
trademark—a long, bushy tail—is ringed in alternating black and white transverse stripes, numbering 12 or
13 white rings and and 13 or 14 black rings, and always ending in a black tip.[5][16][22] Its tail is longer than
its body[22] and is not prehensile. Instead, it is only used for balance, communication, and group cohesion.
[18]
The pelage (fur) is so dense that it can clog electric clippers.[5] The ventral (chest) coat and throat
are white or cream. The dorsal (back) coat varies from gray to rosy-brown, sometimes with a brown pygal
patch around the tail region, where the fur grades to pale gray or grayish brown. The dorsal coloration is
slightly darker around the neck and crown. The hair on the throat, cheeks, and ears is white or off-white and
also less dense, allowing the dark skin underneath to show through.[5][6][18] The muzzle is dark grayish and
the nose is black, and the eyes are encompassed by black triangular patches.[5][18] Facial vibrissae
(whiskers) are developed and found above the lips (mystacal), on the cheeks (genal), and on the eyebrow
(superciliary). Vibrissae are also found slightly above the wrist on the underside of the forearm.[5] The ears
are relatively large compared to other lemurs and are covered in hair, which has only small tufts if any.[5][6]
Although slight pattern variations in the facial region may be seen between individuals, there are no obvious
differences between the sexes.[5]
Unlike most diurnal primates, but like all strepsirhine primates, the Ring-tailed Lemur has a tapetum
lucidum, or reflective layer behind the retina of the eye, that enhances night vision.[23] The tapetum is highly
visible in this species because the pigmentation of the ocular fundus (back surface of the eye), which is
present in—but varies between—all lemurs, is very spotty. The Ring-tailed Lemur also has a rudimentary
foveal depression on the retina. Another shared characteristic with the other strepsirrhine primates is the
rhinarium), a moist, naked, glandular nose supported by the upper jaw and protruding beyond the chin. The
rhinarium continues down where it divides the upper lip. The upper lip is attached to the premaxilla,
preventing the lip from protruding and thus
requiring the lemur to lap water rather than using
suction.[5]
Scent glands on a male: the brachial glands on
the upper chest (left), and antebrachial gland
and spur on the forearm (right)
The skin of the Ring-tailed Lemur is dark gray or black in color, even in places where the fur is white.
It is exposed on the nose, palms, soles, eyelids, lips, and genitalia. The skin is smooth, but the leathery
texture of the hands and feet facilitate terrestrial movement. The anus, located at the joint of the tail, is
covered covered when the tail is lowered. The area around the anus (circumanal area) and the perineum are
covered in fur. In males, the scrotum lacks fur, is covered in small, horny spines, and the two sacs of the
scrotum are divided. The penis is nearly cylindrical in shape is covered in small spines, as well as two pairs
of larger spines on both sides. Males have a relatively small baculum (penis bone) compared to their size.
The scrotum, penis, and prepuce are usually coated with a foul-smelling secretion. Females have a thick,
elongated clitoris that protrudes from the labia of the vulva. The opening of the urethra is closer to the clitoris
than the vagina, forming a "drip tip."[5]
Females have two pairs of mammary glands (four nipples), but only one pair is functional.[5][6] The
pair anterior pair (closest to the head) are very close to the axillae (armpit).[5] Furless scent glands are
present on both males and females. Both genders have small, dark antebrachial (forearm) glands measuring
1 cm long and located on the inner surface of the forearm nearly 25 cm (9.8 in) above the wrist joint.[5][6][18]
(This trait is shared between both the Lemur and Hapalemur genera.[18]) The gland is soft and
compressible, bears fine dermal ridges (like fingerprints), and is connected to the palm by a fine, 2 mm–high,
hairless strip.[5] However, only the male has a horny spur that overlays this scent gland.[5][6][18] The spur
develops with age through the accumulation of secretions from an underlying gland that may connect through
the skin through as many as a thousand minuscule ducts. The males also have brachial (arm) glands on the
axillary surface of their shoulders (near the armpit). The brachial gland is larger than the antebrachial gland,
covered in short hair around the periphery, and has a naked crescent-shaped orifice near the center. The
gland secretes a foul-smelling, brown, sticky substance.[5] The brachial gland is barely developed if present
at all in females.[5][6] Both genders also have apocrine and sebaceous glands in their genital or perianal
regions,[24] which care covered in fur.[6]
Its fingers are slender, padded, mostly lacking webbing, and semi-dexterous with flat, human-like
nails. The thumb is both short and widely separated from the other fingers. Despite being set at a right angle
to the palm, the thumb is not opposable since the ball of the joint is fixed in place. As with all strepsirrhines,
the hand is ectaxonic (the axis passes through the fourth digit) rather than mesaxonic (the axis passing
through the third digit) as seen in monkeys and apes. The fourth digit is the longest, and only slightly longer
than the second digit. Likewise, the fifth digit is only slightly longer than the second. The palms are long and
leathery,[5] and like other primates, they have dermal ridges to improve grip.[25] The feet are semi-
digitigrade and more specialized than the hands. The big toe is opposable and is smaller than the big toe of
other lemurs, which are more arboreal. The second toe is toe is short, has a small terminal pad, and has a
toilet-claw (sometimes referred to as a grooming claw) specialized for personal grooming, specifically to rake
through fur that is unreachable by the mouth.[5] The toilet-claw is a trait shared among nearly all prosimian
primates.[26] Unlike other lemurs, the Ring-tailed Lemur's heel is not covered by fur.[5]
Like other lemurs, the
Ring-tailed lemur has a
claw-like nail (toilet-
claw) on their second
toe (left) and dermal
ridges on its hands to
improve its grip
(center). Unlike other
lemurs, it lacks fur on
its heel (right).
[edit] Dentition
The front, lower dentition includes a toothcomb (4 incisors and 2 canine teeth), while the first
premolars resemble canines.
The Ring-tailed Lemur has a dentition of , meaning that on each side of the jaw it has
two incisors, one canine tooth, three premolars, and three molar teeth.[5] Its deciduous dentition is
.[27] The permanent teeth erupts in the following order: m 1/1 (first molars), i 2/2 (first incisors), i
3/3 (second incisors), C1 (upper canines), m 2/2 (second molars), c1 (lower canines), m 3/3 (third molars), p
4/4 (third premolars), p 3/3 (second premolars), p 2/2 (first premolars).[5]
Its lower incisors (i1 and i2) are long, narrow, and finely spaced while pointing almost straight forward
in the mouth (procumbent). Together with the incisor-shaped (incisiform) lower canines (c1), which are
slightly larger and also procumbent, form a structure called a toothcomb,[5] a trait unique to nearly all
strepsirrhine primates.[28] The toothcomb is used during oral grooming, which involves licking and tooth-
scraping. It may also be used for grasping small fruits, removing leaves from the stem when eating, and
possibly scraping sap and gum from tree bark. The toothcomb is kept clean using a sublingual organ—a thin,
flat, fibrous plate that covers a large part of the base of the tongue. The first lower premolar (p2) following the
toothcomb is shaped like a canine (caniniform) and occludes the upper canine, essentially filling the role of
the incisiform lower canine. There is also a diastema (gap) between the second and third premolars (p2 and
p3).[5]
The upper incisors are small, with the first incisors (I1) space widely from each other, yet closely to
the second incisors (I2). Both are compressed between buccolingually (between the cheek and the tongue).
The upper canines (C1) are long, have a broad base, and curve down and back (recurved). The upper
canines exhibit slight sexual dimorphism, with males exhibiting slightly larger canines than females. Both
sexes use them in combat by slashing with them. There is a small diastema between the upper canine and
the first premolar (P2), which is smaller and more caniniform than the other premolars. Unlike other lemurs,
the first two upper molars (M1 and M2) have prominent lingual cingulae, yet do not have a protostyle.[5]
[edit] Ecology
The Ring-tailed Lemur can leap from tree to tree.

The Ring-tailed Lemur is diurnal and semi-terrestrial. It is the most terrestrial of lemur species,
spending as much as 33% of its time on the ground. However it is still considerably arboreal, spending 23%
of its time in the mid-level canopy, 25% in the upper-level canopy, 6% in the emergent layer and 13% in
small bushes. Troop travel is 70% terrestrial.[29]
Troop size, home range, and population density vary by region and food availability. Troops typically
range in size from 6 to 25, although troops with over 30 individuals have been recorded.[16] The average
troop contains 13 to 15 individuals.[16] Home range sizes varies between 6 and 35 hectares (15 and 86
acres).[30] Troops of the Ring-tailed Lemur will maintain a territory, but overlap is often high. When
encounters occur, they are agonistic, or hostile in nature. A troop will usually occupy the same part of its
range for three or four days before moving. When it does move, the average traveling distance is 1 km (0.62
mi).[29] Population density ranges from 100 individuals per 1 km2 (0.39 sq mi) in dry forests to 250–
600 individuals per km2 in gallery and secondary forests.[22]
The Ring-tailed Lemur has both native and introduced predators. Native predators include the Fossa
(Cryptoprocta ferox), the Madagascar Harrier-hawk (Polyboroides radiatus), the Madagascar Buzzard (Buteo
brachypterus) and the Madagascar Ground Boa (Boa madagascariensis). Introduced predators include the
Small Indian Civet (Viverricula indica), the Domestic Cat and the Domestic Dog.[22]
[edit] Geographic range and habitat
The Ring-tailed Lemur can be found in several protected areas, including Isalo National Park in
Madagascar.
Endemic to southern and southwestern Madagascar, the Ring-tailed Lemur ranges further into
highland areas than other lemurs. It inhabits deciduous forests, dry scrub, montane humid forests, and
gallery forests (forests along riverbanks). It strongly favors gallery forests, but such forests have now been
cleared from much of Madagascar in order to create pasture for livestock.[22][18] Depending on location,
temperatures within its geographic range can vary from −12 °C (10 °F)[21] at Andringitra Massif to 48 °C
(118 °F) in the spiny forests of Beza Mahafaly Special Reserve.[31]
This species is found as far east as Tôlanaro, inland towards the mountains of Andringitra on the
southeastern plateau, among the spiny forests of the southern part of the island,[22][18] and north along the
west coast to the town of Belo sur Mer.[5] Historically, the northern limits of its range in the west extended to
the Morondava River near Morondava. It can still be found in Kirindy Mitea National Park, just south of
Morondava, though at very low densities. It does not occur in Kirindy Forest Reserve, north of Morondava. Its
distribution throughout the rest of its range is very spotty, with population densities varying widely.[18]
The Ring-tailed Lemur can be easily seen in five national parks in Madagascar: Andohahela National
Park, Andringitra National Park, Isalo National Park, Tsimanampetsotse National Park, and Zombitse-
Vohibasia National Park. It can also be found in Beza-Mahafaly Special Reserve, Kalambatritra Special
Reserve, Pic d'Ivohibe Special Reserve, Amboasary Sud, Berenty Private Reserve, Anja Community
Reserve, and marginally at Kirindy Mitea National Park. Unprotected forests that the species has been
reported in include Ankoba, Ankodida, Anjatsikolo, Anbatotsilongolongo, Bereny, Mahazoarivo, Masiabiby,
and Mikea.[18]
Within the protected regions it is known to inhabit, the Ring-tailed Lemur is sympatric (shares its
range) with as many as 24 species of lemur, covering every living genus except Allocebus, Indri, and
Varecia. Historically, the species used to be sympatric with the critically endangered Southern Black-and-
white Ruffed Lemur (Varecia variegata editorum), which was once found at Andringitra National Park;
however, no sightings of the ruffed lemur have been reported in recent years.[32]
List of species sympatric with the Ring-tailed Lemur[32]
• Verreaux's Sifaka • Greater Dwarf Lemur • Southern Lesser Bam
(Propithecus verreauxi) (Cheirogaleus major) Lemur
• Milne-Edwards' Sifaka • White-footed Sportive (Hapalemur meridionalis)
(Propithecus edwardsi) Lemur • Greater Bamboo Lem
• Peyrieras' Woolly Lemur (Lepilemur leucopus) (Prolemur simus)
(Avahi peyrierasi) • Small-toothed Sportive • Common Brown Lemu
• Gray Mouse Lemur Lemur (Eulemur fulvus)
(Microcebus murinus) (Lepilemur microdon) • Red-fronted Lemur
• Brown Mouse Lemur • Petter's Sportive (Eulemur rufifrons)
(Microcebus rufus) Lemur • Red-bellied Lemur
• Reddish-gray Mouse (Lepilemur petteri) {Eulemur rubriventer)
Lemur • Red-tailed Sportive • Collared Brown Lemu
(Microcebus griseorufus) Lemur (Eulemur collaris)
• Aye-aye (Lepilemur ruficaudatus) • Coquerel's Giant Mou
(Daubentonia madagascariensis) • Wright's Sportive Lemur
• Fat-tailed Dwarf Lemur Lemur (Mirza coquereli)
(Cheirogaleus medius) (Lepilemur wrightae) • Pale Fork-marked Lem
• Hubbard's Sportive (Phaner pallescens)[N 6]
Lemur
(Lepilemur hubbardorum)
• Golden Bamboo Lemur
(Hapalemur aureus)
In western Madagascar, sympatric Ring-tailed Lemurs and Red-fronted Lemurs (Eulemur rufifrons)
have been studied together. Little interaction takes place between the two species. While the diets of the two
species overlap, they eat in different proportions since the Ring-tailed Lemur has a more varied diet and
spends more time on the ground.[29]
[edit] Behavior
[edit] Diet
The Ring-tailed Lemur is an opportunistic omnivore primarily eating fruits and leaves, particularly
those of the tamarind tree (Tamarindus indica), known natively as kily.[22][29] When available, tamarind
makes up as much as 50% of the diet, especially during the dry, winter season.[22] The Ring-tailed Lemur
eats from as many as three dozen different plant species, and its diet includes flowers, herbs, bark and sap.
It has been observed eating decayed wood, earth, spider webs, insect cocoons, arthropods (spiders,
caterpillars, cicadas and grasshoppers) and small vertebrates (birds and chameleons).[22] During the dry
season it becomes increasingly opportunistic.
[edit] Social systems

Troops are classified as multi-male groups, with a matriline as the core group.[33] As with most
lemurs, females socially dominate males in all circumstances, including feeding priority. Dominance is
enforced by lunging, chasing, cuffing, grabbing and biting. Young females do not always inherit their mother's
rank and young males leave the troop between three and five years of age.[29][33] Both sexes have separate
dominance hierarchies; females have a distinct hierarchy while male rank is correlated with age. Each troop
has one to three central, high-ranking adult males who interact with females more than other group males
and lead the troop procession with high-ranking females. Recently transferred males, old males or young
adult males that have not yet left their natal group are often lower ranking. Staying at the periphery of the
group they tend to be marginalized from group activity.[33]
The Ring-tailed Lemur will sit facing the sun to warm itself in the mornings.
For males, social structure changes can be seasonal. During the six month period between
December and May a few males immigrate between groups. Established males transfer every 3.5 years,[29]
although young males may transfer every 1.4 years. Group fission occurs when groups get too large and
resources become scarce.[33]
In the mornings the Ring-tailed Lemur sunbathes to warm itself. It faces the sun sitting in what is
frequently described as a "sun-worshipping" posture or Lotus position. However, it sits with its legs extended
outward, not cross-legged, and will often support itself on nearby branches. Sunning is often a group activity,
particularly during the cold mornings. At night, troops will split into sleeping parties huddling closely together
to keep warm.[33] A group of huddled Ring-tailed Lemurs is referred to as a lemur ball.
Despite being quadrupedal the Ring-tailed Lemur can rear up and balance on its hind legs, usually
for aggressive displays. When threatened the Ring-tailed Lemur may jump in the air and strike out with its
short nails and sharp upper canine teeth in a behaviour termed jump fighting. This is extremely rare outside
of the breeding season when tensions are high and competition for mates is intense. Other aggressive
behaviours include a threat-stare, used to intimidate or start a fight, and a submissive gesture known as
pulled-back lips.[33]
Border disputes with rival troops occur occasionally and it is the dominant female's responsibility to
defend the troop's home range. Agonistic encounters include staring, lunging approaches and occasional
physical aggression, and conclude with troop members retreating toward the center of the home range.[33]
[edit] Olfactory communication

Male Ring-tailed Lemurs will scent-mark saplings and branches by spur-marking.
Olfactory communication is critically important for prosimians like the Ring-tailed Lemur. Males and
females scent mark both vertical and horizontal surfaces at the overlaps in their home ranges using their
anogenital scent glands. The Ring-tailed Lemur will perform a handstand to mark vertical surfaces, grasping
the highest point with its feet while it applies its scent.[33] Use of scent marking varies by age, sex and social
status.[34] Male lemurs use their antebrachial and brachial glands to demarcate territories and maintain
intragroup dominance hierarchies. The thorny spur that overlays the antebrachial gland on each wrist is
scraped against tree trunks to create grooves anointed with their scent. This is known as spur-marking.[33]
In displays of aggression, males engage in a social display behaviour called stink fighting, which
involves impregnating their tails with secretions from the antebrachial and brachial glands and waving the
scented tail at male rivals.[35] During the mating season, males wave their scented tails at females as a form
of sexual overture; this usually results in the female cuffing or biting the male and elicits subordinate
vocalizations from the would-be paramour.
Ring-tailed Lemurs have also been shown to mark using urine. Behaviorally, there is a difference
between regular urination, where the tail is slightly raised and a stream of urine is produced, and the urine
marking behavior, where the tail is held up in display and only a few drops of urine are used.[36][37] The
urine-marking behavior is typically used by females to mark territory, and has been observed primarily at the
edges of the troop's territory and in areas where other troops may frequent.[38] The urine marking behavior
also is most frequent during the mating season, and may play a role in reproductive communication between
groups.[36]
[edit] Auditory communication

Main article: Ring-tailed Lemur vocalizations
The Ring-tailed Lemur is one of the most vocal primates and has a complex array of distinct
vocalizations used to maintain group cohesion during foraging and alert group members to the presence of a
predator. Calls range from simple to complex. An example of a simple call is the purr ( listen (help·info)),
which expresses contentment. A complex call is the sequence of clicks, close-mouth click series (CMCS),
open-mouth click series (OMCS) and yaps ( listen (help·info)) used during predator mobbing. Some calls
have variants and undergo transitions between variants, such as an infant "whit" (distress call) transitioning
from one variant to another ( listen (help·info)).[39]
The most commonly heard vocalizations are the moan ( listen (help·info)) (low-to-moderate
arousal, group cohesion), early-high wail ( listen (help·info)) (moderate-to-high arousal, group cohesion),
and clicks ( listen (help·info)) ("location marker" to draw attention).[39]
[edit] Breeding and reproduction
In the wild, females typically give birth to a single offspring.

The Ring-tailed Lemur is polygynous, although the dominant male in the troop typically breeds with
more females than other males. Fighting is most common during the breeding season.[40] A receptive
female may initiate mating by presenting her backside, lifting her tail and looking at the desired male over her
shoulder. Males may inspect the female's genitals to determine receptiveness. Females typically mate within
their troop, but may seek outside males.[29]
The breeding season runs from mid-April to mid-May. Estrus lasts 4 to 6 hours,[16] and females mate
with multiple males during this period.[29] Within a troop, females stagger their receptivity so that each
female comes into season on a different day during the breeding season, reducing competition for male
attention.[41] Gestation lasts for about 135 days, and parturition occurs in September or occasionally
October. In the wild one offspring is the norm, although twins may occur. Ring-tailed Lemur infants have a
birth weight of 70 g (2.5 oz) and are carried ventrally (on the chest) for the first 1 to 2 weeks, then dorsally (on
the back).[16]
The young lemurs begin to eat solid food after two months and are fully weaned after five months.
Sexual maturity is reached between 2.5 and 3 years.[40] Male involvement in infant rearing is limited,
although the entire troop, regardless of age or sex, can be seen caring for the young. Alloparenting between
troop females has been reported. Kidnapping by females and infanticide by males also occur occasionally.
[33] Due to harsh environmental conditions, predation and accidents such as falls infant mortality can be as
high as 50% within the first year and as few as 30% may reach adulthood.[16] The longest-lived Ring-tailed
Lemur in the wild was a female at the Berenty Reserve who lived for 20&nbps;years.[22] In the wild, females
rarely live past the age of 16, whereas the life expectancy of males is not known due to their social structure.
The longest-lived male was reported to be 15&nbps;years old. The maximum lifespan reported in captivity
was 27 years.[5]
[edit] Cognitive abilities and tool use
Historically, the studies of learning and cognition in non-human primates have focused on simians
(monkeys and apes), while strepsirrhine primates, such as the Ring-tailed Lemur and its allies, have been
overlooked and popularly dismissed as unintelligent.[42] A couple of factors stemming from early
experiments have played a role in the development of this assumption. First, the experimental design of older
tests may have favored the natural behavior and ecology of simians over that of strepsirrhines, making the
experimental tasks inappropriate for lemurs. For example, simians are known for their manipulative play with
non-food objects, whereas lemurs are only known to manipulate non-food objects in captivity.[43] This
behaviour is usually connected with food association. Also, lemurs are known to displace objects with their
nose or mouth more so than with their hands.[42] Therefore, an experiment requiring a lemur to manipulate
an object without prior training would favor simians over strepsirrhines. Second, individual Ring-tailed Lemurs
accustomed to living in a troop may not respond well to isolation for laboratory testing. Past studies have
reported hysterical behaviour in such scenarios.[44] As a result of these early studies lemurs were often
omitted from further research.
The notion that lemurs are unintelligent has been perpetuated by the view that the neocortex ratio (as
a measure of brain size) indicates intelligence.[45] In fact, primatologist Alison Jolly noted early in her
academic career that some lemur species, such as the Ring-tailed Lemur, have evolved a social complexity
similar to that of cercopithecine monkeys, but not the corresponding intelligence.[46] After years of
observations of wild Ring-tailed Lemur populations at the Berenty Reserve in Madagascar and as well as
baboons in Africa, she more recently concluded that this highly social lemur species does not demonstrate
the equivalent social complexity of cercopithecine monkeys, despite general appearances.[47]
Regardless, research has continued to illuminate the complexity of the lemur mind, with emphasis on
the cognitive abilities of the Ring-tailed Lemur. As early as the mid-1970s, studies had demonstrated that
they could be trained through operant conditioning using standard schedules of reinforcement. The species
has been shown to be capable of learning pattern, brightness and object discrimination, skills common
among vertebrates. The Ring-tailed Lemur has also been shown to learn a variety of complex tasks often
equaling, if not exceeding, the performance of simians.[42]
More recently, research at the Duke Lemur Center has shown that the Ring-tailed Lemur can
organize sequences in memory and retrieve ordered sequences without language.[48] The experimental
design demonstrated that the lemurs were using internal representation of the sequence to guide their
responses and not simply following a trained sequence, where one item in the sequence cues the selection
of the next.[48] But this is not the limit of the Ring-tailed Lemur's reasoning skills. Another study, performed at
the Myakka City Lemur Reserve, suggests that this species along with several other closely related lemur
species understand simple arithmetic operations.[49]
Since tool use is considered to be a key feature of primate intelligence, the apparent lack of this
behavior in wild lemurs, as well as the lack of non-food object play, has helped reinforce the perception that
lemurs are less intelligent than their simian cousins.[43] However, another study at the Myakka City Lemur
Reserve examined the representation of tool functionality in both the Ring-tailed Lemur and the Common
Brown Lemur and discovered that, like monkeys, they utilized tools with functional properties (e.g., tool
orientation or ease of use) instead of tools with nonfunctional features (e.g., color or texture). Although the
Ring-tailed Lemur may not use tools in the wild, it can not only be trained to use a tool, but will preferentially
select tools based on their functional qualities. Therefore, the conceptual competence to utilize a tool may
have been present in the common primate ancestor, even though the use of tools may not have appeared
until much later.[50]
[edit] Conservation status
In addition to being listed as Near Threatened in 2008 by the IUCN,[2] the Ring-tailed Lemur has
been listed since 1977 by CITES under Appendix I,[3] which makes trade of wild-caught specimens illegal.
Although there are more endangered species of lemur, the Ring-tailed Lemur is considered a flagship
species due to its recognizability.[51]
Ring-tailed lemurs are a common sight at Berenty Private Reserve in southern Madagascar.
Three factors threaten Ring-tailed Lemurs. First and foremost is habitat destruction. Starting nearly
2,000 years ago with the introduction of humans to the island, forests have been cleared to produce pasture
and agricultural land.[51] Extraction of hardwoods for fuel and lumber, as well mining and overgrazing, have
also taken their toll. Today, it is estimated that 90% of Madagascar's original forest cover has been lost.[52]
Rising populations have created even greater demand in the southwest portion of the island for fuel wood,
charcoal, and lumber. Fires from the clearing of grasslands, as well as slash-and-burn agriculture destroy
forests. Another threat to the species is harvesting either for food (bush meat) or pets. Finally, periodic
drought common to southern Madagascar can impact populations already in decline. In 1991 and 1992, for
example, a severe drought caused an abnormally high morality rate among infants and females at the Beza
Mahafaly Special Reserve. Two years later, the population had declined by 31% and took nearly four years
to start to recover.[51]
The Ring-tailed Lemur resides in several protected areas within its range, each offering varying
levels of protection. At the Beza Mahafaly Special Reserve, a holistic approach to in situ conservation has
been taken. Not only does field research and resource management involve international students and local
people (including school children), livestock management is used at the peripheral zones of the reserve and
ecotourism benefits the local people.[51]
Outside of its diminishing habitat and other threats, the Ring-tailed Lemur reproduces readily and has
fared well in captivity. For this reason, along with its popularity, it has become the most populous lemur in
zoos worldwide, with more than 2500 in captivity as of 2009. It is also the most common of all captive
primates.[18] Ex situ facilities actively involved in the conservation of the Ring-tailed Lemur include the Duke
Lemur Center in Durham, NC, the Lemur Conservation Foundation in Myakka City, FL and the Madagascar
Fauna Group headquartered at the Saint Louis Zoo. Due to the high success of captive breeding,
reintroduction is a possibility if wild populations were to crash. Although experimental releases have met
success on St. Catherines Island in Georgia, demonstrating that captive lemurs can readily adapt to their
environment and exhibit a full range of natural behaviors, captive release is not currently being considered.
[51]
Ring-tailed Lemur populations can also benefit from drought intervention, due to the availability of
watering troughs and introduced fruit trees, as seen at the Berenty Private Reserve in southern Madagascar.
[51] However, these interventions are not always seen favorably, since natural population fluctuations are not
permitted. The species is thought to have evolved its high fecundity due to its harsh environment;[51]
therefore, interfering with this natural cycle could significantly impact the gene pool.
[edit] Cultural references

The Ring-tailed Lemur is known locally in Malagasy as hira (pronounced [ˈhirə] or colloquially [ˈir]) or
maky (pronounced [maki̥] , and spelled maki in French). Being the most widely recognized endemic primate
on the island, it has been selected as the symbol for Madagascar National Parks (formerly known as
ANGAP).[18] The Maki brand, which started by selling t-shirts in Madagascar and now sells clothing across
the Indian Ocean islands, is named after the this lemur due to its popularity, despite the fact that the
company's logo portrays the face of a sifaka and its name uses the French spelling.[53]
The first mention of the Ring-tailed Lemur in Western literature came in 1625 when English traveler
and writer Samuel Purchas described them as being comparable in size to a monkey and having a fox-like
long tail with black and white rings.[5]
It has been popularized in Western culture by the Animal Planet television series Lemur Kingdom
(United States) and Lemur Street (United Kingdom), as well as by the character King Julien in the animated
Madagascar film series and spin-off TV series. Lemur Street depicts real events in the lives of wild Ring-
tailed Lemurs, whereas the Madagascar films and spin-off series depict anthropomorphic representations,
such as lemurs talking, singing, and dancing. The Ring-tailed Lemur was also the focus of the 1996 Nature
documentary A Lemur's Tale, which was filmed at the Berenty Reserve and followed a troop of lemurs. The
troop included a unique infant named Sapphire, who was nearly albino, with white fur, bright blue eyes, and
the characteristic ringed tail.
This species also played a role in the 1997 comedy film Fierce Creatures, starring John Cleese, who
has a passion for lemurs.[54][55] Cleese later hosted the 1998 BBC documentary In the Wild: Operation
Lemur with John Cleese, which tracked the progress of a reintroduction of Black-and-white Ruffed Lemurs
back into the Betampona Reserve in Madagascar. The project had been partly funded by Cleese's donation
of the proceeds from the London premier of Fierce Creatures.[55][56]
[edit] Footnotes
1. ^ The genus name Prosimia was declared unavailable by the International Commission on
Zoological Nomenclature in 1998.[5]
2. ^ Type species was designated as Catta mococo (= Lemur catta Linnaeus, 1758).[5]
3. ^ Type species was designated as Maki mococo (= Lemur catta Linnaeus, 1758).[5][6]
4. ^ The synonym Mococo is sometimes omitted because it was technically a vernacular term
for the genus Prosimia.[6] René Primevère Lesson named the type species for this genus as
Prosimia catta (= Lemur catta Linnaeus, 1758) in the same year (1878).[5]
5. ^ Muirhead (1819) credited the name Maki mococo to Anselme Gaëtan Desmarest (1817),
although it was actually used as a vernacular name.[5][6]
6. ^ The Pale Fork-marked Lemur (Phaner pallescens) found at Zombitse-Vohibasia National
Park may instead be a new species.[32]
[edit] References
1. ^ a b c d e Groves, Colin P. (16 November 2005). "Lemur catta". In Wilson, Don E., and
Reeder, DeeAnn M., eds. Mammal Species of the World: A Taxonomic and Geographic Reference
(3rd ed.). Baltimore: Johns Hopkins University Press, 2 vols. (2142 pp.). p. 117. ISBN 978-0-8018-
8221-0. OCLC 62265494. http://www.bucknell.edu/msw3/browse.asp?id=12100056.
2. ^ a b Andrainarivo, C., Andriaholinirina, V. N., Feistner, A., Felix, T., Ganzhorn, J., Garbutt,
N., Golden, C., Konstant, B., Louis Jr., E., Meyers, D., Mittermeier, R. A., Perieras, A., Princee, F.,
Rabarivola, J. C., Rakotosamimanana, B., Rasamimanana, H., Ratsimbazafy, J., Raveloarinoro, G.,
Razafimanantsoa, A., Rumpler, Y., Schwitzer, C., Sussman, R., Thalmann, U., Wilmé, L. & Wright, P.
(2008). 'Lemur catta'. In: IUCN 2008. IUCN Red List of Threatened Species. Downloaded on 1
January 2009.
3. ^ a b UNEP-WCMC. "CITES species database: Lemur catta". UNEP-WCMC Species
Database. http://www.unep-wcmc.org/isdb/CITES/Taxonomy/tax-species-
result.cfm/isdb/CITES/Taxonomy/tax-species-result.cfm?
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4. ^ Mittermeier et al. 2006, p. 238
5. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al am an ao ap aq Wilson,
D.E.; Hanlon, E. (2010). "Lemur catta (Primates: Lemuridae)" (PDF). Mammalian Species 42 (854):
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7. ^ a b Garbutt 2007, pp. 85–86
8. ^ Blunt & Stearn 2002, p. 252
9. ^ Flower 1891, p. 682
10.^ Nield 2007, p. 41
12.^ a b Horvath, J.E.; Weisrock, D.W.; Embry, S.L.; Fiorentino, I.; Balhoff, J.P.; Kappeler, P.;
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27.^ Ankel-Simons 2007, pp. 224–283
28.^ Cuozzo & Yamashita 2006, pp. 67–96
29.^ a b c d e f g h Sussman 1999, pp. 154–173
30.^ Gould & Sauther 2007, p. 53
31.^ Sussman, R.W. (1991). "Demography and social organization of free-ranging Lemur catta
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32.^ a b c Mittermeier et al. 2010, Appendix C
33.^ a b c d e f g h i j Cawthon Lang, K.A. (2005-09-21). "Primate Factsheets: Ring-tailed lemur
(Lemur catta) Behavior". Wisconsin Primate Research Center (WPRC).
http://pin.primate.wisc.edu/factsheets/entry/ring-tailed_lemur/behav. Retrieved 2008-09-23.
34.^ Gouzoules & Gouzoules 2007, p. 624
35.^ Rowe 1996, p. 38
36.^ a b Palagi, E.; Dapporto, L.; Borgognini-Tarli, S.M. (2005). "The neglected scent: on the
marking function of urine in Lemur catta" (PDF). Behavioral Ecology and Sociobiology 58: 437–445.
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of design features" (PDF). Annales Zoologici Fennici 43: 280–284. http://www.sekj.org/PDF/anz43-
free/anz43-280.pdf.
38.^ Palagi, E.; Norscia, I. (2005). "American Journal of Physical Anthropology". Multimodal
Signaling in Wild Lemur catta: Economic Design and Territorial Function of Urine Marking 139: 182–
192. doi:10.1002/ajpa.20971.
39.^ a b Macedonia, J.M. (1993). "The Vocal Repertoire of the ring-tailed Lemur ( Lemur catta)".
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04-18.
41.^ Lemur Street. Narrator: Martin Shaw. BBC. Animal Planet. No. 6 ("Home Alone"), series 1.
8:40 minutes in.
42.^ a b c Ehrlich, E.; Fobes, J.L.; King, J.E. (1976). "Prosimian learning capacities". Journal of
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43.^ a b Jolly, A. (1964). "Prosimians' manipulation of simple object problems". Animal
Behaviour 12 (4): 560–570. doi:10.1016/0003-3472(64)90080-6.
44.^ Hosey, G.R. (2000). "A glimpse into the lemur mind" (PDF). Proceedings of the 2nd Annual
Symposium on Zoo Research: 5–10.
http://www.biaza.org.uk/resources/library/images/ARSP2.pdf#page=11.
45.^ Dunbar, R.I.M. (1998). "The Social Brain Hypothesis" (PDF). Evolutionary Anthropology 6
(4): 178–190. doi:10.1002/(SICI)1520-6505(1998)6:5<178::AID-EVAN5>3.0.CO;2-8.
http://eebweb.arizona.edu/Faculty/Dornhaus/courses/materials/papers/Dunbar%20social%20brain
%20size%20group%20review.pdf.
46.^ Jolly, A. (1966). "Lemur Social Behavior and Primate Intelligence". Science 153 (3735):
501–506. doi:10.1126/science.153.3735.501.
47.^ Jolly, A. (1998). "Pair-Bonding, Female Aggression and the Evolution of Lemur Societies".
Folia Primatologica 69 (Suppl. 1): 1–13. doi:10.1159/000052693.
48.^ a b Merritt, D.; MacLean, E.L.; Jaffe, S.; Brannon, E.M. (2007). "A Comparative Analysis of
Serial Ordering in Ring-Tailed Lemurs (Lemur catta)" (PDF). Journal of Comparative Psychology 121
(4): 363–371. doi:10.1037/0735-7036.121.4.363. PMID 18085919. PMC 2953466.
http://www.duke.edu/web/mind/level2/faculty/liz/Publications/Merritt,%20MacLean,%20Jaffe%20and
%20Brannon%20(2007).pdf.
49.^ Santos, L.R.; Barnes, J.L.; Mahajan, N. (2005). "Expectations about numerical events in
four lemur species (Eulemur fulvus, Eulemur mongoz, Lemur catta and Varecia rubra)" (PDF).
Animal Cognition 8 (4): 253–262. doi:10.1007/s10071-005-0252-4. PMID 15729569.
http://www.yale.edu/caplab/Main/Publications_files/santosetal.lemurnumber.pdf.
50.^ Santos, L.R.; Mahajan, N.; Barnes, J.L. (2005). "How Prosimian Primates Represent Tools:
Experiments With Two Lemur Species (Eulemur fulvus and Lemur catta)" (PDF Reprint). Journal of
Comparative Psychology 119 (4): 394–403. doi:10.1037/0735-7036.119.4.394. PMID 16366773.
http://www.yale.edu/caplab/Main/Publications_files/santosetal.lemurtools.jcp.pdf.
51.^ a b c d e f g Cawthon Lang, K.A. (2005-09-21). "Primate Factsheets: Ring-tailed lemur
(Lemur catta) Conservation". Wisconsin Primate Research Center (WPRC).
http://pin.primate.wisc.edu/factsheets/entry/ring-tailed_lemur/cons. Retrieved 2008-09-23.
52.^ Mittermeier et al. 2006, p. 57
53.^ Mittermeier et al. 2010, Chapter 5
54.^ Hans ten Cate (13 June 2002). "John Cleese Visits Lemurs at San Francisco Zoo".
PythOnline's Daily Llama. Archived from the original on 28 December 2010.
http://www.webcitation.org/5vKSV7UX3. Retrieved 28 December 2010.
55.^ a b John Cleese (host). (1998). In the Wild: Operation Lemur with John Cleese . [DVD].
Tigress Productions Ltd for BBC. Archived from the original on 28 December 2010.
http://www.webcitation.org/5vKSlRiBi. Retrieved 28 December 2010.
56.^ Duke University (12 October 1998). "Four More Lemurs To Be Released Into Madagascar
Jungle This Fall". Science Daily. Archived from the original on 28 December 2010.
http://www.webcitation.org/5vKScRzZ5. Retrieved 28 December 2010.
Books cited
• Ankel-Simons, F. (2007). Primate Anatomy (3rd ed.). Academic Press. ISBN 0-12-372576-3.
• Blunt, W.; Stearn, W.T. (2002). Linnaeus: the compleat naturalist. Princeton University Press.
ISBN 978-0691096360. http://books.google.com/?
id=m6lsDLevuJ4C&pg=PA252&dq=Linnaeus+from+his+general+reading+of+Virgil+and+Ovid&q=Lin
naeus%20from%20his%20general%20reading%20of%20Virgil%20and%20Ovid.
• Flower, W.H.; Lydekker, R. (1891). An introduction to the study of mammals living and
extinct. A. and C. Black. p. 682. ISBN 978-1110768578. http://books.google.com/?
id=B3crAAAAYAAJ&pg=PA682&dq=lemurs+called+Linnaeus&q=lemurs%20called%20Linnaeus.
• Garbutt, N. (2007). Mammals of Madagascar, A Complete Guide. A&C Black Publishers.
ISBN 978-0-300-12550-4.
• Campbell, C.; Fuentes, A.; MacKinnon, K. et al., eds (2007). Primates in Perspective. Oxford
University Press. ISBN 978-0-19-517133-4.
• Gould, L.; Sauther, M. (2007). "Lemuriformes". p. 53.
• Gouzoules, H.; Gouzoules, S. (2007). "The Conundrum of Communication". p. 624.
• Gould, L.; Sauther, M.L., eds (2006). Lemurs: Ecology and Adaptation. Springer. ISBN 978-
0387-34585-7. http://books.google.com/?id=nsBtrhsMU5EC&printsec=frontcover&q.
• Cuozzo, F.P.; Yamashita, N. (2006). "Chapter 4: Impact of Ecology on the Teeth of
Extant Lemurs: A Review of Dental Adaptations, Function, and Life History". pp. 67–96.
• Jolly, A.; Sussman, R.W.; Koyama, N. et al., eds (2006). Ringtailed Lemur Biology. Springer.
ISBN 0-387-32669-3.
• Goodman, S.M.; Rakotoarisoa, S.V.; Wilmé, L. (2006). "The Distribution and
Biogeography of the Ringtailed Lemur ( Lemur catta ) in Madagascar". pp. 3–15.
• Mittermeier, R.A.; Louis, E.E.; Richardson, M.; Schwitzer, C.; Langrand, O.; Rylands, A.B.;
Hawkins, F.; Rajaobelina, S. et al. (2010). Lemurs of Madagascar. Illustrated by S.D. Nash (3rd ed.).
Conservation International. ISBN 978-1-934151-23-5.
• Mittermeier, R.A.; Konstant, W.R.; Hawkins, F.; Louis, E.E.; Langrand, O.; Ratsimbazafy, J.;
Rasoloarison, R.; Ganzhorn, J.U. et al. (2006). Lemurs of Madagascar. Illustrated by S.D. Nash (2nd
ed.). Conservation International. ISBN 1-881173-88-7.
• Nield, T. (2007). Supercontinent: Ten Billion Years in the Life of Our Planet . Harvard
University Press. ISBN 978-0674026599. http://books.google.com/?
id=If3wYQtpECkC&pg=PA41&dq=%22The+ring-tailed+lemur+was+first+described+and+named
%22&q=%22The%20ring-tailed%20lemur%20was%20first%20described%20and%20named%22.
• Rowe, N. (1996). The Pictorial Guide to the Living Primates . Pogonias Press. ISBN 0-
9648825-0-7.
• Sussman, R.W. (1999). Primate Ecology and Social Structure . Volume 1: Lorises, Lemurs
and Tarsiers. Pearson Custom Pub.. pp. 154–173. ISBN 0-536-02256-9.
• Tattersall, I. (1982). The Primates of Madagascar. Columbia University Press. pp. 43–46.
ISBN 0-231-04704-5.

Wikimedia Commons has media related to: Lemur catta
Wikispecies has information related to: Lemur
Wikispecies has information related to: Ring-tailed Lemur
• Official Ring-tailed Lemur SSP

• ARKive – images and movies of the Ring-tailed Lemur (Lemur catta)
• Primate Info Net Lemur catta Factsheet
• Duke Lemur Center: Ring-tailed Lemurs
• Lemur Conservation Foundation: Ring-tailed Lemurs
• Ring-tailed Lemur, Lemur catta fact sheet (San Diego Zoo Library)
• ISIS Species Holdings for Lemur catta
• Nature: A Lemur's Tale
• PBS: In the Wild: Operation Lemur with John Cleese
[hide]v · d · eExtant species of family Lemuridae
Kingdom: Animalia · Phylum: Chordata · Class: Mammalia · Order: Primates · Suborder: Strepsirhini
Lemur Ring-tailed Lemur (L. catta)
Black Lemur (E. macaco) · Common Brown Lemur (E. fulvus) · Sanford's
Eulemur Brown Lemur (E. sanfordi) · White-headed Lemur (E. albifrons) · Red-fronted Lemur
(True lemurs) (E. rufifrons) · Red Lemur (E. rufus) · Collared Brown Lemur (E. collaris) · Gray-headed
Lemur (E. cinereiceps) · Mongoose Lemur (E. mongoz) · Crowned Lemur (E.
coronatus) · Red-bellied Lemur (E. rubriventer) · Sclater's Lemur (E. flavifrons)
Varecia
(Ruffed lemurs) Black-and-white Ruffed Lemur (V. variegata) · Red Ruffed Lemur (V. rubra)
Hapalemur Eastern Lesser Bamboo Lemur (H. griseus) · Western Lesser Bamboo Lemur
(Bamboo lemurs) (H. occidentalis) · Southern Lesser Bamboo Lemur (H. meridionalis) · Lac Alaotra
Gentle Lemur (H. alaotrensis) · Golden Bamboo Lemur (H. aureus)
Prolemur Greater Bamboo Lemur (P. simus)
Category
Retrieved from "http://en.wikipedia.org/wiki/Ring-tailed_Lemur"
Categories: IUCN Red List near threatened species | Lemurs | Monotypic mammal genera | Animals
described in 1758
W000
Introduction to evolution

This article is an accessible, non-technical introduction to the subject. For the main encyclopedia
article, see Evolution.
.
Representative sequence but not a "straight-line"
evolution of the horse. Reconstruction, left forefoot
skeleton (third digit emphasized yellow) and cross
section of molar tooth of selected prehistoric
horses.
Overview
Life forms reproduce to make offspring.
The offspring differs from the parent in

minor random ways.
If the differences are helpful, the offspring

is more likely to survive and reproduce.
This means that more offspring in the next

generation will have the helpful difference.
These differences accumulate resulting in

changes within the population.
Over time, populations branch off to

become new species as they become
geographically separated and genetically isolated.
This process is responsible for the many

diverse life forms in the world today.
Haeckel's Paleontological Tree of
Vertebrates (c. 1879).
The evolutionary history of species has been
described as a "tree", with many branches arising
from a single trunk. While Haeckel's tree is
somewhat outdated, it illustrates clearly the
principles that more complex modern
reconstructions can obscure.
Evolution is the process of change in all forms of life over generations, and evolutionary biology is the
study of how evolution occurs.
The biodiversity of life evolves by means of natural selection, mutations and genetic drift. The
process of natural selection is based on three conditions. First, every individual is supplied with hereditary
material in the form of genes that are received from their parents then passed on to their offspring. Second,
organisms tend to produce more offspring than the environment can support. Third, there are variations
among offspring as a consequence of either the random introduction of new genes via mutations or
reshuffling of existing genes during sexual reproduction.[1][2][3]
Natural selection will occur when these facts of nature (heredity, overproduction of offspring, and
variation) hold true. Natural selection means individuals do not have equal chances of reproductive success.
As a consequence, some individuals produce more offspring and thus have a higher degree of fitness. Traits
that ensure organisms are better adapted to their living conditions become more common in descendant
populations.[2][3] For this reason, populations will never remain exactly the same over successive
generations. The forces of evolution are most evident when populations become isolated, either through
geographic distance or by mechanisms that prevent genetic exchange. Over time, isolated populations can
branch off into new species.[4][5]
Random genetic drift describes another natural process that regulates the evolution of minor
mutations in the genes, leading to changes in allele frequencies over time. These smaller mutations occur
with regular frequency in and among populations. The vast majority of genetic mutations neither assist,
change the appearance of, nor bring harm to individuals. These mutated genes are neutrally sorted among
populations and survive across generations by chance alone. When species migrate they carry different
genetic varieties to different places. When organisms mate they exchange genetic material and new
individuals are born.
The outward expression of each unique genetic mixture in different environments, the phenotype,
comprises a diversity of traits that can be measured and observed as individuals grow and develop. In
contrast to genetic drift, natural selection is not a random process because it acts on traits that become
adaptive for their functional utilities that are necessary for survival.[6] Natural selection and random genetic
drift are forever constant and dynamic parts of life. More than 99.9% of all species have become extinct since
life began over 3.5 billion years ago. Evolution is more death than survival and over time this has shaped the
branching structure in the tree of life.[7]
The modern understanding of evolution began with the 1859 publication of Charles Darwin's On the
Origin of Species. In addition, Gregor Mendel's work with plants helped to explain the hereditary patterns of
genetics.[8] Fossil discoveries in paleontology, advances in population genetics and a global network of
scientific research have provided further details into the mechanisms of evolution. Scientists now have a
good understanding of the origin of new species (speciation) and have observed the speciation process in
the laboratory and in the wild. Evolution is the principal theory that biologists use to understand life and is
used in many disciplines, including medicine, psychology, conservation biology, anthropology, forensics,
agriculture and other social-cultural applications.
Contents
[hide]
• 1 Darwin's idea: evolution by natural selection
• 2 Source of variation
• 3 Modern synthesis
• 4 Evidence for evolution
• 4.2 Comparative anatomy
• 4.2.1 Taxonomy
• 4.2.2 Embryology
• 4.2.3 Vestigial structures
• 4.2.4 Convergent evolution
• 4.3 Molecular biology
• 4.4 Co-evolution
• 4.5 Artificial selection
• 5 Species
• 6 Different views on the mechanism of evolution
• 6.1 Rate of change
• 6.2 Unit of change
• 7 Summary
• 8 See also
• 9 Notes
[edit] Darwin's idea: evolution by natural selection
Further information: Common descent
In the 19th century, natural history collections and museums were a popular pastime. The European
expansion and naval expeditions employed naturalists and curators of grand museums showcasing
preserved and live specimens of the varieties of life. Charles Darwin was an English graduate who was
educated and trained in the disciplines of natural history science. Such natural historians would collect,
catalogue, describe and study the vast collections of specimens stored and managed by curators at these
museums. Charles Darwin served as a ship's naturalist on board the HMS Beagle, assigned to a five-year
research expedition around the world. During his voyage, Darwin observed and collected an abundance of
organisms, being very interested in the diverse forms of life along the coasts of South America and the
neighboring Galapagos Islands.[9][10]
Charles Darwin gained extensive experience as he collected and studied the natural history of life
forms from distant places. Through his studies, Darwin formulated the idea that each species had developed
from ancestors with similar features. In 1838, he described how a process he called natural selection would
make this happen.[11]
Darwin's idea of how evolution works relied on the following observations:[12]
• If all the individuals of a species reproduced successfully, the population of that species
would increase uncontrollably.
• Populations tend to remain about the same size from year to year.
• Environmental resources are limited.
• No two individuals in a given species are exactly alike.
• Much of this variation in a population can be passed on to offspring.
Charles Darwin proposed the theory of evolution by natural selection.
Darwin noted that orchids exhibited a variety of complex adaptations to ensure pollination; all derived
from basic floral parts.
Darwin deduced that since organisms produce more offspring than their environment could possibly
support, there must be a competitive struggle for survival—only a few individuals can survive out of each
generation. Darwin realized that it was not chance alone that determined survival. Instead, survival depends
on the traits of each individual and if these traits aid or hinder survival and reproduction. Well-adapted, or
"fit", individuals are likely to leave more offspring than their less well-adapted competitors. Darwin realized
that the unequal ability of individuals to survive and reproduce could cause gradual changes in the
population. Traits that help an organism survive and reproduce would accumulate over generations. On the
other hand, traits that hinder survival and reproduction would disappear. Darwin used the term natural
selection to describe this process.[13]
Natural selection is commonly equated with survival of the fittest, but this expression originated in
Herbert Spencer's Principles of Biology in 1864, after Charles Darwin published his original works. Survival
of the fittest describes the process of natural selection incorrectly, because natural selection is not only about
survival and it is not always the fittest that survives.[14]
Observations of variations in animals and plants formed the basis of the theory of natural selection.
For example, Darwin observed that orchids and insects have a close relationship that allows the pollination of
the plants. He noted that orchids have a variety of structures that attract insects, so that pollen from the
flowers gets stuck to the insects’ bodies. In this way, insects transport the pollen from a male to a female
orchid. In spite of the elaborate appearance of orchids, these specialized parts are made from the same
basic structures that make up other flowers. In Fertilisation of Orchids Darwin proposed that the orchid
flowers did not represent the work of an ideal engineer, but were adapted from pre-existing parts, through
natural selection.[15]
Darwin was still researching and experimenting with his ideas on natural selection when he received
a letter from Alfred Wallace describing a theory very similar to his own. This led to an immediate joint
publication of both theories. Both Wallace and Darwin saw the history of life like a family tree, with each fork
in the tree’s limbs being a common ancestor. The tips of the limbs represented modern species and the
branches represented the common ancestors that are shared amongst many different species. To explain
these relationships, Darwin said that all living things were related, and this meant that all life must be
descended from a few forms, or even from a single common ancestor. He called this process descent with
modification.[12]
Darwin published his theory of evolution by natural selection in On the Origin of Species in 1859. His
theory means that all life, including humanity, is a product of continuing natural processes. The implication
that all life on Earth has a common ancestor has met with objections from some religious groups who believe
even today that the different types of life are due to special creation.[16] Their objections are in contrast to
the level of support for the theory by more than 99 percent of those within the scientific community today.[17]
[edit] Source of variation

Darwin’s theory of natural selection laid the groundwork for modern evolutionary theory, and his
experiments and observations showed that the organisms in populations varied from each other, that some
of these variations were inherited, and that these differences could be acted on by natural selection.
However, he could not explain the source of these variations. Like many of his predecessors, Darwin
mistakenly thought that heritable traits were a product of use and disuse, and that features acquired during
an organism's lifetime could be passed on to its offspring. He looked for examples, such as large ground
feeding birds getting stronger legs through exercise, and weaker wings from not flying until, like the ostrich,
they could not fly at all.[18] This misunderstanding was called the inheritance of acquired characters and was
part of the theory of transmutation of species put forward in 1809 by Jean-Baptiste Lamarck. In the late 19th
century this theory became known as Lamarckism. Darwin produced an unsuccessful theory he called
pangenesis to try to explain how acquired characteristics could be inherited. In the 1880s August
Weismann's experiments indicated that changes from use and disuse could not be inherited, and
Lamarckism gradually fell from favor.[19]
The missing information needed to help explain how new features could pass from a parent to its
offspring was provided by the pioneering genetics work of Gregor Mendel. Mendel’s experiments with several
generations of pea plants demonstrated that inheritance works by separating and reshuffling hereditary
information during the formation of sex cells and recombining that information during fertilization. This is like
mixing different hands of cards, with an organism getting a random mix of half of the cards from one parent,
and half of the cards from the other. Mendel called the information factors; however, they later became
known as genes. Genes are the basic units of heredity in living organisms. They contain the information that
directs the physical development and behavior of organisms.
Genes are made of DNA, a long molecule that carries information. This information is encoded in the
sequence of nucleotides in the DNA, just as the sequence of the letters in words carries information on a
page. The genes are like short instructions built up of the "letters" of the DNA alphabet. Put together, the
entire set of these genes gives enough information to serve as an "instruction manual" of how to build and
run an organism. The instructions spelled out by this DNA alphabet can be changed, however, by mutations,
and this may alter the instructions carried within the genes. Within the cell, the genes are carried in
chromosomes, which are packages for carrying the DNA, with the genes arranged along them like beads on
a string. It is the reshuffling of the chromosomes that results in unique combinations of genes in offspring.
Although such mutations in DNA are random, natural selection is not a process of chance: the
environment determines the probability of reproductive success. The end products of natural selection are
organisms that are adapted to their present environments. Natural selection does not involve progress
towards an ultimate goal. Evolution does not strive for more advanced, more intelligent, or more
sophisticated life forms.[20] For example, fleas (wingless parasites) are descended from a winged, ancestral
scorpionfly, and snakes are lizards that no longer require limbs - although pythons still grow tiny structures
that are the remains of their ancestor's hind legs.[21][22] Organisms are merely the outcome of variations
that succeed or fail, dependent upon the environmental conditions at the time.
Rapid environmental changes typically cause extinctions.[23] Of all species that have existed on
Earth, 99.9 percent are now extinct.[24] Since life began on Earth, five major mass extinctions have led to
large and sudden drops in the variety of species. The most recent, the Cretaceous–Tertiary extinction event,
occurred 65 million years ago, and has attracted more attention than all others because it killed the
dinosaurs.[25]
[edit] Modern synthesis
Further information: Modern evolutionary synthesis
The modern evolutionary synthesis was the outcome of a merger of several different scientific fields
into a cohesive understanding of evolutionary theory. In the 1930s and 1940s, efforts were made to merge
Darwin's theory of natural selection, research in heredity, and understandings of the fossil records into a
unified explanatory model.[26] The application of the principles of genetics to naturally occurring populations,
by scientists such as Theodosius Dobzhansky and Ernst Mayr, advanced understanding of the processes of
evolution. Dobzhansky's 1937 work Genetics and the Origin of Species was an important step in bridging the
gap between genetics and field biology. Mayr, on the basis of an understanding of genes and direct
observations of evolutionary processes from field research, introduced the biological species concept, which
defined a species as a group of interbreeding or potentially interbreeding populations that are reproductively
isolated from all other populations. The paleontologist George Gaylord Simpson helped to incorporate fossil
research, which showed a pattern consistent with the branching and non-directional pathway of evolution of
organisms predicted by the modern synthesis.
The modern synthesis emphasizes the importance of populations as the unit of evolution, the central
role of natural selection as the most important mechanism of evolution, and the idea of gradualism to explain
how large changes evolve as an accumulation of small changes over long periods of time.
[edit] Evidence for evolution
During the voyage of the Beagle, naturalist Charles Darwin collected fossils in South America, and
found fragments of armor which he thought were like giant versions of the scales on the modern armadillos
living nearby. On his return, the anatomist Richard Owen showed him that the fragments were from gigantic
extinct glyptodons, related to the armadillos. This was one of the patterns of distribution that helped Darwin to
develop his theory.[11]
Further information: Evidence of common descent
Scientific evidence for evolution comes from many aspects of biology, and includes fossils,
homologous structures, and molecular similarities between species' DNA.

Research in the field of paleontology, the study of fossils, supports the idea that all living organisms
are related. Fossils provide evidence that accumulated changes in organisms over long periods of time have
led to the diverse forms of life we see today. A fossil itself reveals the organism's structure and the
relationships between present and extinct species, allowing paleontologists to construct a family tree for all of
the life forms on Earth.[27]
Modern paleontology began with the work of Georges Cuvier (1769–1832). Cuvier noted that, in
sedimentary rock, each layer contained a specific group of fossils. The deeper layers, which he proposed to
be older, contained simpler life forms. He noted that many forms of life from the past are no longer present
today. One of Cuvier’s successful contributions to the understanding of the fossil record was establishing
extinction as a fact. In an attempt to explain extinction, Cuvier proposed the idea of “revolutions” or
catastrophism in which he speculated that geological catastrophes had occurred throughout the Earth’s
history, wiping out large numbers of species.[28] Cuvier's theory of revolutions was later replaced by
uniformitarian theories, notably those of James Hutton and Charles Lyell who proposed that the Earth’s
geological changes were gradual and consistent.[29] However, current evidence in the fossil record supports
the concept of mass extinctions. As a result, the general idea of catastrophism has re-emerged as a valid
hypothesis for at least some of the rapid changes in life forms that appear in the fossil records.
A very large number of fossils have now been discovered and identified. These fossils serve as a
chronological record of evolution. The fossil record provides examples of transitional species that
demonstrate ancestral links between past and present life forms.[30] One such transitional fossil is
Archaeopteryx, an ancient organism that had the distinct characteristics of a reptile (such as a long, bony tail
and conical teeth) yet also had characteristics of birds (such as feathers and a wishbone). The implication
from such a find is that modern reptiles and birds arose from a common ancestor.[31]
[edit] Comparative anatomy

Further information: Convergent evolution and Divergent evolution
The comparison of similarities between organisms of their form or appearance of parts, called their
morphology, has long been a way to classify life into closely related groups. This can be done by comparing
the structure of adult organisms in different species or by comparing the patterns of how cells grow, divide
and even migrate during an organism's development.
[edit] Taxonomy
Taxonomy is the branch of biology that names and classifies all living things. Scientists use
morphological and genetic similarities to assist them in categorizing life forms based on ancestral
relationships. For example, orangutans, gorillas, chimpanzees, and humans all belong to the same
taxonomic grouping referred to as a family – in this case the family called Hominidae. These animals are
grouped together because of similarities in morphology that come from common ancestry (called homology).
[32]
Strong evidence for evolution comes from the analysis of homologous structures: structures in
different species that no longer perform the same task but which share a similar structure.[33] Such is the
case of the forelimbs of mammals. The forelimbs of a human, cat, whale, and bat all have strikingly similar
bone structures. However, each of these four species' forelimbs performs a different task. The same bones
that construct a bat's wings, which are used for flight, also construct a whale's flippers, which are used for
swimming. Such a "design" makes little sense if they are unrelated and uniquely constructed for their
particular tasks. The theory of evolution explains these homologous structures: all four animals shared a
common ancestor, and each has undergone change over many generations. These changes in structure
have produced forelimbs adapted for different tasks.[34]
[edit] Embryology
In some cases, anatomical comparison of structures in the embryos of two or more species provides
evidence for a shared ancestor that may not be obvious in the adult forms. As the embryo develops, these
homologies can be lost to view, and the structures can take on different functions. Part of the basis of
classifying the vertebrate group (which includes humans), is the presence of a tail (extending beyond the
anus) and pharyngeal slits. Both structures appear during some stage of embryonic development but are not
always obvious in the adult form.[35]
Because of the morphological similarities present in embryos of different species during
development, it was once assumed that organisms re-enact their evolutionary history as an embryo. It was
thought that human embryos passed through an amphibian then a reptilian stage before completing their
development as mammals. Such a re-enactment, (often called Recapitulation theory), is not supported by
scientific evidence. What does occur, however, is that the first stages of development are similar in broad
groups of organisms.[36] At very early stages, for instance, all vertebrates appear extremely similar, but do
not exactly resemble any ancestral species. As development continues, specific features emerge from this
basic pattern.
[edit] Vestigial structures
Homology includes a unique group of shared structures referred to as vestigial structures. Vestigial
refers to anatomical parts that are of minimal, if any, value to the organism that possesses them. These
apparently illogical structures are remnants of organs that played an important role in ancestral forms. Such
is the case in whales, which have small vestigial bones that appear to be remnants of the leg bones of their
ancestors which walked on land.[37] Humans also have vestigial structures, including the ear muscles, the
wisdom teeth, the appendix, the tail bone, body hair (including goose bumps), and the semilunar fold in the
corner of the eye.[38]
[edit] Convergent evolution
The bird and the bat wing are examples of convergent evolution.
A bat is a mammal and its forearm bones have been adapted for flight.
Anatomical comparisons can be misleading, as not all anatomical similarities indicate a close
relationship. Organisms that share similar environments will often develop similar physical features, a
process known as convergent evolution. Both sharks and dolphins have similar body forms, yet are only
distantly related – sharks are fish and dolphins are mammals. Such similarities are a result of both
populations being exposed to the same selective pressures. Within both groups, changes that aid swimming
have been favored. Thus, over time, they developed similar appearances (morphology), even though they
are not closely related.[39]
[edit] Molecular biology
A section of DNA
Every living organism (with the possible exception of RNA viruses) contains molecules of DNA,
which carries genetic information. Genes are the pieces of DNA that carry this information, and they influence
the properties of an organism. Genes determine an individual's general appearance and to some extent their
behavior. If two organisms are closely related, their DNA will be very similar.[40] On the other hand, the more
distantly related two organisms are, the more differences they will have. For example, brothers are closely
related and have very similar DNA, while cousins share a more distant relationship and have far more
differences in their DNA. Similarities in DNA are used to determine the relationships between species in
much the same manner as they are used to show relationships between individuals. For example, comparing
chimpanzees with gorillas and humans shows that there is as much as a 96 percent similarity between the
DNA of humans and chimps. Comparisons of DNA indicate that humans and chimpanzees are more closely
related to each other than either species is to gorillas.[41][42]
The field of molecular systematics focuses on measuring the similarities in these molecules and
using this information to work out how different types of organisms are related through evolution. These
comparisons have allowed biologists to build a relationship tree of the evolution of life on Earth.[43] They
have even allowed scientists to unravel the relationships between organisms whose common ancestors lived
such a long time ago that no real similarities remain in the appearance of the organisms.
[edit] Co-evolution
Co-evolution is a process in which two or more species influence the evolution of each other. All
organisms are influenced by life around them; however, in co-evolution there is evidence that genetically
determined traits in each species directly resulted from the interaction between the two organisms.[40]
An extensively documented case of co-evolution is the relationship between Pseudomyrmex, a type
of ant, and the acacia, a plant that the ant uses for food and shelter. The relationship between the two is so
intimate that it has led to the evolution of special structures and behaviors in both organisms. The ant
defends the acacia against herbivores and clears the forest floor of the seeds from competing plants. In
response, the plant has evolved swollen thorns that the ants use as shelter and special flower parts that the
ants eat.[44] Such co-evolution does not imply that the ants and the tree choose to behave in an altruistic
manner. Rather, across a population small genetic changes in both ant and tree benefited each. The benefit
gave a slightly higher chance of the characteristic being passed on to the next generation. Over time,
successive mutations created the relationship we observe today.
[edit] Artificial selection
The results of artificial selection: a Chihuahua mix and a Great Dane.

Artificial selection is the controlled breeding of domestic plants and animals. Humans determine
which animal or plant will reproduce and which of the offspring will survive; thus, they determine which genes
will be passed on to future generations. The process of artificial selection has had a significant impact on the
evolution of domestic animals. For example, people have produced different types of dogs by controlled
breeding. The differences in size between the Chihuahua and the Great Dane are the result of artificial
selection. Despite their dramatically different physical appearance, they and all other dogs evolved from a
few wolves domesticated by humans in what is now China less than 15,000 years ago.[45]
Artificial selection has produced a wide variety of plants. In the case of maize (corn), recent genetic
evidence suggests that domestication occurred 10,000 years ago in central Mexico.[46] Prior to
domestication, the edible portion of the wild form was small and difficult to collect. Today The Maize Genetics
Cooperation • Stock Center maintains a collection of more than 10,000 genetic variations of maize that have
arisen by random mutations and chromosomal variations from the original wild type.[47]
In artificial selection the new breed or variety that emerges is the one with random mutations
attractive to humans, while in natural selection the surviving species is the one with random mutations useful
to it in its non-human environment. In both natural and artificial selection the variations are a result of random
mutations, and the underlying genetic processes are essentially the same.[48] Darwin carefully observed the
outcomes of artificial selection in animals and plants to form many of his arguments in support of natural
selection.[49] Much of his book On the Origin of Species was based on these observations of the many
varieties of domestic pigeons arising from artificial selection. Darwin proposed that if humans could achieve
dramatic changes in domestic animals in short periods, then natural selection, given millions of years, could
produce the differences seen in living things today.
[edit] Species
Further information: Species, Speciation, and Phylogenetics
There are numerous species of cichlids that demonstrate dramatic variations in morphology.
Given the right circumstances, and enough time, evolution leads to the emergence of new species.
Scientists have struggled to find a precise and all-inclusive definition of species. Ernst Mayr (1904–2005)
defined a species as a population or group of populations whose members have the potential to interbreed
naturally with one another to produce viable, fertile offspring. (The members of a species cannot produce
viable, fertile offspring with members of other species).[50] Mayr's definition has gained wide acceptance
among biologists, but does not apply to organisms such as bacteria, which reproduce asexually.
Speciation is the lineage-splitting event that results in two separate species forming from a single
common ancestral population.[13] A widely accepted method of speciation is called allopatric speciation.
Allopatric speciation begins when a population becomes geographically separated.[33] Geological
processes, such as the emergence of mountain ranges, the formation of canyons, or the flooding of land
bridges by changes in sea level may result in separate populations. For speciation to occur, separation must
be substantial, so that genetic exchange between the two populations is completely disrupted. In their
separate environments, the genetically isolated groups follow their own unique evolutionary pathways. Each
group will accumulate different mutations as well as be subjected to different selective pressures. The
accumulated genetic changes may result in separated populations that can no longer interbreed if they are
reunited.[13] Barriers that prevent interbreeding are either prezygotic (prevent mating or fertilization) or
postzygotic (barriers that occur after fertilization). If interbreeding is no longer possible, then they will be
considered different species.[51]
Usually the process of speciation is slow, occurring over very long time spans; thus direct
observations within human life-spans are rare. However speciation has been observed in present day
organisms, and past speciation events are recorded in fossils.[52][53][54] Scientists have documented the
formation of five new species of cichlid fishes from a single common ancestor that was isolated fewer than
5000 years ago from the parent stock in Lake Nagubago.[55] The evidence for speciation in this case was
morphology (physical appearance) and lack of natural interbreeding. These fish have complex mating rituals
and a variety of colorations; the slight modifications introduced in the new species have changed the mate
selection process and the five forms that arose could not be convinced to interbreed.[56]
[edit] Different views on the mechanism of evolution
James Hutton
Stephen Jay Gould
Richard Dawkins
The theory of evolution is widely accepted among the scientific community, serving to link the diverse
specialty areas of biology.[17] Evolution provides the field of biology with a solid scientific base. The
significance of evolutionary theory is best described by the title of a paper by Theodosius Dobzhansky (1900–
1975), published in American Biology Teacher; "Nothing in Biology Makes Sense Except in the Light of
Evolution".[57] Nevertheless, the theory of evolution is not static. There is much discussion within the
scientific community concerning the mechanisms behind the evolutionary process. For example, the rate at
which evolution occurs is still under discussion. In addition, there are conflicting opinions as to which is the
primary unit of evolutionary change – the organism or the gene.
[edit] Rate of change
Two views exist concerning the rate of evolutionary change. Darwin and his contemporaries viewed
evolution as a slow and gradual process. Evolutionary trees are based on the idea that profound differences
in species are the result of many small changes that accumulate over long periods.
The view that evolution is gradual had its basis in the works of the geologist James Hutton (1726–
1797) and his theory called "gradualism". Hutton's theory suggests that profound geological change was the
cumulative product of a relatively slow continuing operation of processes which can still be seen in operation
today, as opposed to catastrophism which promoted the idea that sudden changes had causes which can no
longer be seen at work. A uniformitarian perspective was adopted for biological changes. Such a view can
seem to contradict the fossil record, which shows evidence of new species appearing suddenly, then
persisting in that form for long periods. The paleontologist Stephen Jay Gould (1941–2002) developed a
model that suggests that evolution, although a slow process in human terms, undergoes periods of relatively
rapid change over only a few thousand or million years, alternating with long periods of relative stability, a
model called "punctuated equilibrium" which explains the fossil record without contradicting Darwin's ideas.
[58]
[edit] Unit of change

A common unit of selection in evolution is the organism. Natural selection occurs when the
reproductive success of an individual is improved or reduced by an inherited characteristic, and reproductive
success is measured by the number of an individual's surviving offspring. The organism view has been
challenged by a variety of biologists as well as philosophers. Richard Dawkins (born 1941) proposes that
much insight can be gained if we look at evolution from the gene's point of view; that is, that natural selection
operates as an evolutionary mechanism on genes as well as organisms.[59] In his 1976 book The Selfish
Gene, he explains:
“ Individuals are not stable things, they are fleeting. Chromosomes too are
shuffled to oblivion, like hands of cards soon after they are dealt. But the cards
themselves survive the shuffling. The cards are the genes. The genes are not destroyed
by crossing-over; they merely change partners and march on. Of course they march on.
That is their business. They are the replicators and we are their survival machines.
When we have served our purpose we are cast aside. But genes are denizens of
geological time: genes are forever.[60] ”
Others view selection working on many levels, not just at a single level of organism or gene; for
example, Stephen Jay Gould called for a hierarchical perspective on selection.[61]
[edit] Summary
Several basic observations establish the theory of evolution, which explains the variety and
relationship of all living things. There are genetic variations within a population of individuals. Some
individuals, by chance, have features that allow them to survive and thrive better than their kind. The
individuals that survive will be more likely to have offspring of their own. The offspring might inherit the useful
feature.
Evolution is not a random process. While mutations are random, natural selection is not. Evolution is
an inevitable result of imperfectly copying, self-replicating organisms reproducing over billions of years under
the selective pressure of the environment. The outcome of evolution is not a perfectly designed organism.
The outcome is simply an individual that can survive better and reproduce more successfully than its
neighbors in a particular environment. Fossils, the genetic code, and the peculiar distribution of life on Earth
provide a record of evolution and demonstrate the common ancestry of all organisms, both living and long
dead. Evolution can be directly observed in artificial selection, the selective breeding for certain traits of
domestic animals and plants. The diverse breeds of cats, dogs, horses, and agricultural plants serve as
examples of evolution.
Although some groups raise objections to the theory of evolution, the evidence of observation and
experiments that has been gathered for over a hundred years by thousands of scientists supports evolution.
[16] The result of four billion years of evolution is the diversity of life around us, with an estimated 1.75 million
different species in existence today.[5][62]
[edit] See also

Book:Evolution

• Creation-evolution controversy
• Evidence of common descent
• Evolution as theory and fact
• Level of support for evolution
• Misconceptions about evolution
[edit] Notes
1. ^ Darwin, Charles (1859). On the Origin of Species (1st ed.). London: John Murray. p. 1.
http://darwin-online.org.uk/content/frameset?itemID=F373&viewtype=text&pageseq=16. . Related
earlier ideas were acknowledged in Darwin, Charles (1861). On the Origin of Species (3rd ed.).
London: John Murray. xiii. http://darwin-online.org.uk/content/frameset?
2. ^ a b Gould, Stephen J. (2002). The Structure of Evolutionary Theory. Harvard University
Press. pp. 1433. ISBN 0674006135, 9780674006133.
3. ^ a b Gregory, T. R. (2009). "Understanding Natural Selection: Essential Concepts and
Common Misconceptions". Evolution: Education and Outreach 2 (2): 156–175. doi:10.1007/s12052-
009-0128-1. http://www.springerlink.com/content/2331741806807x22/.
4. ^ "An introduction to evolution" (web resource). Understanding Evolution: your one-stop
source for information on evolution. The University of California Museum of Paleontology, Berkeley.
2008. http://evolution.berkeley.edu/evolibrary/article/0_0_0/evo_02. Retrieved 2008-01-23
5. ^ a b Cavalier-Smith T (2006). "Cell evolution and Earth history: stasis and revolution". Philos
PMC 1578732. http://www.ncbi.nlm.nih.gov/pubmed/16754610. Retrieved 2008-01-24.
6. ^ Garvin-Doxas, K.; Klymkowsky, M. W. (2008). "Understanding Randomness and its Impact
on Student Learning: Lessons Learned from Building the Biology Concept Inventory (BCI)". CBE Life
Sci Educ. 7 (2): 227–233. doi:10.1187/cbe.07-08-0063. PMID 18519614. PMC 2424310.
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2424310/.
7. ^ Raup, D. M. (1992). Extinction: bad genes or bad luck.. New York, W. W.: Norton and Co..
pp. 210. ISBN 978-0393309270. http://books.google.com/?
id=8klou91MwJoC&printsec=frontcover&dq=Extinction:
+bad+genes+or+bad+luck&cd=1#v=onepage&q=.
8. ^ Rhee, Sue Yon (1999). "Gregor Mendel". Access Excellence. National Health Museum.
http://www.accessexcellence.org/RC/AB/BC/Gregor_Mendel.html. Retrieved 2008-01-05.
9. ^ Farber, P. L. (2000). Finding Order in Nature: The Natualist Tradition from Linnaeus to E.
O. Wilson. Baltimore and London: The Johns Hopkins University Press. pp. 136. ISBN 0-8018-6389-
9. http://books.google.com/?id=tyG4pfKJ8WEC&printsec=frontcover&dq=Finding+Order+in+Nature:
+The+Natualist+Tradition+from+Linnaeus+to+E.+O.+Wilson&cd=1#v=onepage&q=.
10.^ Watson, J. D. (2005). Darwin, The Indelible Stamp: The Evolution of an Idea . Philadelphia
and London: Running Press. pp. 1257. ISBN 13-978-0-7624-2136-7. http://books.google.com/?
id=LEWfWf0mUJIC&printsec=frontcover&dq=Darwin:+The+Indelible+Stamp:
+The+Evolution+of+an+Idea&cd=1#v=onepage&q=.
11.^ a b Eldredge, Niles (Spring 2006). "Confessions of a Darwinist". The Virginia Quarterly
Review: 32–53. http://www.vqronline.org/articles/2006/spring/eldredge-confessions-darwinist/.
12.^ a b Wyhe, John van (2002). "Charles Darwin: gentleman naturalist". The Complete Work of
Charles Darwin Online. University of Cambridge. http://darwin-online.org.uk/darwin.html. Retrieved
2008-01-16.
13.^ a b c Quammen, David (2004). "Was Darwin Wrong?". National Geographic Magazine.
National Geographic. http://ngm.nationalgeographic.com/ngm/0411/feature1/fulltext.html#top.
14.^ Futuyma, D. J. (2005). The Nature of Natural Selection. Ch. 8, pages 93-98 in Cracraft, J.
and Bybee R. W. (Eds.) Evolutionary Science and Society: Educating a New Generation. American
Institute of Biological Sciences.
15.^ Wyhe, John van (2002). "Fertilisation of Orchids". The Complete Works of Charles Darwin.
University of Cambridge. http://darwin-
online.org.uk/EditorialIntroductions/Freeman_FertilisationofOrchids.html. Retrieved 2008-01-07.
16.^ a b DeVries A (2004). "The enigma of Darwin". Clio Med 19 (2): 136–55. PMID 6085987.
17.^ a b Delgado, Cynthia (2006). "Finding the Evolution in Medicine". NIH Record (National
Institutes of Health). http://nihrecord.od.nih.gov/newsletters/2006/07_28_2006/story03.htm.
18.^ (Darwin 1872, p. 108.) Effects of the increased Use and Disuse of Parts, as controlled by
Natural Selection
19.^ Ghiselin, Michael T. (September/October 1994). "Nonsense in schoolbooks: 'The Imaginary
Lamarck'". The Textbook Letter. The Textbook League. http://www.textbookleague.org/54marck.htm.
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and the Evolution of Animal Design (2nd ed.). Oxford: Blackwell Publishing. ISBN 1-4051-1950-0
• Darwin, Charles (1872). The Origin of Species (6th ed.). London: John Murray. http://darwin-
online.org.uk/content/frameset?itemID=F391&viewtype=text&pageseq=1
• Dawkins, Richard (1976). The Selfish Gene (1st ed.). Oxford University Press. pp. 33.
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Ed
• Diamond, Jared (1992). The Third Chimpanzee: the evolution and future of the human
animal. New York: HarperCollins. ISBN 0060183071
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New York: W.W, Norton & Company. ISBN 0393308197
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ISBN 0517703939
• Lyell, Charles (1830). Principles of geology. New York: Penguin Books. ISBN 014043528X.
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• Mayr, Ernst (1970). Populations, Species, and Evolution . Cambridge, MA: Belknap Press of
Harvard University Press. ISBN 0674690109
• Mayr, Ernst (2001). What evolution is. New York: Basic Books. ISBN 0-465-04425-5
• Tattersall, Ian (1995). The Fossil Trail: How We Know What We Think We Know About
Human Evolution. New York: Oxford University Press. ISBN 0195061012
• Weichert, Charles; Presch, William (1975). Elements of Chordate Anatomy. New York:
McGraw-Hill. ISBN 0070690081
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from and where we're going. [DVD]. South Burlington, VT: WGBH Boston / PBS television series
Nova. ASIN B00005RG6J. http://www.pbs.org/wgbh/evolution/index.html. Retrieved 2008-01-24. -
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2010-10-30
[edit] Related information

Processes of
Population genetic
Evolutionary
developmental
concepts

Modes of

Retrieved from "http://en.wikipedia.org/wiki/Introduction_to_evolution"
Categories: Biological evolution | Biology theories

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Evolution
Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation
Introduction
Evidence
History
Level of support
Modern synthesis
Social effect
Theory and fact
Cladistics
Ecological genetics
Molecular evolution
Phylogenetics
Population genetics
Systematics
A large body of evidence of common descent of living things has been discovered by scientists
working in a variety of fields over many years. This evidence has demonstrated and verified the occurrence
of evolution and provided a wealth of information on the natural processes by which the variety and diversity
of life on Earth developed. This evidence supports the modern evolutionary synthesis, the current scientific
theory that explains how and why life changes over time. Evolutionary biologists document the fact of
common descent: making testable predictions, testing hypotheses, and developing theories that illustrate and
describe its causes.
Comparison of the genetic sequence of organisms has revealed that organisms that are
phylogenetically close have a higher degree of sequence similarity than organisms that are phylogenetically
distant. Further evidence for common descent comes from genetic detritus such as pseudogenes, regions of
DNA that are orthologous to a gene in a related organism, but are no longer active and appear to be
undergoing a steady process of degeneration.
Fossils are important for estimating when various lineages developed in geologic time. As
fossilization is an uncommon occurrence, usually requiring hard body parts and death near a site where
sediments are being deposited, the fossil record only provides sparse and intermittent information about the
evolution of life. Evidence of organisms prior to the development of hard body parts such as shells, bones
and teeth is especially scarce, but exists in the form of ancient microfossils, as well as impressions of various
soft-bodied organisms. The comparative study of the anatomy of groups of animals reveals structural
features that are fundamentally similar or homologous, clearly demonstrating phylogenetic and ancestral
relationships with other organism, most especially when compared with fossils of ancient extinct organisms.
Vestigial structures and comparisons in embryonic development are largely a contributing factor in
anatomical resemblance in concordance with common descent. Since metabolic processes do not leave
fossils, research into the evolution of the basic cellular processes is done largely by comparison of existing
organisms’ physiology and biochemistry. Many lineages diverged at different stages of development, so it is
possible to determine when certain metabolic processes appeared by comparing the traits of the
descendants of a common ancestor. Universal biochemical organization and molecular variance patterns in
all organisms also show a direct correlation with common descent.
Further evidence comes from the field of biogeography because evolution with common descent
provides the best and most thorough explanation for a variety of facts concerning the geographical
distribution of plants and animals across the world. This is especially obvious in the field of island
biogeography. Combined with the theory of plate tectonics common descent provides a way to combine facts
about the current distribution of species with evidence from the fossil record to provide a logically consistent
explanation of how the distribution of living organisms has changed over time.
The development and spread of antibiotic resistant bacteria, like the spread of pesticide resistant
forms of plants and insects provides evidence that evolution due to natural selection is an ongoing process in
the natural world. Alongside this, are observed instances of the separation of populations of species into sets
of new species (speciation). Speciation has been observed directly and indirectly in the lab and in nature.
Multiple forms of such have been described and documented as examples for individual modes of speciation.
Furthermore, evidence of common descent extends from direct laboratory experimentation with the artificial
selection of organisms—historically and currently—and other controlled experiments involving many of the
topics in the article. This article explains the different types of evidence for evolution with common descent
along with many specialized examples of each.
Contents
[hide]
• 1 Evidence from comparative physiology and biochemistry
• 1.1 Genetics
• 1.1.1 Universal biochemical organisation
and molecular variance patterns
• 1.1.2 DNA sequencing
• 1.1.3 Proteins
• 1.1.4 Pseudogenes
• 1.1.5 Other mechanisms
• 1.2 Specific examples
• 1.2.1 Chromosome 2 in humans
• 1.2.2 Cytochrome c
• 1.2.3 Recent African origin of modern
humans
• 2 Evidence from comparative anatomy
• 2.1 Nested hierarchies and classification
• 2.2 Homologous structures and divergent (adaptive)
evolution
• 2.3 Vestigial structures
• 2.4 Evolutionary developmental biology and
embryonic development
[edit] Evidence from comparative physiology and biochemistry
See also: Archaeogenetics, Common descent, Last universal ancestor, and Most recent common
ancestor, Timeline of evolution, Timeline of human evolution, and Universal Code (Biology)
[edit] Genetics
While on board HMS Beagle, Charles Darwin collected numerous specimens, many new to science,
which supported his later theory of evolution by natural selection.
Although it has only recently become available, one of the strongest evidences for common descent
comes from the study of gene sequences. Comparative sequence analysis examines the relationship
between the DNA sequences of different species,[1] producing several lines of evidence that confirm
Darwin's original hypothesis of common descent. If the hypothesis of common descent is true, then species
that share a common ancestor will have inherited that ancestor's DNA sequence. Notably they will have
inherited mutations unique to that ancestor. More closely-related species will have a greater fraction of
identical sequence and will have shared substitutions when compared to more distantly-related species.
The simplest and most powerful evidence is provided by phylogenetic reconstruction. Such
reconstructions, especially when done using slowly-evolving protein sequences, are often quite robust and
can be used to reconstruct a great deal of the evolutionary history of modern organisms (and even in some
instances such as the recovered gene sequences of mammoths, Neanderthals or T. rex, the evolutionary
history of extinct organisms). These reconstructed phylogenies recapitulate the relationships established
through morphological and biochemical studies. The most detailed reconstructions have been performed on
the basis of the mitochondrial genomes shared by all eukaryotic organisms, which are short and easy to
sequence; the broadest reconstructions have been performed either using the sequences of a few very
ancient proteins or by using ribosomal RNA sequence.
This evidence does not support the rival hypothesis that genetic similarity of two species is the
product of common functional or structural requirements, and not common descent[ citation needed] (for
example, if there is one best way to produce a hoof, all hoofed creatures will share a genetic basis even if
they are not related). However, phylogenetic relationships also extend to a wide variety of nonfunctional
sequence elements, including repeats, transposons, pseudogenes, and mutations in protein-coding
sequences that do not result in changes in amino-acid sequence. While a minority of these elements might
later be found to harbor function, in aggregate they demonstrate that identity must be the product of common
descent rather than common function.
[edit] Universal biochemical organisation and molecular variance patterns
All known extant organisms are based on the same fundamental biochemical organisation: genetic
information encoded as nucleic acid (DNA, or RNA for viruses), transcribed into RNA, then translated into
proteins (that is, polymers of amino acids) by highly conserved ribosomes. Perhaps most tellingly, the
Genetic Code (the "translation table" between DNA and amino acids) is the same for almost every organism,
meaning that a piece of DNA in a bacterium codes for the same amino acid as in a human cell. ATP is used
as energy currency by all extant life. A deeper understanding of developmental biology shows that common
morphology is, in fact, the product of shared genetic elements.[ citation needed] For example, although
camera-like eyes are believed to have evolved independently on many separate occasions,[ citation needed]
they share a common set of light-sensing proteins (opsins), suggesting a common point of origin for all
sighted creatures.[2] Another noteworthy example is the familiar vertebrate body plan, whose structure is
controlled by the homeobox (Hox) family of genes.
[edit] DNA sequencing

Comparison of the DNA sequences allows organisms to be grouped by sequence similarity, and the
resulting phylogenetic trees are typically congruent with traditional taxonomy, and are often used to
strengthen or correct taxonomic classifications. Sequence comparison is considered a measure robust
enough to be used to correct erroneous assumptions in the phylogenetic tree in instances where other
evidence is scarce. For example, neutral human DNA sequences are approximately 1.2% divergent (based
on substitutions) from those of their nearest genetic relative, the chimpanzee, 1.6% from gorillas, and 6.6%
from baboons.[3] Genetic sequence evidence thus allows inference and quantification of genetic relatedness
between humans and other apes.[4][5] The sequence of the 16S ribosomal RNA gene, a vital gene encoding
a part of the ribosome, was used to find the broad phylogenetic relationships between all extant life. The
analysis, originally done by Carl Woese, resulted in the three-domain system, arguing for two major splits in
the early evolution of life. The first split led to modern Bacteria and the subsequent split led to modern
Archaea and Eukaryotes.
[edit] Proteins
The proteomic evidence also supports the universal ancestry of life. Vital proteins, such as the
ribosome, DNA polymerase, and RNA polymerase, are found in everything from the most primitive bacteria
to the most complex mammals. The core part of the protein is conserved across all lineages of life, serving
similar functions. Higher organisms have evolved additional protein subunits, largely affecting the regulation
and protein-protein interaction of the core. Other overarching similarities between all lineages of extant
organisms, such as DNA, RNA, amino acids, and the lipid bilayer, give support to the theory of common
descent. Phylogenetic analyses of protein sequences from various organisms produce similar trees of
relationship between all organisms.[6] The chirality of DNA, RNA, and amino acids is conserved across all
known life. As there is no functional advantage to right- or left-handed molecular chirality, the simplest
hypothesis is that the choice was made randomly by early organisms and passed on to all extant life through
common descent. Further evidence for reconstructing ancestral lineages comes from junk DNA such as
pseudogenes, "dead" genes which steadily accumulate mutations.[7]
[edit] Pseudogenes
Pseudogenes, also known as Junk DNA, or dead DNA, are the extra DNA in an organism's genome
that do not get transcribed into RNA to produce protein. This is an example of a vestige since reproducing
these genes uses energy and is a waste, yet 99% of the human genome is these (1% working DNA).[8] A
pseudogene is produced when a gene that is no longer used accumulates mutations that make it non-
functional. But since it is not transcribed, it does not affect fitness when it disappears, unless it has provided
some new beneficial function as non-coding DNA.
[edit] Other mechanisms

There is also a large body of molecular evidence for a number of different mechanisms for large
evolutionary changes, among them: genome and gene duplication, which facilitates rapid evolution by
providing substantial quantities of genetic material under weak or no selective constraints; horizontal gene
transfer, the process of transferring genetic material to another cell that is not an organism's offspring,
allowing for species to acquire beneficial genes from each other; and recombination, capable of reassorting
large numbers of different alleles and of establishing reproductive isolation. The Endosymbiotic theory
explains the origin of mitochondria and plastids (e.g. chloroplasts), which are organelles of eukaryotic cells,
as the incorporation of an ancient prokaryotic cell into ancient eukaryotic cell. Rather than evolving
eukaryotic organelles slowly, this theory offers a mechanism for a sudden evolutionary leap by incorporating
the genetic material and biochemical composition of a separate species. Evidence supporting this
mechanism has recently been found in the protist Hatena: as a predator it engulfs a green algae cell, which
subsequently behaves as an endosymbiont, nourishing Hatena, which in turn loses its feeding apparatus and
behaves as an autotroph.[9][10]
Since metabolic processes do not leave fossils, research into the evolution of the basic cellular
processes is done largely by comparison of existing organisms. Many lineages diverged when new metabolic
processes appeared, and it is theoretically possible to determine when certain metabolic processes
appeared by comparing the traits of the descendants of a common ancestor or by detecting their physical
manifestations. As an example, the appearance of oxygen in the earth's atmosphere is linked to the evolution
of photosynthesis.
[edit] Specific examples
[edit] Chromosome 2 in humans

Main article: Chromosome 2 (human)
Further information: Genes of the Chromosome 2 fusion site in chimpanzees
Fusion of ancestral chromosomes left distinctive remnants of telomeres, and a vestigial centromere
Clear evidence for the evolution of Homo sapiens from a common ancestor with chimpanzees is the
number of chromosomes in human as compared to all other members of Hominidae. All Hominidae with the
exception of humans have 24 pairs of chromosomes. Humans have only 23 pairs. Human chromosome 2 is
widely accepted to be a result of an end-to-end fusion of two ancestral chromosomes.[11][12]
The evidence for this includes:
• The correspondence of chromosome 2 to two ape chromosomes. The closest human
relative, the common chimpanzee, has near-identical DNA sequences to human chromosome 2, but
they are found in two separate chromosomes. The same is true of the more distant gorilla and
orangutan.[13][14]
• The presence of a vestigial centromere. Normally a chromosome has just one centromere,
but in chromosome 2 there are remnants of a second centromere.[15]
• The presence of vestigial telomeres. These are normally found only at the ends of a
chromosome, but in chromosome 2 there are additional telomere sequences in the middle.[16]
Chromosome 2 thus presents very strong evidence in favour of the common descent of humans and
other apes. According to researcher J. W. IJdo, "We conclude that the locus cloned in cosmids c8.1 and
c29B is the relic of an ancient telomere-telomere fusion and marks the point at which two ancestral ape
chromosomes fused to give rise to human chromosome 2."[16]
[edit] Cytochrome c
Main article: Cytochrome c
A classic example of biochemical evidence for evolution is the variance of the ubiquitous (i.e. all
living organisms have it, because it performs very basic life functions) protein Cytochrome c in living cells.
The variance of cytochrome c of different organisms is measured in the number of differing amino acids,
each differing amino acid being a result of a base pair substitution, a mutation. If each differing amino acid is
assumed to be the result of one base pair substitution, it can be calculated how long ago the two species
diverged by multiplying the number of base pair substitutions by the estimated time it takes for a substituted
base pair of the cytochrome c gene to be successfully passed on. For example, if the average time it takes
for a base pair of the cytochrome c gene to mutate is N years, the number of amino acids making up the
cytochrome c protein in monkeys differ by one from that of humans, this leads to the conclusion that the two
species diverged N years ago.
The primary structure of cytochrome c consists of a chain of about 100 amino acids. Many higher
order organisms possess a chain of 104 amino acids.[17]
The cytochrome c molecule has been extensively studied for the glimpse it gives into evolutionary
biology. Both chicken and turkeys have identical sequence homology (amino acid for amino acid), as do pigs,
cows and sheep. Both humans and chimpanzees share the identical molecule, while rhesus monkeys share
all but one of the amino acids:[18] the 66th amino acid is isoleucine in the former and threonine in the latter.
[17]
What makes these homologous similarities particularly suggestive of common ancestry in the case of
cytochrome C, in addition to the fact that the phylogenies derived from them match other phylogenies very
well, is the high degree of functional redundancy of the cytochrome C molecule. The different existing
configurations of amino acids do not significantly affect the functionality of the protein, which indicates that
the base pair substitutions are not part of a directed design, but the result of random mutations that aren't
subject to selection.[19]
[edit] Recent African origin of modern humans

Main article: Recent single-origin hypothesis
See also: Human mitochondrial DNA haplogroup and Human Y-chromosome DNA haplogroup
Mathematical models of evolution, pioneered by the likes of Sewall Wright, Ronald Fisher and J. B.
S. Haldane and extended via diffusion theory by Motoo Kimura, allow predictions about the genetic structure
of evolving populations. Direct examination of the genetic structure of modern populations via DNA
sequencing has recently allowed verification of many of these predictions. For example, the Out of Africa
theory of human origins, which states that modern humans developed in Africa and a small sub-population
migrated out (undergoing a population bottleneck), implies that modern populations should show the
signatures of this migration pattern. Specifically, post-bottleneck populations (Europeans and Asians) should
show lower overall genetic diversity and a more uniform distribution of allele frequencies compared to the
African population. Both of these predictions are borne out by actual data from a number of studies.[20]
[edit] Evidence from comparative anatomy

Comparative study of the anatomy of groups of animals or plants reveals that certain structural
features are basically similar. For example, the basic structure of all flowers consists of sepals, petals,
stigma, style and ovary; yet the size, colour, number of parts and specific structure are different for each
individual species.
[edit] Nested hierarchies and classification

Taxonomy is based on the fact that all organisms are related to each other in nested hierarchies
based on shared characteristics. Most existing species can be organized rather easily in a nested
hierarchical classification. This is evident from the Linnaean classification scheme. Based on shared derived
characters, closely related organisms can be placed in one group (such as a genus), several genera can be
grouped together into one family, several families can be grouped together into an order, etc.[21] The
existence of these nested hierarchies was recognized by many biologists before Darwin, but he showed that
his theory of evolution with its branching pattern of common descent could explain them.[21][22] Darwin
described how common descent could provide a logical basis for classification:[23]
“ All the foregoing rules and aids and difficulties in classification are explained, if I ”
do not greatly deceive myself, on the view that the natural system is founded on
descent with modification; that the characters which naturalists consider as showing
true affinity between any two or more species, are those which have been inherited from
a common parent, and, in so far, all true classification is genealogical; that community of
descent is the hidden bond which naturalists have been unconsciously seeking, ...
—Charles Darwin, On the Origin of Species, page 577
[edit] Homologous structures and divergent (adaptive) evolution

If widely separated groups of organisms are originated from a common ancestry, they are expected
to have certain basic features in common. The degree of resemblance between two organisms should
indicate how closely related they are in evolution:
• Groups with little in common are assumed to have diverged from a common ancestor much
earlier in geological history than groups which have a lot in common;
• In deciding how closely related two animals are, a comparative anatomist looks for structures
that are fundamentally similar, even though they may serve different functions in the adult. Such
structures are described as homologous and suggest a common origin.
• In cases where the similar structures serve different functions in adults, it may be necessary
to trace their origin and embryonic development. A similar developmental origin suggests they are
the same structure, and thus likely to be derived from a common ancestor.
When a group of organisms share a homologous structure which is specialized to perform a variety
of functions in order to adapt different environmental conditions and modes of life are called adaptive
radiation. The gradual spreading of organisms with adaptive radiation is known as divergent evolution.
[edit] Vestigial structures
Main article: Vestigiality
See also: Human vestigiality
A strong and direct evidence for common descent comes from vestigial structures.[24] Rudimentary
body parts, those that are smaller and simpler in structure than corresponding parts in the ancestral species,
are called vestigial organs. They are usually degenerated or underdeveloped. The existence of vestigial
organs can be explained in terms of changes in the environment or modes of life of the species. Those
organs are thought to be functional in the ancestral species but are now either nonfunctional or repurposed.
Examples are the pelvic girdles of whales, haltere (hind wings) of flies and mosquitos, wings of flightless
birds such as ostriches, and the leaves of some xerophytes (e.g. cactus) and parasitic plants (e.g. dodder). It
must be noted, however, that vestigial structures may have had their original function replaced with another.
For example, the halteres in dipterists help balance the insect while in flight and the wings of ostriches are
used in mating rituals.
“ The most reasonable conclusion to draw is that these creatures descended
from creatures in which these parts were functional, which in turn indicates that most (or
indeed all) creatures descended from common ancenstors. ”
—Natan Slifkin, The Challenge of Creation, page 262
[edit] Evolutionary developmental biology and embryonic development

Main article: Evolutionary developmental biology
See also: Embryogenesis
Evolutionary developmental biology is the biological field that compares the developmental process
of different organisms to determine ancestral relationships between species. A large variety of organism’s
genomes contain a small fraction of genes that control the organisms development. Hox genes are an
example of these types of nearly universal genes in organisms pointing to an origin of common ancestry.
Embryological evidence comes from the development of organisms at the embryological level with the
comparison of different organisms embryos similarity. Remains of ancestral traits often appear and disappear
in different stages of the embryological development process. Examples include such as hair growth and loss
(lanugo) during human development;[25] the appearance of transitions from fish to amphibians to reptiles
and then to mammals in all mammal embryos; development and degeneration of a yolk sac; terrestrial frogs
and salamanders passing through the larval stage within the egg—with features of typically aquatic larvae—but
hatch ready for life on land;[26] and the appearance of gill-like structures (pharyngeal arch) in vertebrate
embryo development including. Note that in fish the arches become gills while in humans for example,
become the pharynx.
[edit] Atavisms
Main article: Atavism
An atavism is an evolutionary throwback, such as traits reappearing which had disappeared
generations ago.[27] Atavisms occur because genes for previously existing phenotypical features are often
preserved in DNA, even though the genes are not expressed in some or most of the organisms possessing
them.[28] Some examples of this are hind-legged snakes or whales;[27][29][30] the extra toes of ungulates
that do not even reach the ground,[31] chicken's teeth,[32] reemergence of sexual reproduction in Hieracium
pilosella and Crotoniidae;[33] and humans with tails,[27] extra nipples,[29] and large canine teeth.[29]
Figure 5a: The principle of homology illustrated by the adaptive radiation of the forelimb of mammals.
All conform to the basic pentadactyl pattern but are modified for different usages. The third metacarpal is
shaded throughout; the shoulder is crossed-hatched.
Figure 5b: Illustration of the Eoraptor lunensis pelvis of the saurischian order and the Lesothosaurus
diagnosticus pelvis of the ornithischian order in the Dinosauria superorder. The parts of the pelvis show
modification over time. The cladogram is shown to illustrate the distance of divergence between the two
species.
Figure 5c: Adaptation of insect mouthparts: a, antennae; c, compound eye; lb, labrium; lr, labrum;
md, mandibles; mx, maxillae.
[edit] Pentadactyl limb

Further information: Evolution of mammals
The pattern of limb bones called pentadactyl limb is an example of homologous structures (Fig. 5a).
It is found in all classes of tetrapods (i.e. from amphibians to mammals). It can even be traced back to the
fins of certain fossil fishes from which the first amphibians are thought to have evolved such as tiktaalik. The
limb has a single proximal bone (humerus), two distal bones (radius and ulna), a series of carpals (wrist
bones), followed by five series of metacarpals (palm bones) and phalanges (digits). Throughout the
tetrapods, the fundamental structures of pentadactyl limbs are the same, indicating that they originated from
a common ancestor. But in the course of evolution, these fundamental structures have been modified. They
have become superficially different and unrelated structures to serve different functions in adaptation to
different environments and modes of life. This phenomenon is clearly shown in the forelimbs of mammals.
For example:
• In the monkey, the forelimbs are much elongated to form a grasping hand for climbing and
swinging among trees.
• In the pig, the first digit is lost, and the second and fifth digits are reduced. The remaining two
digits are longer and stouter than the rest and bear a hoof for supporting the body.
• In the horse, the forelimbs are adapted for support and running by great elongation of the
third digit bearing a hoof.
• The mole has a pair of short, spade-like forelimbs for burrowing.
• The anteater uses its enlarged third digit for tearing down ant hills and termite nests.
• In the whale, the forelimbs become flippers for steering and maintaining equilibrium during
swimming.
• In the bat, the forelimbs have turned into wings for flying by great elongation of four digits,
while the hook-like first digit remains free for hanging from trees.
[edit] Pelvic structure of dinosaurs

See also: Evolution of dinosaurs and Evolution of birds
Similar to the pentadactyl limb in mammals, the earliest dinosaurs split into two distinct orders—the
saurischia and ornithischia. They are classified as one or the other in accordance with what the fossils show.
In Figure 5b, it is clear that early saurischians resembled early ornithischians. The pattern of the pelvis in all
species of dinosaurs is an example of homologous structures. Each order of dinosaur has slightly differing
pelvis bones providing evidence of common descent. Additionally, modern birds show a similarity to ancient
saurischian pelvic structures therefore indicating the evolution of birds from dinosaurs.
[edit] Insect mouthparts

The basic structures of mouthparts are the same in different species of insects, including a labrum
(upper lip), a pair of mandibles, a hypopharynx (floor of mouth), a pair of maxillae, and a labium. Evolution
has caused enlargement and modification of these structures in some species, while it has caused the
reduction and loss of them in other species. The modifications enable the insects to exploit a variety of food
materials (Fig. 5c):
(A) Primitive state — biting and chewing: e.g. grasshopper. Strong mandibles and maxillae for
manipulating food. (B) Ticking and biting: e.g. honey bee. Labium long to lap up nectar; mandibles chew
pollen and mould wax. (C) Sucking: e.g. butterfly. Labrum reduced; mandibles lost; maxillae long forming
sucking tube. (D) Piercing and sucking, e.g.. female mosquito. Labrum and maxillae form tube; mandibles
form piercing stylets; labrum grooved to hold other parts.
[edit] Other arthropod appendages

Insect mouthparts and antennae are considered homologues of insect legs. Parallel developments
are seen in some arachnids: The anterior pair of legs may be modified as analogues of antennae, particularly
in whip scorpions, which walk on six legs. These developments provide support for the theory that complex
modifications often arise by duplication of components, with the duplicates modified in different directions.
[edit] Recurrent laryngeal nerve in giraffes
The path of the recurrent laryngeal nerve in giraffes. The laryngeal nerve is compensated for by
subsequent tinkering from natural selection.
The recurrent laryngeal nerve is a fourth branch of the vagus nerve, which is a cranial nerve. In
mammals, its path is extraordinarily long. As a part of the vagus nerve, it comes from the brain, passes
through the neck down to heart, rounds the dorsal aorta and returns up to the larynx, again through the neck.
This path is suboptimal even for humans, but for giraffes it becomes even more suboptimal. Due to
the lengths of their necks, the recurrent laryngeal nerve may be up to 4m long (13 ft), despite its optimal route
being a distance of just several inches.
The indirect route of this nerve is the result of evolution of mammals from fish, which had no neck
and had a relatively short nerve that innervated one gill slit and passed near the gill arch. Since then, gills
have evolved into lungs and the gill arch has become the dorsal aorta in mammals.[34][35]
[edit] Route of the vas deferens
Route of the vas deferens from the testis to the penis

Similar to the laryngeal nerve in giraffes, the vas deferens is part of the male anatomy of many
vertebrates; it transports sperm from the epididymis in anticipation of ejaculation. In humans, the vas
deferens routes up from the testicle, looping over the ureter, and back down to the urethra and penis. It has
been suggested that this is due to the descent of the testicles during the course of human evolution—likely
associated with temperature. As the testicles descended, the vas deferens lengthened to accommodate the
accidental “hook” over the ureter.[35][36]
[edit] Evidence from paleontology

An insect trapped in amber.
When organisms die, they often decompose rapidly or are consumed by scavengers, leaving no
permanent evidences of their existence. However, occasionally, some organisms are preserved. The
remains or traces of organisms from a past geologic age embedded in rocks by natural processes are called
fossils. They are extremely important for understanding the evolutionary history of life on Earth, as they
provide direct evidence of evolution and detailed information on the ancestry of organisms. Paleontology is
the study of past life based on fossil records and their relations to different geologic time periods.
For fossilization to take place, the traces and remains of organisms must be quickly buried so that
weathering and decomposition do not occur. Skeletal structures or other hard parts of the organisms are the
most commonly occurring form of fossilized remains (Paul, 1998), (Behrensmeyer, 1980) and (Martin, 1999).
There are also some trace "fossils" showing moulds, cast or imprints of some previous organisms.
As an animal dies, the organic materials gradually decay, such that the bones become porous. If the
animal is subsequently buried in mud, mineral salts will infiltrate into the bones and gradually fill up the pores.
The bones will harden into stones and be preserved as fossils. This process is known as petrification. If dead
animals are covered by wind-blown sand, and if the sand is subsequently turned into mud by heavy rain or
floods, the same process of mineral infiltration may occur. Apart from petrification, the dead bodies of
organisms may be well preserved in ice, in hardened resin of coniferous trees (amber), in tar, or in anaerobic,
acidic peat. Fossilization can sometimes be a trace, an impression of a form. Examples include leaves and
footprints, the fossils of which are made in layers that then harden.
Fossil trilobite. Trilobites were hard-shelled arthropods, related to living horseshoe crabs and spiders,
that first appeared in significant numbers around 540 mya, dying out 250 mya.
It is possible to find out how a particular group of organisms evolved by arranging its fossil records in
a chronological sequence. Such a sequence can be determined because fossils are mainly found in
sedimentary rock. Sedimentary rock is formed by layers of silt or mud on top of each other; thus, the resulting
rock contains a series of horizontal layers, or strata. Each layer contains fossils which are typical for a
specific time period during which they were made. The lowest strata contain the oldest rock and the earliest
fossils, while the highest strata contain the youngest rock and more recent fossils.
A succession of animals and plants can also be seen from fossil records. By studying the number
and complexity of different fossils at different stratigraphic levels, it has been shown that older fossil-bearing
rocks contain fewer types of fossilized organisms, and they all have a simpler structure, whereas younger
rocks contain a greater variety of fossils, often with increasingly complex structures.[ citation needed]
In the past, geologists could only roughly estimate the ages of various strata and the fossils found.
They did so, for instance, by estimating the time for the formation of sedimentary rock layer by layer. Today,
by measuring the proportions of radioactive and stable elements in a given rock, the ages of fossils can be
more precisely dated by scientists. This technique is known as radiometric dating.
Throughout the fossil record, many species that appear at an early stratigraphic level disappear at a
later level. This is interpreted in evolutionary terms as indicating the times at which species originated and
became extinct. Geographical regions and climatic conditions have varied throughout the Earth's history.
Since organisms are adapted to particular environments, the constantly changing conditions favoured
species which adapted to new environments through the mechanism of natural selection.
[edit] Extent of the fossil record

See also: Transitional fossil and List of transitional fossils
Charles Darwin collected fossils in South America, and found fragments of armor which he thought
were like giant versions of the scales on the modern armadillos living nearby. The anatomist Richard Owen
showed him that the fragments were from gigantic extinct glyptodons, related to the armadillos. This was one
of the patterns of distribution that helped Darwin to develop his theory.[37]
Cynognathus, a Eucynodont, one of a grouping of Therapsids ("mammal-like reptiles") that is

ancestral to all modern mammals.
Despite the relative rarity of suitable conditions for fossilization, approximately 250,000 fossil species
are known.[38] The number of individual fossils this represents varies greatly from species to species, but
many millions of fossils have been recovered: for instance, more than three million fossils from the last Ice
Age have been recovered from the La Brea Tar Pits in Los Angeles.[39] Many more fossils are still in the
ground, in various geological formations known to contain a high fossil density, allowing estimates of the total
fossil content of the formation to be made. An example of this occurs in South Africa's Beaufort Formation
(part of the Karoo Supergroup, which covers most of South Africa), which is rich in vertebrate fossils,
including therapsids (reptile/mammal transitional forms).[40] It has been estimated that this formation
contains 800 billion vertebrate fossils.[41]
[edit] Limitations
The fossil record is an important source for scientists when tracing the evolutionary history of
organisms. However, because of limitations inherent in the record, there are not fine scales of intermediate
forms between related groups of species. This lack of continuous fossils in the record is a major limitation in
tracing the descent of biological groups. Furthermore, there are also much larger gaps between major
evolutionary lineages.[citation needed] When transitional fossils are found that show intermediate forms in
what had previously been a gap in knowledge, they are often popularly referred to as "missing links".
There is a gap of about 100 million years between the beginning of the Cambrian period and the end
of the Ordovician period. The early Cambrian period was the period from which numerous fossils of sponges,
cnidarians (e.g., jellyfish), echinoderms (e.g., eocrinoids), molluscs (e.g., snails) and arthropods (e.g.,
trilobites) are found. The first animal that possessed the typical features of vertebrates, the Arandaspis, was
dated to have existed in the later Ordovician period. Thus few, if any, fossils of an intermediate type between
invertebrates and vertebrates have been found, although likely candidates include the Burgess Shale animal,
Pikaia gracilens, and its Maotianshan shales relatives, Myllokunmingia, Yunnanozoon, Haikouella lanceolata,
and Haikouichthys.[citation needed]
Some of the reasons for the incompleteness of fossil records are:[ citation needed]
• In general, the probability that an organism becomes fossilized is very low;
• Some species or groups are less likely to become fossils because they are soft-bodied;
• Some species or groups are less likely to become fossils because they live (and die) in
conditions that are not favourable for fossilization;
• Many fossils have been destroyed through erosion and tectonic movements;
• Most fossils are fragmentary;
• Some evolutionary change occurs in populations at the limits of a species' ecological range,
and as these populations are likely to be small, the probability of fossilization is lower (see
punctuated equilibrium);
• Similarly, when environmental conditions change, the population of a species is likely to be
greatly reduced, such that any evolutionary change induced by these new conditions is less likely to
be fossilized;
• Most fossils convey information about external form, but little about how the organism
functioned;
• Using present-day biodiversity as a guide, this suggests that the fossils unearthed represent
only a small fraction of the large number of species of organisms that lived in the past.

[edit] Evolution of the horse

Main article: Evolution of the horse
Evolution of the horse showing reconstruction of the fossil species obtained from successive rock
strata. The foot diagrams are all front views of the left forefoot. The third metacarpal is shaded throughout.
The teeth are shown in longitudinal section.
Due to an almost-complete fossil record found in North American sedimentary deposits from the
early Eocene to the present, the horse provides one of the best examples of evolutionary history (phylogeny).
This evolutionary sequence starts with a small animal called Hyracotherium (commonly referred to as
Eohippus) which lived in North America about 54 million years ago, then spread across to Europe and Asia.
Fossil remains of Hyracotherium show it to have differed from the modern horse in three important respects:
it was a small animal (the size of a fox), lightly built and adapted for running; the limbs were short and
slender, and the feet elongated so that the digits were almost vertical, with four digits in the forelimbs and
three digits in the hindlimbs; and the incisors were small, the molars having low crowns with rounded cusps
covered in enamel.
The probable course of development of horses from Hyracotherium to Equus (the modern horse)
involved at least 12 genera and several hundred species. The major trends seen in the development of the
horse to changing environmental conditions may be summarized as follows:
• Increase in size (from 0.4 m to 1.5 m — from 15in to 60in);
• Lengthening of limbs and feet;
• Reduction of lateral digits;
• Increase in length and thickness of the third digit;
• Increase in width of incisors;
• Replacement of premolars by molars; and
• Increases in tooth length, crown height of molars.
Fossilized plants found in different strata show that the marshy, wooded country in which
Hyracotherium lived became gradually drier. Survival now depended on the head being in an elevated
position for gaining a good view of the surrounding countryside, and on a high turn of speed for escape from
predators, hence the increase in size and the replacement of the splayed-out foot by the hoofed foot. The
drier, harder ground would make the original splayed-out foot unnecessary for support. The changes in the
teeth can be explained by assuming that the diet changed from soft vegetation to grass. A dominant genus
from each geological period has been selected to show the slow alteration of the horse lineage from its
ancestral to its modern form.
[edit] Evidence from geographical distribution

Data about the presence or absence of species on various continents and islands (biogeography)
can provide evidence of common descent and shed light on patterns of speciation.
[edit] Continental distribution

All organisms are adapted to their environment to a greater or lesser extent. If the abiotic and biotic
factors within a habitat are capable of supporting a particular species in one geographic area, then one might
assume that the same species would be found in a similar habitat in a similar geographic area, e.g. in Africa
and South America. This is not the case. Plant and animal species are discontinuously distributed throughout
the world:
• Africa has Old World monkeys, apes, elephants, leopards, giraffes, and hornbills.
• South America has New World monkeys, cougars, jaguars, sloths, llamas, and toucans.
• Deserts in North and South America have native cacti, but deserts in Africa, Asia, and
Australia have succulent native euphorbs that resemble cacti but are very different, even though in
some cases cacti have done very well (for example in Australian deserts) when introduced by
humans.[42]
Even greater differences can be found if Australia is taken into consideration, though it occupies the
same latitude as much of South America and Africa. Marsupials like kangaroos, bandicoots, and quolls make
up about half of Australia's indigenous mammal species.[43] By contrast, marsupials are today totally absent
from Africa and form a smaller portion of the mammalian fauna of South America, where opossums, shrew
opossums, and the monito del monte occur. The echidnas and platypus, the only living representatives of
primitive egg-laying mammals (monotremes), can be found only in Australia and are totally absent in the rest
of the world. On the other hand, Australia is missing many groups of placental mammals that are common on
other continents (carnivorans, artiodactyls, shrews, squirrels, lagomorphs), although it does have indigenous
bats and murine rodents; many other placentals, such as rabbits and foxes, have been introduced there by
humans.
Other animal distribution examples include bears, located on all continents excluding Africa and
Australia, and the polar bear only located nearest to the North Pole. Penguins are located only around the
South Pole despite similar weather conditions at the North Pole. Families of sirenians are distributed
exclusively around the earth’s waters, where manatees are located in western Africa waters, northern South
American waters, and West Indian waters only while the related family, the Dugongs, are located only in
Oceanic waters north of Australia, and the coasts surrounding the Indian Ocean.
The same kinds of fossils are found from areas known to be adjacent to one another in the past but
which, through the process of continental drift, are now in widely divergent geographic locations. For
example, fossils of the same types of ancient amphibians, arthropods and ferns are found in South America,
Africa, India, Australia and Antarctica, which can be dated to the Paleozoic Era, at which time these regions
were united as a single landmass called Gondwana.[44] Sometimes the descendants of these organisms can
be identified and show unmistakable similarity to each other, even though they now inhabit very different
regions and climates.
[edit] Island biogeography
Four of the 13 finch species found on the Galápagos Archipelago, are thought to have evolved by an
adaptive radiation that diversified their beak shapes to adapt them to different food sources.
[edit] Types of species found on islands
Evidence from island biogeography has played an important and historic role in the development of
evolutionary biology. For purposes of biogeography, islands are divided into two classes. Continental islands
are islands like Great Britain, and Japan that have at one time or another been part of a continent. Oceanic
islands, like the Hawaiian islands, the Galapagos islands and St. Helena, on the other hand are islands that
have formed in the ocean and never been part of any continent. Oceanic islands have distributions of native
plants and animals that are unbalanced in ways that make them distinct from the biotas found on continents
or continental islands. Oceanic islands do not have native terrestrial mammals (they do sometimes have bats
and seals), amphibians, or fresh water fish. In some cases they have terrestrial reptiles (such as the iguanas
and giant tortoises of the Galapagos islands) but often (for example Hawaii) they do not. This despite the fact
that when species such as rats, goats, pigs, cats, mice, and cane toads, are introduced to such islands by
humans they often thrive. Starting with Charles Darwin, many scientists have conducted experiments and
made observations that have shown that the types of animals and plants found, and not found, on such
islands are consistent with the theory that these islands were colonized accidentally by plants and animals
that were able to reach them. Such accidental colonization could occur by air, such as plant seeds carried by
migratory birds, or bats and insects being blown out over the sea by the wind, or by floating from a continent
or other island by sea, as for example by some kinds of plant seeds like coconuts that can survive immersion
in salt water, and reptiles that can survive for extended periods on rafts of vegetation carried to sea by
storms.[45]
[edit] Endemism
Many of the species found on remote islands are endemic to a particular island or group of islands,
meaning they are found nowhere else on earth. Examples of species endemic to islands include many
flightless birds of New Zealand, lemurs of Madagascar, the Komodo dragon of Komodo ,[46] the Dragon’s
blood tree of Socotra ,[47] Tuatara of New Zealand,[48][49] and others. However many such endemic
species are related to species found on other nearby islands or continents; the relationship of the animals
found on the Galapagos Islands to those found in South America is a well-known example.[45] All of these
facts, the types of plants and animals found on oceanic islands, the large number of endemic species found
on oceanic islands, and the relationship of such species to those living on the nearest continents, are most
easily explained if the islands were colonized by species from nearby continents that evolved into the
endemic species now found there.[45]
Other types of endemism do not have to include, in the strict sense, islands. Islands can mean
isolated lakes or remote and isolated areas. Examples of these would include the highlands of Ethiopia, Lake
Baikal, Fynbos of South Africa, forests of New Caledonia, and others. Examples of endemic organisms living
in isolated areas include the Kagu of New Caledonia,[50] cloud rats of the Luzon tropical pine forests of the
Philippines,[51][52] the boojum tree (Fouquieria columnaris) of the Baja California peninsula,[53] the Baikal
Seal[54] and the omul of Lake Baikal.
[edit] Adaptive radiations

Oceanic islands are frequently inhabited by clusters of closely related species that fill a variety of
ecological niches, often niches that are filled by very different species on continents. Such clusters, like the
Finches of the Galapagos, Hawaiian honeycreepers, members of the sunflower family on the Juan
Fernandez Archipelago and wood weevils on St. Helena are called adaptive radiations because they are best
explained by a single species colonizing an island (or group of islands) and then diversifying to fill available
ecological niches. Such radiations can be spectacular; 800 species of the fruit fly family Drosophila, nearly
half the world's total, are endemic to the Hawaiian islands. Another illustrative example from Hawaii is the
Silversword alliance, which is a group of thirty species found only on those islands. Members range from the
Silverswords that flower spectacularly on high volcanic slopes to trees, shrubs, vines and mats that occur at
various elevations from mountain top to sea level, and in Hawaiian habitats that vary from deserts to
rainforests. Their closest relatives outside Hawaii, based on molecular studies, are tarweeds found on the
west coast of North America. Interestingly, these tarweeds have sticky seeds that facilitate distribution by
migrant birds. Continental islands have less distinct biota, but those that have been long separated from any
continent also have endemic species and adaptive radiations, such as the 75 lemur species of Madagascar,
and the eleven extinct moa species of New Zealand.[45][55]
[edit] Ring Species

Main article: Ring species
In biology, a ring species is a connected series of neighboring populations that can interbreed with
relatively closely related populations, but for which there exist at least two "end" populations in the series that
are too distantly related to interbreed. Often such non-breeding-though-genetically-connected populations
co-exist in the same region thus creating a "ring". Ring species provide important evidence of evolution in
that they illustrate what happens over time as populations genetically diverge, and are special because they
represent in living populations what normally happens over time between long deceased ancestor
populations and living populations. If any of the populations intermediate between the two ends of the ring
were gone they would not be a continuous line of reproduction and each side would be a different species.
[56][57]
Figure 6a: A dymaxion map of the world showing the distribution of present species of camelid. The
solid black lines indicate migration routes and the blue represents current camel locations.
Figure 6b: Current distribution of Glossopteris placed on a Permian map showing the connection of
the continents. (1, South America; 2, Africa; 3, Madagascar; 4, India; 5, Antarctica; and 6, Australia)
Figure 6c: Present day distribution of marsupials. (Distribution shown in blue. Introduced areas
shown in green.)
[edit] Migration, isolation, and distribution of the Camel

The history of the camel provides an example of how fossil evidence can be used to reconstruct
migration and subsequent evolution. The fossil record indicates that the evolution of camelids started in
North America (see figure 6a), from which 6 million years ago they migrated across the Bering Strait into Asia
and then to Africa, and 3.5 million years ago through the Isthmus of Panama into South America. Once
isolated, they evolved along their own lines, giving rise to the Bactrian camel and Dromedary in Asia and
Africa and the llama and its relatives in South America. Camelids then went extinct in North America at the
end of the last ice age.[58]
[edit] Distribution of Glossopteris
The combination of continental drift and evolution can sometimes be used to make predictions about
what will be found in the fossil record. Glossopteris is an extinct species of seed fern plants from the
Permian. Glossopteris appears in the fossil record around the beginning of the Permian on the ancient
continent of Gondwana.[59] Continental drift explains the current biogeography of the tree. Present day
Glossopteris fossils are found in Permian strata in southeast South America, southeast Africa, all of
Madagascar, northern India, all of Australia, all of New Zealand, and scattered on the southern and northern
edges of Antarctica. During the Permian, these continents were connected as Gondwana (see figure 6b) in
agreement with magnetic striping, other fossil distributions, and glacial scratches pointing away from the
temperate climate of the South Pole during the Permian.[45]
[edit] Distribution of marsupials

The history of marsupials also provides an example of how the theories of evolution and continental
drift can be combined to make predictions about what will be found in the fossil record. The earliest marsupial
fossils are about 80 million years old and found in North America; by 40 million years ago fossils show that
they could be found throughout South America, but there is no evidence of them in Australia, where they now
predominate, until about 30 million years ago. The theory of evolution predicts that the Australian marsupials
must be descended from the older ones found in the Americas. The theory of continental drift says that
between 30 and 40 million years ago South America and Australia were still part of the Southern hemisphere
super continent of Gondwana and that they were connected by land that is now part of Antarctica. Therefore
combining the two theories scientists predicted that marsupials migrated from what is now South America
across what is now Antarctica to what is now Australia between 40 and 30 million years ago. This hypothesis
led paleontologists to Antarctica to look for marsupial fossils of the appropriate age. After years of searching
they found, starting in 1982, fossils on Seymour Island off the coast of the Antarctic Peninsula of more than a
dozen marsupial species that lived 35-40 million years ago.[42]
[edit] Evidence from observed natural selection

Examples for the evidence for evolution often stems from direct observation of natural selection in
the field and the laboratory. Scientists have observed and documented a multitude of events where natural
selection is in action. The most well known examples are antibiotic resistance in the medical field along with
better-known laboratory experiments documenting evolution's occurrence. Natural selection is tantamount to
common descent in the fact that long-term occurrence and selection pressures can lead to the diversity of life
on earth as found today. All adaptations—documented and undocumented changes concerned—are caused
by natural selection (and a few other minor processes). The examples below are only a small fraction of the
actual experiments and observations.
[edit] Specific examples of natural selection in the lab and in the field
[edit] Antibiotic and pesticide resistance

Main article: Antibiotic resistance
The development and spread of antibiotic resistant bacteria, like the spread of pesticide resistant
forms of plants and insects is evidence for evolution of species, and of change within species. Thus the
appearance of vancomycin resistant Staphylococcus aureus, and the danger it poses to hospital patients is a
direct result of evolution through natural selection. The rise of Shigella strains resistant to the synthetic
antibiotic class of sulfonamides also demonstrates the generation of new information as an evolutionary
process.[60] Similarly, the appearance of DDT resistance in various forms of Anopheles mosquitoes, and the
appearance of myxomatosis resistance in breeding rabbit populations in Australia, are all evidence of the
existence of evolution in situations of evolutionary selection pressure in species in which generations occur
rapidly.
[edit] E. coli long-term evolution experiment

Main article: E. coli long-term evolution experiment
See also: Experimental evolution
Experimental evolution uses controlled experiments to test hypotheses and theories of evolution. In
one early example, William Dallinger set up an experiment shortly before 1880, subjecting microbes to heat
with the aim of forcing adaptive changes. His experiment ran for around seven years, and his published
results were acclaimed, but he did not resume the experiment after the apparatus failed.[61]
The E. coli long-term evolution experiment that began in 1988 under the leadership of Richard Lenski
is still in progress, and has shown adaptations including the evolution of a strain of E. coli that was able to
grow on citric acid in the growth media.
[edit] Humans
Natural selection is being observed in contemporary human populations, with recent findings
demonstrating the population which is at risk of the severe debilitating disease kuru has significant over-
representation of an immune variant of the prion protein gene G127V versus non-immune alleles. Scientists
postulate one of the reasons for the rapid selection of this genetic variant is the lethality of the disease in non-
immune persons.[62][63] Other reported evolutionary trends in other populations include a lengthening of the
reproductive period, reduction in cholesterol levels, blood glucose and blood pressure.[64]
[edit] Lactose intolerance in humans

Lactose intolerance is the inability to metabolize lactose, because of a lack of the required enzyme
lactase in the digestive system. The normal mammalian condition is for the young of a species to experience
reduced lactase production at the end of the weaning period (a species-specific length of time). In humans, in
non-dairy consuming societies, lactase production usually drops about 90% during the first four years of life,
although the exact drop over time varies widely.[65] However, certain human populations have a mutation on
chromosome 2 which eliminates the shutdown in lactase production, making it possible for members of these
populations to continue consumption of fresh milk and other dairy products throughout their lives without
difficulty. This appears to be an evolutionarily recent adaptation to dairy consumption, and has occurred
independently in both northern Europe and east Africa in populations with a historically pastoral lifestyle.[66]
[edit] Nylon-eating bacteria

Nylon-eating bacteria are a strain of Flavobacterium that is capable of digesting certain byproducts of
nylon 6 manufacture. There is scientific consensus that the capacity to synthesize nylonase most probably
developed as a single-step mutation that survived because it improved the fitness of the bacteria possessing
the mutation. This is seen as a good example of evolution through mutation and natural selection that has
been observed as it occurs.[67][68][69][70]
[edit] Peppered moth

Main article: Peppered moth evolution
One classic example of adaptation in response to selection pressure is the case of the peppered
moth. The color of the moth has gone from light to dark to light again over the course of a few hundred years
due to the appearance and later disappearance of pollution from the Industrial Revolution in England.
[edit] Radiotrophic fungus

Recently evolved Radiotrophic fungi are fungi which appear to use the pigment melanin to convert
gamma radiation into chemical energy for growth[71][72] and were first discovered in 2007 as black molds
growing inside and around the Chernobyl Nuclear Power Plant.[71] Research at the Albert Einstein College
of Medicine showed that three melanin-containing fungi, Cladosporium sphaerospermum, Wangiella
dermatitidis, and Cryptococcus neoformans, increased in biomass and accumulated acetate faster in an
environment in which the radiation level was 500 times higher than in the normal environment.
[edit] Urban wildlife

Urban wildlife is wildlife that is able to live or thrive in urban environments. These types of
environments can exert selection pressures on organism, often leading to new adaptations. For example, the
weed Crepis sancta, found in France, has two types of seed, heavy and fluffy. The heavy ones land nearby to
the parent plant, whereas the fluffy seeds float further away on the wind. In urban environments, seeds that
float far will often land on infertile concrete. Within about 5-12 generations, the weed has been found to
evolve to produce significantly more heavy seeds than its rural relatives do.[73][74] Other examples of urban
wildlife are rock pigeons and species of crows adapting to city environments around the world; African
penguins in Simons Town; baboons in South Africa; and a variety of insects living in human habitations.
[edit] Evidence from observed speciation
Speciation is the evolutionary process by which new biological species arise. Speciation can occur
from a variety of different causes and are classified in various forms (e.g. allopatric, sympatric,
polyploidization, etc.). Scientists have observed numerous examples of speciation in the laboratory and in
nature, however, evolution has produced far more species than an observer would consider necessary. For
example, there are well over 350,000 described species of beetles.[75] Great examples of observed
speciation come from the observations of island biogeography and the process of adaptive radiation, both
explained in an earlier section. The examples shown below provide strong evidence for common descent and
are only a small fraction of the instances observed.
[edit] Blackcap
The bird species, Sylvia atricapillab, commonly referred to as Blackcaps, lives in Germany and flies
southwest to Spain while a smaller group flies northwest to Great Britain during the winter. The smaller
blackcap population only recently rerouted to Britain. Gregor Rolshausen from the University of Freiburg
found that the genetic separation of the two populations is already in progress. The differences found have
arisen in about 30 generations. With DNA sequencing, the individuals can be assigned to a correct group
with an 85% accuracy. Stuart Bearhop from the University of Exeter reported that birds wintering in England
tend to mate only among themselves, and not usually with those wintering in the Mediterranean.[76] It is still
inference to say that the populations will become two different species, but experts deduce that it is expected
due to the continued genetic and geographic separation.[77]
[edit] Drosophila melanogaster
A common fruit fly (Drosophila melanogaster).

William R. Rice and George W. Salt found experimental evidence of sympatric speciation in the
common fruit fly. They collected a population of Drosophila melanogaster from Davis, California and placed
the pupae into a habitat maze. Newborn flies had to investigate the maze to find food. The flies had three
choices to take in finding food. Light and dark (phototaxis), up and down (geotaxis), and the scent of
acetaldehyde and the scent of ethanol (chemotaxis) were the three options. This eventually divided the flies
into 42 spatio-temporal habitats.
They then cultured two strains that chose opposite habitats. One of the strains emerged early,
immediately flying upward in the dark attracted to the acetaldehyde. The other strain emerged late and
immediately flew downward, attracted to light and ethanol. Pupae from the two strains were then placed
together in the maze and allowed to mate at the food site. They then were collected. A selective penalty was
imposed on the female flies that switched habitats. This entailed that none of their gametes would pass on to
the next generation. After 25 generations of this mating test, it showed reproductive isolation between the two
strains. They repeated the experiment again without creating the penalty against habitat switching and the
result was the same; reproductive isolation was produced.[78][79][80]
[edit] Hawthorn fly

One example of evolution at work is the case of the hawthorn fly, Rhagoletis pomonella, also known
as the apple maggot fly, which appears to be undergoing sympatric speciation.[81] Different populations of
hawthorn fly feed on different fruits. A distinct population emerged in North America in the 19th century some
time after apples, a non-native species, were introduced. This apple-feeding population normally feeds only
on apples and not on the historically preferred fruit of hawthorns. The current hawthorn feeding population
does not normally feed on apples. Some evidence, such as the fact that six out of thirteen allozyme loci are
different, that hawthorn flies mature later in the season and take longer to mature than apple flies; and that
there is little evidence of interbreeding (researchers have documented a 4-6% hybridization rate) suggests
that this is occurring.[82]
[edit] London Underground mosquito

The London Underground mosquito is a species of mosquito in the genus Culex found in the London
Underground. It is thought to have evolved from the overground species Culex pipiens recently.
This mosquito, although first discovered in the London Underground system, has been found in
underground systems around the world. It is suggested that it may have adapted to human-made
underground systems since the last century from local above-ground Culex pipiens,[83] although more recent
evidence suggests that it is a southern mosquito variety related to Culex pipiens that has adapted to the
warm underground spaces of northern cities.[84]
The evidence for this mosquito being a different species from Culex pipiens comes from research by
Kate Byne and Richard Nichols. The species have very different behaviours,[85] are extremely difficult to
mate,[83] and with different allele frequency, consistent with genetic drift during a founder event.[86] More
specifically, this mosquito, Culex pipiens molestus, breeds all-year round, is cold intolerant, and bites rats,
mice, and humans, in contrast to the above ground species Culex pipiens that is cold tolerant, hibernates in
the winter, and bites only birds. When the two varieties were cross-bred the eggs were infertile suggesting
reproductive isolation.[83][85]
The fundamental results still stands: the genetic data indicate that the molestus form in the London
Underground mosquito appeared to have a common ancestry, rather than the population at each station
being related to the nearest above-ground population (i.e. the pipiens form). Byrne and Nichols' working
hypothesis was that adaptation to the underground environment had occurred locally in London once only.
These widely separated populations are distinguished by very minor genetic differences, which
suggest that the molestus form developed recently: a single mtDNA difference shared among the
underground populations of ten Russian cities;[87] a single fixed microsatellite difference in populations
spanning Europe, Japan, Australia, the middle East and Atlantic islands.[84]
[edit] Madeira House Mouse

The Madeira House mouse is a new species of mouse descended from the house mouse ( Mus
Musculus) that went through a recent speciation event.[88]
[edit] Mollies
The Shortfin Molly—Poecilia Mexicana—is a small fish that lives in the Sulfur Caves of Mexico.
Michael Tobler from the Texas A&M University has studied the fish for years and found that two distinct
populations of mollies—the dark interior fish and the bright surface water fish—are becoming more genetically
divergent.[89] The populations have no obvious barrier separating the two; however, it was found that the
mollies are hunted by a large water bug ( Belostoma spp). Tobler collected the bug and both types of mollies,
placed them in large plastic bottles, and put them back in the cave. After a day, it was found that, in the light,
the cave-adapted fish endured the most damage, with four out of every five stab-wounds from the water bugs
sharp mouthparts. In the dark, the situation was the opposite. The mollies’ senses can detect a predator’s
threat in their own habitats, but not in the other ones. Moving from one habitat to the other significantly
increases the risk of dying. Tobler plans on further experiments, but it is believed that it is a good example of
the rise of a new species.[90]
[edit] Thale cress
Arabidopsis thaliana (colloquially known as thale cress, mouse-ear cress or Arabidopsis).

Kirsten Bomblies et al. from the Max Planck Institute for Developmental Biology discovered that two
genes passed down by each parent of the thale cress plant, Arabidopsis thaliana. When the genes are
passed down, it ignites a reaction in the hybrid plant that turns its own immune system against it. In the
parents, the genes were not detrimental, but they evolved separately to react defectively when combined.[91]
To test this, Bomblies crossed 280 genetically different strains of Arabidopsis in 861 distinct ways
and found that 2 per cent of the resulting hybrids were necrotic. Along with allocating the same indicators, the
20 plants also shared a comparable collection of genetic activity in a group of 1,080 genes. In almost all of
the cases, Bomblies discovered that only two genes were required to cause the autoimmune response.
Bomblies looked at one hybrid in detail and found that one of the two genes belonged to the NB-LRR class, a
common group of disease resistance genes involved in recognizing new infections. When Bomblies removed
the problematic gene, the hybrids developed normally.[91]
Over successive generations, these incompatibilities could create divisions between different plant
strains, reducing their chances of successful mating and turning distinct strains into separate species.[92]
[edit] Interspecies fertility or hybridization

Understood from laboratory studies and observed instances of speciation in nature, finding species
that are able reproduce successfully or create hybrids between two different species infers that their
relationship is close. In conjunction with this, hybridization has been found to be a precursor to the creation of
new species by being a source of new genes for a species. The examples provided are only a small fraction
of the observed instances of speciation through hybridization. It is worth noting that plants are often subject
to the creation of a new species though hybridization.
[edit] Polar bear

See also: Polar bear#Taxonomy and evolution and Grizzly–polar bear hybrid
A specific example of large-scale evolution is the polar bear (Ursus maritimus), which though clearly
related to the brown bear (Ursus arctos) by virtue of the fact that though separate species they can still
interbreed and produce fertile offspring[93] it has also obviously acquired significant physiological differences
from the brown bear. These differences allow the polar bear to comfortably survive in conditions that the
brown could not including the ability to swim sixty miles or more at a time in freezing waters, blend in—the
change in fur color—and to stay warm in the arctic environment. Additionally, the elongation of the neck
makes it easier to keep their heads above water while swimming and the oversized webbed feet that act as
paddles when swimming. The polar bear has also evolved small papillae and vacuole-like suction cups on
the soles to make them less likely to slip on the ice, feet covered with heavy matting to protect the bottoms
from intense cold and provide traction, smaller ears to reduce the loss of heat, eyelids that act like
sunglasses, to accommodate their all-meat diet, a large stomach capacity to enable opportunistic feeding,
and the ability to fast for up to nine months while recycling their urea.[94][95]
[edit] Raphanobrassica
Raphanobrassica includes all intergeneric hybrids between the genera Raphanus (radish) and
Brassica (cabbages, etc.).[96][97]
The Raphanobrassica is an allopolyploid cross between the radish (Raphanus sativus) and cabbage
(Brassica oleracea). Plants of this parentage are now known as radicole. Two other fertile forms of
Raphanobrassica are known. Raparadish, an allopolyploid hybrid between Raphanus sativus and Brassica
rapa is grown as a fodder crop. "Raphanofortii" is the allopolyploid hybrid between Brassica tournefortii and
Raphanus caudatus.
The Raphanobrassica is a fascinating plant, because—in spite of its hybrid nature—it is not sterile.
This has led some botanists to propose that the accidental hybridization of a flower by pollen of another
species in nature could be a mechanism of speciation common in higher plants.
[edit] Salsify
Purple Salsify, Tragopogon porrifolius

Salsifies are one example where hybrid speciation has been observed. In the early 20th century,
humans introduced three species of goatsbeard into North America. These species, the western salsify ( T.
dubius), the meadow salsify (T. pratensis), and the oyster plant (T. porrifolius), are now common weeds in
urban wastelands. In the 1950s, botanists found two new species in the regions of Idaho and Washington,
where the three already known species overlapped. One new species, Tragopogon miscellus, is a tetraploid
hybrid of T. dubius and T. pratensis. The other species, Tragopogon mirus, is also an allopolyploid, but its
ancestors were T. dubius and T. porrifolius. These new species are usually referred to as "the Ownbey
hybrids" after the botanist who first described them. The T. mirus population grows mainly by reproduction of
its own members, but additional episodes of hybridization continue to add to the T. mirus population.[citation
needed]
[edit] Welsh groundsel

Welsh groundsel is an allopolyploid, a plant which contains sets of chromosomes originating from
two different species. Its ancestor was Senecio × baxteri, an infertile hybrid which can arise spontaneously
when the closely-related groundsel (Senecio vulgaris) and Oxford ragwort (Senecio squalidus) grow
alongside each other. Sometime in the early 20th century, an accidental doubling of the number of
chromosomes in an S. × baxteri plant led to the formation of a new fertile species.[98][99]
[edit] York groundsel

The York groundsel (Senecio eboracensis) is a hybrid species of the self-incompatible Senecio
squalidus (also known as Oxford ragwort) and the self-compatible Senecio vulgaris (also known as Common
groundsel). Like S. vulgaris, S. eboracensis is self-compatible, however, it shows little or no natural crossing
with its parent species, and is therefore reproductively isolated, indicating that strong breed barriers exist
between this new hybrid and its parents.
It is thought to have resulted from a backcrossing of the F1 hybrid of its parents to S. vulgaris. S.
vulgaris is native to Britain, while S. squalidus was introduced from Sicily in the early 18th century; therefore,
S. eboracensis has speciated from those two species within the last 300 years.
Other hybrids descended from the same two parents are known. Some are infertile, such as S. x
baxteri. Other fertile hybrids are also known, including S. vulgaris var. hibernicus, now common in Britain,
and the allohexaploid S. cambrensis, which according to molecular evidence probably originated
independently at least three times in different locations. Morphological and genetic evidence support the
status of S. eboracensis as separate from other known hybrids.[100]
[edit] Evidence from artificial selection
The Chihuahua mix and Great Dane illustrate the range of sizes among dog breeds.
Artificial selection demonstrates the diversity that can exist among organisms that share a relatively
recent common ancestor. In artificial selection, one species is bred selectively at each generation, allowing
only those organisms that exhibit desired characteristics to reproduce. These characteristics become
increasingly well-developed in successive generations. Artificial selection was successful long before science
discovered the genetic basis. Examples of artificial selection would be dog breeding, genetically modified
food, flower breeding, cultivation of foods such as wild cabbage,[101] and others.
[edit] Evidence from computation and mathematical iteration

"It has taken more than five decades, but the electronic computer is now powerful enough to simulate
evolution"[102] assisting bioinformatics in its attempt to solve biological problems.
Computer science allows the iteration of self changing complex systems to be studied, allowing a
mathematical understanding of the nature of the processes behind evolution; providing evidence for the
hidden causes of known evolutionary events. The evolution of specific cellular mechanisms like
spliceosomes that can turn the cell's genome into a vast workshop of billions of interchangeable parts that
can create tools that create tools that create tools that create us can be studied for the first time in an exact
way.
For example, Christoph Adami et al. make this point in Evolution of biological complexity :
To make a case for or against a trend in the evolution of complexity in biological evolution,
complexity needs to be both rigorously defined and measurable. A recent information-theoretic (but
intuitively evident) definition identifies genomic complexity with the amount of information a sequence
stores about its environment. We investigate the evolution of genomic complexity in populations of
digital organisms and monitor in detail the evolutionary transitions that increase complexity. We show
that, because natural selection forces genomes to behave as a natural "Maxwell Demon," within a
fixed environment, genomic complexity is forced to increase.[103]
For example, David J. Earl and Michael W. Deem make this point in Evolvability is a selectable trait:
Not only has life evolved, but life has evolved to evolve. That is, correlations within protein
structure have evolved, and mechanisms to manipulate these correlations have evolved in tandem.
The rates at which the various events within the hierarchy of evolutionary moves occur are not random
or arbitrary but are selected by Darwinian evolution. Sensibly, rapid or extreme environmental change
leads to selection for greater evolvability. This selection is not forbidden by causality and is strongest
on the largest-scale moves within the mutational hierarchy. Many observations within evolutionary
biology, heretofore considered evolutionary happenstance or accidents, are explained by selection for
evolvability. For example, the vertebrate immune system shows that the variable environment of
antigens has provided selective pressure for the use of adaptable codons and low-fidelity polymerases
during somatic hypermutation. A similar driving force for biased codon usage as a result of
productively high mutation rates is observed in the hemagglutinin protein of influenza A.[104]
"Computer simulations of the evolution of linear sequences have demonstrated the importance of
recombination of blocks of sequence rather than point mutagenesis alone. Repeated cycles of point
mutagenesis, recombination, and selection should allow in vitro molecular evolution of complex sequences,
such as proteins."[105] Evolutionary molecular engineering, also called directed evolution or in vitro
molecular evolution involves the iterated cycle of mutation, multiplication with recombination, and selection of
the fittest of individual molecules (proteins, DNA, and RNA). Natural evolution can be relived showing us
possible paths from catalytic cycles based on proteins to based on RNA to based on DNA.[105][106][107]
[108]
[edit] See also

• Nothing in Biology Makes Sense Except in the Light of Evolution
Book:Evolution
[edit] References
1. ^ Mount DM. (2004). Bioinformatics: Sequence and Genome Analysis (2nd ed.). Cold Spring
Harbor Laboratory Press: Cold Spring Harbor, NY.. ISBN 0-87969-608-7.
2. ^ http://www.imls.uzh.ch/research/noll/publ/Dev_Cell_2003_5_773_785.pdf
3. ^ Two sources:
Chen FC, Li WH (February 2001). "Genomic divergences between humans and other hominoids and
the effective population size of the common ancestor of humans and chimpanzees". Am J Hum
Genet. 68 (2): 444–56. doi:10.1086/318206. PMID 11170892.
Cooper GM, Brudno M, Green ED, Batzoglou S, Sidow A (May 2003). "Quantitative estimates of
sequence divergence for comparative analyses of mammalian genomes". Genome Res. 13 (5): 813–
20. doi:10.1101/gr.1064503. PMID 12727901. PMC 430923.
http://www.genome.org/cgi/content/full/13/5/813.
4. ^ The picture labeled "Human Chromosome 2 and its analogs in the apes" in the article
Comparison of the Human and Great Ape Chromosomes as Evidence for Common Ancestry is
literally a picture of a link in humans that links two separate chromosomes in the nonhuman apes
creating a single chromosome in humans. It is considered a missing link, and the ape-human
connection is of particular interest. Also, while the term originally referred to fossil evidence, this too
is a trace from the past corresponding to some living beings which when alive were the physical
embodiment of this link.
5. ^ The New York Times report Still Evolving, Human Genes Tell New Story, based on A Map
of Recent Positive Selection in the Human Genome, states the International HapMap Project is
"providing the strongest evidence yet that humans are still evolving" and details some of that
evidence.
6. ^ "Converging Evidence for Evolution." Phylointelligence: Evolution for Everyone. Web. 26
Nov. 2010. <http://phylointelligence.org/combined.html>.
7. ^ Petrov DA, Hartl DL (2000). "Pseudogene evolution and natural selection for a compact
genome". J Hered. 91 (3): 221–7. doi:10.1093/jhered/91.3.221. PMID 10833048.
http://jhered.oxfordjournals.org/cgi/pmidlookup?view=long&pmid=10833048.
8. ^ http://www.genomesize.com/
9. ^ Okamoto N, Inouye I (2005). "A secondary symbiosis in progress". Science 310 (5746):
287. doi:10.1126/science.1116125. PMID 16224014.
10.^ Okamoto N, Inouye I (October 2006). "Hatena arenicola gen. et sp. nov., a katablepharid
undergoing probable plastid acquisition". Protist 157 (4): 401–19. doi:10.1016/j.protis.2006.05.011.
PMID 16891155.
11.^ Human Chromosome 2 is a fusion of two ancestral chromosomes by Alec MacAndrew;
accessed 18 May 2006.
12.^ Evidence of Common Ancestry: Human Chromosome 2 (video) 2007
13.^ Yunis and Prakash; Prakash, O (1982). "The origin of man: a chromosomal pictorial
legacy". Science 215 (4539): 1525–1530. doi:10.1126/science.7063861. PMID 7063861.
14.^ Human and Ape Chromosomes; accessed 8 September 2007.
15.^ Avarello et al.; Pedicini, A; Caiulo, A; Zuffardi, O; Fraccaro, M (1992). "Evidence for an
ancestral alphoid domain on the long arm of human chromosome 2". Human Genetics 89 (2): 247–9.
doi:10.1007/BF00217134. PMID 1587535.
16.^ a b IJdo et al.; Baldini, A; Ward, DC; Reeders, ST; Wells, RA (1991). "Origin of human
chromosome 2: an ancestral telomere-telomere fusion". Proceedings of the National Academy of
Sciences 88 (20): 9051–5. doi:10.1073/pnas.88.20.9051. PMID 1924367.
17.^ a b Amino acid sequences in cytochrome c proteins from different species, adapted from
Strahler, Arthur; Science and Earth History, 1997. page 348.
18.^ Lurquin PF, Stone L (2006). Genes, Culture, and Human Evolution: A Synthesis. Blackwell
Publishing, Incorporated. pp. 79. ISBN 1-4051-5089-0. http://books.google.com/?
id=zdeWdF_NQhEC&pg=PA79&lpg=PA79&dq=chimpanzee+rhesus+cytochrome+c.
19.^ 29+ Evidences for Macroevolution; Protein functional redundancy, Douglas Theobald,
Ph.D.
20.^ Pallen, Mark (2009). Rough Guide to Evolution. Rough Guides. pp. 200–206. ISBN 978-1-
85828-946-5.
21.^ a b http://www.talkorigins.org/faqs/comdesc/section1.html#nested_hierarchy
22.^ Coyne, Jerry A. (2009). Why Evolution is True. Viking. pp. 8–11. ISBN 978-0-670-02053-9.
23.^ Charles Darwin (1859). On the Origin of Species. John Murray. p. 420.
24.^ Natan Slifkin (2006). The Challenge of Creation.... Zoo Torah. pp. 258–9. ISBN 1-933143-
15-0.
25.^ http://www.springerlink.com/content/mr73x80w2x7744jp/fulltext.pdf
26.^ Douglas J. Futuyma (1998). Evolutionary Biology (3rd ed.). Sinauer Associates Inc..
pp. 122. ISBN 0-87893-189-9.
27.^ a b c TalkOrigins Archive. "29+ Evidences for Macroevolution: Part 2".
http://www.talkorigins.org/faqs/comdesc/section2.html#atavisms. Retrieved 2006-11-08.
28.^ http://animals.howstuffworks.com/animal-facts/atavism.htm
29.^ a b c http://universe-review.ca/I10-10-snake.jpg
30.^ http://www.edwardtbabinski.us/whales/atavisms.html
31.^ Tyson, Reid; Graham, John P.; Colahan, Patrick T.; Berry, Clifford R. (July 2004). "Skeletal
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• Biological science, Oxford, 2002.
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ISBN 978-0-19-923084-6.
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Selection or the Preservation of Favoured Races in the Struggle for Life . London: John Murray,
Albemarle Street. 502 pages. Reprinted: Gramercy (May 22, 1995). ISBN 0-517-12320-7
• Dawkins, Richard (2009). The Greatest Show on Earth: The Evidence for Evolution . Bantam
Press. ISBN 978-1-4165-9478-9.
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ISBN 0-691-08387-8.
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Massachusetts. (dated 1998, published 1997) ISBN 0-87893-189-9
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everyday life. Cambridge, UK: Cambridge University Press. ISBN 0-521-33115-3.
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• TalkOrigins Archive - 29+ Evidences for Macroevolution: The Scientific Case for Common
Descent
• TalkOrigins Archive - Transitional Vertebrate Fossils FAQ
• Understanding Evolution: Your one-stop source for information on evolution
• National Academy Press: Teaching About Evolution and the Nature of Science
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• Evolution News from Genome News Network (GNN)
• Evolution by Natural Selection — An introduction to the logic of the theory of evolution by
natural selection
• 15 Evolutionary Gems
Processes of
Population genetic
Evolutionary
developmental
concepts

Modes of

Retrieved from "http://en.wikipedia.org/wiki/Evidence_of_common_descent"
Categories: Creationist objections to evolution | Evolutionary biology

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Biological life cycle

(Redirected from Biological life history)
A life cycle is a period involving all different generations of a species succeeding each other through
means of reproduction, whether through asexual reproduction or sexual reproduction (a period from one
generation of organisms to the same identical). For example, a complex life cycle of Fasciola hepatica
includes three different multicellular generations: 1) "adult" hermaphroditic; 2) sporocyst; 3) redia.
In regard to changes of ploidy, there are 3 types of cycles:
• haplontic life cycle
• diplontic life cycle
• diplobiontic life cycle (also referred to as diplohaplontic, haplodiplontic, or dibiontic life cycle)
These three types of cycles feature alternating haploid and all germinates. To return to a haploid
stage, meiosis must occur (see Cell division). The cycles differ in the product of meiosis, and whether mitosis
(growth) occurs. Zygotic and gametic meioses have one mitotic stage and form: during the n phase in zygotic
meiosis and during the 2n phase in gametic meiosis. Therefore, zygotic and gametic meiosis are collectively
term haplobiontic (single mitosis per phase). Sporic meiosis, on the other hand, has two mitosis events
(diplobiontic): one in each phase.
Contents
[hide]
• 1 Haplontic life cycle
• 2 Diplontic life cycle
• 3 Haplodiplontic life cycle
• 4 Life history
• 5 Life history theory
• 6 See also
• 7 References
[edit] Haplontic life cycle
Zygotic meiosis
A zygotic meiosis is a meiosis of a zygote immediately after karyogamy, which is the fusion of two
cell nuclei. This way, the organism ends its diploid phase and produces several haploid cells. These cells
divide mitotically to form either larger, multicellular individuals, or more haploid cells. Two opposite types of
gametes (e.g., male and female) from these individuals or cells fuse to become a zygote.
In the whole cycle, zygotes are the only diploid cell; mitosis occurs only in the haploid phase.
The individuals or cells as a result of mitosis are haplonts, hence this life cycle is also called
haplontic life cycle. Haplonts are:
• Most fungi
• Some green algae
• Many protozoa
[edit] Diplontic life cycle
Gametic meiosis
In gametic meiosis, instead of immediately dividing meiotically to produce haploid cells, the zygote
divides mitotically to produce a multicellular diploid individual or a group of more unicellular diploid cells.
Cells from the diploid individuals then undergo meiosis to produce haploid cells or gametes. Haploid cells
may divide again (by mitosis) to form more haploid cells, as in many yeasts, but the haploid phase is not the
predominant life cycle phase. In most diplonts, mitosis occurs only in the diploid phase, i.e. gametes usually
form quickly and fuse to produce diploid zygotes.
In the whole cycle, gametes are usually the only haploid cells, and mitosis usually occurs only in the
diploid phase.
The diploid multicellular individual is a diplont, hence a gametic meiosis is also called a diplontic life
cycle. Diplonts are:
• Animals
• Some brown algae
• Some fungi, e.g. brewer's yeast
[edit] Haplodiplontic life cycle
Sporic meiosis
In sporic meiosis (also commonly known as intermediary meiosis), the zygote divides mitotically to
produce a multicellular diploid "sporophyte". The sporophyte creates spores via meiosis which also then
divide mitotically producing haploid individuals called "gametophytes". Gametophytes now produce gametes
via mitosis. In many plants the gametophyte is not only small-sized but also short-lived.
In the whole cycle, gametes are usually the only haploid cells, and mitosis usually occurs only in the
diploid phase.
Haplodiplonts are:
• Most plants
• Some fungi
[edit] Life history

The term "Life history" is better than "life cycle" especially for the red algae (Rhodophyta). In the
Rhodophyceae there are generally three stages and thus "alternation of generations" is not suitable.[1]
[edit] Life history theory

In animal and human biology, life history theory is a method of understanding evolved behaviors and
strategies to optimize reproductive success.
[edit] See also

Strobilanthes callosus, a 8 years haplodiplontic
[edit] References
1. ^ Dixon, P.S. 1973. Biology of the Rhodophyta. Oliver & oyd

Wikimedia Commons has media related to: Life cycles
Retrieved from "http://en.wikipedia.org/wiki/Biological_life_cycle"
Categories: Biology | Reproduction

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18S ribosomal RNA


18S ribosomal RNA (abbreviated 18S rRNA) is a part of the ribosomal RNA. The S in 18S represents
Svedberg units. 18S rRNA is a component of the small eukaryotic ribosomal subunit (40S). 18S rRNA is the
structural RNA for the small component of eukaryotic cytoplasmic ribosomes, and thus one of the basic
components of all eukaryotic cells.
It is the eukaryotic nuclear homologue of 16S ribosomal RNA, which exists only in Prokaryotes.
18S rRNA data is widely used in molecular analysis to reconstruct the evolutionary history of
organisms, especially invertebrates, as its slow evolutionary rate makes it suitable to reconstruct ancient
divergences.
[edit] Uses in phylogeny

The small subunit (SSU) 18S rRNA gene is one of the most frequently used genes in phylogenetic
studies and an important marker for random target polymerase chain reaction (PCR) in environmental
biodiversity screening.[1] In general, rRNA gene sequences are easy to access due to highly conserved
flanking regions allowing for the use of universal primers.[1] Their repetitive arrangement within the genome
provides excessive amounts of template DNA for PCR, even in smallest organisms. The 18S gene is part of
the ribosomal functional core and is exposed to similar selective forces in all living beings. Thus, when the
first large-scale phylogenetic studies based on 18S sequences were published - first and foremost phylogeny
of the animal kingdom by Field et al. (1988)[2] - the gene was celebrated as the prime candidate for
reconstructing the metazoan tree of life.[1] And in fact, 18S sequences later provided evidence for the
splitting of Ecdysozoa and Lophotrochozoa, thus contributing to the most recent revolutionary change in our
understanding of metazoan relationships.[1]
During recent years and with increased numbers of taxa included into molecular phylogenies,
however, two problems became apparent. First, there are prevailing sequencing impediments in
representatives of certain taxa, such as the mollusk classes Solenogastres and Tryblidia, selected bivalve
taxa, and the enigmatic crustacean class Remipedia.[1] Failure to obtain 18S sequences of single taxa is
considered a common phenomenon but is rarely ever reported.[1] Secondly, in contrast to initially high
hopes, 18S cannot resolve nodes at all taxonomic levels and its efficacy varies considerably among clades.
This has been discussed as an effect of rapid ancient radiation within short periods. Multigene analyses are
currently thought to give more reliable results for tracing deep branching events in Metazoa but 18S still is
extensively used in phylogenetic analyses.[1]
[edit] References
This article incorporates CC-By-2.0 text from the reference.[1]
1. ^ a b c d e f g h Meyer A., Todt C., Mikkelsen N. T. & Lieb B. (2010). "Fast evolving 18S rRNA
sequences from Solenogastres (Mollusca) resist standard PCR amplification and give new insights
into mollusk substitution rate heterogeneity". BMC Evolutionary Biology 10: 70. doi:10.1186/1471-
2148-10-70
2. ^ Field K. G., Olsen G. J., Lane D. J., Giovannoni S. J., Ghiselin M. T., Raff E. C., Pace N. R.
& Raff R. A. (1988). "Molecular phylogeny of the animal kingdom". Science 239(4841): 748-753.
doi:10.1126/science.3277277.
[hide]v · d · eRibosomal RNA / ribosome subunits
Prokaryotes (70S) Large: 50S (5S, 23S)

Small: 30S (16S)
Eukaryotes (80S) Large: 60S (5S, 5.8S, 28S)

Cytoplasmic
Small: 40S (18S)
MT-RNR1, 12S · MT-RNR2,
Mitochondrial
16S
B bsyn: dna (repl, cycl, reco, repr) · tscr (fact, tcrg, nucl, rnat, rept, ptts) · tltn (risu, pttl, nexn) · dnab,
rnab/runp · stru (domn, 1°, 2°, 3°, 4°)
Retrieved from "http://en.wikipedia.org/wiki/18S_ribosomal_RNA"
Categories: Ribosomal RNA

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Hans-Dieter Sues

Hans-Dieter Sues is a German-born paleontologist who is Senior Scientist and Curator of Vertebrate
Paleontology at the National Museum of Natural History of the Smithsonian Institution in Washington, DC. He
received his education at Johannes Gutenberg-Universitaet, University of Alberta, and Harvard University
(Ph.D., 1984). Before assuming his present position, Sues worked at the Royal Ontario Museum and the
University of Toronto and at the Carnegie Museum of Natural History in Pittsburgh. He is interested in the
diversity, paleoecology, and evolutionary history of Mesozoic tetrapods, especially archosaurian reptiles and
cynodont therapsids, and the history of biology and paleontology. Sues has discovered a number of new
dinosaurs and other extinct terrestrial vertebrates in Mesozoic continental strata around the world. He has
authored or co-authored over 100 articles and book chapters on vertebrate paleontology and paleoecology.
Sues has edited Evolution of Herbivory in Terrestrial Vertebrates (Cambridge Univ. Press, 2000) and co-
edited Terrestrial Ecosystems through Time (with A. K. Behrensmeyer et al.; Univ. of Chicago Press, 1992),
In the Shadow of the Dinosaurs: Early Mesozoic Tetrapods (with N. C. Fraser; Cambridge Univ. Press,
1994), and Major Transitions in Vertebrate Evolution (with J. S. Anderson; Indiana Univ. Press, 2007). He is
also active in promoting the value of natural history collections for addressing major questions in current
science. Sues was elected a Fellow of the Royal Society of Canada in 2003 and a Fellow of the American
Association for the Advancement of Science in 1998.[1][2]
1. ^ Who's Who in Canada. Univ. of Toronto Press, Toronto, 1997-present.
2. ^ http://www.mnh.si.edu/about/sues.html
Retrieved from "http://en.wikipedia.org/wiki/Hans-Dieter_Sues"
Categories: American paleontologists | Harvard University alumni | German immigrants to the United
States | Living people
W000
Geologic time scale


This clock representation shows some of the major units of geological time and definitive events of
Earth history. The Hadean eon represents the time before fossil record of life on Earth; its upper boundary is
now regarded as 4.0 Ga.[1] Other subdivisions reflect the evolution of life; the Archean and Proterozoic are
both eons, the Palaeozoic, Mesozoic and Cenozoic are eras of the Phanerozoic eon. The two million year
Quaternary period, the time of recognizable humans, is too small to be visible at this scale.
The geologic time scale provides a system of chronologic measurement relating stratigraphy to time
that is used by geologists, paleontologists and other earth scientists to describe the timing and relationships
between events that have occurred during the history of the Earth. The table of geologic time spans
presented here agrees with the dates and nomenclature proposed by the International Commission on
Stratigraphy, and uses the standard color codes of the United States Geological Survey.[citation needed]
Evidence from radiometric dating indicates that the Earth is about 4.570 billion years old. The
geological or deep time of Earth's past has been organized into various units according to events which took
place in each period. Different spans of time on the time scale are usually delimited by major geological or
paleontological events, such as mass extinctions. For example, the boundary between the Cretaceous period
and the Paleogene period is defined by the Cretaceous–Tertiary extinction event, which marked the demise of
the dinosaurs and of many marine species. Older periods which predate the reliable fossil record are defined
by absolute age.
Each era on the scale is separated from the next by a major event or change.
Contents
[hide]
• 1 Terminology
• 2 Graphical timelines
• 3 History of the time scale and names
• 4 Table of geologic time
• 5 See also
• 6 References and footnotes
[edit] Terminology
e h
Units in geochronology and stratigraphy[2]
Segments of rock (strata) in Periods of time in
Notes
chronostratigraphy geochronology
4 total, half a billion years or

Eonothem Eon more
Erathem Era 12 total, several hundred

million years
System Period
Series Epoch tens of millions of years
Stage Age millions of years
smaller than an age/stage,

Chronozone Chron not used by the ICS timescale
The largest defined unit of time is the supereon, composed of eons. Eons are divided into eras,
which are in turn divided into periods, epochs and ages. The terms eonothem, erathem, system, series, and
stage are used to refer to the layers of rock that correspond to these periods of geologic time.
Geologists qualify these units as Early, Middle, and Late when referring to time, and Lower, Mid, and
Upper when referring to the corresponding rocks. For example, the Lower Jurassic Series in geochronology
corresponds to the Early Jurassic Epoch in chronostratigraphy.[3] The adjectives are capitalized when the
subdivision is formally recognized, and lower case when not; thus "early Miocene" but "Early Jurassic."
Geologic units from the same time but different parts of the world often look different and contain
different fossils, so the same period was historically given different names in different locales. For example,
in North America the Lower Cambrian is called the Waucoban series that is then subdivided into zones
based on succession of trilobites. In East Asia and Siberia, the same unit is split into Alexian, Atdabanian,
and Botomian stages. A key aspect of the work of the International Commission on Stratigraphy is to
reconcile this conflicting terminology and define universal horizons that can be used around the world.[4]
[edit] Graphical timelines
The second and third timelines are each subsections of their preceding timeline as indicated by
asterisks.
Millions of Years
The Holocene (the latest epoch) is too short to be shown clearly on this timeline.
[edit] History of the time scale and names

Main articles: History of geology and History of paleontology
Animation showing Earth's palaeogeographic reconstruction beginning from early Cambrian period.
Diagram of geological time scale, where the past is toward the bottom of the spiral
In classical antiquity, Aristotle saw that fossil seashells from rocks were similar to those found on the
beach and inferred that the fossils were once part of living animals. He reasoned that the positions of land
and sea had changed over long periods of time. Leonardo da Vinci concurred with Aristotle's view that fossils
were the remains of ancient life.[5]
The 11th-century Persian geologist Avicenna (Ibn Sina) examined various fossils and inferred that
they originated from the petrifaction of plants and animals.[6] He also first proposed one of the principles
underlying geologic time scales: the law of superposition of strata. While discussing the origins of mountains
in The Book of Healing in 1027, he outlined the principle as follows:
It is also possible that the sea may have happened to flow little by little over the land consisting
of both plain and mountain, and then have ebbed away from it. ... It is possible that each time
the land was exposed by the ebbing of the sea a layer was left, since we see that some
mountains appear to have been piled up layer by layer, and it is therefore likely that the clay
from which they were formed was itself at one time arranged in layers. One layer was formed
first, then at a different period, more layers were formed and piled, upon the first, and so on.
Over each layer there spread a substance of different material, which formed a partition
between it and the next layer; but when petrification took place something occurred to the
partition which caused it to break up and disintegrate from between the layers [possibly referring
to unconformity]. ... As to the beginning of the sea, its clay is either sedimentary or primeval, the
latter not being sedimentary. It is probable that the sedimentary clay was formed by the
disintegration of the strata of mountains. Such is the formation of mountains.[7]
Later in the 11th century, the Chinese naturalist, Shen Kuo (1031–1095), also recognized the concept
of 'deep time'.[8]
The principles underlying geologic (geological) time scales were later laid down by Nicholas Steno in
the late 17th century. Steno argued that rock layers (or strata) are laid down in succession, and that each
represents a "slice" of time. He also formulated the law of superposition, which states that any given stratum
is probably older than those above it and younger than those below it. While Steno's principles were simple,
applying them to real rocks proved complex. Over the course of the 18th century geologists realized that:
1. Sequences of strata were often eroded, distorted, tilted, or even inverted after deposition;
2. Strata laid down at the same time in different areas could have entirely different
appearances;
3. The strata of any given area represented only part of the Earth's long history.
A comparative geological timescale

The first serious attempts to formulate a geological time scale that could be applied anywhere on
Earth were made in the late 18th century. The most influential of those early attempts (championed by
Abraham Werner, among others) divided the rocks of the Earth's crust into four types: Primary, Secondary,
Tertiary, and Quaternary. Each type of rock, according to the theory, formed during a specific period in Earth
history. It was thus possible to speak of a "Tertiary Period" as well as of "Tertiary Rocks." Indeed, "Tertiary"
(now Paleocene-Pliocene) and "Quaternary" (now Pleistocene-Holocene) remained in use as names of
geological periods well into the 20th century.
The Neptunist theories popular at this time (expounded by Werner) proposed that all rocks had
precipitated out of a single enormous flood. A major shift in thinking came when James Hutton presented his
Theory of the Earth; or, an Investigation of the Laws Observable in the Composition, Dissolution, and
Restoration of Land Upon the Globe before the Royal Society of Edinburgh in March and April 1785. It has
been said that "as things appear from the perspective of the twentieth century, James Hutton in those
reading became the founder of modern geology"[9] Hutton proposed that the interior of the Earth was hot,
and that this heat was the engine which drove the creation of new rock: land was eroded by air and water and
deposited as layers in the sea; heat then consolidated the sediment into stone, and uplifted it into new lands.
This theory was dubbed "Plutonist" in contrast to the flood-oriented theory.
The identification of strata by the fossils they contained, pioneered by William Smith, Georges
Cuvier, Jean d'Omalius d'Halloy, and Alexandre Brogniart in the early 19th century, enabled geologists to
divide Earth history more precisely. It also enabled them to correlate strata across national (or even
continental) boundaries. If two strata (however distant in space or different in composition) contained the
same fossils, chances were good that they had been laid down at the same time. Detailed studies between
1820 and 1850 of the strata and fossils of Europe produced the sequence of geological periods still used
today.
The process was dominated by British geologists, and the names of the periods reflect that
dominance. The "Cambrian," (the Roman name for Wales) and the "Ordovician," and "Silurian", named after
ancient Welsh tribes, were periods defined using stratigraphic sequences from Wales.[10] The "Devonian"
was named for the English county of Devon, and the name "Carboniferous" was simply an adaptation of "the
Coal Measures," the old British geologists' term for the same set of strata. The "Permian" was named after
Perm, Russia, because it was defined using strata in that region by Scottish geologist Roderick Murchison.
However, some periods were defined by geologists from other countries. The "Triassic" was named in 1834
by a German geologist Friedrich Von Alberti from the three distinct layers (Latin trias meaning triad) —red
beds, capped by chalk, followed by black shales— that are found throughout Germany and Northwest Europe,
called the 'Trias'. The "Jurassic" was named by a French geologist Alexandre Brogniart for the extensive
marine limestone exposures of the Jura Mountains. The "Cretaceous" (from Latin creta meaning 'chalk') as a
separate period was first defined by Belgian geologist Jean d'Omalius d'Halloy in 1822, using strata in the
Paris basin[11] and named for the extensive beds of chalk (calcium carbonate deposited by the shells of
marine invertebrates).
British geologists were also responsible for the grouping of periods into Eras and the subdivision of
the Tertiary and Quaternary periods into epochs.
When William Smith and Sir Charles Lyell first recognized that rock strata represented successive
time periods, time scales could be estimated only very imprecisely since various kinds of rates of change
used in estimation were highly variable. While creationists had been proposing dates of around six or seven
thousand years for the age of the Earth based on the Bible, early geologists were suggesting millions of
years for geologic periods with some even suggesting a virtually infinite age for the Earth. Geologists and
paleontologists constructed the geologic table based on the relative positions of different strata and fossils,
and estimated the time scales based on studying rates of various kinds of weathering, erosion,
sedimentation, and lithification. Until the discovery of radioactivity in 1896 and the development of its
geological applications through radiometric dating during the first half of the 20th century (pioneered by such
geologists as Arthur Holmes) which allowed for more precise absolute dating of rocks, the ages of various
rock strata and the age of the Earth were the subject of considerable debate.
The first geologic time scale was eventually published in 1913 by the British geologist Arthur Holmes.
[12] He greatly furthered the newly created discipline of geochronology and published the world renowned
book The Age of the Earth in 1913 in which he estimated the Earth's age to be at least 1.6 billion years.[13]
In 1977, the Global Commission on Stratigraphy (now the International Commission on Stratigraphy)
started an effort to define global references (Global Boundary Stratotype Sections and Points) for geologic
periods and faunal stages. The commission's most recent work is described in the 2004 geologic time scale
of Gradstein et al.[14] A UML model for how the timescale is structured, relating it to the GSSP, is also
available.[15]
[edit] Table of geologic time

The following table summarizes the major events and characteristics of the periods of time making
up the geologic time scale. As above, this time scale is based on the International Commission on
Stratigraphy. (See lunar geologic timescale for a discussion of the geologic subdivisions of Earth's moon.)
This table is arranged with the most recent geologic periods at the top, and the most ancient at the bottom.
The height of each table entry does not correspond to the duration of each subdivision of time.
The content of the table is based on the current official geologic time scale of the International
Commission on Stratigraphy,[16] with the epoch names altered to the early/late format from lower/upper as
recommended by the ICS when dealing with chronostratigraphy.[3]
v · d · e Geologic time scale[hide]
Supereon Eon Era Period[17] Epoch Ag
Phanerozoic Cenozoic[19] Quaternary Holocene None
chrons:
Subatlantic
Subboreal
Atlantic
Boreal
Preboreal
Pleistocene Taran
(Tyrrhenian
Stage/Eemian
onian)
Ionian
Calab
Gelas
Piace
ncan
Pliocene
Zancl
Neogene
Miocene Messi
Torton
Serra
Langh
Burdig
Aquita
Paleogene Chatti
Oligocene
Rupel
Eocene Priabo
Barton
Luteti
Ypres
Paleocene Thane
Selan
Dania
Mesozoic Cretaceous Maas
Camp
Santo
Late
Conia
Turon
Cenom
Early Albian
Aptian
Barre
Haute
Valan
Berria
Tithon
Late Kimm
Oxfor
Callov
Batho
Jurassic Middle
Bajoc
Aalen
Toarc
Early Pliens
Sinem
Hettan
Triassic Rhaet
Late Norian
Carnia
Ladin
Middle
Anisia
Early Olene
Indua
Paleozoic Permian Chang
Lopingian
Wuch
Capita
Wordi
Guadalupian ian
Roadi
n
Cisuralian Kungu
Artins
Sakm
Carbon- Gzhel
iferous[22]/ Late
Pennsyl- Kasim
vanian
Middle Mosco
Early Bashk
Carbon- Late Serpu
iferous[22]/
Missis- Middle Viséa
sippian
Early Tourn
Devonian Fame
Late
Frasn
Giveti
Middle
Eifelia
Early Emsia
Pragia
Lochk
Silurian no fau
Pridoli
stages defined
Ludfo
Ludlow/Cayu
gan
Gorst
Wenlock Home
portian
Shein
Tonawandan
Telych
rian
Aeron
Llandovery/
Alexandrian
Rhudd
Hirnan
Late
other
stages
Ordovician
Darriw
Middle
other
stages
Arenig
Early
Trema
Cambrian other
stages
Furongian Paibia
/
Ayusokkanian
Aksayan
other
Middle
stages/Alberta
Early other
stages/
Waucoban/To
Atdabanian/B
Good fossils of the first mult
worldwide in seas. Simple trace foss
etc. First sponges and trilobitomorph
Ediacaran jellied creatures shaped like bags, d
Orogeny in North America. Aravalli R
Beginning of Petermann Orogeny on
Orogeny in Antarctica, 633–620 Ma.
Possible "Snowball Earth" p

Neo-
Cryogenian landmass begins to break up. Late R
proterozoic[24]
Precam- Proter- tapers off.
brian[23] ozoic[24] Rodinia supercontinent pers
eukaryotes. First radiation of dinofla
tapers off in North America. Pan-Afr
Tonian Nimrod Orogeny in Antarctica, 1000
- 850 Ma), Gascoyne Complex, Wes
laid down on Australian Continent, b
(Delamerian Orogeny) in that contin
Narrow highly metamorphic

Meso-
Stenian Late Ruker / Nimrod Orogeny in Ant
proterozoic[24]
Orogeny (c. 1080 Ma), Musgrave Bl
Platform covers continue to
Ectasian
seas. Grenville Orogeny in North Am
Platform covers expand. Ba

Calymmian Northern Australia, and Isan Orogen
Queensland
First complex single-celled l

primordial supercontinent. Kimban O
Statherian Yapungku Orogeny on Yilgarn crato
Orogeny, 1680–1620 Ma, on the Gas
Kararan Orogeny (1650-Ma), Gawle
The atmosphere become ox

Paleo- asteroid impacts. Much orogeny. Pe
proterozoic[24] Orosirian Orogenies in North America. Early R
Ma. Glenburgh Orogeny, Glenburgh
1920 Ma. Kimban Orogeny, Gawler
Rhyacian Bushveld Formation forms.
Oxygen catastrophe: bande

Siderian
Orogeny on Australian Continent, G
Archean[24] Neoarchean[2 Stabilization of most modern cratons; possible m

4] 2650 ± 150 Ma. Abitibi greenstone belt in present-day On
stablizes by 2600 Ma.
First stromatolites (probably colonial cyanobacte

Mesoarchean[
Orogeny in Antarctica. Blake River Megacaldera Comple
24]
and Quebec, ends by roughly 2696 Ma.
First known oxygen-producing bacteria. Oldest d

Paleoarchean[
Earth (such as the Canadian Shield and the Pilbara Crato
24]
period[26]. Rayner Orogeny in Antarctica.
Eoarchean[24] Simple single-celled life (probably bacteria and a
Early Indirect photosynthetic evidence (e.g., kerogen) o

Imbrian[24][28] end of the Late Heavy Bombardment of the inner solar sy
Nectarian[24] This unit gets its name from the lunar geologic tim
[28] other major lunar basins form by large impact events.
Hadean
[24][27] Oldest known rock (4030 Ma)[29]. The first life fo
Basin
evolve around 4000 Ma after the Late Heavy Bombardme
Groups[24][28]
Antarctica, 4000 ± 200 Ma.
Oldest known mineral (Zircon, 4406 ± 8 Ma[30]).

Cryptic[24][28]
from giant impact. Formation of Earth (4567.17 to 4570 M
[edit] See also
• Age of the Earth
• Anthropocene/Homogenocene
• Deep time
• Geological history of Earth
• Geology of Mars/areology
• Graphical timeline of the universe
• History of the Earth
• Logarithmic timeline
• Lunar geologic timescale/selenological timescale
• Natural history
• New Zealand geologic time scale
• Timeline of the Big Bang
• Timeline of the geologic history of the United States
• Timeline of the Precambrian
[edit] References and footnotes
1. ^ International Commission on Stratigraphy 2008:http://www.stratigraphy.org/column.php?
id=Chart/Time%20Scale, retrieved 9 March 2009.
2. ^ International Commission on Stratigraphy. "International Stratigraphic Chart".
http://www.stratigraphy.org/upload/ISChart2009.pdf. Retrieved 2009-09-25.
3. ^ a b International Commission on Stratigraphy. "Chronostratigraphic Units." International
Stratigraphic Guide. Accessed 14-DEC-2009. [1]
4. ^ Statutes of the International Commission on Stratigraphy, retrieved 26 November 2009
5. ^ Correlating Earth's History, Paul R. Janke
6. ^ Rudwick, M. J. S. (1985). The Meaning of Fossils: Episodes in the History of
Palaeontology. University of Chicago Press. p. 24. ISBN 0226731030.
7. ^ Quoted in The contribution of Ibn Sina (Avicenna) to the development of the Earth
Sciences, among other sources
8. ^ Sivin, Nathan (1995). Science in Ancient China: Researches and Reflections. Brookfield,
Vermont: Ashgate Publishing Variorum series. III, 23–24.
9. ^ John McPhee, Basin and Range, New York:Farrar, Straus and Giroux, 1981, pp.95-100.
10.^ John McPhee, Basin and Range, pp.113-114.
11.^ (in Russian) Great Soviet Encyclopedia (3rd ed.). Moscow: Sovetskaya Enciklopediya.
1974. vol. 16, p. 50.
12.^ Geologic Time Scale
13.^ How the discovery of geologic time changed our view of the world, Bristol University
14.^ Felix M. Gradstein, James G. Ogg, Alan G. Smith (Editors); A Geologic Time Scale 2004,
Cambridge University Press, 2005, (ISBN 0-521-78673-8)
15.^ Cox & Richard, A formal model for the geologic time scale and global stratotype section
and point, compatible with geospatial information transfer standards, Geosphere, volume 1, pp 119-
137, Geological Society of America, 2005
16.^ International Commission on Stratigraphy. "International Stratigraphic Chart".
http://www.stratigraphy.org/upload/ISChart2009.pdf. Retrieved 2009-09-25.
17.^ Paleontologists often refer to faunal stages rather than geologic (geological) periods. The
stage nomenclature is quite complex. For an excellent time-ordered list of faunal stages, see "The
Paleobiology Database". http://flatpebble.nceas.ucsb.edu/cgi-bin/bridge.pl?action=startScale.
18.^ a b Dates are slightly uncertain with differences of a few percent between various sources
being common. This is largely due to uncertainties in radiometric dating and the problem that
deposits suitable for radiometric dating seldom occur exactly at the places in the geologic column
where they would be most useful. The dates and errors quoted above are according to the
International Commission on Stratigraphy 2004 time scale. Dates labeled with a * indicate
boundaries where a Global Boundary Stratotype Section and Point has been internationally agreed
upon: see List of Global Boundary Stratotype Sections and Points for a complete list.
19.^ a b Historically, the Cenozoic has been divided up into the Quaternary and Tertiary sub-
eras, as well as the Neogene and Paleogene periods. The 2009 version of the ICS time chart
recognizes a slightly extended Quaternary as well as the Paleogene and a truncated Neogene, the
Tertiary having been demoted to informal status.
20.^ a b c d e f For more information on this, see the following articles: Earth's atmosphere,
carbon dioxide, Carbon dioxide in the Earth's atmosphere, Image:Phanerozoic_Carbon_Dioxide.png,
Image:65 Myr Climate Change.png, Image:Five Myr Climate Change.png, and Template:DF
temperature
21.^ The start time for the Holocene epoch is here given as 11,430 years ago ± 130 years (that
is, between 9610 BC-9560 BC and 9350 BC-9300 BC). For further discussion of the dating of this
epoch, see Holocene.
22.^ a b In North America, the Carboniferous is subdivided into Mississippian and Pennsylvanian
Periods.
23.^ The Precambrian is also known as Cryptozoic.
24.^ a b c d e f g h i j k l m n The Proterozoic, Archean and Hadean are often collectively referred to
as the Precambrian Time or sometimes, also the Cryptozoic.
25.^ a b c d e f g h i j k l Defined by absolute age (Global Standard Stratigraphic Age).
26.^ The age of the oldest measurable craton, or continental crust, is dated to 3600–3800 Ma
27.^ Though commonly used, the Hadean is not a formal eon and no lower bound for the
Archean and Eoarchean have been agreed upon. The Hadean has also sometimes been called the
Priscoan or the Azoic. Sometimes, the Hadean can be found to be subdivided according to the lunar
geologic time scale. These eras include the Cryptic and Basin Groups (which are subdivisions of the
pre-Nectarian era), Nectarian, and Early Imbrian units.
28.^ a b c d These unit names were taken from the Lunar geologic timescale and refer to
geologic events that did not occur on Earth. Their use for Earth geology is unofficial.
29.^ Bowring, Samuel A.; Williams, Ian S. (1999). "Priscoan (4.00–4.03 Ga) orthogneisses from
northwestern Canada". Contributions to Mineralogy and Petrology 134: 3.
doi:10.1007/s004100050465. The oldest rock on Earth is the Acasta Gneiss, and it dates to 4.03 Ga,
located in the Northwest Territories of Canada.
30.^ http://www.geology.wisc.edu/%7Evalley/zircons/Wilde2001Nature.pdf
• NASA: Geologic Time
• GSA: Geologic Time Scale
• British Geological Survey: Geological Timechart
• GeoWhen Database
• International Commission on Stratigraphy Time Scale
• CHRONOS
• National Museum of Natural History - Geologic Time
• SeeGrid: Geological Time Systems Information model for the geologic time scale
• Exploring Time from Planck Time to the lifespan of the universe
• [2] Gradstein, Felix M. et al. (2004) A new Geologic Time Scale, with special reference to
Precambrian and Neogene, Episodes, Vol. 27, no. 2 June 2004 (pdf)
• Lane, Alfred C,, and Marble, John Putman 1937. Report of the Committee on the
measurement of geologic time
Hadean


[show] Phanerozoic (542 – 0 Mya)
[show] Paleozoic (542 – 251 Mya)
521 Mya)
Cambrian
(542 – 488.3 Mya)
Age 10* (492 – 488.3 Mya)

Floian (478.6 – 471.8 Mya)
Silurian
(426.2 – 422.9 Mya)
(443.7 – 416 Mya)
418.7 Mya)

(416 – 359.2 Mya) (391.8 – 385.3 Mya)
(374.5 – 359.2 Mya)

Carboniferous
(359.2 – 299 Mya)
(303.4 – 299 Mya)
Cisuralian (299 – 270.6 Mya): Asselian (299 – 294.6 Mya) · Sakmarian
(294.6 – 284.4 Mya) · Artinskian (284.4 – 275.6 Mya) · Kungurian (275.6 – 270.6
Mya)
Permian
(299 – 251 Mya)
[show] Mesozoic (251 – 65.5 Mya)
(249.5 – 245.9 Mya)
(251 – 199.6 Mya) 228.7 Mya)
203.6 Mya) · Rhaetian (203.6 – 199.6 Mya)
Jurassic Early Jurassic (199.6 – 175.6 Mya): Hettangian (199.6 – 196.5 Mya) ·
(199.6 – 145.5 Mya) Sinemurian (196.5 – 189.6 Mya) · Pliensbachian (189.6 – 183 Mya) · Toarcian (183
– 175.6 Mya)
Middle Jurassic (175.6 – 161.2 Mya): Aalenian (175.6 – 171.6 Mya) · Bajocian
(171.6 – 167.7 Mya) · Bathonian (167.7 – 164.7 Mya) · Callovian (164.7 – 161.2
Mya)

[show] Cenozoic (65.5 – 0 Mya)
Paleogene Paleocene (65.5 – 55.8 Mya): Danian (65.5 – 61.1 Mya) · Selandian (61.1 –
(65.5 – 23.03 Mya) 58.7 Mya) · Thanetian (58.7 – 55.8 Mya)
Eocene (55.8 – 33.9 Mya): Ypresian (55.8 – 48.6 Mya) · Lutetian (48.6 – 40.4 Mya) ·
Bartonian (40.4 – 37.2 Mya) · Priabonian (37.2 – 33.9 Mya)
Mya)
Miocene (23.03 – 5.332 Mya): Aquitanian (23.03 – 20.43 Mya) · Burdigalian

Neogene
(23.03 – 2.588 Mya)
Mya)

Quaternary
(2.588 – 0 Mya)
Dryas* (12.8 – 11.4 kya)
February 2008.
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W000
Law of Complexity/Consciousness
This article relies largely or entirely upon a single source. Please help improve this
article by introducing appropriate citations to additional sources. (January 2011)
This article needs references that appear in reliable third-party publications.
Primary sources or sources affiliated with the subject are generally not sufficient for a
Wikipedia article. Please add more appropriate citations from reliable sources. (January
2011)
The Law of Complexity/Consciousness is the tendency in matter to become more complex over time
and at the same time to become more conscious. The law was first formulated by Jesuit priest and
paleontologist Pierre Teilhard de Chardin.
Teilhard holds that at all times and everywhere, matter is endeavoring to complexify upon itself, as
observed in the evolutionary history of the earth. Matter complexified from inanimate matter, to plant-life, to
animal-life, to human-life. Or, from the geosphere, to the biosphere, to the noosphere (of which humans
represented, because of their possession of a consciousness which reflects upon themselves). As evolution
rises through the geosphere, biosphere, and noosphere, matter continues to rise in a continual increase of
both complexity and consciousness.
For Teilhard, the Law of Complexity/Consciousness continues to run today in the form of the
socialization of mankind. The closed and circular surface of the earth contributes to the increased
compression (socialization) of mankind. As human beings continue to come into closer contact with one
another, their methods of interaction continue to complexify in the form of better organized social networks,
which contributes to an overall increase in consciousness, or the noosphere.
Teilhard imagines a critical threshold, Omega Point, in which mankind will have reached its highest
point of complexification (socialization) and thus its highest point of consciousness. At this point
consciousness will rupture through time and space and assert itself on a higher plane of existence from
which it can not come back.
Interestingly, for Teilhard, because the Law of Complexity/Consciousness runs everywhere and at all
times, and because of the immensity of both time and space in outer space, and the immensity of the
chances for matter to find the right conditions to complexify upon itself, it is highly probable that life exists,
has existed, and will exist in the universe apart from our earth.
[edit] See also

• Autopoesis
• Fifth force
• Involution (esoterism)
• Moore's law
• Spiral Dynamics
• Technological singularity
[edit] Quotes
"The more complex a being is, so our Scale of Complexity tells us, the more it is centered upon itself
and therefore the more aware does it become. In other words, the higher the degree of complexity in a living
creature, the higher its consciousness; and vice versa. The two properties vary in parallel and
simultaneously. If we depict them in diagrammatic form, they are equivalent and interchangeable."
--Pierre Teilhard de Chardin, The Future of Man, p.111
"For its reflective and inventive forward spring it is in some sort necessary that Life, duplicating its
evolutionary motive center, should henceforth be sustained by two centers of action, separate and conjoined,
one of consciousness and the other of complexity.... In hominised evolution the Physical and the Psychic, the
Without and the Within, Matter and Consciousness, are all found to be functionally linked in one tangible
process."
--Pierre Teilhard de Chardin, The Future of Man, p.209
Retrieved from "http://en.wikipedia.org/wiki/Law_of_Complexity/Consciousness"
Categories: Holism | Integral thought | Phenomenology | Metaphysical cosmology

W000
Timeline of plant evolution


This article attempts to place key plant innovations in a geological context. It concerns itself only with
novel adaptations and events that had a major ecological significance, not those that are of solely
anthropological interest. The timeline displays a graphical representation of the adaptations; the text attempts
to explain the nature and robustness of the evidence.
Life on Earth
-4500 —
–
-4000 —
–
-3500 —
–
-3000 —
–
-2500 —
–
-2000 —
–
-1500 —
–
-1000 —
–
-500 —
–
0—
Life?
Photosynthesis?
Eukaryotes
Complex multicellular life
Animals
Land plants
"Dinosaurs"
Mammals
Flowers
←
Formation
of Earth
←
Meteorite bombardment
←
Atmospheric oxygen
←
Ediacara biota
←
Cambrian explosion
←
Modern-looking
humans
Dates prior to 1 billion years ago are speculative.
Plant evolution is an aspect of the study of biological evolution, involving predominantly evolution of
plants suited to live on land, greening of various land masses by the filling of their niches with land plants,
and diversification of groups of land plants.
Contents
[hide]
• 1 Earliest classifiable plants
• 2 Palæozoic flora
• 2.1 Cambrian flora
• 2.2 Ordovician flora
• 2.3 Silurian flora
• 2.4 Devonian flora
• 2.5 Carboniferous Flora
• 2.6 Permian flora
• 3 Mesozoic flora
• 3.1 Triassic flora
• 3.2 Jurassic flora
• 3.3 Cretaceous flora
• 4 Cenozoic flora
• 5 Species Differentiation
• 6 See also
• 8 References
[edit] Earliest classifiable plants
In the strictly modern sense, the name plant refers to the biological classification kingdom Plantae.
However, other photosynthetic organisms, including protists, green algae, and cyanobacteria have
evolutionary significance to modern plants. While this article is directly about the evolutionary history of the
Plant kingdom, these other organisms provide clues to the evolution of all photosynthetic organisms. All of
these organisms - plants, green algae, and the protists - are primary photosynthetic eukaryotic organisms.
Scientists start the search for fossil evidence of plants with indirect evidence for their presence, the
evidence of photosynthesis in the geological record. The evidence for photosynthesis in the rock record is
varied, but primary evidence comes from around 3000 Ma, in rock records and fossil evidence of
cyanobacteria, photosynthesizing prokaryotic organisms. Cyanobacteria use water as a reducing agent,
thereby producing atmospheric oxygen as a byproduct, and profoundly changing the early reducing
atmosphere of the earth to one in which modern aerobic organisms eventually evolved. This oxygen liberated
by cyanobacteria then oxidized dissolved iron in the oceans, the iron precipitated out of the sea water, and
fell to the ocean floor to form sedimentary layers of oxidized iron called Banded Iron Formations (BIFs).
These BIFs are part of the geological record of evidence for the evolutionary history of plants by identifying
when photosynthesis originated. This also provides deep time constraints upon when enough oxygen could
have been available in the atmosphere to produce the ultraviolet blocking stratospheric ozone layer. The
oxygen concentration in the ancient atmosphere subsequently rose, acting as a poison for anaerobic
organisms, and resulting in a highly oxidizing atmosphere, and opening up niches on land for occupation by
aerobic organisms.
Evidence for the cyanobacteria also comes from the presence of stromatolites in the fossil record
deep into the Precambrian. Stromatolites are layered structures thought to have been formed by the trapping,
binding, and cementation of sedimentary grains by microorganisms, such as cyanobacteria. The direct
evidence for cyanobacteria is less certain than the evidence for their presence as primary producers of
atmospheric oxygen. Modern day stromatoloid structures containing cyanobacteria can be found on the west
coast of Australia.
Chloroplasts in eukaryotic plants evolved from an endosymbiotic relationship between cyanobacteria
and other prokaryotic organisms producing the lineage that eventually led to photosynthesizing eukaryotic
organisms in marine and freshwater environments. These earliest photosynthesizing single-celled autotrophs
later led to organisms such as Charophyta, a group of freshwater green algae.
Key innovations in
early plant evolution
-460 —
–
-450 —
–
-440 —
–
-430 —
–
-420 —
–
-410 —
–
-400 —
–
-390 —
–
-380 —
–
-370 —
–
-360 —
O
S
D
Pragian
Emsian
Eifelian
Givetian
Frasnian
Famennian
Lochkovian
Wenlock
Llandovery
Pridoli
Ludlow
Sandbian
Katian
Hirnantian
←
Spore tetrads
←
Rhynie chert
←
Sporangia[1]
←
Stems[1]
←
Roots[1]
←
Seeds[1]
←
Wood[1]
←
Leaves[1]
[edit] Palæozoic flora

[edit] Cambrian flora
Early plants were small, unicellular or filamentous, composed mostly of soft body tissues, with simple
branching. The identification of plant tissues in Cambrian strata is an uncertain area in the evolutionary
history of plants because of the small and soft-bodied nature of these plants. It is also difficult in a fossil of
this age to distinguish among various similar appearing groups with simple branching patterns, and not all of
these groups are plants. One exception to the uncertainty of fossils from this age is the calcareous green
algae, Dasycladales found in the fossil record since the middle Cambrian. This algae does not belong to the
lineage that is ancestral to the land plants. Other major groups of green algae had been established by this
time. Generally it is accepted that there were no land plants with vascular tissues at this time although some
Biologists believe that the molecular clock points to an earlier Cambrian or perhaps Precambrian origin
because the molecular clock states that land plants appeared around 480–440 Ma and fungi appeared on
land around 1 Ga but however there is debate over whether the fossil evidence supports this interpretation of
the molecular clock.
[edit] Ordovician flora

The evidence for plant evolutionary history changes dramatically in the Ordovician with the first
extensive appearance of spores in the fossil record (Cambrian spores have been found, also). The first
terrestrial plants appeared in the form of tiny plants resembling liverworts when, around the Middle
Ordovician, evidence for the beginning of the terrestrialization of the land is found.[2] These early plants did
not have conducting tissues, severely limiting their size. They were, in effect, tied to wet terrestrial
environments by their inability to conduct water, like extant liverworts, hornworts, and mosses, although they
reproduced with spores, important dispersal units that have hard protective outer coatings, allowing for their
preservation in the fossil record, in addition to protecting the future offspring against the desiccating
environment of life on land. With spores, plants on land could have sent out large numbers of spores that
could grow into an adult plant when sufficient environmental moisture was present.
[edit] Silurian flora
Artist's impression of Cooksonia

The first fossil records of vascular plants, that is, land plants with vascular tissues, appeared in the
Silurian period. The earliest known representatives of this group are Cooksonia (mostly from the northern
hemisphere) and Baragwanathia (from Australia). A primitive Silurian land plant with xylem and phloem but
no differentiation in root, stem or leaf, was much-branched Psilophyton, reproducing by spores and breathing
through stomata on every surface, and probably photosynthesizing in every tissue exposed to light.
Rhyniophyta and primitive lycopods were other land plants that first appear during this period.
[edit] Devonian flora

By the Devonian Period, life was well underway in its colonization of the land. The bacterial and algal
mats were joined early in the period by primitive plants that created the first recognizable soils and harbored
some arthropods like mites, scorpions and myriapods. Early Devonian plants did not have roots or leaves like
the plants most common today, and many had no vascular tissue at all. They probably spread largely by
vegetative growth, and did not grow much more than a few centimeters tall.
By the Late Devonian, forests of small, primitive plants existed: lycophytes, sphenophytes, ferns, and
progymnosperms had evolved. Most of these plants have true roots and leaves, and many were quite tall.
The tree-like ancestral fern Archaeopteris and the giant cladoxylopsid trees grew as a large tree with true
wood. These are the oldest known trees of the world's first forests. Prototaxites was the fruiting body of an
enormous fungus that stood more than 8 meter tall. By the end of the Devonian, the first seed-forming plants
had appeared. This rapid appearance of so many plant groups and growth forms has been called the
"Devonian Explosion". The primitive arthropods co-evolved with this diversified terrestrial vegetation
structure. The evolving co-dependence of insects and seed-plants that characterizes a recognizably modern
world had its genesis in the late Devonian. The development of soils and plant root systems probably led to
changes in the speed and pattern of erosion and sediment deposition.
The 'greening' of the continents acted as a carbon dioxide sink, and atmospheric levels of this
greenhouse gas may have dropped. This may have cooled the climate and led to a massive extinction event.
see Late Devonian extinction.
Also in the Devonian, both vertebrates and arthropods were solidly established on the land.
[edit] Carboniferous Flora
Stigmaria, a fossil tree root. Upper Carboniferous of northeastern Ohio.

External mold of Lepidodendron from the Upper Carboniferous of Ohio.
Early Carboniferous land plants were very similar to those of the preceding Latest Devonian, but new
groups also appeared at this time.
The main Early Carboniferous plants were the Equisetales (Horse-tails), Sphenophyllales
(scrambling plants), Lycopodiales (Club mosses), Lepidodendrales (scale trees), Filicales (Ferns),
Medullosales (previously included in the "seed ferns", an artificial assemblage of a number of early
gymnosperm groups) and the Cordaitales. These continued to dominate throughout the period, but during
late Carboniferous, several other groups, Cycadophyta (cycads), the Callistophytales (another group of "seed
ferns"), and the Voltziales (related to and sometimes included under the conifers), appeared.
The Carboniferous lycophytes of the order Lepidodendrales, which are cousins (but not ancestors) of
the tiny club-moss of today, were huge trees with trunks 30 meters high and up to 1.5 meters in diameter.
These included Lepidodendron (with its fruit cone called Lepidostrobus), Halonia, Lepidophloios and
Sigillaria. The roots of several of these forms are known as Stigmaria.
The fronds of some Carboniferous ferns are almost identical with those of living species. Probably
many species were epiphytic. Fossil ferns and "seed ferns" include Pecopteris, Cyclopteris, Neuropteris,
Alethopteris, and Sphenopteris; Megaphyton and Caulopteris were tree ferns.
The Equisetales included the common giant form Calamites, with a trunk diameter of 30 to 60 cm and
a height of up to 20 meters. Sphenophyllum was a slender climbing plant with whorls of leaves, which was
probably related both to the calamites and the lycopods.
Cordaites, a tall plant (6 to over 30 meters) with strap-like leaves, was related to the cycads and
conifers; the catkin-like inflorescence, which bore yew-like berries, is called Cardiocarpus. These plants were
thought to live in swamps and mangroves. True coniferous trees ( Walchia, of the order Voltziales) appear
later in the Carboniferous, and preferred higher drier ground.
[edit] Permian flora

The Permian began with the Carboniferous flora still flourishing. About the middle of the Permian
there was a major transition in vegetation. The swamp-loving lycopod trees of the Carboniferous, such as
Lepidodendron and Sigillaria, were replaced by the more advanced conifers, which were better adapted to
the changing climatic conditions. Lycopods and swamp forests still dominated the South China continent
because it was an isolated continent and it sat near or at the equator. Oxygen levels were probably high
there. The Permian saw the radiation of many important conifer groups, including the ancestors of many
present-day families. The ginkgos and cycads also appeared during this period. Rich forests were present in
many areas, with a diverse mix of plant groups. The gigantopterids thrived during this time; some of these
may have been part of the ancestral flowering plant lineage, though flowers evolved only considerably later.
[edit] Mesozoic flora
[edit] Triassic flora
On land, the holdover plants included the lycophytes, the dominant cycads, ginkgophyta
(represented in modern times by Ginkgo biloba) and glossopterids. The spermatophytes, or seed plants
came to dominate the terrestrial flora: in the northern hemisphere, conifers flourished. Glossopteris (a seed
fern) was the dominant southern hemisphere tree during the Early Triassic period.
[edit] Jurassic flora

The arid, continental conditions characteristic of the Triassic steadily eased during the Jurassic
period, especially at higher latitudes; the warm, humid climate allowed lush jungles to cover much of the
landscape.[3] Conifers dominated the flora, as during the Triassic; they were the most diverse group and
constituted the majority of large trees. Extant conifer families that flourished during the Jurassic included the
Araucariaceae, Cephalotaxaceae, Pinaceae, Podocarpaceae, Taxaceae and Taxodiaceae.[4] The extinct
Mesozoic conifer family Cheirolepidiaceae dominated low latitude vegetation, as did the shrubby
Bennettitales.[5] Cycads were also common, as were ginkgos and tree ferns in the forest. Smaller ferns were
probably the dominant undergrowth. Caytoniaceous seed ferns were another group of important plants
during this time and are thought to have been shrub to small-tree sized.[6] Ginkgo-like plants were
particularly common in the mid- to high northern latitudes. In the Southern Hemisphere, podocarps were
especially successful, while Ginkgos and Czekanowskiales were rare.[7],[5]
[edit] Cretaceous flora
Artist's restoration of Archaeamphora longicervia, the earliest known carnivorous plant

Flowering plants, also known as angiosperms, spread during this period, although they did not
become predominant until near the end of the period (Campanian age). Their evolution was aided by the
appearance of bees; in fact angiosperms and insects are a good example of coevolution. The first
representatives of many modern trees, including figs, planes and magnolias, appeared in the Cretaceous. At
the same time, some earlier Mesozoic gymnosperms, like Conifers continued to thrive, although other taxa
like Bennettitales died out before the end of the period.
[edit] Cenozoic flora

The Cenozoic is just as much the age of savannas, or the age of co-dependent flowering plants and
insects. At 35 Ma, grasses evolved from among the angiosperms. About ten thousand years ago, humans in
the Fertile Crescent of the Middle East develop agriculture. Plant domestication begins with cultivation of
Neolithic founder crops. This process of food production, coupled later with the domestication of animals
caused a massive increase in human population that has continued to the present. In Jericho (modern
Israel), there is a settlement with about 19,000 people. At the same time, Sahara is green with rivers, lakes,
cattles, crocodiles and monsoons. At 8 ka, Common (Bread) wheat ( Triticum aestivum) originates in
southwest Asia due to hybridisation of emmer wheat with a goat-grass, Aegilops tauschii. At 6.5 ka, two rice
species are domesticated: Asian rice, Oryza sativa, and African rice Oryza glaberrima.
[edit] Species Differentiation

• Development of rooted plants
• Flowering plants vs. Conifers
• Ferns and other primitive plants
• Borderline species such as coliform protists
[edit] See also
• Plant
• Flora
• Paleobotany
• Plant evolutionary developmental biology

• Interactive Plant Evolution Timeline - from the University of Cambridge Ensemble Project
[edit] References
1. ^ a b c d e f Labandeira, C. (2007). "The origin of herbivory on land: Initial patterns of plant
tissue consumption by arthropods". Insect Science 14 (4): 259-275. doi:10.1111/j.1744-
7917.2007.00152.x.
3. ^ Haines, Tim. 2000. Walking with Dinosaurs: A Natural History , (New York: Dorling
Kindersley Publishing, Inc.) ISBN 0-563-38449-2. Page 65.
4. ^ Behrensmeyer, Anna K., Damuth, J.D., DiMichele, W.A., Potts, R., Sues, H.D. & Wing, S.L.
(eds.). 1992. Terrestrial Ecosystems through Time: the Evolutionary Paleoecology of Terrestrial
Plants and Animals, (Chicago & London: University of Chicago Press), ISBN 0-226-04154-9 (cloth),
ISBN 0-226-04155-7 (paper). Page 349.
5. ^ a b Behrensmeyer et al., 1992, 352
6. ^ Behrensmeyer et al., 1992, 353
7. ^ Haines, 2000.

Plant physiology



Transpiration
Sporophyte


morphology terms
Category · Portal
Retrieved from "http://en.wikipedia.org/wiki/Timeline_of_plant_evolution"
Categories: Evolution by taxon | Biology timelines | Botany | Plant taxonomy | Prehistoric plants |
Plants
W000
History of botany

Some traditional tools of botanical science
The history of botany examines the human effort to understand life on Earth by tracing the historical
development of the discipline of botany—that part of natural science dealing with organisms traditionally
treated as plants.
Rudimentary botanical science began with empirically-based plant lore passed from generation to
generation in the oral traditions of paleolithic hunter-gatherers. The first written records of plants were made
in the Neolithic Revolution about 10,000 years ago as writing was developed in the settled agricultural
communities where plants and animals were first domesticated. The first writings that show human curiosity
about plants themselves, rather than the uses that could be made of them, appears in the teachings of
Aristotle's student Theophrastus at the Lyceum in ancient Athens in about 350 BC; this is considered the
starting point for modern botany. In Europe, this early botanical science was soon overshadowed by a
medieval preoccupation with the medicinal properties of plants that lasted more than 1000 years. During this
time, the medicinal works of classical antiquity were reproduced in manuscripts and books called herbals. In
China and the Arab world, the Greco-Roman work on medicinal plants was preserved and extended.
In Europe the Renaissance of the 14th–17th centuries heralded a scientific revival during which
botany gradually emerged from natural history as an independent science, distinct from medicine and
agriculture. Herbals were replaced by Floras: books that described the native plants of local regions. The
invention of the microscope stimulated the study of plant anatomy, and the first carefully designed
experiments in plant physiology were performed. With the expansion of trade and exploration beyond
Europe, the many new plants being discovered were subjected to an increasingly rigorous process of
naming, description, and classification.
Progressively more sophisticated scientific technology has aided the development of contemporary
botanical offshoots in the plant sciences, ranging from the applied fields of economic botany (notably
agriculture, horticulture and forestry), to the detailed examination of the structure and function of plants and
their interaction with the environment over many scales from the large-scale global significance of vegetation
and plant communities (biogeography and ecology) through to the small scale of subjects like cell theory,
molecular biology and plant biochemistry.
Contents
[hide]
• 1 Introduction
• 2 Ancient and medieval knowledge
• 2.1 Plant lore and plant selection
• 2.2 Early botany
• 2.3 Theophrastus and the origin of botanical science
• 2.4 Ancient Rome
• 2.5 Medicinal plants of the early Middle Ages
• 2.6 The Age of Herbals
• 3 1550–1800 The European Renaissance and after
• 3.1 Botanical gardens and herbaria
• 3.2 From Herbal to Flora
• 3.3 Botanical exploration
• 3.4 Classification and morphology
• 3.5 Anatomy
• 3.6 Physiology
• 3.7 Plant sexuality
• 4 Nineteenth century foundations of modern botany
• 4.1 Plant geography and ecology
• 4.2 Anatomy
• 4.3 Water relations
History of science
[edit] Introduction
Main article: Outline of botany
Botany (Greek Βοτάνη - grass, fodder; Medieval Latin botanicus –
herb, plant)[1] and zoology are, historically, the core disciplines of biology
whose history is closely associated with the natural sciences chemistry,
physics and geology. A distinction can be made between botanical science
in a pure sense, as the study of plants themselves, and botany as applied
science, which studies the human use of plants. Early natural history
divided pure botany into three main streams morphology-classification,
anatomy and physiology – that is, external form, internal structure, and
functional operation.[2] The most obvious topics in applied botany are
horticulture, forestry and agriculture although there are many others like Background
weed science, plant pathology, floristry, pharmacognosy, economic botany
Theories/sociology
and ethnobotany which lie outside modern courses in botany. Since the
origin of botanical science there has been a progressive increase in the Historiography
scope of the subject as technology has opened up new techniques and
Pseudoscience
areas of study. Modern molecular systematics, for example, entails the
principles and techniques of taxonomy, molecular biology, computer By era
science and more.
In early cultures
Within botany there are a number of sub-disciplines that focus on
particular plant groups, each with their own range of related studies in Classical Antiquity
(anatomy, morphology etc.). Included here are: phycology (algae), In the Middle Ages
pteridology (ferns), bryology (mosses and liverworts) and palaeobotany
(fossil plants) and their histories are treated elsewhere (see side bar). To In the Renaissance
Scientific Revolution
Romanticism in science
By culture
Islamic
this list can be added mycology, the study of fungi, which were once treated as plants, but are now ranked as
a unique kingdom.
[edit] Ancient and medieval knowledge

Main article: Neolithic Revolution
Nomadic hunter-gatherer societies passed on, by oral tradition, what they knew (their empirical
observations) about the different kinds of plants that they used for food, shelter, poisons, medicines, for
ceremonies and rituals etc. The uses of plants by these pre-literate societies influenced the way the plants
were named and classified—their uses were embedded in folk-taxonomies, the way they were grouped
according to use in everyday communication.[3] The nomadic life-style was drastically changed when settled
communities were established in about twelve centres around the world during the Neolithic Revolution
which extended from about 10,000 to 2500 years ago depending on the region. With these communities
came the development of the technology and skills needed for the domestication of plants and animals and
the emergence of the written word provided evidence for the passing of systematic knowledge and culture
from one generation to the next.[4]
[edit] Plant lore and plant selection

Further information: Cultigen taxonomy and Herbal
A Sumerian harvester's sickle dated to 3000 BC
During the Neolithic Revolution plant knowledge increased most obviously through the use of plants
for food and medicine. All of today's staple foods were domesticated in prehistoric times as a gradual process
of selection of higher-yielding varieties took place, possibly unknowingly, over hundreds to thousands of
years. Legumes were cultivated on all continents but cereals made up most of the regular diet: rice in East
Asia, wheat and barley in the Middle east, and maize in Central and South America. By Greco-Roman times
popular food plants of today, including grapes, apples, figs, and olives, were being listed as named varieties
in early manuscripts.[5] Botanical authority William Stearn has observed that "cultivated plants are mankind’s
most vital and precious heritage from remote antiquity ”.[6]
It is also from the Neolithic, in about 3000 BCE, that we glimpse the first known illustrations of
plants[7] and read descriptions of impressive gardens in Egypt.[8] However protobotany, the first pre-
scientific written record of plants, did not begin with food; it was born out of the medicinal literature of Egypt,
China, Mesopotamia and India.[9] Botanical historian Alan Morton notes that agriculture was the occupation
of the poor and uneducated, while medicine was the realm of socially influential shamans, priests,
apothecaries, magicians and physicians, who were more likely to record their knowledge for posterity.[10]
[edit] Early botany

Ancient India
An early example of ancient Indian plant classification is found in the Rigveda, a collection of Vedic
Sanskrit hymns from about 3700–3100 BP. Plants are divided into vṛska (trees), osadhi (herbs useful to
humans) and virudha (creepers), with further subdivisions. The sacred Hindu text Atharvaveda divides plants
into eight classes: visakha (spreading branches), manjari (leaves with long clusters), sthambini (bushy
plants), prastanavati (which expands); ekasṛnga (those with monopodial growth), pratanavati (creeping
plants), amsumati (with many stalks), and kandini (plants with knotty joints). The Taittiriya Samhita classifies
the plant kingdom into vṛksa, vana and druma (trees), visakha (shrubs with spreading branches), sasa
(herbs), amsumali (spreading plant), vratati (climber), stambini (bushy plant), pratanavati (creeper), and
alasala (spreading on the ground). Other examples of early Indian taxonomy include Manusmriti, the Law
book of Hindus, which classifies plants into eight major categories. Elaborate taxonomies also occur in the
Charaka Samhitā, Sushruta Samhita and Vaisesika.[11]
Ancient China
In ancient China lists of different plants and herb concoctions for pharmaceutical purposes date back
to at least the time of the Warring States (481 BC-221 BC). Many Chinese writers over the centuries
contributed to the written knowledge of herbal pharmaceutics. The Han Dynasty (202 BC-220 AD) includes
the notable work of the Huangdi Neijing and the famous pharmacologist Zhang Zhongjing. There were also
the 11th century scientists and statesmen Su Song and Shen Kuo who compiled learned treatises on natural
history, emphasising herbal medicine.[12]
[edit] Theophrastus and the origin of botanical science

Main article: Theophrastus
"School of Athens"
Fresco in Apostolic Palace, Rome, Vatican City, by Raphael 1509-1510
Ancient Athens, of the 6th century BCE, was the busy trade centre at the confluence of Egyptian,
Mesopotamian and Minoan cultures at the height of Greek colonisation of the Mediterranean. The
philosophical thought of this period ranged freely through many subjects. Empedocles (490–430 BCE)
foreshadowed Darwinian evolutionary theory in a crude formulation of the mutability of species and natural
selection.[13] The physician Hippocrates (460–370 BCE) avoided the prevailing superstition of his day and
approached healing by close observation and the test of experience. At this time a genuine non-
anthropocentric curiosity about plants emerged. The major works written about plants extended beyond the
description of their medicinal uses to the topics of plant geography, morphology, physiology, nutrition, growth
and reproduction.[14]
Foremost among the scholars studying botany was Theophrastus of Eressus (Greek: Θεόφραστος;
c. 371–287 BCE) who has been frequently referred to as the ”Father of Botany”. He was a student and close
friend of Aristotle (384–322 BCE) and succeeded him as head of the Lyceum (an educational establishment
like a modern university) in Athens with its tradition of peripatetic philosophy. Aristotle's special treatise on
plants — θεωρία περὶ φυτῶν — is now lost, although there are many botanical observations scattered
throughout his other writings (these have been assembled by Christian Wimmer in Phytologiae Aristotelicae
Fragmenta, 1836) but they give little insight into his botanical thinking.[15] The Lyceum prided itself in a
tradition of systematic observation of causal connections, critical experiment and rational theorizing.
Theophrastus challenged the superstitious medicine employed by the physicians of his day, called rhizotomi,
and also the control over medicine exerted by priestly authority and tradition.[16] Together with Aristotle he
had tutored Alexander the Great whose military conquests were carried out with all the scientific resources of
the day, the Lyceum garden probably containing many botanical trophies collected during his campaigns as
well as other explorations in distant lands.[17] It was in this garden where he gained much of his plant
knowledge.[18]
Statue of Theophrastus 371–287 BC
"Father of Botany"
Palermo Botanic Gardens
Theophrastus's major botanical works[19] were the Enquiry into Plants (Historia Plantarum) and
Causes of Plants (Causae Plantarum) which were his lecture notes for the Lyceum.[20] The opening
sentence of the Enquiry reads like a botanical manifesto: “We must consider the distinctive characters and
the general nature of plants from the point of view of their morphology, their behaviour under external
conditions, their mode of generation and the whole course of their life ”. The Enquiry is 9 books of "applied"
botany dealing with the forms and classification of plants and economic botany, examining the techniques of
agriculture (relationship of crops to soil, climate, water and habitat) and horticulture. He described some 500
plants in detail, often including descriptions of habitat and geographic distribution, and he recognised some
plant groups that can be recognised as modern-day plant families. Some names he used, like Crataegus,
Daucus and Asparagus have persisted until today. His second book Causes of Plants covers plant growth
and reproduction (akin to modern physiology).[21] Like Aristotle he grouped plants into "trees",
"undershrubs", "shrubs" and "herbs" but he also made several other important botanical distinctions and
observations. He noted that plants could be annuals, perennials and biennials, they were also either
monocotyledons or dicotyledons and he also noticed the difference between determinate and indeterminate
growth and details of floral structure including the degree of fusion of the petals, position of the ovary and
more.[22][23] These lecture notes of Theophrastus comprise the first clear exposition of the rudiments of
plant anatomy, physiology, morphology and ecology — presented in a way that would not be matched for
another eighteen centuries.[24]
Meanwhile the study of medicinal plants was not being neglected and a full synthesis of ancient
Greek pharmacology was compiled in Materia Medica c. 60 CE by Pedanius Dioscorides (c. 40-90 CE) who
was a Greek physician with the Roman army. This work proved to be the definitive text on medicinal herbs,
both oriental and occidental, for fifteen hundred years until the dawn of the European Renaissance being
slavishly copied again and again throughout this period.[25] Though rich in medicinal information with
descriptions of about 600 medicinal herbs, the botanical content of the work was extremely limited.[26]
[edit] Ancient Rome

Main article: Roman agriculture
The Romans contributed little to the foundations of botanical science laid by the ancient Greeks, but
made a sound contribution to our knowledge of applied botany as agriculture. In works titled De Re Rustica
the principles and practice of agriculture were discussed by writers Marcus Porcius Cato (234–149 BCE),
Marcus Varro (116–27 BCE) and, in particular, Columella (4–70 CE).[27] Roman encyclopaedist Pliny the
Elder (23–79 CE) deals with plants in Books 12 to 26 of his 37-volume highly influential work Naturalis
Historia in which he frequently quotes Theophrastus but with a lack of botanical insight although he does,
nevertheless, draw a distinction between true botany on the one hand, and farming and medicine on the
other.[28]
It is estimated that at the time of the Roman Empire between 1300 and 1400 plants had been
recorded in the West.[29]
[edit] Medicinal plants of the early Middle Ages

Further information: Herbalism, Chinese medicine, Byzantine medicine, and Islamic medicine
An Arabic copy of Avicenna's Canon of Medicine dated 1593
In Western Europe, after Theophrastus, botany passed through a bleak period of 1800 years when
little progress was made and, indeed, many of the early insights were lost. As Europe entered the Middle
Ages, a period of disorganised feudalism and indifference to learning, China, India and the Arab world
enjoyed a golden age. Chinese philosophy had followed a similar path to that of the ancient Greeks. The
Chinese dictionary-encyclopaedia Erh Ya probably dates from about 300 BCE and describes about 334
plants classed as trees or shrubs, each with a common name and illustration. Between 100 and 1700 CE
many new works on pharmaceutical botany were produced including encyclopaedic accounts and treatises
compiled for the Chinese imperial court. These were free of superstition and myth with carefully researched
descriptions and nomenclature; they included cultivation information and notes on economic and medicinal
uses — and even elaborate monographs on ornamental plants. But there was no experimental method and no
analysis of the plant sexual system, nutrition, or anatomy.[30]
The 400-year period from the 9th to 13th centuries CE was the Islamic Renaissance, a time when
Islamic culture and science thrived. Greco-Roman texts were preserved, copied and extended although new
texts always emphasised the medicinal aspects of plants. Kurdish biologist Ābu Ḥanīfah Āḥmad ibn Dawūd
Dīnawarī (828–896 CE) is known as the founder of Arabic botany; his Kitâb al-nabât (‘Book of Plants’)
describes 637 species, discussing plant development from germination to senescence and including details
of flowers and fruits.[31] The Mutazilite philosopher and physician Ibn Sina (Avicenna) (c. 980–1037 CE) was
another influential figure, his The Canon of Medicine being a landmark in the history of medicine treasured
until the Enlightenment.[32] In the early 13th century, the Andalusian-Arabian biologist Abu al-Abbas al-
Nabati developed an early scientific method for botany, introducing empirical and experimental techniques in
the testing, description and identification of numerous materia medica, and separating unverified reports from
those supported by actual tests and observations.[33] His student Ibn al-Baitar (circa, 1188–1248) was an
eminent Arab scientist. His Kitab al-Jami fi al-Adwiya al-Mufrada was a pharmacopoeia describing 1400
species, 300 discovered by himself. Translated into Latin in 1758 this was used in Europe until the early 19th
century[34] as a critical summing up of centuries of Arab pharmacology.[35]
In India simple artificial plant classification systems of the Rigveda, Atharvaveda and Taittiriya
Samhita became more botanical with the work of Parashara (c.400–500 CE), the author of Vṛksayurveda (the
science of life of trees). He made close observations of cells and leaves and divided plants into Dvimatrka
(Dicotyledons) and Ekamatrka (Monocotyledons). The dicotyledons were further classified into groupings
(ganas) akin to modern floral families: Samiganiya (Fabaceae), Puplikagalniya (Rutaceae), Svastikaganiya
(Cruciferae), Tripuspaganiya (Cucurbitaceae), Mallikaganiya (Apocynaceae), and Kurcapuspaganiya
(Asteraceae).[36][37] Important medieval Indian works of plant physiology include the Prthviniraparyam of
Udayana, Nyayavindutika of Dharmottara, Saddarsana-samuccaya of Gunaratna, and Upaskara of
Sankaramisra.
[edit] The Age of Herbals

Main article: Herbal
Dioscorides', De Materia Medica, Byzantium, 15th century.

In the European Middle Ages of the 15th and 16th centuries the lives of European citizens were
based around agriculture but when printing arrived, with movable type and woodcut illustrations, it was not
treatises on agriculture that were published, but lists of medicinal plants with descriptions of their properties
or "virtues". These first plant books, known as herbals showed that botany was still a part of medicine, as it
had been for most of ancient history.[32] Authors of herbals were often curators of university gardens,[38]
and most herbals were derivative compilations of classic texts, especially De Materia Medica. However, the
need for accurate and detailed plant descriptions meant that some herbals were more botanical than
medicinal. German Otto Brunfels's (1464–1534) Herbarum Vivae Icones (1530) contained descriptions of
about 47 species new to science combined with accurate illustrations. His fellow countryman Hieronymus
Bock's (1498–1554) Kreutterbuch of 1539 described plants he found in nearby woods and fields and these
were illustrated in the 1546 edition.[39] However, it was Valerius Cordus (1515–1544) who pioneered the
formal botanical description that detailed both flowers and fruits, some anatomy including the number of
chambers in the ovary, and the type of ovule placentation. He also made observations on pollen and
distinguished between inflorescence types.[39] His five-volume Historia Plantarum was published about 18
years after his early death aged 29 in 1561-1563. In Holland Rembert Dodoens (1517–1585), in Stirpium
Historiae (1583), included descriptions of many new species from the Netherlands in a scientific
arrangement[40] and in England William Turner (1515–1568) in his Libellus De Re Herbaria Novus (1538)
published names, descriptions and localities of many native British plants.[41]
Herbals contributed to botany by setting in train the science of plant description, classification, and
botanical illustration. Up to the 17th century botany and medicine were one and the same but those books
emphasising medicinal aspects eventually omitted the plant lore to become modern pharmacopoeias; those
that omitted the medicine became more botanical and evolved into the modern compilations of plant
descriptions we call Floras. These were often backed by specimens deposited in a herbarium which was a
collection of dried plants that verified the plant descriptions given in the Floras. The transition from herbal to
Flora marked the final separation of botany from medicine.[42]
[edit] 1550–1800 The European Renaissance and after

The revival of learning during the European Renaissance renewed interest in plants. The church,
feudal aristocracy and an increasingly influential merchant class that supported science and the arts, now
jostled in a world of increasing trade. Sea voyages of exploration returned botanical treasures to the large
public, private, and newly established botanic gardens, and introduced an eager population to novel crops,
drugs and spices from Asia, the East Indies and the New World.
The number of scientific publications increased. In England, for example, scientific communication
and causes were facilitated by learned societies like Royal Society (founded in 1660) and the Linnaean
Society (founded in 1788): there was also the support and activities of botanical institutions like the Chelsea
Physic Garden, Royal Botanic Gardens Kew, and the Oxford and Cambridge Botanic Gardens, as well as the
influence of renowned private gardens and wealthy entrepreneurial nurserymen.[43] Enlightenment values
were reinvigorating science and in botany a new phase of plant identification, nomenclature and description
began. By the early 17th century the number of plants described in Europe had risen to about 6000.[44]
[edit] Botanical gardens and herbaria
Preparing a herbarium specimen

Further information: Botanical garden, List of botanical gardens, and Herbarium
Public and private gardens have always been strongly associated with the historical unfolding of
botanical science. Early botanical gardens were physic gardens, repositories for the medicinal plants
described in the herbals. As they were generally associated with universities or other academic institutions
the plants were also used for study. The directors of these gardens were eminent physicians with an
educational role as “scientific gardeners” and it was staff of these institutions that produced many of the
published herbals.
A 16th century print of the Botanical Garden of Padova (Garden of the Simples) — the oldest
academic botanic garden that is still in its original location
The botanical gardens of the modern tradition were established in northern Italy, the first being at
Pisa (1544), founded by Luca Ghini (1490–1556). Although part of a medical faculty, the first chair of materia
medica, essentially a chair in botany, was established in Padua in 1533. Then in 1534, Ghini became Reader
in materia medica at Bologna University. Collections of pressed and dried specimens were called a hortus
siccus (garden of dry plants) and the first accumulation of plants in this way (including the use of a plant
press) is attributed to Ghini.[45][46] Buildings called herbaria housed these specimens mounted on card with
descriptive labels. Stored in cupboards in systematic order they could be preserved in perpetuity and easily
transferred or exchanged with other institutions, a taxonomic procedure that is still used today.
By the 18th century the physic gardens had been transformed into "order beds" that demonstrated
the classification systems that were being devised by botanists of the day — but they also had to
accommodate the influx of curious, beautiful and new plants pouring in from voyages of exploration that were
associated with European colonial expansion.
[edit] From Herbal to Flora

Main article: Flora
Plant classification systems of the 17th and 18th centuries now related plants to one-another and not
to man, marking a return to the non-anthropocentric botanical science promoted by Theophrastus over 1500
years before. This approach coupled with the new Linnaean system of binomial nomenclature resulted in
plant encyclopaedias without medicinal information called Floras that meticulously described and illustrated
the plants growing in particular regions.[47] The 17th century also marked the beginning of experimental
botany and application of a rigorous scientific method, while improvements in the microscope launched the
new discipline of plant anatomy whose foundations, laid by the careful observations of Englishman
Nehemiah Grew[48] and Italian Marcello Malpighi, would last for 150 years.[49]
[edit] Botanical exploration

Main article: Plant geography
More new lands were opening up to European colonial powers, the botanical riches being returned to
European botanists for description. This was a romantic era of botanical explorers, intrepid plant hunters and
gardener-botanists. Significant botanical collections came from: the West Indies (Hans Sloane (1660–1753));
China (James Cunningham); the spice islands of the East Indies (Moluccas, George Rumphius (1627–1702));
China and Mozambique (João de Loureiro (1717–1791)); West Africa (Michel Adanson (1727–1806)) who
devised his own classification scheme and forwarded a crude theory of the mutability of species; Canada,
Hebrides, Iceland, New Zealand by Captain James Cook's chief botanist Joseph Banks (1743–1820).[50]
[edit] Classification and morphology

Further information: List of systems of plant taxonomy, Plant taxonomy, and History of plant
systematics
Portrait of Carl Linnaeus by Alexander Roslin, 1775
By the middle of the 18th century the botanical booty resulting from the era of exploration was
accumulating in gardens and herbaria – and it needed to be systematically catalogued. This was the task of
the taxonomists, the plant classifiers.
Plant classifications have changed over time from "artificial" systems based on general habit and
form, to pre-evolutionary "natural" systems expressing similarity using one to many characters, leading to
post-evolutionary "natural" systems that use characters to infer evolutionary relationships.[51]
Italian physician Andrea Caesalpino (1519–1603) studied medicine and taught botany at the
University of Pisa for about 40 years eventually becoming Director of the Botanic Garden of Pisa from 1554
to 1558. His sixteen-volume De Plantis (1583) described 1500 plants and his herbarium of 260 pages and
768 mounted specimens still remains. Caesalpino moved away from characters of form and habit to those of
flower and fruit including the structure and morphology of seeds; he also applied the concept of the genus.
[52] He was the first to try and derive principles of natural classification reflecting the overall similarities
between plants and he produced a classification scheme well in advance of its day.[53] Gaspard Bauhin
(1560–1624) produced two influential publications Prodromus Theatrici Botanici (1620) and Pinax (1623).
These brought order to the 6000 species now described and in the latter he used binomials and synonyms
that may well have influenced Linnaeus's thinking. He also insisted that taxonomy should be based on
natural affinities.[54]
Cover page of Species Plantarum of Carl Linnaeus published in 1753
To sharpen the precision of description and classification Joachim Jung (1587–1657) compiled a
much-needed botanical terminology which has stood the test of time. English botanist John Ray (1623–1705)
built on Jung’s work to establish the most elaborate and insightful classification system of the day.[55] His
observations started with the local plants of Cambridge where he lived, with the Catalogus Stirpium circa
Cantabrigiam Nascentium (1860) which later expanded to his Synopsis Methodica Stirpium Britannicarum,
essentially the first British Flora. Although his Historia Plantarum (1682, 1688, 1704) provided a step towards
a world Flora as he included more and more plants from his travels, first on the continent and then beyond.
He extended Caesalpino’s natural system with a more precise definition of the higher classification levels,
deriving many modern families in the process,and asserted that all parts of plants were important in
classification. He recognised that variation arises from both internal (genotypic) and external environmental
(phenotypic) causes and that only the former was of taxonomic significance. He was also among the first
experimental physiologists. The Historia Plantarum can be regarded as the first botanical synthesis and text
book for modern botany. According to botanical historian Alan Morton, Ray "influenced both the theory and
the practice of botany more decisively than any other single person in the latter half of the seventeenth
century".[56] Ray's family system was later extended by Pierre Magnol (1638–1715) and Joseph de
Tournefort (1656–1708), a student of Magnol, achieved notoriety for his botanical expeditions, his emphasis
on floral characters in classification, and for reviving the idea of the genus as the basic unit of classification.
[57]
Above all it was Swedish Carl Linnaeus (1707–1778) who eased the task of plant cataloguing. He
adopted a sexual system of classification using stamens and pistils as important characters. Among his most
important publications were Systema Naturae (1735), Genera Plantarum (1737), and Philosophia Botanica
(1751) but it was in his Species Plantarum (1753) that he gave every species a binomial thus setting the path
for the future accepted method of designating the names of all organisms. Linnaean thought and books
dominated the world of taxonomy for nearly a century.[58] His sexual system was later elaborated by Bernard
de Jussieu (1699–1777) whose nephew Antoine-Laurent de Jussieu (1748–1836) extended it yet again to
include about 100 orders (present-day families).[59] Frenchman Michel Adanson (1727–1806) in his Familles
des Plantes (1763, 1764), apart from extending the current system of family names, emphasized that a
natural classification must be based on a consideration of all characters, even though these may later be
given different emphasis according to their diagnostic value for the particular plant group. Adanson's method
has, in essence, been followed to this day.[60]
18th century plant taxonomy bequeathed to the 19th century a precise binomial nomenclature and
botanical terminology, a system of classification based on natural affinities, and a clear idea of the ranks of
family, genus and species — although the taxa to be placed within these ranks remains, as always, the
subject of taxonomic research.
[edit] Anatomy
Further information: Microscopy and Plant anatomy
Robert Hooke's microscope which he described in the 1665 Micrographia: he coined the biological
use of the term cell
In the first half of the 18th century botany was beginning to move beyond descriptive science into
experimental science. Although the microscope was invented in 1590 it was only in the late 17th century that
lens grinding by Antony van Leeuwenhoek provided the resolution needed to make major discoveries.
Important general biological observations were made by Robert Hooke (1635–1703) but the foundations of
plant anatomy were laid by Italian Marcello Malpighi (1628–1694) of the University of Bologna in his Anatome
Plantarum (1675) and Royal Society Englishman Nehemiah Grew (1628–1711) in his The Anatomy of Plants
Begun (1671) and Anatomy of Plants (1682). These botanists explored what is now called developmental
anatomy and morphology, by carefully observing, describing and drawing the developmental transition from
seed to mature plant recording stem and wood formation. This work included the discovery and naming of
parenchyma and stomata.[61]
[edit] Physiology
Main article: Plant physiology
In plant physiology research interest was focused on the movement of sap and the absorption of
substances through the roots. Jan Helmont (1577–1644) by experimental observation and calculation, noted
that the increase in weight of a growing plant cannot be derived purely from the soil, and concluded it must
relate to water uptake.[62] Englishman Stephen Hales[63] (1677–1761) established by quantitative
experiment that there is uptake of water by plants and a loss of water by transpiration and that this is
influenced by environmental conditions: he distinguished “root pressure”, “leaf suction” and “imbibition” and
also noted that the major direction of sap flow in woody tissue is upward. His results were published in
Vegetable Staticks (1727) He also noted that “air makes a very considerable part of the substance of
vegetables”.[64] English chemist Joseph Priestly (1733–1804) is noted for his discovery of oxygen (as now
called) and its production by plants. Later Jan Ingenhousz (1730–1799) observed that only in sunlight do the
green parts of plants absorb air and release oxygen, this being more rapid in bright sunlight while, at night,
the air (CO2) is released from all parts. His results were published in Experiments upon vegetables (1779)
and with this the foundations for 20th century studies of carbon fixation were laid. From his observations he
sketched the cycle of carbon in nature even though the composition of carbon dioxide was yet to be resolved.
[65] Studies in plant nutrition had also progressed. In 1804 Nicolas-Théodore de Saussure's (1767–1845)
Recherches Chimiques sur la Végétation was an exemplary study of scientific exactitude that demonstrated
the similarity of respiration in both plants and animals, that the fixation of carbon dioxide includes water, and
that just minute amounts of salts and nutrients (which he analysed in chemical detail from plant ash) have a
powerful influence on plant growth.[66]
[edit] Plant sexuality

Further information: Plant sexuality and Alternation of generations
Diagram showing the sexual parts of a mature flower
It was Rudolf Camerarius (1665–1721) who was the first to establish plant sexuality conclusively by
experiment. He declared in a letter to a colleague dated 1694 and titled De Sexu Plantarum Epistola that “no
ovules of plants could ever develop into seeds from the female style and ovary without first being prepared
by the pollen from the stamens, the male sexual organs of the plant".[67]
Much was learned about plant sexuality by unravelling the reproductive mechanisms of mosses,
liverworts and algae. In his Vergleichende Untersuchungen of 1851 Wilhelm Hofmeister (1824–1877) starting
with the ferns and bryophytes demonstrated that the process of sexual reproduction in plants entails an
“alternation of generations” between sporophytes and gametophytes.[68] This initiated the new field of
comparative morphology which, largely through the combined work of William Farlow (1844–1919),
Nathanael Pringsheim (1823–1894), Frederick Bower, Eduard Strasburger and others, established that an
"alternation of generations" occurs throughout the plant kingdom.[69]
Some time later the German academic and natural historian Joseph Kölreuter (1733–1806) extended
this work by noting the function of nectar in attracting pollinators and the role of wind and insects in
pollination. He also produced deliberate hybrids, observed the microscopic structure of pollen grains and how
the transfer of matter from the pollen to the ovary inducing the formation of the embryo.[70]
Angiosperm (flowering plant) life cycle showing alternation of generations
One hundred years after Camerarius, in 1793, Christian Sprengel (1750–1816) broadened the
understanding of flowers by describing the role of nectar guides in pollination, the adaptive floral mechanisms
used for pollination, and the prevalence of cross-pollination, even though male and female parts are usually
together on the same flower.[71]
[edit] Nineteenth century foundations of modern botany
In about the mid-19th century scientific communication changed. Until this time ideas were largely
exchanged by reading the works of authoritative individuals who dominated in their field: these were often
wealthy and influential "gentlemen scientists". Now research was reported by the publication of “papers” that
emanated from research “schools” that promoted the questioning of conventional wisdom. This process had
started in the late 18th century when specialist journals began to appear.[72] Even so, botany was greatly
stimulated by the appearance of the first “modern” text book, Matthias Schleiden's (1804–1881) Grundzuge
der Wissenschaftlichen, published in English in 1849 as Principles of Scientific Botany.[73] By 1850 an
invigorated organic chemistry had revealed the structure of many plant constituents.[74] Although the great
era of plant classification had now passed the work of description continued. Augustin de Candolle (1778–
1841) succeeded Antoine-Laurent de Jussieu in managing the botanical project Prodromus Systematis
Naturalis Regni Vegetabilis (1824–1841) which involved 35 authors: it contained all the dicotyledons known in
his day, some 58000 species in 161 families, and he doubled the number of recognized plant families, the
work being completed by his son Alphonse (1806–1893) in the years from 1841 to 1873.[75]
[edit] Plant geography and ecology

Further information: Ecology and Plant community
Alexander von Humboldt 1769–1859 painted by Joseph Stieler in 1843
The opening of the 19th century was marked by an increase in interest in the connection between
climate and plant distribution. Carl Willdenow (1765–1812) examined the connection between seed dispersal
and distribution, the nature of plant associations and the impact of geological history. He noticed the
similarities between the floras of N America and N Asia, the Cape and Australia, and he explored the ideas of
“centre of diversity" and "centre of origin”. German Alexander von Humbolt (1769–1859) and Frenchman
Aime Bonpland (1773–1858) published a massive and highly influential 30 volume work on their travels;
Robert Brown (1773–1852) noted the similarities between the floras of S Africa, Australia and India, while
Joakim Schouw (1789–1852) explored more deeply than anyone else the influence on plant distribution of
temperature, soil factors, especially soil water, and light, work that was continued by Alphonse de Candolle
(1806–1893).[76] Joseph Hooker (1817–1911) pushed the boundaries of floristic studies with his work on
Antarctica, India and the Middle East with special attention to endemism. August Grisebach (1814–1879) in
Die Vegetation der Erde (1872) examined physiognomy in relation to climate and in America geographic
studies were pioneered by Asa Gray (1810–1888).[77]
Physiological plant geography, perhaps more familiarly termed ecology, emerged out of floristic
biogeography in the late 19th century as environmental influences on plants received greater recognition.
Early work in this area was synthesised by Danish professor Eugenius Warming (1841–1924) in his book
Plantesamfund (Ecology of Plants, generally taken to mark the beginning of modern ecology) including new
ideas on plant communities, their adaptations and environmental influences. This was followed by another
grand synthesis, the Pflanzengeographie auf Physiologischer Grundlage of Andreas Schimper (1856–1901)
in 1898 (published in English in 1903 as Plant-geography upon a physiological basis translated by W. R.
Fischer, Oxford: Clarendon press, 839 pp.)[78]
[edit] Anatomy
Further information: Plant anatomy and Cell theory
Plant cells with visible chloroplasts
During the 19th century German scientists led the way towards a unitary theory of the structure and
life-cycle of plants. Following improvements in the microscope at the end of the 18th century, Charles Mirbel
(1776–1854) in 1802 published his Traité d'Anatomie et de Physiologie Végétale and Johann Moldenhawer
(1766–1827) published Beyträge zur Anatomie der Pflanzen (1812) in which he describes techniques for
separating cells from the middle lamella. He identified vascular and parenchymatous tissues, described
vascular bundles, observed the cells in the cambium, and interpreted tree rings. He found that stomata were
composed of pairs of cells, rather than a single cell with a hole.[79]
Anatomical studies on the stele were consolidated by Carl Sanio (1832–1891) who described the
secondary tissues and meristem including cambium and its action. Hugo von Mohl (1805–1872) summarized
work in anatomy leading up to 1850 in Die Vegetabilische Zelle (1851) but this work was later eclipsed by the
encyclopaedic comparative anatomy of Heinrich Anton de Bary in 1877. An overview of knowledge of the
stele in root and stem was completed by Van Tieghem (1839–1914) and of the meristem by Karl Nägeli
(1817–1891). Studies had also begun on the origins of the carpel and flower that continue to the present day.
[80]
[edit] Water relations

Main article: Transpiration
The riddle of water and nutrient transport through the plant remained. Physiologist Von Mohl
explored solute transport and the theory of water uptake by the roots using the concepts of cohesion,
transpirational pull, capillarity and root pressure.[74] German dominance in the field of physiology was
underlined by the publication of the definitive textbook on plant physiology synthesising the work of this
period, Sach's Vorlesungen über Pflanzenphysiologie of 1882. There were, however, some advances
elsewhere such as the early exploration of geotropism (the effect of gravity on growth) by Englishman
Thomas Knight, and the discovery and naming of osmosis by Frenchman Henri Dutrochet (1776–1847).[81]
[edit] Cytology
Main article: Cell theory
The cell nucleus was discovered by Robert Brown in 1831. Demonstration of the cellular composition
of all organisms, with each cell possessing all the characteristics of life, is attributed to the combined efforts
of botanist Matthias Schleiden and zoologist Theodor Schwann (1810–1882) in the early 19th century
although Moldenhawer had already shown that plants were wholly cellular with each cell having its own wall
and Julius von Sachs had shown the continuity protoplasm between cell walls.[82]
From 1870 to 1880 it became clear that cell nuclei are never formed anew but always derived from
the substance of another nucleus. In 1882 Flemming observed the longitudinal splitting of chromosomes in
the dividing nucleus and concluded that each daughter nucleus received half of each of the chromosomes of
the mother nucleus: then by the early 20th century it was found that the number of chromosomes in a given
species is constant. With genetic continuity confirmed and the finding by Eduard Strasburger that the nuclei
of reproductive cells (in pollen and embryo) have a reducing division (halving of chromosomes, now known
as meiosis) the field of heredity was opened up. By 1926 Thomas Morgan was able to outline a theory of the
gene and its structure and function. The form and function of plastids received similar attention, the
association with starch being noted at an early date.[83] With observation of the cellular structure of all
organisms and the process of cell division and continuity of genetic material, the analysis of the structure of
protoplasm and the cell wall as well as that of plastids and vacuoles – what is now known as cytology, or cell
theory became firmly established.
Later, the cytological basis of the gene-chromosome theory of heredity extended from about 1900–
1944 and was initiated by the rediscovery of Gregor Mendel's (1822–1884) laws of plant heredity first
published in 1866 in Experiments on Plant Hybridization and based on cultivated pea, Pisum sativum: this
heralded the opening up of plant genetics. The cytological basis for gene-chromosome theory was explored
through the role of polyploidy and hybridization in speciation and it was becoming better understood that
interbreeding populations were the unit of adaptive change in biology.[84]
[edit] Developmental morphology and evolution

Main article: Evolution
Until the 1860s it was believed that species had remained unchanged through time: each biological
form was the result of an independent act of creation and therefore absolutely distinct and immutable. But the
hard reality of geological formations and strange fossils needed scientific explanation. Charles Darwin’s
Origin of Species (1859) replaced the assumption of constancy with the theory of descent with modification.
Phylogeny became a new principle as "natural" classifications became classifications reflecting, not just
similarities, but evolutionary relationships. Wilhelm Hofmeister established that there was a similar pattern of
organization in all plants expressed through the alternation of generations and extensive homology of
structures.[85]
Polymath German intellect Johann Goethe (1749–1832) had interests and influence that extended
into botany. In Die Metamorphose der Pflanzen (1790) he provided a theory of plant morphology (he coined
the word "morphology") and he included within his concept of “metamorphosis” modification during evolution,
thus linking comparative morphology with phylogeny. Though the botanical basis of his work has been
challenged there is no doubt that he prompted discussion and research on the origin and function of floral
parts.[86] His theory probably stimulated the opposing views of German botanists Alexander Braun (1805–
1877) and Matthias Schleiden who applied the experimental method to the principles of growth and form that
were later extended by Augustin de Candolle (1778–1841).[87]
[edit] Carbon fixation (photosynthesis)

Further information: Soil plant atmosphere continuum and Photosynthesis
Photosynthesis splits water to liberate O2 and fixes CO2 into sugar
At the start of the 19th century the idea that plants could synthesise almost all their tissues from
atmospheric gases had not yet emerged. The energy component of photosynthesis, the capture and storage
of the Sun’s radiant energy in carbon bonds (a process on which all life depends) was first elucidated in 1847
by Mayer, but the details of how this was done would take many more years.[88] Chlorophyll was named in
1818 and its chemistry gradually determined, to be finally resolved in the early 20th century. The mechanism
of photosynthesis remained a mystery until the mid-19th century when Sachs, in 1862, noted that starch was
formed in green cells only in the presence of light and in 1882 he confirmed carbohydrates as the starting
point for all other organic compounds in plants.[89] The connection between the pigment chlorophyll and
starch production was finally made in 1864 but tracing the precise biochemical pathway of starch formation
did not begin until about 1915.
[edit] Nitrogen fixation

Main article: Nitrogen fixation
Significant discoveries relating to nitrogen assimilation and metabolism, including ammonification,
nitrification and nitrogen fixation (the uptake of atmospheric nitrogen by symbiotic soil microorganisms) had
to wait for advances in chemistry and bacteriology in the late 19th century and this was followed in the early
20th century by the elucidation of protein and amino-acid synthesis and their role in plant metabolism. With
this knowledge it was then possible to outline the global nitrogen cycle.[90]
[edit] Twentieth century
Thin layer chromatography is used to separate components of chlorophyll

20th century science grew out of the solid foundations laid by the breadth of vision and detailed
experimental observations of the 19th century. A vastly increased research force was now rapidly extending
the horizons of botanical knowledge at all levels of plant organization from molecules to global plant ecology.
There was now an awareness of the unity of biological structure and function at the cellular and biochemical
levels of organisation. Botanical advance was closely associated with advances in physics and chemistry
with the greatest advances in the 20th century mainly relating to the penetration of molecular organization.
[91] However, at the level of plant communities it would take until mid century to consolidate work on ecology
and population genetics.[92] By 1910 experiments using labelled isotopes were being used to elucidate plant
biochemical pathways, to open the line of research leading to gene technology. On a more practical level
research funding was now becoming available from agriculture and industry.
[edit] Molecules
Main article: Molecular biology
In 1903 Chlorophylls a and b were separated by thin layer chromatography then, through the 1920s
and 1930s, biochemists, notably Hans Krebs (1900–1981) and Carl (1896–1984) and Gerty Cori (1896–1957)
began tracing out the central metabolic pathways of life. Between the 1930s and 1950s it was determined
that ATP, located in mitochondria, was the source of cellular chemical energy and the constituent reactions of
photosynthesis were progressively revealed. Then, in 1944 DNA was extracted for the first time.[93] Along
with these revelations there was the discovery of plant hormones or “growth substances”, notably auxins,
(1934) gibberellins (1934) and cytokinins (1964)[94] and the effects of photoperiodism, the control of plant
processes, especially flowering, by the relative lengths of day and night.[95]
Following the establishment of Mendel’s laws, the gene-chromosome theory of heredity was
confirmed by the work of August Weismann who identified chromosomes as the hereditary material. Also, in
observing the halving of the chromosome number in germ cells he anticipated work to follow on the details of
meiosis, the complex process of redistribution of hereditary material that occurs in the germ cells. In the
1920s and 1930s population genetics combined the theory of evolution with Mendelian genetics to produce
the modern synthesis. By the mid-1960s the molecular basis of metabolism and reproduction was firmly
established through the new discipline of molecular biology. Genetic engineering, the insertion of genes into
a host cell for cloning, began in the 1970s with the invention of recombinant DNA techniques and its
commercial applications applied to agricultural crops followed in the 1990s. There was now the potential to
identify organisms by molecular “fingerprinting” and to estimate the times in the past when critical
evolutionary changes had occurred through the use of “molecular clocks”.
[edit] Computers, electron microscopes and evolution

Further information: Ultrastructure and Palynology
Electron microscope constructed by Ernst Ruska in 1933
Increased experimental precision combined with vastly improved scientific instrumentation was
opening up exciting new fields. In 1936 Alexander Oparin (1894–1980) demonstrated a possible mechanism
for the synthesis of organic matter from inorganic molecules. In the 1960s it was determined that the Earth’s
earliest life-forms treated as plants, the cyanobacteria known as stromatolites, dated back some 3.5 billion
years.[96]
Mid-century transmission and scanning electron microscopy presented another level of resolution to
the structure of matter, taking anatomy into the new world of “ultrastructure”.[97]
New and revised “phylogenetic” classification systems of the plant kingdom were produced, perhaps
the most notable being that of August Eichler (1839–1887), and the massive 23 volume Die natürlichen
Pflanzenfamilien of Adolf Engler (1844–1930) & Karl Prantl (1849–1893) published over the period 1887 and
1915. Taxonomy based on gross morphology was now being supplemented by using characters revealed by
pollen morphology, embryology, anatomy, cytology, serology, macromolecules and more.[98] The
introduction of computers facilitated the rapid analysis of large data sets used for numerical taxonomy (also
called taximetrics or phenetics). The emphasis on truly natural phylogenies spawned the disciplines of
cladistics and phylogenetic systematics. The grand taxonomic synthesis An Integrated System of
Classification of Flowering Plants (1981) of American Arthur Cronquist (1919–1992) was superseded when, in
1998, the Angiosperm Phylogeny Group published a phylogeny of flowering plants based on the analysis of
DNA sequences using the techniques of the new molecular systematics which was resolving questions
concerning the earliest evolutionary branches of the angiosperms (flowering plants). The exact relationship of
fungi to plants had for some time been uncertain. Several lines of evidence pointed to fungi being different
from plants, animals and bacteria – indeed, more closely related to animals than plants. In the 1980s-90s
molecular analysis revealed an evolutionary divergence of fungi from other organisms about 1 billion years
ago – sufficient reason to erect a unique kingdom separate from plants.[99]
[edit] Biogeography and ecology

Main article: Biogeography
Map of terrestrial biomes classified by vegetation type
The publication of Alfred Wegener's (1880–1930) theory of continental drift 1912 gave additional
impetus to comparative physiology and the study of biogeography while ecology in the 1930s contributed the
important ideas of plant community, succession, community change, and energy flows.[100] From 1940 to
1950 ecology matured to become an independent discipline as Eugene Odum (1913–2002) formulated many
of the concepts of ecosystem ecology, emphasising relationships between groups of organisms (especially
material and energy relationships) as key factors in the field. Building on the extensive earlier work of
Alphonse de Candolle, Nikolai Vavilov (1887–1943) from 1914 to 1940 produced accounts of the geography,
centres of origin, and evolutionary history of economic plants.[101]
[edit] Twenty-first century
In reviewing the sweep of botanical history it is evident that, through the power of the scientific
method, most of the basic questions concerning the structure and function of plants have, in principle, been
resolved. Now the distinction between pure and applied botany becomes blurred as our historically
accumulated botanical wisdom at all levels of plant organisation is needed (but especially at the molecular
and global levels) to improve human custodianship of planet earth. The most urgent unanswered botanical
questions now relate to the role of plants as primary producers in the global cycling of life's basic ingredients:
energy, carbon, hydrogen, oxygen, and nitrogen, and ways that our plant stewardship can help address the
global environmental issues of resource management, conservation, human food security, biologically
invasive organisms, carbon sequestration, climate change, and sustainability.[102]
[edit] See also

• History of plant systematics
• History of phycology
• List of botanists
• List of botanists by author abbreviation
[edit] References
1. ^ Morton 1981, p. 49
2. ^ Sachs 1890, p. v
3. ^ Walters 1981, p. 3
4. ^ Morton 1981, p. 2
5. ^ See Stearn 1986
6. ^ Stearn 1965, pp. 279–91, 322–41
7. ^ Reed 1942, p. 3
8. ^ Morton 1981, p. 5
9. ^ Reed 1942, pp. 7–29
10.^ Morton 1981, p. 15
11.^ Morton 1981, p. 12
12.^ See Needham et al. 1986.
13.^ Morton 1981, p. 23
14.^ Morton 1981, p. 25
15.^ Vines in Oliver 1913, p. 8
16.^ Morton 1981, pp. 29–43
17.^ Singer 1923, p. 98
18.^ Reed 1942, p. 34
19.^ Writings on plants
20.^ Morton 1981, p. 42
21.^ Reed 1942, p. 37
22.^ See Thanos 2005
23.^ Morton 1981, pp. 36–43
24.^ Harvey-Gibson 1919, p. 9
25.^ Singer 1923, p. 101
26.^ Morton 1981, p. 68
27.^ Morton 1981, p. 69
28.^ Morton 1981, pp. 70–1
29.^ See Sengbusch
30.^ Morton 1981, pp. 58–64
31.^ Fahd 1996, p. 815
32.^ a b Morton 1981, p. 82
33.^ Huff 2003, p. 218
34.^ See Boulanger 2002
35.^ Morton 1981, p. 94
36.^ Tiwari, Lalit (24 June 2003). "Ancient Indian Botany and Taxonomy". The Infinity
Foundation. http://www.infinityfoundation.com/mandala/t_es/t_es_tiwar_botany_frameset.htm.
Retrieved 15 December 2009.
37.^ Majumdar 1982, pp. 356–411
38.^ Sachs 1890, p. 19
39.^ a b Reed 1942, p. 65
40.^ Reed 1942, p. 68
41.^ Arber 1986, pp. 119–124
42.^ Arber in Oliver 1913, pp. 146–246
43.^ Henrey 1975, pp. 631–46
44.^ Morton 1981, p. 145
45.^ Sachs 1890, p. 18
46.^ Morton 1981, pp. 120–4
47.^ Arber 1986, p. 270
48.^ Arber 1913, pp. 44–64
49.^ Morton 1981, pp. 178–80
50.^ Reed 1942, pp. 110–1
51.^ Woodland 1991, pp. 372–408
52.^ Reed 1942, pp. 71–3
53.^ Morton 1981, pp. 130–40
54.^ Morton 1981, pp. 147–8
55.^ Reed 1942, pp. 82–3
56.^ Morton 1981, pp. 196–216
57.^ Woodland 1991, pp. 372–375
58.^ Stafleu 1971, p. 79
59.^ Reed 1942, p. 102
60.^ Morton 1981, pp. 301–11
61.^ Reed 1942, pp. 88–9
62.^ Reed 1942, p. 91
63.^ Darwin in Oliver 1913, pp. 65–83
64.^ Morton 1981, p. 250
65.^ Reed 1942, p. 107
66.^ Morton 1981, p. 338
67.^ Reed 1942, p. 96
68.^ Reed 1942, p. 138
69.^ Reed 1942, p. 140
70.^ Reed 1942, p. 97
71.^ Reed 1942, p. 98
72.^ Reynolds Green 1909, p. 502
73.^ Morton 1981, p. 377
74.^ a b Morton 1981, p. 388
75.^ Morton 1981, p. 372
76.^ Morton 1981, p. 364
77.^ Morton 1981, p. 413
78.^ Reed 1942, pp. 126–33
79.^ Morton 1981, pp. 368–370
80.^ Morton 1981, pp. 386–395
81.^ Morton 1981, pp. 390–1
82.^ Morton 1981, pp. 381–2
83.^ Reed 1942, pp. 154–75
84.^ Morton 1981, p. 453
85.^ Reynolds Green 1909, pp. 7–10, 501
86.^ Morton 1981, pp. 343–6
87.^ Morton 1981, pp. 371–3
88.^ Reed 1942, p. 207
89.^ Reed 1942, p. 197
90.^ Reed 1942, pp. 214–40
91.^ Morton 1981, p. 448
92.^ Morton 1981, p. 451
93.^ Morton 1981, p. 460
94.^ Morton 1981, p. 461
95.^ Morton 1981, p. 464
96.^ Morton 1981, p. 454
97.^ Morton 1981, p. 459
98.^ Morton 1981, p. 456
99.^ See Bruns 2006
100.^ Morton 1981, p. 457
101.^ See de Candolle 2006
102.^ Botanical Society of America
[edit] Bibliography
• Arber, Agnes (1986). Herbals, Their Origin and Evolution, a Chapter in the History of Botany
1470–1670. Cambridge: Cambridge University Press. ISBN 0521338794. (First published in 1912.)
• Botanical Society of America. "Botany for the Next Millennium: I. The intellectual: evolution,
development, ecosystems". http://www.botany.org/bsa/millen/mil-chp1.html#Evolution. Retrieved
2009-12-17 . (Report from the Botanical Society of America.)
• Boulanger, Diane (2002). "The Islamic Contribution to Science, Mathematics and
Technology". OISE Papers in STSE Education 3.
• Bruns, Tom (2006). "Evolutionary biology: a kingdom revised". Nature 443 (7113): 758–61.
doi:10.1038/443758a. PMID 17051197.
• de Candolle, Alphonse (2006). Origin of Cultivated Plants. Glacier National Park Montana:
Kessinger Publishing. ISBN 9781428609464. http://books.google.com/?
id=a242a8XfLycC&dq=de+candolle+origin+of+cultivated+plants&printsec=frontcover&q=. Retrieved
2009-12-14. (First edition in French published in 1882.)
• Denham, Tim et al.; Haberle, SG; Lentfer, C; Fullagar, R; Field, J; Therin, M; Porch, N;
Winsborough, B (2003). "Origins of Agriculture at Kuk Swamp in the Highlands of New Guinea".
• Fahd, Toufic (1996). "Botany and agriculture". In Morelon, Régis; Rashed, Roshdi.
Encyclopedia of the History of Arabic Science. 3. London: Routledge. ISBN 0415124107.
• Greene, Edward Lee & Egerton, Frank N. (ed.) (1983a). Landmarks of Botanical History:
Part 1. Stanford: Stanford University Press. ISBN 9780804710756. ; originally published as Greene,
Edward L. (1909). Landmarks of Botanical History 1. Prior to 1562 A.D. Washington: Smithsonian
Institution. OCLC 174698401.
• Greene, Edward Lee & Egerton, Frank N. (ed.) (1983b). Landmarks of Botanical History:
Part 2. Stanford: Stanford University Press. ISBN 9780804710756.
• Harvey-Gibson, Robert J. (1919). Outlines of the History of Botany. London: A. & C. Black.
ISBN 0405138776.
• Helmsley, Alan R.; Poole, Imogen (eds) (2004). The Evolution of Plant Physiology: From
Whole Plants to Ecosystems. London: Elsevier Academic Press. ISBN 0123395526.
• Henrey, Blanche (1975). British botanical and horticultural literature before 1800 (Vols 1–3) .
Oxford: Oxford University Press. ISBN 0192115480.
• Huff, Toby (2003). The Rise of Early Modern Science: Islam, China, and the West .
• Majumdar, G. P. (1982). "Studies in History of Science in India". In Chattopadhyaya,
Debiprasad. The history of botany and allied sciences in India (c. 2000 B.C. to 100 A.D.) . Asha Jyoti,
New Delhi: Editorial Enterprise.
• Morton, Alan G. (1981). History of Botanical Science: An Account of the Development of
Botany from Ancient Times to the Present Day. London: Academic Press. ISBN 0125083823.
• Meyer, Ernst H.F. (1854–57). Geschichte der Botanik. Köningsberg: Verlag de Gebrűder
Bornträger. http://books.google.com/?
id=ukECAAAAYAAJ&pg=PP3&dq=intitle:Geschichte+intitle:der+intitle:Botanik+inauthor:Meyer.
• Needham, Joseph; Lu, Gwei-djen & Huang, Hsing-Tsung (1986). Science and Civilisation in
China, Vol. 6 Part 1 Botany. Cambridge: Cambridge University Press.
• Needham, Joseph & Lu, Gwei-Djen (2000). Sivin, Nathan. ed. Science and Civilisation in
China, Vol. 6 Part 6 Medicine. Cambridge: Cambridge University Press.
• Oliver, Francis W.(ed) (1913). Makers of British Botany. Cambridge: Cambridge University
Press. http://en.wikisource.org/wiki/Makers_of_British_botany. Retrieved 2010-01-12.
• Reynolds Green, Joseph (1909). History of Botany 1860–1900. Oxford: Clarendon Press.
• Reed, Howard S. (1942). A Short History of the Plant Sciences. New York: Ronald Press.
• Sachs, Julius von (1890). History of Botany (1530–1860). trans. Garnsey, H.E.F. & Balfour,
I.B.. Oxford: Clarendon Press.
• Sengbusch, Peter. "Botany: The History of a Science". Botany online.
http://www.biologie.uni-hamburg.de/b-online/e01/01.htm. Retrieved 2009-12-11.
• Singer, Charles (1923). "Herbals". The Edinburgh Review 237: 95–112.
• Stafleu, Frans A. (1971). Linnaeus and the Linnaeans. Utrecht: International Association of
Plant Taxonomy. ISBN 9060460642.
• Stearn, William T. (1965). "The Origin and Later Development of Cultivated Plants". Journal
of the Royal Horticultural Society 90: 279–291, 322–341.
• Stearn, William T. (1986). "Historical Survey of the Naming of Cultivated Plants". Acta
Horticulturae 182: 18–28.
• Thanos, Costas A. (2005). "The Geography of Theophrastus' Life and of his Botanical
Writings (Περι Φυτων)". In Karamanos, A.J. & Thanos, C.A. (eds). Biodiversity and Natural Heritage
in the Aegean, Proceedings of the Conference 'Theophrastus 2000' (Eressos - Sigri, Lesbos, July 6–
8, 2000). Athens: Fragoudis. pp. 23–45. http://www.biology.uoa.gr/~cthanos/Papers/Theophrastus
%20Geography.pdf. Retrieved 2009-11-11.
• Vavilov, Nicolai I. (1951). trans. K. Starr Chester. "The Origin, Variation, Immunity and
Breeding of Cultivated Plants". Chronica Botanica 13: 1–366.
• Vavilov, Nicolai I. (1992). Origin and Geography of Cultivated Plants . Cambridge: Cambridge
University Press. ISBN 0521404274.
• Walters, Stuart M. (1981). The Shaping of Cambridge Botany. Cambridge: Cambridge
University Press. ISBN 0521237955.
• Winterborne, Jeffrey (2005). Hydroponics: indoor horticulture. Guildford: Pukka Press.
ISBN 9780955011207. http://www.hydroponicist.com/. Retrieved 2009-12-14.
• Woodland, Dennis W. (1991). Contemporary Plant Systematics. New Jersey: Prentice Hall.
ISBN 0205121829.

• Johnson, Dale E. (1985). "Literature on the history of botany and botanic gardens 1730–1840:
A bibliography". Huntia 6 (1): 1–121.
[show]v · d · eBotany

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[hide]v · d · eHistory of botany

Agriculture · Bryology · Cell theory · Cladistics ·
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disciplines Natural history · Numerical taxonomy · Paleobotany · Palynology ·
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Institutions Orto botanico di Padova · Orto botanico di Pisa · Rothamsted
Experimental Station · Royal Botanic Gardens, Kew
Publications Historia Plantarum of Theophrastus and Causes of Plants

c. 200 BC · Materia Medica c. 60 AD · Naturalis Historia 77-79
AD · Herbarum Vivae Icones 1530 · Libellus De Re Herbaria
Novus 1538 · Kreutterbuch 1539 · Historia Plantarum of John Ray
1561-1563 · De Plantis 1583 · Stirpium Historiae 1583 ·
Prodromus Theatrici Botanici 1620 · Pinax 1623 · Anatome
Plantarum 1675 · Anatomy of Plants 1682 · Historia Plantarum
1682–1704 · De Sexu Plantarum Epistola 1694 · Vegetable
Staticks 1727 · Systema Naturae 1735 · Genera Plantarum 1737 ·
Philosophia Botanica 1751 · Species Plantarum 1753 · Familles
des Plantes 1763-4 · Experiments Upon Vegetables 1779 · Die
Metamorphose der Pflantzen 1790 · Traité d'Anatomie et de
Physiologie Végétale 1802 · Recherches Chimiques sur la
Végétation 1804 · Beyträge zur Anatomie der Pflanzen 1812 ·
Prodromus Systematis Naturalis Regni Vegetabilis 1824-1841 ·
Die Vegetabilische Zelle 1851 · Vergleichende Untersuchungen
1851 · On the Origin of Species 1859 · Experiments on Plant
Hybridization 1862 · Die Vegetation der Erde 1872 · Prodromus
Systematis Naturalis Regni Vegetabilis 1873 · Plantesamfund
1895 · Pflanzengeographie auf Physiologischer Grundlage 1898 ·
Variation and Evolution in Plants 1950 · Ontogeny and Phylogeny
1977 · An Integrated System of Classification of Flowering Plants
1981
Theories Alternation of generations · Biogeographical realm ·

and concepts Biological classification · Cell theory · Center of diversity ·
Spontaneous generation
Influential Theophrastus c. 371 – c. 287 BC · Pliny the Elder 23–79

figures AD · Pedanius Dioscorides c. 40-90 AD · Otto Brunfels 1464-
1534 · Hieronymus Bock 1498-1554 · Valerius Cordus 1515-1544 ·
William Turner 1515-1568 · Rembert Dodoens 1517-1585 ·
Andrea Cesalpino 1519-1603 · Gaspard Bauhin 1560–1624 ·
Joachim Jungius 1587–1657 · John Ray 1623–1705 · Nehemiah
Grew 1628–1711 · Marcello Malpighi 1628–1694 · Joseph Pitton de
Tournefort 1656–1708 · Rudolf Jakob Camerarius 1665–1721 ·
Stephen Hales 1677–1761 · Bernard de Jussieu 1699–1777 ·
Carolus Linnaeus 1707–1778 · Michel Adanson 1727–1806 · Jan
Ingenhousz 1730–1799 · Joseph Banks 1743–1820 · Johann
Wolfgang von Goethe 1749–1832 · Carl Ludwig Willdenow 1765–
1812 · Nicolas-Théodore de Saussure 1767–1845 · Alexander von
Humboldt 1769–1859 · Aimé Bonpland 1773–1858 · Joakim
Frederik Schouw 1789–1852 · Matthias Jakob Schleiden 1804–
1881 · Alexander Braun 1805–1877 · Asa Gray 1810–1888 · August
Grisebach 1814–1879 · Joseph Hooker 1817–1911 · Gregor
Mendel 1822–1884 · Nathanael Pringsheim 1823–1894 · Wilhelm
Hofmeister 1824–1877 · Julius von Sachs 1832–1897 · Eugenius
Warming 1841–1924 · William Gilson Farlow 1844–1919 · Andreas
Franz Wilhelm Schimper 1856–1901 · Nikolai Vavilov 1887–1943 ·
G. Ledyard Stebbins 1906–2000 · Eugene Odum 1913–2002 ·
Arthur Cronquist 1919–1992
Related History of agricultural science · History of agriculture ·

topics History of biochemistry · History of biotechnology · History of
ecology · History of evolutionary thought · History of genetics ·
History of geology · History of medicine · History of molecular
biology · History of molecular evolution · History of paleontology ·
History of phycology · History of plant systematics · History of
science · Natural philosophy · Philosophy of biology · Timeline of
biology and organic chemistry · Herbal · Botanical garden


Institutions Zoologica • Marine Biological Laboratory • Rockefeller University •
Biology
Experiments Griffith's experiment • Miller-Urey experiment • Luria–

Hershey–Chase experiment • Meselson–Stahl experiment • Crick,



Oparin • Alexander Fleming • J. B. S. Haldane • Sewall Wright • R.


Retrieved from "http://en.wikipedia.org/wiki/History_of_botany"
Categories: Botany | History of biology | History of science

W000
Evolution of birds

Archaeopteryx at Paläontologisches Museum München
see also: Evolution of reptiles
The evolution of birds is thought to have begun in the Jurassic Period, with the earliest birds derived
from theropod dinosaurs. Birds are categorized as a biological class, Aves. The earliest known species of
class Aves is Archaeopteryx lithographica, from the Late Jurassic period, though Archaeopteryx is not
commonly considered to have been a true bird. Modern phylogenies place birds in the dinosaur clade
Theropoda. According to the current consensus, Aves and a sister group, the order Crocodilia, together are
the sole living members of an unranked "reptile" clade, the Archosauria.
Phylogenetically, Aves is usually defined as all descendants of the most recent common ancestor of
a specific modern bird species (such as the House Sparrow, Passer domesticus), and either Archaeopteryx,
[1] or some prehistoric species closer to Neornithes (to avoid the problems caused by the unclear
relationships of Archaeopteryx to other theropods).[2] If the latter classification is used then the larger group
is termed Avialae. Currently, the relationship between dinosaurs, Archaeopteryx, and modern birds is still
under debate.
Contents
[hide]
• 1 Origins
• 2 Adaptive radiation of birds
• 3 Classification of modern species
• 4 Current evolutionary trends in birds
• 5 References
[edit] Origins
Main article: Origin of birds
See also: Avicephala
There is significant evidence that birds evolved from theropod dinosaurs, specifically, that birds are
members of Maniraptora, a group of theropods which includes dromaeosaurs and oviraptorids, among
others.[3] As more non-avian theropods that are closely related to birds are discovered, the formerly clear
distinction between non-birds and birds becomes less so. Recent discoveries in northeast China (Liaoning
Province), demonstrating that many small theropod dinosaurs had feathers, contribute to this ambiguity.[4]
The basal bird Archaeopteryx, from the Jurassic, is well-known as one of the first "missing links" to
be found in support of evolution in the late 19th century, though it is not considered a direct ancestor of
modern birds. Confuciusornis is another early bird; it lived in the Early Cretaceous. Both may be predated by
Protoavis texensis, though the fragmentary nature of this fossil leaves it open to considerable doubt whether
this was a bird ancestor. Other Mesozoic birds include the Confuciusornis, the Enantiornithes, Yanornis,
Ichthyornis, Gansus, and the Hesperornithiformes - a group of flightless divers resembling grebes and loons.
The recently (2002) discovered dromaeosaur Cryptovolans (which may be a Microraptor) was capable of
powered flight, possessed a sternal keel and had ribs with uncinate processes. In fact, Cryptovolans makes a
better "bird" than Archaeopteryx which lacks some of these modern bird features. Because of this, some
paleontologists have suggested that dromaeosaurs are actually basal birds whose larger members are
secondarily flightless, i.e. that dromaeosaurs evolved from birds and not the other way around. Evidence for
this theory is currently inconclusive, but digs continue to unearth fossils (especially in China) of the strange
feathered dromaeosaurs. At any rate, it is fairly certain that flight utilizing feathered wings existed in the mid-
Jurassic theropods and was "tried out" in several lineages and variants by the mid-Cretaceous, such as in
Confuciusornis. This latter species had some peculiar features. For example, its vestigial tail was unfit for
steering, and its wing shape seems rather specialized although the arm skeleton was still quite
"dinosaurian").
Although ornithischian (bird-hipped) dinosaurs share the same hip structure as birds, birds actually
originated from the saurischian (lizard-hipped) dinosaurs if the dinosaurian origin theory is correct. They thus
arrived at their hip structure condition independently. In fact, a bird-like hip structure also developed a third
time among a peculiar group of theropods, the Therizinosauridae.
An alternate theory to the dinosaurian origin of birds, espoused by a few scientists, notably Larry
Martin and Alan Feduccia, states that birds (including maniraptoran "dinosaurs") evolved from early
archosaurs like Longisquama.[5] This theory is contested by most other paleontologists and experts in
feather development and evolution.[6]
[edit] Adaptive radiation of birds

Modern birds are classified in Neornithes, which are now known to have evolved into some basic
lineages by the end of the Cretaceous (see Vegavis). The Neornithes are split into the paleognaths and
neognaths.
Paleognathae
The paleognaths include the tinamous (found only in Central and South America) and the ratites which
nowadays are found almost exclusively on the Southern Hemisphere. The ratites are large flightless birds,
and include ostriches, rheas, cassowaries, kiwis and emus. A few scientists propose that the ratites
represent an artificial grouping of birds which have independently lost the ability to fly in a number of
unrelated lineages[citation needed]; in any case, the available data regarding their evolution is still very
confusing.
Neognathae
The basal divergence from the remaining Neognathes was that of the Galloanserae, the superorder
containing the Anseriformes (ducks, geese and swans), and the Galliformes (chickens, turkeys, pheasants,
and their allies).
The dates for the splits are a matter of considerable debate amongst scientists. It is agreed that the
Neornithes evolved in the Cretaceous and that the split between the Galloanserae and the other neognaths -
the Neoaves - occurred before the K-T extinction event, but there are different opinions about whether the
radiation of the remaining neognaths occurred before or after the extinction of the other dinosaurs.[7] This
disagreement is in part caused by a divergence in the evidence, with molecular dating suggesting a
Cretaceous radiation, a small and equivocal neoavian fossil record from Cretaceous, and most living families
turning up during the Paleogene. Attempts made to reconcile the molecular and fossil evidence have proved
controversial.[7][8]
On the other hand, two factors must be considered: First, molecular clocks cannot be considered
reliable in the absence of robust fossil calibration, whereas the fossil record is naturally incomplete. Second,
in reconstructed phylogenetic trees, the time and pattern of lineage separation corresponds to the evolution
of the characters (such as DNA sequences, morphological traits etc.) studied, not to the actual evolutionary
pattern of the lineages; these ideally should not differ by much, but may well do so in practice.
Considering this, it is easy to see that fossil data, compared to molecular data, tends to be more
accurate in general, but also to underestimate divergence times: morphological traits, being the product of
entire developmental genetics networks, usually only start to diverge some time after a lineage split would
become apparent in DNA sequence comparison - especially if the sequences used contain many silent
mutations.
[edit] Classification of modern species

See also: Sibley-Ahlquist taxonomy and dinosaur classification
The phylogenetic classification of birds is a contentious issue. Sibley & Ahlquist's Phylogeny and
Classification of Birds (1990) is a landmark work on the classification of birds (although frequently debated
and constantly revised). A preponderance of evidence suggests that most modern bird orders constitute
good clades. However, scientists are not in agreement as to the precise relationships between the orders;
evidence from modern bird anatomy, fossils and DNA have all been brought to bear on the problem but no
strong consensus has emerged. As of the mid-2000s, new fossil and molecular data provide an increasingly
clear picture of the evolution of modern bird orders, and their relationships. For example, the Charadriiformes
seem to consititute an ancient and distinct lineage, while the Mirandornithes and Cypselomorphae are
supported by a wealth of anatomical and molecular evidence. Our understanding of the interrelationships of
lower level taxa also continues to increase, particularly in the massively diverse perching bird order
Passeriformes.
On June 27, 2008, the largest study of bird genetics was published. It overturns several hypothesized
relationships, and will likely necessitate a wholesale restructuring of the avian phylogenetic tree.[9]
[edit] Current evolutionary trends in birds

See also: Bird conservation
Evolution generally occurs at a scale far too slow to be witnessed by humans. However, bird species
are currently going extinct at a far greater rate than any possible speciation or other generation of new
species. The disappearance of a population, subspecies, or species represents the permanent loss of a
range of genes.
Another concern with evolutionary implications is a suspected increase in hybridization. This may
arise from human alteration of habitats enabling related allopatric species to overlap. Forest fragmentation
can create extensive open areas, connecting previously isolated patches of open habitat. Populations that
were isolated for sufficient time to diverge significantly, but not sufficient to be incapable of producing fertile
offspring may now be interbreeding so broadly that the integrity of the original species may be compromised.
For example, the many hybrid hummingbirds found in northwest South America may represent a threat to the
conservation of the distinct species involved.[10]
Several species of birds have been bred in captivity to create variations on wild species. In some
birds this is limited to color variations, while others are bred for larger egg or meat production, for
flightlessness or other characteristics.
Some species, like the rock pigeon or several species of crows have been successful living in man
made environments. Because these new habitats are different from their far less numerous "natural" habitats,
these species are to a certain extent displaying evolutionary adaptations to living close to man, including
color changes, increased memory, and enhanced intelligence.[ citation needed]
[edit] References
1. ^ Padian K & Chiappe LM (1997). "Bird Origins". In Currie PJ & Padian K. Encyclopedia of
Dinosaurs. San Diego: Academic Press. pp. 41–96.
2. ^ Gauthier, J (1986). "Saurischian Monophyly and the origin of birds". In Padian K. The
Origin of Birds and the Evolution of Flight. Mem. California Acad. Sci 8 . pp. 1–55.
3. ^ Hou L,Martin M, Zhou Z & Feduccia A, (1996) "Early Adaptive Radiation of Birds: Evidence
from Fossils from Northeastern China" Science 274(5290): 1164-1167 Abstract
4. ^ Norell, M & Ellison M (2005) Unearthing the Dragon, The Great Feathered Dinosaur
Discovery Pi Press, New York, ISBN 0-13-186266-9
5. ^ Feduccia A, Lingham-Soliar T, Hinchliffe JR (2005) "Do feathered dinosaurs exist? Testing
the hypothesis on neontological and paleontological evidence" Journal of Morphology 266(2): 125-
166
6. ^ Prum R (2003) "Are Current Critiques Of The Theropod Origin Of Birds Science? Rebuttal
To Feduccia 2002" Auk 120(2) 550-561
7. ^ a b Ericson PGP, Anderson CL, Britton T, Elzanowski A, Johansson US, Kallersjo M,
Ohlson JI, Parsons TJ, Zuccon D, Mayr G (2006)"Diversification of Neoaves: integration of molecular
sequence data and fossils" Biology Letters 2(4): 543-547
8. ^ Brown JW, Payne RB, & Mindell DP (2007) "Nuclear DNA does not reconcile ‘rocks’ and
‘clocks’ in Neoaves: a comment on Ericson et al. Biology Letters 3(3): 257-259
9. ^ Science Daily, June 27, 2008
10.^ Fjeldså, Jon; Niels Krabbe. (1990). Birds of the High Andes: A Manual to the Birds of the
Temperate Zone of the Andes and Patagonia, South America. Apollo Books. ISBN 8788757161.
[hide]v · d · eBirds (class: Aves)
Anatomy Bird anatomy • Flight • Eggs • Feathers • Plumage • Beak • Vision • Dactyly •
Preen gland
Behaviour Singing • Intelligence • Migration • Incubation • Brood parasites • Nesting •

Hybrids
Evolution Evolution of birds • Origin of birds • Darwin's finches • Seabirds
Superorder: Palaeognathae
Subclass:
Neornithes Struthioniformes (ratites) • Tinamiformes (tinamous)
(29 orders of
Superorder: Neognathae
Accipitriformes • Anseriformes (waterfowl) • Apodiformes (swifts and

hummingbirds) • Caprimulgiformes (nightjars and relatives) • Cariamae (seriemas and
relatives) • Charadriiformes (gulls and relatives) • Ciconiiformes (storks) • Coliiformes
(mousebirds) • Columbiformes (doves and pigeons) • Coraciiformes (kingfishers and
modern birds) relatives) • Cuculiformes (cuckoos and relatives) • Falconiformes (falcons and
relatives) • Galliformes (gamebirds) • Gaviiformes (loons or divers) • Gruiformes
(cranes and relatives) • Passeriformes (perching birds) • Pelecaniformes (pelicans
and relatives) • Phaethontiformes (tropicbirds) • Phoenicopteriformes (flamingos) •
Piciformes (woodpeckers and relatives) • Podicipediformes (grebes) •
Procellariiformes (albatrosses and petrels) • Psittaciformes (parrots) • Pteroclidiformes
(sandgrouses) • Sphenisciformes (penguins) • Strigiformes (owls) • Trogoniformes
(trogons and quetzals)
Fossil birds Archaeopteryx • Enantiornithes • Hesperornithes
Birds and Ringing • Ornithology • Bird collections • Birdwatching • Bird feeding •

humans Conservation • Aviculture • Waterfowl hunting • Pigeon racing
Lists Families and orders • Genera • Lists by region • Extinct birds • Late Quaternary
prehistoric birds • Sibley-Ahlquist taxonomy
Category • Portal
Processes of
Population genetic
Evolutionary
developmental
concepts

Modes of

Retrieved from "http://en.wikipedia.org/wiki/Evolution_of_birds"
Categories: Bird evolution

W000
Life

For other uses, see Life (disambiguation).
Life
Life
Domains and Kingdoms
Life on Earth:
• Non-cellular life (viruses) [note 1]
• Cellular life
• Bacteria
• Archaea
• Eukarya
• Protista
• Fungi
• Plantae
• Animalia
Life (cf. biota) is a characteristic that distinguishes objects that have signaling and self-sustaining
processes (biology) from those that do not,[1][2] either because such functions have ceased (death), or else
because they lack such functions and are classified as inanimate.[3]
In biology, the science of living organisms, life is the condition which distinguishes active organisms
from inorganic matter.[4] Living organisms undergo metabolism, maintain homeostasis, possess a capacity
to grow, respond to stimuli, reproduce and, through natural selection, adapt to their environment in
successive generations. More complex living organisms can communicate through various means.[1][5] A
diverse array of living organisms (life forms) can be found in the biosphere on Earth, and the properties
common to these organisms—plants, animals, fungi, protists, archaea, and bacteria—are a carbon- and water-
based cellular form with complex organization and heritable genetic information.
In philosophy and religion, the conception of life and its nature varies. Both offer interpretations as to
how life relates to existence and consciousness, and both touch on many related issues, including life
stance, purpose, conception of a god or gods, a soul or an afterlife.
Contents
[hide]
• 1 Early theories about life
• 1.1 Materialism
• 1.2 Hylomorphism
• 1.3 Vitalism
• 2 Definitions
• 2.1 Biology
• 2.1.1 Proposed
• 2.1.2 Viruses
• 2.2 Biophysics
• 2.3 Living systems theories
• 2.3.1 Gaia hypothesis
• 2.3.2 Nonfractionability
• 2.3.3 Life as a property of ecosystems
• 3 Origin
• 4 Conditions for life
• 4.1 Range of tolerance
• 4.2 Extremophiles
• 4.3 Chemical element requirements
• 5 Classification of life
• 6 Extraterrestrial life
[edit] Early theories about life
[edit] Materialism
Plant life
Herds of zebra and impala gathering on the Masai Mara plain
An aerial photo of microbial mats around the Grand Prismatic Spring of Yellowstone National Park.
Some of the earliest theories of life were materialist, holding that all that exists is matter, and that all
life is merely a complex form or arrangement of matter. Empedocles (430 BC) argued that every thing in the
universe is made up of a combination of four eternal "elements" or "roots of all": earth, water, air, and fire. All
change is explained by the arrangement and rearrangement of these four elements. The various forms of life
are caused by an appropriate mixture of elements. For example, growth in plants is explained by the natural
downward movement of earth and the natural upward movement of fire.[6]
Democritus (460 BC), the disciple of Leucippus, thought that the essential characteristic of life is
having a soul (psyche). In common with other ancient writers, he used the term to mean the principle of living
things that causes them to function as a living thing. He thought the soul was composed of fire atoms,
because of the apparent connection between life and heat, and because fire moves.[7] He also suggested
that humans originally lived like animals, gradually developing communities to help one another, originating
language, and developing crafts and agriculture.[8]
In the scientific revolution of the 17th century, mechanistic ideas were revived by philosophers like
Descartes.
[edit] Hylomorphism
Hylomorphism is the theory (originating with Aristotle (322 BC)) that all things are a combination of
matter and form. Aristotle was one of the first ancient writers to approach the subject of life in a scientific way.
Biology was one of his main interests, and there is extensive biological material in his extant writings.
According to him, all things in the material universe have both matter and form. The form of a living thing is its
soul (Greek psyche, Latin anima). There are three kinds of souls: the "vegetative soul" of plants, which
causes them to grow and decay and nourish themselves, but does not cause motion and sensation; the
"animal soul" which causes animals to move and feel; and the rational soul which is the source of
consciousness and reasoning which (Aristotle believed) is found only in man.[9] Each higher soul has all the
attributes of the lower one. Aristotle believed that while matter can exist without form, form cannot exist
without matter, and therefore the soul cannot exist without the body.[10]
Consistent with this account is a teleological explanation of life. A teleological explanation accounts
for phenomena in terms of their purpose or goal-directedness. Thus, the whiteness of the polar bear's coat is
explained by its purpose of camouflage. The direction of causality is the other way round from materialistic
science, which explains the consequence in terms of a prior cause. Modern biologists now reject this
functional view in terms of a material and causal one: biological features are to be explained not by looking
forward to future optimal results, but by looking backwards to the past evolutionary history of a species,
which led to the natural selection of the features in question.
[edit] Vitalism
Vitalism is the belief that the life-principle is essentially immaterial. This originated with Stahl (17th
century), and held sway until the middle of the 19th century. It appealed to philosophers such as Henri
Bergson, Nietzsche, Wilhelm Dilthey, anatomists like Bichat, and chemists like Liebig.
Vitalism underpinned the idea of a fundamental separation of organic and inorganic material, and the
belief that organic material can only be derived from living things. This was disproved in 1828 when Friedrich
Wöhler prepared urea from inorganic materials. This so-called Wöhler synthesis is considered the starting
point of modern organic chemistry. It is of great historical significance because for the first time an organic
compound was produced from inorganic reactants.
Later, Helmholtz, anticipated by Mayer, demonstrated that no energy is lost in muscle movement,
suggesting that there were no vital forces necessary to move a muscle. These empirical results led to the
abandonment of scientific interest in vitalistic theories, although the belief lingered on in non-scientific
theories such as homeopathy, which interprets diseases and sickness as caused by disturbances in a
hypothetical vital force or life force.
[edit] Definitions
It is still a challenge for scientists and philosophers to define life in unequivocal terms.[11][12][13]
Defining life is difficult—in part—because life is a process, not a pure substance.[14] Any definition must be
sufficiently broad to encompass all life with which we are familiar, and it should be sufficiently general that,
with it, scientists would not miss life that may be fundamentally different from life on Earth.[15]
[edit] Biology
Since there is no unequivocal definition of life, the current understanding is descriptive, where life is a
characteristic of organisms that exhibit all or most of the following phenomena:[14][16]
1. Homeostasis: Regulation of the internal environment to maintain a constant state; for
example, electrolyte concentration or sweating to reduce temperature.
2. Organization: Being structurally composed of one or more cells, which are the basic units of
life.
3. Metabolism: Transformation of energy by converting chemicals and energy into cellular
components (anabolism) and decomposing organic matter (catabolism). Living things require energy
to maintain internal organization (homeostasis) and to produce the other phenomena associated with
life.
4. Growth: Maintenance of a higher rate of anabolism than catabolism. A growing organism
increases in size in all of its parts, rather than simply accumulating matter.
5. Adaptation: The ability to change over a period of time in response to the environment. This
ability is fundamental to the process of evolution and is determined by the organism's heredity as well
as the composition of metabolized substances, and external factors present.
6. Response to stimuli: A response can take many forms, from the contraction of a unicellular
organism to external chemicals, to complex reactions involving all the senses of multicellular
organisms. A response is often expressed by motion, for example, the leaves of a plant turning
toward the sun (phototropism) and by chemotaxis.
7. Reproduction: The ability to produce new individual organisms, either asexually from a single
parent organism, or sexually from two parent organisms.
[edit] Proposed
To reflect the minimum phenomena required, some have proposed other biological definitions of life:
• Living things are systems that tend to respond to changes in their environment, and inside
themselves, in such a way as to promote their own continuation.[ citation needed]
• A network of inferior negative feedbacks (regulatory mechanisms) subordinated to a superior
positive feedback (potential of expansion, reproduction).[17]
• A systemic definition of life is that living things are self-organizing and autopoietic (self-
producing). Variations of this definition include Stuart Kauffman's definition as an autonomous agent
or a multi-agent system capable of reproducing itself or themselves, and of completing at least one
thermodynamic work cycle.[18]
• Life is a self-sustained chemical system capable of undergoing Darwinian evolution.[19]
• Things with the capacity for metabolism and motion.[14]
• Life is a delay of the spontaneous diffusion or dispersion of the internal energy of the
biomolecules towards more potential microstates.[20]
• Living beings are thermodynamic systems that have an organized molecular structure.[20]
[edit] Viruses
Viruses are most often considered replicators rather than forms of life. They have been described as
"organisms at the edge of life,"[21] since they possess genes, evolve by natural selection,[22] and replicate
by creating multiple copies of themselves through self-assembly. However, viruses do not metabolize and
require a host cell to make new products. Virus self-assembly within host cells has implications for the study
of the origin of life, as it may support the hypothesis that life could have started as self-assembling organic
molecules.[23][24]
[edit] Biophysics
Biophysicists have also commented on the nature and qualities of life forms—notably that they
function on negative entropy.[25][26] In more detail, according to physicists such as John Bernal, Erwin
Schrödinger, Eugene Wigner, and John Avery, life is a member of the class of phenomena which are open or
continuous systems able to decrease their internal entropy at the expense of substances or free energy taken
in from the environment and subsequently rejected in a degraded form (see: entropy and life).[27][28][29]
[edit] Living systems theories

Some scientists have proposed in the last few decades that a general living systems theory is
required to explain the nature of life.[30] Such a general theory, arising out of the ecological and biological
sciences, attempts to map general principles for how all living systems work. Instead of examining
phenomena by attempting to break things down into component parts, a general living systems theory
explores phenomena in terms of dynamic patterns of the relationships of organisms with their environment.
[31]
[edit] Gaia hypothesis

The idea that the Earth is alive is probably as old as humankind, but the first public expression of it as
a fact of science was by a Scottish scientist, James Hutton. In 1785 he stated that the Earth was a
superorganism and that its proper study should be physiology. Hutton is rightly remembered as the father of
geology, but his idea of a living Earth was forgotten in the intense reductionism of the 19th century.[32] The
Gaia hypothesis, originally proposed in the 1960s by scientist James Lovelock,[33][34] explores the idea that
the life on Earth functions as a single organism which actually defines and maintains environmental
conditions necessary for its survival.[35]
[edit] Nonfractionability
Robert Rosen (1991) built on the assumption that the explanatory powers of the mechanistic
worldview cannot help understand the realm of living systems. One of several important clarifications he
made was to define a system component as "a unit of organization; a part with a function, i.e., a definite
relation between part and whole." From this and other starting concepts, he developed a "relational theory of
systems" that attempts to explain the special properties of life. Specifically, he identified the
"nonfractionability of components in an organism" as the fundamental difference between living systems and
"biological machines."[36]
[edit] Life as a property of ecosystems
A systems view of life treats environmental fluxes and biological fluxes together as a "reciprocity of
influence",[37] and a reciprocal relation with environment is arguably as important for understanding life as it
is for understanding ecosystems. As Harold J. Morowitz (1992) explains it, life is a property of an ecological
system rather than a single organism or species.[38] He argues that an ecosystemic definition of life is
preferable to a strictly biochemical or physical one. Robert Ulanowicz (2009) also highlights mutualism as the
key to understand the systemic, order-generating behavior of life and ecosystems.[39]
[edit] Origin
Main article: Origin of life
For religious beliefs about the creation of life, see Creation myth.
Evidence suggests that life on Earth has existed for about 3.7 billion years.[40] All known life forms
share fundamental molecular mechanisms, and based on these observations, theories on the origin of life
attempt to find a mechanism explaining the formation of a primordial single cell organism from which all life
originates. There are many different hypotheses regarding the path that might have been taken from simple
organic molecules via pre-cellular life to protocells and metabolism. Many models fall into the "genes-first"
category or the "metabolism-first" category, but a recent trend is the emergence of hybrid models that
combine both categories.[41]
There is no scientific consensus as to how life originated and all proposed theories are highly
speculative. However, most currently accepted scientific models build in one way or another on the following
hypotheses:
• The Miller-Urey experiment, and the work of Sidney Fox, suggest that conditions on the
primitive Earth may have favored chemical reactions that synthesized some amino acids and other
organic compounds from inorganic precursors.
• Phospholipids spontaneously form lipid bilayers, the basic structure of a cell membrane.
Life as we know it today synthesizes proteins, which are polymers of amino acids using instructions
encoded by cellular genes—which are polymers of deoxyribonucleic acid (DNA). Protein synthesis also
entails intermediary ribonucleic acid (RNA) polymers. One possibility is that genes came first[42] and then
proteins. Another possibility is that proteins came first[43] and then genes. However, because genes are
required to make proteins, and proteins are required to make genes, the problem of considering which came
first is like that of the chicken or the egg. Most scientists have adopted the hypothesis that because DNA and
proteins function together so intimately, it's unlikely that they arose independently.[44] Therefore, many
scientists consider the possibility, apparently first suggested by Francis Crick,[45] that the first life was based
on the DNA-protein intermediary: RNA.[44] In fact, RNA has the DNA-like properties of information storage
and replication and the catalytic properties of some proteins. Crick and others actually favored the RNA-first
hypothesis[46] even before the catalytic properties of RNA had been demonstrated by Thomas Cech.[47]
A significant issue with the RNA-first hypothesis is that experiments designed to synthesize RNA
from simple precursors have not been nearly as successful as the Miller-Urey experiments that synthesized
other organic molecules from inorganic precursors. One reason for the failure to create RNA in the laboratory
is that RNA precursors are very stable and do not react with each other under ambient conditions. However,
the successful synthesis of certain RNA molecules under conditions hypothesized to exist prior to life on
Earth has been achieved by adding alternative precursors in a specified order with the precursor phosphate
present throughout the reaction.[48] This study makes the RNA-first hypothesis more plausible to many
scientists.[49]
Recent experiments have demonstrated true Darwinian evolution of unique RNA enzymes
(ribozymes) made up of two separate catalytic components that replicate each other in vitro.[50] In describing
this work from his laboratory, Gerald Joyce stated: "This is the first example, outside of biology, of
evolutionary adaptation in a molecular genetic system."[51] Such experiments make the possibility of a
primordial RNA World even more attractive to many scientists.
[edit] Conditions for life

The diversity of life on Earth today is a result of the dynamic interplay between genetic opportunity,
metabolic capability, environmental challenges,[52] and symbiosis.[53][54][55] For most of its existence,
Earth's habitable environment has been dominated by microorganisms and subjected to their metabolism
and evolution. As a consequence of such microbial activities on a geologic time scale, the physical-chemical
environment on Earth has been changing, thereby determining the path of evolution of subsequent life.[52]
For example, the release of molecular oxygen by cyanobacteria as a by-product of photosynthesis induced
fundamental, global changes in the Earth's environment. The altered environment, in turn, posed novel
evolutionary challenges to the organisms present, which ultimately resulted in the formation of our planet's
major animal and plant species. Therefore this "co-evolution" between organisms and their environment is
apparently an inherent feature of living systems.[52]
[edit] Range of tolerance

The inert components of an ecosystem are the physical and chemical factors necessary for life—
energy (sunlight or chemical energy), water, temperature, atmosphere, gravity, nutrients, and ultraviolet solar
radiation protection.[56] In most ecosystems the conditions vary during the day and often shift from one
season to the next. To live in most ecosystems, then, organisms must be able to survive a range of
conditions, called "range of tolerance."[57] Outside of that are the "zones of physiological stress," where the
survival and reproduction are possible but not optimal. Outside of these zones are the "zones of intolerance,"
where life for that organism is implausible. It has been determined that organisms that have a wide range of
tolerance are more widely distributed than organisms with a narrow range of tolerance.[57]
[edit] Extremophiles
Deinococcus radiodurans can resist radiation exposure.

Main article: Extremophile
To survive, some microorganisms can assume forms that enable them to withstand freezing,
complete desiccation, starvation, high-levels of radiation exposure, and other physical or chemical
challenges. Furthermore, some microorganisms can survive exposure to such conditions for weeks, months,
years, or even centuries.[52] Extremophiles are microbial life forms that thrive outside the ranges life is
commonly found in. They also excel at exploiting uncommon sources of energy. While all organisms are
composed of nearly identical molecules, evolution has enabled such microbes to cope with this wide range of
physical and chemical conditions. Characterization of the structure and metabolic diversity of microbial
communities in such extreme environments is ongoing. An understanding of the tenacity and versatility of life
on Earth, as well as an understanding of the molecular systems that some organisms utilize to survive such
extremes, will provide a critical foundation for the search for life beyond Earth.[52]
[edit] Chemical element requirements

All life forms require certain core chemical elements needed for biochemical functioning. This list of
core life elements usually includes carbon, hydrogen, nitrogen, oxygen, phosphorus, and sulfur—the "Big Six"
elemental macronutrients for all organisms[58] —often represented by the acronym CHNOPS. Together these
make up nucleic acids, proteins and lipids, the bulk of living matter.
However, alternative hypothetical types of biochemistry have been proposed which eliminate one or
more of these elements, swap out an element for one not on the list, or change required chiralities or other
chemical properties. For example, the recently discovered GFAJ-1 bacteria in Mono Lake, California may be
able to partially substitute phosphorus with arsenic, which is toxic to most forms of life.[59][60]
[edit] Classification of life

Main article: Biological classification
definition by genus and differentia. Life is divided into domains, which are subdivided into further groups.
Intermediate minor rankings are not shown.
Traditionally, people have divided organisms into the classes of plants and animals, based mainly on
their ability of movement. The first known attempt to classify organisms was conducted by the Greek
philosopher Aristotle (384–322 BC). He classified all living organisms known at that time as either a plant or
an animal. Aristotle distinguished animals with blood from animals without blood (or at least without red
blood), which can be compared with the concepts of vertebrates and invertebrates respectively. He divided
the blooded animals into five groups: viviparous quadrupeds (mammals), birds, oviparous quadrupeds
(reptiles and amphibians), fishes and whales. The bloodless animals were also divided into five groups:
cephalopods, crustaceans, insects (which also included the spiders, scorpions, and centipedes, in addition to
what we now define as insects), shelled animals (such as most molluscs and echinoderms) and "zoophytes."
Though Aristotle's work in zoology was not without errors, it was the grandest biological synthesis of the time
and remained the ultimate authority for many centuries after his death.[61]
The exploration of the American continent revealed large numbers of new plants and animals that
needed descriptions and classification. In the latter part of the 16th century and the beginning of the 17th,
careful study of animals commenced and was gradually extended until it formed a sufficient body of
knowledge to serve as an anatomical basis for classification.
In the late 1740s, Carolus Linnaeus introduced his method, still used, to formulate the scientific name
of every species.[62] Linnaeus took every effort to improve the composition and reduce the length of the
many-worded names by abolishing unnecessary rhetoric, introducing new descriptive terms and defining
their meaning with an unprecedented precision. By consistently using his system, Linnaeus separated
nomenclature from taxonomy. This convention for naming species is referred to as binomial nomenclature.
The fungi were originally treated as plants. For a short period Linnaeus had placed them in the taxon
Vermes in Animalia. He later placed them back in Plantae. Copeland classified the Fungi in his Protoctista,
thus partially avoiding the problem but acknowledged their special status.[63] The problem was eventually
solved by Whittaker, when he gave them their own kingdom in his five-kingdom system. As it turned out, the
fungi are more closely related to animals than to plants.[64]
As new discoveries enabled us to study cells and microorganisms, new groups of life were revealed,
and the fields of cell biology and microbiology were created. These new organisms were originally described
separately in protozoa as animals and protophyta/thallophyta as plants, but were united by Haeckel in his
kingdom protista, later the group of prokaryotes were split off in the kingdom Monera, eventually this kingdom
would be divided in two separate groups, the Bacteria and the Archaea, leading to the six-kingdom system
and eventually to the current three-domain system.[65] The classification of eukaryotes is still controversial,
with protist taxonomy especially problematic.[66]
As microbiology, molecular biology and virology developed, non-cellular reproducing agents were
discovered, such as viruses and viroids. Sometimes these entities are considered to be alive but others
argue that viruses are not living organisms since they lack characteristics such as cell membrane,
metabolism and do not grow or respond to their environments. Viruses can however be classed into
"species" based on their biology and genetics but many aspects of such a classification remain controversial.
[67]
Since the 1960s a trend called cladistics has emerged, arranging taxa in an evolutionary or
phylogenetic tree. It is unclear, should this be implemented, how the different codes will coexist.[68]
Linnaeus Haeckel Chatton Copeland Whittaker Woese et al.
1735[69] 1866[70] 1925[71][72] 1938[63][73] 1969[74] 1977[75][76]
2 kingdoms 3 kingdoms 2 empires 4 kingdoms 5 kingdoms 6 kingdoms
Eubacteria
Prokaryota Mychota Monera
(not Archaebacteria
Protista
treated)
Protoctista Protista Protista
Eukaryota Plantae Plantae Plantae

Vegetabilia Plantae
Protoctista Fungi Fungi
Animalia Animalia Animalia Animalia Animalia
[edit] Extraterrestrial life

Main articles: Extraterrestrial life and astrobiology
Panspermia hypothesis
Earth is the only planet in the universe known to harbor life. The Drake equation, which relates the
number of extraterrestrial civilizations in our galaxy with which we might come in contact, has been used to
discuss the probability of life elsewhere, but scientists disagree on many of the values of variables in this
equation. Depending on those values, the equation may either suggest that life arises frequently or
infrequently.
The region around a main sequence star that could support Earth-like life on an Earth-like planet is
known as the habitable zone. The inner and outer radii of this zone vary with the luminosity of the star, as
does the time interval during which the zone will survive. Stars more massive than the Sun have a larger
habitable zone, but will remain on the main sequence for a shorter time interval during which life can evolve.
Small red dwarf stars have the opposite problem, compounded with higher levels of magnetic activity and the
effects of tidal locking from close orbits. Hence, stars in the intermediate mass range such as the Sun may
possess the optimal conditions for Earth-like life to develop. The location of the star within a galaxy may also
have an impact on the likelihood of life forming.
Panspermia, also called exogenesis, is a hypothesis proposing that life originated elsewhere in the
universe and was subsequently transferred to Earth in the form of spores perhaps via meteorites, comets or
cosmic dust. However, this hypothesis does not help explain the ultimate origin of life.
[edit] Death
Main article: Death
Death is the permanent termination of all vital functions or life processes in an organism or cell.[78]
[79] After death, the remains of an organism become part of the biogeochemical cycle. Organisms may be
consumed by a predator or a scavenger and leftover organic material may then be further decomposed by
detritivores, organisms which recycle detritus, returning it to the environment for reuse in the food chain.
One of the challenges in defining death is in distinguishing it from life. Death would seem to refer to
either the moment at which life ends, or when the state that follows life begins.[80] However, determining
when death has occurred requires drawing precise conceptual boundaries between life and death. This is
problematic, however, because there is little consensus over how to define life. The nature of death has for
millennia been a central concern of the world's religious traditions and of philosophical inquiry. Many religions
maintain faith in either some kind of afterlife, reincarnation, or resurrection.
[edit] Extinction
Main article: Extinction
Extinction is the gradual process by which a group of taxa or species dies out, reducing biodiversity.
[81] The moment of extinction is generally considered to be the death of the last individual of that species.
Because a species' potential range may be very large, determining this moment is difficult, and is usually
done retrospectively after a period of apparent absence. Species become extinct when they are no longer
able to survive in changing habitat or against superior competition. Over the history of the Earth, over 99% of
all the species that have ever lived have gone extinct;[82] however, mass extinctions may have accelerated
evolution by providing opportunities for new groups of organisms to diversify.[83]
[edit] Fossils
Main article: Fossil
Fossils are the preserved remains or traces of animals, plants, and other organisms from the remote
past. The totality of fossils, both discovered and undiscovered, and their placement in fossil-containing rock
formations and sedimentary layers (strata) is known as the fossil record. Such a preserved specimen is
called a "fossil" if it is older than the arbitrary date of 10,000 years ago.[84] Hence, fossils range in age from
the youngest at the start of the Holocene Epoch to the oldest from the Archaean Eon, a few billion years old.
[edit] See also

• Alpha taxonomy • Life form
• Artificial life • Life on Mars
• Biological • Meaning of life
immortality • Nature
• Biology—the study • Non-cellular life
of life • Organic life
• Carbon-based life • Organism
• Cell • Origin of life
• Conway's Game • Personal life
of Life • Phylogenetics
• Entropy and life • Prehistoric life
• Evolutionary • Prion
history of life • Quality of life
• Extraterrestrial life • Silicon-based life
• Extremophile • Synthetic life
• Gaia hypothesis
• Genetics
• Genetic
engineering
• Hierarchy of life
• Kingdom (biology)
[edit] Notes
1. ^ The 'evolution' of viruses and other similar forms is still uncertain. Therefore, this
classification may be paraphyletic because cellular life might have evolved from non-cellular life, or
polyphyletic because the most recent common ancestor might not be included.
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• Kauffman, Stuart. The Adjacent Possible: A Talk with Stuart Kauffman
• Nealson KH, Conrad PG (December 1999). "Life: past, present and future". Philos. Trans. R.
Soc. Lond., B, Biol. Sci. 354 (1392): 1923–39. doi:10.1098/rstb.1999.0532. PMID 10670014.
PMC 1692713. http://journals.royalsociety.org/content/7r10hqn3rp1g1vag/.
• Walker, Martin G. LIFE! Why We Exist...And What We Must Do to Survive Dog Ear
Publishing, 2006, ISBN 1-59858-243-7

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W000
Virus

This article is about the biological agent. For other uses, see Virus (disambiguation).
For a generally accessible and less technical introduction to the topic, see Introduction to viruses.
Viruses
Rotavirus
Virus classification
Group: I–VII
Groups
I: dsDNA viruses
II: ssDNA viruses
III: dsRNA viruses
IV: (+)ssRNA viruses
V: (−)ssRNA viruses
VI: ssRNA-RT viruses
VII: dsDNA-RT viruses
A virus is a small infectious agent that can replicate only inside the living cells of organisms. Most
viruses are too small to be seen directly with a light microscope. Viruses infect all types of organisms, from
animals and plants to bacteria and archaea.[1] Since the initial discovery of the tobacco mosaic virus by
Martinus Beijerinck in 1898,[2] about 5,000 viruses have been described in detail,[3] although there are
millions of different types.[4] Viruses are found in almost every ecosystem on Earth and are the most
abundant type of biological entity.[5][6] The study of viruses is known as virology, a sub-speciality of
microbiology.
Virus particles (known as virions) consist of two or three parts: the genetic material made from either
DNA or RNA, long molecules that carry genetic information; a protein coat that protects these genes; and in
some cases an envelope of lipids that surrounds the protein coat when they are outside a cell. The shapes of
viruses range from simple helical and icosahedral forms to more complex structures. The average virus is
about one one-hundredth the size of the average bacterium.
The origins of viruses in the evolutionary history of life are unclear: some may have evolved from
plasmids – pieces of DNA that can move between cells – while others may have evolved from bacteria. In
evolution, viruses are an important means of horizontal gene transfer, which increases genetic diversity.[7]
Viruses spread in many ways; plant viruses are often transmitted from plant to plant by insects that
feed on sap, such as aphids, while animal viruses can be carried by blood-sucking insects. These disease-
bearing organisms are known as vectors. Influenza viruses are spread by coughing and sneezing. The
norovirus and rotavirus, common causes of viral gastroenteritis, are transmitted by the faecal-oral route and
are passed from person to person by contact, entering the body in food or water. HIV is one of several
viruses transmitted through sexual contact and by exposure to infected blood. Viruses can infect only a
limited range of host cells called the "host range". This can be narrow or, as when a virus is capable of
infecting many species, broad.[8]
Viral infections in animals provoke an immune response that usually eliminates the infecting virus.
Immune responses can also be produced by vaccines, which confer an artificially acquired immunity to the
specific viral infection. However, some viruses including those causing AIDS and viral hepatitis evade these
immune responses and result in chronic infections. Antibiotics have no effect on viruses, but several antiviral
drugs have been developed.
Contents
[hide]
• 1 Etymology
• 2 History
• 3 Origins
• 4 Microbiology
• 4.1 Life properties
• 4.2 Structure
• 4.3 Genome
• 4.4 Replication cycle
• 4.5 Effects on the host cell
• 4.6 Host range
• 5.1 ICTV classification
• 5.2 Baltimore classification
• 6 Viruses and human disease
• 6.1 Epidemiology
• 6.2 Epidemics and pandemics
• 6.3 Cancer
• 6.4 Host defence mechanisms
• 6.5 Prevention and treatment
• 6.5.1 Vaccines
[edit] Etymology
The word is from the Latin virus referring to poison and other noxious substances, first used in
English in 1392.[9] Virulent, from Latin virulentus (poisonous), dates to 1400.[10] A meaning of "agent that
causes infectious disease" is first recorded in 1728,[9] before the discovery of viruses by Dmitry Ivanovsky in
1892. The plural is viruses. The adjective viral dates to 1948.[11] The term virion is also used to refer to a
single infective viral particle.
[edit] History
Main article: History of viruses
Martinus Beijerinck in his laboratory in 1921
Louis Pasteur was unable to find a causative agent for rabies and speculated about a pathogen too
small to be detected using a microscope.[12] In 1884, the French microbiologist Charles Chamberland
invented a filter (known today as the Chamberland filter or Chamberland-Pasteur filter) with pores smaller
than bacteria. Thus, he could pass a solution containing bacteria through the filter and completely remove
them from the solution.[13] In 1892, the Russian biologist Dmitry Ivanovsky used this filter to study what is
now known as the tobacco mosaic virus. His experiments showed that crushed leaf extracts from infected
tobacco plants remain infectious after filtration. Ivanovsky suggested the infection might be caused by a toxin
produced by bacteria, but did not pursue the idea.[14] At the time it was thought that all infectious agents
could be retained by filters and grown on a nutrient medium – this was part of the germ theory of disease.[2]
In 1898, the Dutch microbiologist Martinus Beijerinck repeated the experiments and became convinced that
the filtered solution contained a new form of infectious agent.[15] He observed that the agent multiplied only
in cells that were dividing, but as his experiments did not show that it was made of particles, he called it a
contagium vivum fluidum (soluble living germ) and re-introduced the word virus.[14] Beijerinck maintained
that viruses were liquid in nature, a theory later discredited by Wendell Stanley, who proved they were
particulate.[14] In the same year Friedrich Loeffler and Frosch passed the first animal virus – agent of foot-
and-mouth disease (aphthovirus) – through a similar filter.[16]
In the early 20th century, the English bacteriologist Frederick Twort discovered a group of viruses
that infect bacteria, now called bacteriophages[17] (or commonly phages), and the French-Canadian
microbiologist Félix d'Herelle described viruses that, when added to bacteria on agar, would produce areas of
dead bacteria. He accurately diluted a suspension of these viruses and discovered that the highest dilutions
(lowest virus concentrations), rather than killing all the bacteria, formed discrete areas of dead organisms.
Counting these areas and multiplying by the dilution factor allowed him to calculate the number of viruses in
the original suspension.[18] Phages were heralded as a potential treatment for diseases such as typhoid and
cholera, but their promise was forgotten with the development of penicillin. The study of phages provided
insights into the switching on and off of genes, and a useful mechanism for introducing foreign genes into
bacteria.
By the end of the 19th century, viruses were defined in terms of their infectivity, their ability to be
filtered, and their requirement for living hosts. Viruses had been grown only in plants and animals. In 1906,
Ross Granville Harrison invented a method for growing tissue in lymph, and, in 1913, E. Steinhardt, C.
Israeli, and R. A. Lambert used this method to grow vaccinia virus in fragments of guinea pig corneal tissue.
[19] In 1928, H. B. Maitland and M. C. Maitland grew vaccinia virus in suspensions of minced hens' kidneys.
Their method was not widely adopted until the 1950s, when poliovirus was grown on a large scale for vaccine
production.[20]
Another breakthrough came in 1931, when the American pathologist Ernest William Goodpasture
grew influenza and several other viruses in fertilised chickens' eggs.[21] In 1949, John F. Enders, Thomas
Weller, and Frederick Robbins grew polio virus in cultured human embryo cells, the first virus to be grown
without using solid animal tissue or eggs. This work enabled Jonas Salk to make an effective polio vaccine.
[22]
The first images of viruses were obtained upon the invention of electron microscopy in 1931 by the
German engineers Ernst Ruska and Max Knoll.[23] In 1935, American biochemist and virologist Wendell
Meredith Stanley examined the tobacco mosaic virus and found it was mostly made of protein.[24] A short
time later, this virus was separated into protein and RNA parts.[25] The tobacco mosaic virus was the first to
be crystallised and its structure could therefore be elucidated in detail. The first X-ray diffraction pictures of
the crystallised virus were obtained by Bernal and Fankuchen in 1941. On the basis of her pictures, Rosalind
Franklin discovered the full DNA structure of the virus in 1955.[26] In the same year, Heinz Fraenkel-Conrat
and Robley Williams showed that purified tobacco mosaic virus RNA and its coat protein can assemble by
themselves to form functional viruses, suggesting that this simple mechanism was probably the means
through which viruses were created within their host cells.[27]
The second half of the 20th century was the golden age of virus discovery and most of the 2,000
recognised species of animal, plant, and bacterial viruses were discovered during these years.[28][29] In
1957, equine arterivirus and the cause of Bovine virus diarrhea (a pestivirus) were discovered. In 1963, the
hepatitis B virus was discovered by Baruch Blumberg,[30] and in 1965, Howard Temin described the first
retrovirus. Reverse transcriptase, the key enzyme that retroviruses use to translate their RNA into DNA, was
first described in 1970, independently by Howard Martin Temin and David Baltimore.[31] In 1983 Luc
Montagnier's team at the Pasteur Institute in France, first isolated the retrovirus now called HIV.[32]
[edit] Origins
Viruses are found wherever there is life and have probably existed since living cells first evolved.[33]
The origin of viruses is unclear because they do not form fossils, so molecular techniques have been the
most useful means of investigating how they arose.[34] These techniques rely on the availability of ancient
viral DNA or RNA, but, unfortunately, most of the viruses that have been preserved and stored in laboratories
are less than 90 years old.[35][36] There are three main hypotheses that try to explain the origins of viruses:
[37][38]
Regressive hypothesis
Viruses may have once been small cells that parasitised larger cells. Over time, genes not
required by their parasitism were lost. The bacteria rickettsia and chlamydia are living cells that, like
viruses, can reproduce only inside host cells. They lend support to this hypothesis, as their
dependence on parasitism is likely to have caused the loss of genes that enabled them to survive
outside a cell. This is also called the degeneracy hypothesis,[39][40] or reduction hypothesis.[41]
Cellular origin hypothesis
Some viruses may have evolved from bits of DNA or RNA that "escaped" from the genes of a
larger organism. The escaped DNA could have come from plasmids (pieces of naked DNA that can
move between cells) or transposons (molecules of DNA that replicate and move around to different
positions within the genes of the cell).[42] Once called "jumping genes", transposons are examples of
mobile genetic elements and could be the origin of some viruses. They were discovered in maize by
Barbara McClintock in 1950.[43] This is sometimes called the vagrancy hypothesis,[39][44] or the
escape hypothesis.[41]
Coevolution hypothesis
This is also called the virus-first hypothesis[41] and proposes that viruses may have evolved
from complex molecules of protein and nucleic acid at the same time as cells first appeared on earth
and would have been dependent on cellular life for billions of years. Viroids are molecules of RNA that
are not classified as viruses because they lack a protein coat. However, they have characteristics that
are common to several viruses and are often called subviral agents.[45] Viroids are important
pathogens of plants.[46] They do not code for proteins but interact with the host cell and use the host
machinery for their replication.[47] The hepatitis delta virus of humans has an RNA genome similar to
viroids but has a protein coat derived from hepatitis B virus and cannot produce one of its own. It is
therefore a defective virus and cannot replicate without the help of hepatitis B virus.[48] Similarly, the
virophage 'sputnik' is dependent on mimivirus, which infects the protozoan Acanthamoeba castellanii.
[49] These viruses that are dependent on the presence of other virus species in the host cell are called
satellites and may represent evolutionary intermediates of viroids and viruses.[50][51]
Historically, there were problems with all of these hypotheses: The regressive hypothesis did not
explain why even the smallest of cellular parasites do not resemble viruses in any way. The escape
hypothesis did not explain the complex capsids and other structures on virus particles. And, the virus-first
hypothesis contravened the definition of viruses in that they require host cells.[41] However, viruses are now
recognised as ancient and to have origins that pre-date the divergence of life into the three domains.[52] This
discovery has led modern virologists to reconsider and re-evaluate these three classical hypotheses.[52]
The evidence for an ancestral world of RNA cells[53] and computer analysis of viral and host DNA
sequences are giving a better understanding of the evolutionary relationships between different viruses and
may help identify the ancestors of modern viruses. To date, such analyses have not proved which of these
hypotheses are correct.[53] However, it seems unlikely that all currently known viruses have a common
ancestor and viruses have probably arisen numerous times in the past by one or more mechanisms.[54]
Prions are infectious protein molecules that do not contain DNA or RNA.[55] They cause an infection
in sheep called scrapie and cattle bovine spongiform encephalopathy ("mad cow" disease). In humans they
cause kuru and Creutzfeldt-Jakob disease.[56] They are able to replicate because some proteins can exist in
two different shapes and the prion changes the normal shape of a host protein into the prion shape. This
starts a chain reaction where each prion protein converts many host proteins into more prions, and these new
prions then go on to convert even more protein into prions. Although they are fundamentally different from
viruses and viroids, their discovery gives credence to the idea that viruses could have evolved from self-
replicating molecules.[57]
[edit] Microbiology
[edit] Life properties
Opinions differ on whether viruses are a form of life, or organic structures that interact with living
organisms. They have been described as "organisms at the edge of life",[58] since they resemble organisms
in that they possess genes and evolve by natural selection,[59] and reproduce by creating multiple copies of
themselves through self-assembly. Although they have genes, they do not have a cellular structure, which is
often seen as the basic unit of life. Viruses do not have their own metabolism, and require a host cell to make
new products. They therefore cannot naturally reproduce outside a host cell[60] – although bacterial species
such as rickettsia and chlamydia are considered living organisms despite the same limitation.[61][62]
Accepted forms of life use cell division to reproduce, whereas viruses spontaneously assemble within cells.
They differ from autonomous growth of crystals as they inherit genetic mutations while being subject to
natural selection. Virus self-assembly within host cells has implications for the study of the origin of life, as it
lends further credence to the hypothesis that life could have started as self-assembling organic molecules.[1]
[edit] Structure
Diagram of how a virus capsid can be constructed using multiple copies of just two protein molecules
Viruses display a wide diversity of shapes and sizes, called morphologies. Generally viruses are
much smaller than bacteria. Most viruses that have been studied have a diameter between 10 and 300
nanometres. Some filoviruses have a total length of up to 1400 nm; their diameters are only about 80 nm.[63]
Most viruses cannot be seen with a light microscope so scanning and transmission electron microscopes are
used to visualise virions.[64] To increase the contrast between viruses and the background, electron-dense
"stains" are used. These are solutions of salts of heavy metals, such as tungsten, that scatter the electrons
from regions covered with the stain. When virions are coated with stain (positive staining), fine detail is
obscured. Negative staining overcomes this problem by staining the background only.[65]
A complete virus particle, known as a virion, consists of nucleic acid surrounded by a protective coat
of protein called a capsid. These are formed from identical protein subunits called capsomers.[66] Viruses
can have a lipid "envelope" derived from the host cell membrane. The capsid is made from proteins encoded
by the viral genome and its shape serves as the basis for morphological distinction.[67][68] Virally coded
protein subunits will self-assemble to form a capsid, generally requiring the presence of the virus genome.
Complex viruses code for proteins that assist in the construction of their capsid. Proteins associated with
nucleic acid are known as nucleoproteins, and the association of viral capsid proteins with viral nucleic acid
is called a nucleocapsid. The capsid and entire virus structure can be mechanically (physically) probed
through atomic force microscopy.[69][70] In general, there are four main morphological virus types:
RNA coiled in a helix of repeating protein sub-units
Electron micrograph of icosahedral adenovirus

Herpes viruses have a lipid envelope
Helical
These viruses are composed of a single type of capsomer stacked around a central axis to
form a helical structure, which may have a central cavity, or hollow tube. This arrangement results in
rod-shaped or filamentous virions: these can be short and highly rigid, or long and very flexible. The
genetic material, generally single-stranded RNA, but ssDNA in some cases, is bound into the protein
helix by interactions between the negatively charged nucleic acid and positive charges on the protein.
Overall, the length of a helical capsid is related to the length of the nucleic acid contained within it and
the diameter is dependent on the size and arrangement of capsomers. The well-studied tobacco
mosaic virus is an example of a helical virus.[71]
Icosahedral
Most animal viruses are icosahedral or near-spherical with icosahedral symmetry. A regular
icosahedron is the optimum way of forming a closed shell from identical sub-units. The minimum
number of identical capsomers required is twelve, each composed of five identical sub-units. Many
viruses, such as rotavirus, have more than twelve capsomers and appear spherical but they retain this
symmetry. Capsomers at the apices are surrounded by five other capsomers and are called pentons.
Capsomers on the triangular faces are surrounded by six others and are call hexons.[72]
Envelope
Some species of virus envelop themselves in a modified form of one of the cell membranes,
either the outer membrane surrounding an infected host cell, or internal membranes such as nuclear
membrane or endoplasmic reticulum, thus gaining an outer lipid bilayer known as a viral envelope.
This membrane is studded with proteins coded for by the viral genome and host genome; the lipid
membrane itself and any carbohydrates present originate entirely from the host. The influenza virus
and HIV use this strategy. Most enveloped viruses are dependent on the envelope for their infectivity.
[73]
Complex
These viruses possess a capsid that is neither purely helical, nor purely icosahedral, and that
may possess extra structures such as protein tails or a complex outer wall. Some bacteriophages,
such as Enterobacteria phage T4 have a complex structure consisting of an icosahedral head bound to
a helical tail, which may have a hexagonal base plate with protruding protein tail fibres. This tail
structure acts like a molecular syringe, attaching to the bacterial host and then injecting the viral
genome into the cell.[74]
The poxviruses are large, complex viruses that have an unusual morphology. The viral genome is
associated with proteins within a central disk structure known as a nucleoid. The nucleoid is surrounded by a
membrane and two lateral bodies of unknown function. The virus has an outer envelope with a thick layer of
protein studded over its surface. The whole virion is slightly pleiomorphic, ranging from ovoid to brick shape.
[75] Mimivirus is the largest known virus, with a capsid diameter of 400 nm. Protein filaments measuring
100 nm project from the surface. The capsid appears hexagonal under an electron microscope, therefore the
capsid is probably icosahedral.[76]
Some viruses that infect Archaea have complex structures that are unrelated to any other form of
virus, with a wide variety of unusual shapes, ranging from spindle-shaped structures, to viruses that resemble
hooked rods, teardrops or even bottles. Other archaeal viruses resemble the tailed bacteriophages, and can
have multiple tail structures.[77]
[edit] Genome
Genomic diversity among viruses
Property Parameters
• DNA
• RNA
Nucleic acid
• Both DNA and RNA (at different stages in the
life cycle)
Shape • Linear
• Circular
• Segmented
• Single-stranded
• Double-stranded
Strandedness
• Double-stranded with regions of single-
strandedness
• Positive sense (+)

Sense • Negative sense (−)
• Ambisense (+/−)
An enormous variety of genomic structures can be seen among viral species; as a group they contain
more structural genomic diversity than plants, animals, archaea, or bacteria. There are millions of different
types of viruses,[4] although only about 5,000 of them have been described in detail.[3] A virus has either
DNA or RNA genes and is called a DNA virus or a RNA virus respectively. The vast majority of viruses have
RNA genomes. Plant viruses tend to have single-stranded RNA genomes and bacteriophages tend to have
double-stranded DNA genomes.[78]
Viral genomes are circular, as in the polyomaviruses, or linear, as in the adenoviruses. The type of
nucleic acid is irrelevant to the shape of the genome. Among RNA viruses and certain DNA viruses, the
genome is often divided up into separate parts, in which case it is called segmented. For RNA viruses, each
segment often codes for only one protein and they are usually found together in one capsid. However, all
segments are not required to be in the same virion for the virus to be infectious, as demonstrated by brome
mosaic virus and several other plant viruses.[63]
A viral genome, irrespective of nucleic acid type, is almost always either single-stranded or double-
stranded. Single-stranded genomes consist of an unpaired nucleic acid, analogous to one-half of a ladder
split down the middle. Double-stranded genomes consist of two complementary paired nucleic acids,
analogous to a ladder. The virus particles of some virus families, such as those belonging to the
Hepadnaviridae, contain a genome that is partially double-stranded and partially single-stranded.[78]
For most viruses with RNA genomes and some with single-stranded DNA genomes, the single
strands are said to be either positive-sense (called the plus-strand) or negative-sense (called the minus-
strand), depending on whether or not they are complementary to the viral messenger RNA (mRNA). Positive-
sense viral RNA is in the same sense as viral mRNA and thus at least a part of it can be immediately
translated by the host cell. Negative-sense viral RNA is complementary to mRNA and thus must be
converted to positive-sense RNA by an RNA-dependent RNA polymerase before translation. DNA
nomenclature for viruses with single-sense genomic ssDNA is similar to RNA nomenclature, in that the
coding strand for the viral mRNA is complementary to it (−), and the non-coding strand is a copy of it (+).[78]
However, several types of ssDNA and ssRNA viruses have genomes which are ambisense in that
transcription can occur off both strands in a double-stranded replicative intermediate. Examples include
geminiviruses, which are ssDNA plant viruses and arenaviruses, which are ssRNA viruses of animals.[79]
Genome size varies greatly between species. The smallest viral genomes – the ssDNA circoviruses,
family Circoviridae – code for only two proteins and have a genome size of only 2 kilobases; the largest –
mimiviruses – have genome sizes of over 1.2 megabases and code for over one thousand proteins.[80] RNA
viruses generally have smaller genome sizes than DNA viruses because of a higher error-rate when
replicating, and have a maximum upper size limit.[81] Beyond this limit, errors in the genome when
replicating render the virus useless or uncompetitive. To compensate for this, RNA viruses often have
segmented genomes – the genome is split into smaller molecules – thus reducing the chance that an error in a
single-component genome will incapacitate the entire genome. In contrast, DNA viruses generally have
larger genomes because of the high fidelity of their replication enzymes.[82] Single-strand DNA viruses are
an exception to this rule, however, as mutation rates for these genomes can approach the extreme of the
ssRNA virus case.[83]
How antigenic shift, or reassortment, can result in novel and highly pathogenic strains of human
influenza
Viruses undergo genetic change by several mechanisms. These include a process called genetic
drift where individual bases in the DNA or RNA mutate to other bases. Most of these point mutations are
"silent" – they do not change the protein that the gene encodes – but others can confer evolutionary
advantages such as resistance to antiviral drugs.[84] Antigenic shift occurs when there is a major change in
the genome of the virus. This can be a result of recombination or reassortment. When this happens with
influenza viruses, pandemics might result.[85] RNA viruses often exist as quasispecies or swarms of viruses
of the same species but with slightly different genome nucleoside sequences. Such quasispecies are a prime
target for natural selection.[86]
Segmented genomes confer evolutionary advantages; different strains of a virus with a segmented
genome can shuffle and combine genes and produce progeny viruses or (offspring) that have unique
characteristics. This is called reassortment or viral sex.[87]
Genetic recombination is the process by which a strand of DNA is broken and then joined to the end
of a different DNA molecule. This can occur when viruses infect cells simultaneously and studies of viral
evolution have shown that recombination has been rampant in the species studied.[88] Recombination is
common to both RNA and DNA viruses.[89][90]
[edit] Replication cycle

Viral populations do not grow through cell division, because they are acellular. Instead, they use the
machinery and metabolism of a host cell to produce multiple copies of themselves, and they assemble in the
cell.
A typical virus replication cycle
Some bacteriophages inject their genomes into bacterial cells
The life cycle of viruses differs greatly between species but there are six basic stages in the life cycle
of viruses:[91]
• Attachment is a specific binding between viral capsid proteins and specific receptors on the
host cellular surface. This specificity determines the host range of a virus. For example, HIV infects a
limited range of human leucocytes. This is because its surface protein, gp120, specifically interacts
with the CD4 molecule – a chemokine receptor – which is most commonly found on the surface of
CD4+ T-Cells. This mechanism has evolved to favour those viruses that only infect cells in which
they are capable of replication. Attachment to the receptor can induce the a viral envelope protein to
undergo changes that results in the fusion of viral and cellular membranes, or changes of non-
enveloped virus surface proteins that allow the virus to enter.
• Penetration follows attachment: virions enter the host cell through receptor mediated
endocytosis or membrane fusion. This is often called viral entry. The infection of plant and probably
fungal cells is different from that of animal cells. Plants have a rigid cell wall made of cellulose, and
fungi one of chitin, so most viruses can only get inside the cells after trauma to the cell wall.[92]
However, nearly all plant viruses (such as tobacco mosaic virus) can also move directly from cell to
cell, in the form of single-stranded nucleoprotein complexes, through pores called plasmodesmata.
This process requires movement proteins, which are virus-encoded proteins probably originally
derived from plant proteins, which interact with the plasmodesmatal transport machinery [93]
Bacteria, like plants, have strong cell walls that a virus must breach to infect the cell. However, given
that bacterial cell walls are much less thick than plant cell walls due to their much smaller size, some
viruses have evolved mechanisms that inject their genome into the bacterial cell across the cell wall,
while the viral capsid remains outside.[94]
• Uncoating is a process in which the viral capsid is removed: this may be by degradation by
viral enzymes or host enzymes or by simple dissociation; the end result is the releasing of the viral
genomic nucleic acid.
• Replication of viruses primarily involves multiplication of the genome; however, this involves
synthesis of viral messenger RNA (mRNA) for all viruses except some positive sense RNA viruses,
from "early" genes; viral protein synthesis, possible assembly of viral proteins, then viral genome
replication mediated by early or regulatory protein expression. This may be followed, for complex
viruses with larger genomes, by one or more further rounds of mRNA synthesis: "late" gene
expression is generally of structural or virion proteins.
• Following the structure-mediated self- assembly of the virus particles, some modification of
the proteins often occurs. In viruses such as HIV, this modification (sometimes called maturation)
occurs after the virus has been released from the host cell.[95]
• Viruses can be released from the host cell by lysis, a process that kills the cell by bursting its
membrane and cell wall if present: this is a feature of many bacterial and some animal viruses. Some
viruses undergo a lysogenic cycle where the viral genome is incorporated by genetic recombination
into a specific place in the host's chromosome. The viral genome is then known as a "provirus" or, in
the case of bacteriophages a "prophage".[96] Whenever the host divides, the viral genome is also
replicated. The viral genome is mostly silent within the host; however, at some point, the provirus or
prophage may give rise to active virus, which may lyse the host cells.[97] Enveloped viruses (e.g.,
HIV) typically are released from the host cell by budding. During this process the virus acquires its
envelope, which is a modified piece of the host's plasma or other, internal membrane.[98]
The genetic material within virus particles, and the method by which the material is replicated, varies
considerably between different types of viruses.
DNA viruses
The genome replication of most DNA viruses takes place in the cell's nucleus. If the cell has
the appropriate receptor on its surface, these viruses enter the cell sometimes by direct fusion with the
cell membrane (e.g.: herpesviruses) or – more usually – by receptor-mediated endocytosis. Most DNA
viruses are entirely dependent on the host cell's DNA and RNA synthesising machinery, and RNA
processing machinery; however, viruses with larger genomes may encode much of this machinery
themselves. In eukaryotes the viral genome must cross the cell's nuclear membrane to access this
machinery, while in bacteria it need only enter the cell.[99]
RNA viruses
Replication usually takes place in the cytoplasm. RNA viruses can be placed into four different
groups depending on their modes of replication. The polarity (whether or not it can be used directly by
ribosomes to make proteins) of single-stranded RNA viruses largely determines the replicative
mechanism; the other major criterion is whether the genetic material is single-stranded or double-
stranded. All RNA viruses use their own RNA replicase enzymes to create copies of their genomes.
[100]
Reverse transcribing viruses
These have ssRNA (Retroviridae, Metaviridae, Pseudoviridae) or dsDNA (Caulimoviridae, and
Hepadnaviridae) in their particles. Reverse transcribing viruses with RNA genomes (retroviruses), use
a DNA intermediate to replicate, whereas those with DNA genomes (pararetroviruses) use an RNA
intermediate during genome replication. Both types use a reverse transcriptase, or RNA-dependent
DNA polymerase enzyme, to carry out the nucleic acid conversion. Retroviruses integrate the DNA
produced by reverse transcription into the host genome as a provirus as a part of the replication
process; pararetroviruses do not, although integrated genome copies of especially plant
pararetroviruses can give rise to infectious virus.[101] They are susceptible to antiviral drugs that
inhibit the reverse transcriptase enzyme, e.g. zidovudine and lamivudine. An example of the first type
is HIV, which is a retrovirus. Examples of the second type are the Hepadnaviridae, which includes
Hepatitis B virus.[102]
[edit] Effects on the host cell

The range of structural and biochemical effects that viruses have on the host cell is extensive.[103]
These are called cytopathic effects.[104] Most virus infections eventually result in the death of the host cell.
The causes of death include cell lysis, alterations to the cell's surface membrane and apoptosis.[105] Often
cell death is caused by cessation of its normal activities because of suppression by virus-specific proteins,
not all of which are components of the virus particle.[106]
Some viruses cause no apparent changes to the infected cell. Cells in which the virus is latent and
inactive show few signs of infection and often function normally.[107] This causes persistent infections and
the virus is often dormant for many months or years. This is often the case with herpes viruses.[108][109]
Some viruses, such as Epstein-Barr virus, can cause cells to proliferate without causing malignancy,[110]
while others, such as papillomaviruses, are established causes of cancer.[111]
[edit] Host range

Viruses are by far the most abundant parasites on earth and they have been found to infect all types
of cellular life including animals, plants and bacteria.[3] However, different types of viruses can only infect a
limited range of hosts and many are species-specific. Some, such as smallpox virus for example, can only
infect one species – in this case humans,[112] and are said to have a narrow host range. Other viruses, such
as rabies virus, can infect different species of mammals and are said to have a broad range.[113] The viruses
that infect plants are harmless to animals and most viruses that infect other animals are harmless to humans.
[114] The host range of some bacteriophages is limited to a single strain of bacteria and they can be used to
trace the source of outbreaks of infections by a method called phage typing.[115]
Main article: Virus classification
Classification seeks to describe the diversity of viruses by naming and grouping them on the basis of
similarities. In 1962, André Lwoff, Robert Horne, and Paul Tournier were the first to develop a means of virus
classification, based on the Linnaean hierarchical system.[116] This system bases classification on phylum,
class, order, family, genus, and species. Viruses were grouped according to their shared properties (not
those of their hosts) and the type of nucleic acid forming their genomes.[117] Later the International
Committee on Taxonomy of Viruses was formed. However, viruses are not classified on the basis of phylum
or class, as their small genome size and high rate of mutation makes it difficult to determine their ancestry
beyond Order. As such, the Baltimore Classification is used to supplement the more traditional hierarchy.
[edit] ICTV classification

The International Committee on Taxonomy of Viruses (ICTV) developed the current classification
system and wrote guidelines that put a greater weight on certain virus properties to maintain family
uniformity. A unified taxonomy (a universal system for classifying viruses) has been established. The 7th
lCTV Report formalised for the first time the concept of the virus species as the lowest taxon (group) in a
branching hierarchy of viral taxa.[118] However, at present only a small part of the total diversity of viruses
has been studied, with analyses of samples from humans finding that about 20% of the virus sequences
recovered have not been seen before, and samples from the environment, such as from seawater and ocean
sediments, finding that the large majority of sequences are completely novel.[119]
The general taxonomic structure is as follows:
Order (-virales)
Family (-viridae)
Subfamily (-virinae)
Genus (-virus)
Species (-virus)
In the current (2008) ICTV taxonomy, five orders have been established, the Caudovirales,
Herpesvirales, Mononegavirales, Nidovirales, and Picornavirales. The committee does not formally
distinguish between subspecies, strains, and isolates. In total there are 5 orders, 82 families, 11 subfamilies,
307 genera, 2,083 species and about 3,000 types yet unclassified.[120][121]
[edit] Baltimore classification

Main article: Baltimore classification
The Baltimore Classification of viruses is based on the method of viral mRNA synthesis.
The Nobel Prize-winning biologist David Baltimore devised the Baltimore classification system.[31]
[122] The ICTV classification system is used in conjunction with the Baltimore classification system in
modern virus classification.[123][124][125]
The Baltimore classification of viruses is based on the mechanism of mRNA production. Viruses
must generate mRNAs from their genomes to produce proteins and replicate themselves, but different
mechanisms are used to achieve this in each virus family. Viral genomes may be single-stranded (ss) or
double-stranded (ds), RNA or DNA, and may or may not use reverse transcriptase (RT). Additionally, ssRNA
viruses may be either sense (+) or antisense (−). This classification places viruses into seven groups:
• I: dsDNA viruses (e.g. Adenoviruses, Herpesviruses, Poxviruses)
• II: ssDNA viruses (+)sense DNA (e.g. Parvoviruses)
• III: dsRNA viruses (e.g. Reoviruses)
• IV: (+)ssRNA viruses (+)sense RNA (e.g. Picornaviruses, Togaviruses)
• V: (−)ssRNA viruses (−)sense RNA (e.g. Orthomyxoviruses, Rhabdoviruses)
• VI: ssRNA-RT viruses (+)sense RNA with DNA intermediate in life-cycle (e.g. Retroviruses)
• VII: dsDNA-RT viruses (e.g. Hepadnaviruses)
As an example of viral classification, the chicken pox virus, varicella zoster (VZV), belongs to the
order Herpesvirales, family Herpesviridae, subfamily Alphaherpesvirinae, and genus Varicellovirus. VZV is in
Group I of the Baltimore Classification because it is a dsDNA virus that does not use reverse transcriptase.
[edit] Viruses and human disease

See also: Table of clinically important viruses
Overview of the main types of viral infection and the most notable species involved[126][127]
Examples of common human diseases caused by viruses include the common cold, influenza,
chickenpox and cold sores. Many serious diseases such as ebola, AIDS, avian influenza and SARS are
caused by viruses. The relative ability of viruses to cause disease is described in terms of virulence. Other
diseases are under investigation as to whether they too have a virus as the causative agent, such as the
possible connection between human herpes virus six (HHV6) and neurological diseases such as multiple
sclerosis and chronic fatigue syndrome.[128] There is controversy over whether the borna virus, previously
thought to cause neurological diseases in horses, could be responsible for psychiatric illnesses in humans.
[129]
Viruses have different mechanisms by which they produce disease in an organism, which largely
depends on the viral species. Mechanisms at the cellular level primarily include cell lysis, the breaking open
and subsequent death of the cell. In multicellular organisms, if enough cells die the whole organism will start
to suffer the effects. Although viruses cause disruption of healthy homeostasis, resulting in disease, they may
exist relatively harmlessly within an organism. An example would include the ability of the herpes simplex
virus, which causes cold sores, to remain in a dormant state within the human body. This is called
latency[130] and is a characteristic of the herpes viruses including Epstein-Barr virus, which causes
glandular fever, and varicella zoster virus, which causes chickenpox and shingles. Most people have been
infected with at least one of these types of herpes virus.[131] However, these latent viruses might sometimes
be beneficial, as the presence of the virus can increase immunity against bacterial pathogens, such as
Yersinia pestis.[132]
Some viruses can cause life-long or chronic infections, where the viruses continue to replicate in the
body despite the host's defence mechanisms.[133] This is common in hepatitis B virus and hepatitis C virus
infections. People chronically infected are known as carriers, as they serve as reservoirs of infectious virus.
[134] In populations with a high proportion of carriers, the disease is said to be endemic.[135]
[edit] Epidemiology
Viral epidemiology is the branch of medical science that deals with the transmission and control of
virus infections in humans. Transmission of viruses can be vertical, that is from mother to child, or horizontal,
which means from person to person. Examples of vertical transmission include hepatitis B virus and HIV
where the baby is born already infected with the virus.[136] Another, more rare, example is the varicella
zoster virus, which although causing relatively mild infections in humans, can be fatal to the foetus and newly
born baby.[137]
Horizontal transmission is the most common mechanism of spread of viruses in populations.
Transmission can occur when: body fluids are exchanged during sexual activity, e.g. HIV; blood is
exchanged by contaminated transfusion or needle sharing, e.g. hepatitis C; a child is born to an infected
mother, e.g. hepatitis B; exchange of saliva by mouth, e.g. Epstein-Barr virus; contaminated food or water is
ingested, e.g. norovirus; aerosols containing virions are inhaled, e.g. influenza virus; and insect vectors such
as mosquitoes penetrate the skin of a host, e.g. dengue. The rate or speed of transmission of viral infections
depends on factors that include population density, the number of susceptible individuals, (i.e. those who are
not immune),[138] the quality of health care and the weather.[139]
Epidemiology is used to break the chain of infection in populations during outbreaks of viral diseases.
[140] Control measures are used that are based on knowledge of how the virus is transmitted. It is important
to find the source, or sources, of the outbreak and to identify the virus. Once the virus has been identified, the
chain of transmission can sometimes be broken by vaccines. When vaccines are not available sanitation and
disinfection can be effective. Often infected people are isolated from the rest of the community and those that
have been exposed to the virus placed in quarantine.[141] To control the outbreak of foot and mouth disease
in cattle in Britain in 2001, thousands of cattle were slaughtered.[142] Most viral infections of humans and
other animals have incubation periods during which the infection causes no signs or symptoms.[143]
Incubation periods for viral diseases range from a few days to weeks but are known for most infections.[144]
Somewhat overlapping, but mainly following the incubation period, there is a period of communicability; a
time when an infected individual or animal is contagious and can infect another person or animal.[145] This
too is known for many viral infections and knowledge the length of both periods is important in the control of
outbreaks.[146] When outbreaks cause an unusually high proportion of cases in a population, community or
region they are called epidemics. If outbreaks spread worldwide they are called pandemics.[147]
[edit] Epidemics and pandemics
See also: Spanish flu, AIDS, and Ebola
For more details on this topic, see List of epidemics.
Transmission electron microscope image of a recreated 1918 influenza virus

Native American populations were devastated by contagious diseases, particularly smallpox, brought
to the Americas by European colonists. It is unclear how many Native Americans were killed by foreign
diseases after the arrival of Columbus in the Americas, but the numbers have been estimated to be close to
70% of the indigenous population. The damage done by this disease significantly aided European attempts
to displace and conquer the native population.[148]
A pandemic is a worldwide epidemic. The 1918 flu pandemic, commonly referred to as the Spanish
flu, was a category 5 influenza pandemic caused by an unusually severe and deadly influenza A virus. The
victims were often healthy young adults, in contrast to most influenza outbreaks, which predominantly affect
juvenile, elderly, or otherwise weakened patients.[149]
The Spanish flu pandemic lasted from 1918 to 1919. Older estimates say it killed 40–50 million
people,[150] while more recent research suggests that it may have killed as many as 100 million people, or
5% of the world's population in 1918.[151] Most researchers believe that HIV originated in sub-Saharan
Africa during the 20th century;[152] it is now a pandemic, with an estimated 38.6 million people now living
with the disease worldwide.[153] The Joint United Nations Programme on HIV/AIDS (UNAIDS) and the World
Health Organization (WHO) estimate that AIDS has killed more than 25 million people since it was first
recognised on June 5, 1981, making it one of the most destructive epidemics in recorded history.[154] In
2007 there were 2.7 million new HIV infections and 2 million HIV-related deaths.[155]
Marburg virus
Several highly lethal viral pathogens are members of the Filoviridae. Filoviruses are filament-like
viruses that cause viral hemorrhagic fever, and include the ebola and marburg viruses. The Marburg virus
attracted widespread press attention in April 2005 for an outbreak in Angola. Beginning in October 2004 and
continuing into 2005, the outbreak was the world's worst epidemic of any kind of viral hemorrhagic fever.[156]
[edit] Cancer
For more details on this topic, see Oncovirus.
Viruses are an established cause of cancer in humans and other species. Viral cancers only occur in
a minority of infected persons (or animals). Cancer viruses come from a range of virus families, including
both RNA and DNA viruses, and so there is no single type of "oncovirus" (an obsolete term originally used for
acutely transforming retroviruses). The development of cancer is determined by a variety of factors such as
host immunity[157] and mutations in the host.[158] Viruses accepted to cause human cancers include some
genotypes of human papillomavirus, hepatitis B virus, hepatitis C virus, Epstein-Barr virus, Kaposi's sarcoma-
associated herpesvirus and human T-lymphotropic virus. The most recently discovered human cancer virus
is a polyomavirus (Merkel cell polyomavirus) that causes most cases of a rare form of skin cancer called
Merkel cell carcinoma.[159] Hepatitis viruses can develop into a chronic viral infection that leads to liver
cancer.[160][161] Infection by human T-lymphotropic virus can lead to tropical spastic paraparesis and adult
T-cell leukemia.[162] Human papillomaviruses are an established cause of cancers of cervix, skin, anus, and
penis.[163] Within the Herpesviridae, Kaposi's sarcoma-associated herpesvirus causes Kaposi's sarcoma
and body cavity lymphoma, and Epstein–Barr virus causes Burkitt's lymphoma, Hodgkin’s lymphoma, B
lymphoproliferative disorder and nasopharyngeal carcinoma.[164] Merkel cell polyomavirus closely related to
SV40 and mouse polyomaviruses that have been used as animal models for cancer viruses for over 50
years.[165]
[edit] Host defence mechanisms

See also: Immune system
The body's first line of defence against viruses is the innate immune system. This comprises cells
and other mechanisms that defend the host from infection in a non-specific manner. This means that the cells
of the innate system recognise, and respond to, pathogens in a generic way, but unlike the adaptive immune
system, it does not confer long-lasting or protective immunity to the host.[166]
RNA interference is an important innate defence against viruses.[167] Many viruses have a
replication strategy that involves double-stranded RNA (dsRNA). When such a virus infects a cell, it releases
its RNA molecule or molecules, which immediately bind to a protein complex called dicer that cuts the RNA
into smaller pieces. A biochemical pathway called the RISC complex is activated, which degrades the viral
mRNA and the cell survives the infection. Rotaviruses avoid this mechanism by not uncoating fully inside the
cell and by releasing newly produced mRNA through pores in the particle's inner capsid. The genomic
dsRNA remains protected inside the core of the virion.[168][169]
When the adaptive immune system of a vertebrate encounters a virus, it produces specific antibodies
that bind to the virus and render it non-infectious. This is called humoral immunity. Two types of antibodies
are important. The first, called IgM, is highly effective at neutralizing viruses but is only produced by the cells
of the immune system for a few weeks. The second, called IgG, is produced indefinitely. The presence of IgM
in the blood of the host is used to test for acute infection, whereas IgG indicates an infection sometime in the
past.[170] IgG antibody is measured when tests for immunity are carried out.[171]
Two rotaviruses: the one on the right is coated with antibodies that stop its attaching to cells and
infecting them
A second defence of vertebrates against viruses is called cell-mediated immunity and involves
immune cells known as T cells. The body's cells constantly display short fragments of their proteins on the
cell's surface, and if a T cell recognises a suspicious viral fragment there, the host cell is destroyed by killer T
cells and the virus-specific T-cells proliferate. Cells such as the macrophage are specialists at this antigen
presentation.[172] The production of interferon is an important host defence mechanism. This is a hormone
produced by the body when viruses are present. Its role in immunity is complex, but it eventually stops the
viruses from reproducing by killing the infected cell and its close neighbours.[173]
Not all virus infections produce a protective immune response in this way. HIV evades the immune
system by constantly changing the amino acid sequence of the proteins on the surface of the virion. These
persistent viruses evade immune control by sequestration, blockade of antigen presentation, cytokine
resistance, evasion of natural killer cell activities, escape from apoptosis, and antigenic shift.[174] Other
viruses, called neurotropic viruses, are disseminated by neural spread where the immune system may be
unable to reach them.
[edit] Prevention and treatment

Because viruses use vital metabolic pathways within host cells to replicate, they are difficult to
eliminate without using drugs that cause toxic effects to host cells in general. The most effective medical
approaches to viral diseases are vaccinations to provide immunity to infection, and antiviral drugs that
selectively interfere with viral replication.
[edit] Vaccines
For more details on this topic, see Vaccination.
Vaccination is a cheap and effective way of preventing infections by viruses. Vaccines were used to
prevent viral infections long before the discovery of the actual viruses. Their use has resulted in a dramatic
decline in morbidity (illness) and mortality (death) associated with viral infections such as polio, measles,
mumps and rubella.[175] Smallpox infections have been eradicated.[176] Vaccines are available to prevent
over thirteen viral infections of humans,[177] and more are used to prevent viral infections of animals.[178]
Vaccines can consist of live-attenuated or killed viruses, or viral proteins (antigens).[179] Live vaccines
contain weakened forms of the virus, which do not cause the disease but nonetheless confer immunity. Such
viruses are called attenuated. Live vaccines can be dangerous when given to people with a weak immunity,
(who are described as immunocompromised), because in these people, the weakened virus can cause the
original disease.[180] Biotechnology and genetic engineering techniques are used to produce subunit
vaccines. These vaccines use only the capsid proteins of the virus. Hepatitis B vaccine is an example of this
type of vaccine.[181] Subunit vaccines are safe for immunocompromised patients because they cannot
cause the disease.[182] The yellow fever virus vaccine, a live-attenuated strain called 17D, is probably the
safest and most effective vaccine ever generated.[183]
[edit] Antiviral drugs

For more details on this topic, see Antiviral drug.
Antiviral drugs are often nucleoside analogues, (fake DNA building
blocks), which viruses mistakenly incorporate into their genomes during
replication. The life-cycle of the virus is then halted because the newly
Guanosine The
guanosine
analogue
Aciclovir
synthesised DNA is inactive. This is because these analogues lack the hydroxyl groups, which, along with
phosphorus atoms, link together to form the strong "backbone" of the DNA molecule. This is called DNA
chain termination.[184] Examples of nucleoside analogues are aciclovir for Herpes simplex virus infections
and lamivudine for HIV and Hepatitis B virus infections. Aciclovir is one of the oldest and most frequently
prescribed antiviral drugs.[185] Other antiviral drugs in use target different stages of the viral life cycle. HIV is
dependent on a proteolytic enzyme called the HIV-1 protease for it to become fully infectious. There is a
large class of drugs called protease inhibitors that inactivate this enzyme.
Hepatitis C is caused by an RNA virus. In 80% of people infected, the disease is chronic, and without
treatment, they are infected for the remainder of their lives. However, there is now an effective treatment that
uses the nucleoside analogue drug ribavirin combined with interferon.[186] The treatment of chronic carriers
of the hepatitis B virus by using a similar strategy using lamivudine has been developed.[187]
[edit] Infection in other species

Main article: Animal virology
Viruses infect all cellular life and, although viruses occur universally, each cellular species has its
own specific range that often infect only that species.[188] Some viruses, called satellites, can only replicate
within cells that have already been infected by another virus.[189] Viruses are important pathogens of
livestock. Diseases such as Foot and Mouth Disease and bluetongue are caused by viruses.[190]
Companion animals such as cats, dogs, and horses, if not vaccinated, are susceptible to serious viral
infections. Canine parvovirus is caused by a small DNA virus and infections are often fatal in pups.[191] Like
all invertebrates, the honey bee is susceptible to many viral infections.[192] Fortunately, most viruses co-
exist harmlessly in their host and cause no signs or symptoms of disease.[2]
[edit] Plants
Main article: Plant pathology
Peppers infected by mild mottle virus

There are many types of plant virus, but often they cause only a loss of yield, and it is not
economically viable to try to control them. Plant viruses are often spread from plant to plant by organisms,
known as vectors. These are normally insects, but some fungi, nematode worms and single-celled organisms
have been shown to be vectors. When control of plant virus infections is considered economical, for
perennial fruits for example, efforts are concentrated on killing the vectors and removing alternate hosts such
as weeds.[193] Plant viruses are harmless to humans and other animals because they can reproduce only in
living plant cells.[194]
Plants have elaborate and effective defence mechanisms against viruses. One of the most effective
is the presence of so-called resistance (R) genes. Each R gene confers resistance to a particular virus by
triggering localised areas of cell death around the infected cell, which can often be seen with the unaided eye
as large spots. This stops the infection from spreading.[195] RNA interference is also an effective defence in
plants.[196] When they are infected, plants often produce natural disinfectants that kill viruses, such as
salicylic acid, nitric oxide, and reactive oxygen molecules.[197]
Plant virus particles or virus-like particles (VLPs) have applications in both biotechnology and
nanotechnology. The capsids of most plant viruses are simple and robust structures and can be produced in
large quantities either by the infection of plants or by expression in a variety of heterologous systems. Plant
virus particles can be modified genetically and chemically to encapsulate foreign material and can be
incorporated into supramolecular structures for use in biotechnology.[198]
[edit] Bacteria
Main article: Bacteriophage
Transmission electron micrograph of multiple bacteriophages attached to a bacterial cell wall
Bacteriophages are a common and diverse group of viruses and are the most abundant form of
biological entity in aquatic environments – there are up to ten times more of these viruses in the oceans than
there are bacteria,[199] reaching levels of 250,000,000 bacteriophages per millilitre of seawater.[200] These
viruses infect specific bacteria by binding to surface receptor molecules and then entering the cell. Within a
short amount of time, in some cases just minutes, bacterial polymerase starts translating viral mRNA into
protein. These proteins go on to become either new virions within the cell, helper proteins, which help
assembly of new virions, or proteins involved in cell lysis. Viral enzymes aid in the breakdown of the cell
membrane, and, in the case of the T4 phage, in just over twenty minutes after injection over three hundred
phages could be released.[201]
The major way bacteria defend themselves from bacteriophages is by producing enzymes that
destroy foreign DNA. These enzymes, called restriction endonucleases, cut up the viral DNA that
bacteriophages inject into bacterial cells.[202] Bacteria also contain a system that uses CRISPR sequences
to retain fragments of the genomes of viruses that the bacteria have come into contact with in the past, which
allows them to block the virus's replication through a form of RNA interference.[203][204] This genetic system
provides bacteria with acquired immunity to infection.
[edit] Archaea
Some viruses replicate within archaea: these are double-stranded DNA viruses with unusual and
sometimes unique shapes.[5][77] These viruses have been studied in most detail in the thermophilic
archaea, particularly the orders Sulfolobales and Thermoproteales.[205] Defences against these viruses may
involve RNA interference from repetitive DNA sequences within archaean genomes that are related to the
genes of the viruses.[206][207]
[edit] Role in aquatic ecosystems

Main article: Marine bacteriophage
Viruses are the most abundant biological entity in aquatic environments:[1] a teaspoon of seawater
contains about one million of them.[208] They are essential to the regulation of saltwater and freshwater
ecosystems.[209] Most of these viruses are bacteriophages, which are harmless to plants and animals. They
infect and destroy the bacteria in aquatic microbial communities, comprising the most important mechanism
of recycling carbon in the marine environment. The organic molecules released from the bacterial cells by the
viruses stimulates fresh bacterial and algal growth.[210]
Microorganisms constitute more than 90% of the biomass in the sea. It is estimated that viruses kill
approximately 20% of this biomass each day and that there are fifteen times as many viruses in the oceans
as there are bacteria and archaea. Viruses are the main agents responsible for the rapid destruction of
harmful algal blooms,[211] which often kill other marine life.[212] The number of viruses in the oceans
decreases further offshore and deeper into the water, where there are fewer host organisms.[213]
The effects of marine viruses are far-reaching; by increasing the amount of photosynthesis in the
oceans, viruses are indirectly responsible for reducing the amount of carbon dioxide in the atmosphere by
approximately 3 gigatonnes of carbon per year.[213]
Like any organism, marine mammals are susceptible to viral infections. In 1988 and 2002 thousands
of harbour seals were killed in Europe by phocine distemper virus.[214] Many other viruses, including
caliciviruses, herpesviruses, adenoviruses and parvoviruses, circulate in marine mammal populations.[213]
[edit] Role in evolution

Main article: Horizontal gene transfer
Viruses are an important natural means of transferring genes between different species, which
increases genetic diversity and drives evolution.[7] It is thought that viruses played a central role in the early
evolution, before the diversification of bacteria, archaea and eukaryotes and at the time of the last universal
common ancestor of life on Earth.[215] Viruses are still one of the largest reservoirs of unexplored genetic
diversity on the Earth.[213]
[edit] Applications
[edit] Life sciences and medicine
Scientist studying the H5N1 influenza virus.
Viruses are important to the study of molecular and cellular biology as they provide simple systems
that can be used to manipulate and investigate the functions of cells.[216] The study and use of viruses have
provided valuable information about aspects of cell biology.[217] For example, viruses have been useful in
the study of genetics and helped our understanding of the basic mechanisms of molecular genetics, such as
DNA replication, transcription, RNA processing, translation, protein transport, and immunology.
Geneticists often use viruses as vectors to introduce genes into cells that they are studying. This is
useful for making the cell produce a foreign substance, or to study the effect of introducing a new gene into
the genome. In similar fashion, virotherapy uses viruses as vectors to treat various diseases, as they can
specifically target cells and DNA. It shows promising use in the treatment of cancer and in gene therapy.
Eastern European scientists have used phage therapy as an alternative to antibiotics for some time, and
interest in this approach is increasing, because of the high level of antibiotic resistance now found in some
pathogenic bacteria.[218]
Expression of heterologous proteins by viruses is the basis of several manufacturing processes that
are currently being used for the production of various proteins such as vaccine antigens and antibodies.
Industrial processes have been recently developed using viral vectors and a number of pharmaceutical
proteins are currently in pre-clinical and clinical trials.[219]
[edit] Materials science and nanotechnology

Current trends in nanotechnology promise to make much more versatile use of viruses. From the
viewpoint of a materials scientist, viruses can be regarded as organic nanoparticles. Their surface carries
specific tools designed to cross the barriers of their host cells. The size and shape of viruses, and the number
and nature of the functional groups on their surface, is precisely defined. As such, viruses are commonly
used in materials science as scaffolds for covalently linked surface modifications. A particular quality of
viruses is that they can be tailored by directed evolution. The powerful techniques developed by life sciences
are becoming the basis of engineering approaches towards nanomaterials, opening a wide range of
applications far beyond biology and medicine.[220]
Because of their size, shape, and well-defined chemical structures, viruses have been used as
templates for organizing materials on the nanoscale. Recent examples include work at the Naval Research
Laboratory in Washington, DC, using Cowpea Mosaic Virus (CPMV) particles to amplify signals in DNA
microarray based sensors. In this application, the virus particles separate the fluorescent dyes used for
signalling to prevent the formation of non-fluorescent dimers that act as quenchers.[221] Another example is
the use of CPMV as a nanoscale breadboard for molecular electronics.[222]
[edit] Synthetic viruses

Many viruses can be synthesized de novo (“from scratch”) and the first synthetic virus was created in
2002.[223] Although somewhat of a misconception, it is not the actual virus that is synthesized, but rather its
DNA genome (in case of a DNA virus), or a cDNA copy of its genome (in case of RNA viruses). For many
virus families the naked synthetic DNA or RNA (once enzymatically converted back from the synthetic cDNA)
is infectious when introduced into a cell. That is, they contain all the necessary information to produce new
viruses. This technology is now being used to investigate novel vaccine strategies.[224] The ability to
synthesize viruses has far-reaching consequences, since viruses can no longer be regarded as extinct, as
long as the information of their genome sequence is known and permissive cells are available. Currently, the
full-length genome sequences of 2408 different viruses (including smallpox) are publicly available at an
online database, maintained by the National Institute of Health.[225]
[edit] Weapons
For more details on this topic, see Biological warfare.
The ability of viruses to cause devastating epidemics in human societies has led to the concern that
viruses could be weaponised for biological warfare. Further concern was raised by the successful recreation
of the infamous 1918 influenza virus in a laboratory.[226] The smallpox virus devastated numerous societies
throughout history before its eradication. There are officially only two centers in the world which keep stocks
of smallpox virus – the Russian Vector laboratory, and the United States Centers for Disease Control.[227]
But fears that it may be used as a weapon are not totally unfounded;[227] the vaccine for smallpox has
sometimes severe side-effects – during the last years before the eradication of smallpox disease more people
became seriously ill as a result of vaccination than did people from smallpox[228] — and smallpox vaccination
is no longer universally practiced.[229] Thus, much of the modern human population has almost no
established resistance to smallpox.[227]
[edit] References
[edit] Notes
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15.^ Dimmock p.4–5
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• In 1887, Buist visualised one of the largest, Vaccinia virus, by optical microscopy
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[edit] Bibliography
• Collier, Leslie; Balows, Albert; Sussman, Max (1998) Topley and Wilson's Microbiology and
Microbial Infections ninth edition, Volume 1, Virology, volume editors: Mahy, Brian and Collier, Leslie.
Arnold. ISBN 0-340-66316-2.
• Dimmock, N.J; Easton, Andrew J; Leppard, Keith (2007) Introduction to Modern Virology
sixth edition, Blackwell Publishing, ISBN 1-4051-3645-6.
• Knipe, David M; Howley, Peter M; Griffin, Diane E; Lamb, Robert A; Martin, Malcolm A;
Roizman, Bernard; Straus Stephen E. (2007) Fields Virology, Lippincott Williams & Wilkins. ISBN 0-
7817-6060-7.
• Shors, Teri (2008). Understanding Viruses. Jones and Bartlett Publishers. ISBN 0-7637-
2932-9.

• ViralZone A Swiss Institute of Bioinformatics resource for all viral families, providing general
molecular and epidemiological information
• "A Gazillion Tiny Avatars" , article on viruses by Olivia Judson, NY Times, Dec 15, 2009
• Viruses – an Open Access journal
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W000
Pterodactylus

Pterodactylus
Gnathostomata
Pterodactylus
Temporal range: Late Jurassic, 150.8–148.5 Ma
PreЄ ↓
g
Well preserved specimen, Bürgermeister Müller Museum, Solnhofen
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: †Pterosauria
Suborder: †Pterodactyloidea
(unranked): †Euctenochasmatia
†Pterodactylidae
Family:
Meyer, 1830
†Pterodactylus
Genus:
Cuvier, 1809
†P. antiquus
Species:
(Sömmerring, 1812)
Synonyms
Ornithocephalus antiquus Sömmering, 1812

Ornithocephalus brevirostris Sömmering, 1816
Pterodactylus longirostris Cuvier, 1809
Pterodactylus crocodilocephaloidea Ritgen, 1826
Ornithocephalus kochi Wagner, 1837
Pterodactylus mayeri Muenster, 1842
Pterodactylus scolopaciceps Meyer, 1850
Macrotrachelus longirostris (Cuvier, 1809) Giebel, 1852
Pterodactylus kochi (Wagner, 1837) Meyer, 1859
Pterodactylus spectabilis Meyer, 1861
Diopecephalus kochi (Wagner, 1837) Seeley, 1871
Ptenodracon brevirostris (Sömmering, 1816) Lydekker, 1888
Pterodactylus (pronounced /ˌtɛrəˈdæktɨləs/ TERR-ə-DAK-til-əs) is a genus of pterosaur, the first to be
named and identified as a flying reptile. Its fossil remains have been found primarily in the Solnhofen
limestone of Bavaria, Germany, dated to the late Jurassic Period (early Tithonian), about 150.8-148.5 million
years ago,[1] though more fragmentary remains have been identified from elsewhere in Europe and in Africa.
It was a carnivore and probably preyed upon fish and other small animals. Like all pterosaurs, the wings of
Pterodactylus were formed by a skin and muscle membrane stretching from its elongated fourth finger to its
hind limbs. It was supported internally by collagen fibres and externally by keratinous ridges.
The name derives from the Greek words pteron (πτερόn, meaning 'wing') and daktylos (δάκτυλος,
meaning 'finger') and refers to the way in which the wing is supported by one large finger.
Contents
[hide]
• 1 Description
• 2 Paleobiology
• 2.1 Year classes
• 2.2 Growth and breeding seasons
• 3 History
• 4.1 Species
• 4.2 List of species and synonyms
• 5 References
[edit] Description
Size of the sub-adult holotype (green) and hypothetical adult (gray) in both flying and standing
postures, compared with a human
Pterodactylus is known from over 27 fossil specimens, and though most of those are juveniles, many
preserve complete skeletons.[2] The discovery of several specimens with well-preserved soft tissue traces
has allowed scientists to faithfully reconstruct the life appearance of Pterodactylus. Pterodactylus was a
relatively small pterosaur, with an estimated adult wingspan of about 1.5 meters (5 ft) in P. antiquus.[2] Other
"species" were once thought to be smaller. However, these smaller specimens have been shown to
represent juveniles of Pterodactylus, as well as its contemporary relatives Ctenochasma, Germanodactylus
and Gnathosaurus.[3]
The skulls of adult Pterodactylus were long and narrow with about 90 large, conical teeth. The teeth
extended back from the tips of both jaws, and became smaller farther away from the jaw tips (unlike some
relatives, where teeth were absent in the upper jaw tip and were relatively uniform in size). The teeth
extended farther back into the jaw than in close relatives, as some were present below the front of the
nasoantorbital fenestra, the largest opening in the skull. Unlike related species, the skull and jaws were
straight, not curved upwards.[4] A small, hooked beak was present in the very tips of the jaws, with both
upper and lower hook no larger than the teeth that surrounded them.[5]
The neck was long, and covered in long, bristle-like pycnofibres. A throat pouch extended from about
the middle of the lower jaw to the upper part of the neck.[6]
Reconstructed life appearance

Pterodactylus, like related pterosaurs, had a crest on its skull composed mainly of soft tissues. In
adult Pterodactylus, this crest extended between the back edge of the antorbital fenestra (the largest opening
in the skull) and the back of the skull. The back of the crest extended upward into a backward-curving cone-
shaped structure. The crest was composed mainly of long, hardened fibres (twisted together in a spiral
pattern inside the conical part of the crest), and covered in scales. In at least one specimen of P. longicollum,
the crest had a short bony base, also seen in related pterosaurs like Germanodactylus. Crests have only
been found on large, fully adult specimens of Pterodactylus, indicating that this was a display structure and
only developed when individuals reached maturity.[6]
The wings were long, and the wing membranes appear to have lacked the furry covering of
pycnofibres present in some other pterosaurs (such as Pterorhynchus and Jeholopterus). The wing
membrane extended between the fingers and toes as webbing, and a uropatagium (secondary membrane
between the feet and tail) was present, as well as a propatagium (membrane between the wrist and
shoulder).[6] Both the finger and toe claws were covered in keratin sheaths that extended and curved into
sharp hooks well beyond their bony cores.[5]
[edit] Paleobiology
[edit] Year classes
Juvenile specimen (formerly P. kochi), Museum für Naturkunde, Berlin

Like other pterosaurs (notably Rhamphorhynchus), Pterodactylus specimens can vary considerably
based on age or level of maturity. Both the proportions of the limb bones, size and shape of the skull, and
size and number of teeth changed as the animals grew. Historically, this has led to various growth stages
(including growth stages of related pterosaurs) being mistaken for new species of Pterodactylus. Several
detailed studies using various methods to measure growth curves among known specimens have
demonstrated that there is actually only one valid Pterodactylus species, P. antiquus.[4]
Cast of the sub-adult type specimen of P. antiquus, Carnegie Museum of Natural History
The youngest immature Pterodactylus specimens have a small number of teeth (as few as 15), and
the teeth have a relatively broad base.[2] The teeth of older specimens are both narrower and more
numerous (up to 90 teeth are present in some specimens).[4]
Pterodactylus specimens can be divided into two distinct year classes. In the first year class, the
skulls are only 15-45mm in length. The second year class is characterized by skulls 55-95mm long, but still
immature. These first two size groups were once classified as juveniles and adults of the species P. kochi,
until further study showed that even the supposed "adults" were immature. A third year class is represented
by specimens of the "traditional" P. antiquus, as well as a few isolated, large specimens once assigned to P.
kochi that overlap P. antiquus in size. However, all specimens in this third year class also show sign of
immaturity. Fully mature Pterodactylus specimens remain unknown, or may have been mistakenly classified
as a different genus.[2]
[edit] Growth and breeding seasons
The distinct year classes of Pterodactylus antiquus specimens show that this species, like the
contemporary Rhamphorhynchus muensteri, likely bred seasonally and grew consistently during its lifetime.
A new generation of 1st year class P. antiquus would have been produced seasonally, and reached 2nd-year
size by the time the next generation hatched, creating distinct 'clumps' of similarly-sized and aged individuals
in the fossil record. The smallest size class probably consisted of individuals that had just begun to fly and
were less than one year old.[2][7] The second year class represents individuals one to two years old, and the
rare third year class is composed of specimens over two years old. This growth pattern is similar to modern
crocodilians, rather than the rapid growth of modern birds.[2]
[edit] History
Original copper engraving of the P. antiquus holotype by Egid Verhelst II and published by Cosimo
Collini, 1784
The animal now known as Pterodactylus was the first pterosaur ever to be identified. The first
Pterodactylus specimen was described by the Italian scientist Cosimo Alessandro Collini in 1784, based on a
fossil skeleton unearthed from the Solnhofen limestone of Bavaria. Collini was the curator of the
"Naturalienkabinett", or nature cabinet (a precursor to the modern concept of the natural history museum), in
the palace of Charles Theodore, Elector of Bavaria at Mannheim.[8] The specimen had been given to the
collection by Count Friedrich Ferdinand zu Pappenheim, probably around 1780, having been recovered from
a lithographic limestone quarry in Eichstätt.[9] Collini, however, did not conclude that the specimen was a
flying animal. In fact, Collini could not fathom what kind of animal it might have been, rejecting affinities with
the birds or the bats. He speculated that it may have been a sea creature, not for any anatomical reason, but
because he thought the ocean depths were more likely to have housed unknown types of animals.[10][11]
The idea that pterosaurs were aquatic animals persisted among a minority of scientists as late as 1830, when
the German zoologist Johann Georg Wagler published a text on "amphibians" which included an illustration
of Pterodactylus using its wings as flippers. Wagler went so far as to classify Pterodactylus, along with other
aquatic vertebrates (namely plesiosaurs, ichthyosaurs, and monotremes), in the class Gryphi, between birds
and mammals.[12]
Hermann's original life restoration, the first of any pterosaur, 1800
It was the German/French scientist Johann Hermann who first stated that Pterodactylus used its long
fourth finger to support a wing membrane. In March 1800, Hermann alerted the French scientist George
Cuvier to the existence of Collini's fossil, believing that it had been captured by the occupying armies of
Napoleon and sent to the French collections in Paris (and perhaps to Cuvier himself) as war booty; at the
time special French political commissars systematically seized art treasures and objects of scientific interest.
Hermann sent Cuvier a letter containing his own interpretation of the specimen (though he had not examined
it personally), which he believed to be a mammal, including the first known life restoration of a pterosaur.
Hermann restored the animal with wing membranes extending from the long fourth finger to the ankle and a
covering of fur (neither wing membranes nor fur had been preserved in the specimen). Hermann also added
a membrane between the neck and wrist, as is the condition in bats. Cuvier agreed with this interpretation,
and at Hermann's suggestion, Cuvier became the first to publish these ideas in December 1800 in a very
short description.[11] Cuvier remarked, "[It is not possible to doubt that the long finger served to support a
membrane that, by lengthening the anterior extremity of this animal, formed a good wing.]"[13] However,
contrary to Hermann, Cuvier was convinced the animal was a reptile.
The specimen had not in fact been seized by the French. Rather, in 1802, following the death of
Charles Theodore, it was brought to Munich, where Baron Johann Paul Carl von Moll had obtained a general
exemption of confiscation for the Bavarian collections. Cuvier asked von Moll to study the fossil but was
informed it could not be found. In 1809 Cuvier published a somewhat longer description, naming the animal,
in which he refuted a hypothesis by Johann Friedrich Blumenbach that it would have been a shore bird.
Von Soemmerring's incorrect restoration of P. brevirostris

Contrary to von Moll's report, the fossil was not missing; it was being studied by Samuel Thomas von
Sömmerring, who gave a public lecture about it on 27 December 1810. In January 1811, von Sömmerring
wrote a letter to Cuvier deploring the fact that he had only recently been informed of Cuvier's request for
information. His lecture was published in 1812, and in it von Sömmerring named the species
Ornithocephalus antiquus.[14] The animal was described as being both a mammal, a bat, and a form in
between mammals and birds, i.e. not intermediate in descent but in "affinity" or archetype. Cuvier disagreed,
and the same year in his Ossemens fossiles provided a lengthy description in which he restated that the
animal was a reptile.[15] It was not until 1817 that a second specimen of Pterodactylus came to light, again
from Solnhofen. This tiny specimen was that year described by von Soemmerring as Ornithocephalus
brevirostris (for its short snout, now understood to be a juvenile character), and provided a restoration of the
skeleton, the first one published for any pterosaur.[11] This restoration was very inaccurate, von
Soemmerring mistaking the long metacarpals for the bones of the lower arm, the lower arm for the humerus,
this upper arm for the breast bone and this sternum again for the shoulder blades.[16] Von Soemmerring did
not change his opinion that these forms were bats and this "bat model" for interpreting pterosaurs would
remain influential long after a consensus had been reached around 1860 that they were reptiles. The
standard assumptions were that pterosaurs were quadrupedal, clumsy on the ground, furred, warmblooded
and had a wing membrane reaching the ankle.[17] Some of these elements have been confirmed, some
refuted by modern research, while others remain disputed.
In 1998, the discovery of one specimen assigned to P. kochi shed light on the life appearance of
Pterodactylus, as it preserved unique soft-tissue traits not present in previous fossil skeletons, including long,
bristly pycnofibres (a fur-like body covering known only in pterosaurs) on the neck, details of a urpatagium
(hind wing membrane between the legs and tail) that also stretched between the toes as webbing, and a
pelican-like throat pouch.[6] An additional specimen, studied using ultra-violet light, revealed even more
information on the soft anatomy of Pterodactylus. This specimen (catelogue number JME SOS 4784) showed
that like many other pterosaurs, Pterodactylus had a striated soft-tissue crest on the skull. Soft tissue
impressions also showed unusually long, sharp, and recurved keratin sheaths on its claws. This specimen
was also covered in hair-like pycnofibres, with unusually long pycnofibres covering the back of its neck. The
remains of a small, hooked beak were preserved at the tips of the jaws between its upper and lower front
teeth.[5]
Outdated depiction (left, on branch), with several Rhamphorhynchus flying in the background, by
Heinrich Harder, c. 1920
The genus now known as Pterodactylus was originally named Petro-Dactyle by Cuvier in 1809,[18]
though this was a typographical error, later corrected by him to Ptéro-Dactyle.[11] In 1812, Samuel Thomas
von Sömmerring named the same specimen Ornithocephalus antiquus. As the senior synonym, Cuvier's
name has precedence, so applying modern rules for the combination of the genus name and the specific
name the valid species name became Ptéro-Dactyle antiquus. The genus name was Latinized to the current
Pterodactylus by Constantine Samuel Rafinesque in 1815, which by these rules is the valid name as they do
not allow for diacritics or hyphens. Unaware of Rafinesque's publication Cuvier himself in 1819 again
Latinized the genus,[19] but the specific name he then gave, longirostris, has to give precedence to von
Soemmerring's antiquus. In 1888 Richard Lydekker designated Pterodactylus antiquus the type species. The
original specimen is the holotype of the genus, BSP No. AS.I.739.
Hermann von Meyer, in 1830, used the name Pterodactyli to contain Pterodactylus and other
pterosaurs known at the time. This was emended to the family Pterodactylidae by Prince Charles Lucien
Bonaparte in 1838. This family has more recently been used to refer to many similar species from Germany
and elsewhere, though recent studies suggest it may be a paraphyletic or polyphyletic unnatural grouping
with respect to more advanced members of the Ctenochasmatoidea (or Archaeopterodactyloidea).[20][21]
[edit] Species
P. antiquus, AMNH 1942, a specimen showing muscle impressions

Numerous species have been assigned to Pterodactylus in the years since its discovery. In the first
half of the nineteenth century any new pterosaur species would be named Pterodactylus, which thus became
a typical "waste-basket taxon". Even after clearly different forms had later been given their own generic
name, new species would be created from the very productive late Jurassic German sites, often based on
only slightly different material.
Around 1980, subsequent revisions by Peter Wellnhofer had reduced the number of recognized
species to about half a dozen. Many species assigned to Pterodactylus had been based on juvenile
specimens, and subsequently been recognized as immature individuals of other species or genera. By the
1990s it was understood that this was even true for part of the remaining species. P. elegans, for example,
was found by numerous studies to be an immature Ctenochasma.[4] Another species of Pterodactylus based
on small, immature specimens is P. micronyx. However, it has been difficult to determine exactly of what
genus and species P. micronyx might be the juvenile form. Stéphane Jouve, Christopher Bennett and others
suggested that it probably belongs either to Gnathosaurus subulatus or one of the Ctenochasma species, but
more data and study would be required to determine which one.[3][4]
The only well-known and well-supported species left were P. antiquus and P. kochi. However, most
studies since the 1990s have found little reason to separate even these two, and have treated them as
synonymous.[20][22] In 1996, Bennett suggested that the differences between specimens of P. kochi and P.
antiquus could be explained by differences in age. In a 2004 paper, Jouve used a different method of
analysis and recovered the same result, showing that the "distinctive" features of P. kochi were age-related,
and using mathematical comparison to show that the two forms are different growth stages of the same
species.[4]
A special case is P. longicollum, named by von Meyer in 1854, based on a large specimen with a
long neck and fewer teeth. Many researchers, including David Unwin, have found P. longicollum to be distinct
from P. kochi and P. antiquus. Unwin found P. longicollum to be closer to Germanodactylus and therefore
requiring a new genus name.[20] It has sometimes been placed in the genus Diopecephalus because Harry
Govier Seeley based this genus partly on the P. longicollum material. However, it was shown by Bennett that
the type specimen later designated for Diopecephalus was a fossil belonging to P. kochi, and no longer
thought to be separate from Pterodactylus. Diopecephalus is therefore a synonym of Pterodactylus, and as
such is unavailable for use as a new genus for "P." longicollum.[22]
[edit] List of species and synonyms
During its over 200 year history, the various species of Pterodactylus have gone through a number of
changes in classification, and thus have acquired a large number of synonyms. Additionally, a number of
species assigned to Pterodactylus are based on poor remains that have proven difficult to assign to one
species or another, and are therefore considered nomina dubia ("doubtful names"). The following list includes
names that are based on German material presently, or until recently, thought to be pertaining to
Pterodactylus proper and names based on other material that has as yet not been assigned to other genera.
Historical diagram of a juvenile P. antiquus (formerly Ptenodracon brevirostris)

Specimen assigned to P. longicollum
P. antiquus specimen formerly classified as P. longirostris
Holotype fossil of the dubious species P. grandipelvis
Specimen formerly assigned to P. spectabilis

Name Author Year Status Notes
(von
Pterodactylus (1812) Originally Ptéro-
Sömmerring) Valid, type species
antiquus 1815 dactyle, Cuvier 1809
Rafinesque
Ornithocephalus von Synonym of Objective junior

1812
antiquus Sömmerring Pterodactylus antiquus synonym of P. antiquus
Ornithocephalus von 1816- Synonym of Based on a

brevirostris Sömmerring 17 Pterodactylus antiquus juvenile specimen
Reclassified from
(von
Ptenodracon (1816- Synonym of Pterodactylus
Sömmerring)
brevirostris 17) 1888 Pterodactylus antiquus brevirostris, based on a
Lydekker
juvenile specimen
Name correction
Pterodactylus (von (1816- Synonym of from Ornithocephalus
brevirostris Sömmerring) Oken 17) 1819 Pterodactylus antiquus brevirostris, based on a
juvenile specimen
Pterodactylus Synonym of
Cuvier 1819 -
longirostris Pterodactylus antiquus
Macrotrachelus (Cuvier) (1819) Synonym of Reclassified from
longirostris Giebel 1852 Pterodactylus antiquus Pterodactylus longirostri
Incorrect name
Ornithocephalus (Cuvier) (1819) Synonym of
correction from
longirostris Ritgen 1826 Pterodactylus antiquus
Pterodactylus longirostri
Oken 1825 Nomen nudum
"suevicus" Pterodactylus antiquus
Pterodactylus Ritgen 1826 Synonym of
crocodilocephaloides Pterodactylus antiquus
von Meyer 1861
spectabilis Pterodactylus antiquus
Nomen dubium,
possible synonym of
Pterodactylus grandis Cuvier 1824
Rhamphorhynchus
muensteri
Pterodactylus Valid, may belong
von Meyer 1854
longicollum to a separate genus
Pterodactylus Possible synonym

von Meyer 1852
micronyx of Gnathosaurus subulatus
Ritgen 1826
nettecephaloides Pterodactylus micronyx
Ornithocephalus Synonym of
Wagner 1851
redenbacheri Pterodactylus micronyx
Name correction
Pterodactylus (Wagner) (1851) Synonym of
from Ornithocephalus
redenbacheri Wagner 1861 Pterodactylus micronyx
redenbacheri
Pterodactylus von Meyer 1861 Synonym of
pulchellus Pterodactylus micronyx
Name correction
Synonym of
Pterodactylus kochi (Wagner) (1837) from Ornithocephalus
Pterodactylus antiquus
kochi
Ornithocephalus Synonym of
Wagner 1837
kochi Pterodactylus kochi
(Wagner) (1837) Synonym of Reclassification
Diopecephalus kochi
Seeley 1871 Pterodactylus kochi of Pterodactylus kochi
Synonym of
Pterodactylus meyeri Muenster 1842
Pterodactylus kochi
Incorrect name
Ornithocephalus (Muenster) (1842) Synonym of
correction from
meyeri Wagner 1851 Pterodactylus kochi
Pterodactylus meyeri
von Meyer 1850
scolopaciceps Pterodactylus kochi
Reclassification
Rhamphorhynchus Synonym of
(von Meyer) (1850) of Pterodactylus
scolopaciceps Pterodactylus kochi
scolopaciceps
Pterodactylus von Meyer 1860 Nomen dubium
cerinensis
Pterodactylus
von Meyer 1860 Nomen dubium
grandipelvis
Pterodactylus
Owen 1874 Nomen dubium
manseli
Pterodactylus
Owen 1874 Nomen dubium
pleydelli
Pterodactylus
Sauvage 1873 Nomen dubium
suprajurensis
Pterodactylus arningi Reck 1931 Nomen dubium
Pterodactylus
Reck 1931 Nomen dubium
maximus
[edit] References
1. ^ Schweigert, G. (2007). "Ammonite biostratigraphy as a tool for dating Upper Jurassic
lithographic limestones from South Germany – first results and open questions." Neues Jahrbuch für
Geologie und Paläontologie – Abhandlungen, 245(1): 117-125. doi:10.1127/0077-7749/2007/0245-
0117
2. ^ a b c d e f Bennett, S.C. (1996). "Year-classes of pterosaurs from the Solnhofen Limestone
of Germany: Taxonomic and Systematic Implications". Journal of Vertebrate Paleontology 16 (3):
432–444. doi:10.1080/02724634.1996.10011332.
3. ^ a b Bennett, S.C. (2002). "Soft tissue preservation of the cranial crest of the pterosaur
Germanodactylus from Solnhofen". Journal of Vertebrate Paleontology 22 (1): 43–48.
doi:10.1671/0272-4634(2002)022[0043:STPOTC]2.0.CO;2.
4. ^ a b c d e f Jouve, S. (2004). "Description of the skull of a Ctenochasma (Pterosauria) from
the latest Jurassic of eastern France, with a taxonomic revision of European Tithonian
Pterodactyloidea". Journal of Vertebrate Paleontology 24 (3): 542–554. doi:10.1671/0272-
4634(2004)024[0542:DOTSOA]2.0.CO;2.
5. ^ a b c Frey, E., Tischilinger, H., Buchy, M.-C. and Martill, D.M. (2003). "New specimens of
Pterosauria (Reptilia) with soft partes with implications for pterosaurian anatomy and locomotion. In:
Buffetaut, E. and Mazin, J.-M. (eds.)". Evolution and Palaeobiology of Pterosaurs (London:
Geological Society) 217: 233–266.
6. ^ a b c d Frey, E., and Martill, D.M. (1998). "Soft tissue preservation in a specimen of
Pterodactylus kochi (Wagner) from the Upper Jurassic of Germany". Neues Jahrbuch für Geologie
und Paläontologie, Abhandlungen 210: 421–441.
7. ^ Wellnhofer, P. (1970). Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke
Siiddeutschlands. Bayerische Akademie der Wissenschaften, Mathematisch-Wissenschaftlichen
Klasse, Abhandlungen, 141: 133 pp.
8. ^ Unwin, David M. (2006). The Pterosaurs: From Deep Time. New York: Pi Press. pp. 246.
ISBN ISBN 0-13-146308-X.
9. ^ Brougham, H.P. (1844). Dialogues on instinct; with analytical view of the researches on
fossil osteology. Volume 19 of Knight's weekly vol.
10.^ Collini, C A. (1784). "Sur quelques Zoolithes du Cabinet d’Histoire naturelle de S. A. S. E.
Palatine & de Bavière, à Mannheim." Acta Theodoro-Palatinae Mannheim 5 Pars Physica , pp. 58–
103 (1 plate).
11.^ a b c d Taquet, P., and Padian, K. (2004). "The earliest known restoration of a pterosaur and
the philosophical origins of Cuvier’s Ossemens Fossiles". Comptes Rendus Palevol 3 (2): 157–175.
doi:10.1016/j.crpv.2004.02.002.
12.^ Wagler, J. (1830). Natürliches System der Amphibien Munich, 1830: 1-354.
13.^ Cuvier, G. (1801). "[Reptile volant]. In: Extrait d’un ouvrage sur les espèces de
quadrupèdes dont on a trouvé les ossemens dans l’intérieur de la terre". Journal de Physique, de
Chimie et d’Histoire Naturelle 52: 253–267.
14.^ von Sömmerring, S. T. (1812). "Über einen Ornithocephalus oder über das unbekannten
Thier der Vorwelt, dessen Fossiles Gerippe Collini im 5. Bande der Actorum Academiae Theodoro-
Palatinae nebst einer Abbildung in natürlicher Grösse im Jahre 1784 beschrieb, und welches
Gerippe sich gegenwärtig in der Naturalien-Sammlung der königlichen Akademie der
Wissenschaften zu München befindet", Denkschriften der königlichen bayerischen Akademie der
Wissenschaften, München: mathematisch-physikalische Classe 3: 89–158
15.^ Cuvier, G. (1812). Recherches sur les ossemens fossiles. I ed. p. 24, tab. 31
16.^ Sömmering, T. v. (1817). "Über einen Ornithocephalus brevirostris der Vorwelt". Denkschr.
Kgl. Bayer Akad. Wiss., math.phys. Cl. 6: 89–104.
17.^ Padian, K. (1987). "The case of the bat-winged pterosaur. Typological taxonomy and the
influence of pictorial representation on scientific perception". Pp. 65-81 in: Czerkas, S. J. and Olson,
E. C., eds. Dinosaurs past and present. An exhibition and symposium organized by the Natural
History Museum of Los Angeles County. Volume 2. Natural History Museum of Los Angeles County
and University of Washington Press, Seattle and London
18.^ Cuvier, G. (1809). "Mémoire sur le squelette fossile d’un reptile volant des environs
d’Aichstedt, que quelques naturalistes ont pris pour un oiseau, et dont nous formons un genre de
Sauriens, sous le nom de Petro-Dactyle". Annales du Muséum national d’Histoire Naturelle, Paris 13:
424–437.
19.^ Cuvier, G., 1819, (Pterodactylus longirostris) in Isis von Oken, 1126 und 1788, Jena
20.^ a b c Unwin, D. M., (2003). "On the phylogeny and evolutionary history of pterosaurs." Pp.
139-190. in Buffetaut, E. & Mazin, J.-M., (eds.) (2003). Evolution and Palaeobiology of Pterosaurs.
Geological Society of London, Special Publications 217, London, 1-347.
21.^ Kellner, A.W.A. (2003). "Pterosaur phylogeny and comments on the evolutionary history of
the group." Pp. 105-137 in Buffetaut, E. and Mazin, J.-M., (eds.) (2003), Evolution and Palaeobiology
of Pterosaurs. Geological Society of London, Special Publications 217, London: 1-347.
22.^ a b Bennett, S. Christopher (2006). "Juvenile specimens of the pterosaur Germanodactylus
cristatus, with a review of the genus". Journal of Vertebrate Paleontology 26 (4): 872–878.
doi:10.1671/0272-4634(2006)26[872:JSOTPG]2.0.CO;2.

Palaeontology portal
Wikimedia Commons has media related to: Pterodactylus
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Categories: Jurassic pterosaurs | Ctenochasmatoids | Prehistoric reptiles of Europe | Solnhofen

fauna
W000
Mitochondrion

Two mitochondria from mammalian lung tissue displaying their matrix and membranes as shown by
electron microscopy.
Schematic of typical animal cell, showing subcellular components. Organelles:
(1) nucleolus
(2) nuclear membrane
(3) Ribosomes
(4) Vesicle
(5) Rough endoplasmic reticulum (ER)
(6) Golgi body
(7) Cytoskeleton
(8) Smooth ER
(9) Mitochondria
(13) Centrioles within centrosome
In cell biology, a mitochondrion (plural mitochondria) is a membrane-enclosed organelle found in
most eukaryotic cells.[1] These organelles range from 0.5 to 10 micrometers (μm) in diameter. Mitochondria
are sometimes described as "cellular power plants" because they generate most of the cell's supply of
adenosine triphosphate (ATP), used as a source of chemical energy.[2] In addition to supplying cellular
energy, mitochondria are involved in a range of other processes, such as signaling, cellular differentiation,
cell death, as well as the control of the cell cycle and cell growth.[3] Mitochondria have been implicated in
several human diseases, including mitochondrial disorders[4] and cardiac dysfunction,[5] and may play a role
in the aging process. The word mitochondrion comes from the Greek μίτος or mitos, thread + χονδρίον or
chondrion, granule.
Several characteristics make mitochondria unique. The number of mitochondria in a cell varies
widely by organism and tissue type. Many cells have only a single mitochondrion, whereas others can
contain several thousand mitochondria.[6][7] The organelle is composed of compartments that carry out
specialized functions. These compartments or regions include the outer membrane, the intermembrane
space, the inner membrane, and the cristae and matrix. Mitochondrial proteins vary depending on the tissue
and the species. In humans, 615 distinct types of proteins have been identified from cardiac mitochondria,[8]
whereas in Murinae (rats), 940 proteins encoded by distinct genes have been reported.[9] The mitochondrial
proteome is thought to be dynamically regulated.[10] Although most of a cell's DNA is contained in the cell
nucleus, the mitochondrion has its own independent genome. Further, its DNA shows substantial similarity to
bacterial genomes.[11]
Contents
[hide]
• 1 Structure
• 1.1 Outer membrane
• 1.2 Intermembrane space
• 1.3 Inner membrane
• 1.3.1 Cristae
• 1.4 Matrix
• 2 Organization and distribution
• 3 Function
• 3.1 Energy conversion
• 3.1.1 Pyruvate and the citric acid cycle
• 3.1.2 NADH and FADH2: the electron
transport chain
• 3.1.3 Heat production
• 3.2 Storage of calcium ions
• 3.3 Additional functions
• 4 Origin
• 5 Genome
• 6 Replication and inheritance
• 7 Population genetic studies
• 8 Dysfunction and disease
[edit] Structure
A mitochondrion contains outer and inner membranes composed of phospholipid bilayers and
proteins.[6] The two membranes, however, have different properties. Because of this double-membraned
organization, there are five distinct compartments within the mitochondrion. There is the outer mitochondrial
membrane, the intermembrane space (the space between the outer and inner membranes), the inner
mitochondrial membrane, the cristae space (formed by infoldings of the inner membrane), and the matrix
(space within the inner membrane).
[edit] Outer membrane

Main article: Outer mitochondrial membrane
The outer mitochondrial membrane, which encloses the entire organelle, has a protein-to-
phospholipid ratio similar to that of the eukaryotic plasma membrane (about 1:1 by weight). It contains large
numbers of integral proteins called porins. These porins form channels that allow molecules 5000 Daltons or
less in molecular weight to freely diffuse from one side of the membrane to the other.[6] Larger proteins can
enter the mitochondrion if a signaling sequence at their N-terminus binds to a large multisubunit protein
called translocase of the outer membrane, which then actively moves them across the membrane.[12]
Disruption of the outer membrane permits proteins in the intermembrane space to leak into the cytosol,
leading to certain cell death.[13] The mitochondrial outer membrane can associate with the endoplasmic
reticulum (ER) membrane, in a structure called MAM (mitochondria-associated ER-membrane). This is
important in ER-mitochondria calcium signaling and involved in the transfer of lipids between the ER and
mitochondria.[14]
[edit] Intermembrane space

The intermembrane space is the space between the outer membrane and the inner membrane.
Because the outer membrane is freely permeable to small molecules, the concentrations of small molecules
such as ions and sugars in the intermembrane space is the same as the cytosol.[6] However, large proteins
must have a specific signaling sequence to be transported across the outer membrane, so the protein
composition of this space is different from the protein composition of the cytosol. One protein that is localized
to the intermembrane space in this way is cytochrome c.[13]
[edit] Inner membrane

Main article: Inner mitochondrial membrane
The inner mitochondrial membrane contains proteins with five types of functions:[6]
1. Those that perform the redox reactions of oxidative phosphorylation
2. ATP synthase, which generates ATP in the matrix
3. Specific transport proteins that regulate metabolite passage into and out of the matrix
4. Protein import machinery.
5. Mitochondria fusion and fission protein
It contains more than 151 different polypeptides, and has a very high protein-to-phospholipid ratio
(more than 3:1 by weight, which is about 1 protein for 15 phospholipids). The inner membrane is home to
around 1/5 of the total protein in a mitochondrion.[6] In addition, the inner membrane is rich in an unusual
phospholipid, cardiolipin. This phospholipid was originally discovered in cow hearts in 1942, and is usually
characteristic of mitochondrial and bacterial plasma membranes.[15] Cardiolipin contains four fatty acids
rather than two and may help to make the inner membrane impermeable.[6] Unlike the outer membrane, the
inner membrane doesn't contain porins and is highly impermeable to all molecules. Almost all ions and
molecules require special membrane transporters to enter or exit the matrix. Proteins are ferried into the
matrix via the translocase of the inner membrane (TIM) complex or via Oxa1.[12] In addition, there is a
membrane potential across the inner membrane formed by the action of the enzymes of the electron
transport chain.
[edit] Cristae
Cross-sectional image of cristae in rat liver mitochondrion to demonstrate the likely 3D structure and
relationship to the inner membrane.
Main article: Cristae
The inner mitochondrial membrane is compartmentalized into numerous cristae, which expand the
surface area of the inner mitochondrial membrane, enhancing its ability to produce ATP. For typical liver
mitochondria the area of the inner membrane is about five times greater than the outer membrane. This ratio
is variable and mitochondria from cells that have a greater demand for ATP, such as muscle cells, contain
even more cristae. These folds are studded with small round bodies known as F1 particles or oxysomes.
These are not simple random folds but rather invaginations of the inner membrane, which can affect overall
chemiosmotic function.[16]
One recent mathematical modeling study has suggested that the optical properties of the christae in
filamentous mitochondria may affect the generation and propogation of light within the tissue.[17]
[edit] Matrix
Main article: Mitochondrial matrix
The matrix is the space enclosed by the inner membrane. It contains about 2/3 of the total protein in
a mitochondrion.[6] The matrix is important in the production of ATP with the aid of the ATP synthase
contained in the inner membrane. The matrix contains a highly-concentrated mixture of hundreds of
enzymes, special mitochondrial ribosomes, tRNA, and several copies of the mitochondrial DNA genome. Of
the enzymes, the major functions include oxidation of pyruvate and fatty acids, and the citric acid cycle.[6]
Mitochondria have their own genetic material, and the machinery to manufacture their own RNAs and
proteins (see: protein biosynthesis). A published human mitochondrial DNA sequence revealed 16,569 base
pairs encoding 37 total genes: 22 tRNA, 2 rRNA, and 13 peptide genes.[18] The 13 mitochondrial peptides in
humans are integrated into the inner mitochondrial membrane, along with proteins encoded by genes that
reside in the host cell's nucleus.
[edit] Organization and distribution

Mitochondria are found in nearly all eukaryotes. They vary in number and location according to cell
type. A single mitochondrion is often found in unicellular organisms. Conversely, numerous mitochondria are
found in human liver cells, with about 1000–2000 mitochondria per cell making up 1/5th of the cell volume.[6]
The mitochondria can be found nestled between myofibrils of muscle or wrapped around the sperm
flagellum.[6] Often they form a complex 3D branching network inside the cell with the cytoskeleton. The
association with the cytoskeleton determines mitochondrial shape, which can affect the function as well.[19]
Recent evidence suggests vimentin, one of the components of the cytoskeleton, is critical to the association
with the cytoskeleton.[20]
[edit] Function
The most prominent roles of mitochondria are to produce ATP (i.e., phosphorylation of ADP) through
respiration, and to regulate cellular metabolism.[7] The central set of reactions involved in ATP production
are collectively known as the citric acid cycle, or the Krebs Cycle. However, the mitochondrion has many
other functions in addition to the production of ATP.
[edit] Energy conversion

A dominant role for the mitochondria is the production of ATP, as reflected by the large number of
proteins in the inner membrane for this task. This is done by oxidizing the major products of glucose,
pyruvate, and NADH, which are produced in the cytosol.[7] This process of cellular respiration, also known as
aerobic respiration, is dependent on the presence of oxygen. When oxygen is limited, the glycolytic products
will be metabolized by anaerobic respiration, a process that is independent of the mitochondria.[7] The
production of ATP from glucose has an approximately 13-fold higher yield during aerobic respiration
compared to anaerobic respiration.[21] Recently it has been shown that plant mitochondria can produce a
limited amount of ATP without oxygen by using the alternate substrate nitrite.[22]
[edit] Pyruvate and the citric acid cycle

Main articles: Pyruvate decarboxylation and Citric acid cycle
Each pyruvate molecule produced by glycolysis is actively transported across the inner mitochondrial
membrane, and into the matrix where it is oxidized and combined with coenzyme A to form CO2, acetyl-CoA,
and NADH.[7]
The acetyl-CoA is the primary substrate to enter the citric acid cycle, also known as the tricarboxylic
acid (TCA) cycle or Krebs cycle. The enzymes of the citric acid cycle are located in the mitochondrial matrix,
with the exception of succinate dehydrogenase, which is bound to the inner mitochondrial membrane as part
of Complex II.[23] The citric acid cycle oxidizes the acetyl-CoA to carbon dioxide, and, in the process,
produces reduced cofactors (three molecules of NADH and one molecule of FADH2) that are a source of
electrons for the electron transport chain, and a molecule of GTP (that is readily converted to an ATP).[7]
[edit] NADH and FADH2: the electron transport chain

Main articles: Electron transport chain and Oxidative phosphorylation
Diagram of the electron transport chain in the mitonchondrial intermembrane space
The redox energy from NADH and FADH2 is transferred to oxygen (O2) in several steps via the
electron transport chain. These energy-rich molecules are produced within the matrix via the citric acid cycle
but are also produced in the cytoplasm by glycolysis. Reducing equivalents from the cytoplasm can be
imported via the malate-aspartate shuttle system of antiporter proteins or feed into the electron transport
chain using a glycerol phosphate shuttle.[7] Protein complexes in the inner membrane (NADH
dehydrogenase, cytochrome c reductase, and cytochrome c oxidase) perform the transfer and the
incremental release of energy is used to pump protons (H+) into the intermembrane space. This process is
efficient, but a small percentage of electrons may prematurely reduce oxygen, forming reactive oxygen
species such as superoxide.[7] This can cause oxidative stress in the mitochondria and may contribute to the
decline in mitochondrial function associated with the aging process.[24]
As the proton concentration increases in the intermembrane space, a strong electrochemical gradient
is established across the inner membrane. The protons can return to the matrix through the ATP synthase
complex, and their potential energy is used to synthesize ATP from ADP and inorganic phosphate (P i).[7]
This process is called chemiosmosis, and was first described by Peter Mitchell[25][26] who was awarded the
1978 Nobel Prize in Chemistry for his work. Later, part of the 1997 Nobel Prize in Chemistry was awarded to
Paul D. Boyer and John E. Walker for their clarification of the working mechanism of ATP synthase.[27]
[edit] Heat production

Under certain conditions, protons can re-enter the mitochondrial matrix without contributing to ATP
synthesis. This process is known as proton leak or mitochondrial uncoupling and is due to the facilitated
diffusion of protons into the matrix. The process results in the unharnessed potential energy of the proton
electrochemical gradient being released as heat.[7] The process is mediated by a proton channel called
thermogenin, or UCP1.[28] Thermogenin is a 33kDa protein first discovered in 1973.[29] Thermogenin is
primarily found in brown adipose tissue, or brown fat, and is responsible for non-shivering thermogenesis.
Brown adipose tissue is found in mammals, and is at its highest levels in early life and in hibernating animals.
In humans, brown adipose tissue is present at birth and decreases with age.[28]
[edit] Storage of calcium ions
Mitochondria (M) within a chondrocyte stained for calcium as shown by electron microscopy.
The concentrations of free calcium in the cell can regulate an array of reactions and is important for
signal transduction in the cell. Mitochondria can transiently store calcium, a contributing process for the cell's
homeostasis of calcium.[30] In fact, their ability to rapidly take in calcium for later release makes them very
good "cytosolic buffers" for calcium.[31][32][33] The endoplasmic reticulum (ER) is the most significant
storage site of calcium, and there is a significant interplay between the mitochondrion and ER with regard to
calcium.[34] The calcium is taken up into the matrix by a calcium uniporter on the inner mitochondrial
membrane.[35] It is primarily driven by the mitochondrial membrane potential.[30] Release of this calcium
back into the cell's interior can occur via a sodium-calcium exchange protein or via "calcium-induced-
calcium-release" pathways.[35] This can initiate calcium spikes or calcium waves with large changes in the
membrane potential. These can activate a series of second messenger system proteins that can coordinate
processes such as neurotransmitter release in nerve cells and release of hormones in endocrine cells.
[edit] Additional functions

Mitochondria play a central role in many other metabolic tasks, such as:
• Regulation of the membrane potential[7]
• Apoptosis-programmed cell death[36]
• Calcium signaling (including calcium-evoked apoptosis)[37]
• Cellular proliferation regulation[38]
• Regulation of cellular metabolism[38]
• Certain heme synthesis reactions[39] (see also: porphyrin)
• Steroid synthesis.[31]
Some mitochondrial functions are performed only in specific types of cells. For example,
mitochondria in liver cells contain enzymes that allow them to detoxify ammonia, a waste product of protein
metabolism. A mutation in the genes regulating any of these functions can result in mitochondrial diseases.
[edit] Origin
Main article: Endosymbiotic theory
Mitochondria have many features in common with prokaryotes. As a result, they are believed to be
originally derived from endosymbiotic prokaryotes.
A mitochondrion contains DNA, which is organized as several copies of a single, circular
chromosome. This mitochondrial chromosome contains genes for redox proteins such as those of the
respiratory chain. The CoRR hypothesis proposes that this co-location is required for redox regulation. The
mitochondrial genome codes for some RNAs of ribosomes, and the twenty-two tRNAs necessary for the
translation of messenger RNAs into protein. The circular structure is also found in prokaryotes, and the
similarity is extended by the fact that mitochondrial DNA is organized with a variant genetic code similar to
that of Proteobacteria.[40] This suggests that their ancestor, the so-called proto-mitochondrion, was a
member of the Proteobacteria.[40] In particular, the proto-mitochondrion was probably closely related to the
rickettsia.[41] However, the exact relationship of the ancestor of mitochondria to the alpha-proteobacteria and
whether the mitochondria was formed at the same time or after the nucleus, remains controversial.[42]
The ribosomes coded for by the mitochondrial DNA are similar to those from bacteria in size and
structure.[43] They closely resemble the bacterial 70S ribosome and not the 80S cytoplasmic ribosomes,
which are coded for by nuclear DNA.
The endosymbiotic relationship of mitochondria with their host cells was popularized by Lynn
Margulis.[44] The endosymbiotic hypothesis suggests that mitochondria descended from bacteria that
somehow survived endocytosis by another cell, and became incorporated into the cytoplasm. The ability of
these bacteria to conduct respiration in host cells that had relied on glycolysis and fermentation would have
provided a considerable evolutionary advantage. In a similar manner, host cells with symbiotic bacteria
capable of photosynthesis would have had an advantage. The incorporation of symbiotes would have
increased the number of environments in which the cells could survive. This symbiotic relationship probably
developed 1.7[45]-2[46] billion years ago.
A few groups of unicellular eukaryotes lack mitochondria: the microsporidians, metamonads, and
archamoebae.[47] These groups appear as the most primitive eukaryotes on phylogenetic trees constructed
using rRNA information, which once suggested that they appeared before the origin of mitochondria.
However, this is now known to be an artifact of long-branch attraction—they are derived groups and retain
genes or organelles derived from mitochondria (e.g., mitosomes and hydrogenosomes).[1]
[edit] Genome
Main article: Mitochondrial DNA
The human mitochondrial genome is a circular DNA molecule of about 16 kilobases.[48] It encodes
37 genes: 13 for subunits of respiratory complexes I, III, IV and V, 22 for mitochondrial tRNA (for the 20
standard amino acids, plus an extra gene for leucine and serine), and 2 for rRNA.[48] One mitochondrion can
contain two to ten copies of its DNA.[49]
As in prokaryotes, there is a very high proportion of coding DNA and an absence of repeats.
Mitochondrial genes are transcribed as multigenic transcripts, which are cleaved and polyadenylated to yield
mature mRNAs. Not all proteins necessary for mitochondrial function are encoded by the mitochondrial
genome; most are coded by genes in the cell nucleus and the corresponding proteins are imported into the
mitochondrion.[50] The exact number of genes encoded by the nucleus and the mitochondrial genome differs
between species. In general, mitochondrial genomes are circular, although exceptions have been reported.
[51] In general, mitochondrial DNA lacks introns, as is the case in the human mitochondrial genome;[50]
however, introns have been observed in some eukaryotic mitochondrial DNA,[52] such as that of yeast[53]
and protists,[54] including Dictyostelium discoideum.[55]
In animals the mitochondrial genome is typically a single circular chromosome that is approximately
16-kb long and has 37 genes. The genes while highly conserved may vary in location. Curiously this pattern
is not found in the human body louse (Pediculus humanus). Instead this mitochondrial genome is arranged in
18 minicircular chromosomes each of which is 3–4 kb long and has one to three genes.[56] This pattern is
also found in other sucking lice but not in chewing lice. Recombination has been shown to occur between the
minichromosomes. The reason for this difference is not known.
While slight variations on the standard code had been predicted earlier,[57] none was discovered
until 1979, when researchers studying human mitochondrial genes determined that they used an alternative
code.[58] Many slight variants have been discovered since,[59] including various alternative mitochondrial
codes.[60] Further, the AUA, AUC, and AUU codons are all allowable start codons.
Exceptions to the universal genetic code (UGC) in mitochondria[6]
Organism Codon Standard Novel
AGA, AGG Arginine Stop codon
Mammalian AUA Isoleucine Methionine
UGA Stop codon Tryptophan
AGA, AGG Arginine Serine
Invertebrates AUA Isoleucine Methionine
Yeast AUA Isoleucine Methionine

CUA Leucine Threonine
Some of these differences should be regarded as pseudo-changes in the genetic code due to the
phenomenon of RNA editing, which is common in mitochondria. In higher plants, it was thought that CGG
encoded for tryptophan and not arginine; however, the codon in the processed RNA was discovered to be the
UGG codon, consistent with the universal genetic code for tryptophan.[61] Of note, the arthropod
mitochondrial genetic code has undergone parallel evolution within a phylum, with some organisms uniquely
translating AGG to lysine.[62]
Mitochondrial genomes have far fewer genes than the bacteria from which they are thought to be
descended. Although some have been lost altogether, many have been transferred to the nucleus, such as
the respiratory complex II protein subunits.[48] This is thought to be relatively common over evolutionary
time. A few organisms, such as the Cryptosporidium, actually have mitochondria that lack any DNA,
presumably because all their genes have been lost or transferred.[63] In Cryptosporidium, the mitochondria
have an altered ATP generation system that renders the parasite resistant to many classical mitochondrial
inhibitors such as cyanide, azide, and atovaquone.[63]
[edit] Replication and inheritance

See also: mitochondrial genome
Mitochondria divide by binary fission similar to bacterial cell division; unlike bacteria, however,
mitochondria can also fuse with other mitochondria.[48][64] The regulation of this division differs between
eukaryotes. In many single-celled eukaryotes, their growth and division is linked to the cell cycle. For
example, a single mitochondrion may divide synchronously with the nucleus. This division and segregation
process must be tightly controlled so that each daughter cell receives at least one mitochondrion. In other
eukaryotes (in mammals for example), mitochondria may replicate their DNA and divide mainly in response
to the energy needs of the cell, rather than in phase with the cell cycle. When the energy needs of a cell are
high, mitochondria grow and divide. When the energy use is low, mitochondria are destroyed or become
inactive. In such examples, and in contrast to the situation in many single celled eukaryotes, mitochondria
are apparently randomly distributed to the daughter cells during the division of the cytoplasm.
An individual's mitochondrial genes are not inherited by the same mechanism as nuclear genes. At
fertilization of an egg cell by a sperm, the egg nucleus and sperm nucleus each contribute equally to the
genetic makeup of the zygote nucleus. In contrast, the mitochondria, and therefore the mitochondrial DNA,
usually comes from the egg only. The sperm's mitochondria enter the egg but do not contribute genetic
information to the embryo.[65] Instead, paternal mitochondria are marked with ubiquitin to select them for
later destruction inside the embryo.[66] The egg cell contains relatively few mitochondria, but it is these
mitochondria that survive and divide to populate the cells of the adult organism. Mitochondria are, therefore,
in most cases inherited down the female line, known as maternal inheritance. This mode is seen in most
organisms including all animals. However, mitochondria in some species can sometimes be inherited
paternally. This is the norm among certain coniferous plants, although not in pine trees and yew trees.[67] It
has been suggested that it occurs at a very low level in humans.[68]
Uniparental inheritance leads to little opportunity for genetic recombination between different
lineages of mitochondria, although a single mitochondrion can contain 2–10 copies of its DNA.[49] For this
reason, mitochondrial DNA usually is thought to reproduce by binary fission. What recombination does take
place maintains genetic integrity rather than maintaining diversity. However, there are studies showing
evidence of recombination in mitochondrial DNA. It is clear that the enzymes necessary for recombination
are present in mammalian cells.[69] Further, evidence suggests that animal mitochondria can undergo
recombination.[70] The data are a bit more controversial in humans, although indirect evidence of
recombination exists.[71][72] If recombination does not occur, the whole mitochondrial DNA sequence
represents a single haplotype, which makes it useful for studying the evolutionary history of populations.
[edit] Population genetic studies

Main article: Human mitochondrial genetics
The near-absence of genetic recombination in mitochondrial DNA makes it a useful source of
information for scientists involved in population genetics and evolutionary biology.[73] Because all the
mitochondrial DNA is inherited as a single unit, or haplotype, the relationships between mitochondrial DNA
from different individuals can be represented as a gene tree. Patterns in these gene trees can be used to
infer the evolutionary history of populations. The classic example of this is in human evolutionary genetics,
where the molecular clock can be used to provide a recent date for mitochondrial Eve.[74][75] This is often
interpreted as strong support for a recent modern human expansion out of Africa.[76] Another human
example is the sequencing of mitochondrial DNA from Neanderthal bones. The relatively large evolutionary
distance between the mitochondrial DNA sequences of Neanderthals and living humans has been
interpreted as evidence for lack of interbreeding between Neanderthals and anatomically-modern humans.
[77]
However, mitochondrial DNA reflects the history of only females in a population and so may not
represent the history of the population as a whole. This can be partially overcome by the use of paternal
genetic sequences, such as the non-recombining region of the Y-chromosome.[76] In a broader sense, only
studies that also include nuclear DNA can provide a comprehensive evolutionary history of a population.[78]
[edit] Dysfunction and disease
[edit] Mitochondrial diseases
Main article: Mitochondrial disease
With their central place in cell metabolism, damage — and subsequent dysfunction — in mitochondria
is an important factor in a wide range of human diseases. Mitochondrial disorders often present as
neurological disorders, but can manifest as myopathy, diabetes, multiple endocrinopathy, or a variety of other
systemic manifestations.[79] Diseases caused by mutation in the mtDNA include Kearns-Sayre syndrome,
MELAS syndrome and Leber's hereditary optic neuropathy.[80] In the vast majority of cases, these diseases
are transmitted by a female to her children, as the zygote derives its mitochondria and hence its mtDNA from
the ovum. Diseases such as Kearns-Sayre syndrome, Pearson's syndrome, and progressive external
ophthalmoplegia are thought to be due to large-scale mtDNA rearrangements, whereas other diseases such
as MELAS syndrome, Leber's hereditary optic neuropathy, myoclonic epilepsy with ragged red fibers
(MERRF), and others are due to point mutations in mtDNA.[79]
In other diseases, defects in nuclear genes lead to dysfunction of mitochondrial proteins. This is the
case in Friedreich's ataxia, hereditary spastic paraplegia, and Wilson's disease.[81] These diseases are
inherited in a dominance relationship, as applies to most other genetic diseases. A variety of disorders can
be caused by nuclear mutations of oxidative phosphorylation enzymes, such as coenzyme Q10 deficiency
and Barth syndrome.[79] Environmental influences may interact with hereditary predispositions and cause
mitochondrial disease. For example, there may be a link between pesticide exposure and the later onset of
Parkinson's disease.[82][83]
Other pathologies with etiology involving mitochondrial dysfunction include schizophrenia, bipolar
disorder, dementia, Alzheimer's disease, Parkinson's disease, epilepsy, stroke, cardiovascular disease,
retinitis pigmentosa, and diabetes mellitus.[84][85] A common thread thought to link these seemingly-
unrelated conditions is cellular damage causing oxidative stress. How exactly mitochondrial dysfunction fits
into the etiology of these pathologies is yet to be elucidated.[ citation needed]
[edit] Possible relationships to aging

Given the role of mitochondria as the cell's powerhouse, there may be some leakage of the high-
energy electrons in the respiratory chain to form reactive oxygen species. This can result in significant
oxidative stress in the mitochondria with high mutation rates of mitochondrial DNA.[86] A vicious cycle is
thought to occur, as oxidative stress leads to mitochondrial DNA mutations, which can lead to enzymatic
abnormalities and further oxidative stress. A number of changes occur to mitochondria during the aging
process.[87] Tissues from elderly patients show a decrease in enzymatic activity of the proteins of the
respiratory chain.[88] Large deletions in the mitochondrial genome can lead to high levels of oxidative stress
and neuronal death in Parkinson's disease.[89] Hypothesized links between aging and oxidative stress are
not new and were proposed over 50 years ago;[90] however, there is much debate over whether
mitochondrial changes are causes of aging or merely characteristics of aging. One notable study in mice
demonstrated shortened lifespan but no increase in reactive oxygen species despite increasing mitochondrial
DNA mutations, suggesting that mitochondrial DNA mutations can cause lifespan shortening by other
mechanisms.[91] As a result, the exact relationships between mitochondria, oxidative stress, and aging have
not yet been settled.
[edit] See also
Wikimedia Commons has media related to: Mitochondrion
• Anti-mitochondrial antibodies
• Bioenergetics
• CoRR Hypothesis
• Human mitochondrial genetics
• Mitochondrial permeability transition pore
• Nebenkern
• Oncocyte
• Oncocytoma
• Paternal mtDNA transmission
• Plastid
• Submitochondrial particle
• TIM/TOM Complex
• Chloroplast
• Hydrogenosome
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doi:10.1074/jbc.271.44.27536. PMID 8910339.
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387 (6630): 247. doi:10.1038/387247a0. PMID 9153388.
71.^ Eyre-Walker A, Smith NH, Maynard Smith J (1999-03-07). "How clonal are human
mitochondria?". Proc. Royal Soc. Biol. Sci. (Series B) 266 (1418): 477–483.
doi:10.1098/rspb.1999.0662. PMID 10189711.
72.^ Awadalla P, Eyre-Walker A, Maynard Smith J (24 December 1999). "Linkage
Disequilibrium and Recombination in Hominid Mitochondrial DNA". Science. 286 (5449): 2524–2525.
doi:10.1126/science.286.5449.2524. PMID 10617471.
73.^ Castro JA, Picornell A, Ramon M (1998). "Mitochondrial DNA: a tool for populational
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• Mitochondria Atlas at University of Mainz
• Mitochondria Research Portal at mitochondrial.net
• Mitochondria: Architecture dictates function at cytochemistry.net
• Mitochondria links at University of Alabama
• MIP Mitochondrial Physiology Society
• 3D structures of proteins from inner mitochondrial membrane at University of Michigan
• 3D structures of proteins associated with outer mitochondrial membrane at University of
Michigan
• Mitochondrial Protein Partnership at University of Wisconsin
• Mitochondrion - Cell Centered Database
• Mitochondrion Reconstructed by Electron Tomography at San Diego State University
• Video Clip of Rat-liver Mitochondrion from Cryo-electron Tomography

Endomembrane
Microfilaments · Intermediate filaments · Microtubules · Prokaryotic

cytoskeleton
Cytoskeleton
Myofibril

Other internal
External Undulipodium: Cilium/Flagellum · Axoneme · Radial spoke

[hide]v · d · eMitochondrial proteins
fatty acid degradation (Carnitine palmitoyltransferase I, Long fatty acyl

CoA synthetase)
Outer membrane
tryptophan metabolism (Kynureninase)
monoamine neurotransmitter metabolism (Monoamine oxidase)
Intermembrane
space Adenylate kinase - Creatine kinase
Inner membrane oxidative phosphorylation (Coenzyme Q - cytochrome c reductase,

Cytochrome c, NADH dehydrogenase, Succinate dehydrogenase)
pyrimidine metabolism (Dihydroorotate dehydrogenase)
mitochondrial shuttle (Malate-aspartate shuttle, Glycerol phosphate
shuttle)
other (Glutamate aspartate transporter, Glycerol-3-phosphate
dehydrogenase, ATP synthase, Carnitine palmitoyltransferase II, Uncoupling
protein)
citric acid cycle (Citrate synthase, Aconitase, Isocitrate dehydrogenase,

Oxoglutarate dehydrogenase, Succinyl coenzyme A synthetase, Fumarase,
Malate dehydrogenase)
anaplerotic reactions (Aspartate transaminase, Glutamate
Matrix dehydrogenase, Pyruvate dehydrogenase complex)
urea cycle (Carbamoyl phosphate synthetase I, Ornithine
transcarbamylase, N-Acetylglutamate synthase)
alcohol metabolism (ALDH2)
PMPCB
steroidogenesis (Cholesterol side-chain cleavage enzyme, Steroid 11-

Other/to be beta-hydroxylase, Aldosterone synthase), Frataxin
sorted
Mitochondrial membrane transport protein (Mitochondrial permeability
transition pore, Mitochondrial carrier)
Mitochondrial Complex I (MT-ND1, MT-ND2, MT-ND3, MT-ND4, MT-ND4L, MT-ND5,

DNA MT-ND6) - Complex III (MT-CYB) - Complex IV (MT-CO1, MT-CO2, MT-CO3)
ATP synthase (MT-ATP6, MT-ATP8)
tRNA (MT-TA, MT-TC, MT-TD, MT-TE, MT-TF, MT-TG, MT-TH, MT-TI,
MT-TK, MT-TL1, MT-TL2, MT-TM, MT-TN, MT-TP, MT-TQ, MT-TR, MT-TS1,
MT-TS2, MT-TT, MT-TV, MT-TW, MT-TY)
see also mitochondrial diseases

This article incorporates public domain material from the NCBI document "Science Primer".
Retrieved from "http://en.wikipedia.org/wiki/Mitochondrion"
Categories: Cellular respiration | Organelles

W000
W000
Taxonomy of the Gastropoda

(Bouchet & Rocroi, 2005)

The taxonomy of the Gastropoda as it was revised by Philippe Bouchet and Jean-Pierre Rocroi in
2005 is currently (2011) the most up-to-date overall system for classifying gastropod mollusks. Gastropods
are a taxonomic class of animals which consists of snails and slugs of every kind, from the land, from
freshwater, and from saltwater. Systems of classification such as this one are primarily of value to
malacologists (people who study mollusks) and other biologists. Biological classification schemes are not
merely a convenience, they are an attempt to show the actual phylogeny (the evolutionary relatedness)
within a group of organisms. Thus a taxonomy such as this one can be seen as an attempt to elucidate part
of the tree of life, a phylogenetic tree.
The Bouchet & Rocroi 2005 system of gastropod taxonomy was laid out in a book-length paper
entitled "Classification and Nomenclator of Gastropod Families", which was published in the journal
Malacologia[1] and which was written in collaboration with J. Frýda, B. Hausdorf, W. Ponder, Á. Valdés and
A. Warén. This system supersedes the system of Ponder and Lindberg from 1997. Subsequent revisions by
other authors have been made since the publication of this paper.
The taxonomy set out by Bouchet & Rocroi is considered to be one step closer to representing the
evolutionary history of the phylum Mollusca. This classification system is a hybrid of the pre-existing, more
traditional Linnaean taxonomy along with some more recent far-reaching revisions which are based on
molecular work and use clades as taxa, (see cladistics). In the past, the taxonomy of gastropods was largely
based on the morphological characters of the taxa, such as shell characteristics and the internal anatomy,
including the structure of the radula and details of the reproductive system. Recent advances are based more
on the molecular characteristics of the DNA and RNA. This has made the newer taxa and their hierarchy
controversial, and the debate about these issues is not likely to end soon.
This proposed classification has tried to integrate these recent advances in understanding by using
unranked clades for taxa below the traditional rank of class (class Gastropoda) but above the rank of
superfamily (replacing the ranks subclass, superorder, order, and suborder), while still using the traditional
Linnaean approach for superfamilies and all taxa below the rank of superfamily (i.e., family, subfamily, tribe,
genus, subgenus, and species. The clades have been given names which are similar to, or in some cases
the same as, traditional Linnaean names for taxa. Whenever monophyly (a single lineage) has not yet been
tested and confirmed, or where a traditional taxon of gastropods has now been discovered to be paraphyletic
or polyphyletic (contains more than one lineage), the term "group" or "informal group" has been used.
In this taxonomy 611 valid families are recognized. Of these, 202 families are exclusively fossil, and
this is indicated here with a dagger †. The superfamily names are standardized so that they all end in the
suffix "-oidea"; there are no longer any that end "-acea". The classification of families into subfamilies
however is often not well resolved, and should be regarded as the best possible hypothesis.
The publication Bouchet & Rocroi (2005)[1] also includes a nomenclator of about 2400 suprageneric
taxa (taxa above the level of genus) of gastropods, from the subtribe to the superfamily. A full bibliographic
reference is provided for each taxon, giving the name of the authority, the original publication, the date of that
publication, the type genus for the taxon, its nomenclatural status, and its validity under the rules of the ICZN.
Since the publication of this taxonomic system in 2005, various changes have been published by
other authors, see Changes in the taxonomy of gastropods since 2005.
Contents
[hide]
• 1 Main clades, groups and informal groups
• 2 Taxonomy
• 2.1 Paleozoic molluscs of uncertain systematic
position
• 2.1.1 Uncertain position (Gastropoda or
Monoplacophora)
• 2.1.2 With isostrophically coiled shells of
uncertain position (Gastropoda or Monoplacophora)
• 2.1.3 With anisostrophically coiled shells of
uncertain position (Gastropoda?)
• 2.2 Basal taxa that are certainly Gastropoda
• 2.3 Clade Patellogastropoda
• 2.4 Clade Vetigastropoda
• 2.5 Clade Cocculiniformia
• 2.6 Clade Neritimorpha (= Neritopsina)
• 2.6.1 Clade Cyrtoneritimorpha
• 2.6.2 Clade Cycloneritimorpha
• 2.7 Clade Caenogastropoda
• 2.8 Informal group Architaenioglossa
• 2.9 Clade Sorbeoconcha
[edit] Main clades, groups and informal groups
This information is displayed in the form of a cladogram (an evolutionary tree of descent.) It is worth
bearing in mind however that this taxonomy is provisional: many of the taxa are still only known as "groups"
or "informal groups", and therefore are likely to turn out to be polyphyletic (in other words, they are likely to
contain more than one lineage, and if so, they will be split in the future.)
Neritimorpha
† Paleozoic Neritimorpha of uncertain systematic
position
† Cyrtoneritimorpha
Cycloneritimorpha
Caenogastropoda
Caenogastropoda of uncertain systematic
position
Architaenioglossa
Sorbeoconcha
Hypsogastropoda
Littorinimorpha
Ptenoglossa
Neogastropoda
Heterobranchia
Opisthobranchia
C
T
Nudipleura
Pleurobranchomorpha
Nudibranchia
Euctenidiacea
Dexiarchia
Pseudoeuctenidiacea
Cladobranchia
Euarminida
Dendronotida
Aeolidida
Pulmonata
Basommatophora
Eupulmonata
Systellommatophora
Stylommatophora
Elasmognatha
Orthurethra
Sigmurethra
The diagram above was based on the following information. The list format used below makes clear which
taxa are groups and informal groups rather than clades:
• Paleozoic molluscs of uncertain systematic position
• Clade Patellogastropoda
• Clade Vetigastropoda
• Clade Cocculiniformia
• Clade Neritimorpha
• Paleozoic Neritimorpha of uncertain systematic position
• Clade †Cyrtoneritimorpha
• Clade Cycloneritimorpha
• Clade Caenogastropoda
• Caenogastropoda of uncertain systematic position
• Informal group Architaenioglossa
• Clade Sorbeoconcha
• Clade Hypsogastropoda
• Clade Littorinimorpha
• Informal group Ptenoglossa
• Clade Neogastropoda
• Clade Heterobranchia
• Informal group Lower Heterobranchia
• Informal group Opisthobranchia
• Clade Cephalaspidea
• Clade Thecosomata
• Clade Gymnosomata
• Clade Aplysiomorpha
• Group Acochlidiacea
• Clade Sacoglossa
• Group Cylindrobullida
• Clade Umbraculida
• Clade Nudipleura
• Clade Pleurobranchomorpha
• Clade Nudibranchia
• Clade Euctenidiacea
• Clade Dexiarchia
• Clade Pseudoeuctenidiacea
• Clade Cladobranchia
• Clade Euarminida
• Clade Dendronotida
• Clade Aeolidida
• Informal group Pulmonata
• Informal group Basommatophora
• Clade Eupulmonata
• Clade Systellommatophora
• Clade Stylommatophora
• Clade Elasmognatha
• Clade Orthurethra
• Informal group Sigmurethra
[edit] Taxonomy
In the following more detailed list, indentation is used only for the ranks of superfamily and family.
The clade names are not indented, but their hierarchy is indicated by the size of the font used in their names.
A clearer sense of their actual hierarchy can be drawn from the list immediately above this one.
[edit] Paleozoic molluscs of uncertain systematic position

(existing as fossils only)
[edit] Uncertain position (Gastropoda or Monoplacophora)

• unassigned to superfamily
• † Khairkhaniidae
• † Ladamarekiidae
• † Metoptomatidae
• † Patelliconidae
• † Protoconchoididae
• Archinacelloidea
• † Archinacellidae
• † Archaeopragidae
• Pelagielloidea
• † Pelagiellidae
• † Aldanellidae
• Scenelloidea
• † Scenellidae
• † Coreospiridae
• † Igarkiellidae
• Yochelcionelloidea
• † Yochelcionellidae
• † Stenothecidae
• † Trenellidae
[edit] With isostrophically coiled shells of uncertain position (Gastropoda or

Monoplacophora)
• Bellerophontoidea
• † Bellerophontidae
• † Bucanellidae
• † Bucaniidae
• † Euphemitidae
• † Pterothecidae
• † Sinuitidae
• † Tremanotidae
• † Tropidodiscidae
[edit] With anisostrophically coiled shells of uncertain position (Gastropoda?)

• Euomphaloidea
• † Euomphalidae
• † Helicotomidae
• † Lesueurillidae
• † Omphalocirridae
• † Omphalotrochidae
• Macluritoidea
• † Macluritidae
[edit] Basal taxa that are certainly Gastropoda

(existing as fossils only)
• unasigned to superfamily
• † Anomphalidae
• † Codonocheilidae
• † Crassimarginatidae
• † Holopeidae
• † Isospiridae
• † Opisthonematidae
• † Paraturbinidae
• † Planitrochidae
• † Pragoserpulinidae
• † Pseudophoridae
• † Raphistomatidae
• † Rhytidopilidae
• † Scoliostomatidae
• † Sinuopeidae
• Clisospiroidea
• † Clisospiridae
• † Onychochilidae
• Loxonematoidea
• † Loxonematidae
• † Palaeozygopleuridae
• Ophiletoidea
• † Ophiletidae
• Straparollinoidea
• † Straparollinidae
• Trochonematoidea
• † Trochonematidae
• † Lophospiridae
[edit] Clade Patellogastropoda

• Superfamily Patelloidea
• Family Patellidae
• Superfamily Nacelloidea
• Family Nacellidae
• Superfamily Lottioidea
• Family Lottiidae
• Family Acmaeidae
• Family Lepetidae
• Superfamily Neolepetopsoidea
• Family Neolepetopsidae
• † Family Daminilidae
• † Family Lepetopsidae
[edit] Clade Vetigastropoda

• Not assigned to a superfamily:
• Family Ataphridae
• Family Pendromidae
• † Family Schizogoniidae
• Superfamily †Amberleyoidea
• † Family Amberleyidae
• † Family Nododelphinulidae
• Superfamily †Eotomarioidea
• † Family Eotomariidae
• † Family Gosseletinidae
• † Family Luciellidae
• † Family Phanerotrematidae
• Superfamily Fissurelloidea
• Family Fissurellidae
• Superfamily Haliotoidea
• Family Haliotidae
• † Family Temnotropidae
• Superfamily Lepetelloidea
• Family Lepetellidae
• Family Addisoniidae
• Family Bathyphytophilidae
• Family Caymanabyssiidae
• Family Cocculinellidae
• Family Osteopeltidae
• Family Pseudococculinidae
• Family Pyropeltidae
• Superfamily Lepetodriloidea
• Family Lepetodrilidae
• Family Clypeosectidae
• Family Sutilizonidae
• Superfamily †Murchisonioidea
• † Family Murchisoniidae
• † Family Cheeneetnukiidae
• † Family Hormotomidae
• Superfamily Neomphaloidea
• Family Neomphalidae
• Family Melanodrymiidae
• Family Peltospiridae
• Superfamily Pleurotomarioidea
• Family Pleurotomariidae
• † Family Catantostomatidae
• † Family Kittlidiscidae
• † Family Phymatopleuridae
• † Family Polytremariidae
• † Family Portlockiellidae
• † Family Rhaphischismatidae
• † Family Trochotomidae
• † Family Zygitidae
• Superfamily †Porcellioidea
• † Family Porcelliidae
• † Family Cirridae
• † Family Discohelicidae
• † Family Pavlodiscidae
• Superfamily Scissurelloidea
• Family Scissurellidae
• Family Anatomidae
• Superfamily Seguenzioidea
• Family Seguenziidae
• Family Chilodontidae
• † Family Eucyclidae
• † Family Laubellidae
• Superfamily Trochoidea
• Family Trochidae
• Family Calliostomatidae
• † Family Elasmonematidae
• † Family Eucochlidae
• † Family Microdomatidae
• † Family Proconulidae
• Family Solariellidae
• † Family Tychobraheidae
• † Family Velainellidae
• Superfamily Turbinoidea
• Family Turbinidae
• Family Liotiidae
• Family Phasianellidae
[edit] Clade Cocculiniformia

• Superfamily Cocculinoidea
• Family Cocculinidae
• Family Bathysciadiidae
[edit] Clade Neritimorpha (= Neritopsina)
Contains the Palaeozoic Neritomorpha of uncertain position and the clades Cyrtoneritimorpha and
Cycloneritimorpha
• Unassigned to superfamily
• † Family Craspedostomatidae
• † Family Pragoscutulidae
• Superfamily Nerrhenoidea
• † Family Nerrhenidae
• Superfamily Oriostomatoidea
• † Family Oriostomatidae
• † Family Tubinidae
• Superfamily Palaeotrochoidea
• † Family Palaeotrochidae
• Superfamily Platyceratoidea
• † Family Platyceratidae
[edit] Clade Cyrtoneritimorpha

• † Family Orthonychiidae
• † Family Vltaviellidae
[edit] Clade Cycloneritimorpha
• Superfamily Helicinoidea
• Family Helicinidae
• † Family Dawsonellidae
• † Family Deaniridae
• Family Neritiliidae
• Family Proserpinellidae
• Family Proserpinidae
• Superfamily Hydrocenoidea
• Family Hydrocenidae
• Superfamily Neritoidea
• Family Neritidae
• Family Phenacolepadidae
• † Family Pileolidae
• Superfamily Neritopsoidea
• Family Neritopsidae
• † Family Cortinellidae
• † Family Delphinulopsidae
• † Family Plagiothyridae
• † Family Pseudorthonychiidae
• Family Titiscaniidae
• Superfamily Symmetrocapuloidea
• † Family Symmetrocapulidae
[edit] Clade Caenogastropoda

Contains the Caenogastropoda of uncertain systematic position, the informal group
Architaenioglossa and the clades Sorbeoconcha and Hypsogastropoda
• Caenogastropoda of uncertain systematic position
• † Family Plicatusidae
• † Family Spanionematidae
• † Family Spirostylidae
• Superfamily Acteoninoidea
• † Family Acteoninidae
• † Family Anozygidae
• † Family Soleniscidae
• Superfamily Dendropupoidea
• † Family Dendropupidae
• † Family Anthracopupidae
• Superfamily Paleostyloidea
• † Family Palaeostylidae
• † Family Goniasmatidae
• † Family Pithodeidae
• Superfamily Peruneloidea
• † Family Perunelidae
• † Family Chuchlinidae
• † Family Imoglobidae
• † Family Sphaerodomidae
• Superfamily Pseudomelanioidea
• † Family Pseudomelaniidae
• † Family Trajanellidae
• Superfamily Subulitoidea
• † Family Subulitidae
• † Family Ischnoptygmatidae
• grade Zygopleuroid group - see Changes since 2005
• † Family Zygopleuridae
• Family Abyssochrysidae
• † Family Polygyrinidae
• † Family Protoculidae
• Family Provannidae
• † Family Pseudozygopleuridae
[edit] Informal group Architaenioglossa

• Superfamily Ampullarioidea
• Family Ampullariidae
• † Family Naricopsinidae
• Superfamily Cyclophoroidea
• Family Cyclophoridae
• Family Aciculidae
• Family Craspedopomatidae
• Family Diplommatinidae
• † Family Ferussinidae
• Family Maizaniidae
• Family Megalomastomatidae
• Family Neocyclotidae
• Family Pupinidae
• Superfamily Viviparoidea
• Family Viviparidae
• † Family Pliopholygidae
[edit] Clade Sorbeoconcha

• Not allocated to superfamily
• † Family Acanthonematidae
• † Family Canterburyellidae
• † Family Prisciphoridae
• Superfamily Cerithioidea
• Family Cerithiidae
• Family Batillariidae
• † Family Brachytrematidae
• † Family Cassiopidae
• Family Dialidae
• Family Diastomatidae
• † Family Eustomatidae
• † Family Ladinulidae
• † Family Lanascalidae
• Family Litiopidae
• † Family Maoraxidae
• Family Melanopsidae
• † Family Metacerithiidae
• Family Modulidae
• Family Pachychilidae
• Family Paludomidae
• Family Planaxidae
• Family Pleuroceridae
• † Family Popenellidae
• Family Potamididae
• † Family Procerithiidae - If the genus Argyropeza is placed in the Procerithiidae, then
this family is not all fossil.
• † Family Prostyliferidae
• † Family Propupaspiridae
• Family Scaliolidae
• Family Siliquariidae
• † Family Terebrellidae
• Family Thiaridae
• Family Turritellidae
• Superfamily Campaniloidea
• Family Campanilidae
• Family Ampullinidae
• Family Plesiotrochidae
• † Family Trypanaxidae
[edit] Clade Hypsogastropoda

Contains the clades Littorinimorpha, Neogastropoda and the informal group Ptenoglossa.
• Not allocated to a superfamily
• † Family Coelostylinidae
• † Family Maturifusidae
• † Family Pommerozygiidae
• † Family Settsassiidae
[edit] Clade Littorinimorpha

• Superfamily Calyptraeoidea
• Family Calyptraeidae
• Superfamily Capuloidea
• Family Capulidae
• Superfamily Cingulopsoidea
• Family Cingulopsidae
• Family Eatoniellidae
• Family Rastodentidae
• Superfamliy Cypraeoidea
• Family Cypraeidae
• Family Ovulidae
• Superfamily Ficoidea
• Family Ficidae
• Superfamily Littorinoidea
• Family Littorinidae
• † Family Bohaispiridae
• Family Pickworthiidae
• Family Pomatiidae
• † Family Purpurinidae
• Family Skeneopsidae
• † Family Tripartellidae
• Family Zerotulidae
• Superfamily Naticoidea
• Family Naticidae
• Superfamily Pterotracheoidea
• Family Pterotracheidae
• Family Atlantidae
• † Family Bellerophinidae
• Family Carinariidae
• Superfamily Rissooidea
• Family Rissoidae
• Family Amnicolidae
• Family Anabathridae
• Family Assimineidae
• Family Barleeiidae
• Family Bithyniidae
• Family Caecidae
• Family Calopiidae
• Family Cochliopidae
• Family Elachisinidae
• Family Emblandidae
• Family Epigridae
• Family Falsicingulidae
• Family Helicostoidae
• Family Hydrobiidae
• Family Hydrococcidae
• Family Iravadiidae
• Family Lithoglyphidae
• † Family Mesocochliopidae
• Family Moitessieriidae
• † Family Palaeorissoinidae
• Family Pomatiopsidae
• Family Stenothyridae
• Family Tornidae
• Family Truncatellidae
• Superfamily Stromboidea
• Family Strombidae
• Family Aporrhaidae
• † Family Colombellinidae
• † Family Pugnellidae
• Family Seraphsidae
• Family Struthiolariidae
• † Family Thersiteidae
• † Family Tylostomatidae
• Superfamily Tonnoidea
• Family Tonnidae
• Family Bursidae
• Family Laubierinidae
• Family Personidae
• Family Pisanianuridae
• Family Ranellidae
• Superfamily Vanikoroidea
• Family Vanikoridae
• Family Haloceratidae
• Family Hipponicidae
• † Family Omalaxidae
• Superfamily Velutinoidea
• Family Velutinidae
• Family Triviidae
• Superfamily Vermetoidea
• Family Vermetidae
• Superfamily Xenophoroidea
• Family Xenophoridae
• † Family Lamelliphoridae
[edit] Informal group Ptenoglossa

• Superfamily Epitonioidea
• Family Epitoniidae
• Family Janthinidae
• Family Nystiellidae
• Superfamily Eulimoidea
• Family Eulimidae
• Family Aclididae
• Superfamily Triphoroidea
• Family Triphoridae
• Family Cerithiopsidae
• Family Newtoniellidae
[edit] Clade Neogastropoda

• Unassigned to a superfamily
• † Family Johnwyattiidae
• † Family Perissityidae
• † Family Sarganidae
• † Family Speightiidae
• † Family Taiomidae
• † Family Weeksiidae
• Superfamily Buccinoidea
• Family Buccinidae
• Family Colubrariidae
• Family Columbellidae
• Family Fasciolariidae
• Family Nassariidae
• Family Melongenidae
• Superfamily Muricoidea
• Family Muricidae
• Family Babyloniidae
• Family Costellariidae
• Family Cystiscidae
• Family Harpidae
• Family Marginellidae
• Family Mitridae
• † Family Pholidotomidae
• Family Pleioptygmatidae
• Family Strepsiduridae
• Family Turbinellidae
• Family Volutidae
• Family Volutomitridae
• Superfamily Olivoidea
• Family Olividae
• Family Olivellidae
• Superfamily Pseudolivoidea
• Family Pseudolividae
• Family Ptychatractidae
• Superfamily Conoidea
• Family Conidae
• Family Clavatulidae
• Family Drilliidae
• Family Pseudomelatomidae
• Family Strictispiridae
• Family Terebridae
• Family Turridae
• Superfamily Cancellarioidea
• Family Cancellariidae
[edit] Clade Heterobranchia

Contains the informal groups Heterobranchia, Opisthobranchia and Pulmonata
[edit] Informal group "Lower Heterobranchia" (= Allogastropoda)

• Unassigned to a superfamily
• Family Cimidae
• † Family Dolomitellidae
• † Family Heterosubulitidae
• † Family Kuskokwimiidae
• † Family Misurinellidae
• Family Orbitestellidae
• Family Tjaernoeiidae
• Family Xylodisculidae
• Superfamily Acteonoidea
• Family Acteonidae
• † Family Acteonellidae
• Family Aplustridae
• Family Bullinidae
• † Family Zardinellidae
• Superfamily Architectonicoidea
• Family Architectonicidae
• † Family Amphitomariidae
• † Family Cassianaxidae
• Superfamily Glacidorboidea
• Family Glacidorbidae
• Superfamily Mathildoidea
• Family Mathildidae
• † Family Ampezzanildidae
• † Family Anoptychiidae
• † Family Gordenellidae
• † Family Tofanellidae
• † Family Trachoecidae
• Superfamily Nerineoidea
• † Family Nerineidae
• † Family Ceritellidae
• † Family Nerinellidae
• Superfamily Omalogyroidea
• Family Omalogyridae
• † Family Studraxidae
• Superfamily Pyramidelloidea
• Family Pyramidellidae
• Family Amathinidae
• † Family Heteroneritidae
• Family Murchisonellidae
• Superfamily Ringiculoidea
• Family Ringiculidae
• Superfamily Rissoelloidea
• Family Rissoellidae
• Superfamily Streptacidoidea
• † Family Streptacididae
• † Family Cassianebalidae
• Superfamily Valvatoidea
• Family Valvatidae
• Family Cornirostridae
• Family Hyalogyrinidae
• † Family Provalvatidae
[edit] Informal group Opisthobranchia

Contains the clades Cephalaspidea, Thecosomata, Gymnosomata, Aplysiomorpha, Sacoglossa,
Umbraculida, Nudipleura and the groups Acochlidiacea and Cylindrobullida.
[edit] Clade Cephalaspidea
• Superfamily Bulloidea
• Family Bullidae
• Superfamily Diaphanoidea
• Family Diaphanidae
• Family Notodiaphanidae
• Superfamily Haminoeoidea
• Family Haminoeidae
• Family Bullactidae
• Family Smaragdinellidae
• Superfamily Philinoidea
• Family Philinidae
• Family Aglajidae
• Family Cylichnidae
• Family Gastropteridae
• Family Philinoglossidae
• Family Plusculidae
• Family Retusidae
• Superfamily Runcinoidea
• Family Runcinidae
• Family Ilbiidae
[edit] Clade Thecosomata
• Superfamily Cavolinioidea
• Family Cavoliniidae
• Family Limacinidae
• † Family Sphaerocinidae
• Superfamily Cymbulioidea
• Family Cymbuliidae
• Family Desmopteridae
• Family Peraclidae
[edit] Clade Gymnosomata

• Superfamily Clionoidea
• Family Clionidae
• Family Cliopsidae
• Family Notobranchaeidae
• Family Pneumodermatidae
• Superfamily Hydromyloidea
• Family Hydromylidae
• Family Laginiopsidae
[edit] Clade Aplysiomorpha (= Anaspidea)

• Superfamily Aplysioidea
• Family Aplysiidae
• Superfamily Akeroidea
• Family Akeridae
[edit] Group Acochlidiacea

• Superfamily Acochlidioidea
• Family Acochlidiidae
• Superfamily Hedylopsoidea
• Family Hedylopsidae
• Family Ganitidae
• Family Livorniellidae
• Family Minicheviellidae
• Family Parhedylidae
• Family Tantulidae
• Superfamily Palliohedyloidea
• Family Palliohedylidae
• Superfamily Strubellioidea
• Family Strubelliidae
• Family Pseudunelidae
[edit] Clade Sacoglossa
[edit] Subclade Oxynoacea

• Superfamily Oxynooidea
• Family Oxynoidae
• Family Juliidae
• Family Volvatellidae
[edit] Subclade Placobranchacea

• Superfamily Placobranchoidea
• Family Placobranchidae
• Family Boselliidae
• Family Platyhedylidae
• Superfamily Limapontioidea
• Family Limapontiidae
• Family Caliphyllidae
• Family Hermaeidae
[edit] Group Cylindrobullida

• Superfamily Cylindrobulloidea
• Family Cylindrobullidae
[edit] Clade Umbraculida
• Superfamily Umbraculoidea
• Family Umbraculidae
• Family Tylodinidae
[edit] Clade Nudipleura

[edit] Subclade Pleurobranchomorpha
• Superfamily Pleurobranchoidea
• Family Pleurobranchidae
[edit] Subclade Nudibranchia

Contains the clades Euctinidiacea and Dexiarchia
• Unassigned to superfamily
• Family Rhodopidae
[edit] Clade Euctenidiacea (= Holohepatica)

Contains the subclades Gnathodoridacea and Doridacea
[edit] Subclade Gnathodoridacea

• Superfamily Bathydoridoidea
• Family Bathydorididae
[edit] Subclade Doridacea

• Superfamily Doridoidea
• Family Dorididae
• Family Actinocyclidae
• Family Chromodorididae
• Family Discodorididae
• Superfamily Phyllidioidea
• Family Phyllidiidae
• Family Dendrodorididae
• Family Mandeliidae
• Superfamily Onchidoridoidea
• Family Onchidorididae
• Family Corambidae
• Family Goniodorididae
• Superfamily Polyceroidea (= Phanerobranchiata Non Suctoria)
• Family Polyceridae
• Family Aegiretidae - Aegiretidae is incorrect subsequent spelling of this family in
Bouchet & Rocroi (2005). Correct is Aegiridae.
• Family Gymnodorididae
• Family Hexabranchidae
• Family Okadaiidae
[edit] Clade Dexiarchia (= Actenidiacea)

Contains the clades Pseudoeuctenidiacea and Cladobranchia
[edit] Clade Pseudoeuctenidiacea ( = Doridoxida)

• Superfamily Doridoxoidea
• Family Doridoxidae
[edit] Clade Cladobranchia ( = Cladohepatica)

Contains the subclades Euarminida, Dendronotida and Aeolidida
• Not assigned to a superfamily
• Family Charcotiidae
• Family Dironidae
• Family Dotidae
• Family Embletoniidae
• Family Goniaeolididae
• Family Heroidae
• Family Madrellidae
• Family Pinufiidae
• Family Proctonotidae
[edit] Subclade Euarminida
• Superfamily Arminoidea
• Family Arminidae
• Family Doridomorphidae
[edit] Subclade Dendronotida

• Superfamily Tritonioidea
• Family Tritoniidae
• Family Aranucidae
• Family Bornellidae
• Family Dendronotidae
• Family Hancockiidae
• Family Lomanoridae
• Family Phylliroidae
• Family Scyllaeidae
• Family Tethydidae
[edit] Subclade Aeolidida

• Superfamily Flabellinoidea ( = Pleuroprocta)
• Family Flabellinidae
• Family Notaeolidiidae
• Superfamily Fionoidea
• Family Fionidae
• Family Calmidae
• Family Eubranchidae
• Family Pseudovermidae
• Family Tergipedidae
• Superfamily Aeolidioidea
• Family Aeolidiidae
• Family Facelinidae
• Family Glaucidae
• Family Piseinotecidae
[edit] Informal Group Pulmonata

Contains the informal group Basommatophora and the clade Eupulmonata
[edit] Informal Group Basommatophora

Contains the clade Hygrophila
• Superfamily Amphiboloidea
• Family Amphibolidae
• Superfamily Siphonarioidea
• Family Siphonariidae
• † Family Acroreiidae
[edit] Clade Hygrophila
• Superfamily Chilinoidea
• Family Chilinidae
• Family Latiidae
• Superfamily Acroloxoidea
• Family Acroloxidae
• Superfamily Lymnaeoidea
• Family Lymnaeidae
• Superfamily Planorboidea
• Family Planorbidae
• Family Physidae
[edit] Clade Eupulmonata

Contains the clades Systellommatophora and Stylommatophora
• Superfamily Trimusculoidea
• Family Trimusculidae
• Superfamily Otinoidea
• Family Otinidae
• Family Smeagolidae
• Superfamily Ellobioidea
• Family Ellobiidae
[edit] Clade Systellommatophora (= Gymnomorpha)
• Superfamily Onchidioidea
• Family Onchidiidae
• Superfamily Veronicelloidea
• Family Veronicellidae
• Family Rathouisiidae
[edit] Clade Stylommatophora

Contains the subclades Elasmognatha, Orthurethra and the informal group Sigmurethra
[edit] Subclade Elasmognatha

• Superfamily Succineoidea
• Family Succineidae
• Superfamily Athoracophoroidea
• Family Athoracophoridae
[edit] Subclade Orthurethra

• Superfamily Partuloidea
• Family Partulidae
• Family Draparnaudiidae
• Superfamily Achatinelloidea
• Family Achatinellidae
• Superfamily Cochlicopoidea
• Family Cochlicopidae
• Family Amastridae
• Superfamily Pupilloidea
• Family Pupillidae
• Family Argnidae
• Family Chondrinidae
• † Family Cylindrellinidae
• Family Lauriidae
• Family Orculidae
• Family Pleurodiscidae
• Family Pyramidulidae
• Family Spelaeodiscidae
• Family Strobilopsidae
• Family Valloniidae
• Family Vertiginidae
• Superfamily Enoidea
• Family Enidae
• Family Cerastidae
[edit] Informal Group Sigmurethra

• Superfamily Clausilioidea
• Family Clausiliidae
• † Family Anadromidae
• † Family Filholiidae
• † Family Palaeostoidae
• Superfamily Orthalicoidea
• Family Orthalicidae
• Family Cerionidae
• Family Coelociontidae
• † Family Grangerellidae
• Family Megaspiridae
• Family Placostylidae
• Family Urocoptidae
• Superfamily Achatinoidea
• Family Achatinidae
• Family Ferussaciidae
• Family Micractaeonidae
• Family Subulinidae
• Superfamily Aillyoidea
• Family Aillyidae
• Superfamily Testacelloidea
• Family Testacellidae
• Family Oleacinidae
• Family Spiraxidae
• Superfamily Papillodermatoidea
• Family Papillodermatidae
• Superfamily Streptaxoidea
• Family Streptaxidae
• Superfamily Rhytidoidea
• Family Rhytididae
• Family Chlamydephoridae
• Family Haplotrematidae
• Family Scolodontidae
• Superfamily Acavoidea
• Family Acavidae
• Family Caryodidae
• Family Dorcasiidae
• Family Macrocyclidae
• Family Megomphicidae
• Family Strophocheilidae
• Superfamily Plectopyloidea
• Family Plectopylidae
• Family Corillidae
• Family Sculptariidae
• Superfamily Punctoidea
• Family Punctidae
• † Family Anastomopsidae
• Family Charopidae
• Family Cystopeltidae
• Family Discidae
• Family Endodontidae
• Family Helicodiscidae
• Family Oreohelicidae
• Family Thyrophorellidae
• Superfamily Sagdoidea
• Family Sagdidae
[edit] limacoid clade

• Superfamily Staffordioidea
• Family Staffordiidae
• Superfamily Dyakioidea
• Family Dyakiidae
• Superfamily Gastrodontoidea
• Family Gastrodontidae
• Family Chronidae
• Family Euconulidae
• Family Oxychilidae
• Family Pristilomatidae
• Family Trochomorphidae
• Fossil taxa that probably belong to the superfamily Gastrodontoidea:
• Subfamily † Archaeozonitinae
• Subfamily † Grandipatulinae
• Subfamily † Palaeoxestininae
• Superfamily Parmacelloidea
• Family Parmacellidae
• Family Milacidae
• Family Trigonochlamydidae
• Superfamily Zonitoidea
• Family Zonitidae
• Superfamily Helicarionoidea
• Family Helicarionidae
• Family Ariophantidae
• Family Urocyclidae
• Superfamily Limacoidea
• Family Limacidae
• Family Agriolimacidae
• Family Boettgerillidae
• Family Vitrinidae
[edit] Informal group Sigmurethra continued

Two other superfamilies are part of the clade Sigmurethra, but they are not in the limacoid clade:
• Superfamily Arionoidea
• Family Arionidae
• Family Anadenidae
• Family Ariolimacidae
• Family Binneyidae
• Family Oopeltidae
• Family Philomycidae
• Superfamily Helicoidea
• Family Helicidae
• Family Bradybaenidae
• Family Camaenidae
• Family Cepolidae
• Family Cochlicellidae
• Family Elonidae
• Family Epiphragmophoridae
• Family Halolimnohelicidae
• Family Helicodontidae
• Family Helminthoglyptidae
• Family Humboldtianidae
• Family Hygromiidae
• Family Monadeniidae
• Family Pleurodontidae
• Family Polygyridae
• Family Sphincterochilidae
• Family Thysanophoridae
• Family Trissexodontidae
• Family Xanthonychidae
[edit] See also

• Changes in the taxonomy of gastropods since 2005
[edit] References
1. ^ a b Bouchet P., Rocroi J.-P., Frýda J., Hausdorf B., Ponder W., Valdés Á. & Warén A.
(2005). "Classification and nomenclator of gastropod families ". Malacologia: International Journal of
Malacology (Hackenheim, Germany: ConchBooks) 47 (1-2): 1–397. ISBN 3925919724. ISSN 0076-
2997. http://www.archive.org/details/malacologia47122005inst.

Wikimedia Commons has media related to: Template:Taxonavigation/Taxonomy of
the Gastropoda (Bouchet & Rocroi, 2005)
• Full text of Malacologia vol. 47 (1-2)

Retrieved from "http://en.wikipedia.org/wiki/Taxonomy_of_the_Gastropoda_(Bouchet_
%26_Rocroi,_2005)"
Categories: Gastropods | Animal taxonomy
W000
Woodpecker

For other uses, see Woodpecker (disambiguation).
Woodpeckers, piculets and wrynecks
Hispaniolan Woodpecker
Tapping sound of a woodpecker (help·info)
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Subclass: Neornithes
Infraclass: Neognathae
Superorder: Neoaves
Order: Piciformes
Suborder: Pici
Picidae
Family:
Vigors, 1825
Subfamilies
Jynginae - wrynecks
Nesoctitinae - Antillean Piculet
Picinae - woodpeckers
Picumninae - typical piculets
The woodpeckers, piculets and wrynecks are a family, Picidae, of near-passerine birds. Members of
this family are found worldwide, except for Australia and New Zealand, Madagascar, and the extreme polar
regions. Most species live in forests or woodland habitats, although a few species are known to live in
treeless areas such as rocky hillsides and deserts.
The Picidae are just one of the eight living families in the order Piciformes. Members of the order
Piciformes, such as the jacamars, puffbirds, barbets, toucans and honeyguides, have traditionally been
thought to be very closely related to the woodpeckers, piculets and wrynecks. More recently, DNA sequence
analyses have confirmed this view.[1]
There are about 200 species and about 30 genera in this family. Many species are threatened or
endangered due to loss of habitat or habitat fragmentation. Two species of woodpeckers, the Ivory-billed
Woodpecker and the Imperial Woodpecker, have been considered extinct for about 30 years (there has been
some controversy recently whether these species still exist).
Contents
[hide]
• 1 Description
• 2 Distribution, habitat and movements
• 3 Behavior
• 3.1 Diet and feeding
• 3.2 Breeding
• 4 Systematics and evolution
• 5 List of genera
• 6 See also
• 7 Footnotes
• 8 References
[edit] Description
The stiffened tails of woodpeckers are crucial for their climbing and foraging techniques. The tail is
used as a prop. Here a Black-rumped Flameback rests while foraging using its tail for support.
The smallest woodpecker is the Bar-breasted Piculet, at 7 g and 8 cm (3 1/4 inches). The largest
woodpecker was the Imperial Woodpecker, at an average of 58 cm (23 inches) and probably over 600 g
(1.3 lb). The Ivory-billed Woodpecker is (or was) slightly smaller at 50 cm (20 inches) and a weight of 500 g
(1.1 lb). If both the Ivory-billed and Imperial Woodpeckers are indeed extinct, the largest extant woodpecker
is the Great Slaty Woodpecker of Southeast Asia, at about 50 cm (20 inches) and 450 g (1 lb). A number of
species exhibit sexual dimorphism in size, bill length and weight. In the piculets it is often the females that are
larger, amongst the woodpeckers that show sexual dimorphism it is usually the males that are larger.
Most species possess predominantly white, black and brown, green and red plumage, although
many piculets show a certain amount of grey and olive green. In woodpeckers, many species exhibit patches
of red and yellow on their heads and bellies, and these bright areas are important in signalling. The dark
areas of plumage are often iridescent. Although the sexes of Picidae species tend to look alike, many
woodpecker species have more prominent red or yellow head markings in males than in females.
Woodpeckers use their long tongues to feed, on feeders as well as from holes in wood or soil. This is
a male Hairy Woodpecker.
Members of the family Picidae have strong bills for drilling and drumming on trees and long sticky
tongues for extracting food.[2] Woodpecker bills are typically longer, sharper and stronger than the bills of
piculets and wrynecks; however their morphology is very similar. The bill's chisel-like tip is kept sharp by the
pecking action in birds that regularly use it on wood. Species of woodpecker and flicker that use their bills in
soil or for probing as opposed to regular hammering tend to have longer and more decurved bills. Due to
their smaller bill size, many piculets and wrynecks will forage in decaying wood more often than
woodpeckers. The long sticky tongues, which possess bristles, aid these birds in grabbing and extracting
insects deep within a hole of a tree. It had been reported that the tongue was used to spear grubs, but more
detailed studies published in 2004 have shown that the tongue instead wraps around the prey before being
pulled out.[3]
Many of the foraging, breeding and signaling behaviors of woodpeckers involve drumming and
hammering using the bill.[4] To prevent brain damage from the rapid and repeated decelerations,
woodpeckers have evolved a number of adaptations to protect the brain. These include small brain size, the
orientation of the brain within the skull (which maximises the area of contact between the brain and the skull)
and the short duration of contact. The millisecond before contact with wood a thickened nictitans membrane
closes, protecting the eye from flying debris.[5] The nostrils are also protected; they are often slit-like and
have special feathers to cover them.
Woodpeckers, piculets and wrynecks all possess zygodactyl feet. Zygodactyl feet consist of four
toes, the first and the fourth facing frontward and the second and third facing back. This foot arrangement is
good for grasping the limbs and trunks of trees. Members of this family can walk vertically up a tree trunk,
which is beneficial for activities such as foraging for food or nest excavation. In addition to the strong claws
and feet woodpeckers have short strong legs, this is typical of birds that regularly forage on trunks. The tails
of all woodpeckers except the piculets and wrynecks are stiffened, and when the bird perches on vertical
surfaces, the tail and feet work together to support it. [2]
[edit] Distribution, habitat and movements
The use of cacti for breeding and roosting holes allows some woodpeckers, like this Ladder-backed
Woodpecker, to live in otherwise treeless deserts.
The woodpeckers have a mostly cosmopolitan distribution, although they are absent from
Australasia, Madagascar and Antarctica. They are also absent from the world's oceanic islands, although
many insular species are found on continental islands. The true woodpeckers, subfamily Picinae, are
distributed across the entire range of the woodpeckers. The Picumninae piculets have a pantropical
distribution, with species in Southeast Asia, Africa and the Neotropics, with South America holding the
majority of piculet species. The second piculet subfamily, Nesoctitinae, has a single species, the Antillean
Piculet, which is restricted to the Caribbean island of Hispaniola. The wrynecks (Jynginae) have an
exclusively Old World distribution, with the two species occurring in Europe, Asia and Africa.
Overall the woodpeckers are arboreal birds of wooded habitats. They reach their greatest diversity in
tropical rainforests, but occur in almost all suitable habitats including woodlands, savannahs, scrublands,
bamboo forests. Even grasslands and deserts have been colonised by various species. These habitats are
more easily occupied where a small number of trees exist, or, in the case of desert species like the Gila
Woodpecker, tall cacti are available for nesting in.[6] A number of species are adapted to spending a portion
of their time feeding on the ground, and a very small minority of species have abandoned trees entirely and
nest in holes in the ground. The Ground Woodpecker is one such species, inhabiting the rocky and grassy
hills of South Africa.
Picidae species can either be sedentary or migratory. Many species are known to stay in the same
area year-round while others travel great distances from their breeding grounds to their wintering grounds.
For example, the Eurasian Wryneck breeds in Europe and west Asia and migrates to the Sahel in Africa in
the winter.[7]
Results from the monitoring programs of the Swiss Ornithological Institute show that the breeding
populations of several forest species for which deadwood is an important habitat element (black woodpecker,
great spotted woodpecker, middle spotted woodpecker, lesser spotted woodpecker, green woodpecker,
three-toed woodpecker as well as crested tit, willow tit and Eurasian tree creeper) have increased in the
period 1990 to 2008, although not to the same extent in all species. At the same time the white-backed
woodpecker extended its range in eastern Switzerland. The Swiss National Forest Inventory shows an
increase in the amount of deadwood in forests for the same period. For all the mentioned species, with the
exception of green and middle spotted woodpecker, the growing availability of deadwood is likely to be the
most important factor explaining this population increase.
[edit] Behavior
[[:Image:|Woodpecker]]
[[Image:|220px|noicon|alt=]]
A woodpecker pecking into a tree.
Problems listening to this file? See media help.

The woodpeckers range from highly antisocial solitary species that are aggressive to other members
of their species, to species that live in groups. Group-living species tend to be communal group breeders. In
addition to these species, a number of species may join mixed-species feeding flocks with other
insectivorous birds, although they tend to stay at the edges of these groups. Joining these flocks allows
woodpeckers to decrease anti-predator vigilance and increase their feeding rate.[8] Woodpeckers are
diurnal, roosting at night inside holes. In most species the roost will become the nest during the breeding
season.
[edit] Diet and feeding
Holes bored by woodpeckers feeding, Gatineau Park, Quebec
The diet of woodpeckers consists mainly of insects and their grubs taken from living and dead trees,
and other arthropods, along with fruit from live trees, nuts and sap both from live trees. Their role ecologically
is thereby keeping trees healthy by keeping them from suffering mass infestations. The family is noted for its
ability to acquire wood-boring grubs using their bills for hammering, but overall the family is characterized by
its dietary flexibility, with many species being both highly omnivorous and opportunistic. The insect prey most
commonly taken are insects found inside tree trunks, whether they are alive or rotten wood and in crevices in
bark on trees. These include beetles and their grubs, ants, termites, spiders, and caterpillars. These may be
obtained either by gleaning or more famously by excavating wood. Having hammered a hole into the wood
the prey is excavated by a long barbed tongue. The ability to excavate allows woodpeckers to obtain tree
sap, an important source of food for some species. Most famously the sapsuckers, (genus Sphyrapicus )
feed in this fashion, but the technique is not restricted to these and others such as the Acorn Woodpecker
and White-headed Woodpecker also feed in this way. It was once thought that the technique was restricted to
the New World, but Old World species such as the Arabian Woodpecker and Great Spotted Woodpecker
also feed in this way.[2]
[edit] Breeding
File:Black Woods.jpg
A male Black Woodpecker attending its chicks
All members of the family Picidae nest in cavities. Almost every species nests in tree cavities,
although in deserts some species nest inside holes in cactus and a few species nest in holes dug into the
earth. Woodpeckers and piculets will excavate their own nests, but wrynecks will not. The excavated nest is
usually only lined from the wood chips produced as the hole was made. Many species of woodpeckers
excavate one hole per breeding season, sometimes after multiple attempts. It takes around a month to finish
the job. Abandoned holes are used by other birds and mammals that are secondary cavity nesters.[9]
Because nesting holes are in great demand by other cavity nesters, woodpeckers face competition for the
nesting sites they excavate from the moment the hole becomes usable. This may come from other species of
woodpecker, or other cavity nesting birds like swallows and starlings. Woodpeckers may aggressively harass
potential competitors, and also use other strategies to reduce the chance of being usurped from their nesting
site; for example the Red-crowned Woodpecker digs its nest in the underside of a small branch, which
reduces the chance that a larger species will take it over and expand it.[10]
Members of Picidae are typically monogamous, with a few species breeding cooperatively and some
polygamy reported in a few species.[11] Polyandry, where a female raises two broods with two separate
males, has also been reported in the West Indian Woodpecker.[12] A pair will work together to help build the
nest, incubate the eggs and raise their altricial young. However, in most species the male does most of the
nest excavation and takes the night shift while incubating the eggs. A nest will usually consist of 2-5 round
white eggs. Since these birds are cavity nesters, their eggs do not need to be camouflaged and the white
color helps the parents to see them in dim light. The eggs are incubated for about 11–14 days before the
chicks are born. It takes about 18–30 days before the young are ready to leave the nest.
[edit] Systematics and evolution

The phylogeny has been updated according to new knowledge about convergence patterns and
evolutionary history.[13] Most notably, the relationship of the picine genera has been largely clarified, and it
was determined that the Antillean Piculet is a surviving offshoot of proto-woodpeckers.
The evolutionary history of this group is not well documented, but the known fossils allow some
preliminary conclusions: the earliest known modern picids were piculet-like forms of the Late Oligocene,
about 25 million years ago (mya). By that time, however, the group was already present in the Americas and
Europe, and it is hypothesized that they actually evolved much earlier, maybe as early as the Early Eocene
(50 mya). The modern subfamilies appear to be rather young by comparison; until the mid-Miocene (10-15
mya), all picids seem to have been small or mid-sized birds similar to a mixture between a piculet and a
wryneck. On the other hand, there exists a feather enclosed in fossil amber from the Dominican Republic,
dated to about 25 mya, which seems to indicate that the Nesoctitinae were already a distinct lineage by then.
[14]
Prehistoric representatives of the extant Picidae genera are treated in the genus articles. An
enigmatic form based on a coracoid found in Pliocene deposits of New Providence, Bahamas, has been
described as Bathoceleus hyphalus and probably also is a woodpecker.[15]
[edit] List of genera

Young Eurasian Wryneck (Jynx torquilla), a relative of the woodpeckers.
Ochre-collared Piculet (Picumnus temminckii).

Female Yellow-shafted Flicker (Colaptes auratus auratus)
The famous Ivory-billed Woodpecker (Campephilus principalis) is probably extinct.

FAMILY: PICIDAE
• Basal
• Genus: Palaeopicus (Late Oligocene of France)
• Incertae sedis
• Picidae gen. et sp. indet. (Middle Miocene of New Mexico, USA)
• Picidae gen. et sp. indet. (Late Miocene of Gargano Peninsula, Italy)
• Subfamily: Jynginae - Wrynecks
• Genus: Jynx (2 species)
• Subfamily: Picumninae - Typical piculets
• Genus: Picumnus - American Piculets (c.27 species)
• Genus: Verreauxia - African Piculet (sometimes included in Sasia)
• Genus: Sasia - Asian Piculets (2 species)
• Subfamily: Nesoctitinae
• Genus Nesoctites - Antillean Piculet
• Subfamily: Picinae - Woodpeckers
• Incertae sedis
• Genus: Palaeonerpes (Ogalalla Early Pliocene of Hitchcock County, USA) -
possibly dendropicine
• Genus: Pliopicus (Early Pliocene of Kansas, USA) - possibly dendropicine
• cf. Colaptes DMNH 1262 (Early Pliocene of Ainsworth, USA) - malarpicine?
• Tribe: Dendropicini
• Genus: Melanerpes (some 22 species)
• Genus: Sphyrapicus - sapsuckers (4 species)
• Genus: Xiphidiopicus - Cuban Woodpecker (Placement in Dendropicini
tentative)
• Genus: Dendropicos (15 species)
• Genus: Mesopicos (3 species)
• Genus: Dendrocopos (21 species)
• Genus: Picoides (presently 12 species; maybe only 3 belong here) - this
genus is in need of revision.[16] See the genus article for more.
• Genus: Veniliornis (14 species)
• Tribe: Malarpicini
• Genus: Campethera (12 species)
• Genus: Geocolaptes - Ground Woodpecker
• Genus: Dinopium - malarpicine flamebacks (4 species)
• Genus: Meiglyptes (3 species)
• Genus: Hemicircus (2 species; placement in Malarpicini tentative)
• Genus: Micropternus - Rufous Woodpecker (formerly in Celeus)
• Tribe: Picini (sometimes included in Malarpicini)
• Genus: Picus (c.15 species)
• Genus: Mulleripicus (3 species)
• Genus: Dryocopus (7 species)
• Genus: Celeus (11 species)
• Genus: Piculus (7 species)
• Genus: Colaptes - flickers (about 12 species)
• Tribe: Megapicini
• Genus: Campephilus (11 species, 2 possibly recently extinct)
• Genus: Chrysocolaptes - megapicine flamebacks (2 species)
• Genus: Reinwardtipicus - Orange-backed Woodpecker
• Genus: Blythipicus (2 species)
• Genus: Gecinulus (2 species; placement in Megapicini tentative)
• Genus: Sapheopipo - Okinawa Woodpecker (Placement in Megapicini
tentative)
[edit] See also

• List of bird speices
• Wrynecks
• Piculets
• Sapsuckers
[edit] Footnotes
1. ^ Johansson & Ericson (2003)
2. ^ a b c Winkler, Hans & Christie, David A. (2002), "Family Picidae (Woodpeckers)" in del
Hoyo, J.; Elliot, A. & Sargatal, J. (editors). (2002). Handbook of the Birds of the World. Volume 7:
Jacamars to Woodpeckers. Lynx Edicions. ISBN 8487334377
3. ^ Villard, Pascal; Jacques Cuisin (2004). "How do woodpeckers extract grubs with their
tongues? A study of the Guadeloupe woodpecker (Melanerpes herminieri) in the French Indies". Auk
121 (2): 509–514. doi:10.1642/0004-8038(2004)121[0509:HDWEGW]2.0.CO;2.
4. ^ Gibson L. (2006) "Woodpecker pecking: how woodpeckers avoid brain injury" Journal of
Zoology 270: 462–465 doi:10.1111/j.1469-7998.2006.00166.x
5. ^ Schwab I (2002) "Cure for a headache" British Journal of Ophthalmology 86 : 843
doi:10.1136/bjo.86.8.843
6. ^ Korol, Jerome; Richard Hutto (1984). "Factors Affecting Nest Site Location in Gila
Woodpeckers". Condor 86 (1): 73–78. doi:10.2307/1367350.
http://elibrary.unm.edu/sora/Condor/files/issues/v086n01/p0073-p0078.pdf.
7. ^ Reichlin, Thomas; Michael Schaub, Myles H. M. Menz, Murielle Mermod, Patricia Portner,
Raphaël Arlettaz & Lukas Jenni (2008). "Migration patterns of Hoopoe Upupa epops and Wryneck
Jynx torquilla : an analysis of European ring recoveries". Journal of Ornithology 150: 393.
doi:10.1007/s10336-008-0361-3.
8. ^ Kimberly, Sullivan (1984). "Information Exploitation By Downy Woodpeckers in Mixed-
Species Flocks". Behavior 91 (4): 294–311. doi:10.1163/156853984X00128.
9. ^ Kotaka N & S Matsuoka (2002) "Secondary users of Great Spotted Woodpecker
(Dendrocopos major) nest cavities in urban and suburban forests in Sapporo City, northern Japan" .
Ornithological Science 1 (2): 117-122
10.^ Short, Lester L. (1979). "Burdens of the Picid Hole-Excavating Habit". Wilson Bulletin 91
(1): 16–28. http://elibrary.unm.edu/sora/Wilson/v091n01/p0016-p0028.pdf.
11.^ Wiktander, Ulf; Ola Olsson & Sven G. Nilsson, (2000). "Parental care and social mating
system in the Lesser Spotted Woodpecker Dendrocopos minor". Journal of Avian Biology 31 (4):
447–456. doi:10.1034/j.1600-048X.2000.310003.x.
12.^ Willimont, LA; Jackson, JA and Jackson, BJS (1991). "Classical polyandry in the West
Indian woodpecker on Abaco, Bahamas". Willson Bulletin 103: 124–125.
http://elibrary.unm.edu/sora/Wilson/v103n01/p0124-p0125.pdf.
13.^ Benz et al. (2006), Moore et al. (2006)
14.^ Grimaldi & Case (1995)
15.^ Cracraft & Morony (1969)
16.^ Moore et al. (2006)
[edit] References
• Benz, Brett W.; Robbins, Mark B. & Peterson, A. Townsend (2006): Evolutionary history of
woodpeckers and allies (Aves: Picidae): Placing key taxa on the phylogenetic tree. Mol. Phylogenet.
Evol. 40(2): 389–399. doi:10.1016/j.ympev.2006.02.021 (HTML abstract)
• Cracraft, Joel & Morony, John J. Jr. (1969): A new Pliocene woodpecker, with comments on
the fossil Picidae. American Museum Novitates 2400: 1-8. PDF fulltext
• Gorman, Gerard (2004): Woodpeckers of Europe: A Study of the European Picidae. Bruce
Coleman, UK. ISBN 1 872842 05 4.
• Grimaldi, David A. & Case, Gerard Ramon (1995): A feather in amber from the Upper
Cretaceous of New Jersey. American Museum Novitates 3126: 1-6. PDF fulltext
• Johansson, U.S. & Ericson, G.P. (2003): Molecular support for a sister group relationship
between Pici and Galbulae (Piciformes sensu Wetmore 1960). J. Avian Biol. 34(2): 185-197.
doi:10.1034/j.1600-048X.2003.03103.x PDF fulltext
• Koenig, W.D. & Haydock, J. (1999): Oaks, acorns, and the geographical ecology of acorn
woodpeckers. J. Biogeogr. 26(1): 159-165. doi:10.1046/j.1365-2699.1999.00256.x (HTML abstract)
• Lemaitre, J. & Villard, M.A. (2005): Foraging patterns of pileated woodpeckers in a managed
Acadian forest: a resource selection function. Can. J. Forest Res. 35(10): 2387-2393.
doi:10.1139/x05-148 (HTML abstract)
• Michalek, K.G. & Winkler, H. (2001): Parental care and parentage in monogamous great
spotted woodpeckers (Picoides major) and middle spotted woodpeckers (Picoides medius).
Behaviour 138(10): 1259-1285. doi:10.1163/15685390152822210 (HTML abstract)
• Moore, William S.; Weibel, Amy C. & Agius, Andrea (2006): Mitochondrial DNA phylogeny of
the woodpecker genus Veniliornis (Picidae, Picinae) and related genera implies convergent evolution
of plumage patterns. Biol. J. Linn. Soc. 87(4): 611–624. doi:10.1111/j.1095-8312.2006.00586.x PDF
fulltext
• Stark, R.D.; Dodenhoff, D.J. & Johnson, E.V. (1998): A quantitative analysis of woodpecker
drumming. Condor 100(2): 350-356. DjVu fulltext PDF fulltext
• Villard, P.; Cuisin, J. & Karasov, W.H. (2004). How do woodpeckers extract grubs with their
tongues? A study of the Guadeloupe woodpecker (Melanerpes herminieri) in the French Indies. Auk
121: 509-514. DOI:10.1642/0004-8038(2004)121[0509:HDWEGW]2.0.CO;2 HTML abstract
• Webb, Daniel Matthew & Moore, William S. (2005): A phylogenetic analysis of woodpeckers
and their allies using 12S, Cyt b, and COI nucleotide sequences (class Aves; order Piciformes). Mol.
Phylogenet. Evol. 36(2): 233-248. doi:10.1016/j.ympev.2005.03.015 PDF fulltext
• Wiebe, K.L. & Swift, T.L. (2001): Clutch size relative to tree cavity size in northern flickers. J.
Avian Biol. 32(2): 167. doi:10.1034/j.1600-048X.2001.320210.x (HTML abstract)
• Wiktander, U.; Olsson, O. & Nilsson, S.F. (2000): Parental care and social mating system in
the lesser spotted woodpecker Dendrocopos minor. J. Avian Biol. 31(4): 447. doi:10.1034/j.1600-
048X.2000.310003.x (HTML abstract)
• Yom-Tov, Y. & Ar, A. (1993): Incubation and fledging durations of woodpeckers. Condor
95(2): 282-287. DjVu fulltext PDF fulltext
Wikimedia Commons has media related to: Picidae
• Woodpecker videos, photos & sounds on the Internet Bird Collection
[hide]v · d · eOrder: Piciformes
Suborder Family
Galbulae Galbulidae (jacamars) • Bucconidae (puffbirds, nunbirds and nunlets)
Lybiidae (African barbets) • Megalaimidae (Asian barbets) • Ramphastidae

Pici (toucans) • Semnornithidae (toucan-barbets) • Capitonidae (American barbets) • Picidae
(woodpeckers, piculets and wrynecks) • Indicatoridae (honeyguides)
Retrieved from "http://en.wikipedia.org/wiki/Woodpecker"
Categories: Bird families | Picidae

W000
Winston Ponder

Winston F. Ponder B.Sc, M.Sc, Ph.D, D.Sc. (born about 1944) is a noted malacologist from New
Zealand who has named and described many marine animals, especially micromolluscs.[1] He is a graduate
of the University of Auckland, New Zealand.
Ponder was the principal research scientist in the malacology section of the Australian Museum,
Sydney, Australia. He helped to build up the museum's mollusc collection so that it became one of the most
extensive of its kind in the world. Ponder retired from this post after a long career of more than forty years of
research on molluscs, and he is now an Honorary Fellow.
He is also a managing editor of the Malacological Society of Australasia,[2] the Natural History
Museum of Los Angeles, and the South Australian Museum in Adelaide. He has been the president of the
Society of Australian Systematic Biologists, and he is the managing editor of the journal Molluscan Research.
Early in his career, in 1964, he worked on Antarctic collections together with Richard Dell and Alan
Beu, resulting in a major monograph on the Antarctic bivalves, chitons and scaphopods.
Ponder is the author of more than 100 research publications. Many of these are on the subjects of
the freshwater molluscs of Australia, and on invertebrate conservation. However, his major contribution was
a taxonomy of the Gastropoda, which he published together with David R. Lindberg in 1997 [3] This was the
last major publication on the taxonomy of the Gastropoda that was based on the morphology of snails and
slugs (their internal and external shapes and forms), and did not take into account any analysis of their DNA
or RNA.
In 2008, again with David Lindberg, he edited the book "Phylogeny and Evolution of the Mollusca" [4]
in which 36 experts provided an up-to-date review on the evolutionary history of the Mollusca, based on
reinvestigation of morphological characters, molecular data and the fossil record.
In 2008 he received the prestigious Australian Marine Sciences Association Silver Jubilee Award, for
a lifetime of achievement in research on marine molluscs.[5]
Contents
[hide]
• 1 Some Gastropod taxa named by Ponder
• 1.1 Higher taxa
• 1.2 Superfamilies
• 1.3 Families
• 1.4 Subfamilies
• 1.5 Genera
• 2 Taxa named after Ponder
• 3 Bibliography
• 4 See also
• 5 References
[edit] Some Gastropod taxa named by Ponder

[edit] Higher taxa
• Subclass Eogastropoda Ponder & Lindberg, 1997
• Order Sorbeoconcha Ponder & Lindberg, 1997
• Suborder Hypsogastropoda Ponder & Lindberg, 1997
[edit] Superfamilies
• Superfamily Glacidorboidea Ponder, 1986
[edit] Families
• Eatoniellidae Ponder, 1965
• Rastodentidae Ponder, 1966
• Elachisinidae Ponder, 1985
• Emblandidae Ponder, 1985
• Epigridae Ponder, 1985
• Amathinidae Ponder, 1987
• Calopiidae Ponder, 1999
[edit] Subfamilies
• Subfamily Pelycidiinae Ponder & Hall, 1983
[edit] Genera
• Microestea Ponder, 1965
• Rufodardanula Ponder, 1965
• Rastodens Ponder, 1966
• Rissolitorina Ponder, 1966
• Tridentifera Ponder,1966
• Fictonoba Ponder, 1967
• Pseudodiala Ponder, 1967
• Pseudestea Ponder, 1967
• Pseudoskenella Ponder, 1973
• Lirobarleeia Ponder, 1983
• Kutikina Ponder & Waterhouse, 1997 [1]
• Kessneria Walker & Ponder, 2001
[edit] Taxa named after Ponder

Ponderia Hoaurt, 1986
• Aspella ponderi Radwin & D' Attilio, 1976
• Heliacus cerdaleus ponderi Garrard, 1977
• Limatula (Stabilima) ponderi Fleming, 1978
• Echineulima ponderi Warén, 1980
• Pisinna ponderi Palazzi, 1982
• Notocrater ponderi B. A. Marshall, 1986
• Oliva (Miniaceoliva) caerulea ponderi Petuch & Sargent, 1986
• Sassia (Sassia) ponderi Beu, 1986
• Tritonoharpa ponderi Beu & Maxwell, 1987
• Favartia (Caribiella) ponderi Myers & d'Attilio, 1989
• Amalda (Alcospira) ponderi Ninomiya, 1991
• Choristella ponderi McLean, 1992
• Austrotrochaclis ponderi B. A. Marshall, 1995
• Powellisetia ponderi Numanami, 1996
• Fissidentalium ponderi Lamprell & Healy, 1998.
[edit] Bibliography
• Ponder W. F. & A. Warén (1988). "Classification of the Caenogastropoda and Heterostropha
- A list of the family-group names and higher taxa". Malacological Review Supll. (4): 288–328.
• Clark A., Miller A. C. & Ponder W. F. (2003) Revision of the snail genus Austropyrgus
(Gastropoda: Hydrobiidae). 109 pp.
• Ponder, W. F. (4 July 2003). "Monograph of the Australian Bithyniidae (Caenogastropoda:
Rissooidea)". Zootaxa 230: 1–126. http://www.mapress.com/zootaxa/2003f/z00230f.pdf. - There are
17 newly described species from the genus Gabbia.
• Colgan D. J., Ponder W. F., Beacham E. & Macaranas J. (2006). "Molecular phylogenetics of
Caenogastropoda (Gastropoda: Mollusca)". Molecular Phylogenetics and Evolution 42(3): 717-737.
doi:10.1016/j.ympev.2006.10.009 PDF
• Ponder W. & Lindberg D. R. (2008). Phylogeny and Evolution of the Mollusca. University of
California Press, 469 pp. ISBN 9780520250925.
[edit] See also

• Taxonomy of the Gastropoda (Ponder & Lindberg, 1997)
[edit] References
1. ^ http://amonline.net.au/invertebrates/staff/ponder.htm
2. ^ Malacological Society of Australasia
3. ^ Ponder W. & Lindberg D. R. 1997. Towards a phylogeny of gastropod molluscs; an
analysis using morphological characters. Zoological Journal of the Linnean Society, 119: 83-265.
4. ^ Winston F. Ponder and David R. Lindberg (eds.) (2008). Phylogeny and Evolution of the
Mollusca. University of California Press. pp. 488.
http://www.calnaturalhistory.com/books/pages/10802.php.
5. ^ Recognising research on molluscs

• photo
Persondata
Name Ponder, Winston
Alternative names
Short description
Date of birth 1944
Place of birth
Date of death
Place of death
Retrieved from "http://en.wikipedia.org/wiki/Winston_Ponder"
Categories: Malacologists | Australian biologists | Living people | 1944 births | University of Auckland
alumni
W000
Human

"Homo sapiens" redirects here. For other uses, see Homo sapiens (disambiguation).
This article is about modern humans. For other human species, see Homo. For other uses, see
Human (disambiguation).
Human
Craniata
Human[1]
Temporal range: 0.2–0 Ma
PreЄ ↓
g
Pleistocene – Recent
Human male and female
Conservation status
Least Concern (IUCN 3.1)[2]

Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Family: Hominidae
Genus: Homo
Species: H. sapiens
Subspecies: H. s. sapiens
Homo sapiens sapiens
Linnaeus, 1758
Humans, known taxonomically as Homo sapiens[3][4] (Latin for "wise man" or "knowing man"),[5] are
the only living species in the Homo genus of bipedal primates in Hominidae, the great ape family.
Anatomically modern-appearing humans originated in Africa about 200,000 years ago, reaching full
behavioral modernity around 50,000 years ago.[6]
Humans have a highly developed brain, capable of abstract reasoning, language, introspection, and
problem solving. This mental capability, combined with an erect body carriage that frees the hands for
manipulating objects, has allowed humans to make far greater use of tools than any other living species on
Earth. Other higher-level thought processes of humans, such as self-awareness, rationality, and sapience,[7]
[8][9] are considered to be defining features of what constitutes a "person".[10][11]
Like most higher primates, humans are social animals. However, humans are uniquely adept at
utilizing systems of communication for self-expression, the exchange of ideas, and organization. Humans
create complex social structures composed of many cooperating and competing groups, from families to
nations. Social interactions between humans have established an extremely wide variety of values, social
norms, and rituals, which together form the basis of human society. With individuals widespread in every
continent except Antarctica, humans are a cosmopolitan species. As of January 2011[update], the population
of humans was estimated to be about 6.89 billion.[12]
Humans are noted for their desire to understand and influence their environment, seeking to explain
and manipulate phenomena through science, philosophy, mythology, and religion. This natural curiosity has
led to the development of advanced tools and skills, which are passed down culturally; humans are the only
species known to build fires, cook their food, clothe themselves, and use numerous other technologies. The
study of humans is the scientific discipline of anthropology.
Contents
[hide]
• 1 Etymology
• 2 History
• 2.1 Evolution
• 2.2 Paleolithic
• 2.3 Transition to civilization
• 3 Habitat and population
• 4 Biology
• 4.1 Anatomy
• 4.2 Physiology
• 4.3 Genetics
• 4.4 Life cycle
• 4.5 Diet
• 4.6 Sleep
• 5 Psychology
• 5.1 Consciousness and thought
• 5.2 Motivation and emotion
• 6 Society and culture
• 6.1 Sexuality and love
• 6.2 Gender roles
• 6.3 Society, government, and politics
Etymology
Further information: Man (word) and List of alternative names for the human species
The English adjective human is a Middle English loan from Old French humain, ultimately from Latin
hūmānus, the adjective of homō "man". Use as a noun (with a plural humans) dates to the 16th century.[13]
The native English term man is now often reserved for male adults, but can still be used for "mankind" in
general in Modern English.[14] The word is from Proto-Germanic *mannaz, from a Proto-Indo-European
(PIE) root *man-, cognate to Sanskrit manu-.
The generic name Homo is a learned 18th century derivation from Latin homō "man", ultimately
"earthly being" (Old Latin hemō, cognate to Old English guma "man", from PIE *dʰǵʰemon-, meaning 'earth' or
'ground').[15]
History
Evolution
Further information: Anthropology, Homo (genus), and Timeline of human evolution
Plesiadapis
A reconstruction of Australopithecus afarensis, a human ancestor that had developed bipedalism, but
which lacked the large brain of modern humans
The scientific study of human evolution encompasses the development of the genus Homo, but
usually involves studying other hominids and hominines as well, such as Australopithecus. "Modern humans"
are defined as the Homo sapiens species, of which the only extant subspecies is known as Homo sapiens
sapiens. Homo sapiens idaltu (roughly translated as "elder wise human"), the other known subspecies, is
now extinct.[16] Homo neanderthalensis, which became extinct 30,000 years ago, has sometimes been
classified as a subspecies, "Homo sapiens neanderthalensis"; genetic studies now suggest that the
functional DNA of modern humans and Neanderthals diverged 500,000 years ago.[17] Similarly, the few
specimens of Homo rhodesiensis have also occasionally been classified as a subspecies, but this is not
widely accepted. Anatomically modern humans first appear in the fossil record in Africa about 195,000 years
ago, and studies of molecular biology give evidence that the approximate time of divergence from the
common ancestor of all modern human populations was 200,000 years ago.[18][19][20][21][22] The broad
study of African genetic diversity headed by Dr. Sarah Tishkoff found the San people to express the greatest
genetic diversity among the 113 distinct populations sampled, making them one of 14 "ancestral population
clusters". The research also located the origin of modern human migration in south-western Africa, near the
coastal border of Namibia and Angola.[23]
The evolutionary history of the primates can be traced back 65 million years, as one of the oldest of
all surviving placental mammal groups. The oldest known primate-like mammal species (those of the genus
Plesiadapis) come from North America, but they were widespread in Eurasia and Africa during the tropical
conditions of the Paleocene and Eocene. Molecular evidence suggests that the last common ancestor
between humans and the remaining great apes diverged between 8 and 4 million years ago, first the gorillas,
and then the chimpanzees (genus Pan) split off from the line leading to the humans; the functional human
DNA is approximately 98.4% identical to that of chimpanzees when comparing single nucleotide
polymorphisms (see human evolutionary genetics). Therefore, the closest living relatives of humans are
gorillas and chimpanzees, as they share a common ancestor.[24]
Humans are probably most closely related to two chimpanzee species: Common Chimpanzee and
Bonobo.[24] Full genome sequencing has resulted in the conclusion that "after 6.5 [million] years of separate
evolution, the differences between chimpanzee and human are ten times greater than those between two
unrelated people and ten times less than those between rats and mice".[ attribution needed] Suggested
concurrence between the functional human and chimpanzee DNA sequences range between 95% and 99%;
[25][26][27][28] It has been estimated that the human lineage diverged from that of chimpanzees about five
million years ago, and from that of gorillas about eight million years ago. However, a hominid skull discovered
in Chad in 2001, classified as Sahelanthropus tchadensis, is approximately seven million years old, which
may indicate an earlier divergence.[29]
Human evolution is characterised by a number of important morphological, developmental,
physiological and behavioural changes, which have taken place since the split between the last common
ancestor of humans and chimpanzees. The first major morphological change was the evolution of a bipedal
locomotor adaptation from an arboreal or semi-arboreal one,[30] with all its attendant adaptations, such as a
valgus knee, low intermembral index (long legs relative to the arms), and reduced upper-body strength.
Later, ancestral humans developed a much larger brain – typically 1,400 cm³ in modern humans, over
twice the size of that of a chimpanzee or gorilla. The pattern of human postnatal brain growth differs from that
of other apes (heterochrony), and allows for extended periods of social learning and language acquisition in
juvenile humans. Physical anthropologists[who?] argue that the differences between the structure of human
brains and those of other apes are even more significant than their differences in size.
Other significant morphological changes included the evolution of a power and precision grip,[31] a
reduced masticatory system, a reduction of the canine tooth, and the descent of the larynx and hyoid bone,
making speech possible. An important physiological change in humans was the evolution of hidden oestrus,
or concealed ovulation, which may have coincided with the evolution of important behavioural changes, such
as pair bonding. Another significant behavioural change was the development of material culture, with
human-made objects becoming increasingly common and diversified over time. The relationship between all
these changes is the subject of ongoing debate.[32][33]
The forces of natural selection have continued to operate on human populations, with evidence that
certain regions of the genome display directional selection in the past 15,000 years.[34]
Paleolithic
Further information: Recent African Origin, Archaic Homo sapiens, Upper Paleolithic, and Early
human migrations
Artistic expression appeared in the Upper Paleolithic: The Venus of Dolní Věstonice figurine, one of
the earliest known depictions of the human body, dates to approximately 29,000–25,000 BP (Gravettian).
Anatomically modern humans evolved from archaic Homo sapiens in Africa in the Middle Paleolithic,
about 200,000 years ago. By the beginning of the Upper Paleolithic 50,000 BP (Before Present), full
behavioral modernity, including language, music and other cultural universals had developed.
The out of Africa migration is estimated to have occurred about 70,000 years BP. Modern humans
subsequently spread to all continents, replacing earlier hominids: they inhabited Eurasia and Oceania by
40,000 BP, and the Americas at least 14,500 years BP.[35] A popular theory is that they displaced Homo
neanderthalensis and other species descended from Homo erectus[36] (which had inhabited Eurasia as early
as 2 million years ago) through more successful reproduction and competition for resources.[37] The exact
manner or extent of the coexistence and interaction of these species is unknown and remains a controversial
subject.[citation needed]
Evidence from archaeogenetics accumulating since the 1990s has lent strong support to the "out-of-
Africa" scenario, and has marginalized the competing multiregional hypothesis, which proposed that modern
humans evolved, at least in part, from independent hominid populations.[38]
Geneticists Lynn Jorde and Henry Harpending of the University of Utah propose that the variation in
human DNA is minute compared to that of other species. They also propose that during the Late Pleistocene,
the human population was reduced to a small number of breeding pairs – no more than 10,000, and possibly
as few as 1,000 – resulting in a very small residual gene pool. Various reasons for this hypothetical bottleneck
have been postulated, one being the Toba catastrophe theory.[39]
Transition to civilization
Main articles: Neolithic revolution and Cradle of Civilization
Further information: History of the world
The rise of agriculture, and domestication of animals, led to stable human settlements.
The path followed by humans in the course of history
Until c. 10,000 years ago, most humans lived as hunter-gatherers. They generally lived in small
nomadic groups known as band societies. The advent of agriculture prompted the Neolithic Revolution, when
access to food surplus led to the formation of permanent human settlements, the domestication of animals
and the use of metal tools. Agriculture encouraged trade and cooperation, and led to complex society.
Because of the significance of this date for human society, it is the epoch of the Holocene calendar or Human
Era.
About 6,000 years ago, the first proto-states developed in Mesopotamia, Egypt's Nile Valley and the
Indus Valleys. Military forces were formed for protection, and government bureaucracies for administration.
States cooperated and competed for resources, in some cases waging wars. Around 2,000–3,000 years ago,
some states, such as Persia, India, China, Rome, and Greece, developed through conquest into the first
expansive empires. Influential religions, such as Judaism, originating in West Asia, and Hinduism, a religious
tradition that originated in South Asia, also rose to prominence at this time.
The late Middle Ages saw the rise of revolutionary ideas and technologies. In China, an advanced
and urbanized society promoted innovations and sciences, such as printing and seed drilling. In India, major
advancements were made in mathematics, philosophy, religion and metallurgy. The Islamic Golden Age saw
major scientific advancements in Muslim empires. In Europe, the rediscovery of classical learning and
inventions such as the printing press led to the Renaissance in the 14th and 15th centuries. Over the next
500 years, exploration and colonialism brought great parts of the world under European control, leading to
later struggles for independence. The Scientific Revolution in the 17th century and the Industrial Revolution
in the 18th–19th centuries promoted major innovations in transport, such as the railway and automobile;
energy development, such as coal and electricity; and government, such as representative democracy and
Communism.
With the advent of the Information Age at the end of the 20th century, modern humans live in a world
that has become increasingly globalized and interconnected. As of 2010, almost 2 billion humans are able to
communicate with each other via the Internet,[40] and 3.3 billion by mobile phone subscriptions.[41]
Although interconnection between humans has encouraged the growth of science, art, discussion,
and technology, it has also led to culture clashes and the development and use of weapons of mass
destruction. Human civilization has led to environmental destruction and pollution, producing an ongoing
mass extinction of other forms of life called the holocene extinction event,[42] that may be further accelerated
by global warming in the future.[43]
Habitat and population

Further information: Human migration, Demography, and World population
See also: City, Town, Nomad, Camping, Farm, House, Watercraft, Infrastructure, Architecture,
Building, and Engineering
Humans often live in family-based social structures and create artificial shelter.
Early human settlements, were dependent on proximity to water and, depending on the lifestyle,
other natural resources used for subsistence, such as populations of animal prey for huntinging and arable
land for growing crops and grazing livestock. But humans have a great capacity for altering their habitats by
means of technology; through irrigation, urban planning, construction, transport, manufacturing goods,
deforestation and desertification. Deliberate habitat alteration is often done with the goals of increasing
material wealth, increasing thermal comfort, improving the amount of food available, improving aesthetics, or
improving ease of access to resources or other human settlements. With the advent of large-scale trade and
transport infrastructure, proximity to these resources has become unnecessary, and in many places, these
factors are no longer a driving force behind the growth and decline of a population. Nonetheless, the manner
in which a habitat is altered is often a major determinant in population change.
Technology has allowed humans to colonize all of the continents and adapt to virtually all climates.
Within the last century, humans have explored Antarctica, the ocean depths, and outer space, although
large-scale colonization of these environments is not yet feasible. With a population of over six billion,
humans are among the most numerous of the large mammals. Most humans (61%) live in Asia. The
remainder live in the Americas (14%), Africa (14%), Europe (11%), and Oceania (0.5%).
Human habitation within closed ecological systems in hostile environments, such as Antarctica and
outer space, is expensive, typically limited in duration, and restricted to scientific, military, or industrial
expeditions. Life in space has been very sporadic, with no more than thirteen humans in space at any given
time. Between 1969 and 1972, two humans at a time spent brief intervals on the Moon. As of February 2011,
no other celestial body has been visited by humans, although there has been a continuous human presence
in space since the launch of the initial crew to inhabit the International Space Station on October 31, 2000.
However, other celestial bodies have been visited by human-made objects.
Since 1800, the human population has increased from one billion to over six billion.[44] In 2004,
some 2.5 billion out of 6.3 billion people (39.7%) lived in urban areas, and this percentage is expected to
continue to rise throughout the 21st century. In February 2008, the U.N. estimated that half the world's
population will live in urban areas by the end of the year.[45] Problems for humans living in cities include
various forms of pollution and crime,[46] especially in inner city and suburban slums.
Humans have had a dramatic effect on the environment. As humans are rarely preyed upon, they
have been described as superpredators.[47] Currently, through land development, combustion of fossil fuels
and pollution, humans are thought to be the main contributor to global climate change.[48] Human activity is
believed to be a major contributor to the ongoing Holocene extinction event, which is a form of mass
extinction. If this continues at its current rate it is predicted that it will wipe out half of all species over the next
century.[49][50]
Biology
Main article: Human biology
Anatomy
Main article: Human anatomy
Further information: Human physical appearance and Anatomically modern humans
Basic anatomical features of female and male humans. Note that these models have had body hair
and male facial hair removed and head hair trimmed.
Human body types vary substantially. Although body size is largely determined by genes, it is also
significantly influenced by environmental factors such as diet and exercise. The average height of an adult
human is about 1.5 to 1.8 m (5 to 6 feet) tall, although this varies significantly from place to place and
depending on ethnic origin.[51][52] The average mass of an adult human is 54–64 kg (120–140 lbs) for
females and 76–83 kg (168–183 lbs) for males.[53] Weight can also vary greatly (e.g. obesity). Unlike most
other primates, humans are capable of fully bipedal locomotion, thus leaving their arms available for
manipulating objects using their hands, aided especially by opposable thumbs.
Vitruvian Man, Leonardo da Vinci's image is often used as an implied symbol of the essential
symmetry of the human body, and by extension, of the universe as a whole.
Although humans appear hairless compared to other primates, with notable hair growth occurring
chiefly on the top of the head, underarms and pubic area, the average human has more hair follicles on his or
her body than the average chimpanzee. The main distinction is that human hairs are shorter, finer, and less
heavily pigmented than the average chimpanzee's, thus making them harder to see.[54]
The hue of human skin and hair is determined by the presence of pigments called melanins. Human
skin hues can range from very dark brown to very pale pink. Human hair ranges from white to brown to red to
most commonly black.[55] This depends on the amount of melanin (an effective sun blocking pigment) in the
skin and hair, with hair melanin concentrations in hair fading with increased age, leading to grey or even
white hair. Most researchers believe that skin darkening was an adaptation that evolved as a protection
against ultraviolet solar radiation. However, more recently it has been argued that particular skin colors are
an adaptation to balance folate, which is destroyed by ultraviolet radiation, and vitamin D, which requires
sunlight to form.[56] The skin pigmentation of contemporary humans is geographically stratified, and in
general correlates with the level of ultraviolet radiation. Human skin also has a capacity to darken (sun
tanning) in response to exposure to ultraviolet radiation.[57][58] Humans tend to be physically weaker than
other similarly sized primates, with young, conditioned male humans having been shown to be unable to
match the strength of female orangutans which are at least three times stronger.[59]
The construction of the human pelvis differs from other primates, as do the toes. As a result, humans
are slower for short distances than most other animals, but are among the best long-distance runners in the
animal kingdom.[60] Humans' thinner body hair and more productive sweat glands also helps avoid heat
exhaustion while running for long distances. For this reason persistence hunting was most likely a very
successful strategy for early humans – in this method, prey is chased until it is literally exhausted. This may
have also helped the early human Cro-Magnon population out-compete the Neanderthal population for food.
The otherwise physically stronger Neanderthal would have much greater difficulty hunting in this way, and
much more likely hunted larger game in close quarters. A trade-off for these advantages of the modern
human pelvis is that childbirth is more difficult and dangerous.
The construction of modern human shoulders enable throwing weapons, which also were much more
difficult or even impossible for Neanderthal competitors to use effectively.[61]
Constituents of the human body in a person weighing 60 kg[62]
Constituent Weight Percentage of atoms
Oxygen 38.8 kg 25.5 %
Carbon 10.9 kg 9.5 %
Hydrogen 6.0 kg 63.0 %
Nitrogen 1.9 kg 1.4 %
Other 2.4 kg 0.6 %

Humans have proportionately shorter palates and much smaller teeth than other primates. They are
the only primates to have short, relatively flush canine teeth. Humans have characteristically crowded teeth,
with gaps from lost teeth usually closing up quickly in young specimens. Humans are gradually losing their
wisdom teeth, with some individuals having them congenitally absent.[63]
Physiology
Main article: Human physiology
Human physiology is the science of the mechanical, physical, and biochemical functions of humans
in good health, their organs, and the cells of which they are composed. The principal level of focus of
physiology is at the level of organs and systems. Most aspects of human physiology are closely homologous
to corresponding aspects of animal physiology, and animal experimentation has provided much of the
foundation of physiological knowledge. Anatomy and physiology are closely related fields of study: anatomy,
the study of form, and physiology, the study of function, are intrinsically tied and are studied in tandem as
part of a medical curriculum.
Genetics
Main article: Human genetics
Humans are a eukaryotic species. Each diploid cell has two sets of 23 chromosomes, each set
received from one parent. There are 22 pairs of autosomes and one pair of sex chromosomes. By present
estimates, humans have approximately 20,000–25,000 genes. Like other mammals, humans have an XY sex-
determination system, so that females have the sex chromosomes XX and males have XY. The X
chromosome carries many genes not on the Y chromosome, which means that recessive diseases
associated with X-linked genes, such as haemophilia, affect men more often than women.
Life cycle
A 10 mm human embryo at 5 weeks

The human life cycle is similar to that of other placental mammals. The zygote divides inside the
female's uterus to become an embryo, which over a period of thirty-eight weeks (9 months) of gestation
becomes a fetus. After this span of time, the fully grown fetus is birthed from the woman's body and breathes
independently as an infant for the first time. At this point, most modern cultures recognize the baby as a
person entitled to the full protection of the law, though some jurisdictions extend various levels of personhood
earlier to human fetuses while they remain in the uterus.
Compared with other species, human childbirth is dangerous. Painful labors lasting twenty-four hours
or more are not uncommon and sometimes leads to the death of the mother, or the child.[64] This is because
of both the relatively large fetal head circumference (for housing the brain) and the mother's relatively narrow
pelvis (a trait required for successful bipedalism, by way of natural selection).[65][66] The chances of a
successful labor increased significantly during the 20th century in wealthier countries with the advent of new
medical technologies. In contrast, pregnancy and natural childbirth remain hazardous ordeals in developing
regions of the world, with maternal death rates approximately 100 times more common than in developed
countries.[67]
In developed countries, infants are typically 3–4 kg (6–9 pounds) in weight and 50–60 cm (20–
24 inches) in height at birth.[68] However, low birth weight is common in developing countries, and
contributes to the high levels of infant mortality in these regions.[69] Helpless at birth, humans continue to
grow for some years, typically reaching sexual maturity at 12 to 15 years of age. Females continue to
develop physically until around the age of 18, whereas male development continues until around age 21. The
human life span can be split into a number of stages: infancy, childhood, adolescence, young adulthood,
adulthood and old age. The lengths of these stages, however, have varied across cultures and time periods.
Compared to other primates, humans experience an unusually rapid growth spurt during adolescence, where
the body grows 25% in size. Chimpanzees, for example, grow only 14%, with no pronounced spurt.[70] The
presence of the growth spurt is probably necessary to keep children physically small until they are
psychologically mature. Humans are one of the few species in which females undergo menopause. It has
been proposed that menopause increases a woman's overall reproductive success by allowing her to invest
more time and resources in her existing offspring and/or their children (the grandmother hypothesis), rather
than by continuing to bear children into old age.[71][72]
There are significant differences in life expectancy around the world. The developed world is
generally aging, with the median age around 40 years. In the developing world the median age is between 15
and 20 years. Life expectancy at birth in Hong Kong is 84.8 years for a female and 78.9 for a male, while in
Swaziland, primarily because of AIDS, it is 31.3 years for both sexes.[73] While one in five Europeans is 60
years of age or older, only one in twenty Africans is 60 years of age or older.[74] The number of centenarians
(humans of age 100 years or older) in the world was estimated by the United Nations at 210,000 in 2002.[75]
At least one person, Jeanne Calment, is known to have reached the age of 122 years; higher ages have
been claimed but they are not well substantiated. Worldwide, there are 81 men aged 60 or older for every
100 women of that age group, and among the oldest, there are 53 men for every 100 women.
A selection of different humans at various stages of the human life cycle
Girl (before puberty) Woman of reproductive age Older woman (after menopause)
Boy (before puberty)

Adult man Elderly man
Diet
Main article: Human nutrition
Humans are omnivorous, capable of consuming plant, animal, and inorganic material.[76][77]
Varying with available food sources in regions of habitation, and also varying with cultural and religious
norms, human groups have adopted a range of diets, from purely vegetarian to primarily carnivorous. In
some cases, dietary restrictions in humans can lead to deficiency diseases; however, stable human groups
have adapted to many dietary patterns through both genetic specialization and cultural conventions to use
nutritionally balanced food sources.[78] The human diet is prominently reflected in human culture, and has
led to the development of food science.
Until the development of agriculture approximately 10,000 years ago, Homo sapiens employed a
hunter-gatherer method as their sole means of food collection. This involved combining stationary food
sources (such as fruits, grains, tubers, and mushrooms, insect larvae and aquatic molluscs) with wild game,
which must be hunted and killed in order to be consumed.[79] It has been proposed that members of H.
sapiens have used fire to prepare and cook food since the time of their divergence from Homo rhodesiensis
(which itself had previously speciated from Homo erectus).[80] Around ten thousand years ago, humans
developed agriculture,[81] which substantially altered their diet. This change in diet may also have altered
human biology; with the spread of dairy farming providing a new and rich source of food, leading to the
evolution of the ability to digest lactose in some adults.[82][83] Agriculture led to increased populations, the
development of cities, and because of increased population density, the wider spread of infectious diseases.
The types of food consumed, and the way in which they are prepared, has varied widely by time, location,
and culture.
In general, humans can survive for two to eight weeks without food, depending on stored body fat.
Survival without water is usually limited to three or four days. About 36 million humans starve to death every
year, due to lack of edible materials in their habitats.[84] Childhood malnutrition is also common and
contributes to the global burden of disease.[85] However global food distribution is not even, and obesity
among some human populations has increased rapidly, leading to health complications and increased
mortality in some developed, and a few developing countries. Worldwide over one billion people are obese,
[86] while in the United States 35% of people are obese, leading to this being described as an "obesity
epidemic".[87] Obesity is caused by consuming more calories than are expended, so excessive weight gain
is usually caused by a combination of an energy-dense high fat diet and insufficient exercise.[86]
Sleep
Main article: Sleep
Humans are generally diurnal. The average sleep requirement is between seven and nine continuous
hours a day for an adult and nine to ten hours for a child; elderly people usually sleep for six to seven hours.
Experiencing less sleep than this is common in modern societies; this sleep deprivation can have negative
effects. A sustained restriction of adult sleep to four hours per day has been shown to correlate with changes
in physiology and mental state, including fatigue, aggression, and bodily discomfort.
Psychology
A sketch of the human brain imposed upon the profile of Michelangelo's David
Main article: Psychology
Further information: Human brain and Mind
The human brain, the focal point of the central nervous system in humans, controls the peripheral
nervous system. In addition to controlling "lower", involuntary, or primarily autonomic activities such as
respiration and digestion, it is also the locus of "higher" order functioning such as thought, reasoning, and
abstraction.[88] These cognitive processes constitute the mind, and, along with their behavioral
consequences, are studied in the field of psychology.
Generally regarded as more capable of these higher order activities, the human brain is believed to
be more "intelligent" in general than that of any other known species. Some are capable of creating
structures and using simple tools—mostly through instinct and mimicry—human technology is vastly more
complex, and is constantly evolving and improving through time.
Although being vastly more advanced than many species in cognitive abilities, most of these abilities
are known in primitive form among other species. Modern anthropology has tended to bear out Darwin's
proposition that "the difference in mind between man and the higher animals, great as it is, certainly is one of
degree and not of kind".[89]
Consciousness and thought

Main articles: Consciousness and Cognition
Humans are one of only nine species to pass the mirror test—which tests whether an animal
recognizes its reflection as an image of itself—along with all the great apes (gorillas, chimpanzees,
orangutans, bonobos), Bottlenose dolphins, Asian elephants, European Magpies, and Orcas.[90] Most
human children will pass the mirror test at 18 months old.[91] However, the usefulness of this test as a true
test of consciousness has been disputed, and this may be a matter of degree rather than a sharp divide.
Monkeys have been trained to apply abstract rules in tasks.[92]
The human brain perceives the external world through the senses, and each individual human is
influenced greatly by his or her experiences, leading to subjective views of existence and the passage of
time. Humans are variously said to possess consciousness, self-awareness, and a mind, which correspond
roughly to the mental processes of thought. These are said to possess qualities such as self-awareness,
sentience, sapience, and the ability to perceive the relationship between oneself and one's environment. The
extent to which the mind constructs or experiences the outer world is a matter of debate, as are the
definitions and validity of many of the terms used above. The philosopher of cognitive science Daniel
Dennett, for example, argues that there is no such thing as a narrative centre called the "mind", but that
instead there is simply a collection of sensory inputs and outputs: different kinds of "software" running in
parallel.[93] Psychologist B.F. Skinner argued that the mind is an explanatory fiction that diverts attention
from environmental causes of behavior,[94] and that what are commonly seen as mental processes may be
better conceived of as forms of covert verbal behavior.[95][96]
Humans study the more physical aspects of the mind and brain, and by extension of the nervous
system, in the field of neurology, the more behavioral in the field of psychology, and a sometimes loosely
defined area between in the field of psychiatry, which treats mental illness and behavioral disorders.
Psychology does not necessarily refer to the brain or nervous system, and can be framed purely in terms of
phenomenological or information processing theories of the mind. Increasingly, however, an understanding
of brain functions is being included in psychological theory and practice, particularly in areas such as artificial
intelligence, neuropsychology, and cognitive neuroscience.
The nature of thought is central to psychology and related fields. Cognitive psychology studies
cognition, the mental processes' underlying behavior. It uses information processing as a framework for
understanding the mind. Perception, learning, problem solving, memory, attention, language and emotion are
all well researched areas as well. Cognitive psychology is associated with a school of thought known as
cognitivism, whose adherents argue for an information processing model of mental function, informed by
positivism and experimental psychology. Techniques and models from cognitive psychology are widely
applied and form the mainstay of psychological theories in many areas of both research and applied
psychology. Largely focusing on the development of the human mind through the life span, developmental
psychology seeks to understand how people come to perceive, understand, and act within the world and how
these processes change as they age. This may focus on intellectual, cognitive, neural, social, or moral
development.
Some philosophers divide consciousness into phenomenal consciousness, which is experience itself,
and access consciousness, which is the processing of the things in experience.[97] Phenomenal
consciousness is the state of being conscious, such as when they say "I am conscious." Access
consciousness is being conscious of something in relation to abstract concepts, such as when one says "I
am conscious of these words." Various forms of access consciousness include awareness, self-awareness,
conscience, stream of consciousness, Husserl's phenomenology, and intentionality. The concept of
phenomenal consciousness, in modern history, according to some, is closely related to the concept of qualia.
Social psychology links sociology with psychology in their shared study of the nature and causes of human
social interaction, with an emphasis on how people think towards each other and how they relate to each
other. The behavior and mental processes, both human and non-human, can be described through animal
cognition, ethology, evolutionary psychology, and comparative psychology as well. Human ecology is an
academic discipline that investigates how humans and human societies interact with both their natural
environment and the human social environment.
Motivation and emotion

Main articles: Motivation and Emotion
Motivation is the driving force of desire behind all deliberate actions of humans. Motivation is based
on emotion—specifically, on the search for satisfaction (positive emotional experiences), and the avoidance of
conflict. Positive and negative is defined by the individual brain state, which may be influenced by social
norms: a person may be driven to self-injury or violence because their brain is conditioned to create a
positive response to these actions. Motivation is important because it is involved in the performance of all
learned responses. Within psychology, conflict avoidance and the libido are seen to be primary motivators.
Within economics, motivation is often seen to be based on incentives; these may be financial, moral, or
coercive. Religions generally posit divine or demonic influences.
Happiness, or the state of being happy, is a human emotional condition. The definition of happiness
is a common philosophical topic. Some people might define it as the best condition that a human can have—a
condition of mental and physical health. Others define it as freedom from want and distress; consciousness
of the good order of things; assurance of one's place in the universe or society.
Emotion has a significant influence on, or can even be said to control, human behavior, though
historically many cultures and philosophers have for various reasons discouraged allowing this influence to
go unchecked. Emotional experiences perceived as pleasant, such as love, admiration, or joy, contrast with
those perceived as unpleasant, like hate, envy, or sorrow. There is often a distinction made between refined
emotions that are socially learned and survival oriented emotions, which are thought to be innate. Human
exploration of emotions as separate from other neurological phenomena is worthy of note, particularly in
cultures where emotion is considered separate from physiological state. In some cultural medical theories
emotion is considered so synonymous with certain forms of physical health that no difference is thought to
exist. The Stoics believed excessive emotion was harmful, while some Sufi teachers felt certain extreme
emotions could yield a conceptual perfection, what is often translated as ecstasy.
In modern scientific thought, certain refined emotions are considered a complex neural trait innate in
a variety of domesticated and non-domesticated mammals. These were commonly developed in reaction to
superior survival mechanisms and intelligent interaction with each other and the environment; as such,
refined emotion is not in all cases as discrete and separate from natural neural function as was once
assumed. However, when humans function in civilized tandem, it has been noted that uninhibited acting on
extreme emotion can lead to social disorder and crime.
Society and culture

Human society statistics
World population 6.9 billion
12.7 per km² (4.9 mi²) by total area

Population density
43.6 per km² (16.8 mi²) by land area
Largest Beijing, Bogotá, Buenos Aires, Cairo, Delhi, Dhaka, Guangzhou,

agglomerations Istanbul, Jakarta, Karachi, Kinshasa, Kolkata, Lagos, Lima, London, Los
Angeles, Manila, Mexico City, Moscow, Mumbai, New York City, Osaka, Paris,
Rio de Janeiro, São Paulo, Seoul, Shanghai, Shenzen, Tehran, Tianjin,
Tokyo, Wuhan
Most widely spoken Mandarin Chinese, English, Spanish, Arabic, Hindi, Bengali,
languages[98] Portuguese, Russian, Japanese, German, Javanese, Punjabi, Telugu,
Vietnamese, Marathi, French, Korean, Tamil, Italian, Urdu, Marathi,
Indonesian
Most popular Christianity, Islam, Hinduism, Buddhism, Sikhism, Judaism, Baha'i

religions[99]
$36,356,240 million USD

GDP (nominal)
($5,797 USD per capita)
$51,656,251 million IND

GDP (PPP)
($8,236 per capita)
Main articles: Culture and Society

Humans are social beings. In comparisons with animalia, humans are regarded like the primates for
their social qualities. But beyond any other creature, humans are adept at utilizing systems of communication
for self-expression, the exchange of ideas, and organization, and as such have created complex social
structures composed of many cooperating and competing groups. Human groups range from families to
nations. Social interactions between humans have established an extremely wide variety of values, social
norms, and rituals, which together form the basis of human society.
Culture is defined here as patterns of complex symbolic behavior, i.e. all behavior that is not innate
but which has to be learned through social interaction with others; such as the use of distinctive material and
symbolic systems, including language, ritual, social organization, traditions, beliefs and technology.
Sexuality and love

Main articles: Love and Human sexuality
Human sexuality, besides ensuring biological reproduction, has important social functions: it creates
physical intimacy, bonds, and hierarchies among individuals; and in a hedonistic sense to the enjoyment of
activity involving sexual gratification. Sexual desire, or libido, is experienced as a bodily urge, often
accompanied by strong emotions such as love, ecstasy and jealousy. The extreme importance of sexuality in
the human species can be seen in a number of physical features, among them hidden ovulation, the
evolution of external scrotum and penis suggesting sperm competition, the absence of an os penis,
permanent secondary sexual characteristics, the forming of pair bonds based on sexual attraction as a
common social structure and sexual ability in females outside of ovulation. These adaptations indicate that
the importance of sexuality in humans is on a par with that found in the Bonobo, and that the complex human
sexual behaviour has a long evolutionary history.
Human choices in acting on sexuality are commonly influenced by cultural norms, which vary widely.
Restrictions are often determined by religious beliefs or social customs. The pioneering researcher Sigmund
Freud believed that humans are born polymorphously perverse, which means that any number of objects
could be a source of pleasure. According to Freud, humans then pass through five stages of psychosexual
development (and can fixate on any stage because of various traumas during the process). For Alfred
Kinsey, another influential sex researcher, people can fall anywhere along a continuous scale of sexual
orientation (with only small minorities fully heterosexual or homosexual). Recent studies of neurology and
genetics suggest people may be born predisposed to various sexual tendencies.[100][101]
Gender roles
Main article: Gender role
See also: Gender
The sexual division of humans into male and female has been marked culturally by a corresponding
division of roles, norms, practices, dress, behavior, rights, duties, privileges, status, and power. Cultural
differences by gender have often been believed to have arisen naturally out of a division of reproductive
labor; the biological fact that women give birth led to their further cultural responsibility for nurturing and
caring for children and households. Gender roles have varied historically, and challenges to predominant
gender norms have recurred in many societies. As a whole, patriarchal societies (i.e., in which men hold the
greater degree of economic and political power) have been predominant, and matriarchal or egalitarian
societies less common. [102][103][104][105]
Society, government, and politics
The United Nations complex in New York City, which houses one of the largest political organizations
in the world.
Main articles: Society, Government, Politics, and State (polity)
Society is the system of organizations and institutions arising from interaction between humans. A
state is an organized political community occupying a definite territory, having an organized government, and
possessing internal and external sovereignty. Recognition of the state's claim to independence by other
states, enabling it to enter into international agreements, is often important to the establishment of its
statehood. The "state" can also be defined in terms of domestic conditions, specifically, as conceptualized by
Max Weber, "a state is a human community that (successfully) claims the monopoly of the 'legitimate' use of
physical force within a given territory."[106]
Government can be defined as the political means of creating and enforcing laws; typically via a
bureaucratic hierarchy. Politics is the process by which decisions are made within groups; this process often
involves conflict as well as compromise. Although the term is generally applied to behavior within
governments, politics is also observed in all human group interactions, including corporate, academic, and
religious institutions. Many different political systems exist, as do many different ways of understanding them,
and many definitions overlap. Examples of governments include monarchy, Communist state, military
dictatorship, theocracy, and liberal democracy, the last of which is considered dominant today. All of these
issues have a direct relationship with economics.
Trade and economics

Main articles: Trade and Economics
Buyers and sellers bargain in a market.
Trade is the voluntary exchange of goods and services, and is a form of economics. A mechanism
that allows trade is called a market. The original form of trade was barter, the direct exchange of goods and
services. Modern traders instead generally negotiate through a medium of exchange, such as money. As a
result, buying can be separated from selling, or earning. The invention of money (and later credit, paper
money and non-physical money) greatly simplified and promoted trade. Because of specialization and
division of labor, most people concentrate on a small aspect of manufacturing or service, trading their labour
for products. Trade exists between regions because different regions have an absolute or comparative
advantage in the production of some tradable commodity, or because different regions' size allows for the
benefits of mass production.
Economics is a social science which studies the production, distribution, trade, and consumption of
goods and services. Economics focuses on measurable variables, and is broadly divided into two main
branches: microeconomics, which deals with individual agents, such as households and businesses, and
macroeconomics, which considers the economy as a whole, in which case it considers aggregate supply and
demand for money, capital and commodities. Aspects receiving particular attention in economics are
resource allocation, production, distribution, trade, and competition. Economic logic is increasingly applied to
any problem that involves choice under scarcity or determining economic value. Mainstream economics
focuses on how prices reflect supply and demand, and uses equations to predict consequences of decisions.
Race and ethnicity

Main articles: Race (classification of human beings) and Ethnic group
See also: Race and genetics and Historical definitions of race
A collection of mugshots showing multiple races
Humans often categorize themselves in terms of race or ethnicity, sometimes on the basis of
differences in appearance. Human racial categories have been based on both ancestry and visible traits,
especially facial features, skin color and hair texture. Most current genetic and archaeological evidence
supports a recent single origin of modern humans in East Africa.[107] Current genetic studies have
demonstrated that humans on the African continent are most genetically diverse.[108] However, compared to
the other great apes, human gene sequences are remarkably homogeneous.[109][110][111][112] The
predominance of genetic variation occurs within racial groups, with only 5 to 15% of total variation occurring
between groups.[113] Thus the scientific concept of variation in the human genome is largely incongruent
with the cultural concept of ethnicity or race. Ethnic groups are defined by linguistic, cultural, ancestral,
national or regional ties. Self-identification with an ethnic group is usually based on kinship and descent.
Race and ethnicity are among major factors in social identity giving rise to various forms of identity politics,
e.g., racism. (Humans have turned to violence and social exclusion as part of their identity politics; a side
effect is name-calling, which has produced numerous offensive terms for humans.)
There is no scientific consensus of a list of the human races, and few anthropologists endorse the
notion of human "race".[114] For example, a color terminology for race includes the following in a
classification of human races: Black (e.g. Sub-Saharan Africa), Red (e.g. Native Americans), Yellow (e.g.
East Asians) and White (e.g. Europeans).
Referring to natural species, in general, the term "race" is obsolete, particularly if a species is
uniformly distributed on a territory. In its modern scientific connotation, the term is not applicable to a species
as genetically homogeneous as the human one, as stated in the declaration on race (UNESCO 1950).[115]
Genetic studies have substantiated the absence of clear biological borders, thus the term "race" is rarely
used in scientific terminology, both in biological anthropology and in human genetics.[ citation needed] What
in the past had been defined as "races"—e.g., whites, blacks, or Asians—are now defined as "ethnic groups" or
"populations", in correlation with the field (sociology, anthropology, genetics) in which they are considered.
[citation needed]
War
Main article: War
The atomic bombings of Hiroshima and Nagasaki immediately killed over 120,000 humans.
War is a state of widespread conflict between states or other large groups of humans, which is
characterized by the use of lethal violence between combatants and/or upon civilians. (Humans also engage
in lesser conflicts, such as brawls, riots, revolts, and melees. A revolution may or may not involve warfare.) It
is estimated that during the 20th century between 167 and 188 million humans died as a result of war.[116] A
common perception of war is a series of military campaigns between at least two opposing sides involving a
dispute over sovereignty, territory, resources, religion, or other issues. A war between internal elements of a
state is a civil war.
There have been a wide variety of rapidly advancing tactics throughout the history of war, ranging
from conventional war to asymmetric warfare to total war and unconventional warfare. Techniques include
hand to hand combat, the use of ranged weapons, Naval warfare, and, more recently, air support. Military
intelligence has often played a key role in determining victory and defeat. Propaganda, which often includes
information, slanted opinion and disinformation, plays a key role in maintaining unity within a warring group,
and/or sowing discord among opponents. In modern warfare, soldiers and armoured fighting vehicles are
used to control the land, warships the sea, and aircraft the sky. These fields have also overlapped in the
forms of marines, paratroopers, naval aircraft carriers, and surface-to-air missiles, among others. Satellites in
low Earth orbit have made outer space a factor in warfare as well as it is used for detailed intelligence
gathering, however no known aggressive actions have been taken from space.
Material culture and technology

Main articles: Tool and Technology
An archaic Acheulean stone tool
Stone tools were used by proto-humans at least 2.5 million years ago.[117] The controlled use of fire
began around 1.5 million years ago. Since then, humans have made major advances, developing complex
technology to create tools to aid their lives and allowing for other advancements in culture. Major leaps in
technology include the discovery of agriculture – what is known as the Neolithic Revolution; and the invention
of automated machines in the Industrial Revolution.
Archaeology attempts to tell the story of past or lost cultures in part by close examination of the
artifacts they produced. Early humans left stone tools, pottery, and jewelry that are particular to various
regions and times.
Language
Main article: Language
The capacity humans have to transfer concepts, ideas and notions through speech and writing is
unrivaled in known species. Unlike the call systems of other primates that are closed, human language is far
more open, and gains variety in different situations. The human language has the quality of displacement,
using words to represent things and happenings that are not presently or locally occurring, but elsewhere or
at a different time.[63] In this way data networks are important to the continuing development of language.
The faculty of speech is a defining feature of humanity, possibly predating phylogenetic separation of the
modern population. Language is central to the communication between humans, as well as being central to
the sense of identity that unites nations, cultures and ethnic groups. The invention of writing systems at least
5,000 years ago allowed the preservation of language on material objects, and was a major step in cultural
evolution. The science of linguistics describes the structure of language and the relationship between
languages. There are approximately 6,000 different languages currently in use, including sign languages,
and many thousands more that are considered extinct.
Spirituality and religion

Main articles: Spirituality and Religion
Religion is generally defined as a belief system concerning the supernatural, sacred or divine, and
practices, values, institutions and rituals associated with such belief. Some religions also have a moral code.
The evolution and the history of the first religions have recently become areas of active scientific
investigation.[118][119][120] However, in the course of its development, religion has taken on many forms
that vary by culture and individual perspective. Some of the chief questions and issues religions are
concerned with include life after death (commonly involving belief in an afterlife), the origin of life, the nature
of the universe (religious cosmology) and its ultimate fate (eschatology), and what is moral or immoral. A
common source in religions for answers to these questions are beliefs in transcendent divine beings such as
deities or a singular God, although not all religions are theistic—many are nontheistic or ambiguous on the
topic, particularly among the Eastern religions. Spirituality, belief or involvement in matters of the soul or
spirit, is one of the many different approaches humans take in trying to answer fundamental questions about
humankind's place in the universe, the meaning of life, and the ideal way to live one's life. Though these
topics have also been addressed by philosophy, and to some extent by science, spirituality is unique in that it
focuses on mystical or supernatural concepts such as karma and God.
Although the exact level of religiosity can be hard to measure,[121] a majority of humans professes
some variety of religious or spiritual belief, although some are irreligious: that is lacking or rejecting belief in
the supernatural or spiritual. Other humans have no religious beliefs and are atheists, scientific skeptics,
agnostics or simply non-religious. Humanism is a philosophy which seeks to include all of humanity and all
issues common to humans; it is usually non-religious. Additionally, although most religions and spiritual
beliefs are clearly distinct from science on both a philosophical and methodological level, the two are not
generally considered mutually exclusive; a majority of humans holds a mix of both scientific and religious
views. The distinction between philosophy and religion, on the other hand, is at times less clear, and the two
are linked in such fields as the philosophy of religion and theology.
Philosophy and self-reflection

Main articles: Philosophy and Human self-reflection
See also: Human nature
Statue of Confucius on Chongming Island in Shanghai
Philosophy is a discipline or field of study involving the investigation, analysis, and development of
ideas at a general, abstract, or fundamental level. It is the discipline searching for a general understanding of
reality, reasoning and values. Major fields of philosophy include logic, metaphysics, epistemology,
philosophy of mind, and axiology (which includes ethics and aesthetics). Philosophy covers a very wide
range of approaches, and is used to refer to a worldview, to a perspective on an issue, or to the positions
argued for by a particular philosopher or school of philosophy.
Art, music, and literature

Main articles: Art, Music, and Literature
Allegory of Music (ca. 1594), a painting of a woman writing sheet music by Lorenzo Lippi
Artistic works have existed for almost as long as humankind, from early pre-historic art to
contemporary art. Art is one of the most unusual aspects of human behaviour and a key distinguishing
feature of humans from other species.
As a form of cultural expression by humans, art may be defined by the pursuit of diversity and the
usage of narratives of liberation and exploration (i.e. art history, art criticism, and art theory) to mediate its
boundaries. This distinction may be applied to objects or performances, current or historical, and its prestige
extends to those who made, found, exhibit, or own them. In the modern use of the word, art is commonly
understood to be the process or result of making material works that, from concept to creation, adhere to the
"creative impulse" of human beings. Art is distinguished from other works by being in large part unprompted
by necessity, by biological drive, or by any undisciplined pursuit of recreation.
Music is a natural intuitive phenomenon based on the three distinct and interrelated organization
structures of rhythm, harmony, and melody. Listening to music is perhaps the most common and universal
form of entertainment for humans, while learning and understanding it are popular disciplines. There are a
wide variety of music genres and ethnic musics. Literature, the body of written—and possibly oral—works,
especially creative ones, includes prose, poetry and drama, both fiction and non-fiction. Literature includes
such genres as epic, legend, myth, ballad, and folklore.
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2007. ISBN 0-13-227584-8 pages 757–761.
External links
Find more about Humans on Wikipedia's sister projects:

Wikispecies has information related to: Homo sapiens sapiens
Wikimedia Commons has media related to: Homo sapiens
Mammals portal
• MNSU
• Archaeology Info
• Chororapithecus abyssinicus Possible human-orangutan split 20 million years ago. (Aug 26
2007)
• Homo sapiens - The Smithsonian Institution's Human Origins Program
• Homo sapiens Linnaeus, 1758 at the Encyclopedia of Life
Hominini
Australopithecines


[hide]v · d · eExtant species of family Hominidae (Great apes)
Kingdom: Animalia · Phylum: Chordata · Class: Mammalia · Order: Primates · Suborder: Haplorrhini
Ponginae Pongo Bornean Orangutan (P. pygmaeus) · Sumatran

(Orangutans) Orangutan (P. abelii)
Western Gorilla (G. gorilla) · Eastern Gorilla (G.

Gorilla
beringei)
Homininae Pan Common Chimpanzee (P. troglodytes) · Bonobo (P.

(Chimpanzees) paniscus)
Homo Human (H. s. sapiens)
Category




Hominid
Retrieved from "http://en.wikipedia.org/wiki/Human"

W000
Microorganism
"Microbe" redirects here. For other uses, see Microbe (disambiguation).

A cluster of Escherichia coli Bacteria magnified 10,000 times.
A microorganism (from the Greek: μικρός, mikrós, "small" and ὀργανισμός, organismós, "organism";
also spelt micro-organism, micro organism or microörganism) or microbe is an organism that is unicellular or
lives in a colony of cellular organisms. The study of microorganisms is called microbiology, a subject that
began with Anton van Leeuwenhoek's discovery of microorganisms in 1675, using a microscope of his own
design.
Microorganisms are very diverse; they include bacteria, fungi, archaea, and protists; microscopic
plants (green algae); and animals such as plankton and the planarian. Some microbiologists also include
viruses, but others consider these as non-living.[1][2] Most microorganisms are unicellular (single-celled), but
this is not universal, since some multicellular organisms are microscopic, while some unicellular protists and
bacteria, like Thiomargarita namibiensis, are macroscopic and visible to the naked eye.[3]
Microorganisms live in all parts of the biosphere where there is liquid water, including soil, hot
springs, on the ocean floor, high in the atmosphere and deep inside rocks within the Earth's crust.
Microorganisms are critical to nutrient recycling in ecosystems as they act as decomposers. As some
microorganisms can fix nitrogen, they are a vital part of the nitrogen cycle, and recent studies indicate that
airborne microbes may play a role in precipitation and weather.[4]
Microbes are also exploited by people in biotechnology, both in traditional food and beverage
preparation, and in modern technologies based on genetic engineering. However, pathogenic microbes are
harmful, since they invade and grow within other organisms, causing diseases that kill people, other animals
and plants.[5]
Contents
[hide]
• 1 History
• 1.1 Evolution
• 1.2 Pre-microbiology
• 1.3 History of microorganisms' discovery
• 2 Classification and structure
• 2.1 Prokaryotes
• 2.1.1 Bacteria
• 2.1.2 Archaea
• 2.2 Eukaryotes
• 2.2.1 Protists
• 2.2.2 Animals
• 2.2.3 Fungi
• 2.2.4 Plants
• 3 Habitats and ecology
• 3.1 Extremophiles
• 3.2 Soil microbes
• 3.3 Symbiotic microbes
• 4 Importance
• 4.1 Use in food
• 4.2 Use in water treatment
History
Evolution
Further information: Timeline of evolution
Single-celled microorganisms were the first forms of life to develop on Earth, approximately 3–4
billion years ago.[6][7][8] Further evolution was slow,[9] and for about 3 billion years in the Precambrian eon,
all organisms were microscopic.[10] So, for most of the history of life on Earth the only forms of life were
microorganisms.[11] Bacteria, algae and fungi have been identified in amber that is 220 million years old,
which shows that the morphology of microorganisms has changed little since the Triassic period.[12]
Most microorganisms can reproduce rapidly and microbes such as bacteria can also freely exchange
genes by conjugation, transformation and transduction between widely-divergent species.[13] This horizontal
gene transfer, coupled with a high mutation rate and many other means of genetic variation, allows
microorganisms to swiftly evolve (via natural selection) to survive in new environments and respond to
environmental stresses. This rapid evolution is important in medicine, as it has led to the recent development
of 'super-bugs' — pathogenic bacteria that are resistant to modern antibiotics.[14]
Pre-microbiology
The possibility that microorganisms exist was discussed for many centuries before their actual
discovery in the 17th century. The existence of unseen microbiological life was postulated by Jainism which
is based on Mahavira’s teachings as early as 6th century BCE.[15] Paul Dundas notes that Mahavira
asserted existence of unseen microbiological creatures living in earth, water, air and fire.[16] Jain scriptures
also describe nigodas which are sub-microscopic creatures living in large clusters and having a very short life
and are said to pervade each and every part of universe, even in tissues of plants and flesh of animals.[17]
However, the earliest known idea to indicate the possibility of diseases spreading by yet unseen organisms
was that of the Roman scholar Marcus Terentius Varro in a 1st century BC book titled On Agriculture in which
he warns against locating a homestead near swamps:
“ …and because there are bred certain minute creatures which cannot be seen by
the eyes, which float in the air and enter the body through the mouth and nose and
there cause serious diseases.[18] ”
In The Canon of Medicine (1020), Abū Alī ibn Sīnā (Avicenna) stated that bodily secretion is
contaminated by foul foreign earthly bodies before being infected.[19][unreliable source?] He also
hypothesized that tuberculosis and other diseases might be contagious, i.e. that they were infectious
diseases, and used quarantine to limit their spread.[20][unreliable source?][21][unreliable source?]
When the Black Death bubonic plague reached Andalusia in Spain, in the 14th century, Ibn Khatima
wrote that infectious diseases were caused by contagious "minute bodies" that enter the human body.[19]
[unreliable source?] Later, in 1546, Girolamo Fracastoro proposed that epidemic diseases were caused by
transferable seedlike entities that could transmit infection by direct or indirect contact, or even without contact
over long distances.
All these early claims about the existence of microorganisms were speculative and were not based
on any data or science. Microorganisms were neither proven, observed, nor correctly and accurately
described until the 17th century. The reason for this was that all these early studies lacked the microscope.
History of microorganisms' discovery

See also: History of biology
Antonie van Leeuwenhoek, the first microbiologist and the first to observe microorganisms using a
microscope.
Anton van Leeuwenhoek was one of the first people to observe microorganisms, using a microscope
of his own design, and made one of the most important contributions to biology.[22] Robert Hooke was the
first to use a microscope to observe living things; his 1665 book Micrographia contained descriptions of plant
cells.
Before Leeuwenhoek's discovery of microorganisms in 1675, it had been a mystery why grapes
could be turned into wine, milk into cheese, or why food would spoil. Leeuwenhoek did not make the
connection between these processes and microorganisms, but using a microscope, he did establish that
there were forms of life that were not visible to the naked eye.[23][24] Leeuwenhoek's discovery, along with
subsequent observations by Lazzaro Spallanzani and Louis Pasteur, ended the long-held belief that life
spontaneously appeared from non-living substances during the process of spoilage.
Lazzaro Spallanzani found that boiling broth would sterilise it and kill any microorganisms in it. He
also found that new microorganisms could only settle in a broth if the broth was exposed to the air. Louis
Pasteur expanded upon Spallanzani's findings by exposing boiled broths to the air, in vessels that contained
a filter to prevent all particles from passing through to the growth medium, and also in vessels with no filter at
all, with air being admitted via a curved tube that would not allow dust particles to come in contact with the
broth. By boiling the broth beforehand, Pasteur ensured that no microorganisms survived within the broths at
the beginning of his experiment. Nothing grew in the broths in the course of Pasteur's experiment. This
meant that the living organisms that grew in such broths came from outside, as spores on dust, rather than
spontaneously generated within the broth. Thus, Pasteur dealt the death blow to the theory of spontaneous
generation and supported germ theory.
In 1876, Robert Koch established that microbes can cause disease. He found that the blood of cattle
who were infected with anthrax always had large numbers of Bacillus anthracis. Koch found that he could
transmit anthrax from one animal to another by taking a small sample of blood from the infected animal and
injecting it into a healthy one, and this caused the healthy animal to become sick. He also found that he could
grow the bacteria in a nutrient broth, then inject it into a healthy animal, and cause illness. Based on these
experiments, he devised criteria for establishing a causal link between a microbe and a disease and these
are now known as Koch's postulates.[25] Although these postulates cannot be applied in all cases, they do
retain historical importance to the development of scientific thought and are still being used today.[26]
Classification and structure

Evolutionary tree showing the common ancestry of all three domains of life.[27] Bacteria are colored
blue, eukaryotes red, and archaea green. Relative positions of some phyla are shown around the tree.
Microorganisms can be found almost anywhere in the taxonomic organization of life on the planet.
Bacteria and archaea are almost always microscopic, while a number of eukaryotes are also microscopic,
including most protists, some fungi, as well as some animals and plants. Viruses are generally regarded as
not living and therefore are not microbes, although the field of microbiology also encompasses the study of
viruses.
Prokaryotes
Main article: Prokaryote
Prokaryotes are organisms that lack a cell nucleus and the other membrane bound organelles. They
are almost always unicellular, although some species such as myxobacteria can aggregate into complex
structures as part of their life cycle.
Consisting of two domains, bacteria and archaea, the prokaryotes are the most diverse and
abundant group of organisms on Earth and inhabit practically all environments where some liquid water is
available and the temperature is below +140 °C. They are found in sea water, soil, air, animals'
gastrointestinal tracts, hot springs and even deep beneath the Earth's crust in rocks.[28] Practically all
surfaces which have not been specially sterilized are covered by prokaryotes. The number of prokaryotes on
Earth is estimated to be around five million trillion trillion, or 5 × 10 30, accounting for at least half the biomass
on Earth.[29]
Bacteria
Main article: Bacteria
Staphylococcus aureus bacteria magnified about 10,000x

Bacteria are practically all invisible to the naked eye, with a few extremely rare exceptions, such as
Thiomargarita namibiensis.[30] They lack membrane-bound organelles, and can function and reproduce as
individual cells, but often aggregate in multicellular colonies.[31] Their genome is usually a single loop of
DNA, although they can also harbor small pieces of DNA called plasmids. These plasmids can be transferred
between cells through bacterial conjugation. Bacteria are surrounded by a cell wall, which provides strength
and rigidity to their cells. They reproduce by binary fission or sometimes by budding, but do not undergo
sexual reproduction. Some species form extraordinarily resilient spores, but for bacteria this is a mechanism
for survival, not reproduction. Under optimal conditions bacteria can grow extremely rapidly and can double
as quickly as every 10 minutes.[32]
Archaea
Main article: Archaea
Archaea are also single-celled organisms that lack nuclei. In the past, the differences between
bacteria and archaea were not recognised and archaea were classified with bacteria as part of the kingdom
Monera. However, in 1990 the microbiologist Carl Woese proposed the three-domain system that divided
living things into bacteria, archaea and eukaryotes.[33] Archaea differ from bacteria in both their genetics and
biochemistry. For example, while bacterial cell membranes are made from phosphoglycerides with ester
bonds, archaean membranes are made of ether lipids.[34]
Archaea were originally described in extreme environments, such as hot springs, but have since
been found in all types of habitats.[35] Only now are scientists beginning to realize how common archaea are
in the environment, with crenarchaeota being the most common form of life in the ocean, dominating
ecosystems below 150 m in depth.[36][37] These organisms are also common in soil and play a vital role in
ammonia oxidation.[38]
Eukaryotes
Ostreococcus is the smallest known free living eukaryote with an average size of 0.8 µm
Main article: Eukaryote
Most living things which are visible to the naked eye in their adult form are eukaryotes, including
humans. However, a large number of eukaryotes are also microorganisms. Unlike bacteria and archaea,
eukaryotes contain organelles such as the cell nucleus, the Golgi apparatus and mitochondria in their cells.
The nucleus is an organelle which houses the DNA that makes up a cell's genome. DNA itself is arranged in
complex chromosomes.[39] Mitochondria are organelles vital in metabolism as they are the site of the citric
acid cycle and oxidative phosphorylation. They evolved from symbiotic bacteria and retain a remnant
genome.[40] Like bacteria, plant cells have cell walls, and contain organelles such as chloroplasts in addition
to the organelles in other eukaryotes. Chloroplasts produce energy from light by photosynthesis, and were
also originally symbiotic bacteria.[40]
Unicellular eukaryotes are those eukaryotic organisms that consist of a single cell throughout their
life cycle. This qualification is significant since most multicellular eukaryotes consist of a single cell called a
zygote at the beginning of their life cycles. Microbial eukaryotes can be either haploid or diploid, and some
organisms have multiple cell nuclei (see coenocyte). However, not all microorganisms are unicellular as
some microscopic eukaryotes are made from multiple cells.
Protists
Main article: Protista
Of eukaryotic groups, the protists are most commonly unicellular and microscopic. This is a highly
diverse group of organisms that are not easy to classify.[41][42] Several algae species are multicellular
protists, and slime molds have unique life cycles that involve switching between unicellular, colonial, and
multicellular forms.[43] The number of species of protozoa is uncertain, since we may have identified only a
small proportion of the diversity in this group of organisms.[44][45]
A microscopic mite Lorryia formosa.
Animals
Main article: Micro-animals
Mostly animals are multicellular,[46] but some are too small to be seen by the naked eye.
Microscopic arthropods include dust mites and spider mites. Microscopic crustaceans include copepods and
the cladocera, while many nematodes are too small to be seen with the naked eye. Another particularly
common group of microscopic animals are the rotifers, which are filter feeders that are usually found in fresh
water. Micro-animals reproduce both sexually and asexually and may reach new habitats as eggs that
survive harsh environments that would kill the adult animal. However, some simple animals, such as rotifers
and nematodes, can dry out completely and remain dormant for long periods of time.[47]
Fungi
Main article: Fungus
The fungi have several unicellular species, such as baker's yeast ( Saccharomyces cerevisiae) and
fission yeast (Schizosaccharomyces pombe). Some fungi, such as the pathogenic yeast Candida albicans,
can undergo phenotypic switching and grow as single cells in some environments, and filamentous hyphae in
others.[48] Fungi reproduce both asexually, by budding or binary fission, as well by producing spores, which
are called conidia when produced asexually, or basidiospores when produced sexually.
Plants
Main article: Plant
The green algae are a large group of photosynthetic eukaryotes that include many microscopic
organisms. Although some green algae are classified as protists, others such as charophyta are classified
with embryophyte plants, which are the most familiar group of land plants. Algae can grow as single cells, or
in long chains of cells. The green algae include unicellular and colonial flagellates, usually but not always
with two flagella per cell, as well as various colonial, coccoid, and filamentous forms. In the Charales, which
are the algae most closely related to higher plants, cells differentiate into several distinct tissues within the
organism. There are about 6000 species of green algae.[49]
Habitats and ecology

Microorganisms are found in almost every habitat present in nature. Even in hostile environments
such as the poles, deserts, geysers, rocks, and the deep sea. Some types of microorganisms have adapted
to the extreme conditions and sustained colonies; these organisms are known as extremophiles.
Extremophiles have been isolated from rocks as much as 7 kilometres below the Earth's surface,[50] and it
has been suggested that the amount of living organisms below the Earth's surface may be comparable with
the amount of life on or above the surface.[28] Extremophiles have been known to survive for a prolonged
time in a vacuum, and can be highly resistant to radiation, which may even allow them to survive in space.
[51] Many types of microorganisms have intimate symbiotic relationships with other larger organisms; some
of which are mutually beneficial (mutualism), while others can be damaging to the host organism
(parasitism). If microorganisms can cause disease in a host they are known as pathogens.
Extremophiles
Main article: Extremophile
Extremophiles are microorganisms which have adapted so that they can survive and even thrive in
conditions that are normally fatal to most life-forms. For example, some species have been found in the
following extreme environments:
• Temperature: as high as 130 °C (266 °F),[52] as low as −17 °C (1.4 °F)[53]
• Acidity/alkalinity: less than pH 0,[54] up to pH 11.5[55]
• Salinity: up to saturation[56]
• Pressure: up to 1,000-2,000 atm, down to 0 atm (e.g. vacuum of space)[57]
• Radiation: up to 5kGy[58]
Extremophiles are significant in different ways. They extend terrestrial life into much of the Earth's
hydrosphere, crust and atmosphere, their specific evolutionary adaptation mechanisms to their extreme
environment can be exploited in bio-technology, and their very existence under such extreme conditions
increases the potential for extraterrestrial life.[59]
Soil microbes
The nitrogen cycle in soils depends on the fixation of atmospheric nitrogen. One way this can occur
is in the nodules in the roots of legumes that contain symbiotic bacteria of the genera Rhizobium,
Mesorhizobium, Sinorhizobium, Bradyrhizobium, and Azorhizobium.[60]
Symbiotic microbes
Symbiotic microbes such as fungi and algae form an association in lichen. Certain fungi form
mycorrhizal symbioses with trees that increase the supply of nutrients to the tree.
Importance
Microorganisms are vital to humans and the environment, as they participate in the Earth's element
cycles such as the carbon cycle and nitrogen cycle, as well as fulfilling other vital roles in virtually all
ecosystems, such as recycling other organisms' dead remains and waste products through decomposition.
Microbes also have an important place in most higher-order multicellular organisms as symbionts. Many
blame the failure of Biosphere 2 on an improper balance of microbes.[61]
Use in food
Main article: Fermentation (food)
Microorganisms are used in brewing, winemaking, baking, pickling and other food-making processes.
They are also used to control the fermentation process in the production of cultured dairy products
such as yogurt and cheese. The cultures also provide flavour and aroma, and inhibit undesirable organisms.
[62]
Use in water treatment

Main article: Sewage treatment
Specially-cultured microbes are used in the biological treatment of sewage and industrial waste
effluent, a process known as bioaugmentation.[63]
Use in energy
Main articles: Algae fuel, Cellulosic ethanol, and Ethanol fermentation
Microbes are used in fermentation to produce ethanol,[64] and in biogas reactors to produce
methane.[65] Scientists are researching the use of algae to produce liquid fuels,[66] and bacteria to convert
various forms of agricultural and urban waste into usable fuels.[67]
Use in science
Microbes are also essential tools in biotechnology, biochemistry, genetics, and molecular biology.
The yeasts (Saccharomyces cerevisiae) and fission yeast (Schizosaccharomyces pombe) are important
model organisms in science, since they are simple eukaryotes that can be grown rapidly in large numbers
and are easily manipulated.[68] They are particularly valuable in genetics, genomics and proteomics.[69][70]
Microbes can be harnessed for uses such as creating steroids and treating skin diseases. Scientists are also
considering using microbes for living fuel cells,[71] and as a solution for pollution.[72]
Use in warfare
Main article: Biological warfare
In the Middle Ages, diseased corpses were thrown into castles during sieges using catapults or other
siege engines. Individuals near the corpses were exposed to the deadly pathogen and were likely to spread
that pathogen to others.[73]
Importance in human health

Human digestion
Further information: Human flora#Human bacterial flora and human health
Microorganisms can form an endosymbiotic relationship with other, larger organisms. For example,
the bacteria that live within the human digestive system contribute to gut immunity, synthesise vitamins such
as folic acid and biotin, and ferment complex indigestible carbohydrates.[74]
Diseases and immunology
Main article: Pathogenic microbes
Microorganisms are the cause of many infectious diseases. The organisms involved include
pathogenic bacteria, causing diseases such as plague, tuberculosis and anthrax; protozoa, causing diseases
such as malaria, sleeping sickness and toxoplasmosis; and also fungi causing diseases such as ringworm,
candidiasis or histoplasmosis. However, other diseases such as influenza, yellow fever or AIDS are caused
by pathogenic viruses, which are not usually classified as living organisms and are not therefore
microorganisms by the strict definition. As of 2007, no clear examples of archaean pathogens are known,[75]
although a relationship has been proposed between the presence of some methanogens and human
periodontal disease.[76]
Importance in ecology
Further information: Decomposition
Microbes are critical to the processes of decomposition required to cycle nitrogen and other elements
back to the natural world.
Hygiene
Main article: Hygiene
Hygiene is the avoidance of infection or food spoiling by eliminating microorganisms from the
surroundings. As microorganisms, particularly bacteria, are found practically everywhere, this means in most
cases the reduction of harmful microorganisms to acceptable levels. However, in some cases it is required
that an object or substance be completely sterile, i.e. devoid of all living entities and viruses. A good example
of this is a hypodermic needle.
In food preparation microorganisms are reduced by preservation methods (such as the addition of
vinegar), clean utensils used in preparation, short storage periods or by cool temperatures. If complete
sterility is needed, the two most common methods are irradiation and the use of an autoclave, which
resembles a pressure cooker.
There are several methods for investigating the level of hygiene in a sample of food, drinking water,
equipment etc. Water samples can be filtrated through an extremely fine filter. This filter is then placed in a
nutrient medium. Microorganisms on the filter then grow to form a visible colony. Harmful microorganisms
can be detected in food by placing a sample in a nutrient broth designed to enrich the organisms in question.
Various methods, such as selective media or PCR, can then be used for detection. The hygiene of hard
surfaces, such as cooking pots, can be tested by touching them with a solid piece of nutrient medium and
then allowing the microorganisms to grow on it.
There are no conditions where all microorganisms would grow, and therefore often several different
methods are needed. For example, a food sample might be analyzed on three different nutrient mediums
designed to indicate the presence of "total" bacteria (conditions where many, but not all, bacteria grow),
molds (conditions where the growth of bacteria is prevented by e.g. antibiotics) and coliform bacteria (these
indicate a sewage contamination).
See also
• Bacterium
• Biological warfare
• Culture collection
• Cyanobacteria
• Fungi
• Microbial intelligence
• Nanobacterium
• Petri dish
• Prokaryote
• Protozoa
• Soil contamination
• Staining
• Virus
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External links
• Our Microbial Planet A free poster from the National Academy of Sciences about the positive
roles of microbes.
• "Uncharted Microbial World: Microbes and Their Activities in the Environment" Report from
the American Academy of Microbiology
• Understanding Our Microbial Planet: The New Science of Metagenomics A 20-page
educational booklet providing a basic overview of metagenomics and our microbial planet.
• Tree of Life Eukaryotes
• Microbe News from Genome News Network
• Microbes Patent List Microbes Related Patents
• Medical Microbiology On-line textbook
• Through the microscope: A look at all things small On-line microbiology textbook by Timothy
Paustian and Gary Roberts, University of Wisconsin-Madison
• MicrobeID.com Online Bacteria Identification Key and Probabilistic Identification Databases

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Categories: Microorganisms | Microbiology

W000
List of mammals described in 21st century


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Contents
[hide]
• 1 Living higher taxa
• 2 Living genera
• 3 Living subgenera
• 4 Living species
• 5 Living subspecies
[edit] Living higher taxa

Name Order Author Year Distribution
Luo
Luo, Cifelli & Kielan- & Kielan-Ja
Boreosphenida 2001
Jaworowska Dual origins
mammals. N
Neoceti Cetacea Fordyce & de Muizon 2001 Ford

Muizon, C. 2
history of th
review. Sec
of Tetrapods
Water 169-2
Ford
Muizon, C. 2
history of th
Platanistida Fordyce & de Muizon 2001
review. Sec
of Tetrapods
Water 169-2
D'E
U.F.J., Teta
2007. Defin
D'Elía, Pardiñas, Teta Argentina, of a new trib
Abrotrichini Rodentia 2007
& Patton Bolivia, Chile, Peru rodents (Cri
Sigmodontin
classificatio
Gayana 71(
[edit] Living genera
Name Order Author Year Distribution R
Gonz
Argentina 2000. Un nue
(Misiones), Brazil do roedor sig
(Espirito Santo, de Argentina
Minas Gerais, (Mammalia: R
Juliomys Rodentia González 2000
Parana, Rio de Sigmodontina
Janeiro, Santa Comunicacio
Catarina, Sao Zoologicas d
Paulo) Historia Natu
Montevideo 1
Pipanacoctomys Rodentia Mares, 2000 Argentina Mare

Braun, Barquez & (Catamarca) Braun, J.K., B
Diaz R.M., & Diaz
new genera a
of halophytic
mammals fro
salt flats in A
Occasional P
Museum of T
University, 20
Mare
Braun, J.K., B
R.M., & Diaz
new genera a
Mares,
Argentina of halophytic
Salinoctomys Rodentia Braun, Barquez & 2000
(La Rioja) mammals fro
Diaz
salt flats in A
Occasional P
Museum of T
University, 20
Ande
Yates, T.L. 2
genus and sp
Anderson &
Tapecomys Rodentia 2000 Bolivia phyllotine rod
Yates
Bolivia. Journ
Mammalogy
[4]
Hyladelphys Didelphimorphia Voss, 2001 Brazil, Voss

Lunde & Simmons French Guiana, Lunde, D.P. &
Guyana, Peru N.B. 2001. M
Paracou, Fre
a Neotropica
rainforest fau
Nonvolant sp
Bulletin of the
Museum of N
History 263:1
Voss
Gomez-Lave
Pacheco, V.
Genus for Ae
Voss, fuscatus Alle
Handleyomys Rodentia Gómez-Laverde & 2002 Oryzomys int
Pacheco Thomas, 192
Murid Roden
Andean Clou
American Mu
Novitates 337
Sommeromys Rodentia Musser & 2002 Muss

Durden Durden, L.A.
Sulawesi rod
description o
species of M
(Muridae, Ro
its parasitic n
of sucking lou
Anoplura). Am
Museum Nov
3368:1-50.
[edit] Living subgenera

Name Order Author Year Distribution Referen
Abramov, A
2000. A taxonomic
review of the genus
Cryptomustela Carnivora Abramov 2000 Mustela (Mammalia
Carnivora).
Zoosystematica Ro
8(2):357-364.
Afropipistrellus Chiroptera Thorn, Kock & 2007 Thorn, E., K

Cuisin D. & Cuisin, J. 2007
Status of the Africa
Vesperugo grandid
Dobson, 1876 and
Vesperugo flavesce
Seabra, 1900
(Chiroptera,
Vespertilionidae), w
description of a new
subgenus. Mamma
71(1-2):70-79.
Nadachow
2007. The taxonom
status of Schelkovn
Pine Vole Microtus
Azerbaijan,
Hyrcanicola Rodentia Nadachowski 2007 schelkovnikovi
Iran
(Rodentia, Mamma
Acta zoologica
cracoviensia 50A(1
2):67-72.[6]
Leuconycteris Chiroptera Porter, 2007 Brazil, Porter, C.A

Hoofer, Cline, Colombia, Ecuador, Hoofer, S.R., Cline
Hoffmann & Baker French Guiana, Hoffmann, F.G. & B
Guyana, Peru, R.J. 2007. Molecul
phylogenetics of th
phyllostomid bat ge
Micronycteris with
Venezuela descriptions of two
subgenera. Journa
Mammalogy 88(5):
1215.[7]
Schizonycteris Chiroptera Porter, 2007 Central and Porter, C.A

Hoofer, Cline, South America Hoofer, S.R., Cline
Hoffmann & Baker Hoffmann, F.G. & B
R.J. 2007. Molecul
phylogenetics of th
phyllostomid bat ge
Micronycteris with
descriptions of two
subgenera. Journa
Mammalogy 88(5):
1215.[8]
[edit] Living species

Name Order Author Year Distributio
Argentina
Akodon oenos Rodentia Braun, Mares & Ojeda 2000
(Mendoza)
Argentina
Christoff, Fagundes,
(Misiones), Brazil
Akodon paranaensis Rodentia Sbalqueiro, Mattevi & 2000
(Parana, Rio Grand
Yonenaga-Yassuda
do Sul)
Amblysomus South Africa

Afrosoricida Bronner 2000
robustus (Mpumalanga)
Australia (N
Antechinus
Dasyuromorphia Van Dyck & Crowther 2000 South Wales,
subtropicus
Queensland)
Thalmann &
Avahi unicolor Primates 2000 Madagasca
Geissmann
Brucepattersonius Rodentia Mares & Braun 2000 Argentina

guarani (Misiones)
Brucepattersonius Argentina
Rodentia Mares & Braun 2000
misionensis (Misiones)
Brucepattersonius Argentina
Rodentia Mares & Braun 2000
paradisus (Misiones)
Cheirogaleus
Primates Groves 2000 Madagasca
minusculus
Cheirogaleus ravus Primates Groves 2000 Madagasca
Patton, Da Silva & Brazil

Mesomys occultus Rodentia 2000
Malcolm (Amazonas)
Mico acariensis Primates Van Roosmalen, Van 2000 Brazil

Roosmalen, Mittermeier &
(Amazonas)
Rylands
Van Roosmalen, Van

Brazil
Mico manicorensis Primates Roosmalen, Mittermeier & 2000
(Amazonas)
Rylands
Microcebus berthae Primates Rasoloarison, 2000 Madagasca

Goodman & Ganzhorn
Microcebus Rasoloarison,
Primates 2000 Madagasca
sambiranensis Goodman & Ganzhorn
Rasoloarison,
Microcebus tavaratra Primates 2000 Madagasca
Goodman & Ganzhorn
Myosorex kihaulei Soricomorpha Stanley & Hutterer 2000 Tanzania
Patton, Da Silva &

Neacomys minutus Rodentia 2000 Brazil
Malcolm
Neacomys musseri Rodentia Patton, Da Silva & 2000 Brazil, Peru

Malcolm
Abramov, Averianov &
Nesolagus timminsi Lagomorpha 2000 Laos, Vietn
Tikhonov
Mexico
Notiosorex villai Soricomorpha Carraway & Timm 2000
(Tamaulipas)
Ochotona nigritia Lagomorpha Gong, Wang & Li 2000 China

(Yunnan)
Pipanacoctomys Mares, Braun, Argentina
Rodentia 2000
aureus Barquez & Diaz (Catamarca)
Kruskop &
Plecotus balensis Chiroptera 2000 Ethiopia
Lavrenchenko
Pseudantechinus Dasyuromorphia Coopers, Aplin & 2000 Australia

roryi Adams (Western Australia)
Rhinolophus Kock, Csorba &
Chiroptera 2000 Tanzania
maendeleo Howell
Patton, Da Silva & Brazil (Acre

Rhipidomys gardneri Rodentia 2000
Malcolm Peru
Salinoctomys Mares, Braun, Argentina (L
Rodentia 2000
loschalchalerosorum Barquez & Diaz Rioja)
Tapecomys primus Rodentia Anderson & Yates 2000 Bolivia
Ochoa, Aguilera,
Aepeomys reigi Rodentia 2001 Venezuela
Pacheco & Soriano
Bradypus pygmaeus Pilosa Anderson & Handley 2001 Panama

Cuartas, Muñoz &
Carollia colombiana Chiroptera 2001 Colombia
Míriam González
Brazil (Rio
Ctenomys lami Rodentia De Freitas 2001
Grande do Sul)
Carleton, Goodman &
Eliurus antsingy Rodentia 2001 Madagasca
Rakotondravony
Glauconycteris Cameroon,
Chiroptera Eger & Schlitter 2001
curryae Congo (Dem.Rep.)
Myotis alcathoe Chiroptera Von Helversen & 2001 Bulgaria,

Heller Czech Republic,
France, Germany,
Greece, Hungary,
Montenegro, Serbia
Slovakia, Spain,
Switzerland
Myotis annamiticus Chiroptera Kruskop & Tsytsulina 2001 Vietnam
Brazil
Voss, Lunde &
Neacomys dubosti Rodentia 2001 (Amapa), French
Simmons
Guiana, Suriname
Neacomys paracou Rodentia Voss, Lunde & 2001 Brazil

Simmons (Amapa, Amazonas
Para), French Guia
Guyana, Suriname,
Venezuela
Indonesia
(Irian Jaya, Waigeo
Paranyctimene tenax Chiroptera Bergmans 2001
Papua New Guinea
(New Guinea)
Saccopteryx
Chiroptera Muñoz & Cuartas 2001 Colombia
antioquensis
Spalax carmeli Rodentia Nevo, Ivanitskaya & 2001 Israel

Bailes
Nevo, Ivanitskaya &
Spalax galili Rodentia 2001 Israel
Bailes
Spalax golani Rodentia Nevo, Ivanitskaya & 2001 Israel

Bailes
Nevo, Ivanitskaya &
Spalax judaei Rodentia 2001 Israel
Bailes
Sphiggurus ichillus Rodentia Voss & da Silva 2001 Ecuador

Brazil
Sphiggurus
Rodentia Voss & da Silva 2001 (Amazonas,
roosmalenorum
Rondonia)
Sturnira mistratensis Rodentia Contreras-Vega & 2001 Colombia

Cadena
Sylvilagus
Lagomorpha Durant & Guevara 2001 Venezuela
varynaensis
Argentina
Abrocoma uspallata Rodentia Braun & Mares 2002
(Mendoza)
Bullimus gamay Rodentia Rickart, Heaney & 2002 Philippines

Tabaranza (Camiguin)
Brazil
Van Roosmalen, Van
Callicebus bernhardi Primates 2002 (Amazonas,
Roosmalen & Mittermeier
Rondonia)
Callicebus Primates Van Roosmalen, Van 2002 Brazil

stephennashi Roosmalen & Mittermeier (Amazonas)
Nyctophilus New
Chiroptera Parnaby 2002
nebulosus Caledonia
Aotus
Primates Defler & Bueno 2007 Colombia
jorgehernandezi
Avahi betsileo Primates Andriantompohavana, 2007 Madagasca

Lei, Zaonarivelo, Engberg,
Nalanirina, McGuire, Shore,
Andrianasolo, Herringto,
Brenneman & Louis, Jr.
J.-S. Zhang, Han,
Barbastella
Chiroptera Jones, Lin, J.-P. Zhang, Zhu, 2007 China
beijingensis
Huang & S.-Y. Zhang
S. Meegaskumbura,
M. Meegaskumbura,
Crocidura hikmiya Soricomorpha 2007 Sri Lanka
Pethiyagoda, Manamendra-
Arachchi & Schneider
Jenkins, Abramov,
Crocidura sokolovi Soricomorpha 2007 Vietnam
Rozhnov & Makarova
Jenkins, Abramov,
Crocidura zaitsevi Soricomorpha 2007 Vietnam
Rozhnov & Makarova
Dyacopterus rickarti Chiroptera Helgen, Kock, Gomez 2007

& Ingle
Eliurus danieli Rodentia Carleton & Goodman 2007 Madagasca
Bates, Rossiter, Indonesia

Hipposideros boeadii Chiroptera 2007
Suyanto & Kingston (Sulawesi)
Costa, Pavan, Leite & Brazil (Espi
Juliomys ossitenuis Rodentia 2007
Fagundas Santo, Sao Paulo)
Francis, Kingston & Malaysia

Kerivoula krauensis Chiroptera 2007
Zubaid (Peninsular)
Kerivoula titania Chiroptera Bates, Struebig, 2007 Cambodia,

Hayes, Furey, Mya, Vu, Pham, Laos, Myanmar,
Nguyen, Harrison, Francis & Thailand, Vietnam
Csorba
Craul, Zimmermann,
Lepilemur Rasoloharijaona,
Primates 2007 Madagasca
manasamody Randrianambinina &
Radespiel
Lepilemur otto Primates Craul, Zimmermann, 2007 Madagasca

Rasoloharijaona,
Randrianambinina &
Radespiel
Lonchophylla Colombia,
Chiroptera Woodman 2007
fornicata Ecuador
Lophuromys Rodentia Lavrenchenko, W.N. 2007

chercherensis Verheyen, E. Verheyen,
Hulselmans & Leirs
W.N. Verheyen,
Hulselmans, Dierckx,
Lophuromys kilonzoi Rodentia 2007
Mulungu, Leirs, Corti & E.
Verheyen
Lophuromys Rodentia W.N. Verheyen, 2007

machangui Hulselmans, Dierckx,
Verheyen
W.N. Verheyen,
Lophuromys Hulselmans, Dierckx,
Rodentia 2007
makundii Mulungu, Leirs, Corti & E.
Verheyen

menageshae Verheyen, E. Verheyen,
Hulselmans & Leirs

pseudosikapusi Verheyen, E. Verheyen,
Hulselmans & Leirs
W.N. Verheyen,
Lophuromys sabunii Rodentia 2007
Verheyen
Lophuromys stanleyi Rodentia W.N. Verheyen, 2007

Verheyen
Olivieri, Zimmermann,
Randrianambinina,
Microcebus
Primates Rasoloharijaona, 2007 Madagasca
bongolavensis
Rakotondravony, Guschanski
& Radespiel
Microcebus danfossi Primates Olivieri, Zimmermann, 2007 Madagasca

Randrianambinina,
Rasoloharijaona,
& Radespiel
Olivieri, Zimmermann,
Randrianambinina,
Microcebus
Primates Rasoloharijaona, 2007 Madagasca
lokobensis
& Radespiel
Micronycteris Chiroptera Baker & Fonseca 2007 Ecuador

giovanniae
Miniopterus Chiroptera Goodman, Ryan, 2007 Madagasca
sororculus Maminirina, Fahr, Christidis &
Appleton
Kawada, Shinohara,
Mogera kanoana Soricomorpha 2007 Taiwan
Kobayashi, Harada, Oda & Lin
Monodelphis
Didelphimorphia Solari 2007 Peru
handleyi
Murina tiensa Chiroptera Csorba, Vu, Bates & 2007 Vietnam

Furey
Myzopoda Goodman,
Chiroptera 2007 Madagasca
schliemanni Rakotrondraparany & Kofoky
Van Roosmalen,
Brazil
Pecari maximus Artiodactyla Frenz, Van Hooft, De Iongh & 2007
(Amazonas)
Leirs
Phyllotis anitae Chiroptera Jayat, D'Elía, Pardiñas 2007 Argentina

& Namen
Proedromys China
Rodentia Liu, Sun, Zeng & Zhao 2007
liangshanensis (Sichuan)
Balete, Rickart,
Rhynchomys Philippines
Rodentia Rosell-Ambal, Jansa & 2007
banahao (Luzon)
Heaney
Balete, Rickart,
Philippines
Rhynchomys tapulao Rodentia Rosell-Ambal, Jansa & 2007
(Luzon)
Heaney
Sorex rohweri Soricomorpha Rausch, Feagin & 2007 Canada

Rausch (British Columbia),
USA (Washington)
Spermophilus Gündüz, Jaarola, Tez,
Rodentia 2007 Turkey
taurensis Yeniyurt, Polly & Searle
Özkurt, Sözen, Yiğit,

Spermophilus
Rodentia Kandemir, R. Çolak, 2007 Turkey
torosensis
Gharkheloo & E. Çolak
Thomasomys
Rodentia Salazar-Bravo & Yates 2007 Bolivia
andersoni
Ukraine,
Eptesicus lobatus Chiroptera Zagorodniuk 2009
Rossia
[edit] Living subspecies
Name Order Author Year Distribution
2000. N
system
pp. 174
Arvicola the faun
Rodentia Zagorodnyuk 2000 Ukraine
scherman gutsulius contigu
water vo
species
Publish
Russian
Calomyscus Rodentia Meyer & Malikov 2000 Turkmenistan

elburzensis firiusaensis Malikov
species
mouse-
genus C
(Roden
souther
Zoologi
79(2):21
with En
Malikov
species
mouse-
Calomyscus genus C
Rodentia Meyer & Malikov 2000 Turkmenistan
elburzensis zykovi (Roden
souther
Zoologi
79(2):21
with En
& Raya
African
Civettictis civetta Kock, Künzel & civetta (
Carnivora 2000 Djibouti
pauli Rayaleh Djibouti
subspe
Sencke
80:241-
Eothenomys Rodentia Wang & Li 2000 China

custos changshanensis (Yunnan) Y., 2000
Rodent
Cricetid
Chen, W
Fauna S
Press, B
with En
Y., 2000
Rodent
Eothenomys China Cricetid
Rodentia Wang & Li 2000
custos ninglangensis (Sichuan, Yunnan) Chen, W
Fauna S
Press, B
with En
Y., 2000
Rodent
Eothenomys
Cricetid
melanogaster Rodentia Wang & Li 2000 China
Chen, W
chenduensis
Fauna S
Press, B
with En
Y., 2000
Rodent
Eothenomys
Cricetid
melanogaster Rodentia Wang & Li 2000 China
Chen, W
yingjiangensis
Fauna S
Press, B
with En
Eothenomys olitor Rodentia Wang & Li 2000 China

hypolitor (Yunnan) Y., 2000
Rodent
Cricetid
Chen, W
Fauna S
Press, B
with En
Geomys bursarius Rodentia Elrod, 2000 USA

ozarkensis Zimmermann, Sudman & (Arkansas) Zimmer
Heidt P.D. & H
new sub
gopher
from the
of Arkan
on its hi
biogeog
Mamma
Fariñas
M.E. 20
subspe
Glossophaga Soriano, Fariñas tongued
Chiroptera 2000 Venezuela
longirostris maricelae & Naranjo longiros
enclave
Andes.
Säugeti
374.
2000. A
Mellivora the hon
Carnivora Baryshnikov 2000 Turkmenistan
capensis buechneri capens
Acta Th
55, Mar
Miniopterus Chiroptera Cardinal & 2000 Australia

Cardina
Mitocho
morpho
geograp
(South Australia,
schreibersii bassanii Christidis lineage
Victoria)
bentwin
schreib
Australi
Zoology
Myotis Chiroptera Tsytsulina 2000

mystacinus caucasicus Benda,
K.A. 20
Таксон
группы
group (M
Chiropt
Палеар
revision
mystaci
(Mamm
the wes
Societa
Bohemi
P. & Ts
Таксон
группы
group (M
Chiropt
Myotis
Chiroptera Benda 2000 Палеар
mystacinus occidentalis
revision
mystaci
(Mamm
the wes
Societa
Bohemi
Necromys Rodentia Galliari & 2000 Argentina

obscurus scagliarum Pardiñas (Buenos Aires) Pardiña
Taxono
of the s
of genu
central
Urugua
45(2):21
Niviventer Rodentia Deng & Wang 2000 China

Q. & Wa
Differen
subspe
white-b
confucia
confucianus deqinensis (Yunnan) southwe
descrip
subspe
Resear
Chinese
abstrac
Q. & Wa
Differen
subspe
white-b
Niviventer China confucia
Rodentia Deng & Wang 2000
confucianus yajiangensis (Sichuan) southwe
descrip
subspe
Resear
Chinese
abstrac
Patterso
Geogra
western
Tamias senex Sutton & USA
Rodentia 2000 senex a
pacificus Patterson (California)
two new
Californ
Mamma
[44]
Tamias siskiyou Rodentia Sutton & 2000 USA

humboldtii Patterson (California) Patterso
Geogra
western
senex a
two new
Californ
Mamma
[45]
Molossus Chiroptera Lopez-Gonzalez 2001

currentium robustus & Presley & Presle
Taxono
Molossu
Allen, 1
Molossi
descrip
subspe
Mamma
Strelkov
Russia (Altai, Taxono
Myotis frater Tsytsulina & Irkutsk oblast, frater sp
Chiroptera 2001
eniseensis Strelkov Khakassia, (Vesper
Krasnoyarsk krai) Chiropt
zoologis
26.[46]
Pipistre
meeste
Pipistre
Neoromicia nana South Africa
Chiroptera Kock 2001 Roberts
meesteri (Eastern Cape)
Chiropt
Vespert
Chiropt
130.
2001. N
and tax
Australa
Nyctimene keasti Indonesia
Chiroptera Bergmans 2001 Pteropo
babari (Babar Islands)
Nyctime
Megach
Pteropo
51:119-
Paranyctimene Chiroptera Bergmans 2001 Indonesia

tenax marculus (Irian Jaya, Waigeo) 2001. N
and tax
Australa
Pteropo
Nyctime
Megach
Pteropo
51:119-
Paranyctimene Chiroptera Bergmans 2001 Papua New

tenax tenax Guinea (New Guinea) 2001. N
and tax
Australa
Pteropo
Nyctime
Megach
Pteropo
51:119-
Solenodon Soricomorpha Ottenwalder 2001 Dominican

paradoxus woodi Republic, Haiti 2001. S
biogeog
Indian g
pp. 253
& Sergi
Biogeog
Indies: P
perspec
Florida:
Cephalorhynchus Cetacea Robineau, 2007 Indian Ocean

commersonii Goodall, Pichler & Baker Goodal
kerguelenensis & Baker
Descrip
subspe
Comme
Cephalo
comme
1804), i
coastal
Kergue
Mamma
Perodicticus potto Primates Butynski & De 2007 Kenya

stockleyi Jong Jong, Y
Distribu
Perodic
(Primat
eastern
descrip
subspe
Kenya.
African
96(2):11
Pipistrellus Chiroptera Benda 2007 Cyprus

pygmaeus cyprius V., Horá
Lučan,
2007. B
Chiropt
Mediter
fauna o
records
six spec
island a
new sub
Societa
Bohemi
Retrieved from "http://en.wikipedia.org/wiki/List_of_mammals_described_in_21st_century"
Categories: Lists of animals described in the 21st century

W000
Polyploidy

Polyploidy occurs in cells and organisms when there are more than two paired (homologous) sets of
chromosomes.
Known paleopolyploidy in eukaryotes
Most organisms are normally diploid, meaning they have two sets of chromosomes — one set
inherited from each parent. Polyploidy may occur due to abnormal cell division during metaphase I in
meiosis. It is most commonly found in plants. (Haploidy may also occur as a normal stage in an organism's
life. A haploid has only one set of chromosomes.)
Polyploidy occurs in some animals, such as goldfish,[1] salmon, and salamanders, but is especially
common among ferns and flowering plants (see Hibiscus rosa-sinensis), including both wild and cultivated
species. Wheat, for example, after millennia of hybridization and modification by humans, has strains that are
diploid (two sets of chromosomes), tetraploid (four sets of chromosomes) with the common name of durum or
macaroni wheat, and hexaploid (six sets of chromosomes) with the common name of bread wheat. Many
agriculturally important plants of the genus Brassica are also tetraploids; their relationship is described by the
Triangle of U.
Polyploidy also occurs normally in some animal tissues, such as human muscle tissues.[2] This is
known as endopolyploidy.
The occurrence of polyploidy is a mechanism of speciation and is known to have resulted in new
species of the plant Salsify (Tragopogon).
Speciation via polyploidy: A diploid cell undergoes failed meiosis, producing diploid gametes, which
self-fertilize to produce a tetraploid zygote.
Polyploidy can be induced in plants and cell cultures by some chemicals: the best known is
colchicine, which can result in chromosome doubling, though its use may have other less obvious
consequences as well. Oryzalin also will double the existing chromosome content.
Contents
[hide]
• 1 Polyploid types
• 2 Polyploidy in animals (non-human)
• 3 Polyploidy in humans
• 4 Polyploidy in plants
• 4.1 Polyploid crops
• 4.2 Examples of polyploid crops
• 5 Terminology
• 5.1 Autopolyploidy
• 5.2 Allopolyploidy
• 5.3 Paleopolyploidy
• 5.4 Karyotype
• 5.5 Paralogous
• 5.6 Homologous
• 5.7 Homoeologous
• 5.7.1 Example of homoeologous
chromosomes
• 6 See also
• 7 References
[edit] Polyploid types
Polyploid types are labeled according to the number of chromosome sets in the nucleus:
• triploid (three sets; 3x), for example seedless watermelons, common in the phylum
Tardigrada[3]
• tetraploid (four sets; 4x), for example Salmonidae fish
• pentaploid (five sets; 5x), for example Kenai Birch (Betula papyrifera var. kenaica)
• hexaploid (six sets; 6x), for example wheat, kiwifruit [4]
• octaploid (eight sets; 8x), for example Acipenser (genus of sturgeon fish)
• decaploid (ten sets; 10x), for example certain strawberries
• dodecaploid (twelve sets; 12x), for example the plant Celosia argentea and the amphibian
Xenopus ruwenzoriensis
[edit] Polyploidy in animals (non-human)

Examples in animals are more common in the 'lower' forms[verification needed] such as flatworms,
leeches, and brine shrimp. Polyploid animals are often sterile, so they often reproduce by parthenogenesis.
Polyploid lizards are also quite common and parthenogenetic. Polyploid mole salamanders (mostly triploids)
are all female and reproduce by kleptogenesis,[5] "stealing" spermatophores from diploid males of related
species to trigger egg development but not incorporating the males' DNA into the offspring. While
mammalian liver cells are polyploid, rare instances of polyploid mammals are known, but most often result in
prenatal death.
One of the few known exceptions to this 'rule' is an octodontid rodent of Argentina's harsh desert
regions, known as the Plains Viscacha-Rat (Tympanoctomys barrerae). This rodent is not a rat, but kin to
guinea pigs and chinchillas. Its "new" diploid [2n] number is 102 and so its cells are roughly twice normal
size. Its closest living relation is Octomys mimax, the Andean Viscacha-Rat of the same family, whose 2n =
56. It is surmised that an Octomys-like ancestor produced tetraploid (i.e., 4n = 112) offspring that were, by
virtue of their doubled chromosomes, reproductively isolated from their parents; but that these likely survived
the ordinarily catastrophic effects of polyploidy in mammals by shedding (via translocation or some similar
mechanism) the "extra" set of sex chromosomes gained at this doubling. (The closely related Golden
Vizcacha Rat, 2n = 96, is thought to have arisen via roughly the same process).
[edit] Polyploidy in humans

True polyploidy rarely occurs in humans, although it occurs in some tissues (especially in the liver).
Polyploidy refers to a numerical change in a whole set of chromosomes. Organisms in which a particular
chromosome, or chromosome segment, is under- or overrepresented are said to be aneuploid (from the
Greek words meaning "not," "good," and "fold"). Therefore the distinction between aneuploidy and polyploidy
is that aneuploidy refers to a numerical change in part of the chromosome set, whereas polyploidy refers to a
numerical change in the whole set of chromosomes.<[6]
Polyploidy occurs in humans in the form of triploidy (69,XXX) and tetraploidy (92,XXXX), not to be
confused with 47,XXX or 48,XXXX aneuploidy[clarification needed]. Triploidy, usually due to polyspermy,
occurs in about 2–3% of all human pregnancies and ~15% of miscarriages.[ citation needed] The vast majority
of triploid conceptions end as miscarriage and those that do survive to term typically die shortly after birth. In
some cases survival past birth may occur longer if there is mixoploidy with both a diploid and a triploid cell
population present.
Triploidy may be the result of either digyny (the extra haploid set is from the mother) or diandry (the
extra haploid set is from the father). Diandry is mostly caused by reduplication of the paternal haploid set
from a single sperm, but may also be the consequence of dispermic (two sperm) fertilization of the egg.[7]
Digyny is most commonly caused by either failure of one meiotic division during oogenesis leading to a
diploid oocyte or failure to extrude one polar body from the oocyte. Diandry appears to predominate among
early miscarriages while digyny predominates among triploidy that survives into the fetal period.[ citation
needed] However, among early miscarriages, digyny is also more common in those cases <8.5 weeks
gestational age or those in which an embryo is present. There are also two distinct phenotypes in triploid
placentas and fetuses that are dependent on the origin of the extra haploid set. In digyny there is typically an
asymmetric poorly grown fetus, with marked adrenal hypoplasia and a very small placenta.[citation needed]
In diandry, a partial hydatidiform mole develops.[7] These parent-of-origin effects reflect the effects of
genomic imprinting.[citation needed]
Complete tetraploidy is more rarely diagnosed than triploidy, but is observed in 1–2% of early
miscarriages. However, some tetraploid cells are commonly found in chromosome analysis at prenatal
diagnosis and these are generally considered 'harmless'. It is not clear whether these tetraploid cells simply
tend to arise during in vitro cell culture or whether they are also present in placental cells in vivo. There are,
at any rate, very few clinical reports of fetuses/infants diagnosed with tetraploidy mosaicism.
Mixoploidy is quite commonly observed in human preimplantation embryos and includes
haploid/diploid as well as diploid/tetraploid mixed cell populations. It is unknown whether these embryos fail
to implant and are therefore rarely detected in ongoing pregnancies or if there is simply a selective process
favoring the diploid cells.
[edit] Polyploidy in plants

Polyploidy is pervasive in plants and some estimates suggest that 30–80% of living plant species are
polyploid, and many lineages show evidence of ancient polyploidy (paleopolyploidy) in their genomes.[8][9]
[10] Huge explosions in angiosperm species diversity appear to have coincided with the timing of ancient
genome duplications shared by many species.[11] It has been established that 15% of angiosperm and 31%
of fern speciation events are accompanied by ploidy increase.[12] Polyploid plants can arise spontaneously
in nature by several mechanisms, including meiotic or mitotic failures, and fusion of unreduced (2n) gametes.
[13] Both autopolyploids (e.g. potato[citation needed]) and allopolyploids (e.g. canola, wheat, cotton) can be
found among both wild and domesticated plant species. Most polyploids display heterosis relative to their
parental species, and may display novel variation or morphologies that may contribute to the processes of
speciation and eco-niche exploitation.[13][9] The mechanisms leading to novel variation in newly formed
allopolyploids may include gene dosage effects (resulting from more numerous copies of genome content),
the reunion of divergent gene regulatory hierarchies, chromosomal rearrangements, and epigenetic
remodeling, all of which affect gene content and/or expression levels.[14][15][16] Many of these rapid
changes may contribute to reproductive isolation and speciation.
Lomatia tasmanica is an extremely rare Tasmanian shrub which is triploid and sterile, and
reproduction is entirely vegetative with all plants having the same genetic structure.
There are few naturally occurring polyploid conifers. One example is the giant tree Sequoia
sempervirens or Coast Redwood which is a hexaploid (6x) with 66 chromosomes (2n = 6x = 66), although the
origin is unclear.[17]
[edit] Polyploid crops

Polyploid plants tend to be larger and better at flourishing in early succession habitats such as farm
fields.[citation needed] In the breeding of crops, the tallest and best thriving plants are selected for. Thus,
many crops (and agricultural weeds) may have unintentionally been bred to a higher level of ploidy.
The induction of polyploidy is a common technique to overcome the sterility of a hybrid species
during plant breeding. For example, Triticale is the hybrid of wheat (Triticum turgidum) and rye (Secale
cereale). It combines sought-after characteristics of the parents, but the initial hybrids are sterile. After
polyploidization, the hybrid becomes fertile and can thus be further propagated to become triticale.
In some situations polyploid crops are preferred because they are sterile. For example many
seedless fruit varieties are seedless as a result of polyploidy. Such crops are propagated using asexual
techniques such as grafting.
Polyploidy in crop plants is most commonly induced by treating seeds with the chemical colchicine.
[edit] Examples of polyploid crops

• Triploid crops: apple, banana, citrus, ginger, watermelon [18]
• Tetraploid crops: apple, durum or macaroni wheat, cotton, potato, cabbage, leek, tobacco,
peanut, kinnow, Pelargonium
• Hexaploid crops: chrysanthemum, bread wheat, triticale, oat, kiwifruit [4]
• Octaploid crops: strawberry, dahlia, pansies, sugar cane
Some crops are found in a variety of ploidies: tulips and lilies are commonly found as both diploid
and as triploid; daylilies (Hemerocallis cultivars) are available as either diploid or tetraploid; apples and
kinnows can be diploid, triploid, or tetraploid.
[edit] Terminology
[edit] Autopolyploidy
Autopolyploids are polyploids with multiple chromosome sets derived from a single species.
Autopolyploids can arise from a spontaneous, naturally occurring genome doubling, like the potato.[citation
needed] Others might form following fusion of 2n gametes (unreduced gametes). Bananas and apples can
be found as autotriploids. Autopolyploid plants typically display polysomic inheritance, and are therefore often
infertile and propagated clonally perfect.
[edit] Allopolyploidy
Allopolyploids are polyploids with chromosomes derived from different species. Precisely it is the
result of doubling of chromosome number in an F1 hybrid. Triticale is an example of an allopolyploid, having
six chromosome sets, allohexaploid, four from wheat ( Triticum turgidum) and two from rye (Secale cereale).
Amphidiploid is another word for an allopolyploid. Some of the best examples of allopolyploids come from the
Brassicas, and the Triangle of U describes the relationships among the three common diploid Brassicas ( B.
oleracea, B. rapa, and B. nigra) and three allotetraploids (B. napus, B. juncea, and B. carinata) derived from
hybridization among the diploids.
[edit] Paleopolyploidy
Main article: Paleopolyploidy
Ancient genome duplications probably occurred in the evolutionary history of all life. Duplication
events that occurred long ago in the history of various evolutionary lineages can be difficult to detect because
of subsequent diploidization (such that a polyploid starts to behave cytogenetically as a diploid over time) as
mutations and gene translations gradually make one copy of each chromosome unlike its other copy.
In many cases, these events can be inferred only through comparing sequenced genomes.
Examples of unexpected but recently confirmed ancient genome duplications include baker's yeast
(Saccharomyces cerevisiae), mustard weed/thale cress (Arabidopsis thaliana), rice (Oryza sativa), and an
early evolutionary ancestor of the vertebrates (which includes the human lineage) and another near the origin
of the teleost fishes. Angiosperms (flowering plants) have paleopolyploidy in their ancestry. All eukaryotes
probably have experienced a polyploidy event at some point in their evolutionary history.
[edit] Karyotype
Main article: Karyotype
A karyotype is the characteristic chromosome complement of a eukaryote species.[19][20] The
preparation and study of karyotypes is part of cytology and, more specifically, cytogenetics.
Although the replication and transcription of DNA is highly standardized in eukaryotes, the same
cannot be said for their karotypes, which are highly variable between species in chromosome number and in
detailed organization despite being constructed out of the same macromolecules. In some cases there is
even significant variation within species. This variation provides the basis for a range of studies in what might
be called evolutionary cytology.
[edit] Paralogous
The term is used to describe the relationship among duplicated genes or portions of chromosomes
that derived from a common ancestral DNA. Paralogous segments of DNA may arise spontaneously by
errors during DNA replication, copy and paste transposons, or whole genome duplications.
[edit] Homologous
The term is used to describe the relationship of similar chromosomes that pair at mitosis and
meiosis. In a diploid, one homolog is derived from the male parent (sperm) and one is derived from the
female parent (egg). During meiosis and gametogenesis, homologous chromosomes pair and exchange
genetic material by recombination, leading to the production of sperm or eggs with chromosome haplotypes
containing novel genetic variation.
[edit] Homoeologous
The term homoeologous, also spelled homeologous, is used to describe the relationship of similar
chromosomes or parts of chromosomes brought together following inter-species hybridization and
allopolyploidization, and whose relationship was completely homologous in an ancestral species. In
allopolyploids, the homologous chromosomes within each parental sub-genome should pair faithfully during
meiosis, leading to disomic inheritance; however in some allopolyploids, the homoeologous chromosomes of
the parental genomes may be nearly as similar to one another as the homologous chromosomes, leading to
tetrasomic inheritance (four chromosomes pairing at meiosis), intergenomic recombination, and reduced
fertility.
[edit] Example of homoeologous chromosomes
Durum wheat is the result of the inter-species hybridization of two diploid grass species Triticum
urartu and Aegilops speltoides. Both the diploid ancestors had two sets of 7 chromosomes, which were
similar in terms of size and genes contained on them. Durum wheat contains two sets of chromosomes
derived from Triticum urartu and two sets of chromosomes derived from Aegilops speltoides. Each
chromosome pair derived from the Triticum urartu parent is homoeologous to the opposite chromosome pair
derived from the Aegilops speltoides parent, though each chromosome pair unto itself is homologous.
[edit] See also

• Chromosome
• Karyotype
• Meiosis
• Paleopolyploidy
• Polyploid complex
• Speciation
• Sympatry
[edit] References
1. ^ Ohno S, Muramoto J, Christian L, Atkin NB (1967). "Diploid-tetraploid relationship among
old-world members of the fish family Cyprinidae". Chromosoma 23 (1): 1–9.
doi:10.1007/BF00293307. http://www.springerlink.com/content/n3rr5m124g728271/.
2. ^ Parmacek MS, Epstein JA (July 2009). "Cardiomyocyte renewal". N. Engl. J. Med. 361 (1):
86–8. doi:10.1056/NEJMcibr0903347. PMID 19571289.
3. ^ Bertolani R (2001). "Evolution of the reproductive mechanisms in Tardigrades: a review".
Zoologischer Anzeiger 240 (3-4): 247–252. doi:10.1078/0044-5231-00032.
4. ^ a b The genetic origin of kiwifruit. http://www.actahort.org/books/297/297_5.htm
5. ^ Bogart JP, Bi K, Fu J, Noble DW, Niedzwiecki J (February 2007). "Unisexual salamanders
(genus Ambystoma) present a new reproductive mode for eukaryotes". Genome 50 (2): 119–36.
doi:10.1139/g06-152. PMID 17546077.
6. ^ Griffiths, Anthony J. F. (1999). An Introduction to genetic analysis. San Francisco: W.H.
Freeman. ISBN 0-7167-3520-2.
7. ^ a b Baker, Phil; Monga, Ash; Baker, Philip (2006). Gynaecology by ten teachers. London:
Arnold. ISBN 0-340-81662-7.
8. ^ Meyers LA, Levin DA (June 2006). "On the abundance of polyploids in flowering plants".
Evolution 60 (6): 1198–206. doi:10.1111/j.0014-3820.2006.tb01198.x. PMID 16892970.
9. ^ a b Rieseberg LH, Willis JH (August 2007). "Plant speciation". Science 317 (5840): 910–4.
doi:10.1126/science.1137729. PMID 17702935.
10.^ Otto SP (November 2007). "The evolutionary consequences of polyploidy". Cell 131 (3):
452–62. doi:10.1016/j.cell.2007.10.022. PMID 17981114.
11.^ de Bodt et al. 2005
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2009). "The frequency of polyploid speciation in vascular plants". Proc. Natl. Acad. Sci. U.S.A. 106
(33): 13875–9. doi:10.1073/pnas.0811575106. PMID 19667210.
13.^ a b Comai L (November 2005). "The advantages and disadvantages of being polyploid".
14.^ Osborn TC, Pires JC, Birchler JA, Auger DL, Chen ZJ, Lee HS, Comai L, Madlung A,
Doerge RW, Colot V, Martienssen RA (March 2003). "Understanding mechanisms of novel gene
expression in polyploids". Trends Genet. 19 (3): 141–7. PMID 12615008.
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expression changes in plant polyploids". Bioessays 28 (3): 240–52. doi:10.1002/bies.20374.
PMID 16479580.
16.^ Chen ZJ (2007). "Genetic and epigenetic mechanisms for gene expression and phenotypic
variation in plant polyploids". Annu Rev Plant Biol 58: 377–406.
doi:10.1146/annurev.arplant.58.032806.103835. PMID 17280525.
17.^ Ahuja MR, Neale DB (2002). "Origins of Polyploidy in Coast Redwood ( Sequoia
sempervirens (D. DON) ENDL.) and Relationship of Coast Redwood to other Genera of
Taxodiaceae". Silvae Genetica 51: 2–3.
18.^ Seedless Fruits Make Others Needless
19.^ White M.J.D. 1973. The chromosomes. 6th ed, Chapman & Hall, London. p28
20.^ Stebbins G.L. 1950. Variation and evolution in plants. Chapter XII: The Karyotype.
Columbia University Press N.Y.

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ISBN 100-471-69939-X
• Arabidopsis Genome Initiative (December 2000). "Analysis of the genome sequence of the
flowering plant Arabidopsis thaliana". Nature 408 (6814): 796–815. doi:10.1038/35048692.
PMID 11130711.
• Eakin GS, Behringer RR (December 2003). "Tetraploid development in the mouse". Dev.
Dyn. 228 (4): 751–66. doi:10.1002/dvdy.10363. PMID 14648853.
• Gaeta RT, Pires JC, Iniguez-Luy F, Leon E, Osborn TC (November 2007). "Genomic
changes in resynthesized Brassica napus and their effect on gene expression and phenotype" . Plant
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http://www.plantcell.org/cgi/pmidlookup?view=long&pmid=18024568.
• Gregory, T.R. & Mable, B.K. (2005). Polyploidy in animals. In The Evolution of the Genome
(edited by T.R. Gregory). Elsevier, San Diego, pp. 427–517.
• Jaillon O, Aury JM, Brunet F, et al. (October 2004). "Genome duplication in the teleost fish
Tetraodon nigroviridis reveals the early vertebrate proto-karyotype". Nature 431 (7011): 946–57.
doi:10.1038/nature03025. PMID 15496914.
• Paterson AH, Bowers JE, Van de Peer Y, Vandepoele K (March 2005). "Ancient duplication
of cereal genomes". New Phytol. 165 (3): 658–61. doi:10.1111/j.1469-8137.2005.01347.x.
PMID 15720677.
• Raes J, Vandepoele K, Simillion C, Saeys Y, Van de Peer Y (2003). "Investigating ancient
duplication events in the Arabidopsis genome". J. Struct. Funct. Genomics 3 (1-4): 117–29.
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"The hidden duplication past of Arabidopsis thaliana". Proc. Natl. Acad. Sci. U.S.A. 99 (21): 13627–
32. doi:10.1073/pnas.212522399. PMID 12374856. PMC 129725.
• Soltis DE, Soltis PS, Schemske DW, Hancock JF, Thompson JN, Husband BC, Judd WS
(2007). "Autopolyploidy in angiosperms: have we grossly underestimated the number of species?".
Taxon 56 (1): 13–30.
• Soltis DE, Buggs RJA, Doyle JJ, Soltis PS (2010). "What we still don't know about
polyploidy". Taxon 59: 1387–1403.
• Taylor JS, Braasch I, Frickey T, Meyer A, Van de Peer Y (March 2003). "Genome
duplication, a trait shared by 22000 species of ray-finned fish". Genome Res. 13 (3): 382–90.
doi:10.1101/gr.640303. PMID 12618368. PMC 430266. http://www.genome.org/cgi/pmidlookup?
• Tate, J.A., Soltis, D.E., & Soltis, P.S. (2005). Polyploidy in plants. In The Evolution of the
Genome (edited by T.R. Gregory). Elsevier, San Diego, pp. 371–426.
• Van de Peer Y, Taylor JS, Meyer A (2003). "Are all fishes ancient polyploids?". J. Struct.
Funct. Genomics 3 (1-4): 65–73. doi:10.1023/A:1022652814749. PMID 12836686.
http://www.kluweronline.com/art.pdf?issn=1345-711X&volume=3&page=65.
• Van de Peer Y (2004). "Tetraodon genome confirms Takifugu findings: most fish are ancient
polyploids". Genome Biol. 5 (12): 250. doi:10.1186/gb-2004-5-12-250. PMID 15575976.
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• Van de Peer, Y. and Meyer, A. (2005). Large-scale gene and ancient genome duplications.
In The Evolution of the Genome (edited by T.R. Gregory). Elsevier, San Diego, pp. 329–368
• Wolfe KH, Shields DC (June 1997). "Molecular evidence for an ancient duplication of the
entire yeast genome". Nature 387 (6634): 708–13. doi:10.1038/42711. PMID 9192896.
• Wolfe KH (May 2001). "Yesterday's polyploids and the mystery of diploidization". Nat. Rev.
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• Polyploidy on Kimball's Biology Pages
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W000
Alternation of generations

Alternation of generations (also known as alternation of phases or metagenesis) is a term primarily
used in describing the life cycle of plants (taken here to mean the Archaeplastida). A multicellular diploid
sporophyte, with N paired chromosomes (i.e. 2N in total), alternates with a multicellular haploid gametophyte,
with N unpaired chromosomes. A mature sporophyte produces spores by meiosis, a process which results in
a reduction of the number of chromosomes by a half. Spores germinate and grow into a gametophyte. At
maturity, the gametophyte produces gametes by mitosis. Two gametes (originating from different organisms
of the same species or from the same organism) fuse to produce a zygote, which develops into a diploid
sporophyte. This cycle, from sporophyte to sporophyte (or equally from gametophyte to gametophyte), is the
way in which all land plants and many algae undergo sexual reproduction.
All animals develop differently. A mature animal is diploid and so is, in one sense, equivalent to a
sporophyte. However, an animal directly produces haploid gametes by meiosis. No haploid spores capable
of dividing are produced, so neither is a haploid gametophyte. There is no alternation between diploid and
haploid forms.
Other organisms, such as fungi, can have life cycles in which different kinds of organism alternate.
The term 'alternation of generations' has also been applied to these cases.[ citation needed]
Life cycles, such as those of plants, with alternating haploid and diploid phases can be referred to as
diplohaplontic (the equivalent terms haplodiplontic, diplobiontic or dibiontic are also in use). Life cycles, such
as those of animals, in which there is only a diploid phase are referred to as diplontic. (Life cycles in which
there is only a haploid phase are referred to as haplontic.)
Contents
[hide]
• 1 Definition
• 2 Alternation of generations in plants
• 2.1 Fundamental elements
• 2.2 Variations
• 2.3 A complex life cycle
• 3 Life cycles of different plant groups
• 4 Other groups of organism
• 5 See also
• 6 Notes and References
• 7 Bibilography
[edit] Definition
The discussion of 'alternation of generations' above treats the alternation of a multicellular diploid
form with a multicellular haploid form as the defining characteristic, regardless of whether these forms are
free-living or not.[1] In some species, such as the alga Ulva lactuca, the diploid and haploid forms are indeed
both free-living independent organisms, essentially identical in appearance. The free-swimming gametes
form a zygote which germinates into a diploid sporophyte; the free-swimming spores germinate into a haploid
gametophyte. Alternation of generations is an appropriate term.
However, in other species, either the sporophyte or the gametophyte is very much reduced and is
incapable of free-living. For example, in seed plants, the gametophyte 'generation' develops totally within the
sporophyte which protects and nurtures it, with the sole exception of pollen grains, which are the 'male'
gametophytes, but which have been reduced to only three cells. Here the notion of two generations is less
obvious; as Bateman & Dimichele say "[s]porophyte and gametophyte effectively function as a single
organism".[2] The alternative term 'alternation of phases' may then be more appropriate.[3]
[edit] Alternation of generations in plants

[edit] Fundamental elements
The diagram below shows the fundamental elements of the alternation of generations in plants. It is
vital to have a good understanding of these fundamentals before considering the many variations found in
different groups of plants. Starting from the right of the diagram, the processes involved are as follows:[4]
• Two single-celled haploid gametes, each containing N unpaired chromosomes, fuse to form
a single-celled diploid zygote, which now contains N paired chromosomes, i.e. 2N chromosomes in
total.
• The single-celled diploid zygote germinates, dividing by the normal process (mitosis), which
maintains the number of chromosomes at 2N. The result is a multi-cellular diploid organism, called
the sporophyte (because at maturity it produces spores).
• When it reaches maturity, the sporophyte produces one or more sporangia (singular
sporangium) which are the organs which produce diploid spore mother cells (sporocytes). These
divide by a special process (meiosis) which reduces the number of chromosomes by a half. This
results in four single-celled haploid spores, each containing N unpaired chromosomes.
• The single-celled haploid spore germinates, dividing by the normal process (mitosis), which
maintains the number of chromosomes at N. The result is a multi-cellular haploid organism, called
the gametophyte (because at maturity it produces gametes).
• When it reaches maturity, the gametophyte produces one or more gametangia (singular
gametangium) which are the organs which produce haploid gametes. At least one kind of gamete
possesses some mechanism for reaching another gamete in order to fuse with it.
The 'alternation of generations' in the life cycle is thus between a diploid (2N) generation of
sporophytes and a haploid (N) generation of gametophytes.
Gametophyte of the fern Onoclea sensibilis (the flat thallus at the bottom of the picture) with a
descendant sporophyte beginning to grow from it (the small frond at the top of the picture).
The situation is quite different in all animals, where the fundamental process is that a diploid (2N)
individual directly produces haploid (N) gametes by meiosis. Spores (i.e. haploid cells which are able to
undergo mitosis) are not produced, so neither is a haploid multi-cellular organism. The single-celled gametes
are the only entities which are haploid.
[edit] Variations
The diagram shown above is a good representation of the life cycle of some multi-cellular algae (e.g.
the genus Cladophora) which have sporophytes and gametophytes of very similar, if not identical,
appearance, and which do not have different kinds of spores or gametes.
However, there are many possible variations on the fundamental elements of a life cycle which has
alternation of generations. Each variation may occur separately or in combination, resulting in a bewildering
variety of life cycles. The terms used by botanists in describing these life cycles can be equally bewildering.
As Bateman and Dimichele say "[...] the alternation of generations has become a terminological morass;
often, one term represents several concepts or one concept is represented by several terms."[5]
Possible variations are:
• Relative importance of the sporophyte and the gametophyte.
• Equal (homomorphy or isomorphy).
Filamentous algae of the genus Cladophora, which are predominantly found in fresh water,
have diploid sporophytes and haploid gametophytes which are externally indistinguishable.
[6] No living land plant has equally dominant sporophytes and gametophytes, although some
theories of the evolution of alternation of generations suggest that ancestral land plants did.
• Unequal (heteromorphy or anisomorphy).
Gametophyte of Mnium hornum, a moss.
• Dominant gametophyte (gametophytic).
In liverworts, mosses and hornworts, the dominant form is the haploid gametophyte.
The diploid sporophyte is not capable of an independent existence, gaining most of
its nutrition from the parent gametophyte, and having no chlorophyll when mature.[7]
Sporophyte of Blechnum discolor, a fern.
• Dominant sporophyte (sporophytic).
In ferns, both the sporophyte and the gametophyte are capable of living
independently, but the dominant form is the diploid sporophyte. The haploid
gametophyte is much smaller and simpler in structure. In seed plants, the
gametophyte is even more reduced (at the minimum to only three cells), gaining all
its nutrition from the sporophyte. The extreme reduction in the size of the
gametophyte and its retention within the sporophyte means that when applied to
seed plants the term 'alternation of generations' is somewhat misleading:
"[s]porophyte and gametophyte effectively function as a single organism".[8] Some
authors have preferred the term 'alternation of phases'.[3]
• Differentiation of the gametes.
• Both gametes the same (isogamy).
Like other species of Cladophora, C. callicoma has flagellated gametes which are identical in
appearance and ability to move.[6]
• Gametes of two distinct sizes (anisogamy).
• Both of similar motility.
Species of Ulva, the sea lettuce, have gametes which all have two flagella and so are
motile. However they are of two sizes: larger 'female' gametes and smaller 'male'
gametes.[9]
• One large and sessile, one small and motile (oogamy). The larger sessile
megagametes are eggs (ova), and smaller motile microgametes are sperm
(spermatazoa, spermatozoids). The degree of motility of the sperm may be very
limited (as in the case of flowering plants) but all are able to move towards the
sessile eggs. When (as is almost always the case) the sperm and eggs are produced
in different kinds of gametangia, these are called antheridia (singular antheridium)
and archegonia (singular archegonium) respectively.
Gametophyte of Pellia epiphylla with sporophytes growing from the remains of
archegonia.
• Antheridia and archegonia occur on the same gametophyte, which is
then called monoicous. (Many sources, particularly those concerned with
bryophyes, use the term 'monoecious' for this situation and 'dioecious' for the
opposite.[10][11] Here 'monoecious' and 'dioecious' are used only for
sporophytes.)
The liverwort Pellia epiphylla has the gametophyte as the dominant
generation. It is monoicous: the small reddish antheridia are scattered along
the midrib while the archegonia grow nearer the tip of divisions of the plant.
[12]
• Antheridia and archegonia occur on different gametophytes, which
are then called dioicous.
The moss Mnium hornum has the gametophyte as the dominant generation.
It is dioicous: male plants produce only antheridia in terminal rosettes,
female plants produce only archegonia in the form of stalked capsules.[13]
Seed plants are also dioicous; however, the extreme reduction of the
gametophyte, particularly the microgametophyte, means that the antheridia
and archegonia are microscopic.
• Differentiation of the spores.
• All spores the same size (homospory or isospory).
Horsetails (species of Equisetum) have spores which are all of the same size.[14]
• Spores of two distinct sizes (heterospory or anisospory): larger megaspores and
smaller microspores. When the two kinds of spore are produced in different kinds of
sporangia, these are called megasporangia and microsporangia. A megaspore often (but not
always) develops at the expense of the other three cells resulting from meiosis, which abort.
• Megasporangia and microsporangia occur on the same sporophyte, which is
then called monoecious.
Most flowering plants fall into this category. Thus the flower of a lily contains six
stamens (the microsporangia) which produce microspores which develop into pollen
grains (the microgametophytes), and three fused carpels (the megasporangia) which
produce megaspores which develop into ovules (the megagametophytes). In other
plants, such as hazel, some flowers have only stamens, others only carpels, but the
same plant (i.e. sporophyte) has both kinds of flower and so is monoecious.
Flowers of European Holly, a dioecious species: male above, female below (leaves
cut to show flowers more clearly)
• Megasporangia and microsporangia occur on different sporophytes, which
are then called dioecious.
An individual tree of the European holly (Ilex aquifolium) produces either 'male'
flowers which have only functional stamens (microsporangia) producing microspores
which develop into pollen grains (microgametophytes) or 'female' flowers which have
only functional carpels (megasporangia) producing megaspores which develop into
ovules (megagametophytes).
There are some correlations between these variations, but they are just that, correlations, and not
absolute. For example, in flowering plants, microspores ultimately produce microgametes (sperm) and
megaspores ultimately produce megagametes (eggs). However, in pteridophytes there are groups with
undifferentiated spores but differentiated gametophytes. For example, the fern Ceratopteris thalictrioides has
spores of only one kind, which vary continuously in size. Smaller spores tend to produce gametophytes
which have only sperm-producing antheridia.[15]
[edit] A complex life cycle
The diagram shows the alternation of generations in a species which is heteromorphic, sporophytic,
oogametic, dioicous, heterosporic and dioecious. A seed plant example is a willow tree (genus Salix).[16]
Starting in the centre of the diagram, the processes involved are:
• An immobile egg, typically remaining in the archegonium, fuses with a mobile sperm,
released from an antheridium. The resulting zygote is either 'male' or 'female'.
• A 'male' zygote develops by mitosis into a microsporophyte, which at maturity produces one
or more microsporangia. Microspores develop within the microsporangium by meiosis.
In a willow (like all seed plants) the zygote first develops into a seed within the ovule
(megasporangium). Later the seed is shed and grows into a mature tree. A 'male' willow tree (a
microsporophyte) produces flowers with only stamens, the anthers of which are the microsporangia.
• Microspores germinate producing microgametophytes; at maturity one or more antheridia are
produced. Sperm develop within the antheridia.
In a willow, microspores are not liberated from the anther (the microsporangium), but develop into
pollen grains (microgametophytes) within it. The whole pollen grain is moved (typically by an insect)
to an ovule (megagametophyte), where a sperm is produced which moves down a pollen tube to
reach the egg.
• A 'female' zygote develops by mitosis into a megasporophyte, which at maturity produces
one or more megasporangia. Megaspores develop within the megasporangium; typically one of the
four spores produced by meiosis gains bulk at the expense of the remaining three, which disappear.
'Female' willow trees (megasporophytes) produce flowers with only carpels (the megasporangia).
• Megaspores germinate producing megagametophytes; at maturity one or more archegonia
are produced. Eggs develop within the archegonia.
In a willow, megaspores develop into ovules (megagametophytes) within the carpels
(megasporangia). An archegonium develops within the ovule and produces an egg. All of this
happens within the carpel (the megasporangium). The whole of the gametophytic 'generation'
remains within the protection of the sporophyte except for pollen grains (which have been reduced to
just three cells).
[edit] Life cycles of different plant groups

The term 'plants' is taken here to mean the Archaeplastida, i.e. the glaucophytes, red and green
algae and land plants.
Alternation of generations occurs in almost all multicellular red and green algae, both freshwater
forms (such as Cladophora) and seaweeds (such as Ulva). In most, the generations are homomorphic
(isomorphic) and free-living. Some species of red algae have a complex triphasic alternation of generations,
in which there is a gametophyte phase and two distinct sporophyte phases. For further information, see Red
algae: Reproduction.
Land plants all have heteromorphic (anisomorphic) alternation of generations, in which the
sporophyte and gametophyte are distinctly different. All bryophytes, i.e. liverworts, mosses and hornworts,
have the gametophyte generation as the most conspicuous. As an illustration, consider a monoicous moss.
Antheridia and archegonia develop on the mature plant (the gametophyte). In the presence of water, the
biflagellate sperm from the antheridia swim to the archegonia and fertilisation occurs, leading to the
production of a diploid sporophyte. The sporophyte grows up from the archegonium. Its body comprises a
long stalk topped by a capsule within which spore-producing cells undergo meiosis to form haploid spores.
Most mosses rely on the wind to disperse these spores. For further information, see Liverwort: Life cycle,
Moss: Life cycle, Hornwort: Life cycle.
Diagram of alternation of generations in Moss life cycle Hornwort life cycle

liverworts. diagram diagram
In ferns and their allies, including clubmosses and horsetails, the conspicuous plant observed in the
field is the diploid sporophyte. The haploid spores develop in sori on the underside of the fronds and are
dispersed by the wind (or in some cases, by floating on water). If conditions are right, a spore will germinate
and grow into a rather inconspicuous plant body called a prothallus. The haploid prothallus does not
resemble the sporophyte, and as such ferns and their allies have a heteromorphic alternation of generations.
The prothallus is short-lived, but carries out sexual reproduction, producing the diploid zygote that then grows
out of the prothallus as the sporophyte. For further information, see Fern: Life cycle.
A sporophyte of The underside

Diagram of A gametophyte of a Dicksonia antarctica
(prothallus) of Dicksonia sp. Dicksonia antarctica.
alternation of generations frond showing the sori,
in ferns. or spore-producing
structures.
In the spermatophytes, the seed plants, the sporophyte is the dominant multicellular phase; the
gametophytes are strongly reduced in size and very different in morphology. The entire gametophyte
generation, with the sole exception of pollen grains (microgametophytes), is contained within the sporophyte.
The life cycle of a dioecious flowering plant (angiosperm), the willow, has been outlined in some detail in an
earlier section (A complex life cycle). The life cycle of a gymnosperm is similar. However, flowering plants
have in addition a phenomenon called 'double fertilization'. Two sperm nuclei from a pollen grain (the
microgametophyte), rather than a single sperm, enter the archegonium of the megagametophyte; one fuses
with the egg nucleus to form the zygote, the other fuses with two other nuclei of the gametophyte to form
'endosperm', which nourishes the developing embryo. For further information, see Double fertilization.
Plant ovules
(megagametophytes):
Tip of tulip stamen Gymnosperm ovule on left,
Angiosperm life angiosperm ovule (inside
showing pollen Double fertilization
cycle ovary) on right
(microgametophytes)
[edit] Other groups of organism

Some organisms currently classified in the Chromalveolata, and thus not plants in the sense used
here, exhibit alternation of generations. Kelp are an example of a brown alga with a heteromorphic
alternation of generations. Species from the genus Laminaria have a large sporophytic thallus that produces
haploid spores which germinate to produce free-living microscopic male and female gametophytes.
Foraminifera undergo a heteromorphic alternation of generations between haploid gamont and diploid
agamont forms. The single-celled haploid organism is typically much larger than the diploid organism.
Fungal mycelia are typically haploid. When mycelia of different mating types meet, they produce two
multinucleate ball-shaped cells, which join via a "mating bridge". Nuclei move from one mycelium into the
other, forming a heterokaryon (meaning "different nuclei"). This process is called plasmogamy. Actual fusion
to form diploid nuclei is called karyogamy, and may not occur until sporangia are formed. Karogamy
produces a diploid zygote, which is a short-lived sporophyte that soon undergoes meiosis to form haploid
spores. When the spores germinate, they develop into new mycelia.
The life cycle of slime moulds is very similar to that of fungi. Haploid spores germinate to form swarm
cells or myxamoebae. These fuse in a process referred to as plasmogamy and karyogamy to form a diploid
zygote. The zygote develops into a plasmodium, and the mature plasmodium produces, depending on the
species, one to many fruiting bodies containing haploid spores.
In some animals, there is an alternation between parthenogenic and sexually reproductive phases
(heterogamy). Although in some ways similar to alternation of generations, the genetics of heterogamy is
significantly different.[citation needed]
[edit] See also

• Evolutionary history of plants#life cycles : Evolutionary origin of the alternation of phases
[edit] Notes and References

1. ^ Taylor, Kerp & Hass 2005
2. ^ Bateman & Dimichele 1994, p. 403
3. ^ a b Stewart & Rothwell 1993
4. ^ Unless otherwise indicated, the material in the whole of this section is based on Foster &
Gifford 1974, Sporne 1974a and Sporne 1974b.
6. ^ a b Shyam 1980
7. ^ Watson 1981, p. 2
9. ^ Kirby 2001
10.^ Watson 1981, p. 33
11.^ Bell & Hemsley 2000, p. 104
12.^ Watson 1981, pp. 425–6
13.^ Watson 1981, pp. 287–8
14.^ Bateman & Dimichele 1994, p. 350–1
15.^ Bateman & Dimichele 1994, p. 350–1
16.^ "Willows", Encyclopædia Britannica, XIX (11th ed.), New York: Encyclopædia Britannica,
1911, http://en.wikisource.org/wiki/Page:EB1911_-_Volume_28.djvu/708, retrieved 2011-01-01
[edit] Bibilography
• Bateman, R.M. & Dimichele, W.A. (1994), "Heterospory – the most iterative key innovation in
the evolutionary history of the plant kingdom", Biological Reviews of the Cambridge Philosophical
Society 69: 315–417, http://si-
pddr.si.edu/dspace/bitstream/10088/7107/1/paleo_1994_BatemanDiMichele_Heterospory_BiolRev_
small.pdf, retrieved 2010-12-30
• Bell, P.R. & Hemsley, A.R. (2000), Green Plants: their Origin and Diversity (2nd ed.),
Cambridge, etc.: Cambridge University Press, ISBN 978-0-521-64109-8
• Foster, A.S. & Gifford, E.M. (1974), Comparative Morphology of Vascular Plants (2nd ed.),
San Francisco: W.H. Freeman, ISBN 978-0-7167-0712-7
• Kirby, A. (2001), Ulva, the sea lettuce, Monterey Bay Aquarium Research Institute,
http://www.mbari.org/staff/conn/botany/greens/anna/frontpages/default.htm, retrieved 2011-01-01
• Shyam, R. (1980), "On the life-cycle, cytology and taxonomy of Cladophora callicoma from
India", American Journal of Botany 67 (5): 619–24, http://www.jstor.org/pss/2442655, retrieved 2010-
12-30 (abstract)
• Sporne, K.R. (1974a), The Morphology of Angiosperms, London: Hutchinson, ISBN 978-0-
09-120611-6
• Sporne, K.R. (1974b), The Morphology of Gymnosperms (2nd ed.), London: Hutchinson,
ISBN 978-0-09-077152-3
• Stewart, W.N. & Rothwell, G.W. (1993), Paleobotany and the Evolution of Plants (2nd ed.),
Cambridge, UK: Cambridge University Press, ISBN 978-0-521-38294-6
• Watson, E.V. (1981), British Mosses and Liverworts (3rd ed.), Cambridge, UK: Cambridge
University Press, ISBN 978-0-521-28536-0
• Taylor, T.N.; Kerp, H. & Hass, H. (2005), "Life history biology of early land plants:
understanding the gametophyte phase", Proceedings of the National Academy of Sciences 102 (16):
5892–5897, doi:10.1073/pnas.0501985102

Plant physiology



Transpiration

Sporophyte

morphology terms
Category · Portal
Retrieved from "http://en.wikipedia.org/wiki/Alternation_of_generations"
Categories: Plant reproduction | Reproduction

W000
Pterosaur

"Pterodactyl" redirects here. For other uses, see Pterodactyl (disambiguation).
Pterosaur
Eukaryota
Pterosaurs
Temporal range: Late Triassic–Late Cretaceous, 220–65 Ma
PreЄ
g
Replica Geosternbergia sternbergi skeletons, male (right) and female (left)
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Division: Archosauria
(unranked): Avemetatarsalia
(unranked): Ornithodira
†Pterosauria
Order:
Kaup, 1834
Suborders
†Pterodactyloidea
†Rhamphorhynchoidea *
Pterosaurs (pronounced /ˈtɛrəsɔr/, from the Greek πτερόσαυρος, pterosauros, meaning "winged
lizard", often referred to as pterodactyls, from the Greek πτεροδάκτυλος, pterodaktulos, meaning "winged
finger" /ˌtɛrəˈdæktɨl/) were flying reptiles of the clade or order Pterosauria. They existed from the late Triassic
to the end of the Cretaceous Period (220 to 65.5 million years ago). Pterosaurs are the earliest vertebrates
known to have evolved powered flight. Their wings were formed by a membrane of skin, muscle, and other
tissues stretching from the legs to a dramatically lengthened fourth finger. Early species had long, fully-
toothed jaws and long tails, while later forms had a highly reduced tail, and some lacked teeth. Many sported
furry coats made up of hair-like filaments known as pycnofibres, which covered their bodies and parts of their
wings. Pterosaurs spanned a wide range of adult sizes, from the very small Nemicolopterus to the largest
known flying creatures of all time, including Quetzalcoatlus and Hatzegopteryx.[1][2][3]
Pterosaurs are sometimes referred to in the popular media as dinosaurs, but this is incorrect. The
term "dinosaur" is properly restricted to a certain group of reptiles with a unique upright stance (superorder
Dinosauria, which includes birds), and therefore excludes the pterosaurs, as well as the various groups of
extinct marine reptiles, such as ichthyosaurs, plesiosaurs, and mosasaurs.
Contents
[hide]
• 1 Description
• 1.1 Wings
• 1.1.1 Parts of the pterosaur wing
• 1.2 Skull, teeth and crests
• 1.3 Pycnofibres
• 2 History of discovery
• 3 Paleobiology
• 3.1 Flight
• 3.2 Air sacs and respiration
• 3.3 Nervous system
• 3.4 Ground movement
• 3.5 Natural predators
• 3.6 Reproduction and life history
• 4 Evolution and extinction
• 4.1 Origins
• 4.2 Phylogeny and classification
• 4.3 Extinction
• 4.4 Well-known genera
• 5 In popular culture
• 6 See also
[edit] Description
The anatomy of pterosaurs was highly modified from their reptilian ancestors for the demands of
flight. Pterosaur bones were hollow and air filled, like the bones of birds. They had a keeled breastbone that
was developed for the attachment of flight muscles and an enlarged brain that shows specialised features
associated with flight.[4] In some later pterosaurs, the backbone over the shoulders fused into a structure
known as a notarium, which served to stiffen the torso during flight, and provide a stable support for the
scapula (shoulder blade).
[edit] Wings
Reconstructed wing planform of Quetzalcoatlus compared to the Wandering Albatross and the
Andean Condor (not to scale)
Pterosaur wings were formed by membranes of skin and other tissues. The primary membranes
attached to the extremely long fourth finger of each arm and extended along the sides of the body to the legs.
While historically thought of as simple, leathery structures composed of skin, research has since
shown that the wing membranes of pterosaurs were actually highly complex and dynamic structures suited to
an active style of flight. First, the outer wings (from the wing to the elbow) were strengthened by closely
spaced fibers called actinofibrils.[5] The actinofibrils themselves consisted of three distinct layers in the wing,
forming a crisscross pattern when superimposed on one another. The actual function of the actinofibrils is
unknown, as is the exact material from which they were made. Depending on their exact composition
(keratin, muscle, elastic structures, etc.), they may have been stiffening or strengthening agents in the outer
part of the wing.[6] The wing membranes also contained a thin layer of muscle, fibrous tissue, and a unique,
complex circulatory system of looping blood vessels.[7]
As evidenced by hollow cavities in the wing bones of larger species and soft tissue preserved in at
least one specimen, some pterosaurs extended their system of respiratory air sacs (see Paleobiology section
below) into the wing membrane itself.[8]
[edit] Parts of the pterosaur wing

Sordes, as depicted here, evidences the possibility that pterosaurs had a cruropatagium (a
membrane connecting the legs, but leaving the tail out unlike the chiropteran uropatagium)
The pterosaur wing membrane is divided into three basic units. The first, called the propatagium
("first membrane"), was the forward-most part of the wing and attached between the wrist and shoulder,
creating the "leading edge" during flight. This membrane may have incorporated the first three fingers of the
hand, as evidenced in some specimens.[7] The brachiopatagium ("arm membrane") was the primary
component of the wing, stretching from the highly elongated fourth finger of the hand to the hind limbs
(though where exactly on the hind limbs it anchored is controversial and may have varied between species,
see below). Finally, at least some pterosaur groups had a membrane that stretched between the legs,
possibly connecting to or incorporating the tail, called the uropatagium; the extent of this membrane isn't
certain, as studies on Sordes seem to suggest that it simply connected the legs but did not involve the tail
(rendering it a cruropatagium). It is generally agreed though that non-pterodactyloid pterosaurs had a broader
uro/cruropatagium, with pterodactyloids only having membranes running along the legs; Pteranodon in
particular might have developed/redeveloped an uropatagium, given the structure of the tail.[ citation needed]
Wing anatomy
A bone unique to pterosaurs, known as the pteroid, connected to the wrist and helped to support a
forward membrane (the propatagium) between the wrist and shoulder. Evidence of webbing between the
three free fingers of the pterosaur forelimb suggests that this forward membrane may have been more
extensive than the simple pteroid-to-shoulder connection traditionally depicted in life restorations.[7] The
position of the pteroid bone itself has been controversial. Some scientists, notably David Unwin, have argued
that the pteroid pointed forward, extending the forward membrane.[9] However, this view was strongly refuted
in a 2007 paper by Chris Bennett, who showed that the pteroid did not articulate as previously thought and
could not have pointed forward, but rather inward toward the body as traditionally thought.[10]
Three lines of evidence, morphological, developmental and histological, indicate that the pteroid is a
true bone, rather than ossified cartilage. The origin of the pteroid is unclear: it may be a modified carpal, the
first metacarpal, or a neomorph (new bone).[11]
The pterosaur wrist consists of two inner (proximal) and four outer (distal) carpals (wrist bones),
excluding the pteroid bone, which may itself be a modified distal carpal. The proximal carpals are fused
together into a "syncarpal" in mature specimens, while three of the distal carpals fuse to form a distal
syncarpal. The remaining distal carpal, referred to here as the medial carpal, but which has also been termed
the distal lateral, or pre-axial carpal, articulates on a vertically elongate biconvex facet on the anterior surface
of the distal syncarpal. The medial carpal bears a deep concave fovea that opens anteriorly, ventrally and
somewhat medially, within which the pteroid articulates.[12]
There has been considerable argument among paleontologists about whether the main wing
membranes (brachiopatagia) attached to the hind limbs, and if so, where. Fossils of the rhamphorhynchoid
Sordes,[13] the anurognathid Jeholopterus,[14] and a pterodactyloid from the Santana Formation seem to
demonstrate that the wing membrane did attach to the hindlimbs, at least in some species.[15] However,
modern bats and flying squirrels show considerable variation in the extent of their wing membranes and it is
possible that, like these groups, different species of pterosaur had different wing designs. Indeed, analysis of
pterosaur limb proportions shows that there was considerable variation, possibly reflecting a variety of wing-
plans.[16]
Many if not all pterosaurs also had webbed feet.[17]
[edit] Skull, teeth and crests

Tooth, possibly from Coloborhynchus
Most pterosaur skulls had elongated, beak-like jaws. Some advanced forms were toothless (such as
the pteranodonts and azhdarchids, though most sported a full complement of needle-like teeth.[18] In some
cases, actual keratinous beak tissue has been preserved, though in toothed forms, the beak is small and
restricted to the jaw tips and does not involve the teeth.[19]
Unlike most archosaurs, which have several openings in the skull in front of the eyes, in
pterodactyloid pterosaurs the antorbital opening and the nasal opening was merged into a single large
opening, called the nasoantorbial fenestra. This likely evolved as a weight-saving feature to lighten the skull
for flight.[18]
The crests of three tapejarids. Clockwise from right: Tapejara, Tupandactylus, and "Tapejara"
navigans (not to scale)
Pterosaurs are well known for their often elaborate crests. The first and perhaps best known of these
is the distinctive backward-pointing crest of some Pteranodon species, though a few pterosaurs, such as the
tapejarids and Nyctosaurus sported incredibly large crests that often incorporated keratinous or other soft
tissue extensions of the bony crest base.
Since the 1990s, new discoveries and more thorough study of old specimens have shown that crests
are far more widespread among pterosaurs than previously thought, due mainly to the fact that they were
frequently extended by or composed completely of keratin, which does not fossilize as often as bone.[7] In
the cases of pterosaurs like Pterorhynchus and Pterodactylus, the true extent of these crests has only been
uncovered using ultraviolet photography.[19][20] The discovery of Pterorynchus and Austriadactylus, both
crested "rhamphorhynchoids", showed that even primitive pterosaurs had crests (previously, crests were
thought to be restricted to the more advanced pterodactyloids).[7]
[edit] Pycnofibres
At least some pterosaurs were covered with hair-like filaments known as pycnofibres, similar to but
not homologous (sharing a common structure) with mammalian hair. Pycnofibres were not true hair as seen
in mammals, but a unique structure that developed a similar appearance through convergent evolution.
Although in some cases actinofibrils (internal structural fibres) in the wing membrane have been mistaken for
pycnofibres or true hair, some fossils such as those of Sordes pilosus (which translates as "hairy demon")
and Jeholopterus ninchengensis do show the unmistakable imprints of pycnofibres on the head and body,
not unlike modern-day bats, another example of convergent evolution.[13] The presence of pycnofibres (and
the demands of flight) imply that pterosaurs were endothermic (warm-blooded).
The term "pycnofibre", meaning "dense filament", was first coined in a paper on the soft tissue
impressions of Jeholopterus by palaeontologist Alexander W.A. Kellner and colleagues in 2009.[6]
[edit] History of discovery
Engraving of the original P. antiquus specimen by Egid Verhelst II, 1784

The first pterosaur fossil was described by the Italian naturalist Cosimo Collini in 1784. Collini
misinterpreted his specimen as a seagoing creature that used its long front limbs as paddles.[21] A few
scientists continued to support the aquatic interpretation even until 1830, when the German zoologist Johann
Georg Wagler suggested that Pterodactylus used its wings as flippers.[22] Georges Cuvier first suggested
that pterosaurs were flying creatures in 1801,[23] and coined the name "Ptero-dactyle" 1809 for the specimen
recovered in Germany; however, due to the standardization of scientific names, the official name for this
genus became Pterodactylus, though the name "pterodactyl" continued to be popularly applied to all
members of this first specimen's order.
Since the first pterosaur fossil was discovered in the Late Jurassic Solnhofen limestone in 1784,
twenty-nine kinds of pterosaurs have been found in those deposits alone. A famous early UK find was an
example of Dimorphodon by Mary Anning, at Lyme Regis in 1828. The name Pterosauria was coined by
Johann Jakob Kaup in 1834, though the name Ornithosauria (or "bird lizards", Bonaparte, 1838) was
sometimes used in the early literature.[2]
The three-dimensionally preserved skull of Anhanguera santanae, from the Santana Formation,
Brazil
Most pterosaur fossils are poorly preserved. Their bones were hollow and, when sediments piled on
top of them, the bones were flattened. The best preserved fossils have come from the Araripe Plateau, Brazil.
For some reason, when the bones were deposited, the sediments encapsulated the bones, rather than
crushing them. This created three-dimensional fossils for paleontologists to study. The first find in the Araripe
Plateau was made in 1974.
Most paleontologists now believe that pterosaurs were adapted for active flight, not just gliding as
was earlier believed. Pterosaur fossils have been found on every continent. At least 60 genera of pterosaurs
have been found to date, ranging from the size of a small bird to wingspans in excess of 10 metres (33 ft).
[edit] Paleobiology
[edit] Flight
Skeletal reconstruction of a quadrupedally launching Pteranodon longiceps

The mechanics of pterosaur flight are not completely understood or modeled at this time.[24][25]
Diagrams showing breathing motion (top two) and internal air sac system (bottom two)
Katsufumi Sato, a Japanese scientist, did calculations using modern birds and decided that it is
impossible for a pterosaur to stay aloft.[26] In the book Posture, Locomotion, and Paleoecology of Pterosaurs
it is theorized that they were able to fly due to the oxygen-rich, dense atmosphere of the Late Cretaceous
period.[27] However, one must note both Katsufumi and the authors of Posture, Locomotion, and
Paleoecology of Pterosaurs based their research on the now outdated theories of pterosaurs being seabird-
like, and the size limit doesn't apply to terrestrial pterosaurs like azhdarchids and tapejarids [3] Furthermore,
Darren Naish concluded that atmospheric differences between the present and the Mesozoic weren't needed
for the giant size of pterosaurs.[28]
However, Mark Witton and Mike Habib, of the University of Portsmouth and Johns Hopkins
University, respectively, argue that pterosaurs used a vaulting mechanism to obtain flight.[29] Once in air,
pterosaurs could reach speeds up to 120 kilometres per hour (75 mph) and travel thousands of kilometers.
[29]
[edit] Air sacs and respiration

A 2009 study showed that pterosaurs had a lung-air sac system and a precisely controlled skeletal
breathing pump, which supports a flow-through pulmonary ventilation model in pterosaurs, analogous to that
of birds. The presence of a subcutaneous air sac system in at least some pterodactyloids would have further
reduced the density of the living animal.[8]
[edit] Nervous system

A study of pterosaur brain cavities using X-rays revealed that the animals ( Rhamphorhynchus
muensteri and Anhanguera santanae) had massive flocculi. The flocculus is a brain region that integrates
signals from joints, muscles, skin and balance organs.[4]
The pterosaurs' flocculi occupied 7.5% of the animals' total brain mass, more than in any other
vertebrate. Birds have unusually large flocculi compared with other animals, but these only occupy between 1
and 2% of total brain mass.[4]
The flocculus sends out neural signals that produce small, automatic movements in the eye muscles.
These keep the image on an animal's retina steady. Pterosaurs may have had such a large flocculus
because of their large wing size, which would mean that there was a great deal more sensory information to
process.[4]
[edit] Ground movement
The probable azhdarchid trace fossil Haenamichnus uhangriensis.
Pterosaur's hip sockets are oriented facing slightly upwards, and the head of the femur (thigh bone)
is only moderately inward facing, suggesting that pterosaurs had a semi-erect stance. It would have been
possible to lift the thigh into a horizontal position during flight as gliding lizards do.
There was considerable debate whether pterosaurs ambulated as quadrupeds or as bipeds. In the
1980s, paleontologist Kevin Padian suggested that smaller pterosaurs with longer hindlimbs such as
Dimorphodon might have walked or even run bipedally, in addition to flying, like road runners.[30] However, a
large number of pterosaur trackways were later found with a distinctive four-toed hind foot and three-toed
front foot; these are the unmistakable prints of pterosaurs walking on all fours.[31][32]
Unlike most vertebrates, which walk on their toes with ankles held off the ground (digitigrade), fossil
footprints show that pterosaurs stood with the entire foot in contact with the ground (plantigrade), in a manner
similar to humans and bears. Footprints from azhdarchids show that at least some pterosaurs walked with an
erect, rather than sprawling, posture.[17]
Fossil trackways show that pterosaurs like Quetzalcoatlus northropi were quadrupeds.
Though traditionally depicted as ungainly and awkward when on the ground, the anatomy of at least
some pterosaurs (particularly pterodactyloids) suggests that they were competent walkers and runners.[33]
The forelimb bones of azhdarchids and ornithocheirids were unusually long compared to other pterosaurs,
and in azhdarchids, the bones of the arm and hand (metacarpals) were particularly elongated. Furthermore,
azhdarchid front limbs as a whole were proportioned similarly to fast-running ungulate mammals. Their hind
limbs, on the other hand, were not built for speed, but they were long compared with most pterosaurs, and
allowed for a long stride length. While azhdarchid pterosaurs probably could not run, they would have been
relatively fast and energy efficient.[17]
The relative size of the hands and feet in pterosaurs (by comparison with modern animals such as
birds) may indicate what type of lifestyle pterosaurs led on the ground. Azhdarchid pterosaurs had relatively
small feet compared to their body size and leg length, with foot length only about 25%-30% the length of the
lower leg. This suggests that azhdarchids were better adapted to walking on dry, relatively solid ground.
Pteranodon had slightly larger feet (47% the length of the tibia), while filter-feeding pterosaurs like the
ctenochasmatoids had very large feet (69% of tibial length in Pterodactylus, 84% in Pterodaustro), adapted to
walking in soft muddy soil, similar to modern wading birds.[17]
[edit] Natural predators

Pterosaurs are known to have been eaten by spinosaurids. In the 1 July 2004 edition of Nature,
paleontologist Eric Buffetaut discusses an early Cretaceous fossil of three cervical vertebrae of a pterosaur
with the broken tooth of a spinosaur embedded in it. The vertebrae are known not to have been eaten and
exposed to digestion, as the joints still articulated.[34]
[edit] Reproduction and life history

Fossil pterodactyloid flapling from the Solnhofen Limestone
Very little is known about pterosaur reproduction, and pterosaur eggs are very rare. The first known
pterosaur egg was found in the quarries of Liaoning, the same place that yielded the famous 'feathered'
dinosaurs. The egg was squashed flat with no signs of cracking, so evidently the eggs had leathery shells, as
in modern lizards.[35] This was supported by the description of an additional pterosaur egg belonging to the
genus Darwinopterus, described in 2011, which also had a leathery shell and, also like modern reptiles but
unlike birds, was fairly small compared to the size of the mother.[36] A study of pterosaur eggshell structure
and chemistry published in 2007 indicated that it is likely pterosaurs buried their eggs, like modern crocodile
and turtles. Egg-burying would have been beneficial to the early evolution of pterosaurs, as it allows for more
weight-reducing adaptations, but this method of reproduction also would have put limits on the variety of
environments pterosaurs could live in, and may have disadvantaged them when they began to face
ecological competition from birds.[37]
Wing membranes preserved in pterosaur embryos are well developed, suggesting pterosaurs were
ready to fly soon after birth.[38] Fossils of pterosaurs only a few days to a week old (called flaplings) have
been found, representing several pterosaur families, including pterodactylids, rhamphorhinchids,
ctenochasmatids and azhdarchids.[18] All preserve bones which show a relatively high degree of hardening
(ossification) for their age, and wing proportions similar to adults. In fact, many pterosaur flaplings have been
considered adults and placed in separate species in the past. Additionally, flaplings are normally found in the
same sediments as adults and juveniles of the same species, such as the Pterodactylus and
Rhamphorhynchus flaplings found in the Solnhofen limestone of Germany, and Pterodaustro flaplings from
Brazil. All are found in deep aquatic environment far from shore.[39]
It is not known whether pterosaurs practiced any form of parental care, but their ability to fly as soon
as they emerged from the egg and the numerous flaplings found in environments far from nests and
alongside adults has led most researchers, including Christopher Bennett and David Unwin, to conclude that
the young were only dependent on their parents for a very short period of time, while the wings grew long
enough to fly, and left the nest to fend for themselves within days of hatching.[18] Alternately, they may have
used stored yolk products for nourishment during their first few days of life, as in modern reptiles, rather than
depend on parents for food.[39]
Growth rates of pterosaurs once they hatched varied across different groups. In more primitive, long-
tailed pterosaurs ("rhamphorhynchoids") such as Rhamphorhynchus, the average growth rate during the first
year of life was 130% to 173%, slightly faster than the growth rate of alligators. Growth in these species
slowed after sexual maturity, and it would have taken more than three years for Rhamphorhynchus to attain
maximum size. In contrast, the more advanced, large pterodactyloid pterosaurs such as Pteranodon grew to
adult size within the first year of life. Additionally, pterodactyloids had determinate growth, meaning that the
animals reached a fixed maximum adult size and stopped growing.[39]
[edit] Evolution and extinction
[edit] Origins
Restoration of two Scleromochlus on a tree
Because pterosaur anatomy has been so heavily modified for flight, and immediate "missing link"
predecessors have not so far been described, the ancestry of pterosaurs is not well understood. Several
hypotheses have been advanced, including links to ornithodirans like Scleromochlus, an ancestry among the
basal archosauriforms like Euparkeria, or among the prolacertiformes (which include gliding forms like
Sharovipteryx).[18]
Two researchers, Chris Bennett (1996) and David Peters (2000), have found pterosaurs to be
prolacertiformes or closely related to them. Bennett only recovered pterosaurs as close relatives of the
prolacertiformes after removing characteristics of the hind limb from his analysis, in an attempt to test the
idea that these characters are the result of convergent evolution between pterosaurs and dinosaurs.
However, subsequent analysis by Dave Hone and Michael Benton (2007) could not reproduce this result.
Hone and Benton found pterosaurs to be closely related to dinosaurs even without hind limb characters.
They also criticized previous studies by David Peters, raising "serious questions" about the methods he used
to recover pterosaurs among the prolacertiformes. Hone and Benton concluded that although more primitive
pterosauromorphs are needed to clarify their relationships, pterosaurs are best considered archosaurs, and
specifically ornithodirans, given current evidence. In Hone and Benton's analysis, pterosaurs are either the
sister group of Scleromochlus or fall between it and Lagosuchus on the ornithodiran family tree.[40]
[edit] Phylogeny and classification

For more details on this topic, see List of pterosaur classifications.
Classification of pterosaurs has historically been difficult, because there were many gaps in the fossil
record. Many new discoveries are now filling in these gaps and giving a better picture of the evolution of
pterosaurs. Traditionally, they are organized into two suborders:
• Rhamphorhynchoidea (Plieninger, 1901): A group of early, basal ("primitive") pterosaurs,
many of which had long tails and short metacarpal bones in the wing. They were small, and their
fingers were still adapted to climbing[citation needed]. They appeared in the Late Triassic period, and
lasted until the late Jurassic. Rhamphorhynchoidea is a paraphyletic group (since the pterodactyloids
evolved directly from them and not from a common ancestor), so with the increasing use of cladistics
it has fallen out of favor in most technical literature.
• Pterodactyloidea (Plieninger, 1901): The more derived ("advanced") pterosaurs, with short
tails and long wing metacarpals. They appeared in the middle Jurassic period, and lasted until the
Cretaceous-Tertiary extinction event wiped them out at the end of the Cretaceous.
Zhejiangopterus, an azhdarchid from the Cretaceous of China
Rhamphorhynchus, a rhamphorhynchid from the Jurassic of Germany

Listing of families and superfamilies within Pterosauria, after Unwin 2006 unless otherwise noted.[18]
• ORDER PTEROSAURIA (extinct)
• Suborder Rhamphorhynchoidea *
• Family Anurognathidae
• Family Campylognathoididae
• Family Dimorphodontidae
• Family Rhamphorhynchidae
• Suborder Pterodactyloidea
• Superfamily Ornithocheiroidea
• Family Istiodactylidae
• Family Nyctosauridae
• Family Ornithocheiridae
• Family Pteranodontidae
• Superfamily Ctenochasmatoidea
• Family Ctenochasmatidae
• Family Gallodactylidae
• Family Pterodactylidae
• Superfamily Dsungaripteroidea
• Family Dsungaripteridae
• Family Germanodactylidae
• Superfamily Azhdarchoidea
• Family Azhdarchidae
• Family Chaoyangopteridae[41]
• Family Lonchodectidae
• Family Tapejaridae
The precise relationships between pterosaurs is still unsettled. However, several newer studies are
beginning to make things clearer. Cladogram simplified after Unwin.[42]
Pterosauria
Macronychoptera
Dimorphodonti
Caelidracones
Anurognathidae
Lonchognatha
Campylognathoididae
Breviquartossa
Rhamphorhynchidae
Pterodactyloidea
(see below)
Pterodactyloidea
Ornithocheiroidea
Istiodactylidae
Euornithocheira
Ornithocheiridae
Pteranodontidae
Lophocratia
Ctenochasmatoidea
Gallodactylidae
Euctenochasmatia
Pterodactylus
Lonchodectes
Ctenochasmatidae
Dsungaripteroidea
Germanodactylidae
Dsungaripteridae
Azhdarchoidea
Tapejaridae
Azhdarchidae
[edit] Extinction
It was once thought that competition with early bird species may have resulted in the extinction of
many of the pterosaurs.[citation needed] By the end of the Cretaceous, only large species of pterosaurs are
known. The smaller species seem to have become extinct, their niche filled by birds.[43] However, pterosaur
decline (if actually present) seems unrelated to bird diversity.[44] At the end of the Cretaceous period, the
great extinction which wiped out all non-avian dinosaurs and most avian dinosaurs as well, and many other
animals, seemed to also take the pterosaurs. Alternatively, most pterosaurs may have been specialised for
an ocean-going lifestyle.[citation needed] Consequently, when the K-T mass-extinction severely affected
marine life that most pterosaurs fed on, they went extinct. However, forms like azhdarchids and istiodactylids
were not marine in habits.
[edit] Well-known genera

Main article: List of pterosaurs
Examples of pterosaur genera include:
• Pteranodon was 1.8 metres (six ft) long, with a wingspan of 7.5 m (25 ft), and lived during the
late Cretaceous period.
• Pterodactylus had a wingspan of 50–75 centimetres (20 to –30 inches), and lived during the
late Jurassic on lake shores.
• Pterodaustro was a Cretaceous pterosaur from South America with a wingspan around 1.33
metres and with over 500 tall, narrow teeth, which were presumably used in filter-feeding, much like
modern flamingos. Also like flamingos, this pterosaur's diet may have resulted in the animal having a
pink hue. It was South America's first pterosaur find.
• Quetzalcoatlus had a wingspan of 10–11 metres (33–36 ft), and was among the largest flying
animals ever. It lived during the late Cretaceous period.
• Rhamphorhynchus was a Jurassic pterosaur with a vane at the end of its tail, which may
have acted to stabilise the tail in flight.
[edit] In popular culture
Quetzalcoatlus models in South Bank, created by Mark Witton for the Royal Society's 350th
anniversary
Pterosaurs have been a staple of popular culture for as long as their cousins the dinosaurs, though
they are usually not featured as prominently in films, literature or other art. Additionally, while the depiction of
dinosaurs in popular media has changed radically in response to advances in paleontology, a mainly
outdated picture of pterosaurs has persisted since the mid 20th century.[45]
The number and diversity of pterosaurs in the popular consciousness is also not as high as it has
been historically for dinosaurs. While the generic term "pterodactyl" is often used to describe these creatures,
the animals depicted frequently represent either Pteranodon or Rhamphorhynchus, or a fictionalized hybrid
of the two.[45] Many children's toys and cartoons feature "pterodactyls" with Pteranodon-like crests and long,
Rhamphorhynchus-like tails and teeth, a combination that never existed in nature. However, at least one type
of pterosaur did have at least the Pteranodon-like crest and teeth—for example, the Ludodactylus, a name
that means "toy finger" for its resemblance to old, inaccurate children's toys.[46]
Pterosaurs were first used in fiction in Arthur Conan Doyle's 1912 novel The Lost World, and
subsequent 1925 film adaptation. They have been used in a number of films and television programs since,
including the 1933 film King Kong, and 1966 One Million Years B.C.. In the latter, animator Ray Harryhausen
had to add inaccurate bat-like wing fingers to his stop motion models in order to keep the membranes from
falling apart, though this particular error was common in art even before the film was made. Pterosaurs were
mainly absent from notable film appearances until 2001, with Jurassic Park III. However, paleontologist Dave
Hone has noted that even after the 40 intervening years, the pterosaurs in this film had not been significantly
updated to reflect modern research. Among the errors he noted as persisting from the 1960s to the 2000s
were teeth even in toothless species (the Jurassic Park III pterosaurs were intended to be Pteranodon, which
translates as "toothless wing"), nesting behavior that was known to be inaccurate by 2001, and leathery
wings, rather than the taut membranes of muscle fiber which was actually present and required for pterosaur
flight.[45]
[edit] See also
• List of pterosaurs
• Mary Anning
• Wyvern
• Flying and gliding animals
[edit] References
1. ^ Wang X, Kellner AW, Zhou Z, Campos Dde A (February 2008). "Discovery of a rare
arboreal forest-dwelling flying reptile (Pterosauria, Pterodactyloidea) from China". Proc. Natl. Acad.
2. ^ Lawson DA (March 1975). "Pterosaur from the Latest Cretaceous of West Texas:
Discovery of the Largest Flying Creature". Science 187 (4180): 947–948.
doi:10.1126/science.187.4180.947. PMID 17745279.
3. ^ Buffetaut E, Grigorescu D, Csiki Z (April 2002). "A new giant pterosaur with a robust skull
from the latest cretaceous of Romania". Naturwissenschaften 89 (4): 180–4. doi:10.1007/s00114-002-
0307-1. PMID 12061403.
http://link.springer.de/link/service/journals/00114/bibs/2089004/20890180.htm.
4. ^ a b c d Witmer LM, Chatterjee S, Franzosa J, Rowe T (October 2003). "Neuroanatomy of
flying reptiles and implications for flight, posture and behaviour". Nature 425 (6961): 950–3.
doi:10.1038/nature02048. PMID 14586467.
5. ^ Bennett SC (2000). "Pterosaur flight: the role of actinofibrils in wing function". Historical
Biology 14 (4): 255–84. doi:10.1080/10292380009380572.
http://www.informaworld.com/smpp/content~content=a907600799~db=all.
6. ^ a b Kellner, A.W.A., Wang, X., Tischlinger, H., Campos, D., Hone, D.W.E. and Meng, X.
(2009). "The soft tissue of Jeholopterus (Pterosauria, Anurognathidae, Batrachognathinae) and the
structure of the pterosaur wing membrane." Proceedings of the Royal Society B, published online
before print August 5, 2009, doi:10.1098/rspb.2009.0846
7. ^ a b c d e Naish D, Martill DM (2003). "Pterosaurs — a successful invasion of prehistoric
skies". Biologist 50 (5): 213–6.
8. ^ a b Claessens LP, O'Connor PM, Unwin DM (2009). "Respiratory evolution facilitated the
origin of pterosaur flight and aerial gigantism". PLoS ONE 4 (2): e4497.
http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0004497.
9. ^ Wilkinson MT, Unwin DM, Ellington CP (January 2006). "High lift function of the pteroid
bone and forewing of pterosaurs". Proc. Biol. Sci. 273 (1582): 119–26. doi:10.1098/rspb.2005.3278.
PMID 16519243.
10.^ Bennett SC (2007). "Articulation and Function of the Pteroid Bone of Pterosaurs". Journal
of Vertebrate Paleontology 27 (4): 881–91. doi:10.1671/0272-4634(2007)27[881:AAFOTP]2.0.CO;2.
http://www.bioone.org/perlserv/?request=get-abstract&doi=10.1671%2F0272-
4634(2007)27%5B881%3AAAFOTP%5D2.0.CO%3B2.
11.^ Unwin, D. M.; Frey, E., Martill, D. M., Clarke, J. B., and Riess, J. (1996). "On the nature of
the pteroid in pterosaurs". Proceedings of the Royal Society: Biological Sciences 263 (1366): 45–52.
doi:10.1098/rspb.1996.0008.
12.^ Wilkinson, M.T., Unwin, D.M. and Ellington, C.P. (2006). "High lift function of the pteroid
bone and forewing of pterosaurs." Proceedings of the Royal Society B, 273(1582): 119–126.
doi:10.1098/rspb.2005.3278
13.^ a b Unwin DM, Bakhurina NN (1994). "Sordes pilosus and the nature of the pterosaur flight
apparatus". Nature 371: 62–4. doi:10.1038/371062a0.
14.^ Wang X, Zhou Z, Zhang F, Xu X (2002). "A nearly completely articulated
rhamphorhynchoid pterosaur with exceptionally well-preserved wing membranes and "hairs" from
Inner Mongolia, northeast China". Chinese Science Bulletin 47: 3. doi:10.1360/02tb9054. ISSN 1001-
6538.
15.^ Frey et al., (2003) New specimens of Pterosauria (Reptilia) with soft parts with implications
for pterosaurian anatomy and locomotion Geological Society London Special Publications
16.^ Dyke GJ, Nudds RL, Rayner JM (July 2006). "Limb disparity and wing shape in pterosaurs".
J. Evol. Biol. 19 (4): 1339–42. doi:10.1111/j.1420-9101.2006.01096.x. PMID 16780534.
17.^ a b c d Witton MP, Naish D (2008). "A reappraisal of azhdarchid pterosaur functional
morphology and paleoecology". PLoS ONE 3 (5): e2271. doi:10.1371/journal.pone.0002271.
PMID 18509539. PMC 2386974. http://www.plosone.org/article/info%3Adoi
%2F10.1371%2Fjournal.pone.0002271.
18.^ a b c d e f Unwin, David M. (2006). The Pterosaurs: From Deep Time. New York: Pi Press.
pp. 246. ISBN ISBN 0-13-146308-X.
19.^ a b Frey E, Martill DM (1998). "Soft tissue preservation in a specimen of Pterodactylus
kochi (Wagner) from the Upper Jurassic of Germany". Neues Jahrbuch fu ̈r Geologie und Pala
̈ontologie, Abhandlungen 210: 421–41.
20.^ Czerkas, S.A., and Ji, Q. (2002). A new rhamphorhynchoid with a headcrest and complex
integumentary structures. In: Czerkas, S.J. (Ed.). Feathered Dinosaurs and the Origin of Flight . The
Dinosaur Museum:Blanding, Utah, 15-41. ISBN 1-93207-501-1.
21.^ Collini, C A. (1784). "Sur quelques Zoolithes du Cabinet d’Histoire naturelle de S. A. S. E.
Palatine & de Bavière, à Mannheim." Acta Theodoro-Palatinae Mannheim 5 Pars Physica , pp. 58–
103 (1 plate).
22.^ Wagler, J. (1830). Natürliches System der Amphibien Munich, 1830: 1-354.
23.^ Cuvier, G. (1801). [Reptile volant]. In: Extrait d’un ouvrage sur les espèces de quadrupèdes
dont on a trouvé les ossemens dans l’intérieur de la terre. Journal de Physique, de Chimie et
d’Histoire Naturelle, 52: 253–267.
24.^ Alleyne, R., Pterodactyls Were Too Heavy To Fly, Scientist Claims, Telegraph, Oct 2008
25.^ Powell, D., Were Pterosaurs Too Big To Fly?, Oct 2008
26.^ Alleyne, Richard (2008-10-01). "Pterodactyls were too heavy to fly, scientist claims". The
Daily Telegraph (London). http://www.telegraph.co.uk/earth/main.jhtml?
xml=/earth/2008/10/01/sciptero101.xml. Retrieved 2010-05-22.
27.^ Templin, R. J.; Chatterjee, Sankar (2004). Posture, locomotion, and paleoecology of
pterosaurs. Boulder, Colo: Geological Society of America. p. 60. ISBN 0-8137-2376-0.
http://books.google.com/?id=idta6AVV-tIC&pg=PA60&dq=Pterosaur+oxygen.
28.^ Naish, Darren (February 18, 2009). "Pterosaurs breathed in bird-like fashion and had
inflatable air sacs in their wings". ScienceBlogs.
http://scienceblogs.com/tetrapodzoology/2009/02/pterosaur_breathing_air_sacs.php. Retrieved 20
October 2010
29.^ a b "Did giant pterosaurs vault aloft like vampire bats?". New Scientist.
http://www.newscientist.com/article/dn19724-did-giant-pterosaurs-vault-aloft-like-vampire-bats.html.
30.^ Padian K (1983). "A Functional Analysis of Flying and Walking in Pterosaurs". Paleobiology
9 (3): 218–39.
31.^ Padian K (2003). "Pterosaur Stance and Gait and the Interpretation of Trackways". Ichnos
10 (2-4): 115–126. doi:10.1080/10420940390255501.
32.^ Hwang K, Huh M, Lockley MG, Unwin DM, Wright JL (2002). "New pterosaur tracks
(Pteraichnidae) from the Late Cretaceous Uhangri Formation, southwestern Korea". Geological
Magazine 139 (4): 421–35. doi:10.1017/S0016756802006647.
33.^ Unwin DM (1997). "Pterosaur tracks and the terrestrial ability of pterosaurs". Lethaia 29:
373–86. doi:10.1111/j.1502-3931.1996.tb01673.x.
34.^ Buffetaut E, Martill D, Escuillié F (July 2004). "Pterosaurs as part of a spinosaur diet".
Nature 430 (6995): 33. doi:10.1038/430033a. PMID 15229562.
35.^ Ji Q, Ji SA, Cheng YN, et al (December 2004). "Palaeontology: pterosaur egg with a
leathery shell". Nature 432 (7017): 572. doi:10.1038/432572a. PMID 15577900.
36.^ Lü, J., Unwin, D.M., Deeming, D.C., Jin, X., Liu, Y. and Ji, Q. (2011). "An egg-adult
association, gender, and reproduction in pterosaurs. Science, 331(6015): 321-324.
doi:10.1126/science.1197323
37.^ Grellet-Tinner G, Wroe S, Thompson MB, Ji Q (2007). "A note on pterosaur nesting
behavior". Historical Biology 19 (4): 273–7. doi:10.1080/08912960701189800.
38.^ Wang X, Zhou Z (June 2004). "Palaeontology: pterosaur embryo from the Early
Cretaceous". Nature 429 (6992): 621. doi:10.1038/429621a. PMID 15190343.
39.^ a b c Bennett, S. C. (1995). "A statistical study of Rhamphorhynchus from the Solnhofen
Limestone of Germany: Year-classes of a single large species." Journal of Paleontology, 69: 569-
580.
40.^ Hone, D.W.E. and Benton, M.J. (2007). "An evaluation of the phylogenetic relationships of
the pterosaurs to the archosauromorph reptiles." Journal of Systematic Palaeontology, 5(4): 465-469.
doi:10.1017/S1477201907002064
41.^ Lü, J., Unwin, D.M., Xu, L., and Zhang, X. (2008). "A new azhdarchoid pterosaur from the
Lower Cretaceous of China and its implications for pterosaur phylogeny and evolution."
Naturwissenschaften, 95(9): 891-897. doi:10.1007/s00114-008-0397-5
42.^ Unwin DM (2003). "On the phylogeny and evolutionary history of pterosaurs". In Jean-
michel Mazin; Buffetaut, Eric. Evolution And Palaeobiology of Pterosaurs (Geological Society
Special Publication). London: Geological Society of London. pp. 139–90. ISBN 1-86239-143-2.
43.^ Slack KE, Jones CM, Ando T, et al (June 2006). "Early penguin fossils, plus mitochondrial
genomes, calibrate avian evolution". Mol. Biol. Evol. 23 (6): 1144–55. doi:10.1093/molbev/msj124.
PMID 16533822. http://mbe.oxfordjournals.org/cgi/reprint/23/6/1144.
44.^ Copyrighted excerpt from Richard Butler, Paul Barrett, Steven Nowbath & Paul Upchurch
[1]; might require new link
45.^ a b c Hone, D. (2010). "Pterosaurs In Popular Culture." Pterosaur.net,
http://www.pterosaur.net/popular_culture.php Accessed online 27-August-2010.
46.^ Frey, E., Martill, D., and Buchy, M. (2003). A new crested ornithocheirid from the Lower
Cretaceous of northeastern Brazil and the unusual death of an unusual pterosaur. In: Buffetaut, E.,
and Mazin, J.-M. (eds.). Evolution and Palaeobiology of Pterosaurs. Geological Society Special
Publication 217:56-63. ISBN 1-86239-143-2.

Wikimedia Commons has media related to: Pterosauria
• Pterosaur.net, multi-authored website about all aspects of pterosaur science

• The Pterosaur Database, by Paul Pursglove.
• Mark Witton's Pterosaur Art
• Comments on the phylogeny of the pterodactyloidea , by Alexander W. A. Kellner. (technical)
[hide]v · d · eArchosauromorphs
Kingdom: Animalia · Phylum: Chordata · Class: Sauropsida · Subclass: Diapsida
Primitive Euparkeriidae • Erythrosuchidae • Proterochampsidae •

Archosauromorphs Proterosuchidae • Choristodera • Prolacertiformes • Rhynchosauria •
Trilophosauria • Vancleavea
Crurotarsi Ornithosuchidae • Aetosauria • Phytosauria • Rauisuchia •

Archosaurs Crocodylomorpha • Crocodilia
Avemetatarsalia and Scleromochlus • Pterosauria • Dinosauromorpha • Dinosauria •

Ornithodira Archosaurs Ornithischia • Saurischia • Aves
Avian Archosaurs Avialae • Archaeopteryx • Confuciusornis • Ichthyornis •

Enantiornithes • Hesperornithes • Neornithes • Palaeognathae • Neognathae
Retrieved from "http://en.wikipedia.org/wiki/Pterosaur"
Categories: Pterosaurs | Cretaceous extinctions

W000
Germanodactylus

Germanodactylus
Fossil range: Late Jurassic
Germanodactylus cristatus
Kingdom: Animalia
Phylum: Chordata
Class: Sauropsida
Order: Pterosauria
Suborder: Pterodactyloidea
Superfamily: Dsungaripteroidea
Family: Germanodactylidae
Germanodactylus
Genus:
Yang, 1964
Species
• G. cristatus (Wiman, 1925 (originally Pterodactylus) (type species))

• ?G. rhamphastinus (possibly "Daitingopterus") (Wagner, 1851 (originally
Ornithocephalus)
Germanodactylus ("German finger") is a genus of dsungaripteroid pterodactyloid pterosaur from Late
Jurassic-age rocks of Germany, including the Solnhofen limestone. Its specimens were long thought to
pertain to Pterodactylus. Its head crest is a distinctive feature.
Contents
[hide]
• 1 Description
• 1.1 Head crest
• 3 History
• 4 References
[edit] Description
Restoration of G. rhamphastinus
Germanodactylus is described as being "raven-sized" in weight.[1] G. cristatus had a 0.98 wingspan
(3.2 ft) and a 13 centimeter long (5.1 in) skull, while G. rhamphastinus was somewhat larger, with a
1.08 meter wingspan (3.5 ft) and a skull 21 centimeters long (8.3 in).[2]
[edit] Head crest

Germanodactylus is known for its head crest, which had a bony portion (a low ridge running up the
midline of the skull) and a soft-tissue portion that more than doubled its height. The bony part does not go as
far up the head in G. cristatus as in G. rhamphastinus. The soft-tissue portion was not known early on, being
first described in 2002 by S. Christopher Bennett. It was probably composed of cornified epidermis.
Germanodactylus is the first genus for which a soft-tissue component of the crest is known, but similar
structures were probably widespread among pterosaurs.[3]
This genus is unspecialized compared to the pterosaurs of the Cretaceous, and has had varying
placements in Pterosauria. Yang Zhongjian, who named the genus, gave it its own family
Germanodactylidae.[4] Bennett included the genus in the family Pterodactylidae,[5] and Alexander W.A.
Kellner found it to be related to Pterodactylus in his 2003 phylogenetic analysis.[6] David M. Unwin, on the
other hand, preferred to consider it a basal dsungaripteroid,[7][8] a group that evolved into dedicated
shellfish-eaters.[9]
[edit] History
G. cristatus is based on specimen BSP 1892.IV.1, from the Solnhofen limestone of Eichstätt,
Germany. It was originally described by Plieninger in 1901 as a specimen of Pterodactylus kochi,[10] and
was given its current specific name by Carl Wiman in 1925, meaning "crested" in Latin.[11] Yang Zhongjian
determined that it deserved its own genus in 1964.[4] Second species G. ramphastinus (in 1858 accidentally
revised to rhamphastinus by Christian Erich Hermann von Meyer) was named as a distinct species long
before G. cristatus, described by Johann Andreas Wagner in 1851 as a species of the now deprecated genus
Ornithocephalus. The specific name refers to the toucan, ramphastinos in Greek. It is based on specimen
BSP AS.I.745, a skeleton from the slightly younger Mörnsheimer Limestone of Daiting, Germany.[12] Peter
Wellnhofer added it to Germanodactylus in 1970,[13] although Maisch and his coauthors have suggested
that it deserves its own genus, "Daitingopterus"[14] David M. Unwin has also referred miscellaneous limb
bones and vertebrae from the somewhat older Kimmeridge Clay of Dorset, England to the genus; these finds
at the time marked the earliest appearance of short-tailed pterosaurs in the fossil record.[15]
Germanodactylus ramphastinus
Bennett suggested in 1996 that Germanodactylus represented adults of Pterodactylus,[16] but this
has been rejected by further studies,[1][6][7][17] including his own.[3][5] Bennett's 2006 reappraisal of
Germanodactylus found both species to be valid and included within the genus, with G. cristatus known from
four specimens including two juveniles, and G. rhamphastinus from two specimens. The genus differs from
other pterosaurs by a combination of characteristics including a sharply pointed jaw tip, 4-5 premaxillary
teeth and 8-12 maxillary teeth per side of the upper jaw, robust maxillary teeth that, unlike in Pterodactylus,
are not reduced in size farther from the tip of the jaw, a naso-antorbital fenestra twice the length of the eye
socket, and various proportional differences. G. cristatus differs from G. rhampastinus by having no teeth in
the tip of the jaw and fewer teeth (~13 in each side of the upper jaw and ~12 in the lower versus 16 upper
and 15 lower on each side for G. rhamphastinus).[5]
[edit] References
1. ^ a b Unwin, David M. (2006). The Pterosaurs: From Deep Time. New York: Pi Press. p. 246.
ISBN ISBN 0-13-146308-X.
2. ^ Wellnhofer, Peter (1996) [1991]. The Illustrated Encyclopedia of Pterosaurs. New York:
Barnes and Noble Books. pp. 95–96. ISBN 0-7607-0154-7.
3. ^ a b Bennett, S. Christopher (2002). "Soft tissue preservation of the cranial crest of the
pterosaur Germanodactylus from Solnhofen". Journal of Vertebrate Paleontology 22 (1): 43–48.
doi:10.1671/0272-4634(2002)022[0043:STPOTC]2.0.CO;2.
4. ^ a b Zhongjian, Yang (1964). "On a new pterosaurian from Sinkiang, China". Vertebrata
PalAsiatic 8: 221–255.
5. ^ a b c Bennett, S. Christopher (2006). "Juvenile specimens of the pterosaur
Germanodactylus cristatus, with a revision of the genus". Journal of Vertebrate Paleontology 26 (4):
872–878. doi:10.1671/0272-4634(2006)26[872:JSOTPG]2.0.CO;2.
6. ^ a b Kellner, Alexander W.A. (2003). "Pterosaur phylogeny and comments on the
evolutionary history of the group". In Buffetaut, Eric; and Mazin, Jean-Michel. Evolution and
Palaeobiology of Pterosaurs. Geological Society Special Publication 217. London: The Geological
Society. pp. 105–137. ISBN 1-86239-143-2.
7. ^ a b Unwin, David M. (2003) "On the phylogeny and evolutionary history of pterosaurs", in
Evolution and Palaeobiology of Pterosaurs, 139-190.
8. ^ Unwin, David M. (2006). The Pterosaurs: From Deep Time, 273.
9. ^ Unwin, David M. (2006). The Pterosaurs: From Deep Time, 84-85.
10.^ Plieninger, F. (1901). "Beiträge zur Kenntnis der Flugsaurier". Palaeontographica 48: 65–
90.
11.^ Wiman, Carl (1925). "Über Pterodactylus Westmanni und andere Flugsaurier". Bulletin of
the Geological Institution of the University of Uppsala 20: 1–38.
12.^ Wagner, Johann Andreas (1851). "Beschreibung einer neuen Art von Ornithocephalus,
nebst kritischer Vergleichung der in der k. palaeontologischen Sammlung zu München aufgestellten
Arten aus dieser Gattung" (in German). Abhandlungen der königlichen bayerischen Akademie der
Wissenschaften 6: 1–64.
13.^ Wellnhofer, Peter (1970). "Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke
Süddeutschlands". Abhandlung der Bayerischen Akademie der Wissenschaften 141: 1–133.
14.^ Maisch, M.W.; Matzke, A.T.; and Ge Sun (2004). "A new dsungaripteroid pterosaur from
the Lower Cretaceous of the southern Junggar Basin, north-west China". Cretaceous Research 25:
625–634. doi:10.1016/j.cretres.2004.06.002.
15.^ Unwin, David M. (1988). "A new pterosaur from the Kimmeridge Clay of Kimmeridge,
Dorset". Proceedings of the Dorset Natural History Museum and Archaeological Society 109: 150–
153.
16.^ Bennett, S. Christopher (1996). "Year-classes of pterosaurs from the Solnhofen Limestone
of Germany: taxonomic and systematic implications". Journal of Vertebrate Paleontology 16 (2): 432–
444. doi:10.1080/02724634.1996.10011332.
17.^ Jouve, Stephane (2004). "Description of the skull of a Ctenochasma (Pterosauria) from the
Late Jurassic of eastern France, with a taxonomic revision of European Tithonian Pterodactyloidea".
Journal of Vertebrate Paleontology 24 (3): 542–554. doi:10.1671/0272-
4634(2004)024[0542:DOTSOA]2.0.CO;2.
• Restoration of G. rhamphastinus at The Grave Yard
Retrieved from "http://en.wikipedia.org/wiki/Germanodactylus"
Categories: Dsungaripteroids | Jurassic pterosaurs | Prehistoric reptiles of Europe | Solnhofen fauna

W000
Lonchodectes

Lonchodectes
Temporal range: Early-Late Cretaceous, 94 Ma
PreЄ ↓
g
L. compressirostris attacked by Ornithocheirus
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Pterosauria
Suborder: Pterodactyloidea
Superfamily: ?Azhdarchoidea
Lonchodectidae
Family:
Hooley, 1914
Lonchodectes
Genus:
Hooley, 1914
Species
• L. compressirostris (Owen, 1851) [originally

Ornithocheirus]
See text for dubious species.
Lonchodectes (meaning "lance biter") was a genus of pterosaur from several formations dating to the
Turonian (Late Cretaceous, though possibly as early as the Valanginian) of England, mostly in the area
around Kent. The species belonging to it had been assigned to Ornithocheirus until David Unwin's work of
the 1990s and 2000s, and the genus is not universally accepted as distinct.[1] Several potential species are
known; most are based on scrappy remains, and have gone through several other generic assignments. The
genus is part of the complex taxonomy issues surrounding Early Cretaceous pterosaurs from Brazil and
England, such as Amblydectes, Anhanguera, Coloborhynchus, and Ornithocheirus (and its possible
synonyms Criorhynchus and Tropeognathus).
Contents
[hide]
• 1 History and species
• 3 Paleobiology
• 4 References
[edit] History and species

Numerous species have been referred to this genus over time, and only those more widely
connected with the genus are included here.
The type species, L. compressirostris, is based on BMNH 39410, a partial upper jaw from the
Turonian-age Upper Cretaceous Upper Chalk near Kent. Richard Owen named in 1851 as a species of
Pterodactylus;[2] it was transferred to Ornithocheirus in 1870 by Harry Govier Seeley,[3] before becoming the
type species of Lonchodectes in Reginald Walter Hooley's 1914 review of Ornithocheirus.[4] Confusingly, this
species was also long regarded, incorrectly, as the type species of Ornithocheirus.[5]
Hooley added two other species at this time, both of which had also been originally referred to
Pterodactylus, then to Ornithocheirus: L. giganteus, a Cenomanian-age jaw fragment;[6] and L. daviesii,
another jaw fragment, from an Albian-age formation.[7]
The genus acquired several more former Pterodactylus and Ornithocheirus species in the 1990s-
2000s. L. sagittirostris, based on BMNH R1823, a lower jaw fragment from the ?Valanginian-Hauterivian-age
Lower Cretaceous Hastings Beds of East Sussex,[7] and L. platystomus[3] had been acquired by 2000.[8]
Two additional species based on jaw fragments, both from the Albian-age Cambridge Greensand,[3] were
added by 2003: L. machaerorhynchus and L. microdon,[9] joining L. compressirostris, L. giganteus, L.
platystomus, and L. sagittirostris in his listing of valid species (thus, additional dubious species may also be
referred here).[10]
The genus is poorly known and not universally accepted, as noted above. In Peter Wellnhofer's 1991
The Illustrated Encyclopedia of Pterosaurs the other major recent synopsis of pterosaurs, written before
Unwin's work, the species are included with Ornithocheirus (because of L. compressirostris being thought to
be the type species), and are in fact the main fossils illustrated to represent the genus.[11] Unwin has them in
their own family, Lonchodectidae, which he grouped with the ctenochasmatoids in 2003,[9] and with the
azhdarchoids, including the tapejarids and azhdarchids, in 2006.[10]
[edit] Paleobiology
Unwin considers Lonchodectes to have been a generalist, like a seagull, with its conservative jaws
and teeth (like those of the much older Pterodactylus) and small to medium size; he estimates its maximum
wingspan at about 2 m (6.6 ft). Lonchodectes had long jaws with many short teeth, and the jaws were
compressed vertically, like "a pair of sugar tongs with teeth".[12] At least some of these species had crests
on their lower jaws.[13]
[edit] References
1. ^ Kellner, A.W.A. (2003). Pterosaur phylogeny and comments on the evolutionary history of
the group: In: Buffetaut, E., and Mazin, J.-M. (Eds.). Evolution and Palaeobiology of Pterosaurs.
Geological Society Special Publication 217:105-137. 1-86239-143-2.
2. ^ Owen, R. (1851). Monograph on the fossil Reptilia of the Cretaceous Formations. The
Palaeontographical Society 5(11):1-118.
3. ^ a b c Seeley, H.G. (1870). The Ornithosauria: an Elementary Study of the Bones of
Pterodactyles. Cambridge, 130 pp.
4. ^ Hooley, R.W. (1914). On the Ornithosaurian genus Ornithocheirus with a review of the
specimens from the Cambridge Greensand in the Sedgwick Museum, Cambridge. Annals and
Magazine of Natural History, series 8, 78:529-557.
5. ^ Unwin, David M. (2001). "An overview of the pterosaur assemblage from the Cambridge
Greensand (Cretaceous) of Eastern England". Mitteilungen as dem Museum für Naturkunde, Berlin,
Geowissenschaftliche Reihe 4: 189–222.
6. ^ Bowerbank, J.S. (1846). On a New Species of Pterodactyl. Found in the Upper Chalk of
Kent (P. giganteus). Quarterly Journal of the Geological Society 2:7–9.
7. ^ a b Owen, R. (1874). A Monograph on the Fossil Reptilia of the Mesozoic Formations. 1.
Pterosauria. The Palaeontographical Society Monograph 27:1–14.
8. ^ Unwin, D.M., Lü, J., and Bakhurina, N.N. (2000). On the systematic and stratigraphic
significance of pterosaurs from the Lower Cretaceous Yixian Formation (Jehol Group) of Liaoning,
China. Mitteilungen Museum für Naturkunde der Humboldt-Universität zu Berlin,
Geowissenschaftliche Reihe 3:181–206.
9. ^ a b Unwin, David M. (2003). "On the phylogeny and evolutionary history of pterosaurs". In
Buffetaut, Eric; and Mazin, Jean-Michel, (eds.). Evolution and Palaeobiology of Pterosaurs.
Geological Society Special Publication 217. London: Geological Society. pp. 139–190. ISBN 1-86239-
143-2.
10.^ a b Unwin, D.M. (2006). The Pterosaurs: From Deep Time. Pi Press:New York, p. 273.
ISBN 0-13-146308-X.
11.^ Wellnhofer, Peter (1996) [1991]. The Illustrated Encyclopedia of Pterosaurs. New York:
Barnes and Noble Books. pp. 110–113. ISBN 0-7607-0154-7.
12.^ Unwin, D.M. (2006). The Pterosaurs: From Deep Time. Pi Press:New York, p. 251. ISBN 0-
13-146308-X.
13.^ Unwin, D.M. (2006). The Pterosaurs: From Deep Time. Pi Press:New York, p. 106. ISBN 0-
13-146308-X.

• Upper Cretaceous species of Lonchodectes (scroll down), The Pterosaur Database.
Accessed 2007-02-10
• Upper Cretaceous species of Lonchodectes (scroll down), The Pterosaur Database.
Accessed 2007-02-10
• Re: Pterosaur Help, a posting from George Olshevsky on the Dinosaur Mailing List, which,
although incomplete, should give some idea as to the complexity of the taxonomy here. Additional,
even more dubious species are included. Accessed 2007-02-10
Retrieved from "http://en.wikipedia.org/wiki/Lonchodectes"
Categories: Azhdarchoids | Cretaceous pterosaurs | Prehistoric reptiles of Europe

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List of pterosaur classifications


This list of pterosaur classifications entails the various schemes used to classify pterosaurs
throughout the years and varying views of these animals.
Contents
[hide]
• 1 Classic
• 2 Unwin (2003)
• 3 Kellner (2003)
• 4 Wellnhofer (1991)
• 5 Peters (2007)
• 6 See also
• 7 References
[edit] Classic
• Pterosauria
• Rhamphorhynchoidea
• Pterodactyloidea
[edit] Unwin (2003)

Pterosauria
Macronychoptera
Caelidracones
Lonchognatha
Campylogna
Breviquartossa
Rhamphorhynchidae
Rhamphorhynchinae
Scaphognathinae
Pterodactyloidea
Ornithocheiroidea
Istiodactylidae
Euornithocheira
Ornithocheiridae
Pteranodontidae
Nyctosaurus
Pteranodontinae
Lophocratia
Ctenochasmatoidea
?Feilongus
Gallodactylidae
Euctenochasmatia
Pterodactylus
Lonchodectes
Ctenochasmatidae
Gnathosaurinae
Ctenochasmatinae
Dsungaripteroidea
Germanodactylidae
Dsungaripteridae
Azhdarchoidea
?
Thalassodromeus
Tupuxuara
Neoazhdarchia
Tapejaridae
Azhdarchidae
[edit] Kellner (2003)
Novialoidea
Pterodactyloidea
Archaeopterodactyloidea
Germanodactylidae
Pterodactylidae
Gnathosaurus
Ctenochasmatidae
Feilongus
Gallodactylidae
Dsungaripteroidea
Nyctosaurus
Ornithocheiroidea
Pteranodontoidea
?Eopteranodon
?Cearadactylus
Pteranodontidae
Istiodactylidae
?
Ornithocheiridae
Criorhynchus
Anhangueridae
Tapejaroidea
Dsungaripteridae
Azhdarchoidea
Tapejaridae
Azhdarchidae
[edit] Wellnhofer (1991)
• Pterosauria
• ?Austriadactylus
• Eudimorphodon
• +
• Dimorphodontidae
• +
• ?Rhamphinion
• ?Herbstosaurus
• ?Nesodactylus
• Rhamphorhynchidae
• +
• Dimorphodontidae
• Pterodactyloidea
• ?Dermodactylus
• ?Mesadactylus
• ?Bennettazhia
• ?Istiodactylus
• ?Araripesaurus
• ?Pterodaustridae
• Pterodactylidae
• Gallodactylidae
• Germanodactylidae
• Ctenochasmatidae
• Dsungaripteridae
• Ornithocheiridae
• Tapejaridae
• Anhangueridae
• +
• ?Nyctosaurus
• Pteranodontidae
• Criorhynchidae
• Azhdarchidae
[edit] Peters (2007)
The basal lizard, Huehuecuetzpalli is the most primitive taxon in this newly revealed third squamate
clade between Iguania and Scleroglossa. Two branches arise from it.
Jesairosaurus is basal to the Drepanosauridae. Three distinct specimens of Macrocnemus give rise
to the Tanystropheidae, the Langobardisaurinae and to the Fenestrasauria respectively.
Within the Fenestrasauria Cosesaurus is basal to Sharovipteryx, Saltopus, Longisquama and the
Pterosauria in that order.
The Milan specimen of Eudimorphodon (MPUM 6009) is the most primitive known pterosaur and
Austriadactylus is a sister taxon. One of two major clades, the Dimorphodontoidea, includes the
Dimorphodontidae and the Anurognathidae.
The other major clade includes all other pterosaurs with Eudimorphodon + (Campylognathoides +
Rhamphorhynchus) at its base. Higher dorygnathoids divide into the Dorygnathidae and a clade with Sordes
at its base. In the former two distinct Dorygnathus specimens are basal to Ctenochasma and kin on one
branch and to Quetzalcoatlus and kin on the other.
Within the Sordes-based clade, Pterorhynchus and Scaphognathus crassirostris are basal.
Subsequently two distinct smaller Scaphognathus specimens are basal to two major clades. The first is a
series of tiny pterosaurs, Cycnorhamphus and the Ornithocheiridae. The second includes some tiny
pterosaurs, the Pterodactylidae and the Germanodactylidae. From the latter arise the Dsungaripteria
(Dsungaripteridae + Tapejaridae) and a clade consisting of (Eoazhdarcho + Eopteranodon) + (Pteranodon +
Nyctosaurus).
Thus the former monophyletic "Pterodactyloidea" is revealed to be four distinct clades demonstrating
some convergence. Major clades typically have a spectral series of tiny pterosaurs at their base suggesting
that paedomorphosis was a major factor in pterosaur evolution.
[edit] See also

• Pterosaur
[edit] References
• Pterosauria @ Mikko's Phylogeny Archive
• Kellner, A. W. A., (2003): Pterosaur phylogeny and comments on the evolutionary history of
the group. pp. 105-137. — in Buffetaut, E. & Mazin, J.-M., (eds.) (2003): Evolution and Palaeobiology
of Pterosaurs. Geological Society of London, Special Publications 217, London, 1-347
• Peters, D., (2007): The origin and radiation of the Pterosauria — in Flugsaurier: The
Wellnhofer pterosaur meeting, Munich, 2007, 27-28
• Unwin, D. M., 2003: On the phylogeny and evolutionary history of pterosaurs. pp. 139-190. —
in Buffetaut, E. & Mazin, J.-M., (eds.) (2003): Evolution and Palaeobiology of Pterosaurs. Geological
Society of London, Special Publications 217, London, 1-347
• Wellnhofer, P., (1991): The Illustrated Encyclopedia of Pterosaurs. Salamander Books Ltd.,
London, pp. 192
Retrieved from "http://en.wikipedia.org/wiki/List_of_pterosaur_classifications"
Categories: Pterosaurs | Lists of prehistoric reptiles

W000
Dorygnathus

Dorygnathus
Temporal range: 180 Ma
PreЄ ↓
g
Early Jurassic
Dorygnathus banthensis fossil
Kingdom: Animalia
Phylum: Chordata
Class: Sauropsida
Order: Pterosauria
Suborder: Rhamphorhynchoidea
Family: Rhamphorhynchidae
Dorygnathus
Genus:
Wagner, 1860
Species
Dorygnathus banthensis
(Theodori, 1830) Wagner, 1860 (Type)
Dorygnathus ("spear jaw") was a genus of pterosaur that lived in Europe during the Early Jurassic
period, 180 million years ago when shallow seas flooded much of the continent. It had a short 1.5 meter (five
feet) wingspan, and a relatively small triangular sternum, which is where its flight muscles attached. Its skull
was long and its eye sockets were the largest opening therein. Large curved fangs that "intermeshed" when
the jaws closed featured prominently at the front of the snout while smaller, straighter teeth lined the back.[1]
Having variable teeth, a condition called heterodonty, is rare in modern reptiles but more common in primitive
pterosaurs.[citation needed] The heterodont dentition in Dorygnathus is consistent with a piscivorous (fish-
eating) diet.[1] The fifth digit on the hindlimbs of Dorygnathus was unusually long and oriented to the side. Its
function is not certain, but the toe may have supported a membrane like those supported by its wing-fingers
and pteroids. Dorygnathus was according to David Unwin related to the Late Jurassic pterosaur,
Rhamphorhynchus and was a contemporary of Campylognathoides in Holzmaden and Ohmden.[1]
Contents
[hide]
• 1 Discovery
• 2 Description
• 3 Dorygnathus mistelgauensis
• 4 Phylogeny
• 5 Paleobiology
• 6 See also
• 7 References
[edit] Discovery
A cast in the Urwelt-Museum Hauff at Holzmaden of UUPM R 156, a specimen sold by Bernhard
Hauff to the University of Uppsala in 1925
The first remains of Dorygnathus, isolated bones and jaw fragments from the Schwarzjura, the
Posidonia Shale dating from the Toarcian, were discovered near Banz, Bavaria and in 1830 described by
Carl Theodori as Ornithocephalus banthensis, the specific name referring to Banz.[2][3] The holotype, a
lower jaw, is PSB 757. The fossils were studied by Christian Erich Hermann von Meyer in 1831[4] and again
by Theodori in 1852 when he referred them to the genus Rhamphorhynchus.[5] In this period a close affinity
was assumed with a pterosaur known from Britain, later named Dimorphodon. Some fossils were sent to a
professor of paleontology in Munich named Johann Andreas Wagner. It was he who, having studied new
finds by Alfred Oppel in 1856 and 1858,[6][7] after Richard Owen had named Dimorphodon concluded that
the German type was clearly different and that therefore a new genus of pterosaur should be erected, which
he formally named Dorygnathus in 1860, from Greek dory, "spear" and gnathos, "jaw".[8] Much more
complete remains have been found since in other German locales and especially in Württemberg, including
Holzmaden, Ohmden, and Zell.[1] One specimen, SMNS 81840, has in 1978 been dug up in Nancy, France.
[9] Dorygnathus fossils were often found in the spoil heaps where unusable rock was dumped from slate
quarries worked by local farmers.[10] Most fossils were found in two major waves, one during the twenties,
the other during the eighties of the twentieth century. Since then the rate of discovery has slowed
considerably because the demand for slate has strongly diminished and many small quarries have closed. At
present over fifty specimens have been collected, many of them are preserved in the collection of the State
Museum of Natural History Stuttgart, as by law paleontological finds in Baden-Württemberg are property of
this Bundesland. Due to the excellent preserval of the later found fossils, Dorygnathus has generated much
interest by pterosaur researchers, important studies having been dedicated to the species by Felix
Plieninger,[11] Gustav von Arthaber,[12] and more recently Kevin Padian.[13]
[edit] Description
Dorygnathus banthensis restoration; the tail form is hypothetical

Dorygnathus in general has the build of a basal, i.e. non-pterodactyloid pterosaur: a short neck, a
long tail and short metacarpals — although for a basal pterosaur the neck and metacarpals of Dorygnathus
are again relatively long. The skull is elongated and pointed. The largest known cranium, that of specimen
MBR 1920.16 prepared by Bernard Hauff in 1915 and eventually acquired by the Humboldt Museum in
Berlin, has a length of sixteen centimetres. In the skull the eye socket forms the largest opening, larger than
the fenestra antorbitalis that is clearly separated from the slit-like bony naris. No bony crest is visible on the
rather straight top of the skull or snout. The lower jaws are thin at the back but deeper toward the front where
they fuse into the symphysis ending in a toothless point after which the genus has been named. In MBR
1920.16, the mandibula as a whole has a length of 147 millimetres.[14]
In the lower jaws the first three pairs of teeth are very long, sharp and pointing outwards and
forwards. They contrast with a row of eight or more upright-standing much smaller teeth that gradually
diminish in size towards the back of the lower jaw. No such extreme contrast exists in the upper jaws, but the
four teeth in the premaxilla are longer than the seven in the maxilla that again become smaller posteriorly.
The total number of teeth is thus at least 44. The long upper and lower front teeth interlaced when the beak
was closed; due to their extreme length they then projected considerably beyond the upper and lower
margins of the head.
According to Padian, eight cervical, fourteen dorsal, three or four sacral and twenty-seven or twenty-
eight caudal vertebrae are present. The exceptional fourth sacral is the first of the normal caudal series. The
number of caudals is not certain because their limits are obscured by long thread-like extensions, stiffening
the tail. The cervical vertebrae are rather long and strongly built, their upper surface having a roughly square
cross-section. They carry double-headed thin cervical ribs. The dorsal vertebrae are more rounded with flat
spines; the first three or four carry ribs that contact the sternal ribs; the more posterior ribs contact the
gastralia. The first five or six, rather short, caudal vertebrae form a flexible tail base. To the back the caudals
grow longer and are immobilised by their intertwining extensions with a length of up to five vertebrae which
together surround the caudals with a bony network, allowing the tail to function as a rudder.
The breastbone is triangular and relatively small; Padian has suggested it may have been extended
at its back with a cartilaginous tissue. It is connected to the coracoid which in older individuals is fused to the
longer scapula forming a saddle-shaped shoulder joint. The humerus has a triangular deltopectoral crest and
is pneumatised. The lower arm is 60% longer than the upper arm. From the five carpal bones in the wrist a
short but robust pteroid points towards the neck, in the living animal a support for a flight membrane, the
propatagium. The first three metacarpals are connected to three small fingers, equipped with short but
strongly curved claws; the fourth to the wing finger, in which the second or third phalanx is the longest; the
first or fourth the shortest. The wing finger supports the main flight membrane.
In the pelvis, the ilium, ischium and pubis are fused. The ilium is elongated with a length of six
vertebrae. The lower leg, in which the lower two thirds of the tibia and fibula of adult specimens are fused, is
a third shorter than the thighbone, the head of which makes an angle of 45° with its shaft. The proximal
tarsals are never fused in a separate astragalocalcaneum; a tibiotarsus is formed. The third metatarsal is the
longest; the fifth is connected to a toe of which the second phalanx shows a 45° bend and has a blunt and
broad end; it perhaps supported a membrane between the legs, a cruropatagium.
In some specimens, soft parts have been preserved but these are rare and limited, providing little
information. It is unknown whether the tail featured a vane on its end, as with Rhamphorhynchus. However,
Ferdinand Broili reported the presence of hairs in specimen BSP 1938 I 49,[15] an indication that
Dorygnathus also had fur and an elevated metabolism, as is presently assumed for all pterosaurs.
[edit] Dorygnathus mistelgauensis

In 1971 Rupert Wild described and named a second species: Dorygnathus mistelgauensis,[16]
based on a specimen collected in a brick pit near the railway station of Mistelgau, to which the specific name
refers, by teacher H. Herppich, who donated it to the private collection of Günther Eicken, a local amateur
paleontologist at Bayreuth, where it still resides. As a result the exemplar has no official inventory number.
The fossil comprises a shoulder-blade with wing, a partial leg, a rib and a caudal vertebra. Wild justified the
creation of a new species name by referring to the great size, with an about 50% larger wingspan than with a
typical specimen; the short lower leg and the long wing.
Padian in 2008 pointed out that D. banthensis specimen MBR 1977.21, the largest then known, has
with a wingspan of 169 centimetres an even larger size; that wing and lower leg proportions are rather
variable in D. banthensis and that the geological age is comparable. He concluded that D. mistelgauensis is
a subjective junior synonym of D. banthensis.
[edit] Phylogeny
The affinity between Dorygnathus and Dimorphodon, assumed by early researchers, was largely
based on a superficial resemblance in tooth form. Baron Franz Nopcsa in 1928 assigned the species to the
Rhamphorhynchinae,[17] which was confirmed by Peter Wellnhofer in 1978.[18] Modern exact cladistic
analyses of the relationships of Dorygnathus have not resulted in a consensus. David Unwin in 2003 found
that it belonged to the clade Rhamphorhynchinae,[19] but analyses by Alexander Kellner resulted in a much
more basal position,[20] below Dimorphodon or Peteinosaurus. Padian, using a comparative method, in 2008
concluded that Dorygnathus was close to Scaphognathus and Rhamphorhynchus in the phylogenetic tree
but also that these species were forming a series of successive off-shoots, meaning that they would not be
united in a separate clade. This was again contradicted by the results of a cladistic study by Brian Andres in
2010 showing that Dorygnathus was part of a monophyletic Rhamphorhynchinae.[21] The following
cladogram shows the position of Dorygnathus according to Andres:
Rhamphorhynchinae
Dorygnathus
unnamed
unnamed
Rhamphorhynchus
Cacibupteryx
unnamed
Harpactognathus
unnamed
Angustinaripterus
Sericipterus
[edit] Paleobiology
Dorygnathus is commonly thought to have had a piscivorous way of living, catching fish or other
slippery sea-creatures with its long teeth. This is confirmed by the fact that the fossils have been found in
marine sediments, deposited in the seas of the European Archipelago. In these it is present together with the
pterosaur Campylognathoides that however is much more rare. Very young juveniles of Dorygnathus are
unknown, the smallest discovered specimen having a wingspan of sixty centimetres; perhaps they were
unable to venture far over open sea. Padian concluded that Dorygnathus after a relatively fast growth in its
early years, faster than any modern reptile of the same size, kept slowly growing after having reached sexual
maturity, which would have resulted in exceptionally large individuals with a 1.7 metres wingspan.
On land, Dorygnathus was probably not a good climber; its claws show no special adaptations for
this type of locomotion. According to Padian, Dorygnathus, as a small pterosaur with a long tail, was well
capable of bipedal movement, though its long metacarpals would make him better suited for a quadrupedal
walk than most basal pterosaurs. Most researchers however, today assume quadrupedality for all pterosaurs.
[edit] See also

• Campylognathoides
• Holzmaden
• Rhamphorhynchus
[edit] References
1. ^ a b c d "Dorygnathus." In: Cranfield, Ingrid (ed.). The Illustrated Directory of Dinosaurs and
Other Prehistoric Creatures. London: Salamander Books, Ltd. Pp. 292-295.
2. ^ Theodori, C. (1830). "Knochen vom Pterodactylus aus der Liasformation von Banz",
Frorieps Notizen für Natur- und Heilkunde, n. 632, 101pp
3. ^ Theodori, C. (1831). "Ueber die Knochen vom Genus Pterodactylus aus der Liasformation
der Gegend von Banz", Okens Isis, 3: 276–281
4. ^ Meyer, H. von (1831). "Über Macrospondylus und Pterodactylus", Nova Acta Academia
Caesarae Leopold-Carolina Germania Naturali Curiae, 15: 198–200
5. ^ Theodori, C. (1852). "Ueber die Pterodactylus-Knochen im Lias von Banz", Berichte des
Naturforschenden Vereins Bamberg, 1: 17–44
6. ^ Oppel, A. (1856). "Die Juraformation", Jahreshefte des Vereins für Vaterländische
Naturkunde in Württemberg, 12
7. ^ Oppel, A. (1858). "Die Geognostische Verbreitung der Pterodactylen", Jahreshefte der
Vereins der vaterländische Naturkunde in Württemberg , 1858, Vorträge 8, 55 pp
8. ^ Wagner, A. (1860). "Bemerkungen über die Arten von Fischen und Sauriern, Welche im
untern wie im oberen Lias zugleich vorkommen sollen", Sitzungsberichte der königlichen
Bayerischen Akademie der Wissenschaften, mat.- physikalische Classe, p. 36-52
9. ^ Dominique Delsate & Rupert Wild. (2000). "Première Découverte d'un Reptile volant
determinable (Pterosauria, Dorygnathus cf banthensis) du Toracien inférieur (Jurassique inférieur)
de Nancy (Lorraine, France)", Bulletin de l'Académie et de la Société lorraines des sciences, 2000,
39: 1-4
10.^ Keller, Thomas (1985). "Quarrying and Fossil Collecting in the Posidonienschiefer (Upper
Liassic) around Holzmaden, Germany", Geological Curator, 4(4): 193-198
11.^ Plieninger, F. (1907). "Die Pterosaurier der Juraformation Schwabens", Palaeontographica,
53: 209–313
12.^ Arthaber, G.E. von (1919). "Studien über Flugsaurier auf Grund der Bearbeitung des
Wiener Exemplares von Dorygnathus banthensis Theod. sp.", Denkschriften der Akademie der
Wissenschaften Wien, Mathematisch-naturwissenschaftliche Klasse, 97: 391–464
13.^ Padian, K. & Wild, R. (1992). "Studies of Liassic Pterosauria, I. The holotype and referred
specimens of the Liassic Pterosaur Dorygnathus banthensis (Theodori) in the Petrefaktensammlung
Banz, Northern Bavaria", Palaeontographica Abteilung A, 225: 55-79
14.^ Kevin Padian (2008). The Early Jurassic Pterosaur Dorygnathus Banthensis(Theodori,
1830). Special Papers in Palaeontology No. 80, The Palaeontological Association, London
15.^ Broili, F. (1939) "Ein Dorygnathus mit Hautresten", Sitzungs-Berichte der Bayerischen
Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Abteilung , 1939: 129–132
16.^ Wild, R. (1971). "Dorygnathus mistelgauensis n. sp., ein neuer Flugsaurier aus dem Lias
Epsilon von Mistelgau (Frankischer Jura)" — Geol. Blatter NO-Bayern, 21(4): 178-195
17.^ Nopcsa, F. v. (1928). "The genera of reptiles". Palaeobiologica, 1: 163-188
18.^ Wellnhofer, P. (1978). Pterosauria. Handbuch der Palaeoherpetologie, Teil 19. Gustav
Fischer Verlag, Stuttgart
19.^ Unwin, D. M. (2003). "On the phylogeny and evolutionary history of pterosaurs". Pp. 139-
190 in: Buffetaut, E. and Mazin, J.-M., eds. Evolution and Palaeobiology of Pterosaurs. Geological
Society Special Publications 217. Geological Society of London
20.^ Kellner, A. W. A. (2003). "Pterosaur phylogeny and comments on the evolutionary history of
the group". Pp. 105-137 in: Buffetaut, E. and Mazin, J.-M., eds. Evolution and Palaeobiology of
Pterosaurs. Geological Society Special Publications 217. Geological Society of London
21.^ Brian Andres; James M. Clark & Xu Xing. (2010). "A new rhamphorhynchid pterosaur from
the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs",
Journal of Vertebrate Paleontology, 30(1): 163-187
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Categories: Rhamphorhynchoids | Jurassic pterosaurs | Prehistoric reptiles of Europe | Pterosaur

stubs
W000
Campylognathoides

Campylognathoides
Fossil range: Early Jurassic
Cast of the Pittsburgh specimen of Campylognathoides liasicus
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Pterosauria
Suborder: Rhamphorhynchoidea
Family: Campylognathoididae
Campylognathoidinae
Subfamily:
Kuhn, 1967
Campylognathoides
Genus:
Strand, 1928
Species
• C. zitteli (Plieninger, 1894 [originally Campylognathus

zitteli]) (type)
• C. indicus Jain, 1978
• C. liasicus (Quenstedt, 1858 [originally Pterodactylus
liasicus])
Campylognathoides ("curved jaw", Strand 1928) was a genus of "rhamphorhynchoid" pterosaur,
discovered in the Württemberg Lias deposits, the first specimen consisting of wing fragments. Further better
preserved specimens were found in the Holzmaden shale and it was based on these specimens that Felix
Plieninger erected a new genus.[1]
Contents
[hide]
• 1 Discovery
• 2 Anatomy and gait
• 3 Species
• 4 Phylogeny
• 5 Paleobiology
• 6 See also
• 7 Notes
• 8 References
[edit] Discovery
In 1858 Friedrich August Quenstedt named a new species of Pterodactylus: P. liasicus. It was based
on a fossil, holotype GPIT 9533, consisting of some wing bones, found on the Wittberg near Metzingen in
layers dating from the early Toarcian, about 180 million years old. The specific name referred to the Lias.
Quenstedt thought he had identified long metacarpals in the wing, concluding that the new species was
therefore not belonging to more basal genera, like the long-tailed Rhamphorhynchus.[2]
In 1893 commercial fossil collector Bernhard Hauff sr. discovered a skeleton of a large pterosaur
near Holzmaden. In 1894 Felix Plieninger based a new genus on this specimen: Campylognathus. The
genus name is derived from Greek kampylos, "bent", and gnathos, "jaw", in reference to the bent lower jaw.
The type species is Campylognathus zitteli. The specific name honours Alfred von Zittel.[3] The holotype is
SMNS 9787.
In 1897 Hauff prepared another specimen that eventually in 1903 was acquired by the Carnegie
Museum of Natural History at Pittsburgh. This fossil, CM 11424, is due to its completeness the best source of
information about the genus.
In 1901 Plieninger for the first time studied P. liasicus and discovered that Quenstedt had mistaken
the, in reality short, metacarpal, for a coracoid, meaning it was a basal pterosaur.[4] In 1906 Plieninger
referred P. liasicus and the Pittsburgh specimen to Campylognathus, though not yet establishing the specific
status of each of the three exemplars.[5] In 1907 however, Plieninger recognised a second species of
Campylognathus: C. liasicus, to which CM 11424 was referred also.[6]
Norwegian entomologist Embrik Strand discovered in the 1920s that the name Campylognathus
been previously been used for the African bug Campylognathus nigrensis, a genus of the Heteroptera named
in 1890. As the name was thus preoccupied, he renamed the pterosaur Campylognathoides in 1928.[7]
During the twentieth century new finds have brought the number of known specimens to about a
dozen.
[edit] Anatomy and gait
Restoration of Campylognathoides with a hypothetical tail form

Compared to its contemporary from the same layers Dorygnathus, the snout on this genus is
relatively short, though the skull is still in general elongated, be it much lighter built. The large eye sockets,
placed low in the skull above a narrow jugal, have caused some researchers to speculate that
Campylognathoides had especially acute vision, or possibly even a nocturnal lifestyle. The back of the skull
is relatively high and flat, with a sudden downturn just in front of the eyes. The snout ends in a slender point
curving a bit upwards at its very end. A large part of the snout is occupied by long bony nares. Below them a
small triangular skull opening, the fenestra antorbitalis is present.
Reflecting the more shallow snout, the teeth of Campylognathoides are also short and not at all
laniaries or fang-like as in the markedly heterodont Dorygnathus. They are conical and recurved but have a
broad base with the point bevelled off from the inside forming a sharp and strong cutting surface. In the upper
jaw there are four rather widely spaced teeth in the praemaxilla gradually increasing in size from the front to
the back; the fourth pair of teeth is the largest. Behind them are ten smaller teeth in the maxilla, gradually
decreasing posteriorely. In the lower jaw there are twelve to fourteen teeth present in C. liasicus, sixteen to
nineteen in C. zitteli. The largest total number is thus 66.
According to a study by Kevin Padian there are eight cervical vertebrae, fourteen dorsals, four or five
sacrals and up to 38 caudal vertebrae. The tail base is flexible with about six short vertebrae; behind them
the caudals elongate and are stiffened by very long extensions allowing the tail to function as a rudder.[8]
The sternum of Campylognathoides was a rather large rectangular plate of bone with a short forward
facing crest called a cristospina.[1] The upper arm is short but robust with a square deltopectoral crest. The
lower arm too is short but wing length is considerable due to the hand, which has short metacarpals but a
very long wing finger for a basal pterosaur, of which the second phalanx is the largest. The pteroid is short
and robust.
The pelvis is not very well known. A fossil collector found a well preserved Campylognathoides hip in
a Braunschweig shale quarry in 1986. This pelvis, BSP 1985 I 87, proved to be scientifically significant
because the hip socket was according to Peter Wellnhofer in an upward lateral position, preventing the
animal from being able to orient its legs erectly like in dinosaurs, birds and mammals. This would prove that
Campylognathoides was not well able to walk on its hind legs but must have walked quadrupedally.[9] This
gait posture has been confirmed in other "rhamphorhynchoids" (i.e. basal pterosaurs) as well.[1] However,
Padian in 2009 concluded the opposite, stating that an erected position was necessary to place the feet on
the ground and that, though a quadrupedal gait was possible, a bipedal way of locomotion was a
precondition for a fast gait.[10] This subject remains highly controversial.
The leg is rather short and the feet are small. The fifth toe, often interpreted as carrying a membrane
between the legs, is exceptionally short for a basal pterosaur.
[edit] Species
Three species of Campylognathoides have been named:
• Campylognathoides zitteli (Plieninger, 1894) is the type species.
The larger Holzmaden Campylognathoides, C. zitteli, had a six foot (1.825 metres for SMNS
9787) wingspan.[1]
• Campylognathoides liasicus (Quenstedt, 1858 [originally Pterodactylus liasicus])
C. liasicus had a three foot wingspan, making it smaller than its Holzmaden contemporary, C.
zitteli.[1]
• "Campylognathoides" indicus (Jain, 1974)
C. indicus was described by Sohan Lal Jain on the basis of a fragment of jaw, ISI R. 48,
recovered from Chanda district, India.[11] Kevin Padian considers this a nomen dubium, possibly
based on a fish fossil. That the Kota Formation in which it was found, has since been redated to the
Middle Jurassic or later, seems to preclude any close connection to Campylognathoides, even if it
were a pterosaur.[12]
The distinction between C. liasicus and C. zitteli is problematical. Plieninger merely recognised the
smaller species because he considered its fossil too poor in quality to refer other specimens to. However, in
1925 Swedish researcher Carl Wiman, studying specimen UUPM R157, concluded that a fundamental
morphological difference could distinguish the two species: C. zitteli has a proportionally much longer wing.
In 2008 however, Padian pointed out that this might well have been a matter of ontogenetic development,
larger individuals growing extra large wings to limit the wing load. Other differences, such as the larger
number of teeth in the lower jaw, a longer snout and nares, five instead of four sacrals, perpendicular sacral
ribs and a longer leg, might conceivably also be size-related. Final proof could only be given by a continuous
growth series, as previously has been done in the case of Rhamphorhynchus and Pterodactylus.
Provisionally Padian kept distinguishing two species, but moved two specimens to C. zitteli: SMNS 51100
and GPIT 24470, because of their larger size and morphological similarities.[13]
[edit] Phylogeny
Plieninger in his later publications assigned Campylognathus to the "Rhamphorhynchoidea". As this
suborder is a paraphyletic assemblage of not specially related basal pterosaurs, this classification merely
states the negative fact that it was not a pterodactyloid. A positive determination was first attempted by Baron
Franz Nopcsa who in 1928 assigned the genus to the subfamily Rhamphorhynchinae within the family
Rhamphorhynchidae.[14] After a period in which very little work was done on pterosaur systematics, in 1967
Oskar Kuhn placed Campylognathoides in its own subfamily within the Rhamphorhynchidae, the
Campylognathoidinae.[15] However, in 1974 Peter Wellnhofer concluded that it was placed in a more basal
position in the phylogenetic tree, below the Rhamphorhynchidae.[16] In the early twenty-first century this was
confirmed by the first extensive exact cladistic analyses. In 2003 both David Unwin and Alexander Kellner
introduced a clade Campylognathoididae; within Unwin's terminology this clade is the sister clade of the
Breviquartossa within the Lonchognatha;[17] applying Kellner's terminology it is the most basal off-shoot
within the Novialoidea.[18] There is no material difference between the two positions.
According to the analyses Campylognathoides would be closely related to Eudimorphodon, to which
it is similar in skull, sternum and humerus form. This was confirmed by Padian in 2009, though Padian also
pointed out several basal features present in Eudimorphodon but lacking in Campylognathoides.[19] In 2010
an analysis was published by Brian Andres showing that Eudimorphodon together with Austriadactylus
formed a very basal clade, leaving Campylognathoides as the only known member of the
Campylognathoididae.[20]
[edit] Paleobiology
Traditionally a piscivore lifestyle is attributed to Campylognathoides, as to most pterosaurs; in this
case supported by the provenance of the finds from marine sediments and the very long wings. Padian
however, has suggested that, in view of the stout short teeth, ideal for delivering a piercing bite, the form
might well have been a predator of small terrestrial animals instead.[21] The niche of specialised fish eater
would then have been filled by Dorygnathus which is five times as common in the layers. The area of the
fossil sites was in the early Jurassic located to the northwest of a large island , the Massif of Bohemia,
situated in a shallow gulf of the Tethys Sea.
[edit] See also

• Dorygnathus
• Holzmaden
• Rhamphorhynchus
[edit] Notes
1. ^ a b c d e "Campylognathoides." In: Cranfield, Ingrid (ed.). The Illustrated Directory of
Dinosaurs and Other Prehistoric Creatures. London: Salamander Books, Ltd. Pp. 292-295.
2. ^ Quenstedt, F. A. (1858). "Ueber Pterodactylus liasicus", Jahrbuch des Vereins
vaterländischer Naturkundler in Württemberg, 14:299-336
3. ^ Plieninger, F. (1894). "Campylognathus Zittelli. Ein neuer Flugsaurier aus dem Oberen Lias
Schwabens", Palaeontographica, 41: 193-222
4. ^ Plieninger, F. (1901). "Beiträge zur Kenntnis der Flugsaurier", Palaeontographica, 48: 65–
90
5. ^ Plieninger, F. (1906). "Notizen über Flugsaurier aus dem Lias Schwabens", Centralblatt für
Mineralogie, 10: 290–293
6. ^ Plieninger, F. (1907) "Die Pterosaurier der Juraformation Schwabens", Palaeontographica,
53: 209-313
7. ^ Strand, E. (1928). "Miscellanea nomenclatorica Zoologica et Palaeontologica", Archiv fur
Naturgeschichte, 92: 30-75
8. ^ Padian (2009), pp. 100-101
9. ^ Wellnhofer, P. & Vahldiek, B.-W. (1986). "Ein Flugsaurier-Rest aus dem Posidonienschiefer
(Unter-Toarcium) von Schandelah bei Braunschweig", Paläontologische Zeitschrift , 60: 329-340
10.^ Padian (2009), p. 103
11.^ Jain, S. L. (1974) "Jurassic Pterosaur from India", Journal of the Geological Society of
India, 15 (3): 330-335
12.^ Padian (2009), p. 98
13.^ Padian (2009), p. 71, 105
14.^ Nopcsa, F. v. (1928). "The genera of reptiles". Palaeobiologica, 1: 163-188
15.^ Kuhn, O. (1967). Die fossile Wirbeltierklasse Pterosauria. Krailling: Oeben-Verlag. 52 pp
16.^ Wellnhofer, P. (1974). "Campylognathoides liasicus (Quenstedt), an Upper Liassic
pterosaur from Holzmaden — The Pittsburgh specimen", Annals of Carnegie Museum, 45: 5-34
17.^ Unwin, D. M. (2003). "On the phylogeny and evolutionary history of pterosaurs". Pp. 139-
190 in: Buffetaut, E. and Mazin, J.-M., eds. Evolution and Palaeobiology of Pterosaurs. Geological
Society Special Publications 217. Geological Society of London
18.^ Kellner, A. W. A. (2003). "Pterosaur phylogeny and comments on the evolutionary history of
the group". Pp. 105-137 in: Buffetaut, E. and Mazin, J.-M., eds. Evolution and Palaeobiology of
Pterosaurs. Geological Society Special Publications 217. Geological Society of London
19.^ Padian (2009), pp. 103-104
20.^ Brian Andres; James M. Clark & Xu Xing. (2010). "A new rhamphorhynchid pterosaur from
the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs",
Journal of Vertebrate Paleontology, 30(1): 163-187
21.^ Padian (2009), p. 105
[edit] References
• Dinosaurs and other Prehistoric Creatures, edited by Ingrid Cranfield, 2000 Salamander
Books Ltd pg 285-286.
• Quenstedt, F. A. 1858 "Ueber Pterodactylus liasicus", Jahreshefte des Vereins für
vaterländische Naturkunde in Württemberg 14, 299–310 & pl. 2.
• Plieninger, F. 1907 "Die Pterosaurier der Juraformation Schwabens", Paläontographica 53,
209–313 & pls 14–19.
• Wellnhofer, P. 1974 "Campylognathoides liasicus (Quenstedt), an Upper Liassic pterosaur
from Holzmaden - The Pittsburgh specimen", Ann. Carnegpterus. 45 (2), 5–34.
• Plieninger, F. 1894 "Campylognathus Zitteli, ein neuer Flugsaurier aus dem obersten Lias
Schwabens", Paläontographica 41, 193–222 & pl. 19.
• Jain, 1974, "Jurassic Pterosaur from India", Journal of the Geological Society of India, vol.15,
Pt.3 pp. 330–335
• Padian, K. 2009. The Early Jurassic Pterosaur Dorygnathus banthenis (Theodori, 1830) and
The Early Jurassic Pterosaur Campylognathoides Strand, 1928, Special Papers in Paleontology 80,
Blackwell ISBN 9781405192248

• Anatomy
Retrieved from "http://en.wikipedia.org/wiki/Campylognathoides"
Categories: Jurassic pterosaurs | Rhamphorhynchoids | Prehistoric reptiles of Asia | Prehistoric

reptiles of Europe
W000
Braconidae

Braconidae
Braconidae
A member of the braconid genus Atanycolus.
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Superfamily: Ichneumonoidea
Family: Braconidae
Subfamilies
• Adeliinae
• Agathidinae
• Alysiinae
• Amicrocentrinae
• Aphidiinae
• Apozyginae
• Betylobraconinae
• Blacinae
• Braconinae
• Cardiochilinae
• Cenocoeliinae
• Cheloninae
• Dirrhopinae
• Doryctinae
• Ecnomiinae
• Euphorinae
• Exothecinae
• Gnamptodontinae
• Helconinae
• Histeromerinae
• Homolobinae
• Hormiinae
• Khoikhoiiinae
• Ichneutinae
• Macrocentrinae
• Masoninae
• Mendesellinae
• Mesostoinae
• Meteorideinae
• Meteorinae
• Microgastrinae
• Microtypinae
• Miracinae
• Neoneurinae
• Opiinae
• Orgilinae
• Pselaphaninae
• Rhyssalinae
• Rogadinae
• Sigalphinae
• Telengaiinae
• Trachypetinae
• Vaepellinae
• Ypsistocerinae
• Xiphozelinae
Tomato hornworm caterpillar Manduca quinquemaculata parasitized by Braconidae wasp larvae.

Braconidae is a family of parasitoid wasps and one of the richest families of insects. Between 50,000
and 150,000 species exist worldwide. The species are grouped into about 45 subfamilies and 1,000 genera,
some important ones being: Ademon, Aphanta, Asobara, Bracon hebetor, Cenocoelius, Chaenusa,
Chorebidea, Chorebidella, Chorebus, Cotesia, Dacnusa, Microgaster, Opius, Parapanteles, Phaenocarpa,
Psenobolus.
Contents
[hide]
• 1 Morphology
• 2 Parasitism
• 3 Larval development
• 4 Natural history
• 6 Other characteristics
• 7 References
[edit] Morphology
The morphological variation among braconids is notable. Braconids are often black-brown
(sometimes with reddish markings), though some species exhibit striking coloration and pattern, being parts
of Müllerian mimicry complexes. They have one or no recurrent veins, unlike other members of the
Ichneumonoidea which usually have two. Wing venation patterns are also divergent to apparent randomness.
The antennae have 16 segments or more; the hind trochanters have 2 segments.
Females often have long ovipositors, an organ that largely varies intraspecifically. This variation is
closely related to the host species upon which the wasp deposits its egg. Species that parasitize
microlepidoptera, for instance, have longer ovipositors, presumably to reach the caterpillar through layers of
plant tissue. Some wasps also have long ovipositors because of caterpillar defense mechanisms such as
spines or hairs.
[edit] Parasitism
Braconid parasitoid wasp Apanteles species cocoons on Papilio demoleus caterpillar

Most braconids are primary parasitoids (both external and internal) on other insects, especially upon
the larval stages of Coleoptera, Diptera, and Lepidoptera, but also some hemimetabolous insects like aphids,
Heteroptera or Embiidina. Most species kill their hosts, though some cause the hosts to become sterile and
less active. Endoparasitoid species often display elaborate physiological adaptations to enhance larval
survival within host, such as the co-option of endosymbiotic viruses for compromising host immune defenses.
These polydnaviruses are often used by the wasps instead of a venom cocktail. The DNA of the wasp
actually contains portions that are the templates for the components of the viral particles and they are
assembled in an organ in the female's abdomen known as the calyx.[1] A 2009 study has traced the origins
of these templates to a 100-million-year-old viral infection whose alterations to its host DNA provided the
necessary basis for these virus-like "templates".[2]
These viruses suppress the immune system and allow the parasitoid to grow inside the host
undetected. The exact function and evolutionary history of these viruses are unknown. It is a little surprising
to consider that sequences of polydnavirus genes show the possibility that venom-like proteins are
expressed inside the host caterpillar. It appears that through evolutionary history the wasps have so highly
modified these viruses that they appear unlike any other known viruses today. Because of this highly
modified system of host immunosuppression it is not surprising that there is a high level of parasitoid-host
specificity. It is this specificity that makes Braconids a very powerful and important biological control agent.
Parasitism on adult insects (particularly on Hemiptera and Coleoptera) is also observed. Members of
two subfamilies (Mesostoinae and Doryctinae) are known to form galls on plants.
Exploitation of ant-aphid mutualism by unidentified parasitic wasp: wasp laying eggs in aphid
undisturbed by investigating ant.
[edit] Larval development

Surprisingly, both syncitial and holoblastic cleavage are present, even in closely related taxa.
Larvae can be found on hosts as diverse as aphids, bark beetles, and foliage-feeding caterpillars.
Many species are egg-larval parasitoids; hence they are often utilized as biological pest control agents,
especially against aphids.
[edit] Natural history

The family seems to date from early Cretaceous (provided that Eobracon is properly assigned to this
family). It underwent extensive diversification from mid or late Cretaceous to early Tertiary, correlating with
the radiation of flowering plants and associated herbivores, the main hosts of braconids.
Braconidae is traditionally divided into more than 40 subfamilies. These fall to two major groups,
informally called the cyclostomes and non-cyclostomes. In cyclostome braconids, the labrum and the lower
part of the clypeus are concave with respect to the upper clypeus and the dorsal margin of the mandibles.
These groups may be clades that diverged early in the evolution of braconids.[3]
[edit] Other characteristics
The species Microplitis croceipes possesses an extremely accurate sense of smell and can be
trained for use in narcotics and explosives detection[4].
[edit] References
1. ^ Piper, Ross (2007), Extraordinary Animals: An Encyclopedia of Curious and Unusual
Animals, Greenwood Press.
2. ^ Drezen, Jean-Michel; Lanzrein, Beatrice; Volkoff, Anne-Nathalie; Huguet, Elisabeth;
Dupuy, Catherine; Periquet, Georges; Pfister-Wilhem, Rita; Belghazi, Maya; Heller, Manfred; Roditi,
Isabel; Wincker, Patrick; Bernard-Samain, Sylvie; Gyapay, Gabor; Wetterwald, Christoph; Herbinière,
Juline; Annaheim, Marc; Bézier, Annie (February 13, 2009). "Polydnaviruses of Braconid Wasps
Derive from an Ancestral Nudivirus". Science (Science Magazine) 323 (5916): 926–930.
doi:10.1126/science.1166788. PMID 19213916.
3. ^ Wharton, Robert M. (2000), "Can braconid classification be restructured to facilitate
portrayal of relationships?", in Austin, Andrew D.; Dowton, Mark, Hymenoptera: evolution,
biodiversity, and biological control, 4th International Hymenopterists Conference, Collingwood,
Victoria, Australia: Commonwealth Scientific and Industrial Research Organisation (CSIRO), pp. 143–
153, ISBN 0 643 06610 1
4. ^ The Scoop: Move Over, Rover. November 20, 2001

• Tree of Life Braconidae
• Agathidinae Synopsis Sharkey
• NNM Technical Bulletin Bibliography of Braconidae 1964-2003
Wikimedia Commons has media related to: Braconidae
Wikispecies has information related to: Braconidae
• Cotesia marginiventris on the UF / IFAS Featured Creatures Web site

• Diachasmimorpha longicaudata on the UF / IFAS Featured Creatures Web site
• Meteorus autographae on the UF / IFAS Featured Creatures Web site
• Checklist of British and Irish Braconidae pdf
• Achterberg C. van, C. O'Toole (1993) Annotated catalogue of the types of Braconidae
(Hymenoptera) in the Oxford University Museum Zoologische Verhandelingen, Vol. 287 P. 1-43 PDF
Retrieved from "http://en.wikipedia.org/wiki/Braconidae"
Categories: Braconidae | Biological pest control agents

W000
Listeria monocytogenes

Scanning electron micrograph of Listeria monocytogenes.
Kingdom: Bacteria
Division: Firmicutes
Class: Bacilli
Order: Bacillales
Family: Listeriaceae
Genus: Listeria
Species: L. monocytogenes
Binomial name
(Murray et al. 1926) Pirie 1940
Listeria monocytogenes is a facultative anaerobe, intracellular bacterium that is the causative agent
of Listeriosis. It is one of the most virulent foodborne pathogens with 20 to 30 percent of clinical infections
resulting in death.[1] Responsible for approximately 2,500 illnesses and 500 deaths in the United States
(U.S.) annually, Listeriosis is the leading cause of death among foodborne bacterial pathogens with fatality
rates exceeding even Salmonella and Clostridium botulinum.[2]
L. monocytogenes is a Gram-positive bacterium, in the division Firmicutes, named for Joseph Lister.
Motile via flagella at 30 °C and below but usually not at 37 °C,[3] L. monocytogenes can instead move within
eukaryotic cells by explosive polymerization of actin filaments (known as comet tails or actin rockets).
Studies suggest that up to 10% of human gastrointestinal tracts may be colonized by L.
monocytogenes.[1]
Nevertheless, clinical diseases due to L. monocytogenes are more frequently recognized by
veterinarians, especially as meningo-encephalitis in ruminants. See: listeriosis in animals.
Due to its frequent pathogenicity causing meningitis in newborns (acquired transvaginally), pregnant
mothers are often advised not to eat soft cheeses such as Brie, Camembert, feta and queso blanco fresco,
which may be contaminated with and permit growth of L. monocytogenes.[4] It is the third most common
cause of meningitis in newborns.
More recently, L. monocytogenes has been used as the model organism to illustrate the Patho-
biotechnology concept.
Contents
[hide]
• 2 History
• 3 Pathogenesis
• 3.1 Regulation of pathogenesis
• 3.2 Pathogenicity of Lineages
• 4 Treatment
• 5 Use as a transfection vector
• 5.1 Cancer vaccine
• 6 Detection
• 7 Epidemiology
• 8 Routes of infection
• 9 Infectious cycle
• 10 References
L. monocytogenes is a gram-positive, non-spore forming, motile, facultatively anaerobic, rod-shaped
bacterium. It is catalase-positive, oxidase-negative, and expresses a Beta hemolysin which causes
destruction of red blood cells. This bacterium exhibits characteristic tumbling motility when viewed with light
microscopy.[5] Although L. monocytogenes is actively motile by means of peritrichous flagella at room
temperature (20−25 °C), the organism does not synthesize flagella at body temperatures (37 °C).[6]
The genus Listeria belongs to the Clostridium sub-branch, together with Staphylococcus,
Streptococcus, Lactobacillus and Brochothrix. The genus Listeria includes 6 different species (L.
monocytogenes, L. ivanovii, L. innocua, L. welshimeri, L. seeligeri, and L. grayi). Both L. ivanovii and L.
monocytogenes are pathogenic in mice, but only L. monocytogenes is consistently associated with human
illness.[7] There are 13 serotypes of L. monocytogenes which can cause disease, but more than 90 percent
of human isolates belong to only three serotypes: 1/2a, 1/2b, and 4b. L. monocytogenes serotype 4b strains
are responsible for 33 to 50 percent of sporadic human cases worldwide and for all major foodborne
outbreaks in Europe and North America since the 1980s.[8][9]
[edit] History
L. monocytogenes was first described by E.G.D.Murray in 1926 based on six cases of sudden death
in young rabbits.[10] Murray referred to the organism as Bacterium monocytogenes before J.H. Harvey Pirie
changed the genus name to Listeria in 1940.[11] Although clinical descriptions of L. monocytogenes infection
in both animals and humans were published in the 1920s, not until 1952 in East Germany was it recognized
as a significant cause of neonatal sepsis and meningitis.[12] Listeriosis in adults would later be associated
with patients living with compromised immune systems, such as individuals taking immunosuppressant drugs
and corticosteroids for malignancies or organ transplants, and those with HIV infection.[13]
It wasn't until 1981, however, that L. monocytogenes was identified as a cause of foodborne illness.
An outbreak of listeriosis in Halifax, Nova Scotia involving 41 cases and 18 deaths, mostly in pregnant
women and neonates, was epidemiologically linked to the consumption of coleslaw containing cabbage that
had been treated with L. monocytogenes contaminated raw sheep manure.[14] Since then a number of cases
of foodborne listeriosis have been reported, and L. monocytogenes is now widely recognized as an important
hazard in the food industry.[15]
[edit] Pathogenesis
Main article: Listeriosis
Infection by L. monocytogenes causes the disease listeriosis. The manifestations of listeriosis
include septicemia,[16] meningitis (or meningoencephalitis),[16] encephalitis,[17] corneal ulcer,[18]
pneumonia,[19] and intrauterine or cervical infections in pregnant women, which may result in spontaneous
abortion (2nd/3rd trimester) or stillbirth. Surviving neonates of Fetomaternal Listeriosis may suffer
granulomatosis infantiseptica - pyogenic granulomas distributed over the whole body, and may suffer from
physical retardation. Influenza-like symptoms, including persistent fever, usually precede the onset of the
aforementioned disorders. Gastrointestinal symptoms such as nausea, vomiting, and diarrhea may precede
more serious forms of listeriosis or may be the only symptoms expressed. Gastrointestinal symptoms were
epidemiologically associated with use of antacids or cimetidine. The onset time to serious forms of listeriosis
is unknown but may range from a few days to three weeks. The onset time to gastrointestinal symptoms is
unknown but probably exceeds 12 hours. An early study suggested that L. monocytogenes was unique
among Gram-positive bacteria in that it possessed lipopolysaccharide,[20] which served as an endotoxin.
Later it was found to not be a true endotoxin, Listeria cell walls consistently contain lipoteichoic acids, in
which a glycolipid moiety, such as a galactosyl-glucosyl-diglyceride, is covalently linked to the terminal
phosphomonoester of the teichoic acid. This lipid region anchors the polymer chain to the cytoplasmic
membrane. These lipoteichoic acids resemble the lipopolysaccharides of gram-negative bacteria in both
structure and function, being the only amphipathic polymers at the cell surface.[21][22]
The infective dose of L. monocytogenes varies with the strain and with the susceptibility of the victim.
From cases contracted through raw or supposedly pasteurized milk, one may safely assume that in
susceptible persons, fewer than 1,000 total organisms may cause disease. L. monocytogenes may invade
the gastrointestinal epithelium. Once the bacterium enters the host's monocytes, macrophages, or
polymorphonuclear leukocytes, it becomes blood-borne (septicemic) and can grow. Its presence
intracellularly in phagocytic cells also permits access to the brain and probably transplacental migration to
the fetus in pregnant women. The pathogenesis of L. monocytogenes centers on its ability to survive and
multiply in phagocytic host cells.
[edit] Regulation of pathogenesis

L. monocytogenes can act as a saprophyte or a pathogen depending on its environment. When this
bacteria is present within a host organism quorum sensing causes the up regulation of several virulence
genes. Depending on the location of the bacteria within the host organism different activators up regulate the
virulence genes. SigB, an alternative sigma factor, up regulates Vir genes in the intestines whereas PrfA up
regulates gene expression when the bacteria is present in blood.[23][24][25][26] Little is known about the
mechanism in how this bacteria switches between acting as a saprophyte and a pathogen however, it is
thought that several non-coding RNAs are required to induce this change.
[edit] Pathogenicity of Lineages

L. monocytogenes has three distinct lineages with differing evolutionary histories and pathogenic
potentials.[27] Lineage I strains contain the majority of human clinical isolates and all human epidemic
clones, but are underrepresented in animal clinical isolates.[27] Lineage II strains are overrepresented in
animal cases and underrepresented in human clinical cases as well as more prevalent in environmental and
food samples.[28] Lineage III isolates are very rare but significantly more common in animal isolates than
human.[27]
[edit] Treatment
When listeric meningitis occurs, the overall mortality may reach 70%; from septicemia 50%, from
perinatal/neonatal infections greater than 80%. In infections during pregnancy, the mother usually survives.
Reports of successful treatment with parenteral penicillin or ampicillin exist. Trimethoprim-sulfamethoxazole
has been shown effective in patients allergic to penicillin.
Bacteriophage treatments have been developed by several companies. EBI Food Safety and
Intralytix both have products suitable for treatment of the bacteria. The Food and Drug Administration of the
United States approved a cocktail of six bacteriophages from Intralytix, and a one type phage product from
EBI Food Safety designed to kill the bacteria L. monocytogenes. Uses would potentially include spraying it on
fruits and ready-to-eat meat such as sliced ham and turkey.
[edit] Use as a transfection vector

Because L. monocytogenes is an intracellular parasite, some studies have used this bacteria as a
vector to deliver genes in vitro. Current transfection efficiency remains poor. One example of the successful
use of L. monocytogenes in in vitro transfer technologies is in the delivery of gene therapies for cystic fibrosis
cases.[29]
[edit] Cancer vaccine
A live attenuated L. monocytogenes cancer vaccine named ADXS11-001 is under development as a
possible treatment for cervical carcinoma.[30]
[edit] Detection
Colonies of typical Listeria monocytogenes as they appear when grown on Listeria selective agar
The methods for analysis of food are complex and time-consuming. The present U.S. Food and Drug
Administration (FDA) method, revised in September, 1990, requires 24 and 48 hours of enrichment, followed
by a variety of other tests. Total time to identification takes from 5 to 7 days, but the announcement of
specific nonradiolabled DNA probes should soon allow a simpler and faster confirmation of suspect isolates.
Recombinant DNA technology may even permit 2-to-3 day positive analysis in the future. Currently,
the FDA is collaborating in adapting its methodology to quantitate very low numbers of the organisms in
foods.
[edit] Epidemiology
Researchers have found L. monocytogenes in at least 37 mammalian species, both domesticated
and feral, as well as in at least 17 species of birds and possibly in some species of fish and shellfish.
Laboratories can isolate L. monocytogenes from soil, silage, and other environmental sources. L.
monocytogenes is quite hardy and resists the deleterious effects of freezing, drying, and heat remarkably
well for a bacterium that does not form spores. Most L. monocytogenes are pathogenic to some degree.
[edit] Routes of infection

L. monocytogenes has been associated with such foods as raw milk, pasteurized fluid milk,[31]
cheeses (particularly soft-ripened varieties), ice cream, raw vegetables, fermented raw-meat sausages, raw
and cooked poultry, raw meats (of all types), and raw and smoked fish. Its ability to grow at temperatures as
low as 0°C permits multiplication in refrigerated foods. In refrigeration temperature such as 4°C the amount of
ferric iron promotes the growth of L. monocytogenes.[32]
[edit] Infectious cycle

The primary site of infection is the intestinal epithelium where the bacteria invade non-phagocytic
cells via the "zipper" mechanism. Uptake is stimulated by the binding of listerial internalins (Inl) to host cell
adhesion factors such as E-cadherin or Met. This binding activates certain Rho-GTPases which
subsequently bind and stabilize Wiskott Aldrich Syndrome Protein (WASp). WASp can then bind the Arp2/3
complex and serve as an actin nucleation point. Subsequent actin polymerization extends the cell membrane
around the bacterium, eventually engulfing it. The net effect of internalin binding is to exploit the junction
forming-apparatus of the host into internalizing the bacterium. Note that L. monocytogenes can also invade
phagocytic cells (e.g. macrophages) but only requires internalins for invasion of non-phagocytic cells.
Following internalisation, the bacterium must escape from the vacuole/phagosome before fusion with
a lysosome can occur. Three main virulence factors which allow the bacterium to escape are listeriolysin O
(LLO - encoded by hly) phospholipase A (encoded by plcA) and phospholipase B (plcB).[33][34] Secretion of
LLO and PlcB disrupts the vacuolar membrane and allows the bacterium to escape into the cytoplasm where
it may proliferate.
Once in the cytoplasm, theL. monocytogenes exploits host actin for the second time. ActA proteins
associated with the old bacterial cell pole (being a bacillus, L. monocytogenes septates in the middle of the
cell and thus has one new pole and one old pole) are capable of binding the Arp2/3 complex and thus induce
actin nucleation at a specific area of the bacterial cell surface. Actin polymerization then propels the
bacterium unidirectionally into the host cell membrane. The protrusion which is formed may then be
internalised by a neighbouring cell, forming a double-membrane vacuole from which the bacterium must
escape using LLO and PlcB.
[edit] References
1. ^ a b Ramaswamy V, Cresence VM, Rejitha JS, Lekshmi MU, Dharsana KS, Prasad SP,
Vijila HM. (02 2007). "Listeria – review of epidemiology and pathogenesis." (PDF). J Microbiol
Immunol Infect. 40 (1): 4–13. PMID 17332901.
http://www.sochinf.cl/documentos/infectologia/listeria.pdf. Retrieved 2010-09-05.
2. ^ Dharmarha, Vaishali (December 2008). The majority of deaths from Listeria food poisoning
are in individuals with compromised immune systems: pregnant women, newborns, the elderly, and
the immunosupressed. "A Focus on Listeria Monocytogenes". National Agricultural Library, Food
Safety Research Information Office. Retrieved January 28, 2009.
3. ^ Gründling, A., Burrack, L.S., Bouwer, H.G.A., Higgins, D.E. 2004. Listeria monocytogenes
regulates flagellar motility gene expression through MogR, a transcriptional repressor required for
virulence. Proc. Natl. Acad. Sci. USA. 101:12316–12323.
4. ^ Genigeorgis, C.,Carniciu, M., Dutulescu, D., Farver, T.B. 1991. Growth and survival of
Listeria monocytogenes in market cheeses stored at 4 to 30 degrees C. J. Food Prot. 54(9):662-668.
5. ^ Farber, J. M.; Peterkin, P. I. (September 1991). "Listeria monocytogenes, a food-borne
pathogen". Microbiology and Molecular Biology Reviews 55 (3): 476–511. PMID 1943998.
PMC 372831. http://mmbr.asm.org/cgi/content/short/55/3/476.
6. ^ Todar, K. (2008). "Listeria monocytogenes". Todar's Online Textbook of Bacteriology.
7. ^ Seafood HACCP Alliance (2007). "Compendium of Fish and Fishery Product Processes,
Hazards, and Controls, Chapter 15: Listeria monocytogenes". Seafood Network Information Center.
8. ^ Dharmarha, Vaishali (December 2008). "A Focus on Listeria Monocytogenes". National
Agricultural Library, Food Safety Research Information Office. Retrieved January 28, 2009.
9. ^ Ward, T. J.; Gorski, L.; Borucki, M. K.; Mandrell, R. E.; Hutchins, J.; Pupedis, K. (2004).
"Intraspecific Phylogeny and Lineage Group Identification Based on the prfA Virulence Gene Cluster
of Listeria monocytogenes ". Journal of Bacteriology 186 (15): 4994–5002.
doi:10.1128/JB.186.15.4994-5002.2004. PMID 15262937.
10.^ Murray, E.G.D., Webb, R.E., Swann, M.B.R. 1926. A disease of rabbits characterized by a
large mononuclear leucocytosis, caused by a hitherto undescribed bacillus Bacterium
monocytogenes (n. sp.). J. Pathol. Bacteriol. 29: 407– 439.
11.^ Harvey, P.J.H. 1940. Listeria: change of name for a genus of bacteria. Nature. 145:264
12.^ Potel, J. 1952. Zur Granulomatosis infantiseptica. Zentr. Bakteriol. I. Orig. 158: 329-331
13.^ Schlech, W.F. III. 2001. Foodborne listeriosis. Clin. Infect. Dis. 31: 770-775.
14.^ Schlech, W.F., Lavigne, P.M., Bortolussi, R.A., Allen, A.C., Haldane, E.V., Wort, A.J.,
Hightower, A.W., Johnson, S.E., King, S.H., Nicholls, E.S. and Broome, C.V. 1983. Epidemic
listeriosis—evidence for transmission by food. New Engl. J. Med. 308:203–206.
15.^ Ryser, E.T., Marth, E.H. (Eds.) 1999. Listeria, Listeriosis, and Food. Safety, 2nd edn.
Marcel Dekker, New York.
16.^ a b Gray, M. L., and A. H. Killinger. 1966. Listeria monocytogenes and listeric infection.
Bacteriol. Rev. 30:309-382.
17.^ Armstrong, R. W., and P. C. Fung. 1993. Brainstem encephalitis (Rhombencephalitis) due
to Listeria monocytogenes: case report and review. Clin. Infect. Dis. 16:689-702.
18.^ Holland, S., E. Alfonso, . Gelender, D. Heidegger, A. Mendelsohn, S. Ullman, and D. Miller.
1987. Corneal ulcer due to Listeria monocytogenes. Cornea 6:144-146.
19.^ Whitelock-Jones, L., J. Carswell, and K. C. Rassmussen. 1989. Listeria pneumonia. A case
report. South African Medical Journal 75:188-189.
20.^ Wexler, H., and J. D. Oppenheim. 1979. Isolation, characterization, and biological
properties of an endotoxin-like material from the gram-positive organism Listeria monocytogenes.
Infect. Immun. 23:845-857.
21.^ Fiedler, F (1988). "Biochemistry of the cell surface of Listeria strains: a locating general
view.". Infection 16 Suppl 2: S92–7. PMID 3417357.
22.^ Farber, JM; Peterkin, PI (1991). "Listeria monocytogenes, a food-borne pathogen.".
Microbiological reviews 55 (3): 476–511. PMID 1943998.
23.^ Mengaud, J.; Dramsi, S.; Gouin, E.; Vazquez-Boland, J. A.; Milon, G.; Cossart, P. (1991).
"Pleiotropic control of Listeria monocytogenes virulence factors by a gene that is autoregulated".
Molecular Microbiology 5 (9): 2273–83. doi:10.1111/j.1365-2958.1991.tb02158.x. PMID 1662763.
24.^ Leimeister-Wachter, M. (1990). "Identification of a Gene that Positively Regulates
Expression of Listeriolysin, the Major Virulence Factor of Listeria monocytogenes". Proceedings of
the National Academy of Sciences 87: 8336–40. doi:10.1073/pnas.87.21.8336.
25.^ Garner, M. R.; Njaa, B. L.; Wiedmann, M.; Boor, K. J. (2006). "Sigma B Contributes to
Listeria monocytogenes Gastrointestinal Infection but Not to Systemic Spread in the Guinea Pig
Infection Model". Infection and Immunity 74 (2): 876–86. doi:10.1128/IAI.74.2.876-886.2006.
PMID 16428730.
26.^ Mandin, Pierre; Fsihi, Hafida; Dussurget, Olivier; Vergassola, Massimo; Milohanic, Eliane;
Toledo-Arana, Alejandro; Lasa, Iñigo; Johansson, JöRgen et al. (2005). "VirR, a response regulator
critical for Listeria monocytogenes virulence". Molecular Microbiology 57 (5): 1367–80.
doi:10.1111/j.1365-2958.2005.04776.x. PMID 16102006.
27.^ a b c Jeffers, G. T., J. L. Bruce, P. L. McDonough, J. Scarlett, K. J. Boor, and M. Wiedmann.
2001. Comparative genetic characterization of Listeria monocytogenes isolates from human and
animal listeriosis cases. Microbiology 147:1095-1104.
28.^ Gray, M. J., R. N. Zadoks, E. D. Fortes, B. Dogan, S. Cai, Y. Chen, V. N. Scott et al. 2004.
Listeria monocytogenes isolates from foods and humans form distinct but overlapping populations.
Applied and Environmental Microbiology 70:5833-5841.
29.^ Krusch, S; Domann, E; Frings, M; Zelmer, A; Diener, M; Chakraborty, T; Weiss, S (2002).
"Listeria monocytogenes mediated CFTR transgene transfer to mammalian cells.". The journal of
gene medicine 4 (6): 655–67. doi:10.1002/jgm.313. PMID 12439857.
30.^ Fran Lowry (2008-05-15). "Live Listeria Vaccine Proves Safe Against End-Stage Cervical
Ca in Human Trial". Ob.Gyn. News Vol.43, No.10, page 2.
31.^ Fleming, D. W., S. L. Cochi, K. L. MacDonald, J. Brondum, P. S. Hayes, B. D. Plikaytis, M.
B. Holmes, A. Audurier, C. V. Broome, and A. L. Reingold. 1985. Pasteurized milk as a vehicle of
infection in an outbreak of listeriosis. N. Engl. J. Med. 312:404-407.
32.^ Dykes, G. A., Dworaczek (Kubo), M. 2002. Influence of interactions between temperature,
ferric ammonium citrate and glycine betaine on the growth of Listeria monocytogenes in a defined
medium. Lett Appl Microbiol. 35(6):538-42.
33.^ Schmid, M. W., E. Y. W. Ng, R. Lampidis, M. Emmerth, M. Walcher, J. Kreft, W. Goebel et
al. 2005. Evolutionary history of the genus Listeria and its virulence genes. Systematic and Applied
Microbiology 28:1-18.
34.^ Zhang, C., M. Zhang, J. Ju, J. Nietfeldt, J. Wise, P. M. Terry, M. Olson et al. 2003. Genome
diversification in phylogenetic lineages I and II of Listeria monocytogenes: identification of segments
unique to lineage II populations. The Journal of Bacteriology 185:5573-5584.

Wikispecies has information related to: Listeria monocytogenes
Wikimedia Commons has media related to: Listeria monocytogenes

• U.S. Food and Drug Administration. Foodborne Pathogenic Microorganisms and Natural
Toxins Handbook: Listeria monocytogenes
• Public Health Agency of Canada
[hide]v · d · eFirmicutes (low-G+C) Infectious diseases · Bacterial diseases: G+ (primarily A00–A79, 001–041, 080–
Bacilli
suscep
pneum
(Pneu
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Lactobacillales oralis,
Streptococcus
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Strept
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S. aga
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Strept
BEA+: E
Enterococcus faecalis (Urinary
Enterococcus fae
Bacillales Cg+ S.
(Cat+) (Staphylococca
syndrome, Tox
syndrome, MRS
Staphylococcus
Cg- nov
susceptible (S.
novobiocin resi
saprophyticus)
Bacillus Bacillus
(Anthrax) · Bac
poisoning)
Listeria
Listeria
(Listeriosis)
motile: Clostridium diffic

(Pseudomembranous colitis) · C
Clostridium (spore- botulinum (Botulism) · Clostridiu
forming) (Tetanus)
Clostridia nonmotile: Clostridium
gangrene, Clostridial necrotizin
Peptostreptococcus
Peptostreptococcus ma
(non-spore forming)
Ureaplasma urealyticum (Ur

Mycoplasmataceae infection) · Mycoplasma genitalium ·
Mollicutes pneumoniae (Mycoplasma pneumon
Anaeroplasmatales Erysipelothrix rhusiopathiae

M: bac gr+f/ drug(
BAC t (clas) gr+a(t)/gr- J1p, w, n, m,
p(c)/gr-o vacc)
Retrieved from "http://en.wikipedia.org/wiki/Listeria_monocytogenes"
Categories: Listeriaceae | Foodborne illnesses

W000
Seahorse

This article is about the genus of fish. For the mythological sea-horse, see Hippocamp. For other
uses, see Seahorse (disambiguation).
Seahorses
Temporal range: 23–0 Ma
PreЄ
g
Lower Miocene to Present

Hippocampus sp.
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Subclass: Neopterygii
Infraclass: Teleostei
Order: Syngnathiformes
Family: Syngnathidae
Subfamily: Hippocampinae
Hippocampus
Genus:
Rafinesque, 1810[1]
Species
47, See text for species.

Seahorses compose the fish genus Hippocampus within the family Syngnathidae, in order
Syngnathiformes. Syngnathidae also includes the pipefishes. "Hippocampus" comes from the Ancient Greek
hippos meaning "horse" and kampos meaning “sea monster”.[2]
There are nearly 50 species of seahorse. They are mainly found in shallow tropical and temperate
waters throughout the world. They prefer to live in sheltered areas such as seagrass beds, coral reefs, or
mangroves. Colonies have been found in European waters such as the Thames Estuary.[3] From North
America down to South America there are approximately four species, ranging from the very small (dwarf
seahorses are only about 2.5 centimeters (1 in)) to much larger specimens off the Pacific Coast of Central
America (the foot-long H. ingens). H. erectus are larger seahorses that range from Nova Scotia to around
Uruguay. Three species live in the Mediterranean Sea: H. hippocampus (long snout), H. brevirostris (short
snout) and H. fuscus (immigrated from the Red Sea). These fish form territories, with males staying in about
1 square metre (11 sq ft) of their habitat while females range about one hundred times that area. They bob
around in sea grass meadows, mangrove stands, and coral reefs where they adopt murky brown and gray
patterns to camouflage themselves among the sea grass. During social moments or in unusual surroundings,
seahorses turn bright colors.
Contents
[hide]
• 1 Description
• 2 Evolution and fossil record
• 3 Reproduction
• 3.1 Courtship
• 3.2 Gestation
• 3.3 Birth
• 3.4 Questions surrounding reproductive roles
• 3.5 Monogamy
• 4 Feeding habits
• 5 In aquaria
• 6 Use in Chinese medicine
• 7 Taxonomy
• 7.1 Pygmy seahorses
• 8 Cultural references
• 9 Threats & Future
• 10 References
[edit] Description
Spiny seahorse Hippocampus hystrix from East Timor holding on to soft coral with its prehensile tail
Seahorses are named for their equine profile. Although they are bony fish, they do not have scales,
but rather a thin skin stretched over a series of bony plates arranged in rings throughout their body. Each
species has a distinct number of rings. Seahorses swim upright, another characteristic that is not shared by
their close pipefish relatives, which swim horizontally. Seahorses have a coronet on their head, which is
distinct to each individual, much like a human fingerprint. They swim very poorly by using a dorsal fin, which
they rapidly flutter and pectoral fins, located behind their eyes, which they use to steer. Seahorses have no
caudal fin. Since they are poor swimmers, they are most likely to be found resting, with their prehensile tails
wound around a stationary object. They have long snouts, which they use to suck up food, and eyes that can
move independently of each other, much like a chameleon. Seahorses eat small shrimp, tiny fish,
crustaceans and plankton.
[edit] Evolution and fossil record

Anatomical evidence, supported by molecular, physical, and genetic evidence, demonstrates that
seahorses are highly modified pipefish. The fossil record of seahorses, however, is very sparse. The best
known and best studied fossils are specimens of H. guttulatus (though literature more commonly refers to
them under the synonym of H. ramulosus), from the Marecchia River Formation of Rimini Province, Italy,
dating back to the Lower Pliocene, about 3 million years ago. The earliest known seahorse fossils are of two
pipefish-like species, H. sarmaticus and H. slovenicus from the coprolitic horizon of Tunjice Hills, a middle
Miocene lagerstätte in Slovenia dating back about 13 million years.[4] Molecular dating finds that pipefish
and seahorses separated during the Late Oligocene. This has led to speculation that seahorses evolved in
response to large areas of shallow-water, newly created as the result of tectonic events. The shallow water
allowed the expansion of seagrass habitats that selected for the camouflage offered by the seahorses’
upright posture.[5] These tectonic changes occurred in the Western Pacific Ocean suggesting an origin there
with molecular data suggesting two later and separate invasions of the Atlantic Ocean.[6]
[edit] Reproduction
Pregnant seahorse, New York Aquarium

The male seahorse is equipped with a brood pouch on the ventral, or front-facing, side. When
mating, the female seahorse deposits up to 1,500 eggs in the male's pouch. The male carries the eggs for
anywhere from 9 to 45 days until they emerge, expelling fully-developed, miniature seahorses in the water.
Once the seahorse babies are released into the water, the male's role is done and he offers no further care.
[7]
[edit] Courtship
Before breeding, seahorses court for several days. Scientists believe the courtship behavior
synchronizes the animals' movements so that the male can receive the eggs when the female is ready to
deposit them. During this time they may change color, swim side by side holding tails or grip the same strand
of sea grass with their tails and wheel around in unison in what is known as a “pre-dawn dance". They
eventually engage in a “true courtship dance" lasting about 8 hours, during which the male pumps water
through the egg pouch on his trunk which expands and opens to display its emptiness. When the female’s
eggs reach maturity, she and her mate let go of any anchors and snout-to-snout, drift upward out of the
seagrass, often spiraling as they rise. The female inserts her ovipositor into the male’s brood pouch and
deposits dozens to thousands of eggs. As the female releases her eggs, her body slims while his swells.
Both animals then sink back into the seagrass and she swims away.
[edit] Gestation
West Australian seahorse, H. subelongatus

The male releases his sperm directly into seawater where it fertilizes the eggs[8], which are then
embedded in the pouch wall and become surrounded by a spongy tissue.[9] The male supplies the eggs with
prolactin, the same hormone responsible for milk production in pregnant mammals. The pouch provides
oxygen as well as a controlled environment incubator. The eggs then hatch in the pouch where the salinity of
the water is regulated; this prepares the newborns for life in the sea.[10][11] Throughout gestation, which in
most species requires two to four weeks, his mate visits him daily for “morning greetings”. They interact for
about 6 minutes, reminiscent of courtship. The female then swims away until the next morning, and the male
returns to vacuuming up food through his snout.[10]
Research published in 2007 indicates the male releases sperm into the surrounding sea water during
fertilization, and not directly into the pouch as previously thought.[12]
[edit] Birth
White's seahorse, H. whitei

The number of young released by the male seahorse averages 100-200 for most species, but may
be as low as 5 for the smaller species, or as high as 1,500. When the fry are ready to be born, the male
expels them with muscular contractions. He typically gives birth at night and is ready for the next batch of
eggs by morning when his mate returns. Like almost all other fish species, seahorses do not nurture their
young after birth. Infants are susceptible to predators or ocean currents which wash them away from feeding
grounds or into temperatures too extreme for their delicate bodies. Fewer than .5% of infants survive to
adulthood, explaining why litters are so large. These survival rates are actually fairly high compared to other
fish, because of their protected gestation, making the process worth the great cost to the father. The eggs of
most other fish are abandoned immediately after fertilization.[11]
[edit] Questions surrounding reproductive roles

Reproduction is energetically costly to the male. This brings into question why the sexual role
reversal even takes place. In an environment where one partner incurs more energy costs than the other,
Bateman's principle suggests that the lesser contributor takes the role of the aggressor. Male seahorses are
more aggressive and sometimes “fight” for female attention. According to Amanda Vincent of Project
Seahorse, only males tail-wrestle and snap their heads at each other. This discovery prompted further study
of energy costs. To estimate the female’s direct contribution, researcher Heather D. Masonjones, associate
professor of biology at the University of Tampa, chemically analyzed the energy stored in each egg. To
measure the burden on the male, Masonjones measured its oxygen consumption. By the end of incubation,
the male consumed almost 33% more oxygen than before mating. The study concluded that the female's
energy expenditure while generating eggs is twice that of males during incubation[10] confirming the
standard hypothesis.
Why the male seahorse (and other members of Syngnathidae) carries the offspring through gestation
is unknown, though some researchers[citation needed] believe it allows for shorter birthing intervals, in turn
resulting in more offspring. Given an unlimited number of ready and willing partners, males have the potential
to produce 17 percent more offspring than females in a breeding season. Also, females have “time-outs” from
the reproductive cycle that are 1.2 times longer than those of males. This seems to be based on mate choice,
rather than physiology. When the female’s eggs are ready, she must lay them in a few hours or eject them
into the water column. Making eggs is a huge cost to her physically, since they amount to about a third of her
body weight. To protect against losing a clutch, the female demands a long courtship. The daily greetings
help to cement the bond between the pair.
[edit] Monogamy
One common misconception about seahorses is that they mate for life. Many species of seahorses
form pair bonds that last through at least the breeding season. Some species show a higher level of mate
fidelity than others.[13][14] However, many species readily switch mates when the opportunity arises. H.
abdominalis and H. breviceps have been shown to breed in groups, showing no continuous mate preference.
Many more species mating habits have not been studied, so it is unknown how many species are actually
monogamous, or how long those bonds actually last.[15]
Although monogamy within fish is not common, it does appear to exist for some. In this case, the
mate guarding hypothesis may be an explanation. This hypothesis states “males remain with a single female
because of ecological factors that make male parental care and protection of offspring especially
advantageous.”[16] Because the rates of survival for newborn seahorses are so low, incubation is essential.
Though not proven, males could have taken on this role because of the lengthy period the females require to
produce their eggs. If males incubate while females prepare the next clutch (amounting to 1/3 of body
weight), they can reduce the interval between clutches.
[edit] Feeding habits
Seahorses feed on small crustaceans floating in the water or crawling on the bottom. With excellent
camouflage and a lot of patience, seahorses ambush prey that float within striking range. Mysid shrimp and
other small crustaceans are favorites, but some seahorses have been observed eating other kinds of
invertebrates and even larval fish.
[edit] In aquaria
Seahorses (Hippocampus erectus) at the New England Aquarium, USA.
While many aquarium hobbyists keep seahorses as pets, seahorses collected from the wild tend to
fare poorly in home aquaria. Many eat only live foods such as brine shrimp and are prone to stress, which
damages their immune systems and makes them susceptible to disease.
In recent years, however, captive breeding has become more popular. Such seahorses survive better
in captivity, and are less likely to carry diseases. They eat frozen mysidacea (crustaceans) that are readily
available from aquarium stores,[17] and do not experience the stress of moving out of the wild. Although
captive-bred seahorses are more expensive, they take no toll on wild populations.
Seahorses should be kept in an aquarium to themselves, or with compatible tank-mates. Seahorses
are slow feeders, and fast, aggressive feeders will leave them without food.[17]
Seahorses can co-exist with many species of shrimp and other bottom-feeding creatures. Gobies
also make good tank-mates. Keepers are generally advised to avoid eels, tangs, triggerfish, squid, octopus,
and sea anemones.[18]
Animals sold as "freshwater seahorses" are usually the closely related pipefish, of which a few
species live in the lower reaches of rivers. The supposed true "freshwater seahorse" called H. aimei was not
a real species, but a name sometimes used for Barbour's and Hedgehog seahorses. The latter is a species
that can be found in brackish waters, but not actually a freshwater fish.[19]
[edit] Use in Chinese medicine
Medicinal seahorse.
Seahorses, silkie, and cordyceps after being used to make a broth
Seahorse and scorpion skewers as street food in China
Seahorse populations are thought to have been endangered in recent years by overfishing and
habitat destruction. The seahorse is used in traditional Chinese herbology, and as many as 20 million
seahorses may be caught each year and sold for this purpose.[20] Medicinal seahorses are not readily bred
in captivity as they are susceptible to disease and it is believed that they have different medicinal properties
from aquarium seahorses. Seahorses are also used as medicines by the Indonesians, the Central Filipinos,
and many other ethnic groups around the world.
Import and export of seahorses has been controlled under CITES since May 15, 2004. However,
Indonesia, Japan, Norway, and South Korea have chosen to opt out of the trade rules set by CITES.
The problem may be exacerbated by the growth of pills and capsules as the preferred method of
ingesting medication as they are cheaper and more available than traditional, individually tailored
prescriptions of raw medicinals but the contents are harder to track. Seahorses once had to be of a certain
size and quality before they were accepted by TCM practitioners and consumers. But declining availability of
the preferred large, pale and smooth seahorses has been offset by the shift towards prepackaged medicines,
which make it possible for TCM merchants to sell previously unused juvenile, spiny and dark-coloured
animals. Today almost a third of the seahorses sold in China are prepackaged. This adds to the pressure on
the species.[21]
[edit] Taxonomy
Common seahorse, Hippocampus kuda

• Genus Hippocampus
• Big-belly seahorse, H. abdominalis Lesson, 1827 (New Zealand and south and east
Australia)
• Winged seahorse, H. alatus Kuiter, 2001
• West African seahorse, H. algiricus Kaup, 1856
• Narrow-bellied seahorse, H. angustus Günther, 1870
• Barbour's seahorse, H. barbouri Jordan & Richardson, 1908
• Pygmy seahorse, H. bargibanti Whitley, 1970 West Pacific area (Indonesia,
Philippines, Papua New Guinea, Solomon Islands, etc.)
• False-eyed seahorse, H. biocellatus Kuiter, 2001
• Réunion seahorse, H. borboniensis Duméril, 1870
• Short-head seahorse or knobby seahorse, H. breviceps Peters, 1869 (south and east
Australia)
• Giraffe seahorse, H. camelopardalis Bianconi, 1854
• Knysna seahorse, H. capensis Boulenger, 1900
• H. colemani Kuiter, 2003
• Tiger tail seahorse, H. comes Cantor, 1850
• Crowned seahorse, H. coronatus Temminck & Schlegel, 1850
• Denise's pygmy seahorse, H. denise Lourie & Randall, 2003
• Lined seahorse, H. erectus Perry, 1810 (east coast of the Americas, between Nova
Scotia and Uruguay)
• Fisher's seahorse, H. fisheri Jordan & Evermann, 1903
• Sea pony, H. fuscus Rüppell, 1838 (Indian Ocean)
• Big-head seahorse, H. grandiceps Kuiter, 2001
• Long-snouted seahorse, H. guttulatus Cuvier, 1829
• Eastern spiny seahorse, H. hendriki Kuiter, 2001
• Short-snouted seahorse, H. hippocampus (Linnaeus, 1758) (Mediterranean Sea and
Atlantic Ocean)
• Thorny seahorse, H. histrix Kaup, 1856 (Indian Ocean, Persian Gulf, Red Sea, and
the Far East)
• Pacific seahorse, H. ingens Girard, 1858 (Pacific coast of North, Central and South
America)
• Jayakar's seahorse, H. jayakari Boulenger, 1900
• Collared seahorse, H. jugumus Kuiter, 2001
• Great seahorse, H. kelloggi Jordan & Snyder, 1901
• Common seahorse, H. kuda Bleeker, 1852
• Lichtenstein's seahorse, H. lichtensteinii Kaup, 1856
• Bullneck seahorse, H. minotaur Gomon, 1997
• Japanese seahorse, H. mohnikei Bleeker, 1854
• Monte Bello seahorse, H. montebelloensis Kuiter, 2001
• Northern spiny seahorse, H. multispinus Kuiter, 2001
• H. pontohi Lourie and Kuiter, 2008
• High-crown seahorse, H. procerus Kuiter, 2001
• Queensland seahorse, H. queenslandicus Horne, 2001
• Longsnout seahorse, H. reidi Ginsburg, 1933 (Caribbean coral reefs)
• Satomi's pygmy seahorse, H. satomiae Lourie and Kuiter, 2008
• Half-spined seahorse, H. semispinosus Kuiter, 2001
• H. severnsi Lourie and Kuiter, 2008
• Shiho's seahorse, H. sindonis Jordan & Snyder, 1901
• Hedgehog seahorse, H. spinosissimus Weber, 1913
• West Australian seahorse, H. subelongatus Castelnau, 1873
• Longnose seahorse, H. trimaculatus Leach, 1814
• White's seahorse, H. whitei Bleeker, 1855 (east Australia)
• Zebra seahorse, H. zebra Whitley, 1964
• Dwarf seahorse, H. zosterae Jordan & Gilbert, 1882 (Gulf of Mexico and the
Caribbean)
[edit] Pygmy seahorses
Hippocampus satomiae attached to a coral

Pygmy Seahorses are less than 15 millimeters (0.6 in) tall and 17 millimeters (0.7 in) wide members
of the genus. Previously the term was applied exclusively to the species H. bargibanti but since 1997,
discoveries have made this term obsolete. The species H. minotaur, H. denise, H. colemani, H. pontohi, H.
severnsi and H. satomiae have been described. Other species that are believed to be unclassified have also
been reported in books, dive magazines and on the Internet. They can be distinguished from other species of
seahorse by their 12 trunk rings, low number of tail rings (26–29), the location in which young are brooded in
the trunk region of males and their extremely small size.[22] Molecular analysis (of ribosomal RNA) of 32
Hippocampus species found that H. bargibanti belongs in a separate clade from other members of the genus
and therefore that the species diverged from the other species in the "ancient" past.[23]
Most pygmy seahorses are well camouflaged and live in close association with other organisms
including colonial hydrozoans (Lytocarpus and Antennellopsis), coralline algae (Halimeda) sea fans
(Muricella, Annella, Acanthogorgia). This combined with their small size accounts for why most species have
only been noticed in recent years.[22]
[edit] Cultural references

This "In popular culture" section may contain minor or trivial references. Please
reorganize this content to explain the subject's impact on popular culture rather than simply
listing appearances, and remove trivial references. (November 2009)
'Fucus like seahorse' from Lydekker's The Royal Natural History
A sculpture of a heraldic seahorse that adorned an 18th or 19th century French naval vessel
• In heraldry, a seahorse is depicted as a creature with the foreparts of a horse and the
hindparts of a fish.
[edit] Threats & Future

Most seahorse species are data deficient. This means there is not enough information to make a
proper assessment about their risk of extinction. Because seahorse population is unknown, there is a greater
risk of losing more seahorses because of the lack of information about how many are dying each year, how
many are being born, how many are used for souvenirs, etc. Seahorse habitats are in great danger though.
Coral reefs and seagrass beds are deteriorating, meaning seahorses have fewer places to live. Also as
stated above, seahorses are used to Chinese Medicine and as souvenirs, which definitely cuts their
population down significantly each year. [24]
[edit] References
1. ^ Hippocampus Rafinesque, 1810, WoRMS
2. ^ http://en.wiktionary.org/wiki/hippocampus Hippocampus at Wiktionary
3. ^ "Rare seahorses breeding in Thames". BBC News. 2008-04-07.
http://news.bbc.co.uk/1/hi/england/london/7333980.stm. Retrieved 2009-11-11.
4. ^ Žalohar J., Hitij T., Križnar M. (2009). "Two new species of seahorses (Syngnathidae,
Hippocampus) from the Middle Miocene (Sarmatian) Coprolitic Horizon in Tunjice Hills, Slovenia:
The oldest fossil record of seahorses". Annales de Paléontologie 95 (2): 71–96.
doi:10.1016/j.annpal.2009.03.002.
5. ^ Teske PR, Beheregaray LB (2009). "Evolution of seahorses' upright posture was linked to
Oligocene expansion of seagrass habitats". Biol Lett. 5 (4): 521–3. doi:10.1098/rsbl.2009.0152.
PMID 19451164.
6. ^ Teske PR, Cherry MI, Matthee CA (2004). "The evolutionary history of seahorses
(Syngnathidae: Hippocampus): molecular data suggest a West Pacific origin and two invasions of the
Atlantic Ocean". Mol Phylogenet Evol. 30 (2): 273–86. doi:10.1016/S1055-7903(03)00214-8.
PMID 14715220.
7. ^ Foster S.J, Vincent C.J. (2004). "Life history and ecology of seahorses: implications for
conservation and management". Journal of Fish Biology 65: 1–61. doi:10.1111/j.1095-
8649.2004.00429.x. http://www.seaturtle.org/PDF/Foster_2004_JFishBiol.pdf.
8. ^ Sex and the seahorse
9. ^ "The biology of seahorses: Reproduction". The Seahorse Project.
http://seahorse.fisheries.ubc.ca/biology5.html. Retrieved 2007-05-08.
10.^ a b c Milius, Susan (2000). "Pregnant—and Still Macho - seahorses | Science News | Find
Articles at BNET.com". Findarticles.com.
http://findarticles.com/p/articles/mi_m1200/is_11_157/ai_61291647/pg_1. Retrieved 2009-11-11.
11.^ a b "Seahorse Fathers Take Reins in Childbirth". News.nationalgeographic.com.
http://news.nationalgeographic.com/news/2002/06/0614_seahorse_recov.html. Retrieved 2009-11-
11.
12.^ Connor, Steve (2007-01-19). "Sex and the seahorse - Science, News - Independent.co.uk".
London: News.independent.co.uk.
http://news.independent.co.uk/world/science_technology/article2165477.ece. Retrieved 2009-11-11.
13.^ Kvarnemo C, Moore G.I, Jones A.G, Nelson W.S, Avise J.C. (2000). "Monogamous pair
bonds and mate switching in the Western Australian seahorse Hippocampus subelongatus". J. Evol.
Biol. 13 (6): 882–8. doi:10.1046/j.1420-9101.2000.00228.x. http://www3.interscience.wiley.com/cgi-
bin/fulltext/119188977/PDFSTART.
14.^ Vincent C.J, Sadler L.M (1995). "Faithful pair bonds in wild seahorses, Hippocampus
whitei'". Anim. Behav. 50: 1557–1569. doi:10.1016/0003-3472(95)80011-5.
http://courses.umass.edu/wfcon470/Vincent%20and%20Sadler%2095.pdf.
15.^ "What’s Love Got to Do With It? The Truth About Seahorse Monogamy". fusedjaw.com.
http://www.fusedjaw.com/biology/seahorse-monogamy/. Retrieved 2010-10-09.
16.^ Alcock, John. Animal Behavior (8th ed.). Massachusetts: Sinauer. pp. 370–1.
17.^ a b "Seahorse and Pipefish Foods | Tami Weiss". Fusedjaw.com. 2005-06-25.
http://www.fusedjaw.com/food-and-nutrition/seahorse-foods-and-feeding/. Retrieved 2009-11-11.
18.^ "Seahorse Tankmates | Will Wooten". Fusedjaw.com. 2004-06-25.
http://www.fusedjaw.com/aquariumcare/seahorse-tankmates-whats-safe-whats-not/. Retrieved 2009-
11-11.
19.^ "Hippocampus spinosissimus". Fishbase.
http://fishbase.org/Summary/speciesSummary.php?
ID=25974&genusname=Hippocampus&speciesname=spinosissimus. Retrieved 2009-11-11.
20.^ "Seahorse Crusader Amanda Vincent" on Nova television show
21.^ New Scientist, January 1995 "Can we tame wild medicine"?
22.^ a b Lourie, Sara; Rudie Kuiter (2008). "Three new pygmy seahorse species from Indonesia
(Teleostei: Syngnathidae: Hippocampus)". Zootaxa (Magnolia Press) 1963: 54–68. ISSN 1175-5334.
http://seahorse.fisheries.ubc.ca/pubs/2008/Lourie_and_Kuiter2008.pdf. Retrieved 9 June 2009.
23.^ Teske, Peter; Michael Cherry and Conrad Matthee (February 2004). "The evolutionary
history of seahorses (Syngnathidae: Hippocampus): molecular data suggest a West Pacific origin
and two invasions of the Atlantic Ocean". Molecular Phylogenetics and Evolution 30 (2): 273–286.
doi:10.1016/S1055-7903(03)00214-8. PMID 14715220.
24.^ Lourie, Sarah A., Sarah J. Foster, Ernest W.T. Cooper, and Amanda C.J. Vincent. "A Guide
to the Identification of Seahorses." Project Seahorse Advancing Marine Conservation (2004): 1-120.
Print.

• Amanda C.J. Vincent and Laila M. Sadler (1995). "Faithful pair bonds in wild seahorse,
Hippocampus whitei". Animal Behaviour 50: 1557–69. doi:10.1016/0003-3472(95)80011-5.
• Amanda C.J. Vincent (1995). "A role for daily greetings in maintaining seahorse pair bonds".
Animal Behaviour 49: 258–260. doi:10.1016/0003-3472(95)80178-2.
• Amanda C.J. Vincent (1990). "A seahorse father makes a good mother". Natural History 12:
34–43.
• Amanda C.J. Vincent and Rosie Woodroffe (1994). "Mothers little helpers: patterns of male
care in mammals". Trends in Ecology and Evolution 9: 294–7. doi:10.1016/0169-5347(94)90033-7.
• John Sparks (1999). Battle of the Sexes: The Natural History of Sex. London: BBC Books.
ISBN 0563371455.
• Sara A. Lourie, Amanda C.J. Vincent and Heather J. Hall (1999). Seahorses: An
Identification Guide to the World's Species and their Conversation . London: Project Seahorse.
• Teske P. R., Hamilton H., Matthee C. A. & Barker N. P. (2007). "Signatures of seaway
closures and founder dispersal in the phylogeny of a circumglobally distributed seahorse lineage".
BMC Evolutionary Biology 7: 138. doi:10.1186/1471-2148-7-138. PMID 17697373. PMC 1978501.
http://www.biomedcentral.com/1471-2148/7/138.
Wikimedia Commons has media related to: Seahorse
• Vimeo.com, Video of various seahorse species at the Monterey Bay Aquarium

• ARKive—images and movies of the pygmy seahorse (Hippocampus bargibanti)
• FusedJaw.com, Information about seahorses
• ProjectSeahorse.org, Research and Conservation Group
• Seahorse.org, Aquarium Hobby
• DiveGallery.com, Seahorse Photos
• RobertoSozzani.it, Hippocampus denise Pygmy seahorse photo galleries
• RobertoSozzani.it, Pontohi Pygmy Seahorse
• RobertoSozzani.it, Walea Pygmy Seahorse
• Arnes.si, Fossil Seahorses from Tunjice hills (Slovenia)
• QataVisitor.com n Sea horses in Qatar
Retrieved from "http://en.wikipedia.org/wiki/Seahorse"
Categories: Hippocampus | Syngnathidae | Animals that can change color

W000
Timeline of human evolution


This article is about the timeline of human evolution. For a timeline of general evolution see Timeline
of evolution.
See also Human evolution for more details on this topic.
Haeckel's Paleontological Tree of Vertebrates (c. 1879).
The evolutionary history of species has been described as a "tree", with many branches arising from a single
trunk. While Haeckel's tree is somewhat outdated, it illustrates clearly the principles that more complex
modern reconstructions can obscure.
The timeline of human evolution outlines the major events in the development of human species, and
the evolution of humans' ancestors. It includes a brief explanation of some animals, species or genera, which
are possible ancestors of Homo sapiens sapiens. It does not address the origin of life, which is addressed by
abiogenesis, but presents a possible line of descendants that led to humans. This timeline is based on
studies from paleontology, developmental biology, morphology and from anatomical and genetic data. The
study of human evolution is a major component of anthropology.
Contents
[hide]
• 1 Homo sapiens taxonomy
• 2 Timeline
• 2.1 First living beings
• 2.2 Chordates
• 2.3 Tetrapods
• 2.4 Mammals
• 2.5 Primates
• 2.6 Hominidae
• 2.7 Homo
• 3 See also
• 4 References
[edit] Homo sapiens taxonomy

The cladistic line of descent (taxonomic rank) of Homo sapiens (modern humans) is as follows:
domain: eukaryotes (2,100,000,000 years ago)
kingdom: animalia (590,000,000 years ago)
phylum: chordata (530,000,000 years ago)
subphylum: vertebrata (505,000,000 years ago)
class: mammalia (220,000,000 years ago)
subclass: theriiformes
infraclass: eutheria (125,000,000 years ago)
magnorder: boreoeutheria
superorder: euarchontoglires (supraprimates) (100,000,000 years ago)
order: primates (75,000,000 years ago)
suborder: haplorrhini (tarsiers, monkeys, apes, "dry-nosed" primates) (40,000,000 years ago)
infraorder: simiiformes (simians, "higher" primates)
parvorder: catarrhini ("narrow nosed" primates) (30,000,000 years ago)
superfamily: hominoidea (apes) (28,000,000 years ago)
family: hominidae (great apes) (15,000,000 years ago)
subfamily: homininae (8,000,000 years ago)
tribe: hominini (5,800,000 years ago)
subtribe: hominina (3,000,000 years ago)
genus: homo (2,500,000 years ago)
species: homo sapiens (500,000 years ago)
sub-species: homo sapiens sapiens (200,000 years ago)
[edit] Timeline
[edit] First living beings
Date Event
4000 Ma The earliest life appears.
(million Further information: Origin of life
years ago)
3900 Ma Cells resembling prokaryotes appear. This marks the first appearance of
photosynthesis and therefore the first occurrence of large quantities of oxygen on the
earth.
Further information: Cell_(biology)#Evolution
2500 Ma First organisms to utilize oxygen. By 2400 Ma, in what is referred to as the Great
Oxygenation Event, the pre-oxygen anerobic forms of life were wiped out by the oxygen
consumers.
2100 Ma More complex cells appear: the eukaryotes.

Further information: Eukaryote#Origin and evolution
1200 Ma Sexual reproduction evolves, leading to faster evolution.[1]
900 Ma
Choanoflagellate
The choanoflagellates may look similar to the ancestors of the entire animal
kingdom, and in particular they may be the direct ancestors of Sponges.[2]
Proterospongia (members of the Choanoflagellata) are the best living examples of what
the ancestor of all animals may have looked like.
They live in colonies, and show a primitive level of cellular specialization for
different tasks.
600 Ma It is thought that the earliest multicellular animal was a sponge-like creature.
Sponges are among the simplest of animals, with partially differentiated tissues.
Sponges (Porifera) are the phylogenetically oldest animal phylum extant today.
580 Ma Animal movement may have started with cnidarians. Almost all cnidarians
possess nerves and muscles. Because they are the simplest animals to possess them,
their direct ancestors were very likely the first animals to use nerves and muscles
together. Cnidarians are also the first animals with an actual body of definite form and
shape. They have radial symmetry. The first eyes evolved at this time.
550 Ma
Flatworm
Flatworms are the earliest animals to have a brain, and the simplest animals
alive to have bilateral symmetry. They are also the simplest animals with organs that
form from three germ layers.
540 Ma Acorn worms are considered more highly specialised and advanced than other
similarly shaped worm-like creatures. They have a circulatory system with a heart that
also functions as a kidney. Acorn worms have a gill-like structure used for breathing, a
structure similar to that of primitive fish. Acorn worms are thus sometimes said to be a
link between vertebrates and invertebrates[citation needed].
[edit] Chordates
Date Event
530 Ma
Pikaia
Pikaia is an iconic ancestor of modern chordates and vertebrates.[3] Other, earlier
chordate predecessors include Myllokunmingia fengjiaoa[4], Haikouella lanceolata[5], and
Haikouichthys ercaicunensis[6].
The lancelet, still living today, retains some characteristics of the primitive
chordates. It resembles Pikaia.
Conodont
Conodonts are a famous type of early (495 Mya and later) chordate fossil; they are
the peculiar teeth of an eel-shaped animal characterised by large eyes, fins with fin rays,
chevron-shaped muscles and a notochord. The animal is sometimes called a conodont, and
sometimes a conodontophore (conodont-bearer) to avoid confusion.
505 Ma
Agnatha
The first vertebrates appear: the ostracoderms, jawless fish related to present-day
lampreys and hagfishes. Haikouichthys and Myllokunmingia are examples of these jawless
fish, or Agnatha. (See also prehistoric fish). They were jawless and their internal skeletons
were cartilaginous. They lacked the paired (pectoral and pelvic) fins of more advanced fish.
They were precursors to the Osteichthyes (bony fish).[7]
480 Ma
A Placoderm
The Placodermi were prehistoric fishes. Placoderms were the first of the jawed
fishes, their jaws evolving from the first of their gill arches [8]. Their head and thorax were
covered by articulated armoured plates and the rest of the body was scaled or naked.
410 Ma The first Coelacanth appears[9]; this order of animals had been thought to have no
extant members until living specimens were discovered in 1938. It is often referred to as a
living fossil.
[edit] Tetrapods
Date Event
390 Ma
Panderichthys
Some fresh water lobe-finned fish (Sarcopterygii) develop legs and give rise to the
Tetrapoda.
The first tetrapods evolved in shallow and swampy freshwater habitats.
Primitive tetrapods developed from a lobe-finned fish (an "osteolepid
Sarcopterygian"), with a two-lobed brain in a flattened skull, a wide mouth and a short
snout, whose upward-facing eyes show that it was a bottom-dweller, and which had already
developed adaptations of fins with fleshy bases and bones. The "living fossil" coelacanth is
a related lobe-finned fish without these shallow-water adaptations. These fishes used their
fins as paddles in shallow-water habitats choked with plants and detritus. The universal
tetrapod characteristics of front limbs that bend backward at the elbow and hind limbs that
bend forward at the knee can plausibly be traced to early tetrapods living in shallow water.
[10]
Panderichthys is a 90–130 cm (35–50 in) long fish from the Late Devonian period
(380 Mya). It has a large tetrapod-like head. Panderichthys exhibits features transitional
between lobe-finned fishes and early tetrapods.
Trackway impressions made by something that resembles Ichthyostega's limbs
were formed 390 Ma in Polish marine tidal sediments. This suggests tetrapod evolution is
older than the dated fossils of Panderichthys through to Ichthyostega.
Lungfishes retain some characteristics of the early Tetrapoda. One example is the
Queensland Lungfish.
375 Ma Tiktaalik is a genus of sarcopterygian (lobe-finned) fishes from the late Devonian
with many tetrapod-like features. It shows a clear link between Panderichthys and
Acanthostega.
365 Ma
Acanthostega
Ichthyostega
Acanthostega is an extinct amphibian, among the first animals to have recognizable
limbs. It is a candidate for being one of the first vertebrates to be capable of coming onto
land. It lacked wrists, and was generally poorly adapted for life on land. The limbs could not
support the animal's weight. Acanthostega had both lungs and gills, also indicating it was a
link between lobe-finned fish and terrestrial vertebrates.
Ichthyostega is an early tetrapod. Being one of the first animals with legs, arms, and
finger bones, Ichthyostega is seen as a hybrid between a fish and an amphibian.
Ichthyostega' had legs but its limbs probably weren't used for walking. They may have
spent very brief periods out of water and would have used their legs to paw their way
through the mud.[11]
Amphibia were the first four-legged animals to develop lungs which may have
evolved from Hynerpeton 360 Mya.
Amphibians living today still retain many characteristics of the early tetrapods.
300 Ma
Hylonomus
From amphibians came the first reptiles: Hylonomus is the earliest known reptile. It
was 20 cm (8 in) long (including the tail) and probably would have looked rather similar to
modern lizards. It had small sharp teeth and probably ate millipedes and early insects. It is
a precursor of later Amniotes and mammal-like reptiles. Α-keratin first evolves here which is
used in claws in modern lizards and birds, and hair in mammals.[12]
Evolution of the amniotic egg gives rise to the Amniota, reptiles that can reproduce
on land and lay eggs on dry land. They did not need to return to water for reproduction. This
adaptation gave them the capability to colonize the uplands for the first time.
Reptiles have advanced nervous systems, compared to amphibians. They have
twelve pairs of cranial nerves.
[edit] Mammals
Date Event
256 Ma
Phthinosuchus, an early Therapsid

Shortly after the appearance of the first reptiles, two branches split off. One branch
is the Diapsids, from which come the modern reptiles. The other branch is Synapsida,
which had temporal fenestra, a pair of holes in their skulls behind the eyes, which were
used to increase the space for jaw muscles.
The earliest mammal-like reptiles are the pelycosaurs. The pelycosaurs were the
first animals to have temporal fenestra. Pelycosaurs are not Therapsids but soon they gave
rise to them. The Therapsida were the direct ancestor of mammals.
The therapsids have temporal fenestrae larger and more mammal-like than
pelycosaurs, their teeth show more serial differentiation; and later forms had evolved a
secondary palate. A secondary palate enables the animal to eat and breathe at the same
time and is a sign of a more active, perhaps warm-blooded, way of life.[13]
220 Ma One sub-group of therapsids, the cynodonts evolved more mammal-like

characteristics.
The jaws of cynodonts resemble modern mammal jaws. It is very likely this group of
animals contains a species which is the direct ancestor of all modern mammals.[14]
220 Ma
Repenomamus
From Eucynodontia (cynodonts) came the first mammals. Most early mammals
were small and shrew-like animals that fed on insects. Although there is no evidence in the
fossil record, it is likely that these animals had a constant body temperature, milk glands for
their young. The neocortex region of the brain first evolved in mammals and thus is unique
to them.
125 Ma
Eomaia scansoria
Eomaia scansoria, a eutherian mammal, leads to the formation of modern placental
mammals. It looks like modern dormouse, climbing small shrubs in Liaoning, China.
Monotremes are an egg laying group of mammals represented amongst modern
animals by the platypus and spiny anteaters that split away from the eutherian mammals.
Recent genome sequencing of the platypus indicates that its sex genes are closer to that of
birds. Thus it can be inferred that the first mammals to gain single paired sex determining
genes (XX or XY) evolved at or after this point within the eutherian group.
100 Ma Common genetic ancestor of mice and humans (base of the clade
Euarchontoglires).
[edit] Primates
Date Event
65–85 Ma
Plesiadapis
Carpolestes simpsoni
A group of small, nocturnal and arboreal, insect-eating mammals called the
Euarchonta begins a speciation that will lead to the primate, treeshrew and flying lemur
orders. The Primatomorpha is a subdivision of Euarchonta that includes the primates and
the proto-primate Plesiadapiformes. One of the early proto-primates is Plesiadapis.
Plesiadapis still had claws and the eyes located on each side of the head. Because of this
they were faster on the ground than on the top of the trees, but they began to spend long
times on lower branches of trees, feeding on fruits and leaves. The Plesiadapiformes very
likely contain the species which is the ancestor of all primates.[15]
One of the last Plesiadapiformes is Carpolestes simpsoni. It had grasping digits
but no forward facing eyes.
40 Ma Primates diverge into suborders Strepsirrhini (wet-nosed primates) and

Haplorrhini (dry nosed primates). Strepsirrhini contain most of the prosimians; modern
examples include the lemurs and lorises. The haplorrhines include the three living
groups: prosimian tarsiers, simian monkeys, and apes. One of the earliest haplorrhines is
Teilhardina asiatica, a mouse-sized, diurnal creature with small eyes. The Haplorrhini
metabolism lost the ability to make its own Vitamin C. This means that it and all its
descendants had to include fruit in its diet, where Vitamin C could be obtained externally.
30 Ma
Aegyptopithecus
Haplorrhini splits into infraorders Platyrrhini and Catarrhini. Platyrrhines, New
World monkeys, have prehensile tails and males are color blind. They may have migrated
to South America on a raft of vegetation across the Atlantic ocean (circa 4,500 km,
2,800 mi). Catarrhines mostly stayed in Africa as the two continents drifted apart. One
ancestor of catarrhines might be Aegyptopithecus.
25 Ma
Proconsul
Catarrhini splits into 2 superfamilies, Old World monkeys (Cercopithecoidea) and
apes (Hominoidae). Our trichromatic color vision had its genetic origins in this period.
Proconsul was an early genus of catarrhine primates. They had a mixture of Old
World monkey and ape characteristics. Proconsul's monkey-like features include thin
tooth enamel, a light build with a narrow chest and short forelimbs, and an arboreal
quadrupedal lifestyle. Its ape-like features are its lack of a tail, ape-like elbows, and a
slightly larger brain relative to body size.
Proconsul africanus is a possible ancestor of both great and lesser apes, and
humans.
[edit] Hominidae
Date Event
15 Ma Hominidae (great apes) speciate from the ancestors of the gibbon (lesser apes).
13 Ma Homininae ancestors speciate from the ancestors of the orangutan[16].

Pierolapithecus catalaunicus is believed to be a common ancestor of humans and
the great apes or at least a species that brings us closer to a common ancestor than any
previous fossil discovery.
Pierolapithecus had special adaptations for tree climbing, just as humans and other
great apes do: a wide, flat ribcage, a stiff lower spine, flexible wrists, and shoulder blades
that lie along its back.
10 Ma Hominini speciate from the ancestors of the gorillas.

7 Ma
Sahelanthropus tchadensis
Hominina speciate from the ancestors of the chimpanzees. The latest common
ancestor lived around the time of Sahelanthropus tchadensis, ca. 7 Ma [3]; S. tchadensis is
sometimes claimed to be the last common ancestor of humans and chimpanzees, but this is
disputed. The earliest known human ancestor post-dating the separation of the human and
the chimpanzee lines is Orrorin tugenensis (Millennium Man, Kenya; ca. 6 Ma). Both
chimpanzees and humans have a larynx that repositions during the first two years of life to a
spot between the pharynx and the lungs, indicating that the common ancestors have this
feature, a precursor of speech.
4.4 Ma Ardipithecus is a very early hominin genus (subfamily Homininae). Two species are
described in the literature: A. ramidus, which lived about 4.4 million years ago[17] during the
early Pliocene, and A. kadabba, dated to approximately 5.6 million years ago[18] (late
Miocene). A. ramidus had a small brain, measuring between 300 and 350 cm3. This is about
the same size as modern bonobo and female common chimpanzee brain, but much smaller
than the brain of australopithecines like Lucy (~400 to 550 cm3) and slightly over a fifth the
size of the modern Homo sapiens brain. Ardipithecus was aboreal, meaning it lived largely
in the forest where it competed with other forest animals for food, including the
contemporary ancestor for the chimpanzees. Ardipithecus was likely bipedal as evidenced
by its bowl shaped pelvis, the angle of its foramen magnum and its thinner wrist bones,
though its feet were still adapted for grasping rather than walking for long distances.
3.6 Ma
Australopithecus afarensis
Some Australopithecus afarensis left human-like footprints on volcanic ash in
Laetoli, Kenya (Northern Tanzania) which provides strong evidence of full-time bipedalism.
Australopithecus afarensis lived between 3.9 and 2.9 million years ago. It is thought that A.
afarensis was ancestral to both the genus Australopithecus and the genus Homo.
Compared to the modern and extinct great apes, A. afarensis has reduced canines and
molars, although they are still relatively larger than in modern humans. A. afarensis also has
a relatively small brain size (~380–430 cm³) and a prognathic (i.e. projecting anteriorly) face.
Australopithecines have been found in Savannah environments and likely increased its diet
to include meat from scavenging opportunities. An analysis of Australopithecus africanus
lower vertebrae suggests that females had changes to support bipedalism even while
pregnant.
3.5 Ma Kenyanthropus platyops, a possible ancestor of Homo, emerges from the
Australopithecus genus.
3 Ma The bipedal australopithecines (a genus of the Hominina subtribe) evolve in the
savannas of Africa being hunted by Dinofelis. Loss of body hair takes place in the period 3-2
Ma, in parallel with the development of full bipedalism.
[edit] Homo
Date Event
2.5 Ma
Homo habilis
Appearance of Homo. Homo habilis is thought to be the ancestor of the lankier and
more sophisticated Homo ergaster. Lived side by side with Homo erectus until at least 1.44
Ma, making it highly unlikely that Homo erectus directly evolved out of Homo habilis. First
stone tools, beginning of the Lower Paleolithic.
Further information: Homo rudolfensis
1.8 Ma
A reconstruction of Homo erectus.

Homo erectus evolves in Africa. Homo erectus would bear a striking resemblance to
modern humans, but had a brain about 74 percent of the size of modern man. Its forehead is
less sloping and the teeth are smaller. Other hominid designations such as Homo georgicus,
Homo ergaster, Homo pekinensis, Homo heidelbergensis are often put under the umbrella
species name of Homo erectus.[19] Starting with Homo georgicus found in what is now the
Republic of Georgia dated at 1.8 Ma, the pelvis and backbone grew more human-like and
gave H. georgicus the ability cover very long distances in order to follow herds of other
animals. This is the oldest fossil of a hominid found (so far) outside of Africa. Control of fire
by early humans is achieved 1.5 Ma by Homo ergaster. Homo ergaster reaches a height of
around 1.9 metres (6.2 ft). Evolution of dark skin, which is linked to the loss of body hair in
human ancestors, is complete by 1.2 Ma. Homo pekinensis first appears in Asia around 700
Ka but according to the theory of a recent African origin of modern humans, they could not
be human ancestors, but rather, were just a cousin offshoot species from Homo ergaster.
Homo heidelbergensis was a very large hominid that had a more advanced complement of
cutting tools and may have hunted big game such as horses.
1.2 Ma Homo antecessor is the common genetic ancestor of humans and Neanderthal.[20]
At present estimate, humans have approximately 20,000–25,000 genes and share 99% of
their DNA with the now extinct Neanderthal [21] and 95-99% of their DNA with their closest
living evolutionary relative, the chimpanzees[22][23] The human variant of the FOXP2 gene
(linked to the control of speech) has been found to be identical in Neanderthals.[24] It can
therefore be deduced that Homo antecessor would also have had the human FOXP2 gene.
600 ka
A reconstruction of Homo heidelbergensis
Three 1.5 m (5 ft) tall Homo heidelbergensis left footprints in powdery volcanic ash
solidified in Italy. Homo heidelbergensis is the common ancestor of both Homo
neanderthalensis and Homo sapiens. It is morphologically very similar to Homo erectus but
Homo heidelbergensis had a larger brain-case, about 93% the size of that of Homo sapiens.
The holotype of the species was tall, 1.8 m (6 ft) and more muscular than modern humans.
Beginning of the Middle Paleolithic.
200 ka
Homo sapiens sapiens (Pioneer plaque)
Omo1, Omo2 (Ethiopia, Omo river) are the earliest fossil evidence for archaic Homo
sapiens, evolved from Homo heidelbergensis[citation needed].
160 ka Homo sapiens (Homo sapiens idaltu) in Ethiopia, Awash River, Herto village,
practice mortuary rituals and butcher hippos. Potential earliest evidence of behavioral
modernity consistent with the continuity hypothesis including use of red ochre and fishing.
[25]
150 ka Mitochondrial Eve is a woman that lived in East Africa. She is the statistically
expected most recent female ancestor common to all mitochondrial lineages in humans
alive today. Note that there is no evidence of any characteristic or genetic drift that
significantly differentiated her from the contemporary social group she lived with at the time.
Her ancestors were homo sapiens and her mother had the same mtDNA.
70 ka Appearance of mitochondrial haplogroup L2. Behavioral modernity according to the

"great leap forward" theory.[26]
60 ka Y-chromosomal Adam lives in Africa. He is the most recent common ancestor from
whom all male human Y chromosomes are descended. Appearance of mitochondrial
haplogroups M and N, which participate in the migration out of Africa. Homo sapiens that
leave Africa in this wave start interbreeding with the Neanderthals they encounter.[27][28]
50 ka Migration to South Asia. M168 mutation (carried by all non-African males).

Beginning of the Upper Paleolithic. mt-haplogroups U, K.
40 ka Migration to Australia[29] and Europe (Cro-Magnon).
25 ka The independent Neanderthal lineage dies out. Y-Haplogroup R2; mt-haplogroups

J, X.
12 ka Beginning of the Mesolithic / Holocene. Y-Haplogroup R1a; mt-haplogroups V, T.

Evolution of light skin in Europeans (SLC24A5). Homo floresiensis dies out, leaving Homo
sapiens as the only living species of the genus Homo.
[edit] See also

• Timeline of prehistory
• Graphical timeline of our universe
• History of Earth
• Natural history
• History of the world
• Human evolution
• Human taxonomy
• Homo (genus)
• Most recent common ancestor
• List of human evolution fossils
• Prehistoric amphibian
• Prehistoric fish
• Prehistoric reptile
• The Ancestor's Tale by Richard Dawkins with a timeline comprising 40 rendezvous points
• Timeline of evolution - an explanation of the evolution of a wide variety of animals living
today
• Y-DNA haplogroups by ethnic groups
[edit] References
1. ^ "'Experiments with sex have been very hard to conduct,' Goddard said. 'In an experiment,
one needs to hold all else constant, apart from the aspect of interest. This means that no higher
organisms can be used, since they have to have sex to reproduce and therefore provide no asexual
control.'
Goddard and colleagues instead turned to a single-celled organism, yeast, to test the idea that sex
allows populations to adapt to new conditions more rapidly than asexual populations." Sex Speeds
Up Evolution, Study Finds (URL accessed on January 9, 2005)
2. ^ "Proterospongia is a rare freshwater protist, a colonial member of the Choanoflagellata."
"Proterospongia itself is not the ancestor of sponges. However, it serves as a useful model for what
the ancestor of sponges and other metazoans may have been like."
http://www.ucmp.berkeley.edu/protista/proterospongia.html Berkeley University
3. ^ "Obviously vertebrates must have had ancestors living in the Cambrian, but they were
assumed to be invertebrate forerunners of the true vertebrates — protochordates. Pikaia has been
heavily promoted as the oldest fossil protochordate." Richard Dawkins 2004 The Ancestor's Tale
Page 289, ISBN 0-618-00583-8
4. ^ Shu, D. G.; Luo, H. L.; Conway Morris, S.; Zhang, X. L.; Hu, S. X.; Chen, L.; Han, J.; Zhu,
M. et al. (1999). Nature 402: 42. doi:10.1038/46965. edit
5. ^ Chen, J. Y.; Huang, D. Y.; Li, C. W. (1999). Nature 402: 518. doi:10.1038/990080. edit
6. ^ Shu, D. G.; Morris, S. C.; Han, J.; Zhang, Z. F.; Yasui, K.; Janvier, P.; Chen, L.; Zhang, X.
L. et al. (Jan 2003), "Head and backbone of the Early Cambrian vertebrate Haikouichthys", Nature
421 (6922): 526–529, doi:10.1038/nature01264, ISSN 0028-0836, PMID 12556891,
http://adsabs.harvard.edu/abs/2003Natur.421..526S edit
7. ^ These first vertebrates lacked jaws, like the living hagfish and lampreys. Jawed vertebrates
appeared 100 million years later, in the Silurian.
http://www.ucmp.berkeley.edu/vertebrates/vertintro.html Berkeley University
8. ^ "Bones of first gill arch became upper and lower jaws. " (Image) (URL accessed on
November 16, 2006)
9. ^ A fossil coelacanth jaw found in a stratum datable 410 mya that was collected near Buchan
in Victoria, Australia's East Gippsland, currently holds the record for oldest coelacanth; it was given
the name Eoactinistia foreyi when it was published in September 2006. [1]
10.^ "Lungfish are believed to be the closest living relatives of the tetrapods, and share a
number of important characteristics with them. Among these characters are tooth enamel, separation
of pulmonary blood flow from body blood flow, arrangement of the skull bones, and the presence of
four similarly sized limbs with the same position and structure as the four tetrapod legs."
http://www.ucmp.berkeley.edu/vertebrates/sarco/dipnoi.html Berkeley University
11.^ "the ancestor that amphibians share with reptiles and ourselves? " " These possibly
transitional fossils have been much studied, among them Acanthostega, which seems to have been
wholly aquatic, and Ichthyostega" Richard Dawkins 2004 The Ancestor's Tale page 250, ISBN 0-618-
00583-8
12.^ Eckhart L, Valle LD, Jaeger K, et al. (November 2008). "Identification of reptilian genes
encoding hair keratin-like proteins suggests a new scenario for the evolutionary origin of hair".
23. doi:10.1073/pnas.0805154105. PMID 19001262.
13.^ "In many respects, the pelycosaurs are intermediate between the reptiles and mammals"
http://www.ucmp.berkeley.edu/synapsids/pelycosaurs.html Berkeley University
14.^ "Tlvinaxodon, like any fossil, should be thought of as a cousin of our ancestor, not the
ancestor itself. It was a member of a group of mammal-like reptiles called the cynodonts. The
cynodonts were so mammal-like, it is tempting to call them mammals. But who cares what we call
them? They are almost perfect intermediates." Richard Dawkins 2004 The Ancestor's Tale page 211,
ISBN 0-618-00583-8
15.^ "Fossils that might help us reconstruct what Concestor 8 was like include the large group
called plesiadapi-forms. They lived about the right time, and they have many of the qualities you
would expect of the grand ancestor of all the primates" Richard Dawkins 2004 The Ancestor's Tale
page 136, ISBN 0-618-00583-8
16.^ Raauma, Ryan, Sternera, K., (2005) "Catarrhine primate divergence dates estimated from
complete mitochondrial genomes", Journal of Human Evolution 48: 237-257 [2]
17.^ Perlman, David (July 12, 2001). "Fossils From Ethiopia May Be Earliest Human Ancestor".
National Geographic News.
http://news.nationalgeographic.com/news/2001/07/0712_ethiopianbones.html. Retrieved July 2009.
"Another co-author is Tim D. White, a paleoanthropologist at UC-Berkeley who in 1994 discovered a
pre-human fossil, named Ardipithecus ramidus, that was then the oldest known, at 4.4 million years."
18.^ White, Tim D.; Asfaw, Berhane; Beyene, Yonas; Haile-Selassie, Yohannes; Lovejoy, C.
Owen; Suwa, Gen; WoldeGabriel, Giday (2009). "Ardipithecus ramidus and the Paleobiology of Early
Hominids.". Science 326 (5949): 75–86. doi:10.1126/science.1175802. PMID 19810190.
19.^ NOVA: Becoming Human Part 2 http://video.pbs.org/video/1319997127/
20.^ Green, R. E., Krause, J, Ptak, S. E., Briggs, A. W., Ronan, M. T., Simons, J. F., et al.
(2006) Analysis of one million base pairs of Neanderthal DNA. Nature, 16, 330–336.
http://www.nature.com/nature/journal/v444/n7117/abs/nature05336.html
21.^ "Rubin also said analysis so far suggests human and Neanderthal DNA are some 99.5
percent to nearly 99.9 percent identical." Neanderthal bone gives DNA clues (URL accessed on
November 16, 2006)
22.^ "The conclusion is the old saw that we share 98.5% of our DNA sequence with chimpanzee
is probably in error. For this sample, a better estimate would be that 95% of the base pairs are
exactly shared between chimpanzee and human DNA." Britten, R.J. (2002). "Divergence between
samples of chimpanzee and human DNA sequences is 5%, counting indels". PNAS 99 (21): 13633–5.
doi:10.1073/pnas.172510699. PMID 12368483.
23.^ "...of the three billion letters that make up the human genome, only 15 million--less than 1
percent--have changed in the six million years or so since the human and chimp lineages diverged. "
Pollard, K.S. (2009), "What makes us human?", Scientific American 300-5: 44–49.
24.^ Krause J, Lalueza-Fox C, Orlando L, Enard W, Green RE, Burbano HA, Hublin JJ, Hänni C,
Fortea J, de la Rasilla M, Bertranpetit J, Rosas A, Pääbo S (November 2007). "The derived FOXP2
variant of modern humans was shared with Neandertals". Curr. Biol. 17 (21): 1908–12.
doi:10.1016/j.cub.2007.10.008. PMID 17949978. Lay summary – New York Times (2007-10-19).
25.^ "Schwarz, J". Uwnews.org. 2007-10-17. http://uwnews.org/article.asp?articleID=37362.
26.^ Diamond, Jared (1992). The Third Chimpanzee. Harper Perennial. pp. 47–57. ISBN 978-0-
060-98403-8.
27.^ Richard E. Green et al (2010). "A Draft Sequence of the Neandertal Genome". Science 328
(5979): 710–722. doi:10.1126/science.1188021. PMID 20448178.
28.^ Rincon, Paul (2010-05-06). "Neanderthal genes 'survive in us'". BBC News (BBC).
29.^ Bowler JM, Johnston H, Olley JM, Prescott JR, Roberts RG, Shawcross W, Spooner NA.
(2003). "New ages for human occupation and climatic change at Lake Mungo, Australia.". Nature 421
(6925): 837–40. doi:10.1038/nature01383. PMID 1259451.

• Palaeos
• Berkeley Evolution
• Tree of Life Web Project - explore complete phylogenetic tree interactively
• History of Animal Evolution
• Human Evolution Timeline Interactive by The Smithsonian Institution's Human Origins
Program
Hominini
Australopithecines

Processes of
Population genetic
Evolutionary
developmental
concepts

Modes of

Retrieved from "http://en.wikipedia.org/wiki/Timeline_of_human_evolution"
Categories: Biology timelines | Human evolution

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Microbial phylogenetics

Microbial phylogenetics is the study of the evolutionary relatedness among various groups of
microorganisms. The molecular approach to microbial phylogenetic analysis, pioneered by Carl Woese in the
1970s and leading to the three-domain model (Archaea, Bacteria, Eucaryota), revolutionized our thinking
about evolution in the microbial world. Phylogenetic analysis plays a central role in microbiology and the
emerging fields of comparative genomics and phylogenomics require substantial knowledge and
understanding of phylogenetic analysis and computational methods.[1][2]
Contents
[hide]
• 1 Historical overview
• 2 Methods and programs
• 3 Multilocus sequence analysis
• 4 rRNA and other global markers
• 5 The phyla of prokaryotes
• 6 Horizontal gene transfer
• 7 See also
• 8 References
[edit] Historical overview

When at the end of the 19th century information began to accumulate about the diversity within the
bacterial world, scientists started to include the bacteria in phylogenetic schemes to explain how life on Earth
may have developed. Some of the early phylogenetic trees of the prokaryote world were morphology-based;
others were based on the then-current ideas on the presumed conditions on our planet at the time that life
first developed. Around 1950 many leading microbiologists had become pessimistic with respect to the
possibility of ever reconstructing bacterial phylogeny. The concept of the prokaryote-eukaryote dichotomy did
little to clarify phylogenetic relationships. The developing technology of nucleic acid sequencing, together
with the recognition that sequences of building blocks in informational macromolecules (nucleic acids,
proteins) can be used as 'molecular clocks' that contain historical information, led to the development of the
three-domain model (Archaea - Bacteria - Eucaryota) in the late 1970s, primarily based on small subunit
ribosomal RNA sequence comparisons. The information currently accumulating from complete genome
sequences of an ever increasing number of prokaryotes are now leading to further modifications of our views
on microbial phylogeny.[1]
[edit] Methods and programs

The purpose of phylogenetic analysis is to understand the past evolutionary path of organisms. Even
though we will never know for certain the true phylogeny of any organism, phylogenetic analysis provides
best assumptions, thereby providing a framework for various disciplines in microbiology. Due to the
technological innovation of modern molecular biology and the rapid advancement in computational science,
accurate inference of the phylogeny of a gene or organism seems possible in the near future. There has
been a flood of nucleic acid sequence information, bioinformatic tools and phylogenetic inference methods in
public domain databases, literature and World Wide Web space. Phylogenetic analysis has long played a
central role in basic microbiology, for example in taxonomy and ecology. In addition, more recently emerging
fields of microbiology, including comparative genomics and phylogenomics, require substantial knowledge
and understanding of phylogenetic analysis and computational skills to handle the large-scale data involved.
Methods of phylogenetic analysis and relevant computer software tools lend accuracy, efficiency and
availability to the task.
There are four steps in general phylogenetic analysis of molecular sequences: (i) selection of a
suitable molecule or molecules (phylogenetic marker), (ii) acquisition of molecular sequences, (iii) multiple
sequence alignment (MSA) and (iv) phylogenetic treeing and evaluation. The first step of phylogenetic
analysis is to choose a suitable homologous part of the genomes to be compared. Mechanisms of molecular
evolution include mutations, duplication of genes, reorganization of genomes, and genetic exchanges such
as recombination, reassortment and lateral gene transfer. Although all of this information can be used to infer
phylogenetic relationships of genes or organisms, information on mutations, including substitution, insertion,
and deletion, is most frequently used in phylogeny reconstruction. The aim is to infer a correct organismal
phylogeny, using orthologous genetic loci, in which common ancestry of two sequences can be traced back
to a speciation event. Phylogeny using homologous genetic loci derived by gene duplication (paralogy) or
related through lateral gene transfer (xenology), cannot reflect evolutionary history of organisms.
Once DNA sequence data are generated, they are subjected to a multiple sequence alignment
process. This involves finding homologous sites, that is, positions derived from the same ancestral organism
in the molecules under study. A set of sequences can be aligned with another by introducing "alignment
gaps" (known in brief as "gaps"). In general, multiple sequence alignment starts by aligning a pair of
sequences (pairwise alignment), and is then expanded to multiple sequences using various algorithms.
Many algorithms and computer programs have been developed in the last few decades for multiple
sequence alignment, but the original Clustal series programs are still most widely used and produce
reasonably good quality MSA for small data sets. For a large dataset, such as massive pyrosequencing
reads, the MUSCLE program can generate good compromise between accuracy and speed. The MAFFT
program utilizes several different algorithmic approaches and can be used for either small or very large
datasets. There are also other computer programs developed for general multiple sequence alignment, but
the above three have been most popular and are routinely used in publications in various microbiological
disciplines.[1]
[edit] Multilocus sequence analysis
Multilocus sequence analysis (MLSA) represents the novel standard in microbial molecular
systematics. In this context, MLSA is implemented in a relatively straightforward way, consisting essentially
in the concatenation of several sequence partitions for the same set of organisms, resulting in a
"supermatrix" which is used to infer a phylogeny by means of distance-matrix or optimality criterion-based
methods. This approach is expected to have an increased resolving power due to the large number of
characters analyzed, and a lower sensitivity to the impact of conflicting signals (i.e. phylogenetic
incongruence) that result from eventual horizontal gene transfer events. The strategies used to deal with
multiple partitions can be grouped in three broad categories: the total evidence, separate analysis and
combination approaches. The concatenation approach that dominates MLSAs in the microbial molecular
systematics literature is known to systematists working with plants and animals as the "total molecular
evidence" approach, and has been used to solve difficult phylogenetic questions such as the relationships
among the major groups of cetaceans, that of microsporidia and fungi, or the phylogeny of major plant
lineages. The total molecular evidence approach has been criticized because by directly concatenating all
available sequence alignments, the evidence of conflicting phylogenetic signals in the different data partitions
is lost along with the possibility to uncover the evolutionary processes that gave rise to such contradictory
signals. The nature of these conflicts is varied, but in the microbial world the strongest conflicting signals
often derive from the existence of horizontal gene transfer events in the dataset. If the individuals containing
xenologous loci are not identified and removed from the supermatrix prior to phylogeny inference, the
resulting hypothesis may be strongly distorted, since standard treeing methods assume a single underlying
evolutionary history. Based on these arguments, the conditional data combination strategy is to be generally
preferred in bacterial MLSA.
[edit] rRNA and other global markers
The introduction of comparative rRNA sequence analysis represents a major milestone in the history
of microbiology. The current taxonomy of prokaryotes as well as modern probe and chip based identification
methods are mainly based upon rRNA derived phylogenetic conclusions. Also of importance is single gene
based phylogenetic inference and alternative global markers include elongation and initiation factors, RNA
polymerase subunits, DNA gyrases, heat shock and recA proteins. Although the comparative analyses are
hampered by the generally low phylogenetic information content, and different resolution power, and multiple
copies of the individual markers, the domain and prokaryotic phyla concept is globally supported.
The conserved inserts or deletions (indels) in protein sequences provide particularly useful means for
identifying different groups of microbes in clear molecular terms and for understanding how they have
branched off from a common ancestor. Conserved indels and lineage-specific proteins can be useful for
understanding microbial phylogeny at different phylogenetic depths.
[edit] The phyla of prokaryotes

There is no official classification of prokaryotes. For the higher taxa there even is no official
nomenclature: the rules of the International Code of Nomenclature of Prokaryotes do not cover taxa above
the rank of class. The most commonly accepted division of the prokaryotes in two "subkingdoms" or
"domains" (Bacteria and Archaea) and the classification of their species with validly published names in
respectively 27 and 2 "phyla" or "divisions" (as of November 2009) is primarily based on 16S rRNA sequence
comparisons. This type of classification was adopted in the latest edition of Bergey's Manual of Systematic
Bacteriology. Alternative classifications have been proposed as well, based e.g. on the structure of the cell
wall. Some 16S rRNA sequence-based phyla unite prokaryotes of similar physiological properties (for
example Cyanobacteria, Chlorobi, Thermotogae); others (Euryarchaeota, Proteobacteria, Flavobacteria)
contain organisms with highly disparate lifestyles. Some phyla based on deep 16S rRNA lineages are
currently represented by one or a few species only. Environmental genomics/metagenomics approaches
suggest existence of many more phyla based on the deep lineages of 16S rRNA gene sequences recovered.
[3] To obtain the organisms harboring these sequences and to study their properties is a major challenge of
microbiology today.
[edit] Horizontal gene transfer

Efforts to construct the tree of life take their conceptual motivation from Charles Darwin's theory of
evolution. Until the advent of molecular biology, however, a universal tree of life was well beyond the scope
of the data and methods of traditional organismal phylogeny. The rapid development of these methods and
bodies of genetic sequence from the 1970s onwards resulted in major reclassifications of life and revived
ambitions to represent all organismal lineages by one true tree of life. Subsequent realization of the
significance of lateral gene transfer and other non-vertical processes has subtly reconceptualized and
reoriented attempts to construct this universal phylogeny.
Horizontal gene transfer has affected the formation of groups of organisms. Gene transfer can make
it more difficult to define and determine relationships. In those cases where many genes have been
transferred between preferred partners, the majority of genes in a genome may reflect gene acquisition, and
as a consequence, if a coherent signal is detected, one nevertheless might not be sure that the signal is due
to organismal shared ancestry. However, the presence of a particular transferred gene has been shown, in
several cases, to constitute a shared derived character useful in classification. Gene transfer can put
together new metabolic pathways that open up new ecological niches, and consequently, the transfer of an
adaptive gene might create a new group of organisms.[1]
[edit] See also
• Phylogenetics
• Molecular phylogenetics
• Computational phylogenetics
• History of molecular evolution
[edit] References
1. ^ a b c d Oren, A; Papke, RT (editor) (2010). Molecular Phylogeny of Microorganisms. Caister
Academic Press. ISBN 978-1-904455-67-7.
2. ^ Blum, P (editor) (2010). Archaea: New Models for Prokaryotic Biology . Caister Academic
Press. ISBN 978-1-904455-27-1.
3. ^ Marco, D (editor) (2010). Metagenomics: Theory, Methods and Applications. Caister
Academic Press. ISBN 978-1-904455-54-7.

• Microbial phylogenetics
Retrieved from "http://en.wikipedia.org/wiki/Microbial_phylogenetics"
Categories: Phylogenetics
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Evolution of mammals

Restoration of Thrinaxodon, a member of the cynodont group, which includes the ancestors of
mammals.
The evolution of mammals within the synapsid lineage (mammal-like-reptiles) was a gradual process
that took approximately 70 million years, beginning in the mid-Permian. By the mid-Triassic, there were many
species that looked like mammals, and the first true mammals appeared in the early Jurassic. The earliest
known marsupial, Sinodelphys, appeared 125 million years ago in the early Cretaceous, around the same
time as Eomaia, the first known eutherian (member of placentals' "parent" group); and the earliest known
monotreme, Teinolophos, appeared two million years later. After the Cretaceous-Tertiary extinction wiped out
the non-avian dinosaurs (birds are generally regarded as the surviving dinosaurs) and several other
mammalian groups, placental and marsupial mammals diversified into many new forms and ecological
niches throughout the Tertiary, by the end of which all modern orders had appeared.
From the point of view of phylogenetic nomenclature, mammals are the only surviving synapsids.
The synapsid lineage became distinct from the sauropsid ("reptile") lineage in the late Carboniferous period,
between 320 and 315 million years ago,[1] and were the most common and largest land vertebrates of the
Permian period.[2] But in the Triassic period a previously obscure group of sauropsids, the archosaurs,
became the dominant vertebrates and one archosaur group, the dinosaurs, dominated the rest of the
Mesozoic era. These changes forced the Mesozoic mammaliforms ("nearly mammals") into nocturnal niches,
and may have contributed greatly to the development of mammalian traits such as endothermy, hair and a
large brain. Later in the Mesozoic mammals spread into other ecological niches, for example aquatic, gliding
and even preying on dinosaurs.
Most of the evidence consists of fossils. For many years fossils of Mesozoic mammals and their
immediate ancestors were very rare and fragmentary, but since the mid 1990s there have been many
important new finds, especially in China. The relatively new techniques of molecular phylogenetics have also
shed light on some aspects of mammalian evolution by estimating the timing of important divergence points
for modern species. When used carefully, these techniques often, but not always, agree with the fossil
record.
Although mammary glands are a signature feature of modern mammals, little is known about the
evolution of lactation, and virtually nothing is known about the evolution of another distinctive feature, the
neocortex region of the brain. This is because these soft tissues are not often preserved in the fossil record.
Hence, most study of the evolution of mammals centers around the development of the middle ear bones
from components of the ancestral amniote jaw joint. Other much-studied aspects include the evolution of
erect limb posture, a bony secondary palate, fur and hair, and warm-bloodedness.
Contents
[hide]
• 1 Definition of "mammal"
• 2 The ancestry of mammals
• 2.1 Amniotes
• 2.2 Synapsids
• 3 Therapsids
• 3.1 Therapsid family tree
• 3.2 Biarmosuchia
• 3.3 Dinocephalians
• 3.4 Anomodonts
• 3.5 Theriodonts
• 3.6 Cynodonts
• 4 Triassic takeover
• 5 From cynodonts to true mammals
• 5.1 Many uncertainties
• 5.2 Mammals or mammaliformes?
• 5.3 Family tree — cynodonts to mammals
• 5.4 Multituberculates
• 5.5 Morganucodontidae
• 5.6 Docodonts
• 5.7 Hadrocodium
[edit] Definition of "mammal"
Mammalian and non-mammalian jaws. In the mammal configuration, the quadrate and articular
bones are much smaller and form part of the middle ear. Note that in mammals the lower jaw consists of only
the dentary bone.
Living mammal species can be identified by the presence of milk-producing mammary glands in
females. Other features are required when classifying fossils, since mammary glands and other soft-tissue
features are not visible in fossils.
Paleontologists therefore use a distinguishing feature that is shared by all living mammals (including
monotremes) but is not present in any of the early Triassic therapsids: Mammals use two bones for hearing
that all other amniotes use for eating. The earliest amniotes had a jaw joint composed of the articular (a small
bone at the back of the lower jaw) and the quadrate (a small bone at the back of the upper jaw). All non-
mammalian amniotes use this system including lizards, crocodilians, dinosaurs (and their descendants the
birds), and therapsids. But mammals have a different jaw joint, composed only of the dentary (the lower jaw
bone, which carries the teeth) and the squamosal (another small skull bone). In mammals, the quadrate and
articular bones have become the incus and malleus bones in the middle ear.[3][4]
Mammals also have a double occipital condyle; they have two knobs at the base of the skull that fit
into the topmost neck vertebra, and other vertebrates have a single occipital condyle.[3] But paleontologists
use only the jaw joint and middle ear as criteria for identifying fossil mammals, as it would be confusing if
they found a fossil that had one feature but not the other (e.g. a mammalian jaw and ear but a non-
mammalian single occipital condyle).
Due to the incremental changes in transitional fossils, it has been said
We may again ask the question, What is a mammal? Where we draw the line between reptile
and mammal has no biological significance. It is purely a matter of convenience. There are two
obvious choices, both immediately following a period of rapid evolution that make as definite a
break as we can hope to find.[5]
[edit] The ancestry of mammals

Tetrapods
Amphibians
Amniotes
Sauropsids (including
dinosaurs)
Synapsids
Pelycosaurs
Therapsids
Mammals
[edit] Amniotes
The first fully terrestrial vertebrates were amniotes — their eggs had internal membranes that allowed
the developing embryo to breathe but kept water in. This allowed amniotes to lay eggs on dry land, while
amphibians generally need to lay their eggs in water (a few amphibians, such as the Surinam toad, have
evolved other ways of getting round this limitation). The first amniotes apparently arose in the late
Carboniferous from the ancestral reptiliomorphs.
Within a few million years two important amniote lineages became distinct: mammals' synapsid
ancestors and the sauropsids, from which lizards, snakes, crocodilians, dinosaurs and birds are descended.
[1] The earliest known fossils of synapsids and sauropsids (such as Archaeothyris and Hylonomus resp.)
date from about 320 to 315 million years ago. Unfortunately it is difficult to be sure about when each of them
evolved, since vertebrate fossils from the late Carboniferous are very rare, and therefore the actual first
occurrences of each of these types of animal might have been considerably earlier.[6]
[edit] Synapsids
The original synapsid skull structure has one hole behind each eye, in a fairly low position on the
skull (lower right in this image).
Synapsid skulls are identified by the distinctive pattern of the holes behind each eye, which served
the following purposes:
• made the skull lighter without sacrificing strength.
• saved energy by using less bone.
• probably provided attachment points for jaw muscles. Having attachment points further away
from the jaw made it possible for the muscles to be longer and therefore to exert a strong pull over a
wide range of jaw movement without being stretched or contracted beyond their optimum range.
Early Permian terrestrial fossils indicate that one synapsid group, the pelycosaurs, were the most
common land vertebrates of their time and included the largest land animals of the time.[2]
[edit] Therapsids
Therapsids descended from pelycosaurs in the middle Permian and took over their position as the
dominant land vertebrates. They differ from pelycosaurs in several features of the skull and jaws, including
larger temporal fenestrae and incisors that are equal in size.[7]
The therapsids went through a series of stages, beginning with animals that were very like their
pelycosaur ancestors and ending with some that could easily be mistaken for mammals:[8]
• gradual development of a bony secondary palate. Most books and articles interpret this as a
prequisite for the evolution of mammals' high metabolic rate, because it enabled these animals to eat
and breathe at the same time. But some scientists point out that some modern ectotherms use a
fleshy secondary palate to separate the mouth from the airway, and that a bony palate provides a
surface on which the tongue can manipulate food, facilitating chewing rather than breathing.[9] The
interpretation of the bony secondary palate as an aid to chewing also suggests the development of a
faster metabolism, since chewing makes it possible to digest food more quickly. In mammals the
palate is formed by two specific bones, but various Permian therapsids had other combinations of
bones in the right places to function as a palate.
• the dentary gradually becomes the main bone of the lower jaw.
• progress towards an erect limb posture, which would increase the animals' stamina by
avoiding Carrier's constraint. But this process was erratic and very slow — for example: all
herbivorous therapsids retained sprawling limbs (some late forms may have had semi-erect hind
limbs); Permian carnivorous therapsids had sprawling forelimbs, and some late Permian ones also
had semi-sprawling hindlimbs. In fact, modern monotremes still have semi-sprawling limbs.
• in the Triassic, progress towards the mammalian jaw and middle ear.
• there is plausible evidence of hair in Triassic therapsids, but none for Permian therapsids
(see below).
• some scientists have argued that some Triassic therapsids show signs of lactation (see
below).
[edit] Therapsid family tree

(simplified from;[7] only those that are most relevant to the evolution of mammals are described
below)
Therapsids
Eutherapsida
Dinocephalia
Neotherapsida
Anomodonts
Dicynodonts
Theriodontia
Gorgonopsia
Eutheriodontia
Therocephalia
Cynodontia
(Mammals,
eventually)
Only the dicynodonts, therocephalians and cynodonts survived into the Triassic.
[edit] Biarmosuchia
The Biarmosuchia were the most primitive and pelycosaur-like of the therapsids.
[edit] Dinocephalians
Dinocephalians ("terrible heads") were large, some as large as a rhinoceros, and included both
carnivores and herbivores. Some of the carnivores had semi-erect hindlimbs, but all dinocephalians had
sprawling forelimbs. In many ways they were very primitive therapsids, for example they had no secondary
palate and their jaws were rather "reptilian".[10]
[edit] Anomodonts
Lystrosaurus, one of the few species of dicynodonts that survived the Permian-Triassic extinction
event
The anomodonts ("anomalous teeth") were the most successful of the herbivorous therapsids — one
sub-group, the dicynodonts, survived almost to the end of the Triassic. But anomodonts were very different
from modern herbivorous mammals, as their only teeth were a pair of fangs in the upper jaw and it is
generally agreed that they had beaks like those of birds or ceratopsians.[11]
[edit] Theriodonts
The theriodonts ("beast teeth") and their descendants had jaw joints in which the lower jaw's articular
bone tightly gripped the skull's very small quadrate bone. This allowed a much wider gape, and one group,
the carnivorous gorgonopsians ("gorgon faces"), took advantage of this to develop "sabre teeth". But the
theriodont's jaw hinge had a longer term significance — the much reduced size of the quadrate bone was an
important step in the development of the mammalian jaw joint and middle ear.
The gorgonopsians still had some primitive features: no bony secondary palate (but other bones in
the right places to perform the same functions); sprawling forelimbs; hindlimbs that could operate in both
sprawling and erect postures. But the therocephalians ("beast heads"), which appear to have arisen at about
the same time as the gorgonopsians, had additional mammal-like features, e.g. their finger and toe bones
had the same number of phalanges (segments) as in early mammals (and the same number that primates
have, including humans).[12]
[edit] Cynodonts
Artist's conception of the cynodont Trirachodon within a burrow

The cynodonts, a theriodont group that also arose in the late Permian, include the ancestors of all
mammals — one sub-group, the trithelodonts, is widely regarded as the most likely to contain mammals'
ancestor. Cynodonts' mammal-like features include further reduction in the number of bones in the lower jaw;
a secondary bony palate; cheek teeth with a complex pattern in the crowns; the brain filled the endocranial
cavity.[13]
Multi-chambered burrows have been found, containing as many as 20 skeletons of the Early Triassic
cynodont Trirachodon; the animals are thought to have been drowned by a flash flood. The extensive shared
burrows indicate that these animals were capable of complex social behaviors.[14]
[edit] Triassic takeover

The catastrophic Permian-Triassic mass extinction killed off about 70 percent of terrestrial vertebrate
species, and the majority of land plants. As a result[15]
• Ecosystems and food chains collapsed, and the recovery took about 6 million years.
• The survivors had to re-start the struggle for dominance of their former ecological niches —
even the cynodonts, which had seemed on the way to dominance at the end of the Permian.
But the cynodonts lost out to a previously obscure group of sauropsids, the archosaurs (which
include the ancestors of crocodilians, dinosaurs and birds). This reversal of fortunes is often called the
"Triassic takeover". Several explanations have been offered for it, but the most likely is that the early Triassic
was predominantly arid and therefore archosaurs' superior water conservation gave them a decisive
advantage (all known sauropsids have glandless skins and excrete uric acid, which requires less water to
keep it sufficiently liquid than urea, which marsupial and placental mammals excrete and presumably
therapsids excreted).[8][16] The Triassic takeover was gradual — in the earliest part of the Triassic cynodonts
were the main predators and lystrosaurs were the main herbivores, but by the mid-Triassic archosaurs
dominated all the large carnivore and herbivore niches.
But the Triassic takeover may have been a vital factor in the evolution of cynodonts into mammals.
The cynodonts' descendants were only able to survive as small, mainly nocturnal insectivores.[13] As a
result:
• The therapsid trend towards differentiated teeth with precise occlusion accelerated, because
of the need to hold captured arthropods and crush their exoskeletons.
• Nocturnal life required advances in thermal insulation and temperature regulation to enable
the ancestors of mammals to be active in the cool of the night.[17]
• Acute senses of hearing and smell became vital.
• This accelerated the development of the mammalian middle ear, and therefore of the
mammalian jaw since bones that had been part of the jaw joint became part of the middle
ear.
• The increase in the size of the olfactory and auditory lobes of the brain increased
brain weight as a total percentage of body weight. Brain tissue requires a disproportionate
amount of energy.[18][19] The need for more food to support the enlarged brains increased
the pressures for improvements in insulation, temperature regulation and feeding.
• As a side-effect of the nocturnal life, discerning colors became less important (they lost two
out of four opsins), and this is reflected in the fact that most mammals have poor color vision,
including the "lower primates" such as lemurs.[20]
[edit] From cynodonts to true mammals

[edit] Many uncertainties
While the Triassic takeover probably accelerated the evolution of mammals, it made life more difficult
for paleontologists because good fossils of the nearly-mammals are extremely rare, mainly because they
were mostly smaller than rats:
• They were largely restricted to environments that are less likely to provide good fossils. The
best terrestrial environments for fossilization are floodplains, where seasonal floods quickly cover
dead animals in a protective layer of silt that is later compressed into sedimentary rock. But
floodplains are dominated by medium to large animals, and the Triassic therapsids and near-
mammals could not compete with archosaurs in the medium to large size range.
• Their delicate bones were vulnerable to being destroyed before they could be fossilized — by
scavengers (including fungi and bacteria) and by being trodden on.
• Small fossils are harder to spot and more vulnerable to being destroyed by weathering and
other natural stresses before they are discovered.
In fact it was said as recently as the 1980s that all the Mesozoic fossils of mammals and near-
mammals could be contained in a few shoeboxes — and they were mostly teeth, which are the most durable
of all tissues.[21] Since then, the number of Mesozoic fossil mammals has increased, from 116 genera
known in 1979 to about 310 in 2007, with an increase in quality such that "at least 18 Mesozoic mammals are
represented by nearly complete skeletons".[22]
As a result:
• In many cases it is difficult to assign a Mesozoic mammal or near-mammal fossil to a genus.
• All the available fossils of a genus seldom add up to a complete skeleton, and hence it is
difficult to decide which genera are most like each other and therefore most likely to be closely-
related. In other words, it becomes very difficult to classify them by means of cladistics, which is the
most reliable and least subjective method currently available.
So the evolution of mammals in the Mesozoic is full of uncertainties, although there is no room for
doubt that true mammals did first appear in the Mesozoic.
[edit] Mammals or mammaliformes?
One result of these uncertainties has been a change in the paleontologists' definition of "mammal".
For a long time a fossil was considered a mammal if it met the jaw-ear criterion (the jaw joint consists only of
the squamosal and dentary; and the articular and the quadrate bones have become the middle ear's malleus
and incus). But more recently some paleontologists have usually defined "mammal" as the crown group
mammals, i.e. the last common ancestor of monotremes, marsupials and placentals and all of its
descendants. The need to address the animals that are more mammal-like than cynodonts, but less closely
related to monotremes, marsupials and placentals, lead to erecting the group mammaliformes to
accommodate these primitive forms. Mammaliformes is a paraphyletic taxon, representing the early radiation
of mammals after the jaw-ear criterion.[23] Although this now appears to be the majority approach, some
paleontologists have resisted it because it simply moves most of the problems into the new taxon (a
paraphyletic one at that) without solving the original problem; the Mammaliformes includes some animals
with "mammalian" jaw joints and some with "reptilian" (articular-to-quadrate) jaw joints; and the newer
definition of "mammal" and "mammaliformes" depend on last common ancestors of both groups, which have
not yet been found.[21] Despite these objections, this article follows the majority approach and treats most of
the cynodonts' Mesozoic descendants as mammaliformes.
[edit] Family tree — cynodonts to mammals

(based on Mammaliformes - Palaeos)
Cynodontia
Dvinia
Procynosuchidae
Epicynodontia
Eucynodontia
Probainognathia
Trithelodontidae
Chiniquodontidae
Mammaliaformes
Allotheria
Multituberc
Morganucodontidae
Docodonta
Hadrocodium
Symmetrodonta
Kuehneotheriidae
crown group
Mammals (all
descendants of the last
common ancestor of all
living mammals)
[edit] Multituberculates
Skull of the multituberculate Ptilodus

Multituberculates (named for the multiple tubercles on their "molars") are often called the "rodents of
the Mesozoic" but this is an example of convergent evolution rather than meaning that they are closely
related to the Rodentia. At first sight they look like mammals: their jaw joints consists of only the dentary and
squamosal bones, and the quadrate and articular bones are part of the middle ear; their teeth are
differentiated, occlude and have mammal-like cusps; they have a zygomatic arch; the structure of the pelvis
suggests that they gave birth to tiny helpless young, like modern marsupials. And they lived for over 120
million years (from mid Jurassic, about 160M years ago, to early Oligocene, about 35M years ago), which in
terms of clade longevity would make them the most successful mammaliformes ever. But a closer look
shows that they are very different from modern mammals:[23]
• Their "molars" have two parallel rows of tubercles, unlike the tribosphenic (three-peaked)
molars of early mammals.
• The chewing action is completely different. Mammals chew with a side-to-side grinding
action, which means that usually the molars occlude on only one side at a time. Multituberculates'
jaws were incapable of side-to-side movement and chewed by dragging the lower teeth backwards
against the upper ones as the jaw closed.
• The anterior (forward) part of the zygomatic arch mostly consists of the maxilla (upper
jawbone) rather than the jugal, and the jugal is a small bone in a little slot in the maxillary process
(extension).
• The squamosal does not form part of the braincase.
• The rostrum (snout) is unlike that of mammals, in fact it looks more like that of a pelycosaur
such as Dimetrodon. The multituberculate rostrum is box-like, with the large flat maxillae forming the
sides, the nasal the top, and the tall premaxilla at the front.
Morganucodontidae and other transitional forms had both types of jaw joint: dentary-squamosal
(front) and articular-quadrate (rear).
[edit] Morganucodontidae
The Morganucodontidae first appeared in the late Triassic, about 205M years ago. They are an
excellent example of transitional fossils, since they have both the dentary-squamosal and articular-quadrate
jaw joints.[24] They were also one of the first discovered and most thoroughly studied of the mammaliformes,
since an unusually large number of morganucodont fossils have been found.
[edit] Docodonts
Reconstruction of Castorocauda. Note the fur and the adaptations for swimming (broad, flat tail;
webbed feet) and for digging (robust limbs and claws).
The most notable member of the docodonts is Castorocauda ("beaver tail"), which lived in the mid
Jurassic about 164M years ago and was first discovered in 2004 and described in 2006. Castorocauda was
not a typical docodont (most were omnivores) and not a true mammal, but it is extremely important in the
study of the evolution of mammals because the first find was an almost complete skeleton (a real luxury in
paleontology) and it breaks the "small nocturnal insectivore" stereotype:[25]
• It was noticeably larger than most Mesozoic mammal-like fossils — about 17 in (43 cm) from
its nose to the tip of its 5-inch (130 mm) tail, and may have weighed 500–800 g (18–28 oz).
• It provides the earliest absolutely certain evidence of hair and fur. Previously the earliest was
Eomaia, a true mammal from about 125M years ago.
• It had aquatic adaptations including flattened tail bones and remnants of soft tissue between
the toes of the back feet, suggesting that they were webbed. Previously the earliest known semi-
aquatic mammal-like animals were from the Eocene, about 110M years later.
• Castorocauda's powerful forelimbs look adapted for digging. This feature and the spurs on its
ankles make it resemble the platypus, which also swims and digs.
• Its teeth look adapted for eating fish: the first two molars had cusps in a straight row, which
made them more suitable for gripping and slicing than for grinding; and these molars are curved
backwards, to help in grasping slippery prey.
[edit] Hadrocodium
Hadrocodium skull. The jaw joint is fully mammalian (squamosal-dentary only) and farther forward
than in earlier transitional forms.
The consensus family tree above shows Hadrocodium as an "aunt" of true mammals, while
symmetrodonts and kuehneotheriids are more closely related to true mammals. But fossils of symmetrodonts
and kuehneotheriids are so few and fragmentary that they are poorly understood and may be paraphyletic.
[26] On the other hand there are good fossils of Hadrocodium (about 195M years ago in the very early
Jurassic) and they have some important features: [27]
• The jaw joint consists only of the squamosal and dentary bones, and the jaw contains no
smaller bones to the rear of the dentary, unlike the therapsid design.
• In therapsids and most mammaliformes the eardrum stretched over a trough at the rear of
the lower jaw. But Hadrocodium had no such trough, which suggests its ear was part of the cranium,
as it is in mammals — and hence that the former articular and quadrate had migrated to the middle
ear and become the malleus and incus. On the other hand the dentary has a "bay" at the rear that
mammals lack. This suggests that Hadrocodium's dentary bone retained the same shape that it
would have had if the articular and quadrate had remained part of the jaw joint, and therefore that
Hadroconium or a very close ancestor may have been the first to have a fully mammalian middle ear.
• Therapsids and earlier mammaliforms had their jaw joints very far back in the skull, partly
because the ear was at the rear end of the jaw but also had to be close to the brain. This
arrangement limited the size of the braincase, because it forced the jaw muscles to run round and
over it. Hadrocodium's braincase and jaws were no longer bound to each other by the need to
support the ear, and its jaw joint was further forward. In its descendants or those of animals with a
similar arrangement, the brain case was free to expand without being constrained by the jaw and the
jaw was free to change without being constrained by the need to keep the ear near the brain — in
other words it now became possible for mammal-like animals both to develop large brains and to
adapt their jaws and teeth in ways that were purely specialized for eating.
[edit] The earliest true mammals

This part of the story introduces new complications, since true mammals are the only group that still
has living members:
• One has to distinguish between extinct groups and those that have living representatives.
• One often feels compelled to try to explain the evolution of features that do not appear in
fossils. This endeavor often involves Molecular phylogenetics, a technique that has become popular
since the mid-1980s but is still often controversial because of its assumptions, especially about the
reliability of the molecular clock.
[edit] Family tree of early true mammals
(based on Mammalia: Overview - Palaeos; X marks extinct groups)
Australosphenida
Ausktribosphenidae X
Monotremes
Triconodonta
X
Spalacotheroidea
X
Cladotheria
Dryolestoidea
X
Theria
Metatheria
Eutheria
[edit] Australosphenida and Ausktribosphenidae

Ausktribosphenidae is a group name that has been given to some rather puzzling finds that:[28]
• appear to have tribosphenic molars, a type of tooth that is otherwise known only in placentals
and marsupials.[29]
• come from mid Cretaceous deposits in Australia — but Australia was connected only to
Antarctica, and placentals originated in the northern hemisphere and were confined to it until
continental drift formed land connections from North America to South America, from Asia to Africa
and from Asia to India (the late Cretaceous map at [1] shows how the southern continents are
separated).
• are represented only by skull and jaw fragments, which is not very helpful.
Australosphenida is a group that has been defined in order to include the Ausktribosphenidae and
monotremes. Asfaltomylos (mid- to late Jurassic, from Patagonia) has been interpreted as a basal
australosphenid (animal that has features shared with both Ausktribosphenidae and monotremes; lacks
features that are peculiar to Ausktribosphenidae or monotremes; also lacks features that are absent in
Ausktribosphenidae and monotremes) and as showing that australosphenids were widespread throughout
Gondwanaland (the old Southern hemisphere super-continent).[30]
But recent analysis of Teinolophos suggests Teinolophos (about 115M years ago) was a "crown
group" (advanced and relatively specialised) monotreme, so the basal (most primitive) monotremes must
have appeared considerably earlier; that some alleged Australosphenids were also "crown group"
monotremes (e.g. Steropodon); and that other alleged Australosphenids (e.g. Ausktribosphenos, Bishops,
Ambondro, Asfaltomylos) are therefore more closely related to and possibly members of the Therian
mammals (group that includes marsupials and placentals, see below).[31]
[edit] Monotremes
The earliest known monotreme is Teinolophos, which lived about 123M years ago in Australia.
Recent (2007, published 2008) analysis suggest that it was not a basal (primitive, ancestral) monotreme but
a full-fledged platypus, and therefore that the platypus and echidna lineages diverged considerably earlier
and that basal monotremes were even earlier.[31]
A more recent study (2009), however, has suggested that while Teinolophis was a type of platypus, it
also was a basal monotreme and predated the radiation of modern monotremes. The semi-aquatic lifestyle of
platypuses prevented them from being outcompeted by the marsupials that migrated to Australia millions of
years ago, since joeys need to keep attached to their mothers and would drown if their mothers ventured into
water. Genetic evidence has determined that echidnas diverged from the platypus lineage as recently as 19-
48M when they made their transition from semi-aquatic to terrestrial lifestyle.[32]
Monotremes have some features that may be inherited from the original amniotes:
• they use the same orifice to urinate, defecate and reproduce ("monotreme" means "one
hole") — as lizards and birds also do.
• they lay eggs that are leathery and uncalcified, like those of lizards, turtles and crocodilians.
Unlike in other mammals, female monotremes do not have nipples and feed their young by
"sweating" milk from patches on their bellies.
Of course these features are not visible in fossils, and the main characteristics from paleontologists'
point of view are:[28]
• a slender dentary bone in which the coronoid process is small or non-existent.
• the external opening of the ear lies at the posterior base of the jaw.
• the jugal bone is small or non-existent.
• a primitive pectoral girdle with strong ventral elements: coracoids, clavicles and interclavicle.
Note: therian mammals have no interclavicle.[33]
• sprawling or semi-sprawling forelimbs.
[edit] Theria
Theria ("beasts") is a name applied to the hypothetical group from which both metatheria (which
include marsupials) and eutheria (which include placentals) descended. Although no convincing fossils of
basal therians have been found (just a few teeth and jaw fragments), metatheria and eutheria share some
features that one would expect to have been inherited from a common ancestral group:[34]
Therian form of crurotarsal ankle. Adapted with permission from Palaeos
• no interclavicle.[33]
• coracoid bones non-existent or fused with the shoulder blades to form coracoid processes.
• a type of crurotarsal ankle joint in which: the main joint is between the tibia and astragalus;
the calcaneum has no contact with the tibia but forms a heel to which muscles can attach. (The other
well-known type of crurotarsal ankle is seen in crocodilians and works differently — most of the
bending at the ankle is between the calcaneum and astragalus).
• tribosphenic molars.[29]
Tribosphenic molars have been found in fossils from Madagascar, which indicates that therian
mammals are at least 167 million years old.[35]
[edit] Metatheria
The living Metatheria are all marsupials ("animals with pouches"). A few fossil genera such as the
Mongolian late Cretaceous Asiatherium may be marsupials or members of some other metatherian group(s).
[36][37]
The oldest known marsupial is Sinodelphys, found in 125M-year old early Cretaceous shale in
China's northeastern Liaoning Province. The fossil is nearly complete and includes tufts of fur and imprints of
soft tissues.[38]
Didelphimorphia (common opossums of the Western Hemisphere) first appeared in the late
Cretaceous and still have living representatives, probably because they are mostly semi-arboreal
unspecialized omnivores.[39]
The best-known feature of marsupials is their method of reproduction:
• The mother develops a kind of yolk sack in her womb that delivers nutrients to the embryo.
Embryos of bandicoots, koalas and wombats additionally form placenta-like organs that connect
them to the uterine wall, although the placenta-like organs are smaller than in placental mammals
and it is not certain that they transfer nutrients from the mother to the embryo.[40]
• Pregnancy is very short, typically 4 to 5 weeks. The embryo is born at a very young age of
development, and is usually less than 2 in (5.1 cm) long at birth. It has been suggested that the short
pregnancy is necessary to reduce the risk that the mother's immune system will attack the embryo.
• The newborn marsupial uses its forelimbs (with relatively strong hands) to climb to a nipple,
which is usually in a pouch on the mother's belly. The mother feeds the baby by contracting muscles
over her mammary glands, as the baby is too weak to suck. The newborn marsupial's need to use its
forelimbs in climbing to the nipple has prevented the forelimbs from evolving into paddles or wings
and has therefore prevented the appearance of aquatic or truly flying marsupials (although there are
several marsupial gliders).
Palate of thylacine, showing one of the paired palatal fenestrae (top left), which are a signature
feature of marsupials.
Although some marsupials look very like some placentals (the thylacine or "marsupial wolf" is a good
example), marsupial skeletons have some features that distinguish them from placentals:[41]
• Some, including the thylacine, have 4 molars. No placentals have more than 3.
• All have a pair of palatal fenestrae, window-like openings on the bottom of the skull (in
addition to the smaller nostril openings).
Marsupials also have a pair of marsupial bones (sometimes called "epipubic bones"), which support
the pouch in females. But these are not unique to marsupials, since they have been found in fossils of
multituberculates, monotremes, and even eutherians — so they are probably a common ancestral feature that
disappeared at some point after the ancestry of living placental mammals diverged from that of marsupials.
[42][43] Some researchers think the epipubic bones' original function was to assist locomotion by supporting
some of the muscles that pull the thigh forwards.[44]
[edit] Eutheria
Main article: Eutheria
The living Eutheria ("true beasts") are all placentals. But the earliest known eutherian, Eomaia, found
in China and dated to 125M years ago, has some features that are more like those of marsupials (the
surviving metatherians):[45]
Fossil of Eomaia in the Hong Kong Science Museum.
• Epipubic bones extending forwards from the pelvis, which are not found in any modern
placental, but are found in all other mammals — non-placental eutherians, marsupials, monotremes
and mammaliformes — and even in the cynodont therapsids that are closest to mammals. Their
function is to stiffen the body during locomotion.[46] This stiffening would be harmful in pregnant
placentals, whose abdomens need to expand.[47]
• A narrow pelvic outlet, which indicates that the young were very small at birth and therefore
pregnancy was short, as in modern marsupials. This suggests that the placenta was a later
development.
• 5 incisors in each side of the upper jaw. This number is typical of metatherians, and the
maximum number in modern placentals is 3, except for homodonts such as the armadillo. But
Eomaia's molar to premolar ratio (it has more pre-molars than molars) is typical of eutherians,
including placentals, and not normal in marsupials.
Eomaia also has a Meckelian groove, a primitive feature of the lower jaw that is not found in modern
placental mammals.
These intermediate features are consistent with molecular phylogenetics estimates that the
placentals diversified about 110M years ago, 15M years after the date of the Eomaia fossil.
Eomaia also has many features that strongly suggest it was a climber, including several features of
the feet and toes; well-developed attachment points for muscles that are used a lot in climbing; and a tail that
is twice as long as the rest of the spine.
Placentals' best-known feature is their method of reproduction:
• The embryo attaches itself to the uterus via a large placenta via which the mother supplies
food and oxygen and removes waste products.
• Pregnancy is relatively long and the young are fairly well-developed at birth. In some species
(especially herbivores living on plains) the young can walk and even run within an hour of birth.
It has been suggested that the evolution of placental reproduction was made possible by retroviruses
that:[48]
• make the interface between the placenta and uterus into a syncytium, i.e. a thin layer of cells
with a shared external membrane. This allows the passage of oxygen, nutrients and waste products
but prevents the passage of blood and other cells, which would cause the mother's immune system
to attack the fetus.
• reduce the aggressiveness of the mother's immune system (which is good for the foetus but
makes the mother more vulnerable to infections).
From a paleontologist's point of view, eutherians are mainly distinguished by various features of their
teeth,[49] ankles and feet.[50]
[edit] Expansion of ecological niches in the Mesozoic

There is still some truth in the "small, nocturnal insectivores" stereotype but recent finds, mainly in
China, show that some mammaliforms and true mammals were larger and had a variety of lifestyles. For
example:
• Castorocauda, a member of Docodonta which lived in the middle Jurassic about 164 million
years, was about 42.5 cm (16.7 in) long, weighed 500–800 g (18–28 oz), had limbs that were adapted
for swimming and digging and teeth adapted for eating fish.[25]
• Multituberculates, are allotherians that survived for over 125 million years (from mid Jurassic,
about 160M years ago, to early Oligocene, about 35M years ago) are often called the "rodents of the
Mesozoic", because they had continuously-growing incisors like those of modern rodents.[23]
Repenomamus sometimes preyed on young dinosaurs

• Fruitafossor, from the late Jurassic period about 150 million years ago, was about the size of
a chipmunk and its teeth, forelimbs and back suggest that it broke open the nest of social insects to
prey on them (probably termites, as ants had not yet appeared).[51]
• Volaticotherium, allotherians from the boundary the early Cretaceous about 125M years ago,
is the earliest-known gliding mammal and had a gliding membrane that stretched out between its
limbs, rather like that of a modern flying squirrel. This also suggests it was active mainly during the
day.[52]
• Repenomamus, tricodonts from the early Cretaceous 130 million years ago, was a stocky,
badger-like predator that sometimes preyed on young dinosaurs. Two species have been
recognized, one more than 1 m (39 in) long and weighing about 12–14 kg (26–31 lb), the other less
than 0.5 m (20 in) long and weighing 4–6 kg (8.8–13 lb).[53][54]
[edit] Evolution of major groups of living mammals

There are currently vigorous debates between traditional paleontologists ("fossil-hunters") and
molecular phylogeneticists about how and when the true mammals diversified, especially the placentals.
Generally the traditional paelontologists date the appearance of a particular group by the earliest known
fossil whose features make it likely to be a member of that group, while the molecular phylogeneticists
suggest that each lineage diverged earlier (usually in the Cretaceous) and that the earliest members of each
group were anatomically very similar to early members of other groups and differed only in their genes.
These debates extend to the definition of and relationships between the major groups of placentals — the
controversy about Afrotheria is a good example.
[edit] Fossil-based family tree of placental mammals

Here is a very simplified version of a typical family tree based on fossils, based on Cladogram of
Mammalia - Palaeos. It tries to show the nearest thing there is at present to a consensus view, but some
paleontologists have very different views, for example:[55]
• The most common view is that placentals originated in the southern hemisphere, but some
paleontologists argue that they first appeared in Laurasia (old supercontinent containing modern
Asia, N. America and Europe).
• Paleontologists differ about when the first placentals appeared, with estimates ranging from
20M years before the end of the Cretaceous to just after the end of the Cretaceous. And molecular
biologists argue for a much earlier origin.
• Most paleontologists suggest that placentals should be divided into Xenarthra and the rest,
but a few think these animals diverged later.
For the sake of brevity and simplicity the diagram omits some extinct groups in order to focus on the
ancestry of well-known modern groups of placentals — X marks extinct groups. The diagram also shows the
following:
• the age of the oldest known fossils in many groups, since one of the major debates between
traditional paleontologists and molecular phylogeneticists is about when various groups first became
distinct.
• well-known modern members of most groups.
Eutheria
Epitheria (latest Cretaceous)

Anagalida
Zalambdalestidae X
(late Cretaceous)
Macroscelidea
(late Eocene)
(elephant shrews)
Anagaloidea
X
Glires (early Paleocene)

Lagomorpha
(Eocene)
(rabbits, hares, pikas)
Rodentia (late
Paleocene)
(mice & rats, squirrels,
porcupines)
Archonta
Scandentia (mid
Eocene)
(tree shrews)
Primatomorpha
Plesiadapiformes X
Primates (early Paleocene)
(tarsiers, lemurs, monkeys, apes including humans)
Dermoptera (late Eocene)

(colugos)
Chiroptera (late Paleocene)

(bats)
Carnivora
(early Paleocene)
(cats, dogs, bears,
seals)
Ungulatomorpha (late Cretaceous)
Eparctocyona (late Cretaceous)
(some
extinct groups)
X
Arctostylopida
X (late Paleocene)
Mesonychia
X (mid Paleocene)
(predators /
scavengers, but not
closely related to
modern carnivores)
Cetartiodactyla
Cetacea
(early Eocene)
(whales, dolphins,
porpoises)
Artiodactyla
(early Eocene)
(even-toed
ungulates: pigs,
hippos, camels,
giraffes, cattle,
deer)
Altungulata
Hilalia
X
Perissodactyla (late Paleocene)

(odd-toed ungulates: horses, rhinos, tapirs)
Tubulidentata (early Miocene)

(aardvarks)
Paenungulata ("not quite ungulates")

Hyracoidea (early
Eocene)
(hyraxes)
Sirenia (early
Eocene)
(manatees, dugongs)
Proboscidea (early
Eocene)
(elephants)
This family tree contains some surprises and puzzles. For example:
• The closest living relatives of cetaceans (whales, dolphins, porpoises) are artiodactyls,
hoofed animals, which are almost all pure vegetarians.
• Bats are fairly close relatives of primates.
• The closest living relatives of elephants are the aquatic sirenians, while their next relatives
are hyraxes, which look more like well-fed guinea pigs.
• There is little correspondence between the structure of the family (what was descended from
what) and the dates of the earliest fossils of each group. For example the earliest fossils of
perissodactyls (the living members of which are horses, rhinos and tapirs) date from the late
Paleocene but the earliest fossils of their "sister group" the Tubulidentata date from the early
Miocene, nearly 50M years later. Paleontologists are fairly confident about the family relationships,
which are based on cladistic analyses, and believe that fossils of the ancestors of modern aardvarks
have simply not been found yet.
[edit] Family tree of placental mammals according to molecular
phylogenetics
Molecular phylogenetics uses features of organisms' genes to work out family trees in much the
same way as paleontologists do with features of fossils — if two organisms' genes are more similar to each
other than to those of a third organism, the two organisms are more closely related to each other than to the
third.
Molecular phylogeneticists have proposed a family tree that is very different from the one with which
paleontologists are familiar. Like paleontologists, molecular phylogeneticists have different ideas about
various details, but here is a typical family tree according to molecular phylogenetics:[56][57] Note that the
diagram shown here omits extinct groups, as one cannot extract DNA from fossils.
Eutheria
Atlantogenata ("born round the Atlantic ocean")
Xenarthra
(armadillos,
anteaters, sloths)
Afrotheria
Afroinsectiphilia
(golden moles, tenrecs,
otter shrews)
unnamed
Macroscelidea (elephant
shrews)
Tubulidentata (aardvarks)
Paenungulata ("not quite ungulate

s")
Hyracoidea (hyraxes)
Proboscidea (elephants)
Sirenia (manatees,
dugongs)
Boreoeutheria ("northern true / placental mammals")
Laurasiatheria
Erinaceomorpha (hedgehogs,
gymnures)
Soricomorpha (moles, shrews,

solenodons)
Cetartiodactyla
Pegasoferae
Pholidota (pangolins)
Chiroptera (bats)
Carnivora (cats, dogs, bears,
seals)
Perissodactyla (horses,
rhinos, tapirs).
Euarchontoglires
Glires
Lagomorpha (rabbits, hares, pikas)
Rodentia (late Paleocene)(mice & rats, squirrels,

porcupines)
Euarchonta
Scandentia (tree shrews)
Dermoptera (colugos)
Primates (tarsiers, lemurs, monkeys, apes including

humans)
Here are the most significant of the many differences between this family tree and the one familiar to
paleontologists:
• The top-level division is between Atlantogenata and Boreoeutheria, instead of between
Xenarthra and the rest. But some molecular phylogeneticists have proposed a 3-way top-level split
between Xenarthra, Afrotheria and Boreoeutheria.
• Afrotheria contains several groups that are only distantly related according to the
paleontologists' version: Afroinsectiphilia ("African insectivores"), Tubulidentata (aardvarks, which
paleontologists regard as much closer to odd-toed ungulates than to other members of Afrotheria),
Macroscelidea (elephant shrews, usually regarded as close to rabbits and rodents). The only
members of Afrotheria that paleontologists would regard as closely related are Hyracoidea (hyraxes),
Proboscidea (elephants) and Sirenia (manatees, dugongs).
• Insectivores are split into 3 groups: one is part of Afrotheria and the other two are distinct
sub-groups within Boreoeutheria.
• Bats are closer to Carnivora and odd-toed ungulates than to primates and Dermoptera
(colugos).
• Perissodactyla (odd-toed ungulates) are closer to Carnivora and bats than to Artiodactyla
(even-toed ungulates).
The grouping together of the Afrotheria has some geological justification. All surviving members of
the Afrotheria originate from South American or (mainly) African lineages — even the Indian elephant, which
diverged from an African lineage about 7.6 million years ago.[58] As Pangaea broke up Africa and South
America separated from the other continents less than 150M years ago, and from each other between 100M
and 80M years ago.[59][60] The earliest known eutherian mammal is Eomaia, from about 125M years ago.
So it would not be surprising if the earliest eutherian immigrants into Africa and South America were isolated
there and radiated into all the available ecological niches.
Nevertheless these proposals have been controversial. Paleontologists naturally insist that fossil
evidence must take priority over deductions from samples of the DNA of modern animals. More surprisingly,
these new family trees have been criticised by other molecular phylogeneticists, sometimes quite harshly:[61]
• Mitochondrial DNA's mutation rate in mammals varies from region to region — some parts
hardly ever change and some change extremely quickly and even show large variations between
individuals within the same species.[62][63]
• Mammalian mitochondrial DNA mutates so fast that it causes a problem called "saturation",
where random noise drowns out any information that may be present. If a particular piece of
mitochondrial DNA mutates randomly every few million years, it will have changed several times in
the 60 to 75M years since the major groups of placental mammals diverged.[64]
[edit] Timing of placental evolution

Recent molecular phylogenetic studies suggest that most placental orders diverged about 100M to
85M years ago, but that modern families first appeared in the late Eocene and early Miocene.[65]
Some paleontologists object that no placental fossils have been found from before the end of the
Cretaceous — for example Maelestes gobiensis, from about 75M years ago, is a eutherian but not a true
placental.[66] Many Cretaceous fossil sites contain well-preserved lizards, salamanders, birds, and
mammals, but not the modern forms of mammals. It is likely that they simply did not exist, and that the
molecular clock runs fast during major evolutionary radiations.[67] On the other hand there is fossil evidence
from 85 million years ago of hoofed animals that may be ancestors of modern ungulates.[68]
Fossils of the earliest members of most modern groups date from the Paleocene, a few date from
later and very few from the Cretaceous, before the extinction of the dinosaurs. But some paleontologists,
influenced by molecular phylogenetic studies, have used statistical methods to extrapolate backwards from
fossils of members of modern groups and concluded that primates arose in the late Cretaceous.[69] However
statistical studies of the fossil record confirm that mammals were restricted in size and diversity right to the
end of the Cretaceous, and rapidly grew in size and diversity during the Early Paleocene.[70][71]
[edit] Evolution of mammalian features

[edit] Jaws and middle ears
See also Evolution of mammalian auditory ossicles
Hadrocodium, whose fossils date from the early Jurassic, provides the first clear evidence of fully
mammalian jaw joints and middle ears, in which the jaw joint is formed by the dentary and squamosal bones
while the articular and quadrate move to the middle ear, where they are known as the incus and malleus.
Curiously it is usually classified as a member of the mammaliformes rather than a as a true mammal.
One analysis of the monotreme Teinolophos suggested that this animal had a pre-mammalian jaw
joint formed by the angular and quadrate bones and that the typical mammalian middle ear evolved twice
independently, in monotremes and in therian mammals, but this idea has been disputed.[72] In fact 2 of the
suggestion's authors co-authored a later paper that reinterpreted the same features as evidence that
Teinolophos was a full-fledged platypus, which means it would have had a mammalian jaw joint and middle
ear.[31]
[edit] Milk production (lactation)

It has been suggested that lactation's original function was to keep eggs moist. Much of the argument
is based on monotremes (egg-laying mammals):[73][74][75]
• Monotremes do not have nipples but secrete milk from a hairy patch on their bellies.
• During incubation, monotremes' eggs are covered in a sticky substance whose origin is not
known. Before the eggs are laid, their shells have only three layers. Afterwards a fourth layer
appears, and its composition is different from that of the original three. The sticky substance and the
fourth layer may be produced by the mammary glands.
• If so, that may explain why the patches from which monotremes secrete milk are hairy — it is
easier to spread moisture and other substances over the egg from a broad, hairy area than from a
small, bare nipple.
[edit] Hair and fur

See also: Evolution of hair
The first clear evidence of hair or fur is in fossils of Castorocauda, from 164M years ago in the mid
Jurassic.
From 1955 onwards some scientists have interpreted the foramina (passages) in the maxillae (upper
jaws) and premaxillae (small bones in front of the maxillae) of cynodonts as channels that supplied blood
vessels and nerves to vibrissae (whiskers), and suggested that this was evidence of hair or fur.[76][77] But
foramina do not necessarily show that an animal had vibrissae — for example the modern lizard Tupinambis
has foramina that are almost identical to those found in the non-mammalian cynodont Thrinaxodon.[9][78]
[edit] Erect limbs

The evolution of erect limbs in mammals is incomplete — living and fossil monotremes have sprawling
limbs. In fact some scientists think that the parasagittal (non-sprawling) limb posture is a synapomorphy
(distinguishing characteristic) of the Boreosphenida, a group that contains the Theria and therefore includes
the last common ancestor of modern marsupials and placentals — and therefore that all earlier mammals had
sprawling limbs.[79]
Sinodelphys (the earliest known marsupial) and Eomaia (the earliest known eutherian) lived about
125M years ago, so erect limbs must have evolved before then.
[edit] Warm-bloodedness
"Warm-bloodedness" is a complex and rather ambiguous term, because it includes some or all of the
following:
• Endothermy, i.e. the ability to generate heat internally rather than via behaviors such as
basking or muscular activity.
• Homeothermy, i.e. maintaining a fairly constant body temperature.
• Tachymetabolism, i.e. maintaining a high metabolic rate, particularly when at rest. This
requires a fairly high and stable body temperature, since biochemical processes run about half as
fast if an animal's temperature drops by 10°C; most enzymes have an optimum operating
temperature and their efficiency drops rapidly outside the preferred range.
Since scientists cannot know much about the internal mechanisms of extinct creatures, most
discussion focuses on homeothermy and tachymetabolism.
Modern monotremes have a lower body temperature and more variable metabolic rate than
marsupials and placentals.[80] So the main question is when a monotreme-like metabolism evolved in
mammals. The evidence found so far suggests Triassic cynodonts may have had fairly high metabolic rates,
but is not conclusive.
[edit] Respiratory turbinates
Modern mammals have respiratory turbinates, convoluted structures of thin bone in the nasal cavity.
These are lined with mucous membranes that warm and moisten inhaled air and extract heat and moisture
from exhaled air. An animal with respiratory turbinates can maintain a high rate of breathing without the
danger of drying its lungs out, and therefore may have a fast metabolism. Unfortunately these bones are very
delicate and therefore have not yet been found in fossils. But rudimentary ridges like those that support
respiratory turbinates have been found in Triassic therapsids such as Thrinaxodon and Diademodon, which
suggests that they may have had fairly high metabolic rates. [76] [81][82]
[edit] Bony secondary palate

Mammals have a secondary bony palate, which separates the respiratory passage from the mouth,
allowing them to eat and breathe at the same time. Secondary bony palates have been found in the more
advanced cynodonts and have been used as evidence of high metabolic rates.[76][77][83] But some cold-
blooded vertebrates have secondary bony palates (crocodilians and some lizards), while birds, which are
warm-blooded, do not have them.[9]
[edit] Diaphragm
A muscular diaphragm helps mammals to breathe, especially during strenuous activity. For a
diaphragm to work, the ribs must not restrict the abdomen, so that expansion of the chest can be
compensated for by reduction in the volume of the abdomen and vice versa. The advanced cynodonts have
very mammal-like rib cages, with greatly reduced lumbar ribs. This suggests that these animals had
diaphragms, were capable of strenuous activity for fairly long periods and therefore had high metabolic rates.
[76][77] On the other hand these mammal-like rib cages may have evolved to increase agility.[9] But the
movement of even advanced therapsids was "like a wheelbarrow", with the hindlimbs providing all the thrust
while the forelimbs only steered the animal, in other words advanced therapsids were not as agile as either
modern mammals or the early dinosaurs.[84] So the idea that the main function of these mammal-like rib
cages was to increase agility is doubtful.
[edit] Limb posture

The therapsids had sprawling forelimbs and semi-erect hindlimbs.[77][85] This suggests that
Carrier's constraint would have made it rather difficult for them to move and breathe at the same time, but not
as difficult as it is for animals such as lizards, which have completely sprawling limbs.[86] But cynodonts
(advanced therapsids) had costal plates that stiffened the rib cage and therefore may have reduced sideways
flexing of the trunk while moving, which would have made it a little easier for them to breathe while moving .
[87] These facts suggest that advanced therapsids were significantly less active than modern mammals of
similar size and therefore may have had slower metabolisms.
[edit] Insulation (hair and fur)

Insulation is the "cheapest" way to maintain a fairly constant body temperature, without consuming
energy to produce more body heat. Therefore, possession of hair or fur would be good evidence of
homeothermy but would not be such strong evidence of a high metabolic rate.[88] [89]
The first clear evidence of hair or fur is in fossils of Castorocauda, from 164M years ago in the mid
Jurassic;[25] arguments that advanced therapsids had hair are unconvincing.
[edit] References
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[edit] Bibliography
• Robert L. Carroll, Vertebrate Paleontology and Evolution, W. H. Freeman and Company,
New York, 1988 ISBN 0-716-71822-7. Chapters XVII through XXI
• Nicholas Hotton III, Paul D. MacLean, Jan J. Roth, and E. Carol Roth, editors, The Ecology
and Biology of Mammal-like Reptiles, Smithsonian Institution Press, Washington and London, 1986
ISBN 0-87474-524-1
• T. S. Kemp, The Origin and Evolution of Mammals , Oxford University Press, New York, 2005
ISBN 0-19-850760-7
• Zofia Kielan-Jaworowska, Richard L. Cifelli, and Zhe-Xi Luo, Mammals from the Age of
Dinosaurs: Origins, Evolution, and Structure, Columbia University Press, New York, 2004 ISBN 0-
231-11918-6. Comprehensive coverage from the first mammals up to the time of the K-T mass
extinction.
• Zhe-Xi Luo, "Transformation and diversification in early mammal evolution", Nature volume
450 number 7172 (13 December 2007) pages 1011–1019. doi:10.1038/nature06277. A survey article
with 98 references to the scientific literature.

• The Cynodontia covers several aspects of the evolution of cynodonts into mammals, with
plenty of references.
Retrieved from "http://en.wikipedia.org/wiki/Evolution_of_mammals"
Categories: Evolution by taxon | Mammals | Synapsids | Mammalogy | Paleontology | Mammal

anatomy | Prehistoric mammals
W000
Great American Interchange


Examples of migrant species in both Americas. Olive green silhouettes denote North American
species with South American ancestors; blue silhouettes denote South American species of North American
origin.
The Great American Interchange was an important paleozoogeographic event in which land and
freshwater fauna migrated from North America via Central America to South America and vice versa, as the
volcanic Isthmus of Panama rose up from the sea floor and bridged the formerly separated continents. The
migration peaked dramatically around three million years (Ma) ago (during the Piacenzian, the upper 1 Ma of
the Pliocene).
It resulted in the joining of the Neotropic (roughly South America) and Nearctic (roughly North
America) definitively to form the Americas. The interchange is visible from observation of both stratigraphy
and nature (neontology). Its most dramatic effect is on the zoogeography of mammals but it also gave an
opportunity for weak-flying or flightless birds, reptiles, amphibians, arthropods and even freshwater fish to
migrate.
The occurrence of the interchange was first discussed in 1876 by the "father of biogeography", Alfred
Russel Wallace.[1][note 1] (Wallace had spent 1848-1852 exploring and collecting specimens in the Amazon
Basin.) Others who made significant contributions to understanding the event in the century that followed
include Florentino Ameghino, W. D. Matthew, W. B. Scott, Bryan Patterson, George Gaylord Simpson and S.
David Webb.[2]
Similar interchanges occurred earlier in the Cenozoic, when the formerly isolated land masses of
India[3] and Africa[4] made contact with Eurasia c. 50 and 30 Ma ago, respectively.
Contents
[hide]
• 1 South America's endemic fauna
• 2 Island-hopping ‘waif dispersers’
• 3 The Great American Biotic Interchange
• 4 Reasons for success or failure
• 5 Late Pleistocene extinctions
• 6 South American invasions of North America exclusive of
Central America
• 7 South American invasions that failed to penetrate beyond
Central America
• 8 North American invasions of South America
• 9 See also
• 10 Notes
• 11 References
[edit] South America's endemic fauna
The sabertooth "marsupial" †Thylacosmilus

The litoptern †Macrauchenia patagonica
†Astrapotherium magnum
Myrmecophaga tridactyla, the largest living descendant of South American's early Cenozoic fauna
After the late Mesozoic breakup of Gondwana, South America spent most of the Cenozoic era as an
island continent whose "splendid isolation" allowed its fauna to evolve into many forms found nowhere else
on earth, most of which are now extinct.[5] Its endemic mammals initially consisted of metatherians
(marsupials and sparassodonts), xenarthrans, and a diverse group of native ungulates: notoungulates (the
"southern ungulates"), litopterns, astrapotheres (e.g. Trigonostylops, Astrapotherium), and pyrotheres (e.g.
Pyrotherium).
Marsupials appear to have traveled (via Gondwanan land connections) from South America through
Antarctica to Australia in the late Cretaceous or early Tertiary.[6] One living South American marsupial, the
Monito del Monte, has been shown to be more closely related to Australian marsupials than to other South
American marsupials; however, it is the most basal australidelphian, meaning that this superorder arose in
South America and then colonized Australia after the Monito del Monte split off.[6] A 61-Ma-old platypus-like
monotreme fossil from Patagonia may represent an Australian immigrant. It appears that ratites (relatives of
South American tinamous) migrated by this route around the same time, more likely in the direction from
South America towards Australia/New Zealand.[7] Other taxa that may have dispersed by the same route (if
not by flying or floating across the ocean) are parrots, chelid turtles and (extinct) meiolaniid turtles.
Marsupials present in South America included didelphimorphs (opossums) and several other small
groups; larger predatory relatives of these also existed, like the borhyaenids and the sabertooth
Thylacosmilus (sparassodont metatherians which are no longer considered to be marsupials).[8]
Metatherians were the only South American mammals to specialize as carnivores; their relative
inefficiency created openings for nonmammalian predators to play more prominent roles than usual (similar
to the situation in Australia). Sparassodonts shared the ecological niches for large predators with fearsome
flightless "terror birds" (phorusrhacids), whose closest extant relatives are the seriemas.[9][10] Terrestrial
ziphodont[note 2] sebecid crocodilians were also present at least through the middle Miocene.[11][12][13][14]
Some of South America's aquatic crocodilians, such as Gryposuchus, Mourasuchus and Purussaurus,
reached monstrous sizes, with lengths up to 12 m. Through the skies over late Miocene South America (6 Ma
ago) soared the largest flying bird known, the teratorn Argentavis, with a wing span of 6 m or more, which
may have subsisted in part on the leftovers of Thylacosmilus kills.[15]
Xenarthrans are a curious group of mammals that developed morphological adaptations for
specialized diets very early in their history.[16] In addition to those extant today (armadillos, anteaters and
tree sloths), a great diversity of larger types were present, including pampatheres, the ankylosaur-like
glyptodontids, various ground sloths, some of which reached the size of elephants (e.g. Megatherium), and
even semiaquatic marine sloths.
The notoungulates and litopterns had many strange forms, like Macrauchenia, a camel-like litoptern
with a small proboscis. They also produced a number of familiar-looking body types that represent examples
of parallel or convergent evolution: one-toed Thoatherium had legs like those of a horse, Pachyrukhos
resembled a rabbit, Homalodotherium was a semi-bipedal clawed browser like a chalicothere, and horned
Trigodon looked like a rhino. Both groups started evolving in the Lower Paleocene, possibly from condylarth
stock, diversified, dwindled before the great interchange, and went extinct at the end of the Pleistocene. The
pyrotheres and astrapotheres were also strange but were less diverse and disappeared earlier, well before
the interchange.
The North American fauna was a pretty typical boreoeutherian one (supplemented with Afrotherian
proboscids).
[edit] Island-hopping ‘waif dispersers’
Hydrochoerus hydrochaeris
Saguinus imperator
Geochelone carbonaria
Megalonyx jeffersonii
The invasions of South America started at least 31.5 Ma ago (late Eocene/early Oligocene), when
caviomorph rodents arrived. Their subsequent vigorous diversification displaced some of South America's
small marsupials and gave rise to – among others – capybaras, chinchillas, viscachas, and New World
porcupines. (The independent development of spines by New and Old World porcupines is another example
of parallel evolution.) This invasion most likely came from Africa.[17][18] The crossing from West Africa to the
northeast corner of Brazil was much shorter then due to continental drift, and may have been aided by island-
hopping (e.g. via St. Paul's Rocks, if they were an inhabitable island at the time) and westward oceanic
currents.[19] Crossings of the ocean were accomplished when at least one fertilised female (more commonly
a group of animals) accidentally floated over on driftwood or mangrove rafts. (Island-hopping caviomorphs
would subsequently colonize the West Indies as far as the Bahamas). Over time, some caviomorph rodents
evolved into larger forms that competed with some of the native South American ungulates, which may have
contributed to the gradual loss of diversity suffered by the latter after the early Oligocene.[5]
A little later (at least 25 Ma ago) primates followed, probably from Africa in a fashion similar to that of
the rodents. Primates capable of migrating had to be small. With little effective competition they also
diversified widely, giving rise to the New World monkeys. (Not long after arriving, monkeys apparently most
closely related to titis island-hopped to Cuba, Hispaniola and Jamaica.) The South American caviomorph
rodents and monkeys are both believed to be clades (i.e., monophyletic).
Tortoises also arrived in South America in the Oligocene. It was long thought that they had come
from North America, but a recent comparative genetic analysis concludes that South American members of
Geochelone are actually most closely related to African hingeback tortoises.[note 3][20] Tortoises are aided
in oceanic dispersal by their ability to float with their heads up, and to survive up to six months without food or
water.[20] South American tortoises then went on to colonize the West Indies and Galápagos Islands. Skinks
of the related genera Mabuya and Trachylepis apparently floated across the Atlantic from Africa to South
America and Fernando de Noronha, respectively, during the last 9 Ma.[21]
The earliest mammalian arrival from North America was a carnivorous procyonid that island-hopped
from Central America before a land bridge formed, around 7 Ma ago. This was South America's first
eutherian carnivore. South American procyonids then diversified into forms now extinct (e.g. the "dog-coati"
Cyonasua, which evolved into the bear-like Chapalmalania). However, all extant procyonid genera appear to
have originated in North America.[22] It has been suggested that the first South American procyonids may
have contributed to the extinction of sebecid crocodilians by eating their eggs, but this view has not been
universally viewed as plausible.[a][14] The procyonids were followed to South America by island-hopping
sigmodontine rodents,[23] peccaries and hog-nosed skunks.[24] The oryzomyine tribe of sigmodontine
rodents then went on to colonize the Lesser Antilles up to Anguilla.
Similarly, megalonychid and mylodontid ground sloths island-hopped to North America by 9 Ma ago.
[23] Megalonychids had colonized the Antilles previously, by the early Miocene.[25] (Megatheriid ground
sloths had to wait for the formation of the isthmus, but then sent several lineages north.) Terror birds may
have also island-hopped to North America as early as 5 Ma ago.[26]
The Caribbean islands were populated primarily by species from South America. This was due to the
prevailing direction of oceanic currents, rather than to a competition between North and South American
forms.[27] (Except in the case of Jamaica, oryzomyine rodents of North American origin were able to enter
the region only after invading South America.)
[edit] The Great American Biotic Interchange
†Titanis walleri, the only known North American terror bird
Didelphis virginiana, the only marsupial in temperate North America

The formation of the Isthmus of Panama led to the last and most conspicuous wave, the great
interchange, around 3 Ma ago. This included the immigration of North American ungulates (including
camelids, tapirs, deer and horses), proboscids (gomphotheres), carnivorans (including felids like cougars and
saber-toothed cats, canids, mustelids, procyonids and bears) and a number of types of rodents[note 4] into
South America. The larger members of the reverse migration, besides ground sloths and terror birds, were
glyptodontids, pampatheres, capybaras and the notoungulate Mixotoxodon (the only South American
ungulate known to have invaded Central America).
In general, the initial net migration was symmetrical. Later on, however, the Neotropic species proved
far less successful than the Nearctic. This misfortune happened both ways. Northwardly migrating animals
often were not able to compete for resources as well as the North American species already occupying the
same ecological niches; those that succeeded in becoming established did not diversify much.[28]
Southwardly migrating Nearctic species established themselves in larger numbers and diversified
considerably more,[28] and are thought to have caused the extinction of a large proportion of the South
American fauna. (There were no extinctions in North America obviously attributable to South American
immigrants.) Although terror birds were initially able to invade North America, this success was temporary; all
of the large Neotropic avian and metatherian predators ultimately disappeared.[i] South America's native
ungulates also fared very poorly, with only several of the largest forms, Macrauchenia and a few toxodontids,
withstanding the northern onslaught. (Among the notoungulates, the mesotheriids and hegetotheriids did
manage to survive into the Pleistocene.)[h] Its small marsupials fared better, while the primitive-looking
xenarthrans proved to be surprisingly competitive. The African immigrants, the caviomorph rodents and
platyrrhine monkeys, generally held their own during the interchange, although the largest rodents (e.g. the
dinomyids) disappeared. With the exception of the North American porcupine and several extinct porcupines
and capybaras, however, they did not migrate past Central America.[note 5]
Due in large part to the success of the xenarthrans, one area of South American ecospace the
Nearctic invaders were unable to dominate was the niches for megaherbivores. Before 12,000 years ago,
South America was home to about 25 species of herbivores weighing more than 1000 kg, consisting of
Neotropic ground sloths, glyptodontids and toxodontids, as well as gomphotheres and camelids of Nearctic
origin.[m] Native South American forms made up about 75% of these species. However, none of these
megaherbivores have survived.
The presence of armadillos, opossums and porcupines in North America today is explained by the
Great American Interchange. Opossums and porcupines were among most successful northward migrants,
reaching as far as Canada and Alaska, respectively. While only one example each of xenarthrans,
marsupials and caviomorph rodents currently lives in the United States, 31 species from these taxa are
present in tropical Central America. Most major groups of xenarthrans were present in North America up until
the end-Pleistocene Quaternary extinction event (as a result of at least seven successful invasions of
temperate North America, and at least six more invasions of Central America only). Among the megafauna,
ground sloths were notably successful emigrants; Megalonyx spread as far north as the Yukon[29] and
Alaska,[30] and might well have eventually reached Eurasia if the extinction event had not intervened.
Generally speaking, however, the dispersal and subsequent explosive adaptive radiation of
sigmodontine rodents throughout South America (leading to over 80 currently recognized genera) was vastly
more successful (both spatially and by number of species) than any northward migration of South American
mammals. Other examples of North American mammal groups that diversified conspicuously in South
America include canids and cervids, both of which currently have 4 genera in North America, 2 or 3 in Central
America, and 6 in South America.[k] Although Canis currently ranges only as far south as Panama, South
America still has more extant canid genera than any other continent.
The effect of formation of the isthmus on the marine biota of the area was the inverse of its effect on
terrestrial organisms, a development that has been termed the "Great American Schism". The connections
between the east Pacific Ocean and the Caribbean were severed, setting now-separated populations on
divergent evolutionary paths.[31] Caribbean species also had to adapt to an environment of lower
productivity after the inflow of nutrient-rich water of deep Pacific origin was blocked.[32]
[edit] Reasons for success or failure
A north-south climatic asymmetry in the Americas. Tropical climate zones, which are warm year-
round and moist at least part of the year (blue zones Af, Am and Aw) cover much of South America and
nearly all of Central America, but very little of the rest of North America.
The eventual triumph of the Nearctic migrants was ultimately based on geography, which played into
the hands of the northern invaders in two crucial respects. The first was a matter of climate. Obviously, any
species that reached Panama from either direction had to be able to tolerate moist tropical conditions. Those
migrating southward would then be able to occupy much of South America without encountering climates that
were markedly different. However, northward migrants would have encountered drier and/or cooler
conditions by the time they reached the vicinity of the Trans-Mexican Volcanic Belt. The challenge this
climatic asymmetry (see map on right) presented was particularly acute for Neotropic species specialized for
tropical rainforest environments, who had little prospect of penetrating beyond Central America.
Land areas over which ancestors of Neotropic (green) and Nearctic (red) species could wander via
two-way migrations during the latter part of the Cenozoic. The smaller area available for Neotropic species to
evolve in tended to put them at a competitive disadvantage during the interchange.
The second and more important advantage geography gave to the northerners is related to the land
area available for their ancestors to evolve in. During the Cenozoic, North America was periodically
connected to Eurasia via Beringia, allowing multiple migrations back and forth to unite the faunas of the two
continents.[j] Eurasia was connected in turn to Africa, which contributed further to the species that made their
way to North America. South America, on the other hand, was connected to Antarctica and Australia, two
much smaller continents, only in the earliest part of the Cenozoic, and this land connection does not seem to
have carried much traffic (apparently no mammals other than marsupials and perhaps a few monotremes
ever migrated by this route). Effectively, this means that northern hemisphere species arose over a land area
roughly six times larger than was available to South American species. This calculation may not be entirely
fair, in that migrations between continents would have been more difficult and less frequent than migrations
within South America. Nevertheless, it is clear that North American species were products of a larger and
more competitive arena,[33][34] where evolution would have proceeded more rapidly. They tended to be
more efficient and brainier,[b] generally able to outrun and outwit their South American counterparts. These
advantages can be clearly seen in the cases of ungulates and their predators, where South American forms
were replaced wholesale by the invaders.
Against this backdrop, the ability of South America's xenarthrans to compete effectively against the
northerners represents a special case. The explanation for the xenarthrans' success lies in part in their
idiosyncratic approach to defending against predation, based on possession of body armor and/or formidable
claws. The xenarthrans did not need to be fleet-footed or quick-witted to survive. Such a strategy may have
been forced on them by their low metabolic rate (the lowest among the therians).[35][36] Their low metabolic
rate may in turn have been advantageous in allowing them to subsist on less abundant[37] and/or less
nutritious food sources. Unfortunately, the defensive adaptations of the large xenarthrans would have been
useless against humans armed with spears and other projectiles.
[edit] Late Pleistocene extinctions
Homo sapiens and †Glyptodon
Erethizon dorsatum, the largest surviving Neotropic migrant to temperate North America
Tapirus bairdii, the largest surviving Nearctic migrant to South America
At the end of the Pleistocene epoch, about 12,000 years ago, three dramatic developments occurred
in the Americas at roughly the same time (geologically speaking). Paleoindians invaded and occupied the
New World, the last glacial period came to an end, and a large fraction of the megafauna of both North and
South America went extinct. This wave of extinctions swept off the face of the Earth many of the successful
participants of the Great American Interchange, as well as other species that had not migrated. All the
pampatheres, glyptodontids, ground sloths, equids, proboscids,[38][39][40] dire wolves, lions and Smilodon
species of both continents disappeared. The last of the South and Central American notoungulates and
litopterns died out, as well as North America's giant beavers, dholes, native cheetahs, scimitar cats, and
many of its antilocaprid, bovid, cervid, tapirid and tayassuid ungulates. Some groups disappeared over most
or all of their original range but survived in their adopted homes, e.g. South American tapirs, camelids and
tremarctine bears (cougars and jaguars may have been temporarily reduced to South American ranges also).
Others, such as capybaras, survived in their original range but died out in areas they had migrated to.
Notably, this extinction pulse eliminated all Neotropic migrants to North America larger than about 15 kg (the
size of a big porcupine), and all native South American mammals larger than about 65 kg (the size of a big
capybara or giant anteater). In contrast, the largest surviving native North American mammal, the wood
bison, can exceed 900 kg, and the largest surviving Nearctic migrant to South America, Baird's tapir, can
reach 400 kg.
The near-simultaneity of the megafaunal extinctions with the glacial retreat and the peopling of the
Americas has led to proposals that both climate change and human hunting played a role. Although the
subject is contentious,[41][42][43][44][45] a number of considerations suggest that human activities were
pivotal.[46][47] The extinctions did not occur selectively in the climatic zones that would have been most
affected by the warming trend, and there is no plausible general climate-based megafauna-killing mechanism
that could explain the continent-wide extinctions. The climate change took place worldwide, but had little
effect on the megafauna in areas like Africa and southern Asia, where megafaunal species had coevolved
with humans. Numerous very similar glacial retreats had occurred previously within the ice age of the last
several Ma without ever producing comparable waves of extinction in the Americas or anywhere else. Similar
megafaunal extinctions have occurred on other recently populated land masses (e.g. Australia, Madagascar,
New Zealand, and many smaller islands around the world, such as Cyprus, Crete, Tilos and New Caledonia)
at different times that correspond closely to the first arrival of humans at each location. These extinction
pulses invariably swept rapidly over the full extent of a contiguous land mass, regardless of whether it was an
island or a hemisphere-spanning set of connected continents. This was true despite the fact that all the larger
land masses involved (as well as many of the smaller ones) contained multiple climatic zones that would
have been affected differently by any climate changes ongoing at the time. However, on sizable islands far
enough offshore from newly occupied territory to escape immediate human colonization, megafaunal species
sometimes survived for many thousands of years after they became extinct on the mainland; examples
include meiolaniid turtles on Lord Howe Island and New Caledonia, ground sloths on the Antilles,[48] Steller's
sea cows off the Commander Islands[49] and woolly mammoths on Wrangel Island[50] and Saint Paul
Island.[51] The glacial retreat may have played a primarily indirect role in the extinctions in the Americas by
simply facilitating the movement of humans southeastward from Beringia down to North America. The reason
that a number of groups went extinct in North America but lived on in South America (while there are no
examples of the opposite pattern) appears to be that the dense rainforest of the Amazon basin and the high
peaks of the Andes provided environments that afforded a degree of protection from human predation.[l][note
6]
[edit] South American invasions of North America exclusive of Central

America
Dasypus novemcinctus
†Glyptotherium
The ground sloth †Eremotherium

Extant or extinct (†) North American taxa whose ancestors migrated out of South America:[note 7]
• Cichlids (Cichlidae: e.g. Herichthys cyanoguttatus) – freshwater fish that often tolerate
brackish conditions
• Bufonid Toads (Bufo)[52]
• Hylid Frogs[53]
• Leptodactylid Frogs[54] – as far north as Texas
• Microhylid Frogs[52]
• Virginia Opossum (Didelphis virginiana)
• Armadillos (Dasypus novemcinctus, †D. bellus)
• †Pachyarmatherium leiseyi, an enigmatic armored armadillo relative
• †Pampatheres (Plaina,[55] Holmesina) – large armadillo-like animals
• †Glyptodontids (Glyptotherium)
• †Megalonychid Ground Sloths[note 8] (Pliometanastes, Megalonyx)
• †Mylodontid Ground Sloths (Thinobadistes, Glossotherium,[55] Paramylodon)
• †Megatheriid Ground Sloths (Eremotherium, Nothrotheriops)
• New World porcupines (Erethizon dorsatum, †Erethizon poyeri, †E. kleini)
• Capybaras (†Neochoerus pinckneyi, †N. aesopi)
• Vampire Bats (†Desmodus stocki, †D. archaeodaptes)
• †Dire Wolf (Canis dirus) – which evolved from earlier North American migrants
• Cougar (Puma concolor) – returning from a South American refugium after North American
cougars were extirpated in the Pleistocene extinctions[56]
• †Terror Birds (Phorusrhacidae: Titanis walleri)
• Hummingbirds (Trochilidae)
• Tanagers (Thraupidae) – descended from earlier (perhaps Miocene) North American
migrants[57]
• Tyrant Flycatchers (Tyrannidae)
• Parrots (Arini: Rhynchopsitta pachyrhyncha, †Conuropsis carolinensis)
[edit] South American invasions that failed to penetrate beyond
Central America
Oophaga pumilio
†Mixotoxodon
Tinamus major
Extant or extinct (†) Central American taxa[note 9] whose ancestors migrated out of South America:
[note 7]
• Gonyleptid Harvestmen (Opiliones: Gonyleptidae)
• Electric Knifefishes (Gymnotiformes)
• Caeciliid Caecilians (Caecilia, Dermophis, Gymnopis, Oscaecilia) – snake-like
amphibians[58]
• Poison Dart Frogs (Dendrobatidae)[59]
• Boine Boas (Boidae: Boinae)
• Spectacled Caiman (Caiman crocodilus)[60]
• other Opossums (Didelphidae) – 11 additional extant species, listed on discussion page
• Northern Naked-tailed Armadillo (Cabassous centralis)
• Hoffmann's Two-toed Sloth (Megalonychidae: Choloepus hoffmanni)
• Three-toed Sloths (Bradypodidae: Bradypus variegatus, B. pygmaeus)
• Silky Anteater (Cyclopedidae: Cyclopes didactylus)
• other Anteaters (Myrmecophagidae: Myrmecophaga tridactyla,[o] Tamandua mexicana)
• Rothschild's and Mexican Hairy Dwarf Porcupines (Coendou rothschildi, Sphiggurus
mexicanus)
• other Caviomorph Rodents (Caviomorpha) – 9 additional extant species, listed on discussion
page
• Platyrrhine Monkeys (Platyrrhini) – at least 8 extant species, listed on discussion page[c]
• †Mixotoxodon larensis – a rhino-sized toxodontid notoungulate[n]
• other Vampire Bats (Desmodontinae) – all 3 extant species
• Toucans (Ramphastidae)
• Tinamous (Tinamidae)
• Great Curassow (Crax rubra)
[edit] North American invasions of South America

The camelid Vicugna vicugna
†Cuvieronius, a gomphothere
The coati Nasua nasua

The saber-toothed cat †Smilodon
Extant or extinct (†) South American taxa whose ancestors migrated out of North America
(considered as including Central America):[note 7]
• Lungless Salamanders[d][58] (Bolitoglossa,[61] Oedipina) – only present in northern South
America
• Ranid Frogs[52] – only present in northern South America
• Emydid Turtles (Trachemys) – only present in northern South America
• Geoemydid Turtles (Rhinoclemmys)[62] – only present in northern South America
• Coral Snakes (Leptomicrurus, Micrurus)[63][64]
• South American Rattlesnake (Crotalus durissus)[65]
• Lanceheads (Bothrops)
• Bushmasters (Lachesis)
• other Pit Vipers (Bothriechis schlegelii, Bothriopsis, Porthidium)[66]
• Small-eared Shrews (Cryptotis) – only present in NW South America: Colombia, Venezuela,
Ecuador, Peru
• Geomyid Pocket Gophers (Orthogeomys thaeleri) – one species, in Colombia
• Heteromyid Mice (Heteromys) – only present in NW South America: Colombia, Venezuela,
Ecuador
• Cricetid – primarily Sigmodontine – Rats and Mice (Cricetidae: Sigmodontinae)
• Tree Squirrels (Sciurus, Microsciurus, Sciurillus)
• Cottontail Rabbits (Sylvilagus brasiliensis, S. floridanus, S. varynaensis) – present in northern
and central South America
• Tapirs (Tapirus bairdii, T. pinchaque, T. terrestris)
• Equids (Equus ferus, †Hippidion)[e]
• Peccaries (Tayassu pecari, Catagonus wagneri, Pecari tajacu, P. maximus)
• Deer (Odocoileus, Blastocerus, Ozotoceros, Mazama, Pudu, Hippocamelus)
• Camelids (Lama guanicoe, Vicugna vicugna, †Eulamaops, †Hemiauchenia, †Palaeolama)
• †Gomphotheres (Cuvieronius hyodon, Stegomastodon[f] waringi, S. platensis)[40][note 10] –
elephant relatives
• Otters (Lontra, Pteronura)
• other Mustelids (Mustelinae: Eira, Galictis, Lyncodon, Mustela)
• Hog-Nosed Skunks (Conepatus chinga, C. humboldtii, C. semistriatus)
• Procyonids (Procyon, Nasua, Nasuella, Potos, Bassaricyon, †Cyonasua, †Chapalmalania)
• Short-Faced Bears (Tremarctinae: Tremarctos ornatus, †Arctotherium)[67]
• Wolves (†Canis ambrusteri, †C. dirus)
• Gray Fox[note 11] (Urocyon cinereoargenteus) – only present in NW South America:
Colombia, Venezuela
• other Canids (Atelocynus, Cerdocyon, Lycalopex, Chrysocyon, Speothos)
• small Felids (Leopardus) – all 9 extant species (e.g. L. pardalis, L. wiedii)
• Cougar (Puma concolor) and Jaguarundi (P. yagouaroundi)
• Jaguar (Panthera onca)
• †American Lion (Panthera leo atrox)
• †Scimitar Cats (Homotherium) – known so far only from Venezuela[68][69]
• †Saber-Toothed Cats (Smilodon gracilis, S. fatalis, S. populator)
• Condors (Vultur gryphus, †Dryornis, †Geronogyps, †Wingegyps, †Perugyps)[70][71][72][g]
• Trogons (Trogon)[73]
[edit] See also

• Columbian Exchange
• List of North American mammals
• Mammals of the Caribbean
• List of Central American mammals
• List of South American mammals
• Lists of birds by region
• Lists of reptiles by region
• Lists of amphibians by region
• List of prehistoric mammals
• List of extinct animals of North America
[edit] Notes
1. ^ The entirety of volume 1 and volume 2 of Wallace's book The Geographical Distribution of
Animals is also available online from Google Books.
2. ^ Ziphodont (lateromedially compressed, recurved and serrated) teeth tend to arise in
terrestrial crocodilians because, unlike their aquatic cousins, they are unable to dispatch their prey by
simply holding them underwater and drowning them; they thus need cutting teeth with which to slice
open their victims.
3. ^ North American gopher tortoises are most closely related to the Asian genus Manouria.
4. ^ Of the 6 families of North American rodents that did not originate in South America, only
beavers and mountain beavers failed to migrate to South America. (However, introduced beavers
have become serious pests in Tierra del Fuego.)
5. ^ Of the 11 extant families of South American caviomorph rodents, 5 are present in Central
America; only 2 of these, Erethizontidae and Caviidae, ever reached North America. (The
nutria/coypu has been introduced to a number of North American locales.)
6. ^ A number of recently extinct North American (and in some cases also South American)
taxa such as tapirs, equids, camelids, saiga antelope, proboscids, dholes and lions survived in the
Old World, probably mostly for different reasons – tapirs being a likely exception, since their Old
World representative survived only in the rainforests of Southeast Asia. (Cheetahs in the broadest
sense could be added to this list, although the New and Old World forms are in different genera.) Old
World herbivores may in many cases have been able to learn to be vigilant about the presence of
humans during a more gradual appearance (by development or migration) of advanced human
hunters in their ranges. In the cases of predators, the Old World representatives in at least some
locations would thus have suffered less from extinctions of their prey species. In contrast, the musk
ox represents a rare example of a megafaunal taxon that recently went extinct in Asia but survived in
remote areas of arctic North America (its more southerly-distributed relatives such as Harlan's musk
ox and the shrub ox were less fortunate).
7. ^ a b c This listing currently has fairly complete coverage of nonflying mammals, but only
spotty coverage of other groups. Crossings by nonflying mammals and birds occurred during the last
10 Ma. Crossings by fish, arthropods, waif-dispersing amphibians and reptiles, and flying bats and
birds were made before 10 Ma ago in many cases. Taxa listed as invasive did not necessarily cross
the isthmus themselves; they may have evolved in the adopted land mass from ancestral taxa that
made the crossing.
8. ^ While all megalonychid ground sloths are extinct, extant two-toed tree sloths are from the
same family. Three-toed tree sloths, in contrast, are not closely related to any of the groups of extinct
ground sloths.
9. ^ For the purposes of this article, all northwardly migrating Neotropic taxa that failed to reach
the territory of the continental U.S. will be treated as having only reached Central America. While
Central America is usually defined physiographically as ending at the Isthmus of Tehuantepec, or
less commonly, at the Trans-Mexican Volcanic Belt, most of the taxa that proceeded further but failed
to reach the present Mexican border are or were confined to tropical or subtropical ecozones similar
to those of Central America. Examples include the giant anteater, the grayish mouse opossum, the
lowland paca, Geoffroy's spider monkey and Mixotoxodon.
10.^ Sometimes classified as elephantids rather than as gomphotheres.
11.^ Not to be confused with the South American gray fox.
a. ^ An alternative explanation blames climatic and physiographic changes associated with the
uplift of the Andes.[14]
b. ^ According to data on the EQ (encephalization quotient, a measure of the brain to body size
ratio adjusted for the expected effect of differences in body size) of fossil ungulates compiled by H.
Jerison,[74] North American ungulates showed a trend towards greater EQs going from the
Paleogene to the Neogene periods (average EQs of 0.43 and 0.64, respectively), while the EQs of
South American ungulates were static over the same time interval (average EQ unchanged at 0.48).
[5] This analysis was later criticized.[75] Jerison subsequently presented data suggesting that native
South American ungulates also lagged in the relative size of their neocortex (a measurement not
subject to the vagaries of body mass estimation).[76] It is interesting to note that the late survivor
Toxodon had one of the highest EQ values (0.88) among native Neotropic ungulates.[75]
Jerison also found that Neogene xenarthrans had low EQs, similar to those he obtained for South
American ungulates.[74]
The estimated EQ of Thylacosmilus atrox, 0.41 (based on brain mass = 43.2 g, body mass = 26.4 kg,
[77] and EQ = 43.2/[0.12*26400^(2/3)][76]), is high for a sparassodont,[78] but is lower than that of
modern felids, with a mean value of 0.87.[79] Estimates of 0.38[80] and 0.59[79] have been given for
the EQ of much larger Smilodon fatalis (based on body mass estimates of 330 and 175 kg,
respectively).
c. ^ It has been proposed that monkeys invaded Central America in at least three and probably
four waves, as follows: (1) an initial invasion by A. pigra and S. oerstedii ~ 3 Ma ago; (2) an invasion
by A. palliata (giving rise to A. coibensis), A. geoffroyi and C. capucinus ~ 2 Ma ago; an invasion by
A. zonalis and S. geoffroyi ~ 1 Ma ago; a most recent invasion by A. fusciceps. The species of the
first wave have apparently been out-competed by those of the second, and now have much more
restricted distributions.[81]
d. ^ Salamanders may have dispersed to South America more than 10 Ma ago. Nevertheless,
the salamander fauna of South America, which is restricted to the tropical region, consists of only 2
clades, and has fewer species and is far less diverse than that of much smaller Central America.
Salamanders are believed to have originated in northern Pangea, perhaps not long before it
separated to become Laurasia,[58] and are not present anywhere else in the southern hemisphere
(see the world salamander distribution map). In contrast, caecilians have a mostly Gondwanan
distribution. Apart from a small region of overlap in southern China and northern Southeast Asia,
Central America and northern South America are the only places in the world where salamanders
and caecilians are both present.
e. ^ Hippidion, a relatively short-legged equid that developed in South America after invading
from North America about 2.5 Ma ago, has traditionally been thought to have evolved from
pliohippines.[82][83] However, recent studies of the DNA of Hippidion and other New World
Pleistocene horses indicate that Hippidion is actually a member of Equus, closely related to the
extant horse, E. ferus.[82][83] Another invasion of South America by Equus occurred about one Ma
ago, and this lineage, traditionally viewed as the subgenus Equus (Amerhippus), appears
indistinguishable from E. ferus.[83] Both these lineages went extinct at the end of the Pleistocene,
but E. ferus was reintroduced from Eurasia by Europeans in the 16th century. Note: the authors of
the DNA sequence study of Equus (Amerhippus) use "E. caballus" as an alternative specific name for
"E. ferus".[83]
f. ^ Not to be confused with the American mastodon (†Mammut americanum), a proboscid from
a different family whose remains have been found no further south than Honduras.[84]
g. ^ Condors apparently reached South America by the late Miocene or early Pliocene (4.5 – 6.0
Ma ago), several million years before the formation of the isthmus.[72] Condor-like forms in North
America date back to the Barstovian stage (middle Miocene, 11.8 – 15.5 Ma ago).[71]
h. ^ The native South American ungulates dwindled gradually as North American ungulates
invaded and diversified. The changes in number and composition of South America's ungulate
genera over time are given in the table below.[85] The Quaternary extinction event that delivered the
coup de grâce to the native Neotropic ungulates also dealt a heavy blow to South America's ungulate
immigrants.
Change in number of South American ungulate genera over time
Time interval Source region of genera
Geologic period Range (Ma ago) South America North America
Huayquerian 9.0 — 6.8 13 0
Montehermosan 6.8 — 4.0 12 1
Chapadmalalan 4.0 — 3.0 12 1
Uquian 3.0 — 1.5 5 10
Ensenadan 1.5 — 0.8 3 14
Lujanian 0.8 — 0.011 3 20
Holocene 0.011 — 0 0 11
i. ^ The dog-like borhyaenids were already in decline prior to the main pulse of the interchange,
at a time when Thylacosmilus and phorusrhacids were still common. Suggested reasons for this
decline include competition with phorusrhacids, carnivorous oppossums, or early-arriving procyonids.
[86] However, it is clear that the remaining sparassodonts and most of the phorusrhacids ( Titanis
being an exception) disappeared quickly once canids and felids reached South America.
j. ^ During the Miocene alone, between about 23 and 5 Ma ago, 11 episodes of invasions of
North America from Eurasia have been recognized, bringing a total of 81 new genera into North
America.[28]
k. ^ Including extinct genera, South America has hosted 9 genera of cervids, 9 genera of
mustelids (if skunks are retained in Mustelidae, 8 if not), and 10 genera of canids. However, some of
this diversity of South American forms apparently arose in North or Central America prior to the
interchange.[28]
l. ^ P. S. Martin (2005), p. 175.[46]
m. ^ P. S. Martin (2005), pp. 30–37, 119.[46] The figure of 25 South American megaherbivore
species breaks down as follows: 4 gomphotheres, 2 camelids, 9 ground sloths, 5 glyptodontids, and
5 toxodontids. This can be compared to Africa's present and recent total of 6 megaherbivores: 1
giraffe, 1 hippo, 2 rhinos and 2 elephants (considering the African forest elephant as a separate
species).
n. ^ Mixotoxodon remains have been identified from as far north as Veracruz and Michoacán,
Mexico, with a possible find in Tamaulipas.[87]
o. ^ Fossils of the giant anteater have been found as far north as northwestern Sonora, Mexico.
[88]
[edit] References
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• History of the Fauna of Latin America by George Gaylord Simpson (1950)
Retrieved from "http://en.wikipedia.org/wiki/Great_American_Interchange"
Categories: Biogeography | Natural history of Central America | Natural history of North America |
Natural history of South America | Natural history of the Americas
W000
Hedgehog signaling pathway


In a growing embryo, cells develop differently in the head or tail end of the embryo, the left or right,
and other positions. They also form segments which develop into different body parts. The hedgehog
signaling pathway gives cells information that they need to make the embryo develop properly. Different
parts of the embryo have different concentrations of hedgehog signaling proteins. The pathway also has
roles in the adult. When the pathway malfunctions, it can result in diseases like basal cell carcinoma. [1]
The hedgehog signaling pathway is one of the key regulators of animal development conserved from
flies to humans (meaning it was present in the common ancestor of both). The pathway takes its name from
its polypeptide ligand, an intercellular signaling molecule called Hedgehog ( Hh) found in fruit flies of the
genus Drosophila. Hh is one of Drosophila's segment polarity gene products, involved in establishing the
basis of the fly body plan. The molecule remains important during later stages of embryogenesis and
metamorphosis.
Mammals have three Hedgehog homologues, of which Sonic hedgehog is the best studied. The
pathway is equally important during vertebrate embryonic development. In knockout mice lacking
components of the pathway, the brain, skeleton, musculature, gastrointestinal tract and lungs fail to develop
correctly. Recent studies point to the role of hedgehog signaling in regulating adult stem cells involved in
maintenance and regeneration of adult tissues. The pathway has also been implicated in the development of
some cancers. Drugs that specifically target hedgehog signaling to fight this disease are being actively
developed by a number of pharmaceutical companies.
Contents
[hide]
• 1 Discovery
• 2 Fruit fly
• 2.1 Mechanism
• 2.2 Role
• 3 Annelids
• 4 Vertebrates
• 4.1 Mechanism
• 4.2 Role
• 4.3 Human disease
• 5 Evolution
• 6 See also
• 8 References
[edit] Discovery
Figure 1. Normal and Hedgehog mutant larvae.

In 1995 Edward B. Lewis, Christiane Nüsslein-Volhard and Eric F. Wieschaus were awarded a Nobel
Prize for their work studying genetic mutations in Drosophila embryogenesis.[2] In the 1970s, a fundamental
problem in developmental biology was to understand how a relatively simple egg can give rise to a complex
segmented body plan. Nüsslein- Volhard and Wieschaus attempted to isolate mutations in genes that control
development of the segmented anterior-posterior body axis of the fly;[3] their "saturation mutagenesis"
technique resulted in the discovery of a group of genes involved in the development of body segmentation.
The Drosophila hedgehog gene was identified as one of several genes important for creating the
differences between the anterior and posterior parts of individual body segments. Some hedgehog mutants
result in abnormally-shaped embryos that are unusually short and stubby compared to wild type embryos.
The function of the hedgehog segment polarity gene has been studied in terms of its influence on the
normally polarized distribution of larval cuticular denticles as well as features on adult appendages such as
legs and antennae.[4] Rather than the normal pattern of denticles, hedgehog mutant larvae tend to have
"solid lawns" of denticles (Figure 1). The appearance of the stubby and "hairy" larvae inspired the name
'hedgehog' (see: Hedgehog, the animal).
[edit] Fruit fly
Figure 2. Production of the CiR transcriptional repressor when Hh is not bound to Patched. In the
diagram, "P" represents phosphate.
Figure 3. When Hh is bound to Patched (PTCH), Ci protein is able to act as a transcription factor in
the nucleus.
[edit] Mechanism
Insect cells express a full size zinc-finger transcription factor Cubitus interruptus (Ci), which forms a
complex with the kinesin- like protein Costal-2 (Cos2) and is localized in the cytoplasm bound to cellular
microtubules (Figure 2). The complex targets the 155 kb full length Ci protein for proteosome- dependent
cleavage, which generates a 75 kb fragment (CiR). CiR builds up in the cell and diffuses into the nucleus,
where it acts as a co-repressor for Hh target genes[5]. The steps leading to Ci protein proteolysis include
phosphorylation of Ci protein by several protein kinases; PKA, GSK3β and CK1 (Figure 2)[6]. The Drosophila
protein Slimb is part of an SCF complex that targets proteins for ubiquitylation. Slimb binds to phosphorylated
Ci protein.
In the absence of Hh (Figure 3), a cell-surface transmembrane protein called Patched (PTCH) acts to
prevent high expression and activity of a 7 membrane spanning receptor[7] called Smoothened (SMO).
Patched has sequence similarity to known membrane transport proteins. When extracellular Hh is present
(Figure 3), it binds to and inhibits Patched, allowing Smoothened to accumulate and inhibit the proteolytic
cleavage of the Ci protein. This process most likely involves the direct interaction of Smoothened and Costal-
2 and may involve sequestration of the Ci protein-containing complex to a microdomain where the steps
leading to Ci protein proteolysis are disrupted[5]. The mechanism by which Hh binding to Patched leads to
increased levels of Smoothened is not clear (Step 1 in Figure 3). Following binding of Hh to Patched,
Smoothened levels increase greatly over the level maintained in cells when Patched is not bound to Hh[8]. It
has been suggested that phosphorylation of Smoothened plays a role in Hh-dependent regulation of
Smoothened levels[9].
In cells with Hh-activated Patched (Figure 3), the intact Ci protein accumulates in the cell cytoplasm
and levels of CiR decrease, allowing transcription of some genes such as decapentaplegic (dpp, a member
of the BMP growth factor family). For other Hh-regulated genes, expression requires not only loss of CiR but
also the positive action of uncleaved Ci acting as a transcriptional activator [6]. Costal-2 is normally important
for holding Ci protein in the cytoplasm, but interaction of Smoothened with Costal-2 allows some intact Ci
protein to go to the nucleus. The Drosophila protein Fused (Fu in Figure 3) is a protein kinase that binds to
Costal-2. Fused can inhibit Suppressor of Fused (SUFU), which in turn interacts with Ci to regulate gene
transcription in some cell types[10].
[edit] Role
Figure 4. Interactions between Wingless and Hedgehog.

Hedgehog has roles in larval body segment development and in formation of adult appendages.
During the formation of body segments in the developing Drosophila embryo, stripes of cells that synthesize
the transcription factor Engrailed can also express the cell-to-cell signaling protein Hedgehog (green in
Figure 4). Hedgehog is not free to move very far from the cells that make it and so it only activates a thin
stripe of cells adjacent to the Engrailed-expressing cells. Only cells to one side of the Engrailed-expressing
cells are competent to respond to Hedgehog following interaction of Hh with the receptor protein Patched
(blue in Figure 4).
Cells with Hh-activated Patched receptor synthesize the Wingless protein (red in Figure 4). If a
Drosophila embryo is altered so as to produce Hh in all cells, all of the competent cells respond and form a
broader band of Wingless-expressing cells in each segment. The wingless gene has an upstream
transcription regulatory region that binds the Ci transcription factor in a Hh-dependent fashion resulting in an
increase in wingless transcription (interaction 2 in Figure 3) in a stripe of cells adjacent to the stripe of Hh-
producing cells[11].
Wingless protein acts as an extracellular signal and patterns the adjacent rows of cells by activating
its cell surface receptor Frizzled. Wingless acts on Engrailed-expressing cells to stabilize the stripes of
Engrailed expression. Wingless is a member of the Wnt family of cell-to-cell signaling proteins. The
reciprocal signaling by Hedgehog and Wingless stabilizes the boundary between parasegments (Figure 4,
top). The effects of Wingless and Hedgehog on other stripes of cells in each segment establishes a positional
code that accounts for the distinct anatomical features along the anterior-posterior axis of the segments [12].
The Wingless protein is called "wingless" because of the phenotype of some wingless fly mutants.
Wingless and Hedgehog functioned together during metamorphosis to coordinate wing formation. Hedgehog
is expressed in the posterior part of developing Drosophila limbs. Hedgehog also participates in the
coordination of eye, brain, gonad, gut and tracheal development.
[edit] Annelids
hedgehog is also involved in segmentation in the annelid worms; because parallel evolution seems
unlikely, this suggests a common origin of segmentation between the two phyla.[13] Whilst Hh does not
induce the formation of segments, it seems to act to stabilize the segmented fields once they have appeared.
[13]
[edit] Vertebrates
[edit] Mechanism
Figure 5. Overview of Sonic hedgehog signaling. Click here for a more detailed diagram
Sonic hedgehog (SHH) is the best studied ligand of the vertebrate pathway. Most of what is known
about hedgehog signaling has been established by studying SHH. It is translated as a ~45kDa precursor and
undergoes autocatalytic processing to produce an ~20kDa N-terminal signaling domain (referred to as SHH-
N) and a ~25kDa C-terminal domain with no known signaling role (1 on figure 5). During the cleavage, a
cholesterol molecule is added to the carboxyl end of the N-terminal domain, which is involved in trafficking,
secretion and receptor interaction of the ligand. SHH can signal in an autocrine fashion, affecting the cells in
which it is produced. Secretion and consequent paracrine hedgehog signaling require the participation of
Dispatched protein(2).
When SHH reaches its target cell, it binds to the Patched-1 (PTCH1) receptor(3). In the absence of
ligand, PTCH1 inhibits Smoothened (SMO), a downstream protein in the pathway(4). It has been suggested
that SMO is regulated by a small molecule, the cellular localisation of which is controlled by PTCH[14].
PTCH1 has homology to Niemann-Pick disease, type C1 (NPC1) that is known to transport lipophilic
molecules across a membrane. [15] PTCH1 has a sterol sensing domain (SSD), which has been shown to be
essential for suppression of Smo activity. [16] A current theory suggests that PTCH regulates SMO by
removing oxysterols from SMO. PTCH acts like a sterol pump and removes oxysterols that have been
created by 7-dehydrocholesterol reductase. [17] Upon binding of a Hh protein or a mutation in the SSD of
PTCH the pump is turned off allowing oxysterols to accumulate around SMO.
Suggested regulation pathway for Smo via Hedgehog and Ptch1
This accumulation of sterols allows SMO to become active or stay on the membrane for a longer
period of time. This hypothesis is supported by the existence of a number of small molecule agonists and
antagonists of the pathway that act on SMO. The binding of SHH relieves SMO inhibition, leading to
activation of the GLI transcription factors(5): the activators Gli1 and Gli2 and the repressor Gli3. The
sequence of molecular events that connect SMO to GLIs is poorly understood. Activated GLI accumulates in
the nucleus(6) and controls the transcription of hedgehog target genes(7). PTCH1 has recently been
reported to repress transcription of hedgehog target genes through a mechanism independent of
Smoothened.[18]
In addition to PTCH1, mammals have another hedgehog receptor PTCH2 whose sequence identity
with PTCH1 is 54%.[19] All three mammalian hedgehogs bind both receptors with similar affinity, so PTCH1
and PTCH2 cannot discriminate between the ligands. They do, however, differ in their expression patterns.
PTCH2 is expressed at much higher levels in the testis and mediates desert hedgehog signaling there.[19] It
appears to have a distinct downstream signaling role from PTCH1. In the absence of ligand binding PTCH2
has a decreased ability to inhibit the activity of SMO.[20] Furthermore, overexpression of PTCH2 does not
replace mutated PTCH1 in basal cell carcinoma.[21]
In invertebrates, just as in Drosophila, the binding of hedgehog to PTCH leads to internalisation and
sequestration of the ligand.[22] Consequently in vivo the passage of hedgehog over a receptive field that
expresses the receptor leads to attenuation of the signal, an effect called ligand-dependent antagonism
(LDA). In contrast to Drosophila, vertebrates possess another level of hedgehog regulation through LDA
mediated by Hh-interacting protein 1 (HHIP1). HHIP1 also sequesters hedgehog ligands, but unlike PTCH, it
has no effect on the activity of SMO.[23]
[edit] Role
Members of the hedgehog family play key roles in a wide variety of developmental processes.[12]
One of the best studied examples is the action of Sonic hedgehog during development of the vertebrate limb.
The classic experiments of Saunders and Gasseling in 1968 on the development of the chick limb bud
formed the basis of the morphogen concept. They showed that identity of the digits in the chick limb was
determined by a diffusible factor produced by the zone of polarizing activity (ZPA), a small region of tissue at
the posterior margin of the limb. Mammalian development appeared to follow the same pattern. This
diffusible factor was later shown to be Sonic hedgehog. However, precisely how SHH determines digit
identity remained elusive until recently. The current model, proposed by Harfe et al.[24], states that both the
concentration and the time of exposure to SHH determines which digit the tissue will develop into in the
mouse embryo (figure 6).
Figure 6. Sonic hedgehog specifies digit identity in mammalian development.

Digits V, IV and part of III arise directly from cells that express SHH during embryogenesis. In these
cells SHH signals in an autocrine fashion and these digits develop correctly in the absence of DISP, which is
required for extracellular diffusion of the ligand. These digits differ in the length of time that SHH continues to
be expressed. The most posterior digit V develops from cells that express the ligand for the longest period of
time. Digit IV cells express SHH for a shorter time, and digit III cells shorter still. Digit II develops from cells
that are exposed to moderate concentrations of extracellular SHH. Finally, digit I development does not
require SHH. It is, in a sense, the default program of limb bud cells.
Hedgehog signaling remains important in the adult. Sonic hedgehog has been shown to promote the
proliferation of adult stem cells from various tissues, including primitive hematopoietic cells[25],
mammary[26] and neural[27] stem cells. Activation of the hedgehog pathway is required for transition of the
hair follicle from the resting to the growth phase.[28] Curis Inc. together with Procter & Gamble are
developing a hedgehog agonist to be used as a drug for treatment of hair growth disorders.[29] This failed
due to toxicities found in animal models.[30]
[edit] Human disease

Disruption of hedgehog signaling during embryonic development, either through deleterious mutation
or consumption of teratogens by the gestating mother, can lead to severe developmental abnormalities.
Holoprosencephaly, the failure of the embryonic prosencephalon to divide to form cerebral hemispheres,
occurs with a frequency of about 1 in 16,000 live births and about 1 in 200 spontaneous abortions in humans
and is commonly linked to mutations in genes involved in the hedgehog pathway, including SHH and PTCH.
[31] Cyclopia, one of the most severe defects of holoprosencephaly, results if the pathway inhibitor
cyclopamine is consumed by gestating mammals.[32]
Activation of the hedgehog pathway has been implicated in the development of cancers in various
organs, including brain, lung, mammary gland, prostate and skin. Basal cell carcinoma, the most common
form of cancerous malignancy, has the closest association with hedgehog signaling. Loss-of-function
mutations in Patched and activating mutations in Smoothened have been identified in patients with this
disease.[33] Abnormal activation of the pathway probably leads to development of disease through
transformation of adult stem cells into cancer stem cells that give rise to the tumor. Cancer researchers hope
that specific inhibitors of hedgehog signaling will provide an efficient therapy for a wide range of
malignancies.[34]
Biotech companies are also attempting to turn this pathway on after a patient has a stroke or heart
attack. Since the pathway has been implicated in a number of lethal cancers Curis and Wyeth have devised a
stable hedgehog protein that can cross the blood brain barrier.[35] In pre-clinical animal models it has shown
that the pathway is up regulated upon a stroke or heart attack event. The pathway provides a protective
barrier against cell death and ischemia. Agonizing the pathway this way allows the PTCH to be up regulated
providing a negative feedback system. This might help minimize the side effects.
Targeting the Hedgehog Pathway
The most common way to target this pathway is modulate SMO. Antagonist and agonist of SMO
have already shown to effect the pathway regulation downstream. PTCH[36] and Gli3 (5E1)[37] antibodies
are also a way to regulate the pathway. A downstream effector and strong transcriptional activator siRNA
Gli1 has been used to inhibit cell growth and promote apoptosis[38].
Hedgehog Pathway and Metastasis

Activation of the Hedgehog pathway leads to an increase in Snail protein expression and a decrease
in E-cadherin and Tight Junctions. [39]. Hedgehog signaling also appears to be a crucial regulator of
angiogenesis and thus metastasis. [40].
Hedgehog Pathway and Tumor Regulation
Activation of the Hedgehog pathway leads to an increase in Angiogenic Factors (angiopoietin-1 and
angiopoietin-2)[41], Cyclins (cyclin D1 and B1))[42], anti-apoptotic genes and a decrease in apoptotic genes
(Fas)[43].
Clinical Trials
GDC-0449 in Treating Patients With Locally Advanced or Metastatic Solid Tumors [2]
A Study of Systemic Hedgehog Antagonist With Concurrent Chemotherapy and Bevacizumab As First-Line
Therapy for Metastatic Colorectal Cancer[3]
Video Presentation from AACR.org [4]
[edit] Evolution
Figure 7. Phylogenetic relationship of hedgehog ligands (based on Ingham and McMahon, 2001).
Hedgehog-like genes, 2 Patched homologs and Patched-related genes exist in the worm C. elegans
[44][45]. These genes have been shown to code for proteins that have roles in C. elegans development[44].
The hedgehog-like and Patched-related gene families are very large and function without the need for a
Smoothened homolog, suggesting a distinct pattern of selection for cholesterol modification and sensing
mechanisms in coelomate and pseudo-coelomate lineages[45].
Lancelets, which are primitive chordates, possess only one homologue of Drosophila Hh (figure 7).
Vertebrates, on the other hand, have several hedgehog ligands that fall within three subgroups - desert,
Indian and sonic, each represented by a single mammalian gene. This is probably a consequence of the two
genome duplications that occurred early in the vertebrate evolutionary history.[46] Two such events would
have produced four homologous genes, one of which must have been lost. Desert hedgehogs are the most
closely related to Drosophila Hh. Additional gene duplications occurred within some species[12] such as the
zebrafish Danio rerio, which has an additional tiggywinkle hedgehog gene in the sonic group. Various
vertebrate lineages have adapted hedgehogs to unique developmental processes. For example, a
homologue of the X.laevis banded hedgehog is involved in regeneration of the salamander limb.[47]
shh has undergone accelerated evolution in the primate lineage leading to humans.[48] Dorus et al.
hypothesise that this allowed for more complex regulation of the protein and may have played a role in the
increase in volume and complexity of the human brain.
The frizzled family of WNT receptors have some sequence similarity to Smoothened [49]. However,
G proteins have been difficult to link to the function Smoothened. Smoothened seems to be a functionally
divergent member of the G protein coupled receptor super family. Other similarities between the WNT and
Hh signaling pathways have been reviewed[50]. Nusse observed that, "a signaling system based on lipid-
modified proteins and specific membrane translocators is ancient, and may have been the founder of the Wnt
and Hh signaling systems".
It has been suggested that invertebrate and vertebrate signaling downstream from Smoothened has
diverged significantly[51]. The role of Suppressor of Fused (SUFU) has been enhanced in vertebrates
compared to Drosophila where its role is relatively minor. Costal-2 is particularly important in Drosophila. The
protein kinase Fused is a regulator of SUFU in Drosophila, but may not play a role in the Hh pathway of
vertebrates[52]. In vertebrates, Hh signalling has been heavily coupled to cilia [53]
[edit] See also

• Sonic hedgehog, best studied ligand of the vertebrate pathway
• Smoothened, the conserved GPCR component of the pathway
• Cyclopamine, a small molecule inhibitor of Hh signaling

• http://hedgehog.sfsu.edu (Hedgehog Pathway Database)
• Netpath - A curated resource of signal transduction pathways in humans
[edit] References
1. ^ Kimball's Biology Pages, The Hedgehog Signaling Pathway
2. ^ 1995 Nobel Prize for discovery of the genetic control of early embryonic development
3. ^ Nusslein-Volhard C and Wieschaus E (1980) "Mutations affecting segment number and
polarity in Drosophila" in Nature Volume 287, pages 795-801.Entrez Pubmed 6776413
4. ^ Mohler J (1988) "Requirements for hedgehog, a segmental polarity gene, in patterning
larval and adult cuticle of Drosophila" in Genetics Volume 120, pages 1061-1072.Entrez Pubmed
3147217
5. ^ a b Collins RT and Cohen SM (2005). "A genetic screen in Drosophila for identifying novel
components of the hedgehog signaling pathway". Genetics 170 (1): 173–184.
doi:10.1534/genetics.104.039420. PMID 15744048. Entrez Pubmed 15744048
6. ^ a b Lum L and Beachy PA (2004). "The Hedgehog response network: sensors, switches,
and routers". Science 304 (5678): 1755–1759. doi:10.1126/science.1098020. PMID 15205520.
Entrez Pubmed 15205520
7. ^ Chen W, Ren XR, Nelson CD, Barak LS, Chen JK, Beachy PA, de Sauvage F and
Lefkowitz RJ (2004) "Activity-dependent internalization of smoothened mediated by beta-arrestin 2
and GRK2" in Science Volume 306, pages 2257-2260.Entrez Pubmed 15618519
8. ^ Alcedo J, Zou Y and Noll M (2000) "Posttranscriptional regulation of smoothened is part of
a self-correcting mechanism in the Hedgehog signaling system" in Molecular Cell Volume 6, pages
457-465. Entrez Pubmed 10983991
9. ^ Apionishev S, Katanayeva NM, Marks SA, Kalderon D and Tomlinson A (2005) "Drosophila
Smoothened phosphorylation sites essential for Hedgehog signal transduction" Nature Cell Biology
Volume 7, page 86-92. Entrez Pubmed 15592457
10.^ Ho KS, Suyama K, Fish M and Scott MP (2005) "Differential regulation of Hedgehog target
gene transcription by Costal2 and Suppressor of Fused" in Development Volume 132, pages 1401-
1412. Entrez Pubmed 15750186
11.^ Von Ohlen T, Lessing D, Nusse R and Hooper JE (1997) "Hedgehog signaling regulates
transcription through cubitus interruptus, a sequence-specific DNA binding protein" in Proceedings of
the National Academy of Sciences of the United States of America Volume 94, pages 2404-2409.
Entrez Pubmed 9122207
12.^ a b c Ingham, P.; McMahon, A. (2001). "Hedgehog signaling in animal development:
paradigms and principles". Genes & development 15 (23): 3059–3087. doi:10.1101/gad.938601.
PMID 11731473. edit Entrez Pubmed 11731473
Ingham and McMahon review the roles of hedgehog proteins in development; this table provides a
summary.
13.^ a b Dray, N.; Tessmar-Raible, K.; Le Gouar, M.; Vibert, L.; Christodoulou, F.; Schipany, K.;
Guillou, A.; Zantke, J. et al. (2010). "Hedgehog signaling regulates segment formation in the annelid
Platynereis". Science (New York, N.Y.) 329 (5989): 339–342. doi:10.1126/science.1188913.
PMID 20647470. edit
14.^ Taipale J, Cooper MK, Maiti T, and Beachy PA (2002). "Patched acts catalytically to
suppress the activity of Smoothened". Nature 418 (6900): 892–897. doi:10.1038/nature00989.
12192414. PMID 12192414.
15.^ Davies,J.P,J.P., Chen,F.W., and Ioannou,Y.A (2000). "Transmembrane molecular pump
activity of Niemann-Pick C1 protein.". Science 5500: 2295–2298. 1125140.
16.^ Strutt,H., Thomas,C., Nakano,Y., Stark,D., Neave,B., Taylor,A.M., and Ingham,P.W.
(2001). "Mutations in the sterol-sensing domain of Patched suggest a role for vesicular trafficking in
Smoothened regulation.". Curr Biol. 8: 608–613. 11369206.
17.^ Corcoran RB, Scott MP (2006). "Oxysterols stimulate Sonic hedgehog signal transduction
and proliferation of medulloblastoma cells.". Proc Natl Acad Sci USA 418: 8408–8413. 167075754.
18.^ Rahnama F, Shimokawa T, Lauth M, Finta C, Kogerman P, Teglund S, Toftgard R and
Zaphiropoulos PG (2006). "Inhibition of GLI1 gene activation by Patched1". Biochem J 394 (Pt 1): 19–
26. doi:10.1042/BJ20050941. PMID 16229683. Entrez Pubmed 16229683
19.^ a b Carpenter D, Stone DM, Brush J, Ryan A, Armanini M, Frantz G, Rosenthal A and de
Sauvage FJ (1998). "Characterization of two patched receptors for the vertebrate hedgehog protein
family". PNAS 95 (23): 13630–13634. doi:10.1073/pnas.95.23.13630. PMID 9811851. Entrez
Pubmed 9811851
20.^ Rahnama F, Toftgard R and Zaphiropoulos PG (2004). "Distinct roles of PTCH2 splice
variants in Hedgehog signalling". Biochem J 378(Pt 2): 325–334. Entrez Pubmed 14613484
21.^ Zaphiropoulos PG, Unden AB, Rahnama F, Hollingsworth RE and Toftgard R (1999).
"PTCH2, a novel human patched gene, undergoing alternative splicing and up-regulated in basal cell
carcinomas". Cancer Res 59 (4): 787–792. PMID 10029063. Entrez Pubmed 10029063
22.^ Incardona JP, Lee JH, Robertson CP, Enga K, Kapur RP and Roelink H (2000). "Receptor-
mediated endocytosis of soluble and membrane-tethered Sonic hedgehog by Patched-1". PNAS 97
(22): 12044–12049. doi:10.1073/pnas.220251997. PMID 11027307. Entrez Pubmed 11027307
23.^ Jeong J and McMahon AP (2005). "Growth and pattern of the mammalian neural tube are
governed by partially overlapping feedback activities of the hedgehog antagonists patched 1 and
Hhip1". Development 132 (1): 143–154. doi:10.1242/dev.01566. PMID 15576403. Entrez Pubmed
15576403
24.^ Harfe BD, Scherz PJ, Nissim S, Tian H, McMahon AP and Tabin CJ (2004). "Evidence for
an expansion-based temporal Shh gradient in specifying vertebrate digit identities". Cell 118 (4): 517–
528. doi:10.1016/j.cell.2004.07.024. PMID 15315763. Entrez Pubmed 15315763
25.^ Bhardwaj G, Murdoch B, Wu D, Baker DP, Williams KP, Chadwick K, Ling LE, Karanu FN
and Bhatia M (2001). "Sonic hedgehog induces the proliferation of primitive human haematopoetic
cells via BMP regulation". Nat Immunol 2 (2): 172–180. doi:10.1038/84282. PMID 11175816. Entrez
Pubmed 11175816
26.^ Liu S, Dontu G, Mantle ID, Patel S, Ahn NS, Jackson KW, Suri P and Wicha MS (2006).
"Hedgehog signaling and Bmi-1 regulate self-renewal of normal and malignant human mammary
stem cells". Cancer Res 66 (12): 6063–6071. doi:10.1158/0008-5472.CAN-06-0054.
PMID 16778178. Entrez Pubmed 16778178
27.^ Ahn S and Joyner AL (2005). "In vivo analysis of quiescent adult neural stem cells
responding to Sonic hedgehog". Nature 437 (7060): 894–897. doi:10.1038/nature03994.
28.^ Paladini RD, Saleh J, Qian C, Xu GX and Rubin LL (2005). "Modulation of hair growth with
small molecule agonists of the hedgehog signaling pathway". J Invest Dermatol 125 (4): 638–646.
doi:10.1111/j.0022-202X.2005.23867.x. PMID 16185261. Entrez Pubmed 16185261
29.^ http://www.curis.com/product_detail.php?id=5, retrieved on 25 June 2006.
30.^ [1], retrieved on May 9, 2007.
31.^ OMIM Holoprosencephaly, updated 16 May 2006 Online 'Mendelian Inheritance in Man'
(OMIM) 236100
32.^ Keeler RF (1978). "Cyclopamine and related steroidal alkaloid teratogens: their occurrence,
structural relationship, and biologic effects". Lipids 13 (10): 708–715. doi:10.1007/BF02533750.
33.^ Xie J, Murone M, Luoh SM, Ryan A, Gu Q, Zhang C, Bonifas JM, Lam CW, Hynes M,
Goddard A et al. (1998). "Activating Smoothened mutations in sporadic basal-cell carcinoma".
Nature 391 (6662): 90–92. doi:10.1038/34201. PMID 9422511. Entrez Pubmed 9422511
34.^ Chen JK, Taipale J, Young KE, Maiti T and Beachy PA (2002). "Small molecule modulation
of Smoothened activity". PNAS 99 (22): 14071–14076. doi:10.1073/pnas.182542899.
35.^ Curis pipeline details Stroke and Heart Attack
36.^ Nakamura M, Kubo M, Yanai K, Mikami Y, Ikebe M, Nagai S, Yamaguchi K, Tanaka M,
Katano M. (2007). "Anti-patched-1 antibodies suppress hedgehog signaling pathway and pancreatic
cancer proliferation.". Anticancer Res. (6A): 3743–7.. Entrez Pubmed 17970037
37.^ Hunt R, Bragina O, Drews M, Kasak L, Timmusk S, Valkna A, Kogerman P, Järvekülg L.
(2007). "Generation and characterization of mouse monoclonal antibody 5E1 against human
transcription factor GLI3.". Hybridoma (4): 231–40. Entrez Pubmed 17725385
38.^ Stecca B, Mas C, Ruiz i Altaba A. (2005). "Interference with HH-GLI signaling inhibits
prostate cancer.". Trends Mol Med. (5): 199–203. Entrez Pubmed 15882606
39.^ Xingnan Li, Wentao Deng, Clinton D. Nail, Sarah K. Bailey, Matthias H. Kraus, J. Michael
Ruppert, and Susan M. Lobo-Ruppert1 (2006). "Snail induction is an early response to Gli1 that
determines the efficiency of epithelial transformation". Oncogene. 99 (4): 609–621. Entrez Pubmed
16158046
40.^ vamsidhar velcheti, (2007). "Hedgehog signaling is a potent regulator of angiogenesis in
small cell lung cancer.". Med Hypotheses. 69 (4): 948–949. doi:10.1016/j.mehy.2007.02.021.
41.^ "Sonic hedgehog inversely regulates the expression of angiopoietin-1 and angiopoietin-2 in
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42.^ "Patched1 functions as a gatekeeper by promoting cell cycle progression.". Cancer Res. 99
(5): 2081–8.. 2006. Entrez Pubmed 16489008
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through regulation of Fas expression and apoptosis.". Cancer Res. 99 (5): 7545–52.. 2004. Entrez
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44.^ a b Zugasti O, Rajan J and Kuwabara PE (2005) "The function and expansion of the
Patched- and Hedgehog-related homologs in C. elegans" in Genome Research Volume 15, pages
1402-1410.Entrez Pubmed 16204193
45.^ a b Novatchkova M, Wildpaner M, Schweizer F and Eisenhaber F (2005) "PhyloDome--
visualization of taxonomic distributions of domains occurring in eukaryote protein sequence sets" in
Nucleic Acids Research Volume 33(Web Server issue):W121-5.Entrez Pubmed 15980439
46.^ Wada H, and Makabe K (2006). "Genome duplications of early vertebrates as a possible
chronicle of the evolutionary history of the neural crest". International Journal of Biological Science 2
(3): 133–141. 16763673.
47.^ Stark DR, Gates PB, Brockes JP and Ferretti P (1998). "Hedgehog family member is
expressed throughout regenerating and developing limbs". Dev Dyn 212 (3): 352–363. doi:10.1002/
(SICI)1097-0177(199807)212:3<352::AID-AJA3>3.0.CO;2-G. PMID 9671939. Entrez Pubmed
9671939
48.^ Dorus S, Anderson JR, Vallender EJ, Gilbert SL, Zhang L, Chemnick LG, Ryder OA, Li W
and Lahn BT (2006). "Sonic Hedgehog, a key development gene, experienced intensified molecular
evolution in primates". Human Molecular Genetics 15 (13): 2031–2037. doi:10.1093/hmg/ddl123.
16687440. PMID 16687440.
49.^ Graul RC and Sadee W (2001) "Evolutionary relationships among G protein-coupled
receptors using a clustered database approach" in AAPS PharmSci Volume 3 article E12.Entrez
Pubmed 11741263
50.^ Nusse R (2003) "Wnts and Hedgehogs: lipid-modified proteins and similarities in signaling
mechanisms at the cell surface" in Development Volume 130, pages 5297-5305. Entrez Pubmed
14530294
51.^ Varjosalo M, Li SP and Taipale J (2006) "Divergence of hedgehog signal transduction
mechanism between Drosophila and mammals" in Developmental cell Volume 10, pages 177-
186.Entrez Pubmed 16459297.
52.^ Chen MH, Gao N, Kawakami T and Chuang PT (2005) "Mice deficient in the fused homolog
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53.^ Huangfuand D and Anderson KV (2006) "Signaling from Smo to Ci/Gli: conservation and
divergence of Hedgehog pathways from Drosophila to vertebrates" in Development Volume 133,
pages 3-14. Entrez Pubmed 16339192.
[hide]v · d · eCell physiology: signal transduction / cell signaling
Signaling GPCR (Hedgehog, Wnt) · RTK (TGF beta, MAPK/ERK) · Notch · JAK-STAT ·
pathways Akt/PKB · Fas apoptosis · Hippo · PI3K/AKT/mTOR pathway · Integrin receptors
Hormones · Neurotransmitters/Neuropeptides ·
Receptor ligands
Cytokines · Growth factors
Receptors Cell surface · Intracellular · Co-receptor
Signal transducing adaptor protein: Scaffold

Agents
Intracellular protein
signaling P+Ps 2nd messenger: cAMP-dependent pathway ·
Ca2+ signaling · Lipid signaling · IP3/DAG pathway
Transcription General · Transcription preinitiation complex ·

factors TFIID, TFIIH
By location Intracrine · Autocrine · Juxtacrine · Paracrine · Endocrine
Other Signaling molecule · Ion channel gating · Mechanotransduction ·

concepts Phototransduction · Synaptic transmission
B trdu: iter (nrpl/grfl/cytl/horl), csrc (lgic, enzr, gprc, igsr, intg, nrpr/grfr/cytr), itra (adap, gbpr, mapk),
calc, lipd; path (hedp, wntp, tgfp+mapp, notp, jakp, fsap, hipp, tlrp)
[hide]v · d · eSignaling pathway: hedgehog signaling pathway
Ligands Sonic hedgehog · Indian hedgehog · Desert hedgehog
Receptor Patched · Smoothened
Transcription factor GLI1 · GLI2 · GLI3
Other HHIP · HHAT

calc, lipd; path (hedp, wntp, tgfp+mapp, notp, jakp, fsap, hipp, tlrp)
Retrieved from "http://en.wikipedia.org/wiki/Hedgehog_signaling_pathway"
Categories: Cell signaling | Hedgehog signaling pathway

W000
Evolutionary linguistics

Evolutionary linguistics is the scientific study of the origins and development of language. The main
challenge in this research is the lack of empirical data: spoken language leaves practically no traces. This led
to an abandonment of the field for more than a century.[1] Since the late 1980s, the field has been revived in
the wake of progress made in the related fields of psycholinguistics, neurolinguistics, evolutionary
anthropology and cognitive science.
Contents
[hide]
• 1 History
• 2 Recent developments
• 3 Approaches
• 4 EVOLANG Conference
• 5 Notes
• 6 See also
• 7 References
[edit] History
August Schleicher (1821–1868) and his Stammbaumtheorie are often quoted as the starting point of
evolutionary linguistics. Inspired by the natural sciences, especially biology, Schleicher was the first to
compare languages to evolving species.[2] He introduced the representation of language families as an
evolutionary tree in articles published in 1853. Joseph Jastrow published a gestural theory of the evolution of
language in the seventh volume of Science, 1886.[3]
The Stammbaumtheorie proved to be very productive for comparative linguistics, but didn't solve the
major problem of evolutionary linguistics: the lack of fossil records. The question of the origin of language
was abandoned as unsolvable. Famously, the Société Linguistique de Paris in 1866 refused to admit any
further papers on the subject.
The field has re-appeared in 1988 in the Linguistic Bibliography, as a subfield of psycholinguistics. In
1990, Steven Pinker and Paul Bloom published their paper "Natural Language & Natural Selection" which
strongly argued for an adaptationist approach to language origins. Their paper is often credited with reviving
the interest in evolutionary linguistics. This development was further strengthened by the establishment (in
1996) of a series of conferences on the Evolution of Language (now known as "Evolang"), promoting a
scientific, multidisciplinary approach to the issue, and interest from major academic publishers (e.g., the
Studies in the Evolution of Language series has been appearing with Oxford University Press since 2001)
and scientific journals.
[edit] Recent developments

Evolutionary linguistics as a field is rapidly evolving as a result of developments in neighboring
disciplines. Some of these developments are:
1. Studies of (human) genetics have started to make an impact on theories of language
evolution. The discovery of the FoxP2-gene in an English family with a heritable disorder has led to a
series of discoveries of versions of the gene in mice, songbirds, predators, great apes and even
Neandertals, and reconstructions of the evolutionary history of the gene. Although the interpretation
of these results is still contentious, it is clear that genetics will sooner or later start providing
constraints on plausible theories of language origins.
2. Studies in (behavioral) biology have forced researchers to reassess many claims of uniquely
human abilities for language and speech. For instance, Tecumseh Fitch has argued that the
descended larynx is not unique to humans. Similarly, once held uniquely human traits such as
formant perception, combinatorial phonology and compositional semantics are now thought to be
shared with at least some nonhuman animal species.
3. Computational modelling is now widely accepted as an approach to assure the internal
consistency of language evolution scenarios. Approximately one third of all accepted papers at the
most recent Evolution of Language conference [1] deal with computational models.
[edit] Approaches
One original researcher in the field is Luc Steels, head of the research units of Sony CSL in Paris and
the AI Lab at the Vrije Universiteit Brussel. He and his team are investigating ways in which artificial agents
self-organize languages with natural-like properties and how meaning can co-evolve with language. Their
research is based on the hypothesis that language is a complex adaptive system that emerges through
adaptive interactions between agents and continues to evolve in order to remain adapted to the needs and
capabilities of the agents. This research has been implemented in fluid construction grammar (FCG), a
formalism for construction grammars that has been specially designed for the origins and evolution of
language.
The approach of computational modeling and the use of robotic agents grounded in real life is
claimed to be theory independent. It enables the researcher to find out exactly what cognitive capacities are
needed for certain language phenomena to emerge. It also focuses the researcher in formulating hypotheses
in a precise and exact manner, whereas theoretical models often stay very vague.
Some linguists, such as John McWhorter, have analyzed the evolution and construction of basic
communication methods such as Pidginization and Creolization.[4]
"Nativist" models of "Universal Grammar" are informed by linguistic universals such as the existence
of pronouns and demonstratives, and the similarities in each languages process of nominalization (the
process of verbs becoming nouns) as well as the reverse, the process of turning nouns into verbs.[5] This is
a purely descriptive approach to what we mean by "natural language" without attempting to address its
emergence.
[edit] EVOLANG Conference

The Evolution of Language International Conferences [2][3] have been held biennially since 1996.
1. 1996 Edinburgh: Hurford, J. R., Studdert-Kennedy, M. & Knight C. (eds), Approaches to the
Evolution of Language - Social and Cognitive Bases, Cambridge University Press, 1998.
2. 1998 London: Chris Knight, James R. Hurford and Michael Studdert-Kennedy (eds), The
Evolutionary Emergence of Language: Social function and the origins of linguistic form , Cambridge
University Press,
3. 2000 Paris: J. L. Desalles & L. Ghadakpour (eds.), Proceedings of the 3rd International
Conference on the Evolution of Language
4. 2002 Boston: J. Hurford & T. Fitch (eds.), Proceedings of the 4th International Conference on
the Evolution of Language
5. 2004 Leipzig
6. 2006 Rome: Angelo Cangelosi, Andrew D. M. Smith, Kenny Smith The Evolution of
Language: Proceedings of the 6th International Conference on the Evolution of Language , World
Scientific, ISBN 9812566562.
7. 2008 Barcelona: [4] Andrew D. M. Smith, Kenny Smith, Ramon Ferrer i Cancho "The
Evolution of Language (EVOLANG 7)", World Scientific, ISBN 9812776117.
8. 2010 Utrecht, the Netherlands, April 14–17, 2010. [5]. Andrew D. M. Smith, Marieke
Schouwstra, Bart de Boer, Kenny Smith "The Evolution of Language (EVOLANG 8)", World
Scientific, ISBN 9814295213.
[edit] Notes
1. ^ for about 12 decades, from the 1860s to the 1980s.
2. ^ Taub, Liba. Evolutionary Ideas and "Empirical" Methods: The Analogy Between Language
and Species in the Works of Lyell and Schleicher. British Journal for the History of Science 26, pages
171–193 (1993)
3. ^ Jastrow J (1886). "The Evolution of Language". Science 7 (176S): 555–557.
doi:10.1126/science.ns-7.176S.555. PMID 17778380. http://links.jstor.org/sici?sici=0036-
8075%2818860618%292%3A7%3A176%3C555%3ATEOL%3E2.0.CO%3B2-7.
4. ^ (2002) McWhorter, John. The Power of Babel: The Natural History of Language, Random
House Group.
5. ^ (2005) Deutscher, Guy. The Unfolding of Language, Owl Books.
[edit] See also

• Origin of language
• Biolinguistics
[edit] References
• Cangelosi, A.; Harnad, S. (2001). "The adaptive advantage of symbolic theft over
sensorimotor toil: Grounding language in perceptual categories". Evolution of Communication 4 (1):
117–142. doi:10.1075/eoc.4.1.07can. http://cogprints.org/2036/.
• M. Christiansen and S. Kirby (eds.), Language Evolution, Oxford University Press, New York
(2003), ISBN 978-0199244843.
• Bickerton, D., Symbol and Structure: A Comprehensive Framework for Language
Evolution, pp. 77–93.
• Hurford, J. R., The Language Mosaic and Its Evolution, pp. 38–57.
• Lieberman, P.,Motor Control, Speech, and the Evolution of Language , pp. 252–271.
• Deacon, T. (1997) The symbolic species: the coevolution of language and the brain , Norton,
New York.
• Hauser, M.D. (1996) The evolution of communication , MIT Press, Cambridge, MA.
• Daniel Dor and Jablonka Eva (2001). How language changed the genes. In Tabant J. Ward.
S. (editors). Mouton de Gruyer: Berlin, pp 149–175.
• Dor D. and Jablonka E. (2001) From cultural selection to genetic selection: a framework for
the evolution of language. Selection, 1–3, pp. 33–57.
• Hauser MD, Chomsky N, Fitch WT (2002). "The faculty of language: what is it, who has it,
and how did it evolve?". Science 298 (5598): 1569–79. doi:10.1126/science.298.5598.1569.
PMID 12446899. http://www3.isrl.uiuc.edu/~junwang4/langev/localcopy/pdf/hauser02science.pdf.
• Jackendoff, R. (2002) Foundations of language: brain, meaning, grammar, evolution Oxford
University Press, New York
• Komarova, N.L. (2007). Language and Mathematics: An evolutionary model of grammatical
communication. In: History & Mathematics. Ed. by Leonid Grinin, Victor C. de Munck, and Andrey
Korotayev. Moscow, KomKniga/URSS. pp. 164–179. ISBN 9785484010011.
• Nowak, M.A.; Komarova, N.L. (2001). "Towards an evolutionary theory of language". Trends
in Cognitive Sciences 5 (7): 288–295. doi:10.1016/S1364-6613(00)01683-1. PMID 11425617.
• Pinker, S. (1994) The language instinct, HarperCollins, New York.
• Pinker, S.; Bloom, P. (1990). "Natural language and natural selection". Behavioral and Brain
Sciences 13: 707–784. http://www.bbsonline.org/documents/a/00/00/04/99/index.html.
• Sampson, Geoffrey: Evolutionary Language Understanding, published 1996 by Cassel
(London), ISBN 0304336505
• Steels, Luc (2001) Grounding Symbols through Evolutionary Language Games. In:
Cangelosi A. and Parisi D. (Eds.) Simulating the Evolution of Language Springer.
• Steklis, H.D.; Harnad, S (1976). "From hand to mouth: Some critical stages in the evolution
of language In: Harnad, S., Steklis, H. D. and Lancaster, J., (1976) (Eds) Origins and Evolution of
Language and Speech". Annals of the New York Academy of Sciences 280: 1–914.
• See also the UIUC Language Evolution and Computation Bibliography/Repository (1200+
related references, citations, and fulltext pointers)
• Encyclopedia Americana,Americana Corporation of Canada{1959}-Iceland-Language
• Zuidema, W. H., The Major Transitions in the Evolution of Language , PhD thesis, Theoretical
and Applied Linguistics, University of Edinburgh (2005) [6]
• Johansson, Sverker, Origins of language : constraints on hypotheses, Converging evidence
in language and communication research vol. 5, Amsterdam : Benjamins (2005).
• Mithen, Steven J., The singing neanderthals : the origins of music, language, mind and body
London : Weidenfeld & Nicolson (2005), ISBN 978-0-297-64317-3
• Partha Niyogi, The computational nature of language learning and evolution MIT Press,
Current studies in linguistics 43 (2006).
• A. Carstairs-McCarthy, The evolution of language, Lingua vol. 117, issue 3 (2007, March).
• Bernd Heine, Tania Kuteva, The genesis of grammar : a reconstruction, Oxford University
Press, 2007, ISBN 978-0-19-922776-1, ISBN 978-0-19-922777-8.
• James R. Hurford, Language in the light of evolution, Oxford University Press, Studies in the
evolution of language vol. 1 (2007).
• Atkinson QD, Meade A, Venditti C, Greenhill SJ, Pagel M (2008). "Languages evolve in
punctuational bursts". Science 319 (5863): 588. doi:10.1126/science.1149683. PMID 18239118.

• Elvira, Javier (2009). Evolución lingüística y cambio sintáctico . Fondo Hispánico de
Lingüística y Filología. Bern et al.: Peter Lang. ISBN 9783034303231.
• Harnad, Stevan R.; Steklis, Horst D.; Lancaster, Jane, [editors] (1976). Origins and Evolution
of Language and Speech. Annals of the New York Academy of Sciences, v. 280. New York: New
York Academy of Sciences. ISBN 0890720266.
• Johanson, Donald C.; and Edgar, Blake (2006). From Lucy to Language (Revised, updated,
and expanded ed.). New York, NY: Simon and Schuster. ISBN 0743280644. OCLC 72440476.
• Kenneally, Christine (2007). The First Word: The Search for the Origins of Language. New
York, NY: Viking. ISBN 9780670034901. OCLC 80460757.
• Tallerman, Maggie (2005). Language Origins: Perspectives on Evolution. Oxford, UK; New
York, NY: Oxford University Press. ISBN 0199279047. OCLC 60607214.

• Fluid Construction Grammar
• Language Evolution and Computation Research Unit, University of Edinburgh
• Sony CSL Research
• ARTI Artificial Intelligence Laboratory, Vrije Universiteit Brussel
• ECAgents: The Project on Embodied and Communicating Agents
[hide]v · d · eAnimal communication
Animal training · Animal language · Animal cognition · Bioacoustics ·

Concepts Ethology · Evolutionary linguistics · FOXP2 · Talking animals · Origin of language ·
Human-animal communication
Animal-specific Bird vocalization · Talking birds · Great ape language (Yerkish) · Whale
song · Dog communication
Related topics Category:Talking birds · Category:Apes from language studies
Retrieved from "http://en.wikipedia.org/wiki/Evolutionary_linguistics"
Categories: Evolution of language | Linguistics

W000
Evolution of the horse


This image shows a representative sequence but should not be construed to represent a "straight-
line" evolution of the horse. Reconstruction, left forefoot skeleton (third digit emphasized yellow) and
longitudinal section of molars of selected prehistoric horses
Skeletal evolution
The evolution of the horse pertains to the phylogenetic ancestry of the modern horse from the fox-
sized, forest-dwelling Hyracotherium over geologic time scales. Paleozoologists have been able to piece
together a more complete picture of the modern horse's evolutionary lineage than that of any other animal.
The horse belongs to an order known as Perissodactyla, or "odd-toed ungulates", which all share
hoofed feet and an odd number of toes on each foot, as well as mobile upper lips and a similar tooth
structure. This means that horses share a common ancestry with tapirs and rhinoceroses. The perissodactyls
originally arose in the late Paleocene, less than 10 million years after the Cretaceous-Tertiary extinction
event. This group of animals appears to have been originally specialized for life in tropical forests, but
whereas tapirs and, to some extent, rhinoceroses, retained their jungle specializations, modern horses are
adapted to life on drier land in the much-harsher climatic conditions of the steppes. Other species of Equus
are adapted to a variety of intermediate conditions.
The early ancestors of the modern horse walked on several spread-out toes, an accommodation to
life spent walking on the soft, moist grounds of primeval forests. As grass species began to appear and
flourish, the equids' diets shifted from foliage to grasses, leading to larger and more durable teeth. At the
same time, as the steppes began to appear, the horse's predecessors needed to be capable of greater
speeds to outrun predators. This was attained through the lengthening of limbs and the lifting of some toes
from the ground in such a way that the weight of the body was gradually placed on one of the longest toes,
the third.
Contents
[hide]
• 1 History of research
• 2 Eocene and Oligocene: early equids
• 2.1 Hyracotherium
• 2.2 Orohippus
• 2.3 Epihippus
• 2.4 Mesohippus
• 2.5 Miohippus
• 3 Miocene and Pliocene: true equines
• 3.1 Kalobatippus
• 3.2 Parahippus
• 3.3 Merychippus
• 3.4 Hipparion
• 3.5 Pliohippus
• 3.6 Dinohippus
• 3.7 Plesippus
• 4 Modern horses
• 4.1 Equus
• 4.2 Pleistocene extinctions
• 4.3 Return to the Americas
• 5 Details
[edit] History of research
Restoration of Eurohippus parvulus, Museum für Naturkunde, Berlin

Horses were absent from the Americas until the Spanish brought domestic horses from Europe,
beginning in 1493, and escaped horses quickly established large wild herds. The early naturalist Buffon
suggested in the 1760s that this was an indication of inferiority of fauna in the New World, then later
reconsidered this idea.[1] William Clark's 1807 expedition to Big Bone Lick found "leg and foot bones of the
Horses" which were included with other fossils sent to Thomas Jefferson and evaluated by the anatomist
Caspar Wistar, but neither commented on the significance of this find.[2]
The first equid fossil was found in the gypsum quarries in Montmartre, Paris in the 1820s. The tooth
was sent to the Paris Conservatory, where it was identified by Georges Cuvier who identified it as a browsing
equine related to the tapir.[3] His sketch of the entire animal matched later skeletons found at the site.[4]
During the Beagle survey expedition the young naturalist Charles Darwin had remarkable success
with fossil hunting in Patagonia. On 10 October 1833 at Santa Fe, Argentina, he was "filled with
astonishment" when he found a horse's tooth in the same stratum as fossil giant armadillos, and wondered if
it might have been washed down from a later layer, but concluded that this was "not very probable".[5] After
the expedition returned in 1836, the anatomist Richard Owen confirmed the tooth was from an extinct
species which he subsequently named Equus curvidens, and remarked that "This evidence of the former
existence of a genus, which, as regards South America, had become extinct, and has a second time been
introduced into that Continent, is not one of the least interesting fruits of Mr. Darwin's palæontological
discoveries."[2][6]
In 1848 a study On the fossil horses of America by Joseph Leidy systematically examined
Pleistocene horse fossils from various collections, including that of the Academy of Natural Sciences and
concluded at least two ancient horse species had existed in North America: Equus curvidens and another
which he named Equus americanus. A decade later, however, he found the latter name had already been
taken and renamed it Equus complicatus.[1] In the same year, he visited Europe and was introduced by
Owen to Darwin.[7]
The original sequence of species believed to have evolved into the horse was based on fossils
discovered in North America in the 1870s by paleontologist Othniel Charles Marsh. The sequence, from
Hyracotherium (popularly called Eohippus) to the modern horse (Equus), was popularized by Thomas Huxley
and became one of the most widely-known examples of a clear evolutionary progression. The horse's
evolutionary lineage became a common feature of biology textbooks, and the sequence of transitional fossils
was assembled by the American Museum of Natural History into an exhibit which emphasized the gradual,
"straight-line" evolution of the horse.
Since then, as the number of equid fossils has increased, the actual evolutionary progression from
Hyracotherium to Equus has been discovered to be much more complex and multi-branched than was
initially supposed. The straight, direct progression from the former to the latter has been replaced by a more
elaborate model with numerous branches in different directions, of which the modern horse is only one of
many. It was first recognized by George Gaylord Simpson in 1951[8] that the modern horse was not the
"goal" of the entire lineage of equids,[9] it is simply the only genus of the many horse lineages that has
survived.
Detailed fossil information on the rate and distribution of new equid species has also revealed the
progression between species was not as smooth and consistent as was once believed. Although some
transitions, such as that of Dinohippus to Equus, were indeed gradual progressions, a number of others,
such as that of Epihippus to Mesohippus, were relatively abrupt and sudden in geologic time, taking place
over only a few million years. Both anagenesis (gradual change in an entire population's gene frequency)
and cladogenesis (a population "splitting" into two distinct evolutionary branches) occurred, and many
species coexisted with "ancestor" species at various times. The change in equids' traits was also not always
a "straight line" from Hyracotherium to Equus: some traits reversed themselves at various points in the
evolution of new equid species, such as size and the presence of facial fossae, and it is only in retrospect
that certain evolutionary trends can be recognized.[10]
[edit] Eocene and Oligocene: early equids

[edit] Hyracotherium
The earliest animal to bear recognizably horse-like anatomy was the Hyracotherium ("hyrax-like
beast"). Its scientific name is derived from initial confusion over early partial fossils' relationship with living
species: Richard Owen likened early Hyracotherium fossils "to a hare in one passage and to something
between a hog and a hyrax in another".[11] A later name for the Hyracotherium, "eohippus" ("dawn horse"), is
also popular, though the earlier name takes precedence due to scientific naming conventions.[12][13]
Hyracotherium lived in the Ypresian (early Eocene), about 52 mya (million years ago). It was an
animal approximately the size of a fox (250–450 mm in height), with a relatively short head and neck and a
springy, arched back. It had 44 low-crowned teeth, in the typical arrangement of an omnivorous, browsing
mammal: 3 incisors, 1 canine, 4 premolars, and 3 molars on each side of the jaw. Its molars were uneven,
dull, and bumpy, and used primarily for grinding foliage. The cusps of the molars were slightly connected in
low crests. The Hyracotherium browsed on soft foliage and fruit, probably scampering between thickets in the
mode of a modern muntjac; the Hyracotherium had a small brain, and possessed especially small frontal
lobes.[10]
Hyracotherium, with left forefoot (third metacarpal colored) and tooth (a enamel; b dentin; c cement)
detailed.
Its limbs were decently long relative to its body, already showing the beginnings of adaptations for
running. However, all of the major leg bones were unfused, leaving the legs flexible and rotatable. Its wrist
and hock joints were low to the ground. The forelimbs had developed five toes, out of which only four were
equipped with a small proto-hoof; the large fifth "toe-thumb" was off the ground. The hind limbs had three out
of the five toes equipped with small hooves, while the vestigial first and fifth toes did not touch the ground. Its
feet were padded, much like a dog's, but with the small hooves on each toe in place of claws.
For a span of about 20 million years, the Hyracotherium thrived with few significant evolutionary
changes.[10] The most significant change was in the teeth, which began to adapt to the changing diet of
Hyracotheria, as these early Equidae shifted from a mixed diet of fruits and foliage to one focused
increasingly on browsing foods. During the Eocene, a Hyracotherium species (most likely Hyracotherium
vassacciense) branched out into various new types of Equidae. Thousands of complete, fossilized skeletons
of these animals have been found in the Eocene layers of North American strata, mainly in the Wind River
basin in Wyoming. Similar fossils have also been discovered in Europe, such as Propalaeotherium (which is
not considered ancestral to the modern horse).[14]
[edit] Orohippus
Approximately 50 million years ago, in the early-to-middle Eocene, Hyracotherium smoothly
transitioned into Orohippus over a gradual series of changes.[14] Although its name means "mountain
horse", Orohippus was not a true horse and did not live in the mountains. It resembled Hyracotherium in size,
but had a slimmer body, an elongated head, slimmer forelimbs, and longer hind legs, all of which are
characteristics of a good jumper. Although Orohippus was still pad-footed, the vestigial outer toes of
Hyracotherium were not present in the Orohippus; there were four toes on each forelimb, and three on each
hind leg.
The most dramatic change between Hyracotherium and Orohippus was in the teeth: the first of the
premolar teeth were dwarfed, the last premolar shifted in shape and function into a molar, and the crests on
the teeth became more pronounced. Both of these factors gave the teeth of Orohippus greater grinding
ability, suggesting that Orohippus ate tougher plant material.
[edit] Epihippus
In the mid-Eocene, about 47 million years ago, Epihippus, a genus which continued the evolutionary
trend of increasingly efficient grinding teeth, evolved from Orohippus. Epihippus had five grinding, low-
crowned cheek teeth with well-formed crests. A late species of Epihippus, sometimes referred to as
Duchesnehippus intermedius, had teeth similar to Oligocene equids, although slightly less developed.
Whether Duchesnehippus was a subgenus of Epihippus or a distinct genus is disputed.
[edit] Mesohippus
In the late Eocene and the early stages of the Oligocene epoch (32–24 mya), the climate of North
America became drier, and the earliest grasses began to evolve. The forests were yielding to flatlands,
[citation needed] home to grasses and various kinds of brush. In a few areas these plains were covered in
sand,[citation needed] creating the type of environment resembling the present-day prairies.
In response to the changing environment, the then-living species of Equidae also began to change.
In the late Eocene, they began developing tougher teeth and becoming slightly larger and leggier, allowing
for faster running speeds in open areas, and thus for evading predators in non-wooded areas[ citation
needed]. About 40 mya, Mesohippus ("middle horse") suddenly developed in response to strong new
selective pressures to adapt, beginning with the species Mesohippus celer and soon followed by Mesohippus
westoni.
In the early Oligocene, Mesohippus was one of the more widespread mammals in North America. It
walked on three toes on each of its front and hind feet (the first and fifth toes remained, but were small and
not used in walking). The third toe was stronger than the outer ones, and thus more weighted; the fourth front
toe was diminished to a vestigial nub. Judging by its longer and slimmer limbs, Mesohippus was an agile
animal.
Mesohippus was slightly larger than Epihippus, about 610 mm (24") at the shoulder. Its back was
less arched, and its face, snout, and neck were somewhat longer. It had significantly larger cerebral
hemispheres, and had a small, shallow depression on its skull called a fossa, which in modern horses is quite
detailed. The fossa serves as a useful marker for identifying an equine fossil's species. Mesohippus had six
grinding "cheek teeth", with a single premolar in front—a trait all descendant Equidae would retain.
Mesohippus also had the sharp tooth crests of Epihippus, improving its ability to grind down tough
vegetation.
[edit] Miohippus
Around 36 million years ago, soon after the development of Mesohippus, Miohippus ("lesser horse")
emerged, the earliest species being Miohippus assiniboiensis. Like Mesohippus, Miohippus's evolution was
relatively abrupt, though a few transitional fossils linking the two genera have been found. It was once
believed that Mesohippus had anagenetically evolved into Miohippus by a gradual series of progressions, but
new evidence has shown that Miohippus's evolution was cladogenetic: a Miohippus population split off from
the main Mesohippus genus, coexisted with Mesohippus for around 4 million years, and then over time came
to replace Mesohippus.[15]
Miohippus was significantly larger than its predecessors, and its ankle joints had subtly changed. Its
facial fossa was larger and deeper, and it also began to show a variable extra crest in its upper cheek teeth,
a trait that became a characteristic feature of equine teeth.
Miohippus ushered in a major new period of diversification in Equidae.[13] While Mesohippus died
out in the mid-Oligocene, Miohippus continued to thrive, and in the early Miocene (24–5.3 mya), it began to
rapidly diversify and speciate. It branched out into two major groups, one of which adjusted to the life in
forests once again, while the other remained suited to life on the prairies.[ citation needed]
[edit] Miocene and Pliocene: true equines

[edit] Kalobatippus
Fossil Megahippus mckennai
The forest-suited form was Kalobatippus (or Miohippus intermedius, depending on whether it was a
new genus or species), whose second and fourth front toes were long, well-suited travel on the soft forest
floors. Kalobatippus probably gave rise to Anchitherium, which travelled to Asia via the Bering Strait land
bridge, and from there to Europe.[16] In both North America and Eurasia, larger-bodied genera evolved from
Anchitherium; Sinohippus in Eurasia and Hypohippus and Megahippus in North America.[17] Hypohippus
became extinct by the late Miocene.[18]
[edit] Parahippus
The Miohippus population that remained on the steppes is believed to be ancestral to Parahippus, a
North American animal about the size of a small pony, with a prolonged skull and a facial structure
resembling the horses of today. Its third toe was stronger and larger, and carried the main weight of the body.
Its four premolars resembled the molar teeth and the first were small and almost nonexistent. The incisive
teeth of Parahippus, like those of its predecessors, had a crown as humans do; however, the top incisors had
a trace of a shallow crease marking the beginning of the core/cup.
[edit] Merychippus
Merychippus, an effective grazer and runner.

In the middle of the Miocene epoch, the grazer Merychippus flourished. Merychippus had wider
molars than its predecessors, which are believed to have been used for crunching the hard grasses of the
steppes. The hind legs, which were relatively short, had side toes equipped with small hooves, but they
probably only touched the ground when running.[13] Merychippus radiated into at least 19 additional
grassland species.
[edit] Hipparion
Protohippus simus
Three lineages within Equidae are believed to be descended from the numerous varieties of
Merychippus: Hipparion, Protohippus and Pliohippus. The most different from Merychippus was Hipparion.
The main difference was in the structure of tooth enamel: in comparison with other Equidae, the inside, or
tongue side, had a completely isolated parapet. A complete and well-preserved skeleton of the North
American Hipparion shows an animal the size of a small pony. They were very slim, rather like antelopes,
and were adapted to life on dry prairies. On its slim legs, Hipparion had three toes equipped with small
hooves, but the side toes did not touch the ground.
In North America, Hipparion and its relatives (Cormohipparion, Nannippus, Neohipparion, and
Pseudhipparion), proliferated into many kinds of equids, at least one of which managed to migrate to Asia
and Europe during the Miocene epoch.[19] (European Hipparion differs from American Hipparion in its
smaller body size – the best-known discovery of these fossils was near Athens.)
[edit] Pliohippus
Pliohippus pernix
Pliohippus arose from Callippus in the middle Miocene, around 12 mya. It was very similar in
appearance to Equus, though it had two long extra toes on both sides of the hoof, externally barely visible as
callused stubs. The long and slim limbs of Pliohippus reveal a quick-footed steppe animal.
Until recently, Pliohippus was believed to be the ancestor of present-day horses because of its many
anatomical similarities. However, though Pliohippus was clearly a close relative of Equus, its skull had deep
facial fossae, whereas Equus had no fossae at all. Additionally, its teeth were strongly curved, unlike the very
straight teeth of modern horses. Consequently, it is unlikely to be the ancestor of the modern horse; instead,
it is a likely candidate for the ancestor of Astrohippus.[20]
[edit] Dinohippus
Dinohippus was the most common species of Equidae in North America during the late Pliocene. It
was originally thought that Dinohippus was monodactyl, but a 1981 fossil find in Nebraska shows that some
were tridactyl.
[edit] Plesippus
Mounted skeleton of Hagerman Horse (Equus simplicidens)

Plesippus is often considered an intermediate stage between Dinohippus and the extant genus,
Equus.
The famous fossils found near Hagerman, Idaho were originally thought to be a part of the genus
Plesippus. Hagerman Fossil Beds (Idaho) is a Pliocene site, dating to about 3.5 mya. The fossilized remains
were originally called Plesippus shoshonensis, but further study by paleontologists determined that fossils
represented the oldest remains of the genus Equus.[21] Their estimated average weight was 425 kg, roughly
the size of an Arabian horse.
At the end of the Pliocene, the climate in North America began to cool significantly and most of the
animals were forced to move south. One population of Plesippus moved across the Bering land bridge into
Eurasia around 2.5 Mya.[22]
[edit] Modern horses

[edit] Equus
Skull of a giant extinct horse of the genus Equus, E. eisenmannae

The genus Equus, which includes all extant equines, is believed to have evolved from Dinohippus,
via the intermediate form Plesippus. One of the oldest species is Equus simplicidens, described as zebra-like
with a donkey-shaped head. The oldest material to date is ~3.5 million years old from Idaho, USA. The genus
appears to have spread quickly into the Old World, with the similarly aged Equus livenzovensis documented
from western Europe and Russia.[23]
Molecular phylogenies indicate that the most recent common ancestor of all modern equids
(members of the genus Equus) lived ~5.6 (3.9-7.8) mya. The oldest divergencies are the Asian hemiones
(subgenus E. (Asinus)), including the Kulan, Onager, and Kiang), followed by the African zebras (subgenera
E. (Dolichohippus), and E. (Hippotigris)). All other modern forms including the domesticated horse (and many
fossil Pliocene and Pleistocene forms) belong to the subgenus E. (Equus) which diverged ~4.8 (3.2-6.5)
million years ago.[24]
Pleistocene horse fossils have been assigned to a multitude of species, with over 50 species of
equines described from the Pleistocene of North America alone, although the taxonomic validity of most of
these has been called into question.[25] Recent genetic work on fossils has found evidence for only three
genetically divergent equid lineages in Pleistocene North and South America.[24] These results suggest that
all North American fossils of caballine-type horses (which also include the domesticated horse and
Przewalski's Horse of Europe and Asia), as well as South American fossils traditionally placed in the
subgenus E. (Amerhippus)[26] belong to the same species: E. ferus. Remains attributed to a variety of
species and lumped as New World stilt-legged horses (including E. francisci, E. tau, E. quinni and potentially
N. American Pleistocene fossils previously attributed to E. cf. hemiones, and E. (Asinus) cf. kiang) likely all
belong to a second species endemic to N. America, which despite a superficial resemblance to species in the
subgenus E. (Asinus) (and hence occasionally referred to as North American Ass) is closely related to E.
ferus.[24] Surprisingly, the third species, endemic to S. America, and traditionally referred to as Hippidion,
originally believed to be descended from Pliohippus, was shown to be a third species in the genus Equus,
closely related to the New World stilt-legged horse.[24] The temporal and regional variation in body size and
morphological features within each lineage indicates extraordinary intraspecific plasticity. Such environment-
driven adaptative changes would explain why the taxonomic diversity of Pleistocene equids has been
overestimated on morphoanatomical grounds.[26]
According to these results, it appears that the genus Equus evolved from a Dinohippus-like ancestor
~4-7 mya. It rapidly spread into the Old World and there diversified into the various species of asses and
zebras. A North American lineage of the subgenus E. (Equus) evolved into the New World stilt-legged horse
(NWSLH). Subsequently, populations of this species entered South America as part of the Great American
Interchange shortly after the formation of the Isthmus of Panama and evolved into the form currently referred
to as "Hippidion" ~2.5 million years ago. "Hippidion" is thus unrelated to the morphologically similar
Pliohippus, which presumably went extinct during the Miocene. Both the NWSLH and "Hippidium" show
adaptations to dry, barren ground, whereas the shortened legs of " Hippidion" may have been a response to
sloped terrain.[26] In contrast, the geographic origin of the closely related modern E. ferus is not resolved.
However, genetic results on extant and fossil material of Pleistocene age indicate two clades, potentially
subspecies, one of which had a holarctic distribution spanning from Europe through Asia and across North
America and would become the founding stock of the modern domesticated horse.[27][28] The other
population appears to have been restricted to N. America. One or more N. American populations of E. ferus
entered S. America ~1.0-1.5 million years ago, leading to the forms currently known as " E. (Amerhippus)",
which represent an extinct geographic variant or race of E. ferus, however.
[edit] Pleistocene extinctions

Digs in western Canada have unearthed clear evidence that horses existed in North America until
about 12,000 years ago.[29] However, all Equidae in North America ultimately became extinct. The causes of
this extinction (simultaneous with the extinctions of a variety of other American megafauna) have been a
matter of debate. Given the suddenness of the event and the fact that these mammals had been flourishing
for millions of years previously, something quite unusual must have happened. There are two main
hypotheses. The first attributes extinction to climate change. For example, in Alaska, beginning
approximately 12,500 years ago, the grasses characteristic of a steppe ecosystem gave way to shrub tundra,
which was covered with unpalatable plants.[30][31] Another hypothesis suggests extinction was linked to
overexploitation of naive prey by newly arrived humans. Extinctions were roughly simultaneous with the end
of the most recent glacial advance and the appearance of the big-game-hunting Clovis culture.[32][33]
Several studies have indicated that humans probably arrived in Alaska at the same time or shortly before the
local extinction of horses.[33][34][35] Additionally, it has been proposed that the steppe-tundra vegetation
transition in Beringia may have been a consequence, rather than a cause, of the extinction of megafaunal
grazers.[36]
In Eurasia, horse fossils began occurring frequently again in archaeological sites in Kazakhstan and
the southern Ukraine about 6,000 years ago.[27] From then on, it is probable that domesticated horses as
well as the knowledge of capturing, taming, and rearing horses spread relatively quickly, with wild mares from
several wild populations being incorporated en route.[28]
[edit] Return to the Americas

Horses only returned to the Americas with Christopher Columbus in 1493. These were Iberian horses
first brought to Hispaniola and later to Panama, Mexico, Brazil, Peru, Argentina, and, in 1538, Florida.[37]
The first horses to return to the main continent were 16 specifically identified horses brought by Hernan
Cortes. Subsequent explorers, such as Coronado and De Soto brought ever-larger numbers, some from
Spain and other from breeding establishments set up by the Spanish in the Caribbean. Later, as Spanish
missions were founded on the mainland, horses would eventually be lost or stolen, and proliferated into large
herds of feral horses that became known as mustangs.[citation needed]
The indigenous peoples of the Americas did not have a specific word for horses, and came to refer to
them in various languages as a type of dog or deer (in one case, "elk-dog").[ citation needed]
[edit] Details
[edit] Toes
The ancestors of the horse came to walk only on the end of the third toe and both side toes. Skeletal
remnants show obvious wear on the back of both sides of metacarpal and metatarsal bones, commonly
called the “splint bones”. They are the remnants of the second and the fourth toe. Modern horses retain the
splint bones; it is often believed that they are a useless attachment, but they in fact play an important role in
supporting the carpal joints (front knee) and even the tarsal joints (hock).
[edit] Teeth
Throughout the phylogenetic development, the teeth of the horse underwent significant changes. The
type of the original omnivorous teeth with short, "bumpy" molars, with which the prime members of the
evolutionary line distinguished themselves, gradually changed into the teeth common to herbivorous
mammals. They became long (as much as 100 mm), roughly cubical molars equipped with a flat grinding
surface. In conjunction with the teeth, during the horse’s evolution the elongation of the facial part of the skull
is apparent, and can also be observed in the backward set eyeholes. In addition, the relatively short neck of
the equine ancestors became longer with equal elongation of the legs. Finally, the size of the body grew as
well.
[edit] See also
• Evidence of common descent

• MacFadden, Bruce J (2003 digitization of 1999 reprint). Fossil Horses: Systematics,
Paleobiology, and Evolution of the Family Equidae . Cambridge & New York: Cambridge University
Press. ISBN 0-521-47708-5. http://books.google.com/?
id=K1upTamSEW0C&printsec=frontcover&dq=%22fossil+horses%22&cd=1#v=onepage&q.
Retrieved 6 June 2010
[edit] References
1. ^ a b Academy of Natural Sciences - Joseph Leidy - American Horses
2. ^ a b Academy of Natural Sciences - Thomas Jefferson Fossil Collection - Ancient Horse
Fossils
3. ^ Horse breeding and management, James Warren Evans 1992
4. ^ Knell, Simon J.; Suzanne Macleod; Sheila E. R. Watson; Museum revolutions: how
museums and change and are changed Routledge, 2007, 385 pages ISBN 0-415-44467-5,
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5. ^ 'Filled with astonishment': an introduction to the St. Fe Notebook,
Barlow, Nora (ed. 1945) Charles Darwin and the voyage of the Beagle. London: Pilot Press, p. 210
6. ^ Darwin, C. R. (ed. 1840). Fossil Mammalia Part 1 No. 4 of The zoology of the voyage of
H.M.S. Beagle. By Richard Owen. London: Smith Elder and Co. p. 108–109
7. ^ Academy of Natural Sciences - Joseph Leidy - Leidy and Darwin
8. ^ Simpson, George Gaylord (1951): Horses. Oxford University Press; New Impression
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9. ^ The notion of a goal would contradict modern evolutionary synthesis
10.^ a b c Hunt, Kathleen (1995). Horse Evolution. TalkOrigins Archive.
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11.^ Gould, Stephen Jay (1991). "The Case of the Creeping Fox Terrier Clone" Bully for
Brontosaurus: Reflections in Natural History (pp. 155–167). New York: W.W. Norton & Co.
12.^ Gould, Stephen Jay, op. cit., "Bully for Brontosaurus"
13.^ a b c Fossil Horses In Cyberspace. Florida Museum of Natural History and the National
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14.^ a b MacFadden, B. J. (1976). "Cladistic analysis of primitive equids with notes on other
perissodactyls". Syst. Zool 25 (1): 1–14. doi:10.2307/2412774. http://jstor.org/stable/2412774.
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of Perissodactyls (pp. 142–175). New York: Clarendon Press.
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Nannippus, and Cormohipparion (Mammalia, Equidae) from the Miocene and Pliocene of the New
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20.^ MacFadden, B. J. (1984). "Astrohippus and Dinohippus". J. Vert. Paleon 4 (2): 273–283.
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22.^ Jens Lorenz Franzen: Die Urpferde der Morgenröte. Elsevier, Spektrum Akademischer
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23.^ Azzaroli, A. (1992). "Ascent and decline of monodactyl equids: a case for prehistoric
overkill". Ann. Zool. Finnici 28: 151–163. http://www.sekj.org/PDF/anzf28/anz28-151-163.pdf.
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Pleistocene horses in the New World: a molecular perspective". PLoS Biology 3 (8): e241.
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document&doi=10.1371%2Fjournal.pbio.0030241&ct=1. Retrieved 2008-12-19.
25.^ Azzaroli, A. (1998). "The genus Equus in North America". PalaeontographItal 85: 1–60.
26.^ a b c Orlando, L.; et al. (2008). "Ancient DNA Clarifies the Evolutionary History of American
Late Pleistocene Equids". Journal of Molecular Evolution 66 (5): 533–538. doi:10.1007/s00239-008-
9100-x. PMID 18398561.
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30.^ LeQuire, Elise (2004-01-04). "No Grass, No Horse". The Horse, online edition.
http://www.thehorse.com/ViewArticle.aspx?ID=4849. Retrieved 2009-06-08.
31.^ Guthrie, R. D. (2003-11-13). "Rapid body size decline in Alaskan Pleistocene horses before
extinction". Nature 426: 169–171. doi:10.1038/nature02098. PMID 14614503.
http://www.nature.com/nature/journal/v426/n6963/full/nature02098.html. Retrieved 2010-12-30.
32.^ "Ice Age Horses May Have Been Killed Off by Humans" National Geographic News, May 1,
2006.
33.^ a b Buck, Caitlin E.; Bard, Edouard (2007). "A calendar chronology for Pleistocene
mammoth and horse extinction in North America based on Bayesian radiocarbon calibration".
Quaternary Science Reviews 26 (17-18): 2031. doi:10.1016/j.quascirev.2007.06.013.
34.^ Solow, Andrew; Roberts, David; Robbirt, Karen (May 9, 2006). Haynes, C. Vance. ed. "On
the Pleistocene extinctions of Alaskan mammoths and horses". Proceedings of the National
Academy of Sciences of the United States of America (Proceedings of the National Academy of
Sciences of the United States of America) 103 (19): 7351–3. doi:10.1073/pnas.0509480103.
PMID 16651534. PMC 1464344. http://www.pnas.org/content/103/19/7351.full.
35.^ Guthrie, R. D. (2006-05-11). "New carbon dates link climatic change with human
colonization and Pleistocene extinctions". Nature 441: 207–209. doi:10.1038/nature04604.
2010-12-30.
36.^ Zimov, S. A.; Chuprynin, V. I.; Oreshko, A. P.; Chapin, F. S.; Reynolds, J. F.; Chapin, M. C.
(Nov. 1995). "Steppe-tundra transition: a herbivore-driven biome shift at the end of the Pleistocene".
The American Naturalist 146 (5): 765–794. http://www.jstor.org/stable/2462990.
37.^ Luís, Cristina; et al. (2006). "Iberian Origins of New World Horse Breeds". Journal of
Heredity 97 (2): 107–113. doi:10.1093/jhered/esj020. PMID 16489143.
http://jhered.oxfordjournals.org/cgi/content/full/97/2/107.

• Horse Evolution Over 55 Million Years. Tufts University. January 10, 1998.
http://chem.tufts.edu/science/evolution/HorseEvolution.htm. Retrieved July 11, 2007. : An excerpted
collection of images of horse fossils
[hide]v · d · eEquine
Equine Equine anatomy · Equine nutrition ·

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Hyracotheriinae Epihippus · Haplohippus · Heptaconodon · Hyracotherium · Orohippus ·

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Anchitheriinae Anchitherium · Archaeohippus · Desmatippus · Hypohippus ·

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[hide]v · d · eSpecies and hybrids of genus Equus, both extant and extinct
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complicatus · †Equus conversidens · †Equus crinidens · †Equus cumminsii · †Equus
excelsus · Equus ferus (†Equus ferus ferus, Equus ferus przewalskii · Equus ferus
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parastylidens · †Equus pectinatus · Equus quagga (†Equus quagga quagga) · †Equus
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Retrieved from "http://en.wikipedia.org/wiki/Evolution_of_the_horse"
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Human evolutionary genetics


Human evolutionary genetics studies how one human genome differs from the other, the
evolutionary past that gave rise to it, and its current effects. Differences between genomes have
anthropological, medical and forensic implications and applications. Genetic data can provide important
insight into human evolution.
Contents
[hide]
• 1 Origin of apes
• 2 Cladistics
• 3 Speciation of humans and the African apes
• 3.1 General observations
• 3.2 Divergence times
• 3.3 Divergence times and ancestral effective
population size
• 4 Genetic differences between humans and other great
apes
• 4.1 Gene loss
• 4.1.1 Hair keratin gene KRTHAP1
• 4.1.2 Myosin gene MYH16
• 4.1.3 Other
• 4.2 Gene addition
• 4.3 Selection pressures
• 5 Genetic differences between humans and Neanderthals
• 6 Sequence divergence between humans and apes
• 7 Modern humans
• 7.1 Age of the common ancestor
• 7.2 African origin for modern humans
[edit] Origin of apes
Taxonomic relationships of hominoids

Biologists classify humans, along with only a few other species, as great apes (species in the family
Hominidae). The Hominidae include two distinct species of chimpanzee (the bonobo, Pan paniscus, and the
common chimpanzee, Pan troglodytes), two species of gorilla (the western gorilla, Gorilla gorilla, and the
eastern gorilla, Gorilla graueri), and two species of orangutan (the Bornean orangutan, Pongo pygmaeus,
and the Sumatran orangutan, Pongo abelii).
Apes, in turn, belong to the primates order (>400 species). Data from both mitochondrial DNA
(mtDNA) and nuclear DNA (nDNA) indicates that primates belong to the group of Euarchontoglires, together
with Rodentia, Lagomorpha, Dermoptera, and Scandentia.[1] This is further supported by Alu-like short
interspersed nuclear elements (SINEs) which have been found only in members of the Euarchontoglires.[2]
[edit] Cladistics
A phylogenetic tree like the one shown above is usually derived from DNA or protein sequences from
populations. Often mitochondrial DNA or Y chromosome sequences are used to study ancient human
demographics. These single-locus sources of DNA do not recombine and are almost always inherited from a
single parent, with only one known exception in mtDNA (Schwartz and Vissing 2002). Individuals from the
various continental groups tend to be more similar to one another than to people from other continents. The
tree is rooted in the common ancestor of chimpanzees and humans, which is believed to have originated in
Africa. Horizontal distance in the diagram corresponds to two things:
1. Genetic distance. Given below the diagram, the genetic difference between humans and
chimps is less than 2%,[3] or 20 times larger than the variation among modern humans.
2. Temporal remoteness of the most recent common ancestor. Rough estimates are given
above the diagram, in millions of years. The mitochondrial most recent common ancestor of modern
humans lived roughly 200,000 years ago, latest common ancestors of humans and chimps between
four and seven million years ago.
Chimpanzees and humans belong to different genera, indicated in red. Formation of species and
subspecies is also indicated, and the formation of "races" is indicated in the green rectangle to the right (note
that only a very rough representation of human phylogeny is given). Note that vertical distances are not
meaningful in this representation.
[edit] Speciation of humans and the African apes

The separation of humans from their closest relatives, the African apes (chimpanzees and gorillas),
has been studied extensively for more than a century. Five major questions have been addressed:
• Which apes are our closest ancestors?
• When did the separations occur?
• What was the effective population size of the common ancestor before the split?
• Are there traces of population structure (subpopulations) preceding the speciation or partial
admixture succeeding it?
• What were the specific events (including fusion of chromosomes 2a and 2b) prior to and
subsequent to the separation?
[edit] General observations

As discussed before, different parts of the genome show different sequence divergence between
different hominoids. It has also been shown that the sequence divergence between DNA from humans and
chimpanzees varies greatly. For example the sequence divergence varies between 0% to 2.66% between
non-coding, non-repetitive genomic regions of humans and chimpanzees.[4] Additionally gene trees,
generated by comparative analysis of DNA segments, do not always fit the species tree. Summing up:
• The sequence divergence varies significantly between humans, chimpanzees and gorillas.
• For most DNA sequences, humans and chimpanzees appear to be most closely related, but
some point to a human-gorilla or chimpanzee-gorilla clade.
• The human genome has been sequenced, as well as the chimpanzee genome. Humans
have 23 pairs of chromosomes, while chimpanzees, gorillas, and orangutans have 24. Human
chromosome 2 is a fusion between two chromosomes that remained separate in the other primates.
[5]
[edit] Divergence times

The divergence time of humans from other apes is of great interest. One of the first molecular
studies, published in 1967 measured immunological distances (IDs) between different primates.[6] Basically
the study measured the strength of immunological response that an antigen from one species (human
albumin) induces in the immune system of another species (human, chimpanzee, gorilla and Old World
monkeys). Closely related species should have similar antigens and therefore weaker immunological
response to each other's antigens. The immunological response of a species to its own antigens (e.g. human
to human) was set to be 1. The ID between humans and gorillas was determined to be 1.09, that between
humans and chimpanzees was determined as 1.14. However the distance to six different Old World monkeys
was on average 2.46 indicating that the African apes are far closer related to humans than to monkeys. The
authors consider the divergence time between Old World monkeys and hominoids to be 30 million years ago
(MYA), based on fossil data, and the immunological distance was considered to grow at a constant rate.
They concluded that divergence time of humans and the African apes to be roughly ~5 MYA. That was a
surprising result. Most scientists at that time thought that humans and great apes diverged much earlier (>15
MYA). The gorilla was, in ID terms, closer to human than to chimpanzees, however the difference was so
slight that the trichotomy could not be resolved with certainty. Later studies based on molecular genetics
were able to resolve the trichotomy: chimpanzees are phylogenetically closer to humans than to gorillas.
However, the divergence times estimated later (using much more sophisticated methods in molecular
genetics) do not substantially differ from the very first estimate in 1967.
[edit] Divergence times and ancestral effective population size

The sequences of the DNA segments diverge earlier than the species. A large effective population
size in the ancestral population (left) preserves different variants of the DNA segments (=alleles) for a longer
period of time. Therefore, on average, the gene divergence times (t A for DNA segment A; tB for DNA
segment B) will deviate more from the time the species diverge (t S) compared to a small ancestral effective
population size (right).
Current methods to determine divergence times use DNA sequence alignments and molecular
clocks. Usually the molecular clock is calibrated assuming that the orangutan split from the African apes
(including humans) 12-16 MYA. Some studies also include some old world monkeys and set the divergence
time of them from hominoids to 25-30 MYA. Both calibration points are based on very little fossil data and
have been criticized.[7] If these dates are revised, the divergence times estimated from molecular data will
change as well. However, the relative divergence times are unlikely to change. Even if we can't tell absolute
divergence times exactly, we can be pretty sure that the divergence time between chimpanzees and humans
is about sixfold shorter than between chimpanzees (or humans) and monkeys.
One study (Takahata et al., 1995) used 15 DNA sequence from different regions of the genome from
human and chimpanzee and 7 DNA sequences from human, chimpanzee and gorilla.[8] They determined
that chimpanzees are more closely related to humans than gorillas. Using various statistical methods, they
estimated the divergence time human-chimp to be 4.7 MYA and the divergence time between gorillas and
humans (and chimps) to be 7.2 MYA. Additionally they estimated the effective population size of the common
ancestor of humans and chimpanzees to be ~100,000. This was somewhat surprising since the present day
effective population size of humans is estimate to be only ~10,000. If true that means that the human lineage
would have experienced an immense decrease of its effective population size (and thus genetic diversity) in
its evolution. (see Toba catastrophe theory)
A and B are two different loci. In the upper figure they fit to the species tree. The DNA that is present
in today's gorillas diverged earlier from the DNA that is present in today's humans and chimps. Thus both loci
should be more similar between human and chimp than between gorilla and chimp or gorilla and human. In
the lower graph, locus A has a more recent common ancestor in human and gorilla compared to the chimp
sequence. Whereas chimp and gorilla have a more recent common ancestor for locus B. Here the gene trees
are incongruent to the species tree.
Another study (Chen & Li, 2001) sequenced 53 non-repetitive, intergenic DNA segments from a
human, a chimpanzee, a gorilla, and orangutan.[4] When the DNA sequences were concatenated to a single
long sequence, the generated neighbor-joining tree supported the Homo-Pan clade with 100% bootstrap
(that is that humans and chimpanzees are the closest related species of the four). When three species are
fairly closely related to each other (like human, chimpanzee and gorilla), the trees obtained from DNA
sequence data may not be congruent with the tree that represents the speciation (species tree). The shorter
internodal time span (TIN) the more common are incongruent gene trees. The effective population size (N e)
of the internodal population determines how long genetic lineages are preserved in the population. A higher
effective population size causes more incongruent gene trees. Therefore, if the internodal time span is
known, the ancestral effective population size of the common ancestor of humans and chimpanzees can be
calculated.
When each segment was analyzed individually, 31 supported the Homo-Pan clade, 10 supported the
Homo-Gorilla clade, and 12 supported the Pan-Gorilla clade. Using the molecular clock the authors
estimated that gorillas split up first 6.2-8.4 MYA and chimpanzees and humans split up 1.6-2.2 million years
later (internodal time span) 4.6-6.2 MYA. The internodal time span is useful to estimate the ancestral
effective population size of the common ancestor of humans and chimpanzees.
A parsimonious analysis revealed that 24 loci supported the Homo-Pan clade, 7 supported the
Homo-Gorilla clade, 2 supported the Pan-Gorilla clade and 20 gave no resolution. Additionally they took 35
protein coding loci from databases. Of these 12 supported the Homo-Pan clade, 3 the Homo-Gorilla clade, 4
the Pan-Gorilla clade and 16 gave no resolution. Therefore only ~70% of the 52 loci that gave a resolution
(33 intergenic, 19 protein coding) support the 'correct' species tree. From the fraction of loci which did not
support the species tree and the internodal time span they estimated previously, the effective population of
the common ancestor of humans and chimpanzees was estimated to be ~52 000 to 96 000. This value is not
as high as that from the first study (Takahata), but still much higher than present day effective population size
of humans.[dubious – discuss]
A third study (Yang, 2002) used the same dataset that Chen and Li used but estimated the ancestral
effective population of 'only' ~12,000 to 21,000, using a different statistical method.[9]
[edit] Genetic differences between humans and other great apes

The alignable sequences within genomes of humans and chimpanzees differ by about 35 million
single nucleotide substitutions. Additionally about 3% of the complete genomes differ by deletions, insertions
and duplications.[10]
Since mutation rate is relatively constant, roughly one half of these changes occurred in the human
lineage. Only a very tiny fraction of those fixed differences gave rise to the different phenotypes of humans
and chimpanzees and finding those is a great challenge. The vast majority of the differences are neutral and
do not affect the phenotype.[citation needed]
Molecular evolution may act in different ways, through protein evolution, gene loss, differential gene
regulation and RNA evolution. All are thought to have played some part in human evolution.
[edit] Gene loss
Many different mutations can inactivate a gene, but few will change its function in a specific way.
Inactivation mutations will therefore be readily available for selection to act on. Gene loss could thus be a
common mechanism of evolutionary adaptation (the "less-is-more" hypothesis).[11]
80 genes were lost in the human lineage after separation from the last common ancestor with the
chimpanzee. 36 of those were for olfactory receptors. Genes involved in chemoreception and immune
response are overrepresented.[12] Another study estimated that 86 genes had been lost.[13]
[edit] Hair keratin gene KRTHAP1

A gene for type I hair keratin was lost in the human lineage. Keratins are a major component of hairs.
Humans still have nine functional type I hair keratin genes but the loss of that particular gene may have
caused the thinning of human body hair. The gene loss occurred relatively recently in human evolution—less
than 240,000 years ago.[14]
[edit] Myosin gene MYH16

Stedman et al. (2004) stated that the loss of the sarcomeric myosin gene MYH16 in the human
lineage led to smaller masticatory muscles. They estimated that the mutation that led to the inactivation (a
two base pair deletion) occurred 2.4 million years ago, predating the appearance of Homo ergaster/erectus in
Africa. The period that followed was marked by a strong increase in cranial capacity, promoting speculation
that the loss of the gene may have removed an evolutionary constraint on brain size in the genus Homo.[15]
Another estimate for the loss of the MYH16 gene is 5.3 million years ago, long before Homo
appeared.[16]
[edit] Other
• CASPASE12, a cysteinyl aspartate proteinase[17]
[edit] Gene addition

Segmental duplications (SDs or LCRs) have had roles in creating new primate genes and shaping
human genetic variation.
[edit] Selection pressures

Human accelerated regions are areas of the genome that differ between humans and chimpanzees
to a greater extent than can be explained by genetic drift over the time since the two species shared a
common ancestor. These regions show signs of being subject to natural selection, leading to the evolution of
distinctly human traits. Two examples are HAR1F, which is believed to be related to brain development and
HAR2 (a.k.a HACNS1) that may have played a role in the development of the opposable thumb.
[edit] Genetic differences between humans and Neanderthals

An international group of scientists completed a draft sequence of the Neanderthal genome in May
2010. The results indicate some breeding between humans and Neanderthals as the genomes of non-African
humans have 1-4% more in common with Neanderthals than do the genomes of subsaharan Africans.
Neanderthals and most humans share a lactose-intolerant variant of the lactase gene that encodes an
enzyme that is unable to break down lactose in milk after weaning. Humans and Neanderthals also share the
FOXP2 gene variant associated with brain development and with speech in humans, indicating that
Neanderthals may have been able to speak. Chimps have two amino acid differences in FOXP2 compared
with human and Neanderthal FOXP2.[18][19][20]
[edit] Sequence divergence between humans and apes

The draft sequence of the common chimpanzee genome published in the summer 2005 showed the
regions that are similar enough to be aligned with one another account for 2400 million of the human
genome’s 3164.7 million bases[21] – that is, 75.8% of the genome. This 75.8% of the human genome is
1.23% different from the chimpanzee genome in single nucleotide polymorphisms[21] (changes of single
DNA “letters” in the genome). Another type of difference, called indels (insertions/deletions) account for
another ~3 % difference between the alignable sequences.[21] In addition, variation in copy number of large
segments (> 20 kb) of similar DNA sequence provides a further 2.7% difference between the two species.[22]
Hence the total similarity of the genomes could be as low as about 70%.
The figures above do not take into account differences in the organization of the alignable sequences
within the genomes of humans and chimps. Short stretches of alignable sequence may be in very different
orders and locations within the two genomes. At present we cannot fully assess the difference in structure of
the two genomes, because the human genome was used as a scaffold when the chimpanzee draft genome
was assembled. When genomes are sequenced, relatively short sequences of DNA are produced, and these
sequences have to be fitted together like a jigsaw puzzle. This requires multiple overlapping reads to
accurately assemble the overall sequence. The human genome sequence is relatively accurate, with 8 to 9-
fold coverage, but the chimpanzee draft genome only has 3.6-fold coverage.[21] The human genome was
sequenced using a hierarchical shotgun method which can deal with duplications and difficult-to-assemble
sequences better than the whole genome shotgun method that was used for the chimpanzee draft genome.
The human genome was used as a template for the assembly of the draft chimpanzee genome, on the
assumption that the two genomes would be similar.
Almost half of that 1.23% SNP change belongs to the human at 0.53%, whose genetic variance is
lower than a chimp, and just over half to the chimp at 0.7%. If we also take into account that random "genetic
drift" takes up the bulk of the 0.54% difference, then that percentage difference where SNPs have a potential
positive impact on human abilities, is between 0.01% and 0.02%. The bonobo is a sibling species of common
chimpanzee and is genetically about as different from humans as are common chimps.
Percentage sequence divergence between humans and other hominids[4]
Locus Human-Chimp Human-Gorilla Human-Orangutan
Alu elements 2 - -
Non-coding (Chr. Y) 1.68 ± 0.19 2.33 ± 0.2 5.63 ± 0.35
Pseudogenes
1.64 ± 0.10 1.87 ± 0.11 -
(autosomal)
Pseudogenes (Chr. X) 1.47 ± 0.17 - -
Noncoding (autosomal) 1.24 ± 0.07 1.62 ± 0.08 3.08 ± 0.11
Genes (Ks) 1.11 1.48 2.98
Introns 0.93 ± 0.08 1.23 ± 0.09 -
Xq13.3 0.92 ± 0.10 1.42 ± 0.12 3.00 ± 0.18
Subtotal for X 1.16 ± 0.07 1.47 ± 0.08 -

chromosome
Genes (Ka) 0.8 0.93 1.96
The sequence divergence has generally the following pattern: Human-Chimp < Human-Gorilla <<
Human-Orangutan, highlighting the close kinship between humans and the African apes. Alu elements
diverge quickly due to their high frequency of CpG dinucleotides which mutate roughly 10 times more often
than the average nucleotide in the genome. The mutation rate is higher in the male germ line, therefore the
divergence in the Y chromosome—which is inherited solely from the father—is higher than in autosomes. The
X chromosome is inherited twice as often through the female germ line as through the male germ line and
therefore shows slightly lower sequence divergence. The sequence divergence of the Xq13.3 region is
surprisingly low between humans and chimpanzees.[23]
Mutations altering the amino acid sequence of proteins (Ka) are the least common. In fact ~29% of all
orthologous proteins are identical between human and chimpanzee. The typical protein differs by only two
amino acids.[10]
The measures of sequence divergence shown in the table only take the substitutional differences, for
example from an A (adenine) to a G (guanine), into account. DNA sequences may however also differ by
insertions and deletions (indels) of bases. These are usually stripped from the alignments before the
calculation of sequence divergence is performed. The overall sequence divergence between humans and
chimpanzees for example is close to 5% if indels would be included.
[edit] Modern humans
Map of the migration of modern humans out of Africa, based on mitochondrial DNA. Coloured rings
indicate years before present, in thousands.
Molecular biologists starting with Wesley Brown[24] on mtDNA and Allan Wilson[25] on mtDNA have
produced observations relevant to human evolution.
[edit] Age of the common ancestor

By estimating the rate at which mutations occur in mtDNA, the age of the common ancestral mtDNA
type can be estimated: "the common ancestral mtDNA (type a) links mtDNA types that have diverged by an
average of nearly 0.57%. Assuming a rate of 2%-4% per million years, this implies that the common ancestor
of all surviving mtDNA types existed 140,000-290,000 years ago."[25] This observation is robust, and this
common direct female line ancestor (or mitochondrial most recent common ancestor (mtMRCA)) of all extant
humans has become known as Mitochondrial Eve. The observation that the mtMRCA is the direct matrilineal
ancestor of all living humans does not mean either that she was the first anatomically modern human, nor
that no other female humans lived concurrently with her. Other women would have lived at the same time
and passed nuclear genes down to living humans, but their mitochondrial lineages were lost over time. This
could be due to random events such as producing only male children.
[edit] African origin for modern humans

There is evidence that modern human mtDNA has an African origin: "We infer from the tree of
minimum length... that Africa is a likely source of the human mitochondrial gene pool. This inference comes
from the observation that one of the two primary branches leads exclusively to African mtDNAs... while the
second primary branch also leads to African mtDNAs... By postulating that the common ancestral mtDNA...
was African, we minimize the number of intercontinental migrations needed to account for the geographic
distribution of mtDNA types."[25]
The broad study of African genetic diversity headed by Sarah Tishkoff found the San people to
express the greatest genetic diversity among the 113 distinct populations sampled, making them one of 14
"ancestral population clusters". The research also located the origin of modern human migration in south-
western Africa, near the coastal border of Namibia and Angola.[26]
[edit] Y chromosome findings

The Y chromosome is much larger than mtDNA, and is relatively homogeneous; therefore it has
taken much longer to find distinct lineages and to analyse them. Conversely, because the Y chromosome is
so large by comparison, it holds more genetic information. Y chromosome studies show similar findings to
those made with mtDNA. The estimate for the age of the ancestral Y chromosome for all extant Y
chromosomes is given at about 70,000 years ago and is also placed in Africa; the individual who contributed
this Y chromosomal heritage is sometimes referred to as Y chromosome Adam. The difference in dates
between Y chromosome Adam and mitochondrial Eve is usually attributed to a higher extinction rate for Y
chromosomes due to greater differential reproductive success between individual men, which means that a
small number of very successful men may produce many children, while a larger number of less successful
men will produce far fewer children.
[edit] See also

• The Journey of Man: A Genetic Odyssey
• Chromosome 2 (human)
• Chimpanzee genome project
• Y-DNA haplogroups by ethnic groups
• List of haplogroups of historical and famous figures
• Genetic history of indigenous peoples of the Americas
• Genetic history of the British Isles
• Archaeogenetics of the Near East
• Genetics and archaeogenetics of South Asia
• Genetic history of Europe
• Genetic history of Italy
• Race and genetics
[edit] References
1. ^ Murphy, W.J.; Eizirik, E.; O'Brien, S.J.; Madsen, O.; Scally, M.; Douady, C.J.; Teeling, E.;
Ryder, O.A.; Stanhope, M.J.; de Jong, W.W. & Springer, M.S. (2001). "Resolution of the early
placental mammal radiation using Bayesian phylogenetics". Science 294 (5550): 2348–2351.
doi:10.1126/science.1067179. PMID 11743200.
2. ^ Kriegs, J.O.; Churakov, G.; Kiefmann, M.; Jordan, U.; Brosius, J. & Schmitz, J. (2006).
"Retroposed elements as archives for the evolutionary history of placental mammals". PLoS Biol 4
(4): e91. doi:10.1371/journal.pbio.0040091. PMID 16515367.
3. ^ "Human Chromosome 2." PBS.
4. ^ a b c Chen, F.C. & Li, W.H. (2001). "Genomic divergences between humans and other
hominoids and the effective population size of the common ancestor of humans and chimpanzees".
Am J Hum Genet 68 (2): 444–456. doi:10.1086/318206. PMID 11170892. PMC 1235277.
5. ^ Ken Miller in the Kitzmiller v. Dover trial transcripts.
6. ^ Sarich, V.M. & Wilson, A.C. (1967). "Immunological time scale for hominid evolution".
7. ^ Yoder, A.D. & Yang, Z. (1 July 2000). "Estimation of primate speciation dates using local
molecular clocks". Mol Biol Evol 17 (7): 1081–1090. PMID 10889221.
8. ^ Takahata, N.; Satta, Y. & Klein, J. (1995). "Divergence time and population size in the
lineage leading to modern humans". Theor Popul Biol 48 (2): 198–221. doi:10.1006/tpbi.1995.1026.
PMID 7482371.
9. ^ Yang, Z. (1 December 2002). "Likelihood and Bayes estimation of ancestral population
sizes in hominoids using data from multiple loci" (abstract page). Genetics 162 (4): 1811–1823.
PMID 12524351. PMC 1462394. http://www.genetics.org/cgi/content/abstract/162/4/1811.
10.^ a b Chimpanzee Sequencing & Analysis Consortium (2005). "Initial sequence of the
chimpanzee genome and comparison with the human genome". Nature 437 (7055): 69–87.
doi:10.1038/nature04072. PMID 16136131.
11.^ Olson, M.V. (1999). "When less is more: gene loss as an engine of evolutionary change".
Am J Hum Genet 64 (1): 18–23. doi:10.1086/302219. PMID 9915938. PMC 1377697.
12.^ Wang, X.; Grus, W.E. & Zhang, J. (2006). "Gene losses during human origins". PLoS Biol 4
13.^ "The Evolution of Mammalian Gene Families". Public Library of Science (PLoS) Biology 1:
e85. December 2006.
14.^ Winter, H.; Langbein, L.; Krawczak, M.; Cooper, D.N.; Suarez, L.F.J.; Rogers, M.A.;
Praetzel, S.; Heidt, P.J. & Schweizer, J. (2001). "Human type I hair keratin pseudogene phihHaA has
functional orthologs in the chimpanzee and gorilla: evidence for recent inactivation of the human
gene after the Pan-Homo divergence". Hum Genet 108 (1): 37–42. doi:10.1007/s004390000439.
PMID 11214905.
15.^ Stedman, H.H.; Kozyak, B.W.; Nelson, A.; Thesier, D.M.; Su, L.T.; Low, D.W.; Bridges,
C.R.; Shrager, J.B.; Purvis, N.M. & Mitchell, M.A. (2004). "Myosin gene mutation correlates with
anatomical changes in the human lineage". Nature 428 (6981): 415–418. doi:10.1038/nature02358.
PMID 15042088.
16.^ Perry, G.H.; Verrelli, B.C. & Stone, A.C. (2005). "Comparative analyses reveal a complex
history of molecular evolution for human MYH16". Mol Biol Evol 22 (3): 379–382.
doi:10.1093/molbev/msi004. PMID 15470226.
17.^ Wang, Xiaoxia; Grus, WE; Zhang, J (2006). "Gene Losses during Human Origins". PLoS
Biology 4 (3): e52. doi:10.1371/journal.pbio.0040052. PMID 16464126.
18.^ Saey, Tina H. (2009). "Team decodes Neandertal DNA". Science News 175 (6): 5–6.
19.^ Green, Richard E., et. al. (2010-05-07). "A Draft Sequence of the Neandertal Genome".
20.^ "NEANDERTALS LIVE!". john hawks weblog.
http://johnhawks.net/weblog/reviews/neandertals/neandertal_dna/neandertals-live-genome-
sequencing-2010.html. Retrieved 2010-12-31.
21.^ a b c d The Chimpanzee Sequencing and Analysis Consortium (2005). "Initial sequence of
the chimpanzee genome and comparison with the human genome". Nature 437 (1 September 2005):
69–87. doi:10.1038/nature04072. PMID 16136131.
22.^ Cheng, Z.; Ventura, M.; She, X.; Khaitovich, P.; Graves, T.; Osoegawa, K.; Church, D;
Pieter DeJong, P.; Wilson, R. K.; Paabo, S.; Rocchi, M & Eichler, E. E. (2005). "A genome-wide
comparison of recent chimpanzee and human segmental duplications". Nature 437 (1 September
2005): 88–93. doi:10.1038/nature04000. PMID 16136132.
23.^ Kaessmann, H.; Heissig, F.; von Haeseler, A. & Pääbo, S. (1999). "DNA sequence
variation in a non-coding region of low recombination on the human X chromosome". Nat Genet 22
(1): 78–81. doi:10.1038/8785. PMID 10319866.
24.^ Wesley M. Brown (1980) [1] in Procedings 77: 3605-3609 PubMed
25.^ a b c Rebecca L. Cann, Mark Stoneking, Allan C. Wilson (1987) Mitochondrial DNA and
human evolution in Nature 325: 31-36 PubMed
• Schwartz M, Vissing J (2002) Paternal Inheritance of Mitochondrial DNA. N Engl J Med
347:576-580

• Human Ancestors May Have Interbred With Chimpanzees
Hominini
Australopithecines


[hide]v · d · eHuman genetics
Human evolutionary genetics · Human genetic engineering · Human genetic variation · Human
genetic clustering · Human genome · Human evolution
Archaeogenetics of... Americas · British Isles · Europe · Italy · Near East · South Asia
Retrieved from "http://en.wikipedia.org/wiki/Human_evolutionary_genetics"
Categories: Human genetics | Human evolution | Modern human genetic history

W000
Chloroplast

The simplified internal structure of a chloroplast
Chloroplasts ( /ˈklɒrəplæsts/) are organelles found in plant cells and other eukaryotic organisms
that conduct photosynthesis. Chloroplasts capture light energy to conserve free energy in the form of ATP
and reduce NADP to NADPH through a complex set of processes called photosynthesis.[1]
The word chloroplast (χλωροπλάστης) is derived from the Greek words chloros (χλωρός), which
means green, and plastis (πλάστης), which means "the one who forms". Chloroplasts are members of a class
of organelles known as plastids.
Contents
[hide]
• 1 Evolutionary origin
• 2 Structure
• 3 Transplastomic plants
• 4 See also
• 5 Notes
• 6 References
[edit] Evolutionary origin
Chloroplasts visible in the cells of Plagiomnium affine — Many-fruited Thyme-moss

A model chloroplast
Chloroplasts are one of the many different types of organelles in the plant cell. In general, they are
considered to have originated from cyanobacteria through endosymbiosis. This was first suggested by
Mereschkowsky in 1905[2] after an observation by Schimper in 1883 that chloroplasts closely resemble
cyanobacteria.[3] All chloroplasts are thought to derive directly or indirectly from a single endosymbiotic
event (in the Archaeplastida), except for Paulinella chromatophora, which has recently acquired a
photosynthetic cyanobacterial endosymbiont which is not closely related to chloroplasts of other eukaryotes.
[4] In that they derive from an endosymbiotic event, chloroplasts are similar to mitochondria, but chloroplasts
are found only in plants and protista. The chloroplast is surrounded by a double-layered composite
membrane with an intermembrane space; further, it has reticulations, or many infoldings, filling the inner
spaces. The chloroplast has its own DNA,[5] which codes for redox proteins involved in electron transport in
photosynthesis; this is termed the plastome.[6]
In green plants, chloroplasts are surrounded by two lipid-bilayer membranes. They are believed to
correspond to the outer and inner membranes of the ancestral cyanobacterium.[7] Chloroplasts have their
own genome, which is considerably reduced compared to that of free-living cyanobacteria, but the parts that
are still present show clear similarities with the cyanobacterial genome. Plastids may contain 60-100 genes
whereas cyanobacteria often contain more than 1500 genes.[8] Many of the missing genes are encoded in
the nuclear genome of the host. The transfer of nuclear information has been estimated in tobacco plants at
one gene for every 16000 pollen grains.[9]
In some algae (such as the heterokonts and other protists such as Euglenozoa and Cercozoa),
chloroplasts seem to have evolved through a secondary event of endosymbiosis, in which a eukaryotic cell
engulfed a second eukaryotic cell containing chloroplasts, forming chloroplasts with three or four membrane
layers. In some cases, such secondary endosymbionts may have themselves been engulfed by still other
eukaryotes, thus forming tertiary endosymbionts. In the alga Chlorella, there is only one chloroplast, which is
bell-shaped.
In some groups of mixotrophic protists such as the dinoflagellates, chloroplasts are separated from a
captured alga or diatom and used temporarily. These klepto chloroplasts may only have a lifetime of a few
days and are then replaced.[10]
[edit] Structure
Chloroplasts are observable as flat discs usually 2 to 10 micrometers in diameter and 1 micrometer
thick. In land plants, they are, in general, 5 μm in diameter and 2.3 μm thick. The chloroplast is contained by
an envelope that consists of an inner and an outer phospholipid membrane. Between these two layers is the
intermembrane space. A typical parenchyma cell contains about 10 to 100 chloroplasts.
Chloroplast ultrastructure:
1. outer membrane
2. intermembrane space
3. inner membrane (1+2+3: envelope)
4. stroma (aqueous fluid)
5. thylakoid lumen (inside of thylakoid)
6. thylakoid membrane
7. granum (stack of thylakoids)
8. thylakoid (lamella)
9. starch
10. ribosome
11. plastidial DNA
12. plastoglobule (drop of lipids)
The material within the chloroplast is called the stroma, corresponding to the cytosol of the original
bacterium, and contains one or more molecules of small circular DNA. It also contains ribosomes; however
most of its proteins are encoded by genes contained in the host cell nucleus, with the protein products
transported to the chloroplast.
TEM image of a chloroplast

Within the stroma are stacks of thylakoids, the sub-organelles, which are the site of photosynthesis.
The thylakoids are arranged in stacks called grana (singular: granum).[1] A thylakoid has a flattened disk
shape. Inside it is an empty area called the thylakoid space or lumen. Photosynthesis takes place on the
thylakoid membrane; as in mitochondrial oxidative phosphorylation, it involves the coupling of cross-
membrane fluxes with biosynthesis via the dissipation of a proton electrochemical gradient.
In the electron microscope, thylakoid membranes appear as alternating light-and-dark bands, each
0.01 μm thick. Embedded in the thylakoid membrane are antenna complexes, each of which consists of the
light-absorbing pigments, including chlorophyll and carotenoids, as well as proteins that bind the pigments.
This complex both increases the surface area for light capture, and allows capture of photons with a wider
range of wavelengths. The energy of the incident photons is absorbed by the pigments and funneled to the
reaction centre of this complex through resonance energy transfer. Two chlorophyll molecules are then
ionised, producing an excited electron, which then passes onto the photochemical reaction centre.
Recent studies have shown that chloroplasts can be interconnected by tubular bridges called
stromules, formed as extensions of their outer membranes.[11][12] Chloroplasts appear to be able to
exchange proteins via stromules,[13] and thus function as a network.
[edit] Transplastomic plants

Recently, chloroplasts have caught attention by developers of genetically modified plants. In most
flowering plants, chloroplasts are not inherited from the male parent,[14][15] although in plants such as pines,
chloroplasts are inherited from males.[16] Where chloroplasts are inherited only from the female, transgenes
in these plastids cannot be disseminated by pollen. This makes plastid transformation a valuable tool for the
creation and cultivation of genetically modified plants that are biologically contained, thus posing significantly
lower environmental risks. This biological containment strategy is therefore suitable for establishing the
coexistence of conventional and organic agriculture. While the reliability of this mechanism has not yet been
studied for all relevant crop species, recent results in tobacco plants are promising, showing a failed
containment rate of transplastomic plants at 3 in 1,000,000.[15]
[edit] See also
• Chloroplast membrane
• Inner membrane
• Outer membrane
• Calvin cycle
• Light-dependent reaction
• Light-independent reactions
• Mitochondria
• Hydrogenosome
• CoRR Hypothesis
[edit] Notes
• This article incorporates public domain material from the NCBI document "Science
Primer".
[edit] References
1. ^ a b Campbell, Neil A.; Brad Williamson; Robin J. Heyden (2006). Biology: Exploring Life.
Boston, Massachusetts: Pearson Prentice Hall. ISBN 978-0-13-250882-7.
2. ^ Mereschkowsky C (1905). "Über Natur und Ursprung der Chromatophoren im
Pflanzenreiche". Biol Centralbl 25: 593–604.
3. ^ Schimper AFW (1883). "Über die Entwicklung der Chlorophyllkörner und Farbkörper". Bot.
Zeitung 41: 105–14, 121–31, 137–46, 153–62.
4. ^ Patrick J. Keeling (2004). "Diversity and evolutionary history of plastids and their hosts".
American Journal of Botany 91: 1481–1493. doi:10.3732/ajb.91.10.1481.
http://www.amjbot.org/cgi/content/full/91/10/z1481. [dead link]
5. ^ C.Michael Hogan. 2010. Deoxyribonucleic acid. Encyclopedia of Earth. National Council for
Science and the Environment. eds. S.Draggan and C.Cleveland. Washington DC
6. ^ Krause K (September 2008). "From chloroplasts to "cryptic" plastids: evolution of plastid
genomes in parasitic plants". Curr. Genet. 54 (3): 111–21. doi:10.1007/s00294-008-0208-8.
PMID 18696071.
7. ^ Joyard J Block MA, Douce R (1991). "Molecular aspects of plastid envelope biochemistry.".
Eur J Biochem. 199 (3): 489–509. doi:10.1111/j.1432-1033.1991.tb16148.x. PMID 1868841.
8. ^ Martin W, Rujan T, Richly E (September 2002). "Evolutionary analysis of Arabidopsis,
cyanobacterial, and chloroplast genomes reveals plastid phylogeny and thousands of cyanobacterial
genes in the nucleus". Proc. Natl. Acad. Sci. U.S.A. 99 (19): 12246–51. doi:10.1073/pnas.182432999.
PMID 12218172. PMC 129430. Archived from the original on 2010-11-18.
http://www.webcitation.org/5uKjjsvvn.
9. ^ Huang CY, Ayliffe MA, Timmis JN (March 2003). "Direct measurement of the transfer rate
of chloroplast DNA into the nucleus". Nature 422 (6927): 72–6. doi:10.1038/nature01435.
PMID 12594458.
10.^ Skovgaard A (1998). "Role of chloroplast retention in a marine dinoflagellate". Aquatic
Microbial Ecology 15: 293–301. doi:10.3354/ame015293. Archived from the original on 2010-11-18.
http://www.webcitation.org/5uKjjzB0M.
11.^ Köhler RH, Hanson MR (1 January 2000). "Plastid tubules of higher plants are tissue-
specific and developmentally regulated". J. Cell. Sci. 113 (Pt 1): 81–9. PMID 10591627. Archived
from the original on 2010-11-18. http://www.webcitation.org/5uKjkRKBB.
12.^ Gray JC, Sullivan JA, Hibberd JM, Hansen MR (2001). "Stromules: mobile protrusions and
interconnections between plastids". Plant Biology 3: 223–33. doi:10.1055/s-2001-15204.
13.^ Köhler RH, Cao J, Zipfel WR, Webb WW, Hanson MR (June 1997). "Exchange of protein
molecules through connections between higher plant plastids". Science (journal) 276 (5321): 2039–
42. PMID 9197266. Archived from the original on 2010-11-18.
http://www.webcitation.org/5uKjlSNJo.
14.^ Stegemann S, Hartmann S, Ruf S, Bock R (July 2003). "High-frequency gene transfer from
the chloroplast genome to the nucleus". Proc. Natl. Acad. Sci. U.S.A. 100 (15): 8828–33.
doi:10.1073/pnas.1430924100. PMID 12817081. PMC 166398. Archived from the original on 2010-
11-18. http://www.webcitation.org/5uKjlpWlQ. "most angiosperm species inherit their chloroplasts
maternally".
15.^ a b Ruf S, Karcher D, Bock R (April 2007). "Determining the transgene containment level
provided by chloroplast transformation". Proc. Natl. Acad. Sci. U.S.A. 104 (17): 6998–7002.
doi:10.1073/pnas.0700008104. PMID 17420459. PMC 1849964. Archived from the original on 2010-
11-18. http://www.webcitation.org/5uKjn7jeI.
16.^ Powell W, Morgante M, McDevitt R, Vendramin GG, Rafalski JA (August 1995).
"Polymorphic simple sequence repeat regions in chloroplast genomes: applications to the population
genetics of pines". Proc. Natl. Acad. Sci. U.S.A. 92 (17): 7759–63. doi:10.1073/pnas.92.17.7759.
PMID 7644491. "In the pines, the chloroplast genome is transmitted through pollen".
Wikimedia Commons has media related to: Chloroplasts
• Chloroplast - Cell Centered Database

• Chloroplasts and Photosynthesis: The Role of Light from Kimball's Biology Pages
• Clegg MT, Gaut BS, Learn GH, Morton BR (July 1994). "Rates and patterns of chloroplast
DNA evolution". Proc. Natl. Acad. Sci. U.S.A. 91 (15): 6795–801. doi:10.1073/pnas.91.15.6795.
PMID 8041699.
• 3D structures of proteins associated with thylakoid membrane
• Co-Extra research on chloroplast transformation

Endomembrane
Cytoskeleton Microfilaments · Intermediate filaments · Microtubules · Prokaryotic

cytoskeleton
Myofibril

Other internal
Undulipodium: Cilium/Flagellum · Axoneme · Radial spoke

External
Plant physiology



Transpiration

Sporophyte


morphology terms
Category · Portal
Retrieved from "http://en.wikipedia.org/wiki/Chloroplast"
Categories: Organelles | Photosynthesis

W000
Darwin's Black Box


Darwin's Black Box
paperback edition cover
Author Michael J. Behe
Country United States of America
Language English
Subject(s) Intelligent Design
Publisher Free Press
Publication date September 1996
Media type Hardcover, Paperback
ISBN 0-684-82754-9
ISBN
ISBN 978-0-684-82754-4
OCLC Number 34150540
Dewey Decimal 575 20
LC Classification QH367.3 .B43 1996
Followed by The Edge of Evolution

Darwin's Black Box: The Biochemical Challenge to Evolution (1996, first edition; 2006, second
edition) is a book written by Michael J. Behe and published by Free Press in which he presents his notion of
irreducible complexity and claims that its presence in many biochemical systems indicates therefore that they
must be the result of intelligent design rather than evolutionary processes. In 1993, Behe had written a
chapter on blood clotting in Of Pandas and People, presenting essentially the same arguments but without
the name "irreducible complexity",[1] which he later presented in very similar terms in a chapter in Darwin's
Black Box. Behe later agreed that he had written both and agreed to the similarities when he defended
intelligent design at the Kitzmiller v. Dover Area School District trial.[2][3]
The book has been a source of controversy, as the scientific community at large considers many
criticisms of evolution or proponents of intelligent design to be religious, creationism, and pseudoscience.
Common criticisms were that Behe's ideas are not falsifiable, that his definition of an irreducibly complex
system is ambiguous, and that he ignores previous work in biochemical evolution. Though influential within
the intelligent design movement for several years, the book has lost some of its currency as more and more
examples given by Behe as evidence of irreducible complexity have been shown to be explicable by known
evolutionary mechanisms, something Behe conceded under cross examination while testifying as an expert
witness on behalf of the defendants in Kitzmiller v. Dover Area School District.
Contents
[hide]
• 1 Overview
• 2 Reception
• 3 Views of Creation
• 4 Peer review controversy
• 5 References
[edit] Overview
The "black box" in the title refers to the conceptual tool in which, for one reason or another, the
internal workings of a device are taken for granted, so that its function may be discussed.
Behe begins by reminding the general reader of paradigm shifts in the history of science, in which the
foundations and assumptions of theories are examined, sometimes resulting in the rejection of an entire
theory. Behe suggests that such a paradigm shift in biology (and particularly in evolution) is imminent due to
recent discoveries (circa 1996) in biochemistry. Behe acknowledges acceptance of the Theory of Evolution
by "the great majority" of scientists, and states that "most (though not all) do so based on authority."
Behe states that elucidations of the evolutionary history of various biological features typically
assume the existence of certain abilities as their starting point, such as Charles Darwin's example of a cluster
of light-sensitive spots evolving into an eye via a series of intermediate steps. He then points out that Darwin
dismissed the need to explain the origin of the 'simple' light-sensitive spot, summarizes the modern
understanding of the biochemistry of vision and claims that many other evolutionary explanations face a
similar challenge.
Behe next introduces and defines the concept of irreducible complexity as a system with a series of
parts in which the removal of any part causes the entire system to cease functioning, offering a springloaded-
bar mousetrap as a familiar example. In the following chapters, Behe discusses the apparent irreducible
complexity of several biological systems, including the cilium, the bacterial flagellum, blood clotting, the
immune system and intracellular gated and vesicular transport. Behe claims the underlying complexity and
biochemical mechanisms of the systems are vastly under-appreciated, and identifies other, similar systems.
Behe identifies one of the primary counter-arguments of irreducible complexity, gradual adaptation—
that certain systems may have been co-opted from an original, unrelated role to assume a new function as an
irreducibly complex system. He counter-argues that though it is impossible to consider all possible roles for
any component, it is extremely implausible that components can fortuitously change function within a
complex system and that the focus of the theory changes from making to modifying components and
recounts unsuccessful attempts to discover evolutionary pathways for complex systems within scientific
journals. Behe states that though he did identify assertions that evolution had occurred, he found none that
had been supported by experiment or calculation, and concludes the book by offering intelligent design as a
solution to irreducible complexity.
[edit] Reception
Darwin's Black Box was not well received by the scientific community, which overwhelmingly rejected
Behe's premises and arguments. Kenneth Miller described Behe's argument as an updated version of the
argument from design with reference to biochemistry (which was echoed by other reviewers[4][5]), and also
cites areas in biochemistry and the fossil record which demonstrate currently irreducibly complex systems
evolving. Miller also describes Behe's theory as unfalsifiable, arguing that it arbitrarily ignores evidence that
shows the evolution of a biochemical system.[6] On his blog, PZ Myers described it as "...an example of
pseudoscientific dreck that has been enormously influential."[7] In a review for Nature, Jerry Coyne described
the book hailing from 'populist' creationism that failed to deal with the evidence for evolution honestly. Coyne
also accuses Behe of quote mining and using ad hominem attacks against scientists while 'timidly accepting'
evolution.[8] A review on the pro-evolution website talk.origins, described the book as "...an exposition of the
Frontiers of Ignorance" and that within it systems were labeled "irreducibly complex" if Behe was not able to
envision a simpler system that still worked. The review also stated that the theory was unfalsifiable (echoing
Miller[6]), with faulty logic that worked because Behe did not provide crucial facts that would illustrate its
failings.[9] H. Allen Orr has called Behe's argument in the book "...just plain wrong", arguing that gradual
adaptation could produce irreducibly complex systems. Orr points to examples of gradual adaptation already
known (citing to the work of H. J. Muller in the early 20th century[10]). Behe is also criticized for claiming a
conspiracy of silence among scientists regarding the 'failure of Darwinism'.[4]
Richard Dawkins criticized the book for being logically flawed by setting up a false dichotomy in
which Darwinian evolution is rejected despite what he sees as an enormous amount of positive evidence due
to a single apparent failure to explain irreducible complexity. Dawkins further commented that it was an
argument Darwin himself had anticipated, and that the example of a bacterial flagellum used by Behe had in
fact been refuted by Kenneth R. Miller in Kitzmiller v. Dover Area School District.[11]
Behe has responded to some of these criticisms.[12] The politically conservative magazine National
Review voted Darwin's Black Box one of their top 100 non-fiction books of the century.[13] Part of the panel
was Discovery Institute member George Gilder.[14]
[edit] Views of Creation

In a review of Behe's paper 'Design vs. Randomness in Evolution: Where Do the Data Point?', Denis
Lamoureux criticised Darwin's Black Box as having become central to fundamentalist and evangelical anti-
evolution critiques against biological evolution. Behe supports the historically incorrect misrepresentation that
Darwin's views on the origin of life were atheistic, when On the Origin of Species repeatedly refers to a
Creator in a positive and supportive context as impressing laws on matter. Though Behe has avoided
committing himself to the view that God intervenes directly in nature to create purportedly irreducibly complex
structures, Darwin's Black Box briefly speculates that Divine intervention might have caused the direct
creation of a cell from which all of life evolved, supporting creationist views of miraculous acts of creation, but
ironically echoing Darwin's stated "view of life, with its several powers, having been originally breathed into a
few forms or into one". Behe's claim that the creation of an original first cell represents a "gap" in the laws of
nature needing divine intervention appears to be the problematic God of the gaps position which is subject to
the gaps being filled by scientific discoveries. Behe's thesis that irreducible structures are created in "one fell
swoop" is opposed by other biochemists, including many who are devout Christians, and has, it is claimed by
some, no support from the fossil record something Behe would dispute.[15]
[edit] Peer review controversy
In 2005, while testifying for the defense in the Dover trial, Behe claimed under oath that the book had
received a more thorough peer review than a scholarly article in a refereed journal,[16] a claim which
appears to conflict the facts of the book's peer review.[17] Four of the book's five reviewers (Michael
Atchison, Robert Shapiro, K. John Morrow, and Russell Doolittle) have made statements that contradict or
otherwise do not support Behe's claim of the book passing a rigorous peer review.
Michael Atchison
Atchison has stated that he did not review the book at all, but spent 10 minutes on the phone
receiving a brief overview of the book which he then endorsed without ever seeing the text.[18]
Robert Shapiro
Shapiro has said that he reviewed the book, and while he agreed with some of its analysis of
origin-of-life research, he thought its conclusions are false, though the best explanation of the
argument from design that was available.[19] Had the book been submitted to a peer-reviewed journal
and this comment had appeared, the review provided by Shapiro would have forced the conclusions
regarding intelligent design to be changed or removed.[19]
K. John Morrow
Morrow criticized the book as appalling and unsupported, which contributed to the original
publisher turning down the book for publication.[20]
Russell Doolittle
Doolittle, upon whom Behe based much of his discussion of blood clotting, described it as
misrepresenting many important points and disingenuous,[21] which also contributed to the original
publisher turning down the book for publication.[22]
In the same trial, Behe eventually testified under oath that "There are no peer reviewed articles by
anyone advocating for intelligent design supported by pertinent experiments or calculations which provide
detailed rigorous accounts of how intelligent design of any biological system occurred".[23] The result of the
trial was the ruling that intelligent design is not science and is essentially religious in nature.
[edit] References
1. ^ Nicholas J Matzke (September–October 2004). "Design on Trial in Dover, Pennsylvania |
NCSE". National Center for Science Education. http://ncse.com/rncse/24/5/design-trial-dover-
pennsylvania. Retrieved 2009-07-28. "Even Michael Behe's "irreducible complexity" argument
(though not the signature phrase) appears in print for the first time in the second edition of Pandas"
2. ^ Matzke, Nick (Jan 4, 2009). "God of the Gaps…in your own knowledge. Luskin, Behe, &
blood-clotting". The Panda's Thumb (blog). http://pandasthumb.org/archives/2009/01/god-of-the-
gapsin-your-own-knowledge-luskin-behe-blood-clotting.html. Retrieved 2009-01-05.
3. ^ "Kitzmiller v. Dover: Day 11, AM: Michael Behe".
http://www.talkorigins.org/faqs/dover/day11am.html. Retrieved 2009-07-28.
4. ^ a b Dorit, Robert (1997). "A review of Darwin's Black Box". American Scientist
September/October 1997. http://www.americanscientist.org/bookshelf/pub/a-review-of-darwins-
black-box-the-biochemical-challenge-to-evolution-by-michael-j-behe. Retrieved 2009-03-16.
5. ^ Orr, H. Allen (December 1996/January 1997). "Darwin v. Intelligent Design (Again): The
latest attack on evolution is cleverly argued, biologically informed—and wrong". Boston Review 22 (6).
http://bostonreview.net/BR21.6/orr.html. Retrieved 2007-11-02.
6. ^ a b Miller, Kenneth R. (1996). "Darwin's Black Box, reviewed by Kenneth R. Miller".
Creation/Evolution 16: 36–40. http://www.millerandlevine.com/km/evol/behe-review/index.html.
7. ^ Myers, Paul (2006-11-22). "Bad books" (php). Pharyngula.
http://scienceblogs.com/pharyngula/2006/11/bad_books.php. Retrieved 2007-11-02.
8. ^ Coyne, J.A. (1996). "Darwin's Black Box: The Biochemical Challenge to Evolution by MJ
Behe". Nature 383: 227–227. doi:10.1038/383227b0.
http://www.simonyi.ox.ac.uk/dawkins/WorldOfDawkins-archive/Catalano/box/nature.shtml. [dead link]
9. ^ Robison, Keith (1996-12-11 (most recent update)). "Darwin's Black Box: Irreducible
Complexity or Irreproducible Irreducibility?". talk.origins.
http://www.talkorigins.org/faqs/behe/review.html. Retrieved 2007-11-02.
10.^ Muller, H. J. (1939). "Reversibility in evolution considered from the standpoint of genetics".
Biological Reviews 14: 261–80. doi:10.1111/j.1469-185X.1939.tb00934.x.
11.^ Dawkins, Richard (2007-07-01). "Inferior Design". The New York Times.
http://select.nytimes.com/preview/2007/07/01/books/1154680128921.html. Retrieved 2007-11-02.
12.^ Response to critical reviews by:
• Talk.origins:
• Behe, Michael (1996-08-16). "Behe Responds to Postings in Talk Origins
Newsgroup". http://www.arn.org/docs/behe/mb_toresp.htm. Retrieved 2007-11-02.
• H. Allan Orr:
• Behe, Michael. "The Sterility of Darwinism". Boston Review 22 (1).
http://bostonreview.net/BR22.1/behe.html. Retrieved 2007-11-02.
13.^ "The 100 best non-fiction books of the century". National Review.
http://www.nationalreview.com/100best/100_books.html. Retrieved 2009-07-30.
14.^ "George Gilder". Discovery Institute. http://www.discovery.org/scripts/viewDB/index.php?
command=view&id=10&isFellow=true.
15.^ Denis O. Lamoureux (July 1999). "A Black Box or a Black Hole? A Response to Michael
Behe". Canadian Catholic Review. pp. 67–73. http://www.ualberta.ca/~dlamoure/3Behe.htm.
16.^ Behe, Michael (n.d.). "Federal Rule of Civil Procedure 26: Disclosure of Expert Testimony
of Michael Behe" (pdf). National Center for Science Education. http://ncse.com/webfm_send/309.
17.^ Myers, Paul (2005-10-20). "Behe pwnage". Pharyngula.
http://pharyngula.org/index/weblog/comments/behe_pwnage/. Retrieved 2007-11-02.
18.^ Atchison, Michael (2004-06-11). "Mustard Seeds".
http://www.leaderu.com/real/ri9902/atchison.html. Retrieved 2007-11-02.
19.^ a b Evans, Skip (2005-10-22). "Robert Shapiro on Behe and ID". The Panda's Thumb.
http://pandasthumb.org/archives/2005/10/robert-shapiro.html.
20.^ "Comment on 'Robert Shapiro on Behe and ID'". The Panda's Thumb. 2005-10-24.
http://www.pandasthumb.org/archives/2005/10/robert_shapiro.html#comment-50770. Retrieved
2007-11-02.
21.^ Doolittle, Russell (March/February 1997). "A Delicate Balance". Boston Review.
http://bostonreview.net/BR22.1/doolittle.html. Retrieved 2008-12-12.
22.^ Two of Behe's Reviewers Speak Out at the Wayback Machine.
23.^ Kitzmiller v. Dover Area School District/4:Whether ID Is Science, p88
• "The publisher's webpage for Darwin's Black Box".
http://www.simonsays.com/content/content.cfm?sid=33&pid=407734.
• Scanned copy (not all pages included) and reviews at Google books
• McDonald, John. "A reducibly complex mousetrap".
http://udel.edu/~mcdonald/mousetrap.html.
• Ussery, David (2000-08-10 (last updated)). "A biochemists response to the "biochemical
challenge to evolution". http://www.cbs.dtu.dk/staff/dave/Behe.html. Retrieved 2004-11-02.
• "The Mullerian Two-Step: Add a part, make it necessary or, Why Behe's "Irreducible
Complexity" is silly". http://www.talkorigins.org/faqs/comdesc/ICsilly.html.
Retrieved from "http://en.wikipedia.org/wiki/Darwin%27s_Black_Box"
Categories: 1996 books | Intelligent design books | Biochemistry literature

W000
Dwarf elephant

Dwarf elephants
Temporal range: Pleistocene
Dwarf Elephas skeleton cast.
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Proboscidea
Order:
in part
Dwarf elephants are prehistoric members of the order Proboscidea, that, through the process of
allopatric speciation, evolved to a fraction of the size of their immediate ancestors. Insular dwarfism is a
biological phenomenon by which the size of animals isolated on an island shrinks dramatically over time for
the smaller animals have survived because of the underabundance of food.
Fossil remains of dwarf elephants have been found on the Mediterranean islands of Cyprus, Malta
(at Ghar Dalam), Crete, Sicily, Sardinia, the Cyclades Islands and the Dodecanese Islands. Other islands
where dwarf elephants have been found are Sulawesi, Flores, Timor and other islands of the Lesser Sundas.
The Channel Islands of California once supported a dwarf species descended from Columbian mammoths,
[1] while small races of woolly mammoths were once found on Wrangel Island and Saint Paul Island.
Contents
[hide]
• 1 Mediterranean Islands
• 1.1 Sardinia
• 1.2 Sicily & Malta
• 1.3 Crete
• 1.4 Cyprus
• 1.5 Cyclades Islands
• 1.6 Dodecanese Islands
• 2 Channel Islands of California
• 3 Wrangel Island
• 4 Indonesia
• 4.1 Flores
• 4.2 Sulawesi
• 5 See also
• 6 References
[edit] Mediterranean Islands
Elephas Falconeri
Dwarf elephant skeleton of Malta
Dwarf elephants were, after the Messinian salinity crisis, part of the Pleistocene fauna of all the larger
Mediterranean islands, with the apparent exception of Corsica and the Balearics. Mediterranean dwarf
elephants have generally been considered as paleoloxodontine, derived from the continental Straight-tusked
Elephant, Elephas (Palaeoloxodon) antiquus Falconer & Cautley, 1847. An exception is the dwarf Sardinian
Mammoth, Mammuthus lamarmorae (Major, 1883), the only endemic elephant of the Mediterranean islands
belonging to the mammoth line. A DNA research published in 2006 theorized that the Elephas creticus could
be from the mammoth line too. This old theory, proposed by Dorothea Bate as early as 1905, is not widely
accepted. A scientific study of 2007 demonstrates the mistakes of the DNA research of 2006.[2]
During low sea levels, the Mediterranean islands were colonised again and again, giving rise,
sometimes on the same island, to several species (or subspecies) of different body sizes. These endemic
dwarf elephants were taxonomically different on each island or group of very close islands, like the Cyclades
archipelago.
There are many uncertainties about the time of colonisation, the phylogenetic relationships and the
taxonomic status of dwarf elephants on the Mediterranean islands. Extinction of the insular dwarf elephants
has not been correlated with the arrival in the islands of man. Furthermore, it has been suggested by the
paleontologist Othenio Abel in 1914,[3] that the finding of skeletons of such elephants sparked the idea that
they belonged to giant cyclopses, because the center nasal opening was thought to be a cyclopic eye socket.
[edit] Sardinia
• Mammuthus lamarmorae (Major, 1883)
• Elephas (Palaeoloxodon) antiquus (Acconci, 1881)
• Elephas (Palaeoloxodon) melitensis Falconer, 1868
[edit] Sicily & Malta

• Elephas (Palaeoloxodon) antiquus leonardii Aguirre, 1969
• Elephas (Palaeoloxodon) mnaidriensis (Adams, 1874)
• Elephas (Palaeoloxodon) melitensis Falconer, 1868
• Elephas (Palaeoloxodon) falconeri Busk, 1867
[edit] Crete
Skeleton of a Cretan Dwarf Elephant.

• Elephas (Palaeoloxodon) creticus (Bate, 1907)
• Elephas (Palaeoloxodon) creutzburgi (Kuss, 1965)
• Elephas (Palaeoloxodon) chaniensis (Symeonides et al., 2001)[4]
After DNA research, published in 2006, it has been proposed to rename Elephas (Palaeoloxodon)
creticus into Mammuthus creticus (Bate, 1907). Others proposed (in 2002[ citation needed]) to rename all the
described specimens of larger size under the new subspecies name Elephas antiquus creutzburgi (Kuss,
1965). In a recent study of 2007, it was argued for the groundlessness of the theory by Poulakakis et al. in
2006,[5] showing the weak points of that DNA research.[2]
[edit] Cyprus
• Elephas (Palaeoloxodon) cypriotes Bate, 1903
The Cyprus dwarf elephant survived at least until 11,000 BP. Its estimated body weight was only
200 kg, only 2% of its 10,000 kg ancestor. Molars of this dwarf are reduced to approximately 40% the size of
mainland straight-tusked elephants.
Remains of the species were first discovered and recorded by Dorothea Bate in a cave in the Kyrenia
hills of Cyprus in 1902 and reported in 1903.[6][7]
[edit] Cyclades Islands

Remains of paleoloxodontine elephants have been reported from the islands of Delos, Naxos,
Kythnos, Serifos and Milos. The Delos elephant is of similar size to a small Elephas antiquus, while the
Naxos elephant is of similar size to Elephas melitensis. The remains from Kythnos, Serifos and Milos have
not been described.
[edit] Dodecanese Islands

On the island of Rhodes, bones of an endemic dwarf elephant have been discovered. This elephant
was similar in size to Elephas mnaidriensis.
Two groups of remains of dwarf elephants have been found on the island of Tilos. They are similar in
size to Elephas mnaidriensis and the smaller Elephas falconeri, but the two groups indicate sexual
dimorphism.[8][9] The remains had originally been designated to Palaeoloxodon antiquus falconeri (Busk,
1867). However, this name refers to the dwarf elephants from the island of Malta. As a result, since no
migration route between the two islands can be proved, this name should not be used when referring to the
elephant remnants from Tilos, although some scientists have accepted the temporary use of this name until
further material can be examined.[8]
The Tilos dwarf elephant is the first dwarf elephant whose DNA sequence has been studied. The
results of this research are consistent with previous morphological reports, according to which
Palaeoloxodon is more closely related to Elephas than to Loxodonta or Mammuthus. After the study of new
osteological material <Theodorou et al. 2007> that has been excavated in anatomical connection in the
Charkadio Cave on Tilos island the new species name Elephas tiliensis has been assigned to the Tilos dwarf
elephants. It was the latest paleoloxodontine to survive in Europe. They became extinct just less than 4,000
years BP, so this elephant survived well into the Holocene.
[edit] Channel Islands of California

The Columbian mammoth (Mammuthus columbii) produced a separate, isolated population at the
end of the Pleistocene. One of these isolated groups was formed on the Channel Islands of California, most
likely about 40,000 years ago (although the time of isolation is not fully known). Selective forces on the
Channel Islands resulted in smaller animals, forming a new species, the Pygmy Mammoth Mammuthus
exilis. Channel Islands mammoths ranged from 150–190 cm in shoulder height.
[edit] Wrangel Island

During the last ice age, woolly mammoths (Mammuthus primigenius) lived on Wrangel Island in the
Arctic Ocean. It has been shown that mammoths survived on Wrangel Island until 1700 BCE, the most recent
survival of any known mammoth population. They also survived on Saint Paul Island in the Bering Sea until
6000 BCE.[10] Wrangel Island is thought to have become separated from the mainland by 12,000 years BP.
Survival of a mammoth population may be explained by local geographic, topographic and climatic features,
which entailed preservation of communities of steppe plants, as well as a degree of isolation sufficient to
delay colonization by humans. St. Paul Island shares this characteristic of geographic isolation, implying that
human hunting played a role in the disappearance of the woolly mammoth. Wrangel Island mammoths
ranged from 180–230 cm in shoulder height and were for a time considered "dwarf mammoths".[11] However
this classification has been re-evaluated and since the Second International Mammoth Conference in 1999,
these mammoths are no longer considered to be true "dwarf mammoths".
[edit] Indonesia
On Sulawesi and Flores evidence of a succession of distinct endemic island faunas has been found,
including dwarfed elephants, dating until the Middle Pleistocene. Around the early Middle Pleistocene these
dwarfed elephants were replaced by new immigrants of larger to intermediate sizes.
[edit] Flores
The present understanding of the succession of Stegodon species on Flores is that endemic dwarfs,
represented by the Early Pleistocene species Stegodon sondaarii, became extinct around 840,000 years
ago. These dwarf forms were then replaced by the medium to large-sized Stegodon florensis, a species
closely related to the Stegodon trigonocephalus group found both in Java and in the islands of
biogeographical Wallacea, separated by deep water from the Asian and Australian continental shelves. This
Stegodon species went extinct about 12,000 years ago, presumably because of a volcanic eruption.
[edit] Sulawesi
The dwarfed Stegodon sompoensis lived during the Pleistocene on the island of Sulawesi. They had
a shoulder height of only 1.5m.
[edit] See also

• Pygmy elephant
• Insular dwarfism
[edit] References
Paleontology portal
1. ^ "PYGMY MAMMOTH UPDATE". http://www.nps.gov/archive/chis/pygmy.htm.

2. ^ a b Orlando, L., Pagés, M., Calvignac, S., et al. (2007-02-22). "Does the 43bp sequence
from an 800000 year old Cretan dwarf elephantid really rewrite the textbook on mammoths?".
Biology Letters 3 (1): 57–59. doi:10.1098/rsbl.2006.0536.
3. ^ Abel's surmise is noted by Adrienne Mayor, The First Fossil Hunters: Paleontology in
Greek and Roman Times (Princeton University Press) 2000.
4. ^ Symeonides, N. K.; et al. (2001). "New data on Elephas chaniensis (Vamos cave, Chania,
Crete)". In Cavarretta, Giuseppe (ed.), The World of Elephants - International Congress, Rome 2001,
Rome 2001, 510-513. ISBN 88-8080-025-6
5. ^ Poulakakis, N., Parmakelis, A., Lymberakis, P., Mylonas, M., Zouros, E., Reese, D.,
Glaberman, S. & Caccone, A. 2006. Ancient DNA forces reconsideration of evolutionary history of
Mediterranean pygmy elephantids. Biol. Lett. 2, 451–454.
6. ^ Bate, D. M. A.: Preliminary Note on the Discovery of a Pigmy Elephant in the Pleistocene of
Cyprus in Proceedings of the Royal Society of London Vol. 71 (1902 - 1903), pp. 498-500
7. ^ Dorothea Bate, Cyprus work diary 1901–02, 3 volumes, Natural History Museum's earth
sciences library, palaeontology MSS
8. ^ a b Theodorou, G. (1983). The dwarf elephants of the Charkadio cave on the island of Tilos
(Dodekanese, Greece). Phd Thesis Athens University. p. 321 pp.
9. ^ Theodorou, G.E. (1988). "Environmental factors affecting the evolution of islands
endemics: The Tilos example for Greece". Modern Geology 13: 183–188.
10.^ Guthrie, R. Dale (2004-06-17). "Radiocarbon evidence of mid-Holocene mammoths
stranded on an Alaskan Bering Sea island". Nature (Nature Publishing Group) 429 (6993): 746–749.
doi:10.1038/nature02612. PMID 15201907.
http://www.nature.com/nature/journal/v429/n6993/abs/nature02612.html. Retrieved 2008-12-02.
11.^ Vartanyan, S.L., Garutt, V.E., Sher, A.V. (1993). "Holocene dwarf mammoths from Wrangel
Island in the Siberian Arctic". Nature 362: 337–340. doi:10.1038/362337a0.
Retrieved from "http://en.wikipedia.org/wiki/Dwarf_elephant"
Categories: Prehistoric elephants | Holocene extinctions | History of Sardinia | History of Sicily |

History of Malta | Prehistoric Crete | Prehistoric Cyprus | Cyclades | Dodecanese
W000
Homologous recombination

Figure 1. During meiosis, homologous recombination can produce new combinations of genes as
shown here between similar but not identical copies of human chromosome 1.
Homologous recombination is a type of genetic recombination in which nucleotide sequences are
exchanged between two similar or identical molecules of DNA. It is most widely used by cells to accurately
repair harmful breaks that occur on both strands of DNA, known as double-strand breaks. Homologous
recombination also produces new combinations of DNA sequences during meiosis, the process by which
eukaryotes make gamete cells, like sperm and egg cells in animals. These new combinations of DNA
represent genetic variation in offspring, which in turn enables populations to adapt during the course of
evolution.[1] Homologous recombination is also used in horizontal gene transfer to exchange genetic
material between different strains and species of bacteria and viruses.
Although homologous recombination varies widely among different organisms and cell types, most
forms of it involve the same basic steps. After a double-strand break occurs, sections of DNA around the 5'
ends of the break are cut away in a process called resection. In the strand invasion step that follows, an
overhanging 3' end of the broken DNA molecule then "invades" a similar or identical DNA molecule that is not
broken. After strand invasion, one or two cross-shaped structures called Holliday junctions connect the two
DNA molecules. Depending on how the two junctions are cut by enzymes, the type of homologous
recombination that occurs in meiosis results in either chromosomal crossover or non-crossover. Homologous
recombination that occurs during DNA repair tends to result in non-crossover products, in effect restoring the
damaged DNA molecule as it existed before the double-strand break.
Homologous recombination is conserved across all three domains of life as well as viruses,
suggesting that it is a nearly universal biological mechanism. The discovery of genes for homologous
recombination in protists—a diverse group of eukaryotic microorganisms—has been interpreted as evidence
that meiosis emerged early in the evolution of eukaryotes. Since their dysfunction has been strongly
associated with increased susceptibility to several types of cancer, the proteins that facilitate homologous
recombination are topics of active research. Homologous recombination is also used as a technique in
molecular biology for introducing genetic changes into target organisms. The development of gene targeting
techniques that rely on homologous recombination was the subject of the 2007 Nobel Prize for Physiology or
Medicine.
Contents
[hide]
• 1 History and discovery
• 2 In eukaryotes
• 2.1 Timing within the cell cycle
• 2.2 Models
• 2.2.1 DSBR pathway
• 2.2.2 SDSA pathway
• 2.2.3 SSA pathway
• 2.2.4 BIR pathway
• 3 In bacteria
• 3.1 RecBCD pathway
• 3.2 RecF pathway
• 3.3 Branch migration
• 3.4 Resolution
• 3.5 Facilitating genetic transfer
• 4 In viruses
• 5 Effects of dysfunction
• 6 Evolutionary origins
• 7 Technological applications
• 7.1 Gene targeting
• 7.2 Protein engineering
[edit] History and discovery
Figure 2. An early illustration of crossing over from Thomas Hunt Morgan

In the early 1900s, William Bateson and Reginald Punnett found an exception to one of the principles
of inheritance originally described by Gregor Mendel in the 1860s. In contrast to Mendel's notion that traits
are independently assorted when passed from parent to child—for example that a cat's hair color and its tail
length are inherited independent of each other—Bateson and Punnett showed that certain genes associated
with physical traits can be inherited together, or genetically linked.[2][3] In 1911, after observing that linked
traits could on occasion be inherited separately, Thomas Hunt Morgan suggested that "crossovers" can
occur between linked genes,[4] where one of the linked genes physically crosses over to a different
chromosome. Two decades later, Barbara McClintock and Harriet Creighton demonstrated that chromosomal
crossover occurs during meiosis,[5][6] the process of cell division by which sperm and egg cells are made.
Within the same year as McClintock's discovery, Curt Stern showed that crossing over—later called
"recombination"—could also occur in somatic cells like white blood cells and skin cells that divide through
mitosis.[5][7]
In 1947, the microbiologist Joshua Lederberg showed that bacteria—which had been assumed to
reproduce only asexually through binary fission—are capable of genetic recombination, which is more similar
to sexual reproduction. This work established E. coli as a model organism in genetics,[8] and helped
Lederberg win the 1958 Nobel Prize in Physiology or Medicine.[9] Building on studies in fungi, in 1964 Robin
Holliday proposed a model for recombination in meiosis which introduced key details of how the process can
work, including the exchange of material between chromosomes through Holliday junctions.[10] In 1983,
Jack Szostak and colleagues presented a model now known as the DSBR pathway, which accounted for
observations not explained by the Holliday model.[10][11] During the next decade, experiments in
Drosophila, budding yeast and mammalian cells led to the emergence of other models of homologous
recombination, called SDSA pathways, which do not always rely on Holliday junctions.[10]
[edit] In eukaryotes
Homologous recombination is essential to cell division in eukaryotes like plants, animals, fungi and
protists. In cells that divide through mitosis, homologous recombination repairs double-strand breaks in DNA
caused by ionizing radiation or DNA-damaging chemicals.[12] Left unrepaired, these double-strand breaks
can cause large-scale rearrangement of chromosomes in somatic cells,[13] which can in turn lead to cancer.
[14]
In addition to repairing DNA, homologous recombination also helps produce genetic diversity when
cells divide in meiosis to become specialized gamete cells—sperm or egg cells in animals, pollen or ovules in
plants, and spores in fungi. It does so by facilitating chromosomal crossover, in which regions of similar but
not identical DNA are exchanged between homologous chromosomes.[15][16] This creates new, possibly
beneficial combinations of genes, which can give offspring an evolutionary advantage.[17] Chromosomal
crossover begins when a protein called Spo11 makes a targeted double-strand break in DNA.[18] The sites
of these double-strand breaks often occur at recombination hotspots, regions in chromosomes that are about
1,000–2,000 base pairs in length and have high rates of recombination. The absence of a recombination
hotspot between two genes on the same chromosome often means that those genes will be inherited by
future generations in equal proportion. This represents linkage between the two genes greater than would be
expected from genes that independently assort during meiosis.[19]
[edit] Timing within the cell cycle
Figure 3. Homologous recombination repairs DNA before the cell enters mitosis (M phase). It occurs
only during and shortly after DNA replication, during the S and G2 phases of the cell cycle.
Double-strand breaks can be repaired through homologous recombination or through non-

homologous end joining (NHEJ). NHEJ is a DNA repair mechanism which, unlike homologous
recombination, does not require a long homologous sequence to guide repair. Whether homologous
recombination or NHEJ is used to repair double-strand breaks is largely determined by the phase of cell
cycle. Homologous recombination repairs DNA before the cell enters mitosis (M phase). It occurs during and
shortly after DNA replication, in the S and G2 phases of the cell cycle, when sister chromatids are more
easily available.[20] Compared to homologous chromosomes, which are similar to another chromosome but
often have different alleles, sister chromatids are an ideal template for homologous recombination because
they are an identical copy of a given chromosome. In contrast to homologous recombination, NHEJ is
predominant in the G1 phase of the cell cycle, when the cell is growing but not yet ready to divide. It occurs
less frequently after the G1 phase, but maintains at least some activity throughout the cell cycle. The
mechanisms that regulate homologous recombination and NHEJ throughout the cell cycle vary widely
between species.[21]
Cyclin-dependent kinases (CDKs), which modify the activity of other proteins by adding phosphate
groups to (that is, phosphorylating) them, are important regulators of homologous recombination in
eukaryotes.[21] When DNA replication begins in budding yeast, the cyclin-dependent kinase Cdc28 begins
homologous recombination by phosphorylating the Sae2 protein.[22] After being so activated by the addition
of a phosphate, Sae2 uses its endonuclease activity to make a clean cut near a double-strand break in DNA.
This allows a three-part protein known as the MRX complex to bind to DNA, and begins a series of protein-
driven reactions that exchange material between two DNA molecules.[23]
[edit] Models
Two primary models for how homologous recombination repairs double-strand breaks in DNA are the
DSBR pathway (sometimes called the double Holliday junction model) and the synthesis-dependent strand
annealing (SDSA) pathway.[24] The two pathways are similar in their first several steps. After a double-
strand break occurs, the MRX complex (MRN complex in humans) binds to DNA on either side of the break.
Next a resection, in which DNA around the 5' ends of the break is cut back, is carried out in two distinct steps.
In the first step of resection, the MRX complex recruits the Sae2 protein. The two proteins then trim back the
5' ends on either side of the break to create short 3' overhangs of single-strand DNA. In the second step,
5'→3' resection is continued by the Sgs1 helicase and the Exo1 and Dna2 nucleases. As a helicase, Sgs1
"unzips" the double-strand DNA, while Exo1 and Dna2's nuclease activity allows them to cut the single-
stranded DNA produced by Sgs1.[22]
Figure 4. The DSBR and SDSA pathways follow the same initial steps, but diverge thereafter. The
DSBR pathway most often results in chromosomal crossover (bottom left), while SDSA always ends with
non-crossover products (bottom right).
The RPA protein, which has high affinity for single-stranded DNA, then binds the 3' overhangs.[25]
With the help of several other proteins that mediate the process, the Rad51 protein (and Dmc1, in meiosis)
then forms a filament of nucleic acid and protein on the single strand of DNA coated with RPA. This
nucleoprotein filament then begins searching for DNA sequences similar to that of the 3' overhang. After
finding such a sequence, the single-stranded nucleoprotein filament moves into (invades) the similar or
identical recipient DNA duplex in a process called strand invasion. In cells that divide through mitosis, the
recipient DNA duplex is generally a sister chromatid, which is identical to the damaged DNA molecule and
provides a template for repair. In meiosis, however, the recipient DNA tends to be from a similar but not
necessarily identical homologous chromosome.[24] A displacement loop (D-loop) is formed during strand
invasion between the invading 3' overhang strand and the homologous chromosome. After strand invasion, a
DNA polymerase extends the end of the invading 3' strand by synthesizing new DNA. This changes the D-
loop to a cross-shaped structure known as a Holliday junction. Following this, more DNA synthesis occurs on
the invading strand (i.e., one of the original 3' overhangs), effectively restoring the strand on the homologous
chromosome that was displaced during strand invasion.[24]
[edit] DSBR pathway

After the stages of resection, strand invasion and DNA synthesis, the DSBR and SDSA pathways
become distinct.[24] The DSBR pathway is unique in that the second 3' overhang (which was not involved in
strand invasion) also forms a Holliday junction with the homologous chromosome. The double Holliday
junctions are then converted into recombination products by nicking endonucleases, a type of restriction
endonuclease which cuts only one DNA strand. The DSBR pathway commonly results in crossover, though it
can sometimes result in non-crossover products.[26] Because of this tendency for chromosomal crossover,
the DSBR pathway is a likely model of how homologous recombination occurs during meiosis.[15]
Whether recombination in the DSBR pathway results in chromosomal crossover is determined by
how the double Holliday junction is cut, or "resolved". Chromosomal crossover will occur if one Holliday
junction is cut on the crossing strand and the other Holliday junction is cut on the non-crossing strand (in
Figure 4, along the horizontal purple arrowheads at one Holliday junction and along the vertical orange
arrowheads at the other). Alternatively, if the two Holliday junctions are cut on the crossing strands (along the
horizontal purple arrowheads at both Holliday junctions in Figure 4), then chromosomes without crossover
will be produced.[27]
[edit] SDSA pathway

Homologous recombination via the SDSA pathway occurs in cells that divide through mitosis and
results in non-crossover products. In this model, the invading 3' strand is extended along the recipient DNA
duplex by a DNA polymerase, and is released as the Holliday junction between the donor and recipient DNA
molecules slides in a process called branch migration. The newly synthesized 3' end of the invading strand is
then able to anneal to the other 3' overhang in the damaged chromosome through complementary base
pairing. After the strands anneal, a small flap of DNA can sometimes remain. Any such flaps are removed,
and the SDSA pathway finishes with the resealing, also known as ligation, of any remaining single-stranded
gaps.[28]
[edit] SSA pathway
Figure 5. Recombination via the SSA pathway occurs between two repeat elements (purple) on the
same DNA duplex, and results in deletions of genetic material. (Click to view animated diagram in Safari,
Opera or Chrome web browsers.)
The single-strand annealing (SSA) pathway of homologous recombination repairs double-strand
breaks between two repeat sequences. The SSA pathway is unique in that it does not require a separate
similar or identical molecule of DNA, like the DSBR or SDSA pathways of homologous recombination.
Instead, the SSA pathway only requires a single DNA duplex, and uses the repeat sequences as the identical
sequences that homologous recombination needs for repair. The pathway is relatively simple in concept:
after two strands of the same DNA duplex are cut back around the site of the double-strand break, the two
resulting 3' overhangs then align and anneal to each other, restoring the DNA as a continuous duplex.[28][29]
As DNA around the double-strand break is cut back, the single-stranded 3' overhangs being
produced are coated with the RPA protein, which prevents the 3' overhangs from sticking to themselves.[30]
A protein called Rad52 then binds each of the repeat sequences on either side of the break, and aligns them
to enable the two complementary repeat sequences to anneal.[30] After annealing is complete, leftover non-
homologous flaps of the 3' overhangs are cut away by a set of nucleases, known as Rad1/Rad10, which are
brought to the flaps by the Saw1 and Slx4 proteins.[30][31] New DNA synthesis fills in any gaps, and ligation
restores the DNA duplex as two continuous strands.[32] The DNA sequence between the repeats is always
lost, as is one of the two repeats. The SSA pathway is considered mutagenic since it results in such deletions
of genetic material.[28]
[edit] BIR pathway

During DNA replication, double-strand breaks can sometimes be encountered at replication forks as
DNA helicase unzips the template strand. These defects are repaired in the break-induced replication (BIR)
pathway of homologous recombination. The precise molecular mechanisms of the BIR pathway remain
unclear. Three proposed mechanisms have strand invasion as an initial step, but differ in how they model the
migration of the D-loop and later phases of recombination.[33]
The BIR pathway can also help to maintain the length of telomeres, regions of DNA at the end of
eukaryotic chromosomes, in the absence of (or in cooperation with) telomerase. Without working copies of
the telomerase enzyme, telomeres typically shorten with each cycle of mitosis, which eventually blocks cell
division and leads to senescence. In budding yeast cells where telomerase have been inactivated through
mutations, two types of "survivor" cells have been observed to avoid senescence longer than expected by
elongating their telomeres through BIR pathways.[33]
Maintaining telomere length is critical for cell immortalization, a key feature of cancer. Most cancers
maintain telomeres by upregulating telomerase. However, in several types of human cancer, a BIR-like
pathway helps to sustain some tumors by acting as an alternative mechanism of telomere maintenance.[34]
This fact has lead scientists to investigate whether such recombination-based mechanisms of telomere
maintenance could thwart anti-cancer drugs like telomerase inhibitors.[35]
[edit] In bacteria
Figure 6. Crystal structure of two chains of the RecA protein bound to DNA.[36] A double-strand
break and two adjacent 3' overhangs are visible.
Homologous recombination is a major DNA repair process in bacteria. It is also important for
producing genetic diversity in bacterial populations, although the process differs substantially from meiotic
recombination, which brings about diversity in eukaryotic genomes. Homologous recombination has been
most studied and is best understood for Escherichia coli.[37] Double-strand DNA breaks in bacteria are
repaired by the RecBCD pathway of homologous recombination. Breaks that occur on only one of the two
DNA strands, known as single-strand gaps, are thought to be repaired by the RecF pathway.[38] Both the
RecBCD and RecF pathways include a series of reactions known as branch migration, in which single DNA
strands are exchanged between two intercrossed molecules of duplex DNA, and resolution, in which those
two intercrossed molecules of DNA are cut apart and restored to their normal double-stranded state.
[edit] RecBCD pathway
Figure 7A. Molecular model for the RecBCD pathway of recombination. This model is based on
reactions of DNA and RecBCD with ATP in excess over Mg 2+ ions. Step a: RecBCD binds to a double-
stranded DNA end. Step b: RecBCD unwinds DNA. RecD is a fast helicase on the 5’-ended strand, and
RecB is a slower helicase on the 3’-ended strand (that with an arrowhead).[39] This produces two single-
stranded (ss) DNA tails and one ss loop. The loop and tails enlarge as RecBCD moves along the DNA. Step
c: The two tails anneal to produce a second ss DNA loop, and both loops move and grow. Step d: Upon
reaching the Chi hotspot sequence RecBCD nicks the 3’-ended strand. Further unwinding produces a long
3’-ended ss tail with Chi near its end. Step e: RecBCD loads RecA protein onto the Chi tail. At some
undetermined point, the RecBCD subunits disassemble. Step f: The RecA-ssDNA complex invades an intact
homologous duplex DNA to produce a D-loop, which can be resolved into intact, recombinant DNA in two
ways. Step g: The D-loop is cut and anneals with the gap in the first DNA to produce a Holliday junction.
Resolution of the Holliday junction (cutting, swapping of strands, and ligation) at the open arrowheads by
some combination of RuvABC and RecG produces two recombinants of reciprocal type. Step h: The 3’ end of
the Chi tail primes DNA synthesis, from which a replication fork can be generated. Resolution of the fork at
the open arrowheads produces one recombinant (non-reciprocal) DNA, one parental-type DNA, and one
DNA fragment.
Figure 7B. Beginning of the RecBCD pathway. This model is based on reactions of DNA and
RecBCD with Mg2+ ions in excess over ATP. Step 1: RecBCD binds to a DNA double strand break. Step 2:
RecBCD initiates unwinding of the DNA duplex through ATP-dependent helicase activity. Step 3: RecBCD
continues its unwinding and moves down the DNA duplex, cleaving the 3' strand much more frequently than
the 5' strand. Step 4: RecBCD encounters a Chi sequence and stops digesting the 3' strand; cleavage of the
5' strand is significantly increased. Step 5: RecBCD loads RecA onto the 3' strand. Step 6: RecBCD unbinds
from the DNA duplex, leaving a RecA nucleoprotein filament on the 3' tail.[40]
The RecBCD pathway is the main recombination pathway used in bacteria to repair double-strand breaks in
DNA.[41] These double-strand breaks can be caused by UV light and other radiation, as well as chemical
mutagens. Double-strand breaks may also arise by DNA replication through a single-strand nick or gap. Such
a situation causes what is known as a collapsed replication fork and is fixed by several pathways of
homologous recombination including the RecBCD pathway.[42]
In this pathway, a three-subunit enzyme complex called RecBCD initiates recombination by binding
to a blunt or nearly blunt end of a break in double-strand DNA. After RecBCD binds the DNA end, the RecB
and RecD subunits begin unzipping the DNA duplex through helicase activity. The RecB subunit also has a
nuclease domain, which cuts the single strand of DNA that emerges from the unzipping process. This
unzipping continues until RecBCD encounters a specific nucleotide sequence (5'-GCTGGTGG-3') known as
a Chi site.[41]
Upon encountering a Chi site, the activity of the RecBCD enzyme changes drastically.[43][44] DNA
unwinding pauses for a few seconds and then resumes at roughly half the initial speed. This is likely because
the slower RecB helicase unwinds the DNA after Chi, rather than the faster RecD helicase, which unwinds
the DNA before Chi.[39][45] Recognition of the Chi site also changes the RecBCD enzyme so that it cuts the
DNA strand with Chi and begins loading multiple RecA proteins onto the single-stranded DNA with the newly
generated 3' end. The resulting RecA-coated nucleoprotein filament then searches out similar sequences of
DNA on a homologous chromosome. Upon finding such a sequence, the single-stranded nucleoprotein
filament moves into the homologous recipient DNA duplex in a process called strand invasion.[46] The
invading 3' overhang causes one of the strands of the recipient DNA duplex to be displaced, to form a D-
loop. If the D-loop is cut, another swapping of strands forms a cross-shaped structure called a Holliday
junction.[41] Resolution of the Holliday junction by some combination of RuvABC or RecG can produce two
recombinant DNA molecules with reciprocal genetic types, if the two interacting DNA molecules differ
genetically. Alternatively, the invading 3’ end near Chi can prime DNA synthesis and form a replication fork.
This type of resolution produces only one type of recombinant (non-reciprocal).
[edit] RecF pathway

Further information: RecF pathway
Bacteria appear to use the RecF pathway of homologous recombination to repair single-strand gaps
in DNA. When the RecBCD pathway is inactivated by mutations and additional mutations inactivate the
SbcCD and ExoI nucleases, the RecF pathway can also repair DNA double-strand breaks.[47] In the RecF
pathway the RecQ helicase unwinds the DNA and the RecJ nuclease degrades the strand with a 5’ end,
leaving the strand with the 3’ end intact. RecA protein binds to this strand and is either aided by the RecF,
RecO, and RecR proteins or stabilized by them. The RecA nucleoprotein filament then searches for a
homologous DNA and exchanges places with the identical or nearly identical strand in the homologous DNA.
Although the proteins and specific mechanisms involved in their initial phases differ, the two
pathways are similar in that they both require single-stranded DNA with a 3’ end and the RecA protein for
strand invasion. The pathways are also similar in their phases of branch migration, in which the Holliday
junction slides in one direction, and resolution, in which the Holliday junctions are cleaved apart by enzymes.
[48][49] The alternative, non-reciprocal type of resolution may also occur by either pathway.
[edit] Branch migration

Immediately after strand invasion, the Holliday junction moves along the linked DNA during the
branch migration process. It is in this movement of the Holliday junction that base pairs between the two
homologous DNA duplexes are exchanged. To catalyze branch migration, the RuvA protein first recognizes
and binds to the Holliday junction and recruits the RuvB protein to form the RuvAB complex. Two sets of the
RuvB protein, which each form a ring-shaped ATPase, are loaded onto opposite sides of the Holliday
junction, where they act as twin pumps that provide the force for branch migration. Between those two rings
of RuvB, two sets of the RuvA protein assemble in the center of the Holliday junction such that the DNA at
the junction is sandwiched between each set of RuvA. The strands of both DNA duplexes—the "donor" and
the "recipient" duplexes—are unwound on the surface of RuvA as they are guided by the protein from one
duplex to the other.[50][51]
[edit] Resolution
In the resolution phase of recombination, any Holliday junctions formed by the strand invasion
process are cut, thereby restoring two separate DNA molecules. This cleavage is done by RuvAB complex
interacting with RuvC, which together form the RuvABC complex. RuvC is an endonuclease that cuts the
degenerate sequence 5'-(A/T)TT(G/C)-3'. The sequence is found frequently in DNA, about once every 64
nucleotides.[51] Before cutting, RuvC likely gains access to the Holliday junction by displacing one of the two
RuvA tetramers covering the DNA there.[50] Recombination results in either "splice" or "patch" products,
depending on how RuvC cleaves the Holliday junction.[51] Splice products are crossover products, in which
there is a rearrangement of genetic material around the site of recombination. Patch products, on the other
hand, are non-crossover products in which there is no such rearrangement and there is only a "patch" of
hybrid DNA in the recombination product.[52]
[edit] Facilitating genetic transfer

Homologous recombination is an important method of integrating donor DNA into a recipient
organism's genome in horizontal gene transfer, the process by which an organism incorporates foreign DNA
from another organism without being the offspring of that organism. Homologous recombination requires
incoming DNA to be highly similar to the recipient genome, and so horizontal gene transfer is usually limited
to similar bacteria.[53] Studies in several species of bacteria have established that there is a log-linear
decrease in recombination frequency with increasing difference in sequence between host and recipient
DNA.[54][55][56]
In bacterial conjugation, where DNA is transferred between bacteria through direct cell-to-cell
contact, homologous recombination helps integrate foreign DNA into the host genome via the RecBCD
pathway. The RecBCD enzyme promotes recombination after DNA is converted from single-strand DNA–in
which form it originally enters the bacterium–to double-strand DNA during replication. The RecBCD pathway
is also essential for the final phase of transduction, a type of horizontal gene transfer in which DNA is
transferred from one bacterium to another by a virus. Foreign, bacterial DNA is sometimes misincorporated in
the capsid head of bacteriophage virus particles as DNA is packaged into new bacteriophages during viral
replication. When these new bacteriophages infect other bacteria, DNA from the previous host bacterium is
injected into the new bacterial host as double-strand DNA. The RecBCD enzyme then incorporates this
double-strand DNA into the genome of the new bacterial host.[41]
[edit] In viruses
Homologous recombination occurs in several groups of viruses. In DNA viruses such as herpesvirus,
recombination occurs through a break-and-rejoin mechanism like in bacteria and eukaryotes.[57] There is
also evidence for recombination in some RNA viruses, specifically positive-sense ssRNA viruses like
retroviruses, picornaviruses, and coronaviruses. There is controversy over whether homologous
recombination occurs in negative-sense ssRNA viruses like influenza.[58]
In RNA viruses, homologous recombination can be either precise or imprecise. In the precise type of
RNA-RNA recombination, there is no difference between the two parental RNA sequences and the resulting
crossover RNA region. Because of this, it is often difficult to determine the location of crossover events
between two recombining RNA sequences. In imprecise RNA homologous recombination, the crossover
region has some difference with the parental RNA sequences – caused by either addition, deletion, or other
modification of nucleotides. The level of precision in crossover is controlled by the sequence context of the
two recombining strands of RNA: sequences rich in adenine and uracil decrease crossover precision.[59]
Homologous recombination is important in facilitating viral evolution.[59][60] For example, if the
genomes of two viruses with different disadvantageous mutations undergo recombination, then they may be
able to regenerate a fully functional genome. Alternatively, if two similar viruses have infected the same host
cell, homologous recombination can allow those two viruses to swap genes and thereby evolve more potent
variations of themselves.[60]
[edit] Effects of dysfunction

Without proper homologous recombination, chromosomes often incorrectly align for the first phase of
cell division in meiosis. This causes chromosomes to fail to properly segregate in a process called
nondisjunction. In turn, nondisjunction can cause sperm and ova to have too few or too many chromosomes.
Down's syndrome, which is caused by an extra copy of chromosome 21, is one of many abnormalities that
result from such a failure of homologous recombination in meiosis.[51][61]
Deficiencies in homologous recombination have been strongly linked to cancer formation in humans.
For example, each of the cancer-related diseases Bloom's syndrome, Werner's syndrome and Rothmund-
Thomson syndrome are caused by malfunctioning copies of RecQ helicase genes involved in the regulation
of homologous recombination: BLM, WRN and RECQ4, respectively.[62] In the cells of Bloom's syndrome
patients, who lack a working copy of the BLM protein, there is an elevated rate of homologous recombination.
[63] Experiments in mice deficient in BLM have suggested that the mutation gives rise to cancer through a
loss of heterozygosity caused by increased homologous recombination.[64] A loss in heterozygosity refers to
the loss of one of two versions—or alleles—of a gene. If one of the lost alleles helps to suppress tumors, like
the gene for the retinoblastoma protein for example, then the loss of heterozygosity can lead to cancer.
[65]:1236
Decreased rates of homologous recombination cause inefficient DNA repair,[65]:310 which can also
lead to cancer.[66] This is the case with BRCA1 and BRCA2, two similar tumor suppressor genes whose
malfunctioning has been linked with considerably increased risk for breast and ovarian cancer. Cells missing
BRCA1 and BRCA2 have a decreased rate of homologous recombination and increased sensitivity to
ionizing radiation, suggesting that decreased homologous recombination leads to increased susceptibility to
cancer.[66] Because the only known function of BRCA2 is to help initiate homologous recombination,
researchers have speculated that more detailed knowledge of BRCA2's role in homologous recombination
may be the key to understanding the causes of breast and ovarian cancer.[66]
[edit] Evolutionary origins
Figure 8. Protein domains in homologous recombination-related proteins are conserved across the
three main groups of life: archaea, bacteria and eukaryotes.
Based on the similarity of their amino acid sequences, sets of proteins involved in homologous
recombination are thought to share common evolutionary origins.[67] One such set of proteins is the
RecA/Rad51 protein family, which includes the RecA protein from bacteria, the Rad51 and Dmc1 proteins
from eukaryotes and the RadA and RadB proteins from archaea. These proteins play key roles in the
beginning stages of homologous recombination in the organisms that express them. The proteins in the
RecA/Rad51 protein family share a long conserved region known as the RecA/Rad51 domain. Within this
protein domain are two sequence motifs, Walker A and Walker B. The Walker A and B motifs allow members
of the RecA/Rad51 protein family to engage in ATP hydrolysis,[67] which provides energy for the proteins to
drive reactions in homologous recombination.[68]
Studies modeling the evolutionary relationships between the Rad51, Dmc1 and RadA proteins
indicate that they are monophyletic, or that they share a common molecular ancestor.[67] Within this protein
family, Rad51 and Dmc1 are grouped together in a separate clade from RadA. One of the reasons for
grouping these three proteins together is that they all possess a modified helix-turn-helix motif, which helps
the proteins bind to DNA, toward their N-terminal ends.[67] An ancient gene duplication event of a eukaryotic
RecA gene and subsequent mutation has been proposed as a likely origin of the modern RAD51 and DMC1
genes.[67]
The discovery of Dmc1 in several species of Giardia, one of the earliest protists to diverge as a
eukaryote, suggests that meiotic homologous recombination—and thus meiosis itself—emerged very early in
eukaryotic evolution.[69] In addition to research on Dmc1, studies on the Spo11 protein have provided
information on the origins of meiotic recombination.[70] Spo11 is a type II topoisomerase that initiates
homologous recombination in meiosis by making targeted double-strand breaks in DNA.[18] Phylogenetic
trees based on the sequence of genes similar to SPO11 in animals, fungi, plants, protists and archaea have
led scientists to believe that the version Spo11 currently in eukaryotes emerged in the last common ancestor
of eukaryotes and archaea.[70]
[edit] Technological applications
[edit] Gene targeting
Figure 9. As a developing embryo, this chimeric mouse had the agouti coat color gene introduced
into its DNA via gene targeting. Its offspring are homozygous for the agouti gene.
Main article: Gene targeting
Many methods for introducing DNA sequences into organisms to create recombinant DNA and
genetically modified organisms use the process of homologous recombination.[71] Also called gene
targeting, the method is especially common in yeast and mouse genetics. The gene targeting method in
knockout mice uses mouse embryonic stem cells to deliver artificial genetic material (mostly of therapeutic
interest), which represses the target gene of the mouse by the principle of homologous recombination. The
mouse thereby acts as a working model to understand the effects of a specific mammalian gene. In
recognition of their discovery of how homologous recombination can be used to introduce genetic
modifications in mice through embryonic stem cells, Mario Capecchi, Martin Evans and Oliver Smithies were
awarded the 2007 Nobel Prize for Physiology or Medicine.[72]
Advances in gene targeting technologies which hijack the homologous recombination mechanics of
cells are now leading to the development of a new wave of more accurate, isogenic human disease models.
These engineered human cell models are thought to more accurately reflect the genetics of human diseases
than their mouse model predecessors. This is largely because mutations of interest are introduced into
endogenous genes, just as they occur in the real patients, and because they are based on human genomes
rather than rat genomes. Furthermore, certain technologies enable the knock-in of a particular mutation
rather than just knock-outs associated with older gene targeting technologies.
[edit] Protein engineering

Protein engineering with homologous recombination develops chimeric proteins by swapping
fragments between two parental proteins. These techniques exploit the fact that recombination can introduce
a high degree of sequence diversity while preserving a protein's ability to fold into its tertiary structure, or
three-dimensional shape.[73] This stands in contrast to other protein engineering techniques, like random
point mutagenesis, in which the probability of maintaining protein function declines exponentially with
increasing amino acid substitutions.[74] The chimeras produced by recombination techniques are able to
maintain their ability to fold because their swapped parental fragments are structurally and evolutionarily
conserved. These recombinable "building blocks" preserve structurally important interactions like points of
physical contact between different amino acids in the protein's structure. Computational methods like
SCHEMA and statistical coupling analysis can be used to identify structural subunits suitable for
recombination.[75][76][77]
Techniques that rely on homologous recombination have been used to engineer new proteins.[75] In
a study published in 2007, researchers were able to create chimeras of two enzymes involved in the
biosynthesis of isoprenoids, a diverse class of compounds including hormones, visual pigments and certain
pheromones. The chimeric proteins acquired an ability to catalyze an essential reaction in isoprenoid
biosynthesis—one of the most diverse pathways of biosynthesis found in nature—that was absent in the parent
proteins.[78] Protein engineering through recombination has also produced chimeric enzymes with new
function in members of a group of proteins known as the cytochrome P450 family,[79] which in humans is
involved in detoxifying foreign compounds like drugs, food additives and preservatives.[15]
[edit] Cancer therapy

In 2009, researchers reported trial results of a cancer therapy that exploits deficiencies in
homologous recombination in certain types of cancer.[80][81] The drug, named olaparib, a PARP1 inhibitor,
was shown to shrink or stop the growth of tumors from breast, ovarian and prostate cancers caused by
mutations in the BRCA1 or BRCA2 genes. BRCA1 and BRCA2 are necessary for DNA repair by homologous
recombination. When BRCA1 or BRCA2 are absent, another type of DNA repair mechanism called base-
excision repair usually compensates for the lack of DNA repair by homologous recombination. However,
when the PARP1 protein – which is necessary for base-excision repair – is inhibited, DNA repair is drastically
reduced, and the cell dies.[80]
By stopping DNA repair in such a fashion, olaparib applies the concept of synthetic lethality to
specifically target cancer cells. An article in the New England Journal of Medicine noted that exploiting
synthetic lethality was a new direction in anti-cancer drug development.[80] While PARP1 inhibitors
represent a novel approach to cancer therapy, researchers have cautioned that they may prove insufficient
for treating late-stage, metastatic cancers.[80] Cancer cells can become resistant to a PARP1 inhibitor if they
experience deletions of mutations in BRCA2. This undermines the drug's synthetic lethality by restoring
cancer cells' ability to repair DNA by homologous recombination.[82]
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• Animations – homologous recombination : Animations showing several models of homologous
recombination
• Homologous recombination: Tempy & Trun: Animation of the bacterial RecBCD pathway of
homologous recombination
[hide]v · d · eGenetics: homologous recombination / mobile genetic elements
Primarily
prokaryotic Conjugation · Transduction · Transformation · Transposon
Occurs in Transfection · Chromosomal crossover · Gene conversion · Fusion gene ·

eukaryotes Horizontal gene transfer · Sister chromatid exchange
B bsyn: dna (repl, cycl, reco, repr) · tscr (fact, tcrg, nucl, rnat, rept, ptts) · tltn (risu, pttl, nexn) · dnab,
rnab/runp · stru (domn, 1°, 2°, 3°, 4°)
Retrieved from "http://en.wikipedia.org/wiki/Homologous_recombination"
Categories: Genetics | DNA repair

W000
Alan Feduccia

Alan Feduccia is a paleornithologist, specializing in the origins and phylogeny of birds. He is now
Professor Emeritus at the University of North Carolina. Feduccia's principal authored works include two
books, The Age of Birds[1] and The Origin and Evolution of Birds,[2] and numerous papers in various
ornithological and biological journals. Feduccia is best known for his criticisms of the widely-held view that
birds originated from and are deeply nested within Theropoda, and are therefore living theropod dinosaurs.[3]
[4][5][6] He has argued for an alternative theory in which birds share a common stem-ancestor with theropod
dinosaurs among more basal archosaurian lineages, with birds originating from small arboreal archosaurs in
the Triassic.
Contents
[hide]
• 1 Education
• 2 Academic career and research
• 3 Controversy
• 4 References
[edit] Education
Feduccia graduated with a B.S. from Louisiana State University, taking ornithological expeditions to
Honduras, El Salvador and Peru. He received his M.A. and Ph.D. (1969) from the University of Michigan.
[edit] Academic career and research

Feduccia's research has focused on ornithology, evolutionary biology, vertebrate history and
morphogenesis, and the tempo and mode of the Tertiary vertebrate radiation. His early work in the 1970s
focused on clarification of the evolutionary history of modern birds (Neornithes), focusing, in particular, on the
importance of the identification of conserved morphological characters that might elucidate phylogeny more
readily than more functionally correlated characters. Using this approach, in a series of publications,
Feduccia analyzed the morphology of the bony stapes, the ear ossicle of birds, to help elucidate the
interrelationships of passeriform birds.[7][8][9] This approach was extended to the analysis of non-
passeriform birds as well, including owls[10] and the shoebill, also known as the whalebill (Balaeniceps rex).
[11] Other studies in the 1970s focused on the analysis of the Tertiary avian radiation, with a particular focus
on the origin and relationships of waterfowl Anseriformes. Based on his analysis of the osteology of the
Paleocene and Eocene duck Presbyornis,[12][13][14] represented in large quantities from Eocene deposits
from outcrops of the Green River Formation in Utah and Wyoming, Feduccia concluded that Presbyornis
represents a shorebird-duck mosaic and that waterfowl evolved from shorebirds (Charadriiformes).[15][16]
[17][18][19] This is contrary to the more widely held view that waterfowl are most closely related to chickens,
turkeys, and related fowl (Galliformes),[20][21][22][23][24] but Feduccia argues that this alternative
phylogeny is unsupported by fossil evidence, and he suggests that any similarities between anseriform and
galliform birds are attributable to homoplasy.[19][18] Partly based on his analysis of the osteology of
Presbyornis, Feduccia also argued that flamingos, the phylogenetic relationships of which remain disputed,
with some recent studies suggesting a sister-group relationship with grebes,[25] were actually derived from
shorebirds.[19][26][27][15][16][28][29] Feduccia summarized his position in the second edition of his book
The Origin and Evolution of Birds: "The study of Presbyornis planted the idea that shorebirds are the basic
ancestral stock for both flamingolike birds and the anseriformes, ducks and their allies...".[30] Feduccia's
early work on flamingos and waterfowl contributed to the development of his hypothesis that there was an
explosive Tertiary adaptive radiation of neornithine birds following the extinction event at the end of the
Cretaceous.[19][31][32] According to this hypothesis, modern orders of birds initially radiated principally from
a lineage of "transitional shorebirds", represented by the shorebird form-family Graculavidae,[33] from the
Cretaceous-early Paleogene, that managed to survive the Cretaceous extinction event, perhaps through
eking out a living along marginal shoreline environments. This radiation is hypothesized to have been very
rapid, as many orders of modern birds have fossil representatives from the early Tertiary.[34] Feduccia has
suggested that this rapid adaptive radiation of modern birds, compressed into such a short period of geologic
time, might seriously obscure interordinal relationships and make elucidation of the phylogeny of modern
birds particularly difficult, barring the isolation of conserved characters or mosaic fossils demonstrating
transitional character states bridging extant orders.[34] This reiterates an early theme from his research in
the 1970s, in which Feduccia had repeatedly emphasized the importance of homoplasy in evolution, and its
ability to confound the interpretation of phylogeny. This has also been a theme in his study of flightlessness
in birds, a phenomenon the pervasiveness of which has been stressed in his work, and the mechanisms by
which flight is lost, including heterochrony and differential development.[19] Feduccia has argued against the
monophyly of the Ratitae,[19][35] supporting instead independent derivation of ratite lineages, perhaps from
flying ancestral paleognathous taxa, like the Lithornithiformes.[19]
Feduccia is best known for his criticisms of the hypothesis, accepted by most paleontologists,[5][3][4]
[6] that birds originated from and are deeply nested within Theropoda, and are therefore living theropod
dinosaurs. Feduccia's first contribution relative to the origin and early evolution of birds, and their relationship
with dinosaurs, was a critical review of the evidence then available for dinosaurian endothermy in 1973.[36]
In a 1979 paper, Feduccia and Tordoff argued, against the position taken by John Ostrom,[4] that
Archaeopteryx was capable of powered flight, as indicated by the asymmetrical vanes of its primary feathers,
a feature found only in flying birds.[37] In a paper coauthored with Storrs Olson in the same year, Feduccia
noted that the robust furcula of Archaeopteryx could have served as a site of attachment for a well-developed
M. pectoralis major, the principal depressor of the avian wing, responsible for powering the downstroke
during avian flight.[38] Olson and Feduccia concluded that this provided further evidence for the flight
capability of Archaeopteryx. These initial excursions into the subject, and the argument that Archaeopteryx
was clearly a bird, albeit primitive, were expanded upon in Feduccia's 1980 book, The Age of Birds. Feduccia
here criticized the theropod hypothesis for the origin of birds, but his position was largely agnostic, conceding
that there was evidence in support of both a theropod ancestry of birds and an ancestry from more basal
archosaurs, perhaps similar in overall morphological organization to Euparkeria.[39] Feduccia nevertheless
suggested that on the basis of closer stratigraphic fit, ancestry from basal archosaurs rather than from
coelurosaurian theropods might prove a better phylogenetic hypothesis.[39] He thus, essentially, agreed with
the model for the origin of birds proposed by Gerhard Heilmann in his influential 1926 book The Origin of
Birds.[40] Feduccia also criticized "ground-up" theories for the origin of avian flight, arguing on biophysical
grounds that they were implausible, and noting that in other cases in which flight has developed among
vertebrates it has occurred in an arboreal context. He argued, instead, for a "trees-down" model for the origin
of avian flight due to its lack of the biophysical constraints hindering "ground-up" acquisition of flight and due
to the ability to call upon biologically functional stages, represented by living analogues, at each stage in the
evolution of flight. Feduccia's skepticism about the origin of birds from theropods and a "ground-up" origin of
avian flight, which in the absence of any evidence for small, arboreal theropods seemed a concomitant
requirement of that hypothesis, increased following publication of The Age of Birds, culminating in a series of
publications in the latter half of the 1980s and the early 1990s expanding upon arguments presented in The
Age of Birds. In his 1985 contribution to the Eichstatt Archaeopteryx Conference, a major international
meeting on the interpretation and significance of Archaeopteryx, as well as on the origin and early evolution
of birds and avian flight, held in Eichstatt, Germany, Feduccia criticized hypotheses for the evolution of
feathers in non-aerodynamic contexts in endothermic small theropod dinosaurs. He argued that these
hypotheses failed to account for the elaborate aerodynamic architecture of the feather vane and rachis, and
that thermoregulatory functions would have been adequately served by hair, which is a developmentally
simpler structure.[41] In a 1993 paper, Feduccia analyzed claw curvature arcs in the manual and pedal claws
of Archaeopteryx and other birds, and found that Archaeopteryx clustered with other arboreal birds,
suggesting that it was an arboreal animal rather than a terrestrial cursor or a bird which spent any
considerable time on the ground,[42] as is argued by some other workers.[43][44] In other publications in the
early 1990s, Feduccia expanded on earlier arguments for the evolution of feathers in a primarily aerodynamic
rather than thermoregulatory context.[45][46] In 1996, Feduccia published the first edition (second edition in
1999) of The Origin and Evolution of Birds, a comprehensive review of his research on both early avian
evolution and a synopsis of the history of the Tertiary radiation of modern birds. The book presented a
thorough overview of earlier criticisms of the theropod hypothesis for the origin of birds and a "ground-up"
origin of avian flight, expanded on many of those arguments, and presented a series of new arguments
questioning the hypotheses of homology advanced as evidence for the theropod hypothesis. Feduccia
argued that many of the proposed homologous similarities between theropods and birds were ambiguous,
and that other similarities between birds and theropods could plausibly be explained as homoplasy,
particularly those in the hindlimb and pelvis. Feduccia also focused upon the discrepancy between
embryological evidence identifying the digits of the avian manus as the second, third, and fourth of the
primitively pentadactyl archosaur manus, and paleontological evidence indicating that theropod dinosaurs
primitively reduced their fourth and fifth manual digits, eventually retaining only the first, second, and third
(with further reduction in some groups, like tyrannosaurs. This emerged as a principal argument in
Feduccia's research on the origin of birds, and was the subject of developmental studies of the ostrich
definitively identifying first and fifth digital condensations in the embryonic hand, confirming a pentadactyl
ground state for the avian manus with symmetrical reduction, unlike the situation indicated by paleontological
evidence for theropods.[47][48] This conclusion has been supported by some other workers,[49][50] but
others have proposed various responses (see below, under Controversy).
In recent years, Feduccia has argued that the discovery of spectacular new fossils from the
Cretaceous of China, like Microraptor, and other taxa with unambiguous feathers, like the oviraptorosaur
Caudipteryx, suggest that there might have been an extensive, and hitherto unrecognized radiation of cryptic
avian lineages, some of which rapidly lost flight and secondarily adopted a cursorial lifestyle, converging on
theropods. On this argument, very birdlike groups like Dromaeosauridae and Oviraptorosauria, which are
currently considered by most workers to be theropod dinosaurs, are thought actually to represent avian
lineages, probably more derived than Archaeopteryx, that through homoplasy associated with the loss of
flight and secondary acquisition of cursoriality, converged on theropod dinosaurs. Other lineages, like that
represented by Microraptor and Anchiornis, are hypothesized to be have been flighted. This argument
represents a shift from Feduccia's earlier position in the 1990s, as he acknowledged in a 2002 paper where
he first endorsed this view.[51] Feduccia has expanded upon this argument in subsequent papers.[52][53]
Feduccia has appeared frequently on national TV and radio, including NPR, Voice of America, BBC,
CNN, ABC (Australia), NHK (Japan) and McNeil/Lehrer Report, and is a popular university lecturer. Feduccia
served as Chair of the Department of Biology at Chapel Hill from 1997–2002, and prior to that was Chair of
the Division of Natural Sciences. He is an elected Fellow of the American Ornithologists’ Union and the
American Association for the Advancement of Science.
[edit] Controversy
Feduccia's work has been unusually controversial, perhaps because of his skepticism about the
theropod origin of birds, which is accepted by many biologists and by the lay public in general, and which has
historically been a particularly divisive topic in vertebrate zoology.[54][55] Feduccia's principal academic
work, The Origin and Early Evolution of Birds, was well received by some workers,[56] and was winner of the
Association of American Publishers 1996 award for Excellence in Biology. However, it received very negative
reviews from several paleontologists,[57][58] primarily on account of the book's criticisms of the theropod
hypothesis for the origin of birds. Feduccia has been criticized for failing to use cladistics in his studies of the
origin and the evolution of birds,[59][60] and this criticism has been related to further criticisms of,
particularly, his arguments against the theropod origin of birds, as well as some of his hypotheses on the
phylogeny of modern birds. In response to a 2002 paper by Rick Prum in The Auk, the journal of the
American Ornithologists' Union, in which Prum presented a summary of the current state of the theropod
hypothesis for the origin of birds, and urged its acceptance by and integration within ornithology,[61]
Feduccia responded with criticism of this view, arguing that the origin of birds was a complex and as yet
unresolved problem to which the theropod hypothesis as presently formulated was a simplistic answer,
ignoring contrary evidence.[51] Prum responded to this paper and criticized Feduccia's failure to use
cladistics and to specify an explicit alternative sister-group with which to ally birds.[62] He particularly singled
out Feduccia's adoption of the view that some theropod taxa are actually birds that have been mistaken for
theropods through convergence associated with flight loss and secondary adoption of cursoriality. Prum
argued, finally, that Feduccia's methodology and view of the origin and early evolution of birds are
pseudoscientific.
Several of the arguments about whether similarities between birds and theropods are homologous
that have been advanced by Feduccia have been particularly contentious. One example is identification of
the digits of the avian and theropod hands, and whether, and if so by what mechanism, it might be possible to
explain the discrepancy between the conflicting digital identities of tridactyl theropods and birds. Wagner and
Gauthier proposed that a homeotic frame shift, whereby expression domains for groups of genes like the Hox
d group, were repositioned during limb bud development, resulting in the development of the first, second,
and third digits of the archosaur manus from what were originally condensations for the second, third, and
fourth.[63] This view has been supported by some other workers.[64][65] Feduccia has responded to these
counterarguments, and debate continues.[48][52] Another response to Feduccia's digital homology argument
is the counterargument that evidence from the transitional Limusaurus inextricabilis suggests that theropods
too have the three digits II, III and IV.[66]
[edit] References
1. ^ Feduccia, A. (1980). The Age of Birds (1st ed.). Cambridge: Harvard University Press.
ISBN 0674009754.
2. ^ Feduccia, A. (1999) (paperback). The Origin and Evolution of Birds (2nd ed.). New Haven:
Yale University Press. ISBN 0300078617.
3. ^ a b Ostrom, J. H. (1973). "The ancestry of birds". Nature 242: 136. doi:10.1038/242136a0.
4. ^ a b c Ostrom, J. H. (1976). "Archaeopteryx and the origin of birds". Biological Journal of the
Linnean Society 8: 91–182. doi:10.1111/j.1095-8312.1976.tb00244.x.
5. ^ a b Gauthier, J. A. (1986), "Saurischian monophyly and the origin of birds", in Padian, K.,
The Origin of Birds and the Evolution of Flight. Memoirs of the California Academy of Sciences, 8,
California Academy of Sciences, pp. 1–55, ISBN 780940228146
6. ^ a b Padian, Kevin. (2004). "Basal Avialae". In Weishampel, David B.; Dodson, Peter; &
pp. 210–231. ISBN 0-520-24209-2.
7. ^ Feduccia, A. (1974). "Morphology of the bony stapes in New and Old World suboscines:
new evidence for common ancestry" (PDF). Auk 91: 427–429. JSTOR 4084529.
http://elibrary.unm.edu/sora/Auk/v091n02/p0427-p0429.pdf.
8. ^ Feduccia, A. (1975a). "Morphology of the bony stapes (columella) in the Passeriformes and
related groups: evolutionary implications". University of Kansas Museum of Natural History
Miscellaneous Publications 63: 1–34.
9. ^ Feduccia, A. (1975b). "Morphology of the bony stapes in Menuridae and Acanthisittidae:
evidence for oscine affinities" (PDF). Wilson Bulletin 87 (3): 418–420.
http://elibrary.unm.edu/sora/Wilson/v087n03/p0418-p0420.pdf.
10.^ Feduccia, A.; Ferree, C. E. (1978). "Morphology of the bony stapes (columella) in owls:
evolutionary implications". Proceedings of the Biological Society of Washington 91: 431–438.
11.^ Feduccia, A. (1977c). "The whalebill is a stork". Nature 266: 719–720.
doi:10.1038/266719a0.
12.^ Wetmore, A. (1926). "Fossil birds from the Green River deposits of eastern Utah". Annals
of the Carnegie Museum 16: 391–402. ISSN 0097-4463.
13.^ Olson, S. (1994). "A giant Presbyornis (Aves: Anseriformes) and other birds from the
Paleocene Aquia Formation of Maryland and Virginia". Proceedings of the Biological Society of
Washington 107: 429–435. http://hdl.handle.net/10088/6493.
14.^ Ericson, Per G. P. (1997). "Systematic position of the Paleogene family Presbyornithidae
(Aves: Anseriformes)". Zoological Journal of the Linnean Society 121: 429–483. doi:10.1111/j.1096-
3642.1997.tb01286.x.
15.^ a b Feduccia, A. (1977b). "Hypothetical stages in the evolution of modern ducks and
flamingos". Journal of Theoretical Biology 67 (4): 715–721. doi:10.1016/0022-5193(77)90256-9.
PMID 904341.
16.^ a b Feduccia, A. (1978). "Presbyornis and the evolution of ducks and flamingos". American
Scientist 66 (3): 298–304. http://connection.ebscohost.com/content/article/1022217699.html.
17.^ Feduccia (1980)
18.^ a b Olson, S. L.; Feduccia, A. (1980b). "Presbyornis and the origin of the Anseriformes
(Aves: Charadriomorphae)". Smithsonian Contributions to Zoology 323: 1–24.
19.^ a b c d e f g Feduccia (1999)
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(5198): 637–638. doi:10.1126/science.267.5198.637. PMID 17745839.
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35.^ Feduccia, A. 1985b. The morphological evidence for ratite monophyly: fact or fiction?
Proceedings of the Eighteenth International Ornithological Congress , 184-190.
36.^ Feduccia, A. (1973). "Dinosaurs as reptiles". Evolution 27 (1): 166–169.
37.^ Feduccia, A.; Tordoff, H. B. (1979). "Feathers of Archaeopteryx: asymmetric vanes indicate
aerodynamic function". Science 203 (4384): 1021–1022. doi:10.1126/science.203.4384.1021.
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38.^ Olson, S. L.; Feduccia, A. (1979). "Flight capability and the pectoral girdle of
Archaeopteryx". Nature 278 (5701): 247–248. doi:10.1038/278247a0.
39.^ a b Feduccia (1980, p. 37)
40.^ Heilmann, G. 1926. The Origin of Birds. Witherby: London.
41.^ Feduccia, A. 1985. On why the dinosaur lacked feathers. Pages 75-79 in The Beginnings
of Birds: Proceedings of the International Archaeopteryx Conference Eichstatt 1984 (M. K. Hecht, J.
H. Ostrom, G. Viohl, and P. Wellnhofer, eds.). Freunde des Jura-Museums Eichstatt: Eichstatt.
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Archaeopteryx". Science 259 (5096): 790–793. doi:10.1126/science.259.5096.790. PMID 17809342.
43.^ Elzanowski, A. 2002. Archaeopterygidae (Upper Jurassic of Germany). Pages 129-159 in
Mesozoic Birds: Above the Heads of Dinosaurs (L. M. Chiappe and L. M. Witmer, eds.). University of
California Press
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Archaeopteryx". Zoological Journal of the Linnean Society 149: 97–116. doi:10.1111/j.1096-
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Propithecus (Primates, Lemuridae)". Journal of Theoretical Biology 160: 159–164.
doi:10.1006/jtbi.1993.1010.
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misinterpretation". Courier Forschungsinstitut Senckenberg 181: 65–77.
47.^ Burke, A. C.; Feduccia, A. (1997). "Developmental patterns and the identification of
homologies in the avian hand". Science 278: 666–669. doi:10.1126/science.278.5338.666.
48.^ a b Feduccia, A.; Nowicki, J. (2002). "The hand of birds revealed by early ostrich embryos".
Naturwissenschaften 89 (9): 391–393. doi:10.1007/s00114-002-0350-y. PMID 12435089.
49.^ Kundrát, Martin; Seichert, Václav; Russell, Anthony P.; Smetana, Karel (August 2002).
"Pentadactyl pattern of the avian wing autopodium and pyramid reduction hypothesis" (PDF). Journal
of Experimental Zoology 294 (2): 152–159. doi:10.1002/jez.10140. PMID 12210116.
http://www.bio.ucalgary.ca/contact/faculty/pdf/russell/249.pdf.
50.^ Galis, F.; Kundrat, M.; Sinervo, B. (2003). "An old controversy solved: bird embryos have
five fingers". Trends in Ecology and Evolution 18: 7–9. doi:10.1016/S0169-5347(02)00018-6.
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119: 1187–1201. doi:10.1642/0004-8038(2002)119[1187:BADSAT]2.0.CO;2. JSTOR 4090252.
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Testing the hypothesis on morphological and paleontological evidence". Journal of Morphology 266
(2): 125–166. doi:10.1002/jmor.10382. PMID 16217748.
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Groups of Tetrapods: Controversy and Consensus (eds. H.-P. Schultze and L. Trueb). Cornell
University Press: Ithaca
55.^ Witmer, L. M. 2002. The debate on avian ancestry: phylogeny, function, and fossils. Pages
3-30 in Mesozoic Birds: Above the Heads of Dinosaurs (eds. L. M. Chiappe and L. M. Witmer).
University of California Press: Berkeley.
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University Press, 1996)". American Zoologist 37: 210–211. JSTOR 3883960.
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1996)". American Scientist 85: 178–181.
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Evolution of Birds by Alan Feduccia (Yale University Press, 1996)". Nature 384: 230.
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66.^ Xu, X.; Clark, J.M.; Mo, J.; Choiniere, J.; Forster, C.A.; Erickson, G.M.; Hone, D.W.E.;
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(PDF). Nature 459 (18): 940–944. doi:10.1038/nature08124. PMID 19536256.
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%20ceratosaur%202009.pdf.

• Alan Feduccia at the University of North Carolina
Persondata
Name Feduccia, Alan

Alternative names
Short description
Date of birth
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Categories: American ornithologists | Living people

W000
Bryozoa
Bryozoa
Bryozoa
Temporal range: Upper Cambrian–Recent[1]
PreЄ
g
"Bryozoa", from Ernst Haeckel's Kunstformen der Natur, 1904
Kingdom: Animalia
Lophotrochozoa
Superphylum:
[2]
Bryozoa
Phylum:
Ehrenberg, 1831[3]
Classes
[show]
Gymnolaemata
Cheilostomata
Ctenostomata
[show]
Phylactolaemata
Plumatellida
[show]
Stenolaemata
†Cryptostomida
Cyclostomatida
†Cystoporida
†Fenestrida
†Hederellida
†Trepostomatida
Synonyms
Ectoprocta (Nitsche, 1869)[4]

The Bryozoa, also known as Ectoprocta or commonly as moss animals, are a phylum of aquatic
invertebrate animals. Typically about 0.5 millimetres (0.020 in) long, they are filter feeders that sieve food
particles out of the water using a retractable lophophore, a "crown" of tentacles lined with cilia. Most marine
species live in tropical waters, but a few occur in oceanic trenches, and others are found in polar waters. One
class lives only in a variety of freshwater environments, and a few members of a mostly marine class prefer
brackish water. Over 4,000 living species are known. One genus is solitary and the rest colonial.
The phylum was originally called "Polyzoa", but this term was superseded by "Bryozoa" in 1831.
Another group of animals discovered subsequently, whose filtering mechanism looked similar, was also
included in "Bryozoa" until 1869, when the two groups were noted to be very different internally. The more
recently discovered group were given the name Entoprocta, while the original "Bryozoa" were called
"Ectoprocta". However, "Bryozoa" has remained the more widely used term for the latter group.
Individuals in bryozoan (ectoproct) colonies are called zooids, since they are not fully-independent
animals. All colonies contain autozooids, which are responsible for feeding and excretion. Colonies of some
classes have various types of non-feeding specialist zooids, some of which are hatcheries for fertilized eggs,
and some classes also have special zooids for defense of the colony. The class Cheilostomata have the
largest number of species, possibly because they have the widest range of specialist zooids. A few species
can creep very slowly by using spiny defensive zooids as legs. Autozooids supply nutrients to non-feeding
zooids by channels that vary between classes. All zooids, including those of the solitary species, consist of a
cystid that provides the body wall and produces the exoskeleton and a polypide that contains the internal
organs and the lophophore or other specialist extensions. Zooids have no special excretory organs, and the
polypides of autozooids are scrapped when the polypides become overloaded by waste products; usually the
body wall then grows a replacement polypide. In autozooids the gut is U-shaped, with the mouth inside the
"crown" of tentacles and the anus outside it. Colonies take a variety of forms, including fans, bushes and
sheets. The Cheilostomata produce mineralized exoskeletons and form single-layered sheets that encrust
over surfaces.
Zooids of all the freshwater species are simultaneous hermaphrodites. Although those of many
marine species function first as males and then as females, their colonies always contain a combination of
zooids that are in their male and female stages. All species emit sperm into the water. some also release ova
into the water, while others capture sperm via their tentacles to fertilize their ova internally. In some species
the larvae have large yolks, go to feed, and quickly settle on a surface. Others produce larvae that have little
yolk but swim and feed for a few days before settling. After settling, all larvae undergo a radical
metamorphosis that destroys and rebuilds almost all the internal tissues. Freshwater species also produce
statoblasts that lie dormant until conditions are favorable, which enables a colony's lineage to survive even if
severe conditions kill the mother colony.
Predators of marine bryozoans include nudibranchs (sea slugs), fish, sea urchins, pycnogonids,
crustaceans, mites and starfish. Freshwater bryozoans are preyed on by snails, insects, and fish. In
Thailand, many populations of one freshwater species have been wiped out by an introduced species of
snail. A fast-growing invasive bryozoan off the northeast and northwest coasts of the USA has reduced kelp
forests so much that it has affected local fish and invertebrate populations. Bryozoans have spread diseases
to fish farms and fishermen. Chemicals extracted from a marine bryozoan species have been investigated for
treatment of cancer and Alzheimer's Disease, but analyses have not been encouraging.
Mineralized skeletons of bryozoans appear in rocks from Early Ordovician period, about 480 million
years ago, while all other fossilized phyla appear in earlier periods. This has led researchers to suspect that
bryozoans had arisen earlier but were initially unmineralized, and may have differed significantly from
fossilized and modern forms. Early fossils are mainly of erect forms, but encrusting forms gradually became
dominant. It is uncertain whether the phylum is monophyletic. Bryozoans' evolutionary relationships to other
phyla are also unclear, partly because scientists' view of the family tree of animals is mainly influenced by
better-known phyla. Both morphological and molecular phylogeny analyses disagree over bryozoans'
relationships with entoprocts, about whether bryozoans should be grouped with brachiopods and phoronids
in Lophophorata, and whether bryozoans should be considered protostomes or deuterostomes.
Contents
[hide]
• 1 Description
• 1.1 Distinguishing features
• 1.2 Types of zooid
• 1.2.1 Feeding zooids
• 1.2.2 Avicularia and vibracula
• 1.2.3 Other types of colonial zooid
• 1.3 Colony forms and composition
• 2 Taxonomy and Evolution
• 2.1 Classification and diversity
• 2.3 Evolutionary family tree
• 3 Physiology
• 3.1 Feeding and excretion
• 3.2 Respiration and circulation
• 3.3 Reproduction and life cycles
• 4 Ecology
• 4.1 Habitats and distribution
• 4.2 Interactions with non-human organisms
• 4.3 Interaction with humans
• 5 See also
[edit] Description
Freshwater bryozoan with lophophore extended

Bryozoans, phoronids and brachiopods strain food out of the water by means of a lophophore, a
"crown" of hollow tentacles. Bryozoans form colonies consisting of clones called zooids that are typically
about 0.5 millimetres (0.020 in) long.[5] Phoronids resemble bryozoan zooids but are 2 to 20 centimetres
(0.79 to 7.9 in) long and, although they often grow in clumps, do not form colonies consisting of clones.[6]
Brachiopods, generally thought to be closely related to bryozoans and phoronids, are distinguished by having
shells rather like those of bivalves.[7] All three of these phyla have a coelom, an internal cavity lined by
mesothelium.[5][6][7] Some encrusting bryozoan colonies with mineralized exoskeletons look very like small
corals. However, bryozoan colonies are founded by an ancestrula, which is round rather than shaped like a
normal zooid of that species. On the other hand the founding polyp of a coral has a shape like that of its
daughter polyps, and coral zooids have no coelom or lophophore.[8]
Entoprocts, another phylum of filter-feeders, look rather like bryozoans but their lophophore-like
feeding structure has solid tentacles, their anus lies inside rather than outside the base of the "crown" and
they have no coelom.[9]
Summary of distinguishing features
Other
Other lophophorates[10]
Lophotrochozoa
Bryozoa[5]
(Ectoprocta) Phoronida[ Brachiopoda[ Annelida,
6] 7] Mollusca
Three-part, if One per

Coelom the cavity of the Three-part segment in basic form;
epistome is included merged in some taxa
Uncertain
because
Formation of metamorphosis of
Enterocoely Schizocoely
coelom larvae into adults
makes this impossible
to trace
Lophophore With hollow tentacles none fee
wi
Feeding
From tips to bases of tentacles not applicable
current t
Multiciliated
Yes[11] no[11] Yes[1
cells in epithelium
Position of Varies, none Rear end, but

Outside base of lophophore
anus in some species none in Siboglinidae loph
Colonies of
Sessile species often form clumps, but with no active co-
Colonial clones in most; one clone
operation
solitary genus som
Shape of Round, unlike

not applicable
founder zooid normal zooids[8]
Mineralized Some taxa no Bivalve-like Some sessile

exoskeletons shells annelids build
mineralized tubes;[12]
most molluscs have
shells, but most modern
cephalopods have
internal shells or none.
[13]
[edit] Types of zooid

All bryozoans are colonial except for one genus, Monobryozoon.[14][15] Individual members of a
bryozoan colony are about 0.5 millimetres (0.020 in) long and are known as zooids,[5] since they are not
fully-independent animals.[16] All colonies contain feeding zooids, known as autozooids, and those of some
groups also contain non-feeding specialist heterozooids;[15] colony members are genetically identical and
co-operate, rather like the organs of larger animals.[5] What type of zooid grows where in a colony is
determined by chemical signals from the colony as a whole or sometimes in response to the scent of
predators or rival colonies.[15]
The bodies of all types have two main parts. The cystid consists of the body wall and whatever type
of exoskeleton is secreted by the epidermis. The exoskeleton may be organic (chitin, polysaccharide or
protein) or made of the mineral calcium carbonate. The body wall consists of the epidermis, basal lamina (a
mat of non-cellular material), connective tissue, muscles, and the mesothelium which lines the coelom (main
body cavity)[5] – except that in one class, the mesothelium is split into two separate layers, the inner one
forming a membranous sac that floats freely and contains the coelom, and the outer one attached to the body
wall and enclosing the membranous sac in a pseudocoelom.[17] The other main part of the bryozoan body,
known as the polypide and situated almost entirely within the cystid, contains the nervous system, digestive
system, some specialized muscles and the feeding apparatus or other specialized organs that take the place
of the feeding apparatus.[5]
Pharynx
Invert
[edit] Feeding zooids Retractor

muscle
The most common type of zooid is the feeding
autozooid, in which the polypide bears a "crown" of hollow Ovary
tentacles called a lophophore, which captures food
particles from the water.[15] In all colonies a large Protective
percentage of zooids are autozooids, and some consist covering
entirely of autozooids, some of which also engage in Lophophore's
reproduction.[18] tentacles
The basic shape of the "crown" is a full circle. In
Ganglion
the class Phylactolaemata the crown appears U-shaped,
but this impression is created by a deep dent in the rim of Anus
the crown, which has no gap in the fringe of tentacles.[5]
The sides of the tentacles bear fine hairs called cilia, Coelom
whose beating drives a water current from the tips of the (body cavity)
tentacles to their bases, where it exits. Food particles that
Stomach
collide with the tentacles are trapped by mucus, and further
cilia on the inner surfaces of the tentacles convey the Testis
particles towards the mouth, which lies in the center of the
base of the "crown".[19] The method used by ectoprocts is Funiculus
known as "upstream collecting", as food particles are
captured before they pass through the field of cilia that = Digestive tract
creates the feeding current. This method is also used by
phoronids, brachiopods and pterobranchs.[20] = Gonads
= Retractor muscle
= Outer covering
The lophophore and mouth are mounted on a flexible tube, called the "invert" because it can be
turned inside-out and withdrawn into the polypide,[5] rather like the finger of a rubber glove; in this position
the lophophore lies inside the invert and is folded like the spokes of an umbrella. The invert is withdrawn,
sometimes within 60 milliseconds, by a pair of retractor muscles that are anchored at the far end of the
cystid. Sensors at the tips of the tentacles may check for signs of danger before the invert and lophophore
are fully extended. Extension is driven by an increase in internal fluid pressure, which species with flexible
exoskeletons produce by contracting circular muscles that lie just inside the body wall,[5] while species with a
membranous sac use circular muscles to squeeze this.[17] Some species with rigid exoskeletons have a
flexible membrane that replaces part of the exoskeleton, and transverse muscles anchored on the far side of
the exoskeleton increase the fluid pressure by pulling the membrane inwards.[5] In others there is no gap in
the protective skeleton, and the transverse muscles pull on a flexible sac which is connected to the water
outside by a small pore; the expansion of the sac increases the pressure inside the body and pushes the
invert and lophophore out.[5] In some species the retracted invert and lophophore are protected by an
operculum ("lid"), which is closed by muscles and opened by fluid pressure. In one class, a hollow lobe called
the "epistome" overhands the mouth.[5]
The gut is U-shaped, running from the mouth, in the center of the lophophore, down into the animal's
interior and then back to the anus, which is located on the invert, outside and usually below the lophophore.
[5] A network of strands of mesothelium called "funiculi" ("little ropes"[21]) connects the mesothelium
covering the gut with that lining the body wall. The wall of each strand is made of mesothelium, and
surrounds a space filled with fluid, thought to be blood.[5] A colony's zooids are connected, enabling
autozooids to share food with each other and with any non-feeding heterozooids.[5] The method of
connection varies between the different classes of bryozoans, ranging from quite large gaps in the body walls
to small pores through which nutrients are passed by funiculi.[5][17]
There is a nerve ring round the pharynx (throat) and a ganglion that serves as a brain to one side of
this. Nerves run from the ring and ganglion to the tentacles and to the rest of the body.[5] Bryozoans have no
specialized sense organs, but cilia on the tentacles act as sensors. Members of the genus Bugula grow
towards the sun, and therefore must be able to detect light.[5] In colonies of some species, signals are
transmitted between zooids through nerves that pass through pores in the body walls, and coordinate
activities such as feeding and the retraction of lophophores.[5]
The solitary individuals of Monobryozoon are autozooids with pear-shaped bodies. The wider ends
have up to 15 short, muscular projections by which the animals anchor themselves to sand or gravel[22] and
pull themselves through the sediments.[23]
[edit] Avicularia and vibracula

Some authorities use the term avicularia to refer to any type of zooid in which the lophophore is
replaced by an extension that serves some protective function,[18] while others restrict the term to those that
defend the colony by snapping at invaders and small predators, killing some and biting the appendages of
others.[5] In some species the snapping zooids are mounted on a peduncle (stalk), their bird-like appearance
responsible for the term – Charles Darwin described these as like "the head and beak of a vulture in
miniature, seated on a neck and capable of movement".[5][18] Stalked avicularia are placed upside-down on
their stalks.[15] The "lower jaws" are modified versions of the opercula that protect the retracted lophophores
in autozooids of some species, and are snapped shut "like a mousetrap" by similar muscles,[5] while the
beak-shaped upper jaw is the inverted body wall.[15] In other species the avicularia are stationary box-like
zooids laid the normal way up, so that the modified operculum snaps down against the body wall.[15] In both
types the modified operculum is opened by other muscles that attach to it,[18] or by internal muscles that
raise the fluid pressure by pulling on a flexible membrane.[5] The actions of these snapping zooids are
controlled by small, highly modified polypides that are located inside the "mouth" and bear tufts of short
sensory cilia.[5][15] These zooids appear in various positions: some take the place of autozooids, some fit
into small gaps between autozooids, and small avicularia may occur on the surfaces of other zooids.[18]
In vibracula, regarded by some as a type of avicularia, the operculum is modified to form a long
bristle that has a wide range of motion. They may function as defenses against predators and invaders, or as
cleaners. In some species that form mobile colonies, vibracula around the edges are used as legs for
burrowing and walking.[5][18]
[edit] Other types of colonial zooid

Kenozooids (from Greek κενος meaning "empty"[24]) consist only of the body wall and funicular
strands crossing the interior,[5] and no polypide.[15] In some species they form the stems of branching
structures, while in others they act as spacers that enable colonies to grow quickly in a new direction.[15][18]
Spinozooids form defensive spines, and sometimes appear on top of autozooids. Gonozooids act as
brood chambers for fertilized eggs.[15] Some species have miniature nanozooids with small single-tentacled
polypides, and these may grow on other zooids or within the body walls of autozooids that have degenerated.
[18]
[edit] Colony forms and composition

Although zooids are microscopic, colonies range in size from 1 centimeter (0.39 in) to over 1 meter
(3.3 ft).[5] However, the majority are under 10 centimeters (3.9 in) across.[8] The shapes of colonies vary
widely, depend on the pattern of budding by which they grow, the variety of zooids present and the type and
amount of skeletal material they secrete.[5]
Some marine species are bush-like or fan-like, supported by "trunks" and "branches" formed by
kenozooids, with feeding autozooids growing from these. Colonies of these types are generally
unmineralized but may have exoskeletons made of chitin.[5] Others look like small corals, producing heavy
lime skeletons.[25] Many species form colonies which consist of sheets of autozooids. These sheets may
form leaves, tufts or, in the genus Thalmoporella, structures that resemble an open head of lettuce.[5]
The most common marine form, however, is encrusting, in which a one-layer sheet of zooids spreads
over a hard surface or over seaweed. Some encrusting colonies may grow to over 50 centimeters (1.6 ft) and
contain about 2,000,000 zooids.[5] These species generally have exoskeletons reinforced with calcium
carbonate, and the openings through which the lophophores protrude are on the top or outer surface.[5] The
moss-like appearance of encrusting colonies is responsible for the phylum's alternative name Bryozoa
(Ancient Greek words βρυος bryos meaning "moss" and ζωον zoon meaning "animal").[26] Large colonies of
encrusting species often have "chimneys", gaps in the canopy of lophophores, through which they swiftly
expel water that has been sieved, and thus avoid re-filtering water that is already exhausted.[27] They are
formed by patches of non-feeding heterozooids.[28] New chimneys appear near the edges of expanding
colonies, at points where the speed of the outflow is already high, and do not change position if the water
flow changes.[29]
Some freshwater species secrete a mass of gelatinous material, up to 1 meter (3.3 ft) in diameter, to
which the zooids stick. Other freshwater species have plant-like shapes with "trunks" and "branches", which
may stand erect or spread over the surface. A few species can creep at about 2 centimeters (0.79 in) per
day.[5]
Each colony grows by asexual budding from a single zooid known as the ancestrula,[5] which is
round rather than shaped like a normal zooid.[8] This occurs at the tips of "trunks" or "branches" in forms that
have this structure. Encrusting colonies grow round their edges. In species with calcareous exoskeletons,
these do not mineralize until the zooids are fully grown. Colony lifespans range from one to about 12 years,
and the short-lived species pass though several generations in one season.[5]
Species that produce defensive zooids do so only when threats have already appeared, and may do
so within 48 hours.[15] The theory of "induced defenses" suggests that production of defenses is expensive
and that colonies which defend themselves too early or too heavily will have reduced growth rates and
lifespans. This "last minute" approach to defense is feasible because the loss of zooids to a single attack is
unlikely to be significant.[15] Colonies of some encrusting species also produce special heterozooids to limit
the expansion of other encrusting organisms, especially other bryozoans. In some cases this response is
more belligerent if the opposition is smaller, which suggests that zooids on the edge of a colony can
somehow sense the size of the opponent. Some species consistently prevail against certain others, but most
turf wars are indecisive and the combatants soon turn to growing in uncontested areas.[15] Bryozoans
competing for territory do not use the sophisticated techniques employed by sponges or corals, possibly
because the shortness of bryozoan lifespans makes heavy investment in turf wars unprofitable.[15]
[edit] Taxonomy and Evolution

The phylum was originally called "Polyzoa", but this name was soon replaced by Ehrenberg's term
"Bryozoa".[30][31] The name "Bryozoa" was originally applied only to the animals also known as
"Ectoprocta", in which the anus lies outside the "crown" of tentacles (based on the Ancient Greek prefix ἐκτο
meaning "outside" and word πρωκτος meaning "anus").[32] After the discovery of the Entoprocta, in which
the anus lies within a "crown" of tentacles (based on the Ancient Greek prefix ἐντο meaning "inside" and
word πρωκτος meaning "anus"[33]), the name "Bryozoa" was used at phylum level to include the two classes
Ectoprocta and Entoprocta.[34] However, in 1869 Hinrich Nitsche regarded the two groups as quite distinct
for a variety of reasons, and coined the name "Ectoprocta" for Ehrenberg's "Bryozoa".[4][35] Despite their
apparently similar methods of feeding, they differed markedly anatomically; in addition to the different
positions of the anus, ectoprocts have hollow tentacles and a coelom, while entoprocts have solid tentacles
and no coelom. Hence the two groups are now widely regarded as separate phyla, and the name "Bryozoa"
is now synonymous with "Ectoprocta".[34] This has remained the majority view ever since, although most
publications have preferred the name "Bryozoa" rather than "Ectoprocta".[31] Nevertheless some notable
scientists have continued to regard the "Ectoprocta" and Entoprocta as close relatives and group them under
"Bryozoa".[35]
The ambiguity about the scope of the name "Bryozoa" led to proposals in the 1960s and 1970s that it
should be avoided and the unambiguous term "Ectoprocta" should be used.[36] However, the change would
have made it harder to find older works about in which the phylum was called "Bryozoa", and the desire to
avoid ambiguity, if applied consistently to all classifications, would have necessitated renaming of several
other phyla and many lower-level groups.[30] In practice, zoological naming of split or merged groups of
animals is complex and not completely consistent.[37] Works since 2000 have used various names to
resolve the ambiguity, including: "Bryozoa",[5][8] "Ectoprocta",[11][15] "Bryozoa (Ectoprocta)",[17] and
"Ectoprocta (Bryozoa)".[38] Some have used more than one approach in the same work.[39]
The common name "moss animals" is based on the Greek βρυόν (moss) and ζῷα (animals), and
refers to the mossy appearance of encrusting species.[40]
[edit] Classification and diversity

Counts of formally described species range between 4,000 and 4,500.[41] The Gymnolaemata and
especially Cheilostomata have the greatest numbers of species, possibly because of their wide range of
specialist zooids.[15] Under the Linnaean system of classification, which is still used as a convenient way to
label groups of organisms,[42] living members of the phylum Bryozoa are divided into:[5][15]
Class Phylactolaemata Stenolaemata Gymnolaemata
Order Plumatellida[43] Cyclostomata Ctenostomata Cheilostomata
Environments Freshwater Marine Mostly marine
Lip-like
epistome overhanging Yes none
mouth
Gelatinous
Colony Erect,
masses or tubular Erect or encrusting[45]
shapes encrusting or free-living
branching structures[44]
Chitin,
Gelatinous or
Exoskeleton gelatinous or
membranous; Mineralized Mineralized
material membranous;
unmineralized
unmineralized
none[45]
Operculum None in most Yes (except in
none (except in family
("lid") species genus Bugula)
Eleidae[46])
Shape of U-shaped Circular

lophophore appearance
(except in genus
Fredericella, whose
lophophore is circular)
Compressing
Pulling inwards
the membranous sac
How Compressing the Compressing of a flexible section of
(separate inner layer
lophophore extended whole body wall the whole body wall body wall, or making an
of epithelium that lines
internal sac expand.
the coelom)
Limited Stolons and

Full range of
Types of zooid Autozooids only heterozooids, mainly spines as well as
types
gonozooids[47] autozooids[47]
Bryozoan fossils in an Upper Ordovician oil shale (kukersite), northern Estonia.

Fossils of about 15,000 bryozoan species have been found. Marine taxa with mineralized skeletons
appear in rocks dating from the Arenigian stage of the Early Ordovician period, about 480 million years ago.
[8][48] At this point all the modern orders of stenolaemates were present,[49] and the ctenostome order of
gymnolaemates had appeared by the Middle Ordovician, about 465 million years ago. Other types of filter
feeders appeared around the same time, which suggests that some change made the environment more
favorable for this lifestyle.[8] Fossils of cheilostomates, another order of gymnolaemates, first appear in the
Mid Jurassic, about 172 million years ago, and these have been the most abundant and diverse bryozoans
from the Cretaceous to the present.[8] Evidence compiled from the last 100 million years show that
cheilostomates consistently grew over cyclostomates in territorial struggles, which may help to explain how
cheilostomates replaced cyclostomates as the dominant marine bryozoans.[50] Marine fossils from the
Paleozoic era, which ended 251 million years ago, are mainly of erect forms, those from the Mesozoic are
fairly equally divided by erect and encrusting forms, and more recent ones are predominantly encrusting.[51]
Fossils of the soft, freshwater phylactolaemates are very rare,[8] appear in and after the Late Permian (which
began about 260 million years ago) and consist entirely of their durable statoblasts.[44] There are no known
fossils of freshwater members of other classes.[44]
Since all the other phyla that have left fossils are found in Cambrian rocks, it is surprising that the
earliest bryozoan fossil dates from the Ordovician, which immediately followed the Cambrian. This suggests
that the first bryozoans appeared much earlier and were entirely soft-bodied, and the Ordovician fossils
record the appearance of mineralized skeletons in this phylum.[4] The Early Ordovician fossils may also
represent forms that had already become significantly different from the original members of the phylum.[49]
[edit] Evolutionary family tree

Scientists are divided about whether the Bryozoa (Ectoprocta) are a monophyletic group (whether
they include all and only a single ancestor species and all its descendants), about what are the phylum's
closest relatives in the family tree of animals, and even about whether they should be regarded as members
of the protostomes or deuterostomes, the two major groups that account for all moderately complex animals.
An Upper Ordovician cobble with the edrioasteroid Cystaster stellatus and the thin branching
cyclostome bryozoan Corynotrypa. Kope Formation, northern Kentucky.
The traditional view is that the Bryozoa are a monophyletic group, in which the class
Phylactolaemata is most closely related to Stenolaemata and Ctenostomata, the classes that appear earliest
in the fossil record.[52] However, in 2005 a molecular phylogeny study that focused on phylactolaemates
concluded that these are more closely related to the phylum Phoronida, and especially to the only phoronid
species that is colonial, than they are to the other ectoproct classes. That implies that the Entoprocta are not
monophyletic, as the Phoronida are a sub-group of ectoprocts but the standard definition of Entoprocta
excludes the Phoronida.[52] In 2009 another molecular phylogeny study, using a combination of genes from
mitochondria and the cell nucleus, concluded that Bryozoa is a monophyletic phylum, in other words includes
all the descendants of a common ancestor that is itself a bryozoan. The analysis also concluded that the
classes Phylactolaemata, Stenolaemata and Gymnolaemata are also monophyletic, but could not determine
whether Stenolaemata are more closely related to Phylactolaemata or Gymnolaemata. The Gymnolaemata
are traditionally divided into the soft-bodied Ctenostomata and mineralized Cheilostomata, but the 2009
analysis considered it more likely that neither of these orders is monophyletic and that mineralized skeletons
probably evolved more than once within the early Gymnolaemata.[4]
Bryozoans' relationships with other phyla are uncertain and controversial. Traditional phylogeny,
based on anatomy and on the development of the adult forms from embryos, has produced no enduring
consensus about the position of ectoprocts.[11] Attempts to reconstruct the family tree of animals have
largely ignored ectoprocts and other "minor phyla", which have received little scientific study because they
are generally tiny, have relatively simple body plans, and have little impact on human economies – despite
the fact that the "minor phyla" include most of the variety in the evolutionary history of animals.[53] In the
opinion of Ruth Dewel, Judith Winston and Frank McKinney, "Our standard interpretation of bryozoan
morphology and embryology is a construct resulting from over 100 years of attempts to synthesize a single
framework for all invertebrates," and takes little account of some peculiar features of ectoprocts.[49] In
ectoprocts, all of the larva's internal organs are destroyed during the metamorphosis to the adult form and the
adult's organs are built from the larva's epidermis and mesoderm, while in other bilaterians some organs
including the gut are built from endoderm. In most bilaterian embryos the blastopore, a dent in the outer wall,
deepens to become the larva's gut, but in ectoprocts the blastopore disappears and a new dent becomes the
point from which the gut grows. The ectoproct coelom is formed by neither of the processes used by other
bilaterians, enterocoely, in which pouches that form on the wall of the gut become separate cavities, nor
schizocoely, in which the tissue between the gut and the body wall splits, forming paired cavities.[49]
Molecular phylogeny, which attempts to work out the evolutionary family tree of organisms by
comparing their biochemistry and especially their genes, has done much to clarify the relationships between
the better-known invertebrate phyla.[34] However, the shortage of genetic data about "minor phyla" such as
bryozoans and entoprocts has left their relationships to other groups unclear.[35]
When entoprocts were discovered in the nineteenth century, they and bryozoans (ectoprocts) were
regarded as classes within the phylum Bryozoa, because both groups were sessile animals that filter-fed by
means of a crown of tentacles that bore cilia. However, from 1869 onwards increasing awareness of
differences, including the position of the entoproct anus inside the feeding structure and the difference in the
early pattern of division of cells in their embryos, caused scientists to regard the two groups as separate
phyla,[35] and "Bryozoa" became just an alternative name for ectoprocts, in which the anus is outside the
feeding organ.[34] A series of molecular phylogeny studies from 1996 to 2006 have also concluded that
bryozoans (ectoprocts) and entoprocts are not sister groups.[35] However, two well-known zoologists, Claus
Nielsen and Thomas Cavalier-Smith, maintain on anatomical and developmental grounds that bryozoans and
entoprocts are member of the same phylum, Bryozoa. A molecular phylogeny study in 2007 also supported
this old idea, while its conclusions about other phyla agreed with those of several other analyses.[35]
By 1891 bryozoans (ectoprocts) were grouped with phoronids in a super-phylum called
"Tentaculata". In the 1970s comparisons between phoronid larvae and the cyphonautes larva of some
gymnolaete bryozoans produced suggestions that the bryozoans, most of which are colonial, evolved from a
semi-colonial species of phoronid.[54] Brachiopods were also assigned to the "Tentaculata", which were
renamed Lophophorata as they all use a lophophore for filter feeding.[34] Although the majority of scientists
accept this,[34] Claus Nielsen thinks these similarities are superficial.[11] The Lophophorata are usually
defined as animals with a lophophore, a three-part coelom and a U-shaped gut.[54] In Nielsen's opinion,
phoronids' and brachiopods' lophophores are more like those of pterobranchs,[11] which are members of the
phylum Hemichordata.[55] Bryozoan's tentacles bear cells with multiple cilia, while the corresponding cells of
phoronids', brachiopods' and pterobranchs' lophophores have one cilium per cell; and bryozoan tentacles
have no hemal canal ("blood vessel"), which those of the other three phyla have.[11]
If the grouping of bryozoans with phoronids and brachiopods into Lophophorata is correct, the next
issue is whether the Lophophorata are protostomes, along with most invertebrate phyla, or deuterostomes,
along with chordates, hemichordates and echinoderms. The traditional view was that lophophorates were a
mix of protostome and deuterostome features. Research from the 1970s onwards suggested they were
deuterostomes, because of some features that were thought characteristic of deuterostomes: a three-part
coelom; radial rather than spiral cleavage in the development of the embryo;[34] and formation of the coelom
by enterocoely.[11] However the coelom of ectoproct larvae shows no sign of division into three sections,[54]
and that of adult ectoprocts is different from that of other coelomate phyla as it is built anew from epidermis
and mesoderm after metamorphosis has destroyed the larval coelom.[49]
Molecular phylogeny analyses from 1995 onwards, using a variety of biochemical evidence and
analytical techniques, placed the lophophorates as protostomes and closely related to annelids and molluscs
in a super-phylum called Lophotrochozoa.[34][56] "Total evidence" analyses, which used both morphological
features and a relatively small set of genes, came to various conclusions, mostly favoring a close relationship
between lophophorates and Lophotrochozoa.[56] A study in 2008, using a larger set of genes, concluded that
the lophophorates were closer to the Lophotrochozoa than to deuterostomes, but also that the lophophorates
were not monophyletic. Instead, it concluded that brachiopods and phoronids formed a monophyletic group,
but bryozoans (ectoprocts) were closest to entoprocts, supporting the original definition of "Bryozoa".[56]
[edit] Physiology
[edit] Feeding and excretion
Most species are filter feeders that sieve small particles, mainly phytoplankton (microscopic floating
plants), out of the water.[5] The freshwater species Plumatella emarginata feeds on diatoms, green algae,
cyanobacteria, non-photosynthetic bacteria, dinoflagellates, rotifers, protozoa, small nematodes, and
microscopic crustaceans.[57] While the currents that bryozoans generate to draw food towards the mouth are
well understood, the exact method of capture is still debated. All species also flick larger particles towards the
mouth with a tentacle, and a few capture zooplankton (planktonic animals) by using their tentacles as cages.
In addition the tentacles, whose surface area is increased by microvilli (small hairs and pleats), absorb
organic compounds dissolved in the water.[5] Unwanted particles may be flicked away by tentacles or shut
out by closing the mouth.[5] A study in 2008 showed that both encrusting and erect colonies fed more quickly
and grew faster in gentle than in strong currents.[58]
In some species the first part of the stomach forms a muscular gizzard lined with chitinous teeth that
crush armored prey such as diatoms. Wave-like peristaltic contractions move the food through the stomach
for digestion. The final section of the stomach is lined with cilia (minute hairs) that compress undigested
solids, which then pass through the intestine and out through the anus.[5]
There are no nephridia ("little kidneys") or other excretory organs in bryzoa,[15] and it is thought that
ammonia diffuses out through the body wall and lophophore.[5] More complex waste products are not
excreted but accumulate in the polypide, which degenerates after a few weeks. Some of the old polypide is
recycled, but much of it remains as a large mass of dying cells containing accumulated wastes, and this is
compressed into a "brown body". When the degeneration is complete, the cystid (outer part of the animal)
produces a new polypide, and the brown body remains in the coelom, or in the stomach of the new polypide
and is expelled next time the animal defecates.[5]
[edit] Respiration and circulation

There are no respiratory organs, heart or blood vessels. Instead zooids absorb oxygen and eliminate
carbon dioxide through the body wall and especially the lophophore.[5] The fluid in the coelom transports
gases and nutrients and its circulation is passive, except that some relatively large species use cilia to boost
its speed.[5] The different bryozoan groups use various methods to share nutrients and oxygen between
zooids: some have quite large gaps in the body walls, allowing the coelomic fluid to circulate freely; in others
the funiculi (internal "little ropes"[21]) of adjacent zooids connect via small pores in the body wall.[5][17]
[edit] Reproduction and life cycles
Encrusting cyclostome bryozoans (B), the one on the right showing swollen gonozooids; T =
thecideide brachiopod and S = sabellid worm tube; Jurassic of Poland.
Zooids of all phylactolaemate species are simultaneous hermaphrodites. Although those of many
marine species are protandric, in other words function first as males and then as females, their colonies
contain a combination of zooids that are in their male and female stages. In all species the ovaries develop
on the inside of the body wall, and the testes on the funiculus connecting the stomach to the body wall.[15]
Eggs and sperm are released into the coelom, and sperm exit into the water through pores in the tips of
some of the tentacles, and then are captured by the feeding currents of zooids that are producing eggs.[5]
Some species' eggs are fertilized externally after being released through a pore between two tentacles,
which in some cases is at the tip of a small projection called the "intertentacular organ" in the base of a pair
of tentacles. Others' are fertilized internally, in the intertentacular organ or in the coelom.[5] In ctenostomes
the mother provides a brood chamber for the fertilized eggs, and her polypide disintegrates, providing
nourishment to the embryo. Stenolaemates produce specialized zooids to serve as brood chambers, and
their eggs divide within this to produce up to 100 identical embryos.[15]
The cleavage of bryozoan eggs is biradial, in other words the early stages are bilaterally symmetrical.
It is unknown how the coleom forms, since the metamorphosis from larva to adult destroys all of the larva's
internal tissues. In many animals the blastopore, an opening in the surface of the early embryo, tunnels
through to form the gut. However, in bryozoans the blastopore closes, and a new opening develops to create
the mouth.[5]
Bryozoan larvae vary in form, but all have a band of cilia round the body which enables them to
swim, a tuft of cilia at the top, and an adhesive sac that everts and anchors them when they settle on a
surface.[5] Some gymnolaemate species produce cyphonautes larvae which have little yolk but a well-
developed mouth and gut, and live as plankton for a considerable time before settling. These larvae have
triangular shells of chitin, with one corner at the top and the base open, forming a hood round the downward-
facing mouth.[15] In 2006 it was reported that the cilia of cyphonautes larvae use the same range of
techniques as those of adults to capture food.[59] Species that brood their embryos form larvae that are
nourished by large yolks, have no gut and do not feed, and such larvae quickly settle on a surface.[5] In all
marine species the larvae produce cocoons in which they metamorphose completely after settling: the larva's
epidermis becomes the lining of the coelom, and the internal tissues are converted to a food reserve that
nourishes the developing zooid until it is ready to feed.[5] The larvae of phylactolaemates produce multiple
polypides, so that each new colony starts with several zooids.[5] In all species the founder zooids then grow
the new colonies by budding clones of themselves. In phylactolaemates, zooids die after producing several
clones, so that living zooids are found only round the edges of a colony.[5]
Phylactolaemates also reproduce asexually by a method that enables a colony's lineage to survive
the variable and uncertain conditions of freshwater environments.[15] Throughout summer and autumn they
produce disc-shaped statoblasts, masses of cells that function as "survival pods" rather like the gemmules of
sponges.[5] Statoblasts form on the funiculus connected to the parent's gut, which nourishes them.[15] As
they grow, statoblasts develop protective bivalve-like shells made of chitin. When they mature, some
statoblasts stick to the parent colony, some fall to the bottom ("sessoblasts"), some contain air spaces that
enable them to float ("floatoblasts"),[5] and some remain in the parent's cystid to re-build the colony if it dies.
[15] Statoblasts can remain dormant for considerable periods, and while dormant can survive harsh
conditions such as freezing and desiccation. They can be transported across long distances by animals,
floating vegetation, currents[5] and winds,[15] and even in the guts of larger animals.[60] When conditions
improve, the valves of the shell separate and the cells inside develop into a zooid that tries to form a new
colony. Plumatella emarginata produces both "sessoblasts", which enable the lineage to control a good
territory even if hard times decimate the parent colonies, and "floatoblasts", which spread to new sites. New
colonies of Plumatella repens produce mainly "sessoblasts" while mature ones switch to "floatoblasts".[57] A
study estimated that one group of colonies in a patch measuring 1 square metre (11 sq ft) produced
800,000 statoblasts.[5]
[edit] Ecology
[edit] Habitats and distribution
Most marine species live in tropical waters at depths less than 100 metres (330 ft). However, a few
have been found in deep-sea trenches,[61] especially around cold seeps, and others near the poles.[62][63]
The great majority are sessile. Encrusting forms are much the commonest of these in shallow seas, but erect
forms become more common as the depth increases.[62] A few marine species can move, and an Antarctic
species forms floating colonies.[62]
The phylactolaemates live in all types of freshwater environment – lakes and ponds, rivers and
streams, and estuaries[44] – and are among the most abundant sessile freshwater animals.[52] Some
ctenostomes are exclusively freshwater while others prefer brackish water but can survive in freshwater.[44]
Scientists' knowledge of freshwater bryozoan populations in many parts of the world is incomplete, even in
some parts of Europe. It was long thought that some freshwater species occurred worldwide, but since 2002
all of these have been split into more localized species.[44]
[edit] Interactions with non-human organisms
lacelike Membranipora membranacea

Marine species are common on coral reefs, but seldom a significant proportion of the total biomass.
In temperate waters, the skeletons of dead colonies form a significant component of shell gravels, and live
ones are abundant in these areas.[64] The marine lace-like bryozoan Membranipora membranacea produces
spines in response to predation by several species of nudibranchs ("sea slugs").[65] Other predators on
marine bryozoans include fish, sea urchins, pycnogonids, crustaceans, mites[66] and starfish.[67] In general
marine echinoderms and molluscs eat masses of zooids by gouging pieces of colonies, breaking their
mineralized "houses", while most arthropod predators on bryozoans eat individual zooids.[68]
In freshwater, bryozoans are among the most important filter feeders, along with sponges and
mussels.[69] Freshwater bryozoans are attacked by many predators, including snails, insects, and fish.[57]
In Thailand the introduced species Pomacea canaliculata (golden apple snail), which is generally a
destructive herbivore, has wiped out phylactolaemate populations wherever it has appeared. P. canaliculata
also preys on a common freshwater gymnolaemate, but with less devastating effect. Indigenous snails do not
feed on bryozoans.[70]
Several species of the hydroid family Zancleidae have symbiotic relationships with bryozoans, some
of which are beneficial to the hydroids while others are parasitic. Modifications appear in the shapes of some
these hydroids, for example smaller tentacles or encrustation of the roots by bryozoans.[71] The bryozoan
Alcyonidium nodosum protects the whelk Burnupena papyracea against predation by the powerful and
voracious rock lobster Jasus lalandii. While whelk shells encrusted by the bryozoans are stronger than those
without this reinforcement, chemical defenses produced by the bryozoans are probably the more significant
deterrent.[72]
Some phylactolaemate species are parasitized by a group of myxozoa that have also been found to
cause Proliferative Kidney Disease, which is often fatal in salmonid fish,[73] and has severely reduced wild
fish populations in Europe and North America.[44]
Membranipora membranacea, whose colonies feed and grow exceptionally fast in a wide range of
current speeds, was first noticed in the Gulf of Maine in 1987 and quickly became the most abundant
organism living on kelps.[58] This invasion reduced the kelp population by breaking their fronds,[5] so that its
place as the dominant "vegetation" in some areas was taken by another invader, the large alga Codium
fragile tomentosoides.[58] These changes reduced the area of habitat available for local fish and
invertebrates. M. membranacea has also invaded the northwest coast of the U.S.A.[5] A few freshwater
species have been also found thousands of kilometers from their native ranges. Some may have been
transported naturally as statoblasts. Others more probably were spread by humans, for example on imported
water plants or as stowaways on ships.[60]
[edit] Interaction with humans
Fish farms and hatcheries have lost stock to proliferative kidney disease, which is apparently caused
by one or more myxozoans that also parasitize bryozoans.[73]
Fishermen in the North Sea have had to find other work because of a form of eczema (a skin
disease) known as "Dogger Bank itch",[62] caused by contact with bryozoans that have stuck to nets and
lobster pots.[74]
Marine bryozoans are often responsible for biofouling on ships' hulls, on docks and marinas, and on
offshore structures. They are among the first colonizers of new or recently-cleaned structures.[64]
Freshwater species are occasional nuisances in water pipes, drinking water purification equipment, sewage
treatment facilities, and the cooling pipes of power stations.[44][75]
A group of chemicals called bryostatins can be extracted from the marine bryozoan Bugula neritina.
In 2001 pharmaceutical company GPC Biotech licensed Bryostatin 1 from Arizona State University for
commercial development as a treatment for cancer. GPC Biotech canceled development in 2003, saying that
Bryostatin 1 showed little effectiveness and some toxic side-effects.[76] In January 2008 a clinical trial was
submitted to the United States National Institutes of Health to measure the safety and effectiveness of
Bryostatin 1 in the treatment of Alzheimer's Disease. However, no participants had been recruited by the end
of December 2008, when the study was scheduled for completion.[77] About 1 tonne (160 st) of bryozoans
must processed to extract 1 gram (0.035 oz) of bryostatin. As a result, synthetic equivalents have been
developed that are simpler to produce and apparently at least as effective.[78]
[edit] See also
• International Bryozoology Association
[edit] References
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Wikimedia Commons has media related to: Bryozoa
• Index to Bryozoa Bryozoa Home Page, was at RMIT; now bryozoa.net

• Other Bryozoan WWW Resources
• International Bryozoology Association official website
• Bryozoan Introduction
• The Phylum Ectoprocta (Bryozoa)
• Phylum Bryozoa at Wikispecies
• Bryozoans in the Connecticut River
• Bryozoa Fact Sheet
[hide]v · d · eExtant phyla of kingdom Animalia by subkingdom
Parazoa Porifera (Calcarea, Demospongiae, Hexactinellida) · Placozoa (Trichoplax)
Mesozoa Orthonectida · Rhombozoa
Eumetazoa Radiata Ctenophora · Cnidaria (Anthozoa, Hydrozoa, Scyphozoa, Cubozoa, Stauroz
Bilateria Protostomia Cycloneuralia: Scalidophora (K

Loricifera, Priapulida) · Nematoida (Ne
Ecdysozoa Nematomorpha)
Panarthropoda: Onychophora
Arthropoda
Spiralia Platyzoa Platy

Gastrotricha
Gnat
Rotifera · Aca
Gnathostomu
Micrognathoz
Cycliophora
Troch
(Sipuncula, N
Mollusca, An
Lophotrochozoa
Lophophorata
Entoprocta, P
Brachiopoda
Hemichordata · Echinoderma
Ambulacraria
Xenoturbellida
Deuterostomia
Craniata (Vertebrata, Myxini
Chordata
Cephalochordata · Tunicata
Basal/disputed Acoelomorpha (Acoela, Nemertodermatida) · Chae
Retrieved from "http://en.wikipedia.org/wiki/Bryozoa"
Categories: Bryozoans
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Evolutionary history of living beings

Evolutionary history of life

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Research and history

Evolutionary biology fields

History of Earth and its life

[edit] Earliest evidence for life on Earth

[edit] Origins of life on Earth

[edit] Life "seeded" from elsewhere

[edit] Independent emergence on Earth

[edit] Replication first: RNA world

[edit] Membranes first: Lipid world

Cross-section through a liposome.

[edit] Environmental and evolutionary impact of microbial mats

[edit] Diversification of eukaryotes Archaeplastida (Land plants

[edit] Multicellular organisms and sexual reproduction

[edit] Evolution of sexual reproduction

[edit] Fossil evidence for multicellularity and sexual reproduction

Acanthodians were among the earliest vertebrates with jaws.[119]

[edit] Evolution of soil

Lichens growing on concrete

[edit] Plants and the Late Devonian wood crisis

[edit] Land vertebrates

Acanthostega changed views about the early evolution of tetrapods.[140]

[edit] Dinosaurs, birds and mammals

One possible family tree of flowering plants.[165]

Another possible family tree.[166]

[edit] Social insects

[edit] Mass extinctions

Apparent extinction intensity, i.e. the fraction of

[edit] The present

[edit] See also

Books are collections of articles that can be downloaded or ordered in print.

Evidence of common descent

Taxa evolution Birds · Butterflies · Cephalopods · Dinosaurs · Dolphins and whales ·

History of Charles Darwin · On the Origin of Species ·

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

[show]v · d · eElements of nature

Universe Space · Time · Matter · Energy

Environment Ecology · Ecosystem · Wilderness

Biology · Eukaryota (Plants/Flora, Animals/Fauna, Fungi, Protista) ·

[show]v · d · eEarth-related topics

Geography Continents · Geology of solar terrestrial planets · Timezones · Degree

Ecology Earth Day · Millennium Ecosystem Assessment

In fiction Hollow Earth · A Journey to the Center of the Earth

Teleology Creationism · Creation myth

Earth sciences portal · Solar System portal

[edit] External links

Categories: History by topic | Paleontology | Evolutionary biology

Evolutionary history of plants

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[edit] Changing life cycles

[edit] Water transport

[edit] Diversity in meristem architectures

[edit] Role of the KNOX-family genes

Complex leaves of C. hirsuta are a result of KNOX gene expression

[edit] Tree form

The fossil seed Trigonocarpus

The pollen bearing organs of the early "flower" Crossotheca

The inflorescences of the Bennettitales are strikingly similar to flowers

[edit] Concentrating carbon

[edit] Evolutionary record