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Overview 1
Introduction to evolution 1
Evolution 16
Evolution as theory and fact 47
Evolutionary history of life 53
Timeline of evolution 83

History 92
History of evolutionary thought 92
Lamarckism 114
Saltationism 121
Orthogenesis 122
On the Origin of Species 126
Darwinism 154
The Genetical Theory of Natural Selection 158
Neo-Darwinism 160
Modern evolutionary synthesis 162

Base concepts 172

Heredity 172
Fitness 176
Common descent 179
Evidence of common descent 183

Mechanisms 212
Adaptation 212
Genetic drift 223
Gene flow 233
Mutation 236
Natural selection 246
Speciation 260

Phylogenetics 268
Phylogenetics 268
Cladistics 274
Cladogram 284
Molecular phylogenetics 287

Fields 290
Evolutionary developmental biology 290
Molecular evolution 296
Human evolution 299
Evolutionary psychology 316

Controversy and social impacts 332

Creation–evolution controversy 332
Objections to evolution 359
Creationism 384
Intelligent design 406
Social effect of evolutionary theory 436

Article Sources and Contributors 444
Image Sources, Licenses and Contributors 454

Article Licenses
License 458


Introduction to evolution

Artist's depiction of a T. rex.

Natural selection does not lead to perfection; dramatic changes in the environment often lead to mass extinctions, as in the case of the dinosaurs
nearly 65 million years ago.

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somewhat outdated, it illustrates clearly the principles that more complex modern reconstructions can obscure.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
Introduction to evolution 2

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.

Darwin's idea: evolution by natural selection

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
• 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.
Introduction to evolution 3

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
Introduction to evolution 4

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]

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 necessarily 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
Introduction to evolution 5

attracted more attention than all others because it killed the dinosaurs.[25]

Modern 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.

Evidence for evolution

Scientific evidence for evolution comes from many aspects of
biology, and includes fossils, homologous structures, and
molecular similarities between species' DNA.

Fossil record
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 During the voyage of the Beagle, naturalist Charles
of Cuvier’s successful contributions to the understanding of the Darwin collected fossils in South America, and found
fragments of armor which he thought were like giant
fossil record was establishing extinction as a fact. In an attempt to
versions of the scales on the modern armadillos living
explain extinction, Cuvier proposed the idea of “revolutions” or nearby. On his return, the anatomist Richard Owen
catastrophism in which he speculated that geological catastrophes showed him that the fragments were from gigantic
had occurred throughout the Earth’s history, wiping out large extinct glyptodons, related to the armadillos. This was
one of the patterns of distribution that helped Darwin to
numbers of species.[28] Cuvier's theory of revolutions was later [11]
develop his theory.
replaced by uniformitarian theories, notably those of James Hutton
Introduction to evolution 6

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]

Comparative anatomy
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.

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]

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.
Introduction to evolution 7

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]

Convergent evolution

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
The bird and the bat wing are examples of convergent
pressures. Within both groups, changes that aid swimming have
evolution. been favored. Thus, over time, they developed similar appearances
(morphology), even though they are not closely related.[39]

A bat is a mammal and its forearm bones

have been adapted for flight.
Introduction to evolution 8

Molecular biology
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.
A section of DNA

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.
Introduction to evolution 9

Artificial selection
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
The results of artificial selection: a Chihuahua mix and
15,000 years ago.[45]
a Great Dane.

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.

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
There are numerous species of cichlids that with members of other species).[50] Mayr's definition has gained
demonstrate dramatic variations in morphology. 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
Introduction to evolution 10

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]

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
Introduction to evolution 11

organism or the gene.

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]

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.

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]

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.
Introduction to evolution 12

Although some groups raise objections to the theory of evolution, the evidence of observation and experiments 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]

See also
• Creation-evolution controversy
• Evidence of common descent
• Evolution as theory and fact
• Level of support for evolution
• Misconceptions about evolution

[1] Darwin, Charles (1859). On the Origin of Species (http:/ / darwin-online. org. uk/ content/ frameset?itemID=F373& viewtype=text&
pageseq=16) (1st ed.). London: John Murray. p. 1. .. Related earlier ideas were acknowledged in Darwin, Charles (1861). On the Origin of
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[2] Gould, Stephen J. (2002). The Structure of Evolutionary Theory. Harvard University Press. pp. 1433. ISBN 0674006135, 9780674006133.
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[16] DeVries A (2004). "The enigma of Darwin". Clio Med 19 (2): 136–55. PMID 6085987.
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increased Use and Disuse of Parts, as controlled by Natural Selection
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Introduction to evolution 13

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[32] (Diamond 1992, p. 16)
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[35] (Weichert & Presch 1975, p. 8)
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(pdf). TalkOrigins Archive. The TalkOrigins Foundation. . Retrieved 2008-01-27.
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(web resource). The Evidence for Evolution (http:/ / www. txtwriter. com/ backgrounders/ index. html). 'On Science' column in St. Louis Post
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[40] Kennedy, Donald; (Working group on teaching evolution) (1998). "Teaching about evolution and the nature of science" (http:/ / www. nap.
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[41] Lovgren, Stefan (2005-08-31). "Chimps, Humans 96 Percent the Same, Gene Study Finds" (http:/ / news. nationalgeographic. com/ news/
2005/ 08/ 0831_050831_chimp_genes. html). National Geographic News. National Geographic. . Retrieved 2007-12-23.
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[49] Wilner A. (2006). "Darwin's artificial selection as an experiment". Stud Hist Philos Biol Biomed Sci. 37 (1): 26–40.
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darwin. php?page=2). Smithsonian Magazine. Smithsonian Institution. . Retrieved 2007-08-31.
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• Carroll, SB; Grenier, J; Weatherbee, SD (2000). From DNA to Diversity: Molecular Genetics and the Evolution
of Animal Design (2nd ed.). Oxford: Blackwell Publishing. ISBN 1-4051-1950-0
• Darwin, Charles (1872). [[On the Origin of Species|The Origin of Species (
frameset?itemID=F391&viewtype=text&pageseq=1)]] (6th ed.). London: John Murray
• Dawkins, Richard (1976). The Selfish Gene (
Richard-Dawkins-The-Selfish-Gene-Original-Ed) (1st ed.). Oxford University Press. pp. 33. ISBN 0192860925
• Diamond, Jared (1992). The Third Chimpanzee: the evolution and future of the human animal. New York:
HarperCollins. ISBN 0060183071
• Gould (a), Stephen Jay (1981). The Panda's Thumb: More Reflections in Natural History. New York: W.W,
Norton & Company. ISBN 0393308197
• Gould (b), Stephen Jay (1995). Dinosaur in a Haystack. New York: Harmony Books. ISBN 0517703939
• Lyell, Charles (1830). Principles of geology ( New
York: Penguin Books. ISBN 014043528X
• 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
Introduction to evolution 15

Further reading
• Liam Neeson (narrator). (2001-11-20) (web resource). Evolution: a journey into where we're from and where
we're going ( [DVD]. South Burlington, VT: WGBH Boston
/ PBS television series Nova. ASIN: B00005RG6J. Retrieved 2008-01-24. - Age level: Grade 7+
• Horvitz, Leslie Alan (2002). The complete idiot's guide to evolution. Indianapolis: Alpha Books.
ISBN 0028642260.
• Charlesworth, Deborah; Charlesworth, Brian (2003). Evolution: a very short introduction. Oxford: Oxford
University Press. ISBN 0192802518.
• Sis, Peter (2003). The tree of life: a book depicting the life of Charles Darwin, naturalist, geologist & thinker.
New York: Farrar Straus Giroux. ISBN 0-374-45628-3.
• Thomson, Keith Stewart (2005). Fossils: a very short introduction. Oxford: Oxford University Press.
ISBN 0192805045.
• Greg Krukonis (2008). Evolution For Dummies (For Dummies (Math & Science)). For Dummies.
ISBN 0-470-11773-7.
• Darwin, Charles (2008). Quammen, David. ed. On the Origin of Species: The Illustrated Edition. Sterling.
ISBN 1402756399
• Pallan, Mark (2009). The Rough Guide to Evolution. Rough Guides. ISBN 1858289467
• Zimmer, Carl (2009). The Tangled Bank: An Introduction to Evolution. Roberts and Company Publishers.
ISBN 0981519474
• Ellis, R. John (2010). How Science Works: Evolution. Springer. ISBN 9048131820

External links
• Brain, Marshall. "How Stuff Works: Evolution Library" (
htm) (web resource). Retrieved 2008-01-24
• Carl Sagan. (2006-07-06) (Google video). Carl Sagan on evolution (
videoplay?docid=-522726029201501667&q=carl+sagan). [streaming video]. Google. Retrieved 2008-01-24.
• Carl Sagan. (2006-10-21) (Youtube video). Theory of Evolution Explained (
watch?v=E1Y5zMo74cY). [streaming video]. Youtube. Retrieved 2008-01-24.
• "Evolution Education Wiki: EvoWiki" ( (web resource). Retrieved 2008-01-24
• "The Big Picture on Evolution (PDF)" (
@msh_publishing_group/documents/web_document/wtd026042.pdf). The Big Picture Series (http://www. Wellcome Trust.
January 2007. Retrieved 2008-01-23
• "The Talk Origins Archive: Exploring the Creation/Evolution Controversy" ( (web
resource). Retrieved 2008-01-24
• (web resource) Understanding Evolution: your one-stop source for information on evolution (http://evolution. The University of California Museum of Paleontology,
Berkeley. Retrieved 2008-01-24
• "University of Utah Genetics Learning Center animated tour of the basics of genetics" (http://learn.genetics. (web resource). Retrieved 2008-01-24
• "Introduction To Evolution" ( (web resource). Retrieved
Evolution 16

Evolution (also known as biological, genetic or organic evolution) is the change in the inherited traits of a
population of organisms through successive generations.[1] This change results from interactions between processes
that introduce variation into a population, and other processes that remove it. As a result, variants with particular
traits become more, or less, common. A trait is a particular characteristic—anatomical, biochemical or
behavioural—that is the result of gene–environment interaction.
The main source of variation is mutation, which introduces genetic changes. These changes are heritable (can be
passed on through reproduction), and may give rise to alternative traits in organisms. Another source of variation is
genetic recombination, which shuffles the genes into new combinations which can result in organisms exhibiting
different traits. Under certain circumstances, variation can also be increased by the transfer of genes between
species,[2] [3] and by the extremely rare, but significant, wholesale incorporation of genomes through
endosymbiosis.[4] [5]
Two main processes cause variants to become more common or rarer in a population. One is natural selection,
through which traits that aid survival and reproduction become more common, while traits that hinder survival and
reproduction become rarer. Natural selection occurs because only a small proportion of individuals in each
generation will survive and reproduce, since resources are limited and organisms produce many more offspring than
their environment can support. Over many generations, heritable variation in traits is filtered by natural selection and
the beneficial changes are successively retained through differential survival and reproduction. This iterative process
adjusts traits so they become better suited to an organism's environment: these adjustments are called adaptations.[6]
However, not all change is adaptive. Another cause of evolution is genetic drift, which leads to random changes in
how common traits are in a population. Genetic drift is most important when traits do not strongly influence
survival—particularly so in small populations, in which chance plays a disproportionate role in the frequency of
traits passed on to the next generation.[7] [8] Genetic drift is important in the neutral theory of molecular evolution,
and plays a role in the molecular clocks that are used in phylogenetic studies.
A key process in evolution is speciation, in which a single ancestral species splits and diversifies into multiple new
species. There are several modes through which this occurs. Ultimately, all living (and extinct) species are descended
from a common ancestor via a long series of speciation events. These events stretch back in a diverse "tree of life"
which has grown over the 3.5 billion years during which life has existed on Earth.[9] [10] [11] [12] This 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.
Evolutionary biologists document the fact that evolution occurs, and also develop and test theories which explain its
causes. The study of evolutionary biology began in the mid-nineteenth century, when research into the fossil record
and the diversity of living organisms convinced most scientists that species changed over time.[13] The mechanism
driving these changes remained unclear until the theory of natural selection was independently proposed by Charles
Darwin and Alfred Wallace. In 1859, Darwin's seminal work On the Origin of Species brought the new theory of
evolution by natural selection to a wide audience,[14] leading to the overwhelming acceptance of evolution among
scientists.[15] [16] [17] [18] In the 1930s, Darwinian natural selection became understood in combination with
Mendelian inheritance, forming the modern evolutionary synthesis,[19] which connected the substrate of evolution
(inherited genetics) and the mechanism of evolution (natural selection). This powerful explanatory and predictive
theory has become the central organizing principle of modern biology, directing research and providing a unifying
explanation for the history and diversity of life on Earth.[16] [17] [20] Evolution is applied and studied in fields as
diverse as agriculture, anthropology, conservation biology, ecology, medicine, paleontology, philosophy, and
psychology along with other specific topics in the previous listed fields.
Evolution 17

History of evolutionary thought

The roots of naturalistic thinking on biology can be dated to at least the 6th
century BCE, with the Greek philosopher Anaximander.[21] However, the
growth of modern biology out of natural history is fairly recent. The word
evolution (from the Latin evolutio, meaning "to unroll like a scroll") appeared
in English in the 17th century. As biological knowledge grew 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.[22] The ideas
of the biologist Jean-Baptiste Lamarck about transmutation of species
influenced radicals, but were rejected by mainstream scientists. Charles
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
Around 1854 Charles Darwin began
theory, and both were presented to the Linnean Society of London in separate
writing out what became On the Origin
papers.[23] At the end of 1859, Darwin's publication of On the Origin of of Species.
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.[24] Like Lamarck, he still thought that parents passed on
adaptations acquired during their lifetimes,[25] a theory which was subsequently dubbed Lamarckism.[26] In the
1880s, August Weismann's experiments indicated that changes from use and disuse were not heritable, and
Lamarckism gradually fell from favour.[27] [28] 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.[29] When Mendel's work was rediscovered in 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
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.[31] 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.[19]
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.[19] 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
Evolution 18

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.[32] [33]

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.[34] Inherited traits are
controlled by genes and the complete set of genes within an organism's
genome is called its genotype.[35]

The complete set of observable traits that make up the structure and
behavior of an organism is called its phenotype. These traits come from the
interaction of its genotype with the environment.[36] 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.[37]

Heritable traits are passed from one generation to the next via DNA, a
molecule that encodes genetic information.[35] 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 DNA structure. Bases are in the center,
sequence of letters spelling out a sentence. Before a cell divides, the DNA surrounded by phosphate–sugar chains in a
double helix.
is copied, so that each of the resulting two cells will inherit the DNA

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.

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.[38] [39] The study of such complex traits is a major area of
current genetic research. Another unsolved question in genetics is whether or not epigenetics is important in
evolution. Epigenetics is when a trait is inherited without there being any change in gene sequences.[40]
Evolution 19

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.[39] 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.[41]
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 hybridization in plants.[42] Despite the constant
introduction of variation through these processes, most of the genome of a species is identical in all individuals of
that species.[43] 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.[44]

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.[45] [46] [47] 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 percent of these mutations having damaging effects,
and the remainder being either neutral or weakly beneficial.[48] Due to the damaging
effects that mutations can have on cells, organisms have evolved mechanisms such as
DNA repair to remove mutations.[45] 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.[49] Viruses that use RNA as their genetic material have rapid mutation
rates,[50] 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.[51]

Duplication of part of a Mutations can involve large sections of a chromosome becoming duplicated (usually by
chromosome genetic recombination), which can introduce extra copies of a gene into a genome.[52]
Extra copies of genes are a major source of the raw material needed for new genes to
evolve. This is important because most new genes evolve within gene families from pre-existing genes that share
common ancestors.[54] 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.[55] 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.[56] [57] Other types of mutation can
even create entirely new genes from previously noncoding DNA.[58] [59] 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.[60] [61] When new genes are assembled from shuffling pre-existing parts, domains act as modules
Evolution 20

with simple independent functions, which can be mixed together creating new combinations with new and complex
functions.[62] 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.[63]
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.[64] 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.[65]
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.[66] 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.[67] 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

Sex and recombination

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.[68] 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.[69] Sex usually increases genetic variation and may increase
the rate of evolution.[70] [71] However, asexuality is advantageous in some environments as it can evolve in
previously sexual animals.[72] Here, asexuality might allow the two sets of alleles in their genome to diverge and
gain different functions.[73]
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.[74] 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.[75]
When alleles cannot be separated by recombination – such as in mammalian Y chromosomes, which pass intact from
fathers to sons – harmful mutations accumulate.[76] [77] By breaking up allele combinations, sexual reproduction
allows the removal of harmful mutations and the retention of beneficial mutations.[78] 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.[78] The reasons for the evolution of sexual reproduction are therefore unclear and
this question is still an active area of research in evolutionary biology,[79] [80] that has prompted ideas such as the
Red Queen hypothesis.[81]
Evolution 21

Population genetics

White peppered moth

Black morph in peppered moth evolution

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.[82] 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 fertilization.[83] Such a population is said to be in Hardy-Weinberg equilibrium; it is not
Evolution 22

Gene flow
Gene flow is the exchange of genes between
populations, which are usually of the same species.[86]
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.
When they mature, male lions leave the pride where they were born
and take over a new pride to mate, causing gene flow between As barriers to reproduction between two diverging
prides. 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.[87]

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.[88] 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.[89] The importance of hybridization in creating new
species of animals is unclear, although cases have been seen in many types of animals,[90] with the gray tree frog
being a particularly well-studied example.[91]

Hybridization 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.[92] [93] 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.[94] Polyploids also have more genetic diversity, which allows them to avoid
inbreeding depression in small populations.[95]
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.[96] 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.[97] 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.[98] [99] 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.[100] Viruses can
also carry DNA between organisms, allowing transfer of genes even across biological domains.[101] Large-scale gene
transfer has also occurred between the ancestors of eukaryotic cells and prokaryotes, during the acquisition of
chloroplasts and mitochondria.[102]
Evolution 23

The two main mechanisms that produce evolution are natural selection and genetic drift. Natural selection is the
process which favors 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.[103] 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.[104] 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.[41]

Natural selection
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 Natural selection of a population for dark colouration.
competitors pass these advantageous traits
on, while traits that do not confer an advantage are not passed on to the next generation.[105]

The central concept of natural selection is the evolutionary fitness of an organism.[106] Fitness is measured by an
organism's ability to survive and reproduce, which determines the size of its genetic contribution to the next
generation.[106] 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.[107] 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.[106]
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".[108] 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).[109] [110]
Evolution 24

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.[111] 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.[105] [112]
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.[113] Traits that evolved through
sexual selection are particularly prominent in males of
A chart showing three types of selection. 1.Disruptive selection
some animal species, despite traits such as cumbersome
2.Stabilizing selection 3.Directional selection
antlers, mating calls or bright colours that attract
predators, decreasing the survival of individual
males.[114] This survival disadvantage is balanced by higher reproductive success in males that show these hard to
fake, sexually selected traits.[115]

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 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."[116] 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

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.[117] [118] None of these are mutually exclusive and
selection can act on multiple levels simultaneously.[119] An example of selection occurring below the level of the
individual organism are genes called transposons, which can replicate and spread throughout a genome.[120]
Selection at a level above the individual, such as group selection, may allow the evolution of co-operation, as
discussed below.[121]
Evolution 25

Genetic drift
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.[122] 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
The time for an allele to become fixed by genetic drift
depends on population size, with fixation occurring
more rapidly in smaller populations.[123] 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.[124]

An example 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.[125]
However, it is usually difficult to measure the relative importance of selection and drift,[126] so the comparative
importance of these two forces in driving evolutionary change is an area of current research.[127] 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.[128] Hence, in this model, most genetic changes in a population are the result of constant mutation
pressure and genetic drift.[129] This form of the neutral theory is now largely abandoned, since it does not seem to fit
the genetic variation seen in nature.[130] [131] 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.[105]

Evolution influences every aspect of the form and behavior of organisms. Most prominent are the specific behavioral
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.
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,
Evolution 26

such as adaptations, within a species or population.[132] In general, macroevolution is regarded as the outcome of
long periods of microevolution.[133] Thus, the distinction between micro- and macroevolution is not a fundamental
one – the difference is simply the time involved.[134] 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.[135] [136] [137]
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.[138] [139] 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.[140] For example, the overwhelming majority of species are microscopic
prokaryotes, which form about half the world's biomass despite their small size,[141] and constitute the vast majority
of Earth's biodiversity.[142] 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.[143] 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.[144] [145]

Adaptation is one of the basic phenomena of biology,[146] and is the process whereby an organism becomes better
suited to its habitat.[147] [148] 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.[149] 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
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.[151]
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.[152]
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.[153] Other striking examples are the bacteria Escherichia coli evolving the ability to use citric acid as a nutrient
in a long-term laboratory experiment,[154] Flavobacterium evolving a novel enzyme that allows these bacteria to
grow on the by-products of nylon manufacturing,[155] [156] and the soil bacterium Sphingobium evolving an entirely
new metabolic pathway that degrades the synthetic pesticide pentachlorophenol.[157] [158] 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).[159] [160]
Evolution 27

Adaptation occurs through the gradual

modification of existing structures.
Consequently, structures with similar
internal organization may have
different functions in related
organisms. This is the result of a single
ancestral structure being adapted to A baleen whale skeleton, a and b label flipper bones, which were adapted from front leg
function in different ways. The bones bones: while c indicates vestigial leg bones, suggesting an adaptation from land to
within bat wings, for example, are very sea.

similar to those in mice feet and

primate hands, due to the descent of all these structures from a common mammalian ancestor.[162] However, since all
living organisms are related to some extent,[163] 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.[164] [165]

During adaptation, some structures may lose their original function and become vestigial structures.[166] 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,[167] the non-functional remains of eyes in blind
cave-dwelling fish,[168] wings in flightless birds,[169] and the presence of hip bones in whales and snakes.[161]
Examples of vestigial structures in humans include wisdom teeth,[170] the coccyx,[166] the vermiform appendix,[166]
and other behavioral vestiges such as goose bumps,[171] and primitive reflexes.[172] [173] [174] [175]
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.[176] 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.[176] Within cells, molecular machines such as the bacterial flagella[177]
and protein sorting machinery[178] evolved by the recruitment of several pre-existing proteins that previously had
different functions.[132] 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.[179] [180]
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.[181] 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.[182]
An area of current investigation in evolutionary developmental biology is the developmental basis of adaptations and
exaptations.[183] This research addresses the origin and evolution of embryonic development and how modifications
of development and developmental processes produce novel features.[184] 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.[185] 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.[186] It is now becoming clear that most alterations in the form of organisms
are due to changes in a small set of conserved genes.[187]
Evolution 28

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.[188] 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
Common garter snake (Thamnophis sirtalis
produced high levels of toxin in the newt and correspondingly high sirtalis) which has evolved resistance to
levels of toxin resistance in the snake.[189] tetrodotoxin in its amphibian prey.

However, not all interactions between species involve conflict.[190] 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.[191] 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.[192]
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
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.[193] 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.[194] Other processes that may promote cooperation include group selection, where
cooperation provides benefits to a group of organisms.[195]
Evolution 29

Speciation is the process where a
species diverges into two or more
descendant species.[196] Evolutionary
biologists view species as statistical
phenomena and not categories or
types. 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. 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. Indeed, the exact
The four mechanisms of speciation. definition of the term "species" is still
controversial, particularly in
[198] [199]
prokaryotes, and this is called the species problem. 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
concerned.[199] Typically the actual focus on biological study is the population, an observable interacting group of
organisms, rather than a species, an observable similar group of individuals.

Speciation has been observed multiple times under both controlled laboratory conditions and in nature.[200] 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.[201] [202] As selection and
drift act independently on populations isolated from the rest of their species, separation may eventually produce
organisms that cannot interbreed.[203]
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.[204]
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.[196]
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.[205] 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.[206]
Evolution 30

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.[207] Generally, sympatric speciation in animals requires
the evolution of both genetic differences and non-random mating, to
allow reproductive isolation to evolve.[208]

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
Geographical isolation of finches on the
common in plants because plants often double their number of Galápagos Islands produced over a dozen new
chromosomes, to form polyploids.[209] This allows the chromosomes species.
from each parental species to form matching pairs during meiosis,
since each parent's chromosomes are represented by a pair already.[210] 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.[211] This happened about 20,000 years ago,[212] and the speciation process has been repeated in the
laboratory, which allows the study of the genetic mechanisms involved in this process.[213] 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.[93]

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.[214] 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.[215]

Extinction is the disappearance of an entire species.
Extinction is not an unusual event, as species regularly
appear through speciation, and disappear through
extinction.[216] Nearly all animal and plant species that have
lived on Earth are now extinct,[217] and extinction appears
to be the ultimate fate of all species.[218] These extinctions
have happened continuously throughout the history of life,
although the rate of extinction spikes in occasional mass
extinction events.[219] The Cretaceous–Tertiary extinction
event, during which the non-avian dinosaurs went extinct, is
Tyrannosaurus rex. Non-avian dinosaurs died out in the
Cretaceous–Tertiary extinction event at the end of the the most well-known, but the earlier Permian–Triassic
Cretaceous period. extinction event was even more severe, with approximately
96 percent of species driven to extinction.[219] 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
Evolution 31

up to 30 percent of species may be extinct by the mid 21st century.[220] Human activities are now the primary cause
of the ongoing extinction event;[221] global warming may further accelerate it in the future.[222]
The role of extinction in evolution is not very well understood and may depend on which type of extinction is
considered.[219] 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).[19] 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.[117] 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.[223]

Evolutionary history of life

Origin of life
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.[224] 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.[225] 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.[226] [227] Consequently, there is no scientific consensus on how life began, but proposals include
self-replicating molecules such as RNA,[228] and the assembly of simple cells.[229]

Common descent
All organisms on Earth are descended
from a common ancestor or ancestral
gene pool.[163] [230] Current species are
a stage in the process of evolution, with
their diversity the product of a long
series of speciation and extinction
events.[231] 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 The hominoids are descendants of a common ancestor.
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.[14] 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.[232] [233]

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.[234] 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
Evolution 32

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.[235] 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.[236] 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.[237]

Evolution of life
Despite the uncertainty on how life
began, it is generally accepted that
prokaryotes inhabited the Earth from
approximately 3–4 billion years
ago.[238] [239] No obvious changes in
morphology or cellular organization
occurred in these organisms over the
next few billion years.[240]

The eukaryotes were the next major

change in cell structure. These came
from ancient bacteria being engulfed
by the ancestors of eukaryotic cells, in
a cooperative association called
Evolutionary tree showing the divergence of modern species from their common ancestor endosymbiosis.[102] [241] The engulfed
in the center.Ciccarelli FD, Doerks T, von Mering C, Creevey CJ, Snel B, Bork P (2006).
bacteria and the host cell then
"Toward automatic reconstruction of a highly resolved tree of life". Science 311 (5765):
1283–87. doi:10.1126/science.1123061. PMID 16513982.  The three domains are
underwent co-evolution, with the
coloured, with bacteria blue, archaea green, and eukaryotes red. bacteria evolving into either
mitochondria or hydrogenosomes.[242]
An independent second engulfment of cyanobacterial-like organisms led to the formation of chloroplasts in algae and
plants.[243] It is unknown when the first eukaryotic cells appeared though they first emerged between 1.6 – 2.7
billion years ago.

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.[238] [244] The evolution of
multicellularity occurred in multiple independent events, in organisms as diverse as sponges, brown algae,
cyanobacteria, slime moulds and myxobacteria.[245]
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.[246] Various triggers for the Cambrian explosion have been proposed, including the accumulation of oxygen
in the atmosphere from photosynthesis.[247] About 500 million years ago, plants and fungi colonised the land, and
were soon followed by arthropods and other animals.[248] Insects were particularly successful and even today make
up the majority of animal species.[249] 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.[142]
Evolution 33

Evolutionary biology, and in particular the understanding of how organisms evolve through natural selection, is an
area of science with many practical applications.[250] 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.[251] 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
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.[253] 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.[254] This helped identify genes required for vision and pigmentation, such as
crystallins and the melanocortin 1 receptor.[255] 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
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.[257] Artificial evolution became a widely recognised optimization 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.[258]
Genetic algorithms in particular became popular through the writing of John Holland.[259] As academic interest
grew, dramatic increases in the power of computers allowed practical applications, including the automatic evolution
of computer programs.[260] 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.[261]

Social and cultural responses

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 centered 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.[19] However, evolution remains a contentious
concept for some theists.[263]

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.[132] [264]
As had been demonstrated by responses to the publication of Vestiges of
the Natural History of Creation in 1844, the most controversial aspect of As evolution became widely accepted in
evolutionary biology is the implication of human evolution that human mental the 1870s, caricatures of Charles Darwin
and moral faculties, which had been thought purely spiritual, are not distinctly with an ape or monkey body symbolised
Evolution 34

separated from those of other animals.[13] In some countries—notably the United States—these tensions between
science and religion have fueled the current creation-evolution controversy, a religious conflict focusing on politics
and public education.[266] While other scientific fields such as cosmology[267] and Earth science[268] 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.[269]
Another example somewhat 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.[270] 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.[271] [272] [273]

See also
• Current research in evolutionary biology

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[245] Kaiser D (2001). "Building a multicellular organism". Annu. Rev. Genet. 35: 103–23. doi:10.1146/annurev.genet.35.102401.090145.
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[246] Valentine JW, Jablonski D, Erwin DH (1 March 1999). "Fossils, molecules and embryos: new perspectives on the Cambrian explosion"
(http:/ / dev. biologists. org/ cgi/ reprint/ 126/ 5/ 851). Development 126 (5): 851–9. PMID 9927587. .
[247] Ohno S (1997). "The reason for as well as the consequence of the Cambrian explosion in animal evolution". J. Mol. Evol. 44 Suppl 1:
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*Valentine J, Jablonski D (2003). "Morphological and developmental macroevolution: a paleontological perspective" (http:/ / www. ijdb. ehu.
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[248] Waters ER (2003). "Molecular adaptation and the origin of land plants". Mol. Phylogenet. Evol. 29 (3): 456–63.
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Evolution 44

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PMID 18177707.
[255] Gross JB, Borowsky R, Tabin CJ (January 2009). "A novel role for Mc1r in the parallel evolution of depigmentation in independent
populations of the cavefish Astyanax mexicanus" (http:/ / www. pubmedcentral. nih. gov/ articlerender. fcgi?tool=pmcentrez&
artid=2603666). PLoS Genet. 5 (1): e1000326. doi:10.1371/journal.pgen.1000326. PMID 19119422. PMC 2603666.
[256] Yergeau DA, Cornell CN, Parker SK, Zhou Y, Detrich HW (July 2005). "bloodthirsty, an RBCC/TRIM gene required for erythropoiesis in
zebrafish". Dev. Biol. 283 (1): 97–112. doi:10.1016/j.ydbio.2005.04.006. PMID 15890331.
[257] Fraser AS (1958). "Monte Carlo analyses of genetic models". Nature 181 (4603): 208–9. doi:10.1038/181208a0. PMID 13504138.
[258] Rechenberg, Ingo (1973) (in German). Evolutionsstrategie – Optimierung technischer Systeme nach Prinzipien der biologischen Evolution
(PhD thesis). Fromman-Holzboog.
[259] Holland, John H. (1975). Adaptation in Natural and Artificial Systems. University of Michigan Press. ISBN 0262581116.
[260] Koza, John R. (1992). Genetic Programming. MIT Press. ISBN 0262111705.
[261] Jamshidi M (2003). "Tools for intelligent control: fuzzy controllers, neural networks and genetic algorithms". Philosophical transactions.
Series A, Mathematical, physical, and engineering sciences 361 (1809): 1781–808. doi:10.1098/rsta.2003.1225. PMID 12952685.
[262] Browne, Janet (2003). Charles Darwin: The Power of Place. London: Pimlico. pp. 376–379. ISBN 0-7126-6837-3.
[263] For an overview of the philosophical, religious, and cosmological controversies, see: Dennett, D (1995). Darwin's Dangerous Idea:
Evolution and the Meanings of Life. Simon & Schuster. ISBN 978-0684824710.
*For the scientific and social reception of evolution in the 19th and early 20th centuries, see: Johnston, Ian C.. "History of Science: Origins of
Evolutionary Theory" (http:/ / records. viu. ca/ ~johnstoi/ darwin/ sect3. htm). And Still We Evolve. Liberal Studies Department, Malaspina
University College. . Retrieved 2007-05-24.
*Bowler, PJ (2003). Evolution: The History of an Idea, Third Edition, Completely Revised and Expanded. University of California Press.
ISBN 978-0520236936.
*Zuckerkandl E (2006). "Intelligent design and biological complexity". Gene 385: 2–18. doi:10.1016/j.gene.2006.03.025. PMID 17011142.
[264] Ross, M.R. (2005). "Who Believes What? Clearing up Confusion over Intelligent Design and Young-Earth Creationism" (http:/ / www.
nagt. org/ files/ nagt/ jge/ abstracts/ Ross_v53n3p319. pdf) (PDF). Journal of Geoscience Education 53 (3): 319. . Retrieved 2008-04-28.
[265] Hameed, Salman (2008-12-12). "Science and Religion: Bracing for Islamic Creationism" (http:/ / helios. hampshire. edu/ ~sahCS/
Hameed-Science-Creationism. pdf). Science 322 (5908): 1637–1638. doi:10.1126/science.1163672. PMID 19074331. . Retrieved 2009.
[266] Spergel D. N.; Scott, EC; Okamoto, S (2006). "Science communication. Public acceptance of evolution". Science 313 (5788): 765–66.
doi:10.1126/science.1126746. PMID 16902112.
[267] Spergel, D. N.; Verde, L.; Peiris, H. V.; Komatsu, E.; Nolta, M. R.; Bennett, C. L.; Halpern, M.; Hinshaw, G. et al. (2003). "First-Year
Wilkinson Microwave Anisotropy Probe (WMAP) Observations: Determination of Cosmological Parameters". The Astrophysical Journal
Supplement Series 148: 175–94. doi:10.1086/377226.
[268] Wilde SA, Valley JW, Peck WH, Graham CM (2001). "Evidence from detrital zircons for the existence of continental crust and oceans on
the Earth 4.4 Gyr ago". Nature 409 (6817): 175–78. doi:10.1038/35051550. PMID 11196637.
[269] Understanding Creationism after Kitzmiller (http:/ / www. bioone. org/ doi/ full/ 10. 1641/ B570313) 2007
[270] On the history of eugenics and evolution, see Kevles, D (1998). In the Name of Eugenics: Genetics and the Uses of Human Heredity.
Harvard University Press. ISBN 978-0674445574.
[271] Darwin strongly disagreed with attempts by Herbert Spencer and others to extrapolate evolutionary ideas to all possible subjects; see
Midgley, M (2004). The Myths we Live By. Routledge. p. 62. ISBN 978-0415340779.
[272] 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.
[273] Gowaty, Patricia Adair (1997). Feminism and evolutionary biology: boundaries, intersections, and frontiers. London: Chapman & Hall.
ISBN 0-412-07361-7.
Evolution 45

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.
History of evolutionary thought
• Darwin, Charles (1859). On the Origin of Species (
frameset?itemID=F373&viewtype=text&pageseq=1) (1st ed.). London: John Murray. ISBN 0801413192.
• 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.
Evolution 46

External links
General information
• Evolution ( on In Our Time at the BBC. ( listen now (http://
• Everything you wanted to know about evolution by New Scientist (
• — How Evolution Works (
• National Academies Evolution Resources (
• Synthetic Theory Of Evolution: An Introduction to Modern Evolutionary Concepts and Theories (http://anthro.
• Understanding Evolution from University of California, Berkeley (
• Evolution of Evolution - 150 Years of Darwin's "On the Origin of Species" (
History of evolutionary thought
• 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
• The Making of the Fittest ( – lecture by Sean B. Carroll
Evolution as theory and fact 47

Evolution as theory and fact

The statement "evolution is both a theory and a fact" is often seen in biological literature.[1] [2] [3] [4] [5] [6] [7]
Evolution is a "theory" in the scientific sense of the term "theory"; it is an established scientific model that explains
observations and makes predictions through mechanisms such as natural selection.
When scientists say "evolution is a fact", they are using one of two meanings of the word "fact". One meaning is
empirical: evolution can be observed through changes in allele frequencies or traits of a population over successive
Another way "fact" is used is to refer to a certain kind of theory, one that has been so powerful and productive for
such a long time that it is universally accepted by scientists. When scientists say evolution is a fact in this sense, they
mean it is a fact that all living organisms have descended from a common ancestor (or ancestral gene pool) [8] even
though this cannot be directly observed. This implies more tangibly that it is a fact that humans share a common
ancestor with other primates.

Evolution, fact and theory

Evolution has been described as "fact and theory", "fact not theory", and "only a theory, not a fact". This illustrates a
confusion in terminology that hampers discussion.[9] [10] The meanings of the terms "evolution", "fact", and "theory"
are described below.

Evolution is usually defined simply as changes in trait or gene frequency in a population of organisms from one
generation to the next. However, "evolution" is often used to include the following additional claims:
1. Differences in trait composition between isolated populations over many generations may result in the origin of
new species.
2. All living organisms alive today have descended from a common ancestor (or ancestral gene pool).
According to Douglas Futuyma:
Biological evolution may be slight or substantial; it embraces everything from slight changes in the proportion
of different alleles within a population (such as those determining blood types) to the successive alterations
that led from the earliest proto-organism to snails, bees, giraffes, and dandelions.[11]
The term "evolution", especially when referred to as a "theory", is also used more broadly to incorporate processes
such as natural selection and genetic drift.

Fact is often used by scientists to refer to experimental or empirical data or objective verifiable observations.[12] [13]
[14] [15]
"Fact" is also used in a wider sense to mean any theory for which there is overwhelming evidence.
A fact is a hypothesis that is so firmly supported by evidence that we assume it is true, and act as if it were
true. —Douglas Futyuma[16]
Evolution is a fact in the sense that it is overwhelmingly validated by the evidence. Frequently, evolution is said to
be a fact in the same way as the Earth revolving around the Sun is a fact.[16] [17] The following quotation from H. J.
Muller, "One Hundred Years Without Darwin Are Enough" explains the point.
There is no sharp line between speculation, hypothesis, theory, principle, and fact, but only a difference along
a sliding scale, in the degree of probability of the idea. When we say a thing is a fact, then, we only mean that
its probability is an extremely high one: so high that we are not bothered by doubt about it and are ready to
act accordingly. Now in this use of the term fact, the only proper one, evolution is a fact.[3]
Evolution as theory and fact 48

The National Academy of Science (U.S.) makes a similar point:

Scientists most often use the word "fact" to describe an observation. But scientists can also use fact to mean
something that has been tested or observed so many times that there is no longer a compelling reason to keep
testing or looking for examples. The occurrence of evolution in this sense is fact. Scientists no longer question
whether descent with modification occurred because the evidence is so strong.[18]
Philosophers of science argue that we do not know anything with absolute certainty: even direct observations may be
"theory laden" and depend on assumptions about our senses and the measuring instruments used. In this sense all
facts are provisional.[1] [19]

A scientific theory is a well-supported body of interconnected statements that explains observations and can be used
to make testable predictions. Scientific theories describe the coherent framework into which observable data fit. The
"theory of evolution" is the framework that best explains observed changes of species over time and best predicts the
new observations that continue to be made in evolutionary biology and related sciences.
The scientific definition of the word "theory" is different from the colloquial sense of the word. Colloquially,
"theory" can mean a conjecture, an opinion, or a speculation that does not have to be based on facts or make testable
predictions. In science, the meaning of theory is more rigorous: a theory must be based on observed facts and make
testable predictions. In science, a current theory is a theory that has no equally acceptable or more acceptable
alternative theory.

Evolution compared with gravity

The application of the terms "fact" and "theory" to evolution is similar to their use in describing gravity.[20]
The most obvious fact of gravity is that objects in our everyday experience tend to fall downwards when not
otherwise prevented from doing so. People throughout history have wondered what causes this effect. Many
explanations have been proposed over the centuries. Aristotle, Galileo, Newton, and Einstein have all developed
useful models of gravity, each of which constitutes a theory of gravity. (Newton, for example, realized that the fact
of gravity can be extended to the tendency of any two masses to attract one another.) The word "gravity", therefore,
can be used to refer to the observed facts (i.e., that masses attract one another) and the theory used to explain the
facts (the reason why masses attract one another). In this way, gravity is both a theory and a fact.
In the study of biological species, the facts include the existence of many different species in existence today, some
very similar to each other and some very dissimilar, the remains of extinct species in the fossil record, and so forth.
In species that rapidly reproduce, for example fruit flies, the process of change from generation to generation — that
is, evolutionary change — has been observed in the laboratory.[21] The observation of fruit fly populations changing
over time is also an example of a fact. So evolution is a fact just as the observations of gravity are a fact.
There have been many attempts to explain these biological observations over the years. Lamarckism,
transmutationism and orthogenesis were all non-Darwinian theories that attempted to explain the observations of
species and fossils, as well as other evidence. However, the modern theory of evolution is the explanation for all
relevant observations regarding the development of life, based on a model that explains all the available data and
observations (and provides testable predictions). Thus, evolution is not only a fact but also a theory, just as gravity is
both a fact and a theory.
Evolution as theory and fact 49

Evolution as theory and fact in the literature

The confusion over the word evolution and the distinction between "fact" and "theory" is largely due to authors using
evolution to refer to three related yet distinct ideas: first, the changes that occur within species over generations;
second, the mechanism thought to drive change; and third, the concept of common descent. However, among
biologists there is a consensus that evolution is a fact:
• American zoologist and paleontologist George Simpson stated that "Darwin... finally and definitely established
evolution as a fact."[22]
• H. J. Muller wrote, "If you like, then, I will grant you that in an absolute sense evolution is not a fact, or rather,
that it is no more a fact than that you are hearing or reading these words."[3]
• Kenneth R. Miller writes, "evolution is as much a fact as anything we know in science."[23]
• Ernst Mayr observed, "The basic theory of evolution has been confirmed so completely that most modern
biologists consider evolution simply a fact. How else except by the word evolution can we designate the sequence
of faunas and floras in precisely dated geological strata? And evolutionary change is also simply a fact owing to
the changes in the content of gene pools from generation to generation."[7]

Evolution as fact and theory

Commonly "fact" is used to refer to the observable changes in organisms' traits over generations while the word
"theory" is reserved for the mechanisms that cause these changes:
• Paleontologist Stephen Jay Gould writes, "Evolution is a theory. It is also a fact. And facts and theories are
different things, not rungs in a hierarchy of increasing certainty. Facts are the world's data. Theories are
structures of ideas that explain and interpret facts. Facts do not go away when scientists debate rival theories to
explain them. Einstein's theory of gravitation replaced Newton's, but apples did not suspend themselves in
mid-air, pending the outcome. And humans evolved from ape-like ancestors whether they did so by Darwin's
proposed mechanism or by some other yet to be discovered."[2]
• Similarly, biologist Richard Lenski says, "Scientific understanding requires both facts and theories that can
explain those facts in a coherent manner. Evolution, in this context, is both a fact and a theory. It is an
incontrovertible fact that organisms have changed, or evolved, during the history of life on Earth. And biologists
have identified and investigated mechanisms that can explain the major patterns of change."[6]
• Biologist T. Ryan Gregory says, "biologists rarely make reference to 'the theory of evolution,' referring instead
simply to 'evolution' (i.e., the fact of descent with modification) or 'evolutionary theory' (i.e., the increasingly
sophisticated body of explanations for the fact of evolution). That evolution is a theory in the proper scientific
sense means that there is both a fact of evolution to be explained and a well-supported mechanistic framework to
account for it."[20]

Evolution as fact not theory

Other commentators, focusing on the changes in species over generations and in some cases common ancestry have
stressed that evolution is a fact to emphasize the weight of supporting evidence while denying it is helpful to use the
term "theory":
• R. C. Lewontin wrote, "It is time for students of the evolutionary process, especially those who have been
misquoted and used by the creationists, to state clearly that evolution is a fact, not theory."[24]
• Douglas Futuyma writes in his Evolutionary Biology book, "The statement that organisms have descended with
modifications from common ancestors—the historical reality of evolution—is not a theory. It is a fact, as fully as
the fact of the earth's revolution about the sun."[11]
• Richard Dawkins says, "One thing all real scientists agree upon is the fact of evolution itself. It is a fact that we
are cousins of gorillas, kangaroos, starfish, and bacteria. Evolution is as much a fact as the heat of the sun. It is
not a theory, and for pity's sake, let's stop confusing the philosophically naive by calling it so. Evolution is a
Evolution as theory and fact 50

• Neil Campbell wrote in his 1990 biology textbook, "Today, nearly all biologists acknowledge that evolution is a
fact. The term theory is no longer appropriate except when referring to the various models that attempt to explain
how life evolves... it is important to understand that the current questions about how life evolves in no way implies
any disagreement over the fact of evolution."[4]

Predictive power
A central tenet in science is that a scientific theory is supposed to have predictive power, and verification of
predictions are seen as an important and necessary support for the theory. The theory of evolution has provided such
predictions. Four examples are:
• Genetic information must be transmitted in a molecular way that will be almost exact but permit slight changes.
Since this prediction was made, biologists have discovered the existence of DNA, which has a mutation rate of
roughly 10−9 per nucleotide per cell division; this provides just such a mechanism.[26]
• Some DNA sequences are shared by very different organisms. It has been predicted by the theory of evolution
that the differences in such DNA sequences between two organisms should roughly resemble both the biological
difference between them according to their anatomy and the time that had passed since these two organisms have
separated in the course of evolution, as seen in fossil evidence. The rate of accumulating such changes should be
low for some sequences, namely those that code for critical RNA or proteins, and high for others that code for less
critical RNA or proteins; but for every specific sequence, the rate of change should be roughly constant over time.
These results have been experimentally confirmed. Two examples are DNA sequences coding for rRNA, which is
highly conserved, and DNA sequences coding for fibrinopeptides (amino acid chains that are discarded during the
formation of fibrin), which are highly non-conserved.[26]
• Prior to 2004, paleontologists had found fossils of amphibians with necks, ears, and four legs, in rock no older
than 365 million years old. In rocks more than 385 million years old they could only find fish, without these
amphibian characteristics. Evolutionary theory predicted that since amphibians evolved from fish, an intermediate
form should be found in rock dated between 365 and 385 million years ago. Such an intermediate form should
have many fish-like characteristics, conserved from 385 million years ago or more, but also have many amphibian
characteristics as well. In 2004, an expedition to islands in the Canadian arctic searching specifically for this
fossil form in rocks that were 375 million years old discovered fossils of Tiktaalik.[27]
• Evolutionary theory predicts that novel inventions can arise, while creationists predict that new "information"
cannot arise, and that the Second Law of Thermodynamics only allows for "information" to be lost.[28] In an
ongoing experiment, Richard Lenski observed that E. coli evolved the ability to metabolize citrate, which
constitutes a novel invention, and an increase in the information of the DNA of the E. coli.[29]

Related concepts and terminology

• Speculative or conjectural explanations are called hypotheses. Well-tested explanations are called theories.
• "Fact" does not mean "absolute certainty". In the words of Stephen J. Gould: In science, "fact" can only mean
"confirmed to such a degree that it would be perverse to withhold provisional assent."[2]
• "Proof" of a theory does not exist in natural sciences. Proof only exists in formal sciences, such as mathematics.
Experimental observation of the predictions made by a hypothesis or theory is called validation.
• A scientific law is a concept related to a scientific theory. Very well-established "theories" that rely on a simple
principle are often called scientific "laws". For example, it is common to encounter reference to "the law of
gravity", "the law of natural selection", or the "laws of evolution."
Evolution as theory and fact 51

See also
• Epistemology
• Evidence of common descent
• Evolution is just a theory, not a fact (in objections to evolution)
• Misconceptions about evolution (in List of misconceptions)
• Theory vs. Fact (in Creation-evolution controversy)

[1] Moran, Laurence (1993-01-22). "Evolution is a Fact and a Theory" (http:/ / www. talkorigins. org/ faqs/ evolution-fact. html). .
Retrieved 2007-10-18.
[2] Gould, Stephen Jay (1981-05-01). "Evolution as Fact and Theory" (http:/ / www. stephenjaygould. org/ library/ gould_fact-and-theory. html).
Discover 2 (5): 34–37. . Reprinted in:
• Vetter, Herbert F. (ed.) (1982). Speak Out Against The New Right. Beacon Press. ISBN 0807004863.
• Gould, Stephen Jay (1994-04-01). Hen's Teeth and Horse's Toes. New York: Norton. ISBN 0393017168.
[3] Muller, H. J. (1959). "One hundred years without Darwin are enough" (http:/ / www. skepticfiles. org/ evolut/ 100pcnts. htm). School Science
and Mathematics 59: 304–305. doi:10.1111/j.1949-8594.1959.tb08235.x. . Reprinted in:
• Zetterberg, Peter (ed.) (1983-05-01). Evolution Versus Creationism: The Public Education Controversy. Phoenix AZ: ORYX Press.
ISBN 0897740610.
[4] Campbell, Neil A.; Reece, Jane B. (2002-02-05). Biology 6th ed.. Benjamin Cummings. p. 1175. ISBN 0805366245.
[5] Dobzhansky, Theodosius (1973-03-01). "Nothing in biology makes sense except in the light of evolution" (http:/ / people. delphiforums. com/
lordorman/ light. htm). American Biology Teacher 35. .

Reprinted in:
• Zetterberg, Peter (ed.) (1983-05-01). Evolution Versus Creationism: The Public Education Controversy. Phoenix AZ: ORYX Press.
ISBN 0897740610.
[6] Lenski, Richard E. (2000). "Evolution: Fact and Theory" (http:/ / www. actionbioscience. org/ evolution/ lenski. html). American Institute of
Biological Sciences. . Retrieved 2007-10-18.
[7] Mayr, Ernst (1988). Toward a New Philosophy of Biology: Observations of an Evolutionist. Cambridge: Harvard University Press.
ISBN 0-674-89666-1.
[8] Cavalier-Smith T (2006). "Cell evolution and Earth history: stasis and revolution" (http:/ / www. pubmedcentral. nih. gov/ articlerender.
fcgi?tool=pmcentrez& artid=1578732). Philos Trans R Soc Lond B Biol Sci 361 (1470): 969–1006. doi:10.1098/rstb.2006.1842.
PMID 16754610. PMC 1578732.
[9] Is "Evolution" a "Theory" or "Fact" or Is This Just a Trivial Game of Semantics? (http:/ / www. discovery. org/ a/ 6401) by Casey Luskin
[10] Committee for Skeptical Inquiry — Evolution & Creationism: Terminology in Conflict (http:/ / www. csicop. org/ specialarticles/ show/
evolution_amp_creationism_terminology_in_conflict/ ) by Richard Joltes
[11] Futuyma, Douglas J. (1997). , Evolutionary Biology, 3rd ed.. Sinauer Associates. p. 751. ISBN 0878931899.
[12] Wordnet entry for phrase "scientific fact" (http:/ / wordnet. princeton. edu/ perl/ webwn?s=scientific fact)
[13] United States National Park Service Glossary (http:/ / www2. nature. nps. gov/ views/ System/ Glossary. htm#F)
[14] Webster's New Millennium Dictionary of English, Preview Edition (v 0.9.6), Copyright © 2003–2006 Lexico Publishing Group, LLC (http:/
/ dictionary. reference. com/ search?q=scientific+ fact& r=66)
[15] Webster's Encyclopedic Unabridged Dictionary of the English Language (1996) gives a third meaning of the word "fact" as (3) A truth
known by actual experience or observation; something known to be true: 'Scientists gather facts about plant growth.'
[16] Hypotheses, Facts, and the Nature of Science, —Douglas Futyuma (http:/ / www. stephenjaygould. org/ library/ futuyma_theory. html)
[17] Guardian article by Richard Dawkins, Jerry Coyne (http:/ / www. guardian. co. uk/ science/ 2005/ sep/ 01/ schools. research)
[18] Science and Creationism: A View from the National Academy of Sciences, Second Edition (1999), National Academy of Sciences (NAS),
National Academy Press, Washington DC, 2006. (http:/ / books. nap. edu/ openbook. php?record_id=6024& page=28)
[19] Wilkins, JS (1997). "Evolution and Philosophy:Is Evolution Science, and What Does 'Science' Mean?" (http:/ / www. talkorigins. org/ faqs/
evolphil/ falsify. html). The TalkOrigins Archive (http:/ / www. talkorigins. org). . Retrieved 2009-08-17.
[20] Gregory, T. Ryan (2007). "Evolution as Fact, Theory, and Path". Evolution: Education and Outreach 1 (1): 46–52.
[21] Dobzhansky T, Pavlovsky O (1971). "Experimentally created incipient species of Drosophila". Nature 230 (5292): 289–92.
doi:10.1038/230289a0. PMID 5549403.
[22] Robinson, B.A. (2005-08-30). "Is the theory of evolution merely a "theory"?" (http:/ / www. religioustolerance. org/ ev_stat. htm). .
Retrieved 2007-10-18.
[23] "Miller, Kenneth S. (2007). Finding Darwin's God: A Scientist's Search for Common Ground Between God and Evolution (P.S.). New York,
N.Y: Harper Perennial. ISBN 0061233501.
Evolution as theory and fact 52

[24] Lewontin, R. C. (1981). "Evolution/Creation Debate: a time for truth". Bioscience 31: 559. Reprinted in:
• Zetterberg, Peter, ed. (1983-05-01). Evolution Versus Creationism: the public education controversy. Phoenix AZ: Oryx Press.
ISBN 0897740610.
[25] Natural History article : The Illusion of Design (http:/ / www. naturalhistorymag. com/ features/ 101500/ the-illusion-of-design) by Richard
[26] Bruce Alberts; Alexander Johnson; Julian Lewis; Martin Raff; Keith Roberts; Peter Walter (March, 2002). Molecular Biology of the Cell
(4th ed.). Routledge. ISBN 0-8153-3218-1.
[27] "Shubin, Neil. (2008). Your Inner Fish. Pantheon. ISBN 9780375424472.
[28] (http:/ / www. talkorigins. org/ faqs/ thermo/ probability. html)
[29] NS:bacteria make major evolutionary shift in the lab (http:/ / www. newscientist. com/ channel/ life/
dn14094-bacteria-make-major-evolutionary-shift-in-the-lab. htm)

• J.P. Franck, et al., "Evolution of a satellite DNA family in tilapia." Annual Meeting Canadian Federation of
Biological Societies. Halifax, (1990).
• M. Losseau-Hoebeke, "The biology of four haplochromine species of Lake Kivu (Zaire) with evolutionary
implications." Thesis, Dept. Ichthyology, Rhodes University, Grahamstown, (1992).

External links
• Not Just a Theory ( Discredits the assertion that evolution is "just a theory",
with an explanation of the meaning of the word 'theory' in a scientific context.
• Talk Origins ( Response to the claim that no examples
of speciation have been observed.
• Glenn Branch; Louise S. Mead (2008-06-06). "“Theory” in Theory and Practice" (
content/fr258627q2x3t378/fulltext.pdf) (pdf). Evo Edu Outreach (2008) 1:287–289. Springer Science +
Business Media. Retrieved 2008-07-21.
Evolutionary history of life 53

Evolutionary history of life

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 3500 [1] 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.[2]
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.[3] The evolution of oxygenic
photosynthesis, around 3500 [1] million years ago, eventually led to the oxygenation of the atmosphere, beginning
around 2400 [4] million years ago.[5] The earliest evidence of eukaryotes (complex cells with organelles), dates from
1850.0 [6] million years ago,[7] [8] and while they may have been present earlier, their diversification accelerated
when they started using oxygen in their metabolism. Later, around 1700 [9] million years ago, multicellular
organisms began to appear, with differentiated cells performing specialised functions.[10]
The earliest land plants date back to around 450.0 [11] million years ago,[12] though evidence suggests that algal scum
formed on the land as early as 1200 [13] million years ago. Land plants were so successful that they are thought to
have contributed to the late Devonian extinction event.[14] Invertebrate animals appear during the Vendian period,[15]
while vertebrates originated about 525 [16] million years ago during the Cambrian explosion.[17]
During the Permian period, synapsids, including the ancestors of mammals, dominated the land,[18] but the
Permian–Triassic extinction event 251.0 [19] million years ago came close to wiping out all complex life.[20] During
the recovery from this catastrophe, archosaurs became the most abundant land vertebrates, displacing therapsids in
the mid-Triassic.[21] One archosaur group, the dinosaurs, dominated the Jurassic and Cretaceous periods,[22] with the
ancestors of mammals surviving only as small insectivores.[23] After the Cretaceous–Tertiary extinction event 65 [24]
million years ago killed off the non-avian dinosaurs[25] mammals increased rapidly in size and diversity.[26] Such
mass extinctions may have accelerated evolution by providing opportunities for new groups of organisms to
Fossil evidence indicates that flowering plants appeared and rapidly diversified in the Early Cretaceous, between 130
million years ago and 90 [29] 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 [30] million years ago. Although early members of this lineage had
chimp-sized brains, there are signs of a steady increase in brain size after about 3 [31] million years ago.

Earliest history of Earth

History of Earth and its life
Evolutionary history of life 54

Millions of years
The oldest meteorite fragments found on Earth are about 4540 [32] million years old, this, coupled primarily with the
dating of ancient lead deposits, has put the estimated age of Earth at around that time.[33] About 40 million years
later a planetoid struck the Earth, throwing into orbit the material that formed the Moon.[34]
Evolutionary history of life 55

Until recently the oldest rocks found on Earth were about 3800 [35] million years old,[33] 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".[36] However analysis of zircons formed 4400 to 4000 [37] 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.[38]
Evidence from the Moon indicates that from 4000 to 3800 [39] 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.[36] [40] While there is no direct evidence of
conditions on Earth 4000 to 3800 [39] million years ago, there is no reason to think that the Earth was not also
affected by this late heavy bombardment.[41] 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.[42]

Earliest evidence for life on Earth

The earliest identified organisms were minute and relatively featureless, 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 3000 [43] million years ago.[44] Other finds in
rocks dated to about 3500 [1] million years ago have been interpreted as bacteria,[45] with geochemical evidence also
seeming to show the presence of life 3800 [35] million years ago.[46] 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.[47] [48] 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. Currently, the oldest unchallenged evidence for life is geochemical
signatures from rocks deposited 3400 [49] million years ago,[44] [50] although these statements have not been
thoroughly examined by critics.

Origins of life on Earth

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.[51] [52]
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.[53] The lack of
fossil or geochemical evidence for Evolutionary tree showing the divergence of modern species from their common ancestor
earlier organisms has left plenty of in the center.Ciccarelli, F.D., Doerks, T., von Mering, C., Creevey, C.J., et al (2006).
scope for hypotheses, which fall into "Toward automatic reconstruction of a highly resolved tree of life". Science 311 (5765):
1283–7. doi:10.1126/science.1123061. PMID 16513982.  The three domains are colored,
two main groups: 1) that life arose
with bacteria blue, archaea green, and eukaryotes red.
spontaneously on Earth or 2) that it

was "seeded" from elsewhere in the universe.

Evolutionary history of life 56

Life "seeded" from elsewhere

The idea that life on Earth was "seeded" from elsewhere in the universe dates back at least to the fifth century
BCE.[54] In the twentieth century it was proposed by the physical chemist Svante Arrhenius,[55] by the astronomers
Fred Hoyle and Chandra Wickramasinghe,[56] and by molecular biologist Francis Crick and chemist Leslie Orgel.[57]
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;[58] by alien
visitors, possibly as a result of accidental contamination by micro-organisms that they brought with them;[57] and
from outside the Solar system but by natural means.[55] [58] 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.[58] Scientists are divided over the likelihood
of life arising independently on Mars,[59] or on other planets in our galaxy.[58]

Independent emergence on Earth

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.[60] Organisms based on alternative biochemistry may however be possible on
other planets.[61]
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.[62] Research on abiogenesis still has a long way to go, since theoretical and
empirical approaches are only beginning to make contact with each other.[63] [64]

Replication first: RNA world

The replicator in virtually all known life is deoxyribonucleic acid. DNA's structure and replication systems are far more complex
than those of the original replicator.

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.[53] 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.[65] 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.[66]
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.[66] [67] [68] Ribozymes remain as the main
components of ribosomes, modern cells' "protein factories".[69]
Although short self-replicating RNA molecules have been artificially produced in laboratories,[70] doubts have been
raised about where natural non-biological synthesis of RNA is possible.[71] 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.[72] [73]
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 cell
Evolutionary history of life 57

components to appear and until then the proto-cells would be confined to the pores.[74]

Metabolism first: Iron-sulfur world

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.[75]

Membranes first: Lipid world

= water-attracting heads of lipid molecules

= water-repellent tails
Cross-section through a liposome.

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.[76] Experiments that 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
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.[77]

The clay theory

RNA is complex and there are doubts about whether it can be produced non-biologically in the wild.[71] 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.[78] Although this idea has not become the scientific consensus, it still has active
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.[80]
A similar hypothesis presents self-replicating iron-rich clays as the progenitors of nucleotides, lipids and amino
Evolutionary history of life 58

Environmental and evolutionary impact of microbial mats

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.[82] 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.[83]

Stromatolites are stubby pillars built as microbes in mats slowly

migrate upwards to avoid being smothered by sediment deposited
on them by water.[82] There has been vigorous debate about the Modern stromatolites in Shark Bay, Western Australia.
validity of alleged fossils from before 3000 [43] million years
ago,[84] with critics arguing that so-called stromatolites could have been formed by non-biological processes.[47] In
2006 another find of stromatolites was reported from the same part of Australia as previous ones, in rocks dated to
3500 [1] million years ago.[85]

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.[83] 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.[86] From this point onwards life itself produced significantly more of the
resources it needed than did geochemical processes.[87] Oxygen is toxic to organisms that are not adapted to it, but
greatly increases the metabolic efficiency of oxygen-adapted organisms.[88] [89] Oxygen became a significant
component of Earth's atmosphere about 2400 [4] million years ago.[90] Although eukaryotes may have been present
much earlier,[91] [92] the oxygenation of the atmosphere was a prerequisite for the evolution of the most complex
eukaryotic cells, from which all multicellular organisms are built.[93] 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.[3]
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.[3]
Evolutionary history of life 59

Diversification of eukaryotes


Archaeplastida (Land plants, green algae, red algae, and glaucophytes)

Bikonta Chromalveolata





Metazoa (Animals)
Opisthokonta Choanozoa

Eumycota (Fungi)

One possible family tree of eukaryotes[94] [95]

Eukaryotes may have been present long before the oxygenation of the atmosphere,[91] but most modern eukaryotes
require oxygen, which their mitochondria use to fuel the production of ATP, the internal energy supply of all known
cells.[93] In the 1970s it was proposed and, after much debate, widely accepted that eukaryotes emerged as a result of
a sequence of endosymbioses between "procaryotes". For example: a predatory micro-organism invaded a large
procaryote, probably an archaean, but the attack was neutralized, and the attacker 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 eliminated unproductive duplication of genetic functions by
re-arranging their genomes, a process which sometimes involved transfer of genes between them.[96] [97] [98] Another
hypothesis proposes that mitochondria were originally sulfur- or hydrogen-metabolising endosymbionts, and became
oxygen-consumers later.[99] On the other hand mitochondria might have been part of eukaryotes' original
There is a debate about when eukaryotes first appeared: the presence of steranes in Australian shales may indicate
that eukaryotes were present 2700 [101] million years ago;[92] however an analysis in 2008 concluded that these
chemicals infiltrated the rocks less than 2200 [102] million years ago and prove nothing about the origins of
eukaryotes.[103] Fossils of the alga Grypania have been reported in 1850.0 [6] million-year-old rocks (originally
dated to 2100 [104] million years ago but later revised[8] ), and indicates that eukaryotes with organelles had already
evolved.[105] A diverse collection of fossil algae were found in rocks dated between 1500 [106] million years ago and
1400 [107] million years ago.[108] The earliest known fossils of fungi date from 1430 [109] million years ago.[110]
Evolutionary history of life 60

Multicellular organisms 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.[112] Multicellularity evolved independently in
organisms as diverse as sponges and other animals, fungi, plants, A slime mold solves a maze. The mold (yellow)
brown algae, cyanobacteria, slime moulds and myxobacteria.[8] explored and filled the maze (left). When the
[113] researchers placed sugar (red) at two separate points,
For the sake of brevity this article focuses on the organisms
the mold concentrated most of its mass there and left
that show the greatest specialization of cells and variety of cell
only the most efficient connection between the two
types, although this approach to the evolution of complexity could [111]
points (right).
be regarded as "rather anthropocentric".[114]

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;[115] the ability to create an internal environment that gives
protection against the external one;[114] and even the opportunity for a group of cells to behave "intelligently" by
sharing information.[111] These features would also have provided opportunities for other organisms to diversify, by
creating more varied environments than flat microbial mats could.[115]

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.[115]
The available evidence indicates that eukaryotes evolved much earlier but remained inconspicuous until a rapid
diversification around 1000 [116] 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.[115]

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.[117] 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.[118]
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.[119]
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.[117] 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.[120] How sexual reproduction evolved and survived is an unsolved
Evolutionary history of life 61

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.[117]
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.[117]
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
Horodyskia apparently re-arranged itself into
reproduction.[121] Other combinations of hypotheses that are fewer but larger main masses as the sediment
inadequate on their own are also being examined.[117] grew deeper round its base.

The following hypotheses attempt to explain how and why sex

• It may have enabled organisms to repair genetic damage.[122] The most primitive form of sex may have been one
organism repairing damaged DNA by replicating an undamaged strand from a similar organism.[123]
• Sexual reproduction may have originated from selfish parasitic genetic elements propagating themselves by
transfer to new hosts.[124]
• It may have evolved from cannibalism, where some of the victim's DNA was incorporated into the cannibal
• Sexual reproduction may have evolved from ancient haloarchaea through a combination of jumping genes, and
swapping plasmids.[125]
• Or it may have evolved as a form of vaccination in which infected hosts exchanged weakened symbiotic copies of
parasitic DNA as protection against more virulent versions. The meiosis stage of sexual reproduction may then
have evolved as a way of removing the symbiotes.[126]
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.[118] However conjugation is not a means of reproduction and
is not limited to members of the same species, and there are cases where bacteria transfer DNA to plants and
animals.[119] Nevertheless it may be an example of the "selfish genetic element" hypothesis, as it transfers DNA by
means of such a "selfish gene", the F-plasmid.[123]
Evolutionary history of life 62

Fossil evidence for multicellularity and sexual reproduction

[8] [127]
Horodyskia may have been an early metazoan, or a colonial foraminiferan

The Francevillian Group Fossil, dated to 2100 [104] million years ago, is the earliest known fossil organism that is
clearly multicellular.[] They may have had differentiated cells.[128] Another early multicellular fossil,
Qingshania,[129] dated to 1700 [9] million years ago, appears to consist of virtually identical cells. The red alga called
Bangiomorpha, dated at 1200 [13] million years ago, is the earliest known organism which certainly has
differentiated, specialized cells, and is also the oldest known sexually-reproducing organism.[115] The 1430 [109]
million-year-old fossils interpreted as fungi appear to have been multicellular with differentiated cells.[110] The
"string of beads" organism Horodyskia, found in rocks dated from 1500 [106] million years ago to 900.0 [130] million
years ago, may have been an early metazoan;[8] however it has also been interpreted as a colonial foraminiferan.[127]

Emergence of animals

Deuterostomes (chordates, hemichordates, echinoderms)

Ecdysozoa (arthropods, nematodes, tardigrades, etc.)

Bilaterians Protostomes
Lophotrochozoa (molluscs, annelids, brachiopods, etc.)


Cnidaria (jellyfish, sea anemones, hydras)

Ctenophora (comb jellies)


Porifera (sponges): Calcarea

Porifera: Hexactinellida & Demospongiae



A family tree of the animals.[131]

Animals are multicellular eukaryotes,[132] and are distinguished from plants, algae, and fungi by lacking cell
walls.[133] All animals are motile,[134] if only at certain life stages. All animals except sponges have bodies
differentiated into separate tissues, including muscles, which move parts of the animal by contracting, and nerve
tissue, which transmits and processes signals.[135]
The earliest widely-accepted animal fossils are rather modern-looking cnidarians (the group that includes jellyfish,
sea anemones and hydras), possibly from around 580 [136] million years ago, although fossils from the Doushantuo
Formation can only be dated approximately. Their presence implies that the cnidarian and bilaterian lineages had
already diverged.[137]
Evolutionary history of life 63

The Ediacara biota, which flourished for the last 40 million years before the start of the Cambrian,[138] 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.[139] Others, however, been
interpreted as early molluscs (Kimberella[140] [141] ), echinoderms (Arkarua[142] ), and arthropods (Spriggina,[143]
Parvancorina[144] ). 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.[145]
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.[146]
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.[146] 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
Opabinia made the largest single
that the Early Cambrian was a uniquely experimental period of animal
contribution to modern interest in the evolution.[148] Later discoveries of similar animals and the development of
Cambrian explosion. new theoretical approaches led to the conclusion that many of the "weird
wonders" were evolutionary "aunts" or "cousins" of modern groups[149] – 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.[150] 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.[151]

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.[146] Other deuterostome groups are
soft-bodied, and most of the significant Cambrian deuterostome Acanthodians were among the earliest vertebrates with
fossils come from the Chengjiang fauna, a lagerstätte in jaws

China.[153] The Chengjiang fossils Haikouichthys and

Myllokunmingia appear to be true vertebrates,[154] and Haikouichthys had distinct vertebrae, which may have been
slightly mineralized.[155] Vertebrates with jaws, such as the Acanthodians, first appeared in the Late Ordovician.[156]

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.[157] [158] Although the earliest good evidence of
land plants and animals dates back to the Ordovician period (488 to 444 [159] million years ago), modern land
ecosystems only appeared in the late Devonian, about 385 to 359 [160] million years ago.[161]
Evolutionary history of life 64

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.[161]
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.[161] Lichens, which are symbiotic combinations of a fungus
(almost always an ascomycete) and one or more photosynthesizers Lichens growing on concrete
(green algae or cyanobacteria),[162] are also important colonizers
of lifeless environments,[161] and their ability to break down rocks contributes to soil formation in situations where
plants cannot survive.[162] The earliest known ascomycete fossils date from 423 to 419 [163] million years ago in the

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.[161] Burrows have been
found in Ordovician sediments, and are attributed to annelids ("worms") or arthropods.[161] [164]

Plants and the Late Devonian wood crisis

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.[157] [165]

Spores of land plants, possibly rather like liverworts, have been

found in Mid Ordovician rocks dated to about 476 [166] million
years ago. In Mid Silurian rocks 430 [167] 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.[165] Reconstruction of Cooksonia, a vascular plant from the
By the Late Devonian 370 [168] 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.[169] In fact they caused a "Late
Devonian wood crisis",[170] because:
• They removed more carbon dioxide from the atmosphere, reducing the greenhouse effect and thus causing an ice
age in the Carboniferous period.[171] In later ecosystems the carbon
Evolutionary history of life 65

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.[172]
• 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
Fossilized trees from the Mid-Devonian Gilboa fossil
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.[158]
Some trace fossils from the Cambrian-Ordovician boundary about 490.0 [173] million years ago are interpreted as the
tracks of large amphibious arthropods on coastal sand dunes, and may have been made by euthycarcinoids,[174]
which are thought to be evolutionary "aunts" of myriapods.[175] Other trace fossils from the Late Ordovician a little
over 445 [176] 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 [177] million years
ago, including signs that some arthropods ate plants.[178] 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.[179]
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.[158]
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.[180] About 99% of modern insect species fly or are descendants of flying species.[181]

Land vertebrates

Acanthostega changed views about the early evolution

of tetrapods
Evolutionary history of life 66

Osteolepiformes ("fish")





"Fish" Tulerpeton

Early amphibians



Family tree of tetrapods[183]

Tetrapods, vertebrates with four limbs, evolved from other rhipidistians over a relatively short timespan during the
Late Devonian, between 370 [168] million years ago and 360 [184] million years ago.[185] 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 [186] 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.[182]
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.[182]
Later discoveries revealed earlier transitional forms between Acanthostega and completely fish-like animals.[187]
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
Evolutionary history of life 67

accepted as early relatives of the amniotes, whose water-proof skins and eggs enable them to live and breed far from

Dinosaurs, birds and mammals

Early synapsids (extinct)

Extinct pelycosaurs

Extinct therapsids
Therapsids Extinct mammaliformes

Anapsids; whether turtles belong here is debated

Captorhinidae and Protorothyrididae

Araeoscelidia (extinct)

Squamata (lizards and snakes)


Extinct archosaurs


Pterosaurs (extinct)


Archosaurs Theropods theropods



Ornithischians (extinct)
Evolutionary history of life 68

Possible family tree of dinosaurs, birds and mammals[189] [190]

Amniotes, whose eggs can survive in dry environments, probably evolved in the Late Carboniferous period, between
330 [191] million years ago and 314 [192] million years ago. The earliest fossils of the two surviving amniote groups,
synapsids and sauropsids, date from around 313 [193] million years ago.[189] [190] The synapsid pelycosaurs and their
descendants the therapsids are the most common land vertebrates in the best-known Permian fossil beds, between
229.0 [194] million years ago and 251.0 [19] 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.[195]
The Permian-Triassic extinction wiped out almost all land vertebrates,[196] as well as the great majority of other
life.[197] During the slow recovery from this catastrophe, estimated to be 30M years,[198] 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,[199] 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 [200] million years ago
and 65 [24] million years ago.[201]
During the Late Jurassic, birds evolved from small, predatory theropod dinosaurs.[202] The first birds inherited teeth
and long, bony tails from their dinosaur ancestors,[202] but some developed horny, toothless beaks by the very Late
Jurassic[203] and short pygostyle tails by the Early Cretaceous.[204]
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.[205] By 195 [206] million years ago in the Early Jurassic there
were animals that were very nearly mammals.[207] Unfortunately there is a gap in the fossil record throughout the
Mid Jurassic.[208] However fossil teeth discovered in Madagascar indicate that true mammals existed at least 167
million years ago.[210] After dominating land vertebrate niches for about 150 million years, the dinosaurs
perished 65 [24] million years ago in the Cretaceous–Tertiary extinction along with many other groups of
organisms.[211] 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,[212] [213] with bats taking to the air within
13 million years,[214] and cetaceans to the sea within 15 million years.[215]
Evolutionary history of life 69

Flowering plants

(flowering plants)


Welwitschia Cycads
(gymnosperm) (gymnosperm)
Gymnosperms Bennettitales


Angiosperms Gnetales
(flowering plants) (gymnosperm)


One possible family tree of flowering


Another possible family tree.

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 [28] million years ago and 90 [29] million years ago,[216] [217] and that their rise
was associated with that of pollinating insects.[217] Among modern flowering plants Magnolias are thought to be
close to the common ancestor of the group.[216] However paleontologists have not succeeded in identifying the
earliest stages in the evolution of flowering plants.[216] [217]

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.[218]
Evolutionary history of life 70

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.[219] 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,[218] which are not haplodiploid.[220]

The earliest fossils of insects have been found in Early Devonian

rocks from about 400 [221] million years ago, which preserve only
a few varieties of flightless insect. The Mazon Creek lagerstätten
These termite mounds have survived a bush fire.
from the Late Carboniferous, about 300 [222] 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.[223]

Modern humans evolved from a lineage of upright-walking apes that has been traced back over 6 [30] million years
ago to Sahelanthropus.[224] The first known stone tools were made about 2.5 [225] million years ago, apparently by
Australopithecus garhi, and were found near animal bones that bear scratches made by these tools.[226] 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.[227] 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.[228] 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.[229]
Evolutionary history of life 71

Mass extinctions

Late D
Millions of years ago
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)

Life on earth has suffered occasional mass extinctions at least since 542 [230] 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.[231] [232]
The fossil record appears to show that the gaps between mass extinctions are becoming longer and the average and
background rates of extinction are decreasing. Both of these phenomena could be explained in one or more
• 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.[234] [235]
• 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.[233]
Evolutionary history of life 72

All genera
"Well-defined" genera
Trend line
"Big Five" mass extinctions
Other mass extinctions
Million years ago
Thousands of genera
Phanerozoic biodiversity as shown by the fossil record

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"[236]
shows a different trend: a fairly swift rise from 542 to 400 [237] 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 [239] million years ago to the present.[236]

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
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 resistant to
antibiotics.[241] [242] There is even evidence that humans are still evolving, and possibly at an accelerating rate over
the last 40,000 years.[243]
Evolutionary history of life 73

See also
• Evolution
• Evolutionary history of plants
• Timeline of evolution
• History of evolutionary thought

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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.

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 (
• — How Evolution Works (
• Synthetic Theory Of Evolution: An Introduction to Modern Evolutionary Concepts and Theories (http://anthro.
History of evolutionary thought
• The Complete Work of Charles Darwin Online (
• Understanding Evolution: History, Theory, Evidence, and Implications (
Timeline of evolution 83

Timeline of 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.

Basic timeline
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.

Detailed timeline
Ma, ("megaannum") means "million years ago". ka means "thousand years ago" and ya means "years ago"

Hadean Eon
3800 Ma and earlier.
Timeline of evolution 84

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. The gravitational pull of the new Moon
stabilises the Earth's fluctuating axis of rotation and sets up the conditions in which life formed.

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 The earliest life appears, possibly derived from self-reproducing RNA molecules.[5] [6] The replication of these organisms
and 3500 Ma 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
[7] [8] [9]
environment conducive to their replication: proto-cells.

3900 Ma Late Heavy Bombardment: peak rate of impact events upon the inner planets by meteors. 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
[10] [11]
vents below the Earth's surface; or life might have been transported to Earth by a meteor.

Somewhere Cells resembling prokaryotes appear.[12] These first organisms are chemoautotrophs: they use carbon dioxide as a carbon source
between 3900 - and oxidize inorganic materials to extract energy. Later, prokaryotes evolve glycolysis, a set of chemical reactions that free the
2500 Ma energy of organic molecules such as glucose and store it in the chemical bonds of ATP. Glycolysis (and ATP) continue to be
[13] [14]
used in almost all organisms, unchanged, to this day.

Archean Eon
3800 Ma – 2500 Ma

Date Event

3500 Lifetime of the last universal ancestor;[15] [16] the split between bacteria and archaea occurs.[17]
Ma [18]
Bacteria develop primitive forms of photosynthesis which at first do not produce oxygen. These organisms generate ATP by exploiting
a proton gradient, a mechanism still used in virtually all organisms.

3000 Photosynthesizing cyanobacteria evolve; they use water as a reducing agent, thereby producing oxygen as waste product.[19] More recent
Ma 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 1000 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 Timeframe of cyanobacteria evolution suggested by more recent research.


Proterozoic Eon
2500 Ma – 542 Ma
Timeline of evolution 85

Date Event

By 1850 Eukaryotic cells appear.[20] [21] Eukaryotes contain membrane-bound organelles with diverse functions, probably derived from
Ma prokaryotes engulfing each other via phagocytosis. (See Endosymbiosis)

By 1200 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 A global glaciation may have occurred.[23] [24] Opinion is divided on whether it increased or decreased biodiversity or the rate of
Ma evolution.[25] [26] [27]

580–542 The Ediacaran biota represent the first large, complex multicellular organisms - although their affinities remain a subject of debate.[28]

580–500 Most modern phyla of animals begin to appear in the fossil record during the Cambrian explosion.[29] [30]

580–540 The accumulation of atmospheric oxygen allows the formation of an ozone layer.[31] This blocks ultraviolet radiation, permitting the
Ma colonisation of the land.[31]

560 Ma Earliest fungi

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.

Paleozoic Era
542 Ma – 251.0 Ma

Date Event

535 Major diversification of living things in the oceans: chordates, arthropods (e.g. trilobites, crustaceans), echinoderms, mollusks,
Ma brachiopods, foraminifers and radiolarians, etc.

530 The first known footprints on land date to 530 Ma, indicating that early animal explorations may have predated the development of
Ma terrestrial plants.[32]

525 Earliest graptolites.


510 First cephalopods (Nautiloids) and chitons.


505 Fossilization of the Burgess Shale.


485 First vertebrates with true bones (jawless fishes).


450 Land arthropod burrows (millipedes) appear, along with the first complete conodonts and echinoids.

440 First agnathan fishes: Heterostraci, Galeaspida, and Pituriaspida.


434 The first primitive plants move onto land,[33] having evolved from green algae living along the edges of lakes.[34] They are accompanied
Ma by fungi, which may have aided the colonisation of land through symbiosis.
Timeline of evolution 86

420 Earliest ray-finned fishes, trigonotarbid arachnids, and land scorpions.


410 First signs of teeth in fish. Earliest nautiid nautiloids, lycophytes, and trimerophytes.

395 First lichens, stoneworts. Earliest harvestman, mites, hexapods (springtails), and ammonoids.

363 By the start of the Carboniferous Period, the Earth begins to be recognisable. Insects roamed the land and would soon take to the skies;
Ma 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 First crabs and ferns. Land flora dominated by seed ferns.

350 First large sharks, ratfishes, and hagfish.


340 Diversification of amphibians.


330 First amniote vertebrates (Paleothyris).


305 Earliest diapsid reptiles (e.g. Petrolacosaurus).


280 Earliest beetles, seed plants and conifers diversify while lepidodendrids and sphenopsids decrease. Terrestrial temnospondyl amphibians
Ma and pelycosaurs (e.g. Dimetrodon) diversify in species.

251.4 The Permian-Triassic extinction event eliminates over 90-95% of marine species. Terrestrial organisms were not as seriously affected as
Ma the marine biota. This "clearing of the slate" may have led to an ensuing diversification, but life on land took 30M years to completely

Mesozoic Era

Date Event

From The Mesozoic Marine Revolution begins: increasingly well-adapted and diverse predators pressurise sessile marine groups; the "balance
251.4 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

Timeline of evolution 87

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., 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.

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.
Timeline of evolution 88

Cenozoic Era
65.5 Ma – present

Date Event

65.5 The Cretaceous–Tertiary extinction event eradicates about half of all animal species, including mosasaurs, pterosaurs, plesiosaurs,
Ma ammonites, belemnites, rudist and inoceramid bivalves, most planktic foraminifers, and all of the dinosaurs excluding their descendants
the birds

From Rapid dominance of conifers and ginkgos in high latitudes, along with mammals becoming the dominant species. First psammobiid
65 Ma 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

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

Timeline of evolution 89

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 The last members of a dwarf race of Woolly Mammoths vanish from Wrangel Island near Alaska

383 ya The last wild Aurochs die out

37 ya The Thylacine goes extinct in a Tasmanian zoo, the last member of the family Thylacinidae

See also
• Evolutionary history of life
• Evolutionary history of plants
• Extinction events
• Geologic time scale
• History of Earth
• Natural history
• Sociocultural evolution
• Timeline of human evolution
• Timeline of plant evolution

Further reading
• The Ancestor's Tale by Richard Dawkins, for a list of ancestors common to humans and other living species

[1] Planetary Science Institute page (http:/ / www. psi. edu/ projects/ moon/ moon. html) 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
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[3] "However, once the Earth cooled sufficiently, sometime in the first 700 million years of its existence, clouds began to form in the atmosphere,
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[4] * The 'PAH World' (http:/ / nai. nasa. gov/ nai2005/ abstracts/ 616 - S. N. Platts PAH_World. doc. pdf)
[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" (http:/ / www. funpecrp. com. br/ gmr/ year2003/ vol4-2/ gmr0070_full_text. htm). Genetic and Molecular Research 2
(4): 366–375. PMID 15011140. . Retrieved 2008-08-30.(also available as PDF (http:/ / www. funpecrp. com. br/ gmr/ year2003/ vol4-2/ / pdf/
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[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.
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[9] Forterre, P., Benachenhou-Lahfa, N., Confalonieri, F., Duguet, M., Elie, C. and Labedan, B. (1992). "The nature of the last universal ancestor
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[10] Steenhuysen, Julie (May 21, 2009). "Study turns back clock on origins of life on Earth" (http:/ / www. reuters. com/ article/ scienceNews/
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[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 (http:/ / news-service. stanford. edu/ news/ 1998/ december2/
marsunder122. html) (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? (http:/ / www. sciam. com/ askexpert_question.
cfm?articleID=000C32DD-60E1-1C72-9EB7809EC588F2D7& catID=3& topicID=3)" Scientific American, October 21, 1999.
[13] Romano AH, Conway T. (1996) Evolution of carbohydrate metabolic pathways. Res Microbiol. 147(6-7):448-55 PMID 9084754
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doi:10.1146/ PMID 6250450.
[15] Doolittle, W. Ford (February, 2000). Uprooting the tree of life (http:/ / web. archive. org/ web/ 20060907081933/ http:/ / shiva. msu.
montana. edu/ courses/ mb437_537_2004_fall/ docs/ uprooting. pdf). 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?" (http:/ / www. pubmedcentral. nih. gov/ articlerender.
fcgi?tool=pmcentrez& artid=2606769). Philos. Trans. R. Soc. Lond., B, Biol. Sci. 363 (1504): 2731–43. doi:10.1098/rstb.2008.0041.
PMID 18468984. PMC 2606769.
[20] Knoll, Andrew H.; Javaux, E.J, Hewitt, D. and Cohen, P. (2006). "Eukaryotic organisms in Proterozoic oceans" (http:/ / www.
pubmedcentral. nih. gov/ articlerender. fcgi?tool=pmcentrez& artid=1578724). Philosophical Transactions of the Royal Society of London,
Part B 361 (1470): 1023–38. doi:10.1098/rstb.2006.1843. PMID 16754612. PMC 1578724.
[21] Fedonkin, M. A. (March 2003). "The origin of the Metazoa in the light of the Proterozoic fossil record" (http:/ / www. vend. paleo. ru/ pub/
Fedonkin_2003. pdf) (PDF). Paleontological Research 7 (1): 9–41. doi:10.2517/prpsj.7.9. . Retrieved 2008-09-02.
[22] Nicholas J. Butterfield, "Bangiomorpha pubescens n. gen., n. sp.: implications for the evolution of sex, multicellularity, and the
Mesoproterozoic/Neoproterozoic radiation of eukaryotes" (http:/ / paleobiol. geoscienceworld. org/ cgi/ content/ abstract/ 26/ 3/ 386)
[23] Hoffman, P.F.; Kaufman, A.J., Halverson, G.P., Schrag, D.P. (1998-08-28). "A Neoproterozoic Snowball Earth" (http:/ / www. sciencemag.
org/ cgi/ content/ full/ 281/ 5381/ 1342?ijkey=48d78da67bab492803c333f50c0dd84fbbef109c). Science 281 (5381): 1342.
doi:10.1126/science.281.5381.1342. PMID 9721097. . Retrieved 2007-05-04. Full online article (pdf 260 Kb) (http:/ / www. snowballearth.
org/ pdf/ Hoffman_Science1998. pdf)
[24] Kirschvink, J.L. (1992). "Late Proterozoic low-latitude global glaciation: The snowball Earth" (http:/ / www. gps. caltech. edu/ ~jkirschvink/
pdfs/ firstsnowball. pdf). In Schopf, JW, and Klein, C. (PDF). The Proterozoic Biosphere: A Multidisciplinary Study. Cambridge University
Press, Cambridge. pp. 51–52. .
[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" (http:/ / www. pnas. org/ cgi/ content/ abstract/ 100/ 8/ 4399). Proceedings of
the National Academy of Sciences 100 (8): 4399–4404. doi:10.1073/pnas.0730560100. PMID 12682298. PMC 153566. . Retrieved
[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" (http:/ / geol. queensu. ca/ people/ narbonne/ recent_pubs1.
html). Department of Geological Sciences and Geological Engineering, Queen's University. . Retrieved 2007-03-10.
[29] The Cambrian Period (http:/ / www. ucmp. berkeley. edu/ cambrian/ camb. html)
Timeline of evolution 91

[30] The Cambrian Explosion – Timing (http:/ / palaeo. gly. bris. ac. uk/ Palaeofiles/ Cambrian/ timing/ timing. html)
[31] Formation of the Ozone Layer (http:/ / jcbmac. chem. brown. edu/ myl/ ct7/ ozone/ ozone_formation. html)
[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 (http:/ / www. innovations-report. com/ html/ reports/ earth_sciences/ report-9641. html)
[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 (http:/ / www. clas. ufl. edu/ users/ pciesiel/ gly3150/ plant. html)
[34] "The land plants evolved from the algae, more specifically green algae, as suggested by certain common biochemical traits" The first land
plants (http:/ / scitec. uwichill. edu. bb/ bcs/ bl14apl/ conq. htm)
[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 (http:/ / www. elasmo-research. org/ education/ evolution/ evol_s_predator. htm)
[36] Sahney, S. and Benton, M.J. (2008). "Recovery from the most profound mass extinction of all time" (http:/ / journals. royalsociety. org/
content/ qq5un1810k7605h5/ fulltext. pdf) (PDF). Proceedings of the Royal Society: Biological 275 (1636): 759. 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 (http:/ / www. mcb.
uct. ac. za/ tutorial/ virorig. html) (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.
[39] The Oldest Homo Sapiens: (http:/ / www. sciencedaily. com/ releases/ 2005/ 02/ 050223122209. htm) - 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" (http:/ / www. stratigraphy. org/ cheu. pdf). International Commission on Stratigraphy. . Retrieved

External links
• Berkeley Evolution (
• Tree of Life Web Project ( - explore complete phylogenetic tree
• 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


History of evolutionary thought

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 Tree of Life as depicted by Ernst Haeckel in The Evolution
of Man (1879) illustrates the 19th-century view that evolution
The debate over Darwin's work led to the rapid acceptance was a progressive process leading towards man.
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
History of evolutionary thought 93

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.


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 (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 Plato (left) and Aristotle (right), a
Christian thought.[5] detail of The School of Athens

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]
History of evolutionary thought 94

Ideas on changing biologic species were expressed by ancient Chinese thinkers such as Zhuangzi (Chuang Tzu), a
Taoist philosopher who lived around the 4th century BC. 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] Humans, nature and the heavens are seen 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]

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]

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]

Middle Ages

Islamic philosophy and the struggle for existence

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]
History of evolutionary thought 95

Christian philosophy and the great chain of being

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
Drawing of the great chain of being from also perfect. There were no empty links in the chain, and no link was
Rhetorica Christiana (1579) by Diego Valades
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]

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]
History of evolutionary thought 96

Renaissance and Enlightenment

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 Pierre Belon compared the skeletons of birds and
during reproduction and accumulating over the course of many humans in his Book of Birds (1555).

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]
History of evolutionary thought 97

Early 19th century

Paleontology and geology

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

Diagram of the geologic timescale from an 1861 first few decades of the 19th century. By the 1840s, the outlines of the
book by Richard Owen showing the appearance geologic timescale were becoming clear, and in 1841 John Phillips
of major animal types
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]
History of evolutionary thought 98

Transmutation of species
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
Diagram from Vestiges of the Natural History of
influenced Grant was the anatomist Étienne Geoffroy Saint-Hilaire, Creation (1844) by Robert Chambers shows a
whose ideas on the unity of various animal body plans and the model of development where fish (F), reptiles
homology of certain anatomical structures would be widely influential (R), and birds (B) represent branches from a path
leading to mammals (M).
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
History of evolutionary thought 99

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]
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
This 1847 diagram by Richard Owen shows his Vestiges of the Natural History of Creation.[44] [45] Although the
conceptual archetype for all vertebrates.
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]

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:
History of evolutionary thought 100

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]

Natural selection
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 Darwin's first sketch of an evolutionary tree from his First Notebook
for 20 years. However he did share them with certain on Transmutation of Species (1837)

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]
History of evolutionary thought 101

1859–1930s: Darwin and his legacy

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

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
Diagram by O.C. Marsh of the evolution of horse feet
and teeth over time as reproduced in T.H Huxley's cornerstone of the program of the X Club to reform and
1876 book Professor Huxley in America 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]
History of evolutionary thought 102

Application to humans
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,
This illustration was the frontispiece of Thomas Henry Huxley's
although geology had shown that the Earth and life book Evidence as to Man's Place in Nature (1863).
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
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]
History of evolutionary thought 103

Alternatives to natural selection

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
This photo from Henry Fairfield Osborn's 1918 book
following.[69] [70]
Origin and Evolution of Life shows models depicting
In the late 19th century, the term neo-Lamarckism came to be
the evolution of Titanothere horns over time, which
associated with the position of naturalists who viewed the
Osborn claimed was an example of an orthogenic trend
in evolution. 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]
History of evolutionary thought 104

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 Diagram from T.H. Morgan's 1919 book The Physical
Basis of Heredity, showing the sex-linked inheritance
Mendelian genetics on the basis that discrete units of heredity,
of the white-eyed mutation in Drosophila
such as genes, could not explain the continuous range of variation melanogaster
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]


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.

History of evolutionary thought 105

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]

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]
History of evolutionary thought 106

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]

1940s–1960s: Molecular biology and 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,
is responsible for 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]

Late 20th century

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.
History of evolutionary thought 107

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]

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 A phylogenetic tree showing the three-domain system. Eukaryotes
[98] are colored red, Archaea green, and Bacteria blue.
genome data. 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]

Microbiology, horizontal gene transfer, and endosymbiosis

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
History of evolutionary thought 108

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]

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.

21st century

Epigenetic inheritance
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
History of evolutionary thought 109

Lamarckian evolution may occur.[125]

Unconventional evolutionary theory

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.

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.

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.

See also
• Faith and rationality
• Galápagos Islands
• The Voyage of the Beagle

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• 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.
History of evolutionary thought 114

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

Lamarckism (or Lamarckian inheritance) is the idea that an organism can pass on characteristics that it acquired
during its lifetime to its offspring (also known as heritability of acquired characteristics or soft inheritance). It is
named after the French biologist Jean-Baptiste Lamarck (1744–1829), who incorporated the action of soft
inheritance into his evolutionary theories. He is often incorrectly cited as the founder of soft inheritance, which
proposes that individual efforts during the lifetime of the organisms were the main mechanism driving species to
adaptation, as they supposedly would acquire adaptive changes and pass them on to offspring. After publication of
Charles Darwin's theory of natural selection, the importance of individual efforts in the generation of adaptation was
considerably diminished. Later, Mendelian genetics supplanted the notion of inheritance of acquired traits,
eventually leading to the development of the modern evolutionary synthesis, and the general abandonment of the
Lamarckian theory of evolution in biology. In a wider context, soft inheritance is of use when examining the
evolution of cultures and ideas, and is related to the theory of Memetics.

Between 1794 and 1796 Erasmus Darwin wrote Zoönomia suggesting
"that all warm-blooded animals have arisen from one living filament...
with the power of acquiring new parts" in response to stimuli, with
each round of "improvements" being inherited by successive
generations. Subsequently Jean-Baptiste Lamarck repeated in his
Philosophie Zoologique of 1809 the folk wisdom that characteristics
which were "needed" were acquired (or diminished) during the lifetime
of an organism then passed on to the offspring. He incorporated this
mechanism into his thoughts on evolution, seeing it as resulting in the
adaptation of life to local environments.

Lamarck founded a school of French Transformationism which

Jean-Baptiste Lamarck
included Étienne Geoffroy Saint-Hilaire, and which corresponded with
a radical British school of anatomy based in the extramural anatomy
schools in Edinburgh which included the surgeon Robert Knox and the comparative anatomist Robert Edmund
Grant. In addition, the Regius Professor of Natural History, Robert Jameson, was the probable author of an
anonymous paper in 1826 praising "Mr. Lamarck" for explaining how the higher animals had "evolved" from the
"simplest worms" – this was the first use of the word "evolved" in a modern sense. As a young student, Charles
Darwin was tutored by Grant, and worked with him on marine creatures.

The Vestiges of the Natural History of Creation, authored by Robert Chambers in St Andrews and published
anonymously in England in 1844, proposed a theory which combined radical phrenology with Lamarckism, causing
political controversy for its radicalism and unorthodoxy, but exciting popular interest and preparing a huge and
prosperous audience for Darwin.
Lamarckism 115

Darwin's Origin of Species proposed natural selection as the main mechanism for development of species, but did
not rule out a variant of Lamarckism as a supplementary mechanism.[1] Darwin called his Lamarckian hypothesis
Pangenesis, and explained it in the final chapter of his book Variation in Plants and Animals under Domestication,
after describing numerous examples to demonstrate what he considered to be the inheritance of acquired
characteristics. Pangenesis, which he emphasised was a hypothesis, was based on the idea that somatic cells would,
in response to environmental stimulation (use and disuse), throw off 'gemmules' or 'pangenes' which travelled around
the body (though not necessarily in the bloodstream). These pangenes were microscopic particles that supposedly
contained information about the characteristics of their parent cell, and Darwin believed that they eventually
accumulated in the germ cells where they could pass on to the next generation the newly acquired characteristics of
the parents. Darwin's half-cousin, Francis Galton carried out experiments on rabbits, with Darwin's cooperation, in
which he transfused the blood of one variety of rabbit into another variety in the expectation that its offspring would
show some characteristics of the first. They did not, and Galton declared that he had disproved Darwin's hypothesis
of Pangenesis, but Darwin objected, in a letter to Nature, that he had done nothing of the sort, since he had never
mentioned blood in his writings. He pointed out that he regarded Pangenesis as occurring in Protozoa and plants,
which have no blood.[2] With the development of the modern synthesis of the theory of evolution and a lack of
evidence for either a mechanism or even the heritability of acquired characteristics, Lamarckism largely fell from
In the 1920s, experiments by Paul Kammerer on amphibians, particularly the midwife toad, appeared to find
evidence supporting Lamarckism, but his specimens with supposedly-acquired black foot-pads were found to have
been tampered with. In The Case of the Midwife Toad Arthur Koestler surmised that the specimens had been faked
by a Nazi sympathiser to discredit Kammerer for his political views.
A form of Lamarckism was revived in the Soviet Union of the 1930s when Trofim Lysenko promoted Lysenkoism
which suited the ideological opposition of Joseph Stalin to Genetics. This ideologically driven research influenced
Soviet agricultural policy which in turn was later blamed for crop failures.
Since 1988 certain scientists have produced work proposing that Lamarckism could apply to single celled organisms.
A version of Lamarckian acquisition in higher order animals is still posited in certain branches of psychology, as, for
example, in the Jungian racial memory.
Neo-Lamarckism is a theory of inheritance based on a modification and extension of Lamarckism, essentially
maintaining the principle that genetic changes can be influenced and directed by environmental factors.

Lamarck's theory
The identification of Lamarckism with the inheritance of acquired
characteristics is regarded by some as an artifact of the subsequent
history of evolutionary thought, repeated in textbooks without analysis.
Stephen Jay Gould wrote that late 19th century evolutionists "re-read
Lamarck, cast aside the guts of it ... and elevated one aspect of the
mechanics - inheritance of acquired characters - to a central focus it
never had for Lamarck himself."[3] He argued that "the restriction of
"Lamarckism" to this relatively small and non-distinctive corner of
Lamarck's thought must be labelled as more than a misnomer, and
truly a discredit to the memory of a man and his much more The evolution of necks is often used as the
example in explanations of Lamarckism.
comprehensive system"[4] . Gould advocated defining "Lamarckism"
more broadly, in line with Lamarck's overall evolutionary theory.

Lamarck incorporated two ideas into his theory of evolution, in his day considered to be generally true:
Lamarckism 116

1. Use and disuse – Individuals lose characteristics they do not require (or use) and develop characteristics that are
2. Inheritance of acquired traits – Individuals inherit the traits of their ancestors.
Examples of what is traditionally called "Lamarckism" would include:
• Giraffes stretching their necks to reach leaves high in trees (especially Acacias), strengthen and gradually
lengthen their necks. These giraffes have offspring with slightly longer necks (also known as "soft inheritance").
• A blacksmith, through his work, strengthens the muscles in his arms. His sons will have similar muscular
development when they mature.
With this in mind, Lamarck has been credited in some textbooks and popular culture with developing two laws:
1. In every animal which has not passed the limit of its development, a more frequent and continuous use of any
organ gradually strengthens, develops and enlarges that organ, and gives it a power proportional to the length of
time it has been so used; while the permanent disuse of any organ imperceptibly weakens and deteriorates it, and
progressively diminishes its functional capacity, until it finally disappears.
2. All the acquisitions or losses wrought by nature on individuals, through the influence of the environment in which
their race has long been placed, and hence through the influence of the predominant use or permanent disuse of
any organ; all these are preserved by reproduction to the new individuals which arise, provided that the acquired
modifications are common to both sexes, or at least to the individuals which produce the young.[5]
In essence, a change in the environment brings about change in "needs" (besoins), resulting in change in behavior,
bringing change in organ usage and development, bringing change in form over time — and thus the gradual
transmutation of the species.
However, as historians of science such as Michael Ghiselin and Stephen Jay Gould have pointed out, none of these
views were original to Lamarck.[6] [7] On the contrary, Lamarck's contribution was a systematic theoretical
framework for understanding evolution. He saw evolution as comprising two processes;
1. Le pouvoir de la vie (a complexifying force) - in which the natural, alchemical movements of fluids would etch
out organs from tissues, leading to ever more complex construction regardless of the organ's use or disuse. This
would drive organisms from simple to complex forms.
2. L'influence des circonstances (an adaptive force) - in which the use and disuse of characters led organisms to
become more adapted to their environment. This would take organisms sideways off the path from simple to
complex, specialising them for their environment.

Current views on "Lamarckism"

Interest in Lamarckism has recently increased, as several studies in the field of epigenetics have highlighted the
possible inheritance of behavioral traits acquired by the previous generation. Some recent notable studies include
those made by the University of Linköping, Sweden, which have looked at foraging behavior in chickens as well as
stress factors[8] . The conclusion of the referenced study is as follows:
Our findings suggest that unpredictable food access caused seemingly adaptive responses in feeding
behavior, which may have been transmitted to the offspring by means of epigenetic mechanisms,
including regulation of immune genes. This may have prepared the offspring for coping with an
unpredictable environment.... Transmissions of information across generations which does not involve
traditional inheritance of DNA-sequence alleles is often referred to as soft inheritance [9] or 'Lamarckian
On a much smaller scale, genetic exchange without reproduction between single-celled organisms has been
well-researched and is termed horizontal gene transfer.
The group of researchers at The University of Linköping again highlighted the apparent link between food intake and
cross-generational inheritance of acquired traits. This link has been shown before in studies of human populations
Lamarckism 117

who have experienced starvation, where epigenetic factors have altered the functioning of genes[10] . These changed
epigenetic factors appear to show traits in the next generation such as an increased occurrence of diabetes. The
process of methylation is thought to be behind such changes.
In October 2010, further evidence linking food intake to traits inherited by the offspring were shown in a study of
rats conducted by several Australian universities[11] . The study strongly suggested that fathers can transfer a
propensity for obesity to their daughters as a result of the fathers' food intake, and not their genetics (or specific
genes), prior to the conception of the daughter. A "paternal high-fat diet" was shown to cause cell dysfunction in the
daughter, which in turn led to obesity for the daughter.
Several historians have argued that Lamarck's name is linked somewhat unfairly to the theory that has come to bear
his name, and that Lamarck deserves credit for being an influential early proponent of the concept of biological
evolution, far more than for the mechanism of evolution, in which he simply followed the accepted wisdom of his
time. Lamarck died 30 years before the first publication of Charles Darwin's Origin of Species. According to Stephen
Jay Gould, if Lamarck had been aware of Darwin's proposed mechanism of natural selection, there is no reason to
assume he would not have accepted it as a more likely alternative to his own mechanism. Note also that Darwin, like
Lamarck, lacked a plausible alternative mechanism of inheritance - the particulate nature of inheritance was only
observed by Gregor Mendel somewhat later, and published in 1866. Its full significance was not appreciated until the
Modern evolutionary synthesis in the early 1920s. An important point in its favour at the time was that Lamarck's
theory contained a mechanism describing how variation is maintained, which Darwin’s own theory lacked.
Several recent studies, one conducted by researchers at MIT and another by researchers at the Tufts University
School of Medicine, have rekindled the debate once again. As reported [12] in MIT's Technology Review in February
2009, "The effects of an animal's environment during adolescence can be passed down to future offspring ... The
findings provide support for a 200-year-old theory of evolution that has been largely dismissed: Lamarckian
evolution, which states that acquired characteristics can be passed on to offspring."

Unlike neo-Darwinism, the term neo-Lamarckism refers more to a loose grouping of largely heterodox theories and
mechanisms that emerged after Lamarck's time, than to any coherent body of theoretical work.
In the 1920s, Harvard University researcher William McDougall studied the abilities of rats to correctly solve mazes.
He found that children of rats that had learned the maze were able to run it faster. The first rats would get it wrong
165 times before being able to run it perfectly each time, but after a few generations it was down to 20. McDougall
attributed this to some sort of Lamarckian evolutionary process.[13] Oscar Werner Tiegs and Wilfred Eade Agar later
showed McDougall's results to be incorrect, caused by poor experimental controls.[14] [15] [16] [17] [18]
At around the same time, Ivan Pavlov, who was also a Lamarckist, claimed to have observed a similar phenomenon
in animals being subject to conditioned reflex experiments. He claimed that with each generation, the animals
became easier to condition. However, Pavlov never suggested a mechanism to explain these observations.
Soma to germ-line feedback
In the 1970s the immunologist Ted Steele, formerly of the University of Wollongong, and colleagues, proposed a
neo-Lamarckian mechanism to try to explain why homologous DNA sequences from the VDJ gene regions of parent
mice were found in their germ cells and seemed to persist in the offspring for a few generations. The mechanism
involved the somatic selection and clonal amplification of newly acquired antibody gene sequences that were
generated via somatic hyper-mutation in B-cells. The mRNA products of these somatically novel genes were
captured by retroviruses endogenous to the B-cells and were then transported through the blood stream where they
could breach the soma-germ barrier and retrofect (reverse transcribe) the newly acquired genes into the cells of the
germ line. Although Steele was advocating this theory for the better part of two decades, little more than indirect
evidence was ever acquired to support it. An interesting attribute of this idea is that it strongly resembles Darwin's
Lamarckism 118

own theory of pangenesis, except in the soma to germ line feedback theory, pangenes are replaced with realistic
Epigenetic inheritance
Forms of 'soft' or epigenetic inheritance within organisms have been suggested as neo-Lamarckian in nature by such
scientists as Eva Jablonka and Marion J. Lamb. In addition to 'hard' or genetic inheritance, involving the duplication
of genetic material and its segregation during meiosis, there are other hereditary elements that pass into the germ
cells also. These include things like methylation patterns in DNA and chromatin marks, both of which regulate the
activity of genes. These are considered "Lamarckian" in the sense that they are responsive to environmental stimuli
and can differentially affect gene expression adaptively, with phenotypic results that can persist for many
generations in certain organisms. Although the reality of epigenetic inheritance is not doubted (as countless
experiments have validated it), its significance to the evolutionary process is uncertain. Most neo-Darwinians
consider epigenetic inheritance mechanisms to be little more than a specialized form of phenotypic plasticity, with
no potential to introduce evolutionary novelty into a species lineage.[20]

Lamarckism and single-celled organisms

While Lamarckism has been discredited as an evolutionary influence for larger lifeforms, some scientists
controversially argue that it can be observed among microorganisms.[21] Whether such mutations are directed or not
also remains in contention. (see Horizontal Gene Transfer)
In 1988, John Cairns at the Radcliffe Infirmary in Oxford, England, and a group of other scientists renewed the
Lamarckian controversy (which at that point had been a dead debate for many years).[22] The group took a mutated
strain of E. coli that was unable to consume the sugar lactose and placed it in an environment where lactose was the
only food source. They observed over time that mutations occurred within the colony at a rate that suggested the
bacteria were overcoming their handicap by altering their own genes. Cairns, among others, dubbed the process
adaptive mutation.
If bacteria that had overcome their own inability to consume lactose passed on this "learned" trait to future
generations, it could be argued as a form of Lamarckism; though Cairns later chose to distance himself from such a
position.[23] More typically, it might be viewed as a form of ontogenic evolution.
There has been some research into Lamarckism and prions. A group of researchers, for example, discovered that in
yeast cells containing a specific prion protein Sup35, the yeast were able to gain new genetic material, some of
which gave them new abilities such as resistance to a particular herbicide. When the researchers mated the yeast cells
with cells not containing the prion, the trait reappeared in some of the resulting offspring, indicating that some
information indeed was passed down, though whether or not the information is genetic is debatable: trace prion
amounts in the cells may be passed to their offspring, giving the appearance of a new genetic trait where there is
Finally, there is growing evidence that cells can activate low-fidelity DNA polymerases in times of stress to induce
mutations. While this does not directly confer advantage to the organism on the organismal level, it makes sense at
the gene-evolution level. While the acquisition of new genetic traits is random, and selection remains Darwinian, the
active process of identifying the necessity to mutate is considered to be Lamarckian.

Lamarckism and societal change

Jean Molino (2000) has proposed that Lamarckian evolution may be accurately applied to cultural evolution. This
was also previously suggested by Peter Medawar (1959) and Conrad Waddington (1961). K. N. Laland and
colleagues have recently suggested that human culture can be looked upon as an ecological niche like phenomena,
where the effects of cultural niche construction are transmissible from one generation to the next. One interpretation
of the Meme theory is that memes are both Darwinian and Lamarckian in nature, as in addition to being subject to
Lamarckism 119

selection pressures based on their ability to differentially influence Human minds, memes can be modified and the
effects of that modification passed on. Richard Dawkins notes (in Blackmore 2000: The Meme machine, page 13),
that Memes can be copied in a Lamarckian way (copying of the product) or in a Weismann-type evolutionary way
(copying of the instruction) which is much more resistant against changes.

See also
• Jean-Baptiste Lamarck
• Baldwinian evolution
• Darwinism
• Epigenetic inheritance
• Epigenetics
• Evolution
• Inheritance of acquired characters
• Lysenkoism
• Memetics
• Obsolete scientific theories
• Orthogenesis
• Marcus Pembrey
• Racial memory
• Ted Steele
• History of evolutionary thought
• Eclipse of Darwinism

[1] Desmond, A. & Moore, J. (1991) Darwin Penguin Books p.617 "Darwin was loathe to let go of the notion that a well-used and strengthened
organ could be inherited"
[2] Charles Darwin (27 April 1871). "Pangenesis" (http:/ / darwin-online. org. uk/ content/ frameset?viewtype=text& itemID=F1751&
pageseq=1). Nature. A Weekly Illustrated Journal of Science. pp. 502–503. . Retrieved 2007-11-08.
[3] Gould, Stephen J. "Shades of Lamarck", reprinted in The Panda's Thumb (1980) pp.65-71. Quote from page 66.
[4] Gould, Stephen J. (2002). The Structure of Evolutionary Theory. Harvard: Belknap Harvard. pp. 177–178. ISBN 0-674-00613-5.
[5] Jean-Baptiste Lamarck Zoological Philosophy trans. Hugh Elliot, U. Chicago Press, 1984, p.113
[6] The Imaginary Lamarck: a look at bogus "history" in schoolbooks (http:/ / www. textbookleague. org/ 54marck. htm) by Michael Ghiselin
[7] Gould, S.J. (2002) The Structure of Evolutionary Theory
[8] http:/ / www. plosone. org/ article/ info:doi/ 10. 1371/ journal. pone. 0006405
[9] Richards EJ (2006) Inherited epigenetic variation–revisiting soft inheritance. Nat Rev Genet 7(5): 395–401.
[10] http:/ / cat. inist. fr/ ?aModele=afficheN& cpsidt=3596539
[11] http:/ / www. nature. com/ nature/ journal/ v467/ n7318/ full/ nature09491. html
[12] http:/ / www. technologyreview. com/ biomedicine/ 22061
[13] McDougall, 1938. British Journal of Psychology 28:321-345
[14] Pantin, C F A (November 1957). "Oscar Werner Tiegs". Biographical Memiors of Fellows of the Royal Society (The Royal Society) 3: 247.
[15] W E Agar, F H Drummond (1935). "First report on a test of McDougall's Lamarckian experiment on the training of rats". Journal of
Experimental Biology 12: 191.
[16] W E Agar, F H Drummond (1942). "Second report on a test of McDougall's Lamarckian experiment on the training of rats". Journal of
Experimental Biology 19: 158.
[17] W E Agar, F H Drummond (1948). "Third report on a test of McDougall's Lamarckian experiment on the training of rats". Journal of
Experimental Biology 25: 103.
[18] W E Agar, F H Drummond, M M Gunson (1954). "Fourth (final) report on a test of McDougall's Lamarckian experiment on the training of
rats". Journal of Experimental Biology 31: 308.
[19] Lamarck's Signature: how retrogenes are changing Darwin's natural selection paradigm. Edward J. Steele, Robyn A. Lindley, Robert V.
Blanden. Perseus Books, 1998
[20] Epigenetic Inheritance and Evolution: The Lamarckian Dimension. Eva Jablonka, Marion J. Lamb. Oxford University Press, 1995
[21] Adaptive mutation Genetics, Vol. 148, April 1998
Lamarckism 120

[22] http:/ / www. mun. ca/ biochem/ courses/ 4103/ topics/ adaptive_mutation. html Adaptive mutation in bacteria
[23] Adaptive mutation in E. coli, Journal of Bacteriology, August 2004, Vol. 186, No. 15
[24] Lamarckism and prions, New Scientist, 21 August 2004, Issue #2461

Further references
• Burkeman, Oliver. Why everything you've been told about evolution is wrong (
science/2010/mar/19/evolution-darwin-natural-selection-genes-wrong), The Guardian, March 19, 2010.
• Desmond, Adrian (1989). The Politics of Evolution: Morphology, Medicine, and Reform in Radical London.
Chicago: University of Chicago Press. ISBN 0-226-14374-0.
• Gould, Stephen J. (2002). The Structure of Evolutionary Theory. Harvard: Belknap Harvard. pp. 170–197 on
Lamarck. ISBN 0-674-00613-5.
• Medawar, Peter (1959). "The threat and the glory". BBC Reith Lectures No. 6.
• Molino, Jean (2000). "Toward an Evolutionary Theory of Music and Language". In Brown, Merker & Wallin
(Eds.), The Origins of Music, ISBN 0-262-23206-5.
• Waddington, Conrad (1961). "The human evolutionary system". In: Michael Banton (Ed.), Darwinism and the
Study of Society. London: Tavistock.
• Cairns, J., J. Overbaugh, and S. Miller. 1988. Nature 335: 142-145
• Culotta, Elizabeth; "A Boost for 'Adaptive' Mutation", Science, 265:318, 1994.
• Vetsigian K, Woese C, Goldenfeld N. 2006. "Collective Evolution and the Genetic Code." PNAS 103:
• Hall Barry G., Adaptive Evolution That Requires Multiple Spontaneous Mutations. I. Mutations Involving an
Insertion Sequence (

External links
• Nonsense in Schoolbooks - The Imaginary Lamarck (
T. Ghiselin recounts Lamarck's times and writings.
• Jean-Baptiste Lamarck : works and heritage ( an English/French web
site edited by Pietro Corsi (Oxford Univ.) and realised by CNRS (France - IT team of CRHST). This web site
contents all books, texts, manuscripts and the lamarck's herbarium.
• Guralnick, Rob, et. al (2006). "Jean-Baptiste Lamarck (1744-1829)" (
lamarck.html). A History of Evolutionary Thought. University of California Museum of Paleontology. Retrieved
3 July 2010.
• The Sins of the Fathers, Take 2 ( "At tributes to Darwin,
Lamarckism — inheritance of acquired traits — will be the skunk at the party." By Sharon Begley, Newsweek.
From the magazine issue dated January 26, 2009.
Saltationism 121

In biology, saltation (from Latin, saltus, "leap") is a sudden change from one generation to the next, that is large, or
very large, in comparison with the usual variation of an organism. The term is used for occasionally hypothesized,
nongradual changes (especially single-step speciation) that are atypical of, or violate, standard concepts - gradualism
- involved in neo-Darwinian evolution.
Saltation does not fit into contemporary evolutionary theory, but there are some prominent proponents, including
Carl Woese. Woese, and colleagues, suggested that the absence of RNA signature continuum between domains of
bacteria, archaea, and eukarya constitutes a primary indication that the three primary organismal lineages
materialized via one or more major evolutionary saltations from some universal ancestral state involving dramatic
change in cellular organization that was significant early in the evolution of life, but in complex organisms gave way
to the generally accepted Darwinian mechanisms.[1]
Polyploidy (most common in plants but not unknown in animals) is considered a type of saltation [2] , even though
most polyploid individuals are sterile. Polyploidy meets the basic criteria of saltation in that a significant change (in
gene numbers) results in speciation in just one generation. Mammalian liver cells are typically polyploidal, but they
are not part of the germ line.

Punctuated Equilibrium
It is a popular misconception that punctuated equilibrium is a saltationist theory, often mistaken for Richard
Goldschmidt's hypothesis of "Hopeful Monsters."[3] However, punctuated equilibrium refers instead to a pattern of
evolution where most speciation occurs relatively rapidly from a geological perspective (tens of thousands of years
instead of millions of years), but through neo-Darwinian evolution, not by saltations.

Pop culture
In popular culture, a form of saltation appears to have emerged from misconceptions over currently accepted theories
of evolution (the X-men and its various spin-offs being the most egregious examples). It was also fictionalized in
Greg Bear's novel, Darwin's Radio.
The Teenage Mutant Ninja Turtles ( and many other, similar science-fiction pieces ) are not examples of
saltationism, however. A saltation would be a substantial change that takes place during reproduction; a case when a
child belongs to a different species than its parents - between generations, not during a generation.

See also
• Catastrophism
• Phyletic gradualism
• Rapid modes of evolution
• The Blind Watchmaker
• History of evolutionary thought
• Eclipse of Darwinism
Saltationism 122

Notes and references

[1] Elijah Roberts, Anurag Sethi†, Jonathan Montoya, Carl R. Woese, and Zaida Luthey-Schulten (May 19, 2008). "Molecular signatures of
ribosomal evolution" (http:/ / www. pnas. org/ content/ 105/ 37/ 13953. full?sid=f3651397-00e9-4a57-802b-f41c6ef6cf5a). Proceedings of the
National Academy of Sciences. .
[2] France Dufresne, Paul D. N. Herbert (1994). "Hybridization and origins of polyploidy" (http:/ / www. jstor. org/ pss/ 49988). Proceedings of
the Royal Society. . Retrieved 2010-05-06.
[3] Gould, Stephen Jay. "Punctuated Equilibrium's Threefold History" (http:/ / www. stephenjaygould. org/ library/ gould_structure. html). The
Structure of Evolutionary Theory. Harvard University Press. pp. 1006–1021. . Retrieved 2008-05-05. "[T]he urban legend rests on the false
belief that ... punctuated equilibrium became a saltational theory wedded to Goldschmidt's hopeful monsters as a mechanism. I have labored to
refute this nonsensical charge from the day I first heard it."

Orthogenesis, orthogenetic evolution, progressive evolution or autogenesis, is the hypothesis that life has an
innate tendency to move in an unilinear fashion due to some internal or external "driving force". The hypothesis is
based on essentialism and cosmic teleology and proposes an intrinsic drive which slowly transforms species. George
Gaylord Simpson (1953) in an attack on orthogenesis called this mechanism "the mysterious inner force".[1] Classic
proponents of orthogenesis have rejected the theory of natural selection as the organising mechanism in evolution,
and theories of speciation for a rectilinear model of guided evolution acting on discrete species with "essences". The
term orthogenesis was popularised by Theodor Eimer, though many of the ideas are much older (Bateson 1909).[2]

No Goal
Orthogenesis does not postulate a "goal" for evolution. Though it proceeds in a linear fashion driven by some
internal mechanism, it does not have a goal.
Many sources mix this heterodox view of evolution with another—- that evolution is proceeding to some long term
or ultimate goal; the result are definitions that state "orthogenesis proposes that evolution moves in a unilinear
fashion towards a perfect goal". While it is true that early and famous examples of orthogenesis often conflated these
two ideas (e.g. Jean-Baptiste Lamarck's theory of evolution), it is important to recognize that these are in fact two
separate ideas that are rejected by mainstream science; the latter idea of goal-oriented evolution is better understood
as a form of teleology.
The distinction can be seen when we recognize that orthogenesis is inherent in the theories of German biologist Ernst
Haeckel and American paleontologist Richard Swann Lull. Both scientists proposed mechanisms whereby evolution
proceeded in unilinear fashion, but neither saw goals (instead they made pseudo-scientific appeals to unknown
genetic driving processes).
This is important because similar flaws occur recurrently at the fringes of science, typically taking the form of
mysterious molecular drives that supposedly are pushing phenotypic evolution in certain directions or forcing the
formation of new species.

The orthogenesis hypothesis had a significant following in the 19th century when a number of evolutionary
mechanisms, such as Lamarckism, were being proposed. Jean-Baptiste Lamarck himself accepted the idea, and it had
a central role in his theory of inheritance of acquired characteristics, the hypothesised mechanism of which
resembled the "mysterious inner force" of orthogenesis. Other proponents of orthogenesis included Leo Berg,
philosopher Henri Bergson and, for a time, the paleontologist Henry Fairfield Osborn. Orthogenesis was particularly
accepted by paleontologists who saw in their fossils a directional change, and in invertebrate paleontology thought
there was a gradual and constant directional change. Those who accepted orthogenesis in this way, however, did not
Orthogenesis 123

necessarily accept that the mechanism that drove orthogenesis was teleological. In fact, Darwin himself rarely used
the term "evolution" now so commonly used to describe his theory, because in Darwin's time, evolution usually was
associated with some sort of progressive process like orthogenesis, and this had been common usage since at least

Comparison of Theories

Comparison of different theories of evolution

Darwinism Orthogenesis Lamarckism

Mechanism Short-sighted Natural Selection sorting Intrinsic drive towards perfection; Intrinsic drive towards perfection and
random genetic variation, no other natural selection unimportant. inheritance of acquired characteristics (both
guidance or aim. Selected traits are Characters produced may be totally are Lamarckian principles); natural selection
adaptive, i.e. have some survival value. non-adaptive, i.e. have no survival adopted by some in latter years.

Common Yes, new species coming into existence No, speciation rejected or considered Depends upon source quoted. Signs that
descent by speciation events. unimportant in long term trends; species shared a common ancestor were
spontaneous generation of new species detected before Darwin, but in absence of a
resulting in parallel evolution. mechanism some still rejected the idea.

Status Prevailing in modified form as modern Refuted by Charles Darwin's Origin of Declined after the Origin, though the
evolutionary synthesis. Species and the modern evolutionary mechanism was not refuted until the modern
synthesis. evolutionary synthesis in which it was
established that the mechanism does not exist.

Collapse of the hypothesis

The orthogenesis hypothesis began to collapse when it became clear that it could not explain the patterns found by
paleontologists in the fossil record, which was non-linear with many complications. The hypothesis was generally
abandoned when no mechanism could be found that would account for the process, and the theory of evolution by
natural selection became the prevailing theory of evolution. The modern evolutionary synthesis, in which the genetic
mechanisms of evolution were discovered, refuted the hypothesis for good. As more was understood about these
mechanisms it became obvious that there was no possible naturalistic way in which the newly discovered mechanism
of heredity could be far-sighted or have a memory of past trends.
The orthogenetic hypothesis, however, died hard. Even Darwin was at first not opposed to orthogenic thinking, as
this quote from the 1911 Encyclopedia Britannica demonstrates:
Darwin and his generation were deeply imbued with the Butlerian tradition, and regarded the organic world as
almost a miracle of adaptation, of the minute dovetailing of structure, function and environment. Darwin
certainly was impressed with the view that natural selection and variation together formed a mechanism, the
central product of which was adaptation. From the Butlerian side, too, came the most urgent opposition to
Darwinism. How is it possible, it was said, that fortuitous variations can furnish the material for the precise
and balanced adaptations that all nature reveals? Selection cannot create the materials on which it is supposed
to operate; the beginnings of new organs, the initial stages of new functions cannot be supposed to have been
useful. Moreover, many naturalists, especially those concerned with palaeontology, pointed to the existence of
orthogenetic series, of long lines of ancestry, which displayed not a sporadic differentiation. in every direction,
but apparently a steady and progressive march in one direction.[4]
Edward Drinker Cope put such a line of argument in the most cogent fashion; the course of evolution,
both in the production of variations and their selection, seemed to him to imply the existence of an
originative, conscious and directive force, for which he invented the term bathmism (Gr. βαθμ, a step or
Orthogenesis 124

beginning). On the other hand, dislike of mystical interpretations of natural facts has driven many
capable naturalists to another extreme and has led them to insist on the all-powerfulness of natural
selection and on the complete indefiniteness of variation. The apparent opposition between the
conflicting schools is more acute than the facts justify.... there is no connection between the appearance
of the variation and the use to which it may be put... in one sense it is a mere coincidence if a particular
variation turn out to be useful. But there are several directions in which the field of variation appears to
be not only limited but defined in a certain direction. Obviously variations depend on the constitution of
the varying organism; a modification, whether it be large or small, is a modification of an already
definite and limited structure.... A continuous environment both from the point of view of production of
variation and selection of variation would appear necessarily to result in a series with the appearance of
orthogenesis. The past history of the organic world displays many successful series and these, as they
have survived, must inevitably display orthogenesis to some extent; but it also displays many failures
which indeed may be regarded as showing that the limitation of variation has been such that the
organisms have lost the possibility of successful response to a new environment.[4]
A few hung on to the orthogenesis hypothesis as late as the 1950s by claiming that the processes of
macroevolution, the long term trends in evolution, were distinct from the processes of microevolution (genetic
variation and natural selection) which were by then well understood and it was known they could not behave
in an orthogenetic manner. Teilhard de Chardin, a Jesuit paleontologist, in The Phenomenon of Man (a book
influential among non-scientists that was published four years after his death in 1959) argued for evolution
aiming for the "Omega Point", while putting man at the center of the universe and accounting for original sin
(Dennett 1995, von Kitzing 1998). The term Chardin used for this was "directed additivity". This form of
orthogenesis has now also been abandoned as more about evolutionary processes has been discovered
(Wilkins 1997). The refutation of orthogenesis had some ramifications in the field of philosophy, as it refuted
the idea of teleology as first postulated by Aristotle and accepted by Immanuel Kant, who had greatly
influenced many scientists. Before the scientific and philosophical revolution that began with Charles Darwin's
ideas, the prevailing philosophy was that the world was teleological and purposeful, and that science was the
study of God's creation. The refutation of these concepts have led to a shift in what science and scientists are
perceived to be.

Modern co-opted usage

Though linear teleological evolution has been refuted, it is not true that evolution never proceeds in a linear way,
reinforcing characteristics, in certain lineages at times, for example, during a period of slow, sustained environmental
change, but such examples are entirely consistent with the modern neo-Darwinian theory of evolution. These
examples have sometimes been referred to as orthogenetic (e.g. by Jacobs et al. 1995 & Woodley 2006) but are not
strictly orthogenetic, and simply appear as linear and constant changes because of environmental and molecular
constraints on the direction of change.
Orthogenesis 125

See also
• Facilitated variation
• Evolution of complexity
• Eclipse of Darwinism
• History of evolutionary thought
• Law of Complexity/Consciousness

[1] George Gaylord Simpson, Life of the Past: An Introduction to Paleontology, Yale University Press, New Haven, 1953, p. 125.
[2] The evolutionary future of man: A biological view of progress (http:/ / www. simonyi. ox. ac. uk/ dawkins/ WorldOfDawkins-archive/
Dawkins/ Work/ Articles/ 1993-09-11future. shtml)
[3] Darwin's Dilemma: The Odyssey of Evolution (http:/ / web. archive. org/ web/ 20051216025526/ http:/ / wordorigins. org/ wordore.
htm#evolution), Stephen Jay Gould, an essay in Ever Since Darwin: Reflections in Natural History, W. W. Norton, 1977, ISBN
[4] The Encyclopaedia Britannica: A Dictionary of Arts, Sciences, Literature and General Information, Eleventh Edition, Copyright in all
countries subscribing the Berne Convention by the Chancellor, Masters and Scholars of the University of Cambridge, Copyright in the United
States of America by the Encyclopaedia Britannica Company, London, May 31, 1911.

1. Bateson, William, 1909. Heredity and variation in modern lights, in Darwin and Modern Science (A.C. Seward
ed.). Cambridge University Press. Chapter V. E-book (
2. Dennett, Daniel, 1995. Darwin's Dangerous Idea. Simon & Schuster.
3. Huxley, Julian, 1942. The Modern Evolutionary Synthesis, London: George Allen and Unwin.
4. Jacobs, Susan C., Allan Larson & James M. Cheverud, 1995. Phylogenetic Relationships and Orthogenetic
Evolution of Coat Color Among Tamarins (Genus Saguinus). Syst. Biol. 44(4):515—532, Abstract (http://
5. Mayr, Ernst, 2002. What Evolution Is, London: Weidenfeld and Nicolson.
6. Simpson, George G., 1957. Life Of The Past: Introduction to Paleontology. Yale University Press, p.119.
7. Wilkins, John, 1997. What is macroevolution?. TalkOrigins Archive (
macroevolution.html) (14:08 UTC, Oct 13 2004)
8. Woodley, Michael A., 2006. The Limits of Ecology: New Perspectives from a Theoretical Borderland. Abramis
Academic Press.
''On the Origin of Species'' 126

On the Origin of Species

On the Origin of Species

The title page of the 1859 edition

of On the Origin of Species
Author Charles Darwin

Country United Kingdom

Language English

Subject(s) Natural selection

Evolutionary biology

Genre(s) science, biology

Publisher John Murray

Publication [2]
24 November 1859

Media type Print (Hardback)

Pages 502


Preceded by On the Tendency of Species to form Varieties; and on the Perpetuation of 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.
''On the Origin of Species'' 127

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.

Summary of Darwin's theory

Darwin's theory of evolution is based on key facts and the inferences
drawn from them, which biologist Ernst Mayr summarised as
• 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
• 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 Darwin pictured shortly before publication
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).


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]
''On the Origin of Species'' 128

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
Cuvier's 1799 paper on living and fossil elephants [8]
helped establish the reality of extinction.

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]

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]
''On the Origin of Species'' 129

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 In mid-July 1837 Darwin started his "B"
notebooks, on the possibility that "one species does change into notebook on Transmutation of Species, and on
another" to explain these findings, and around July sketched a page 36 wrote "I think" above his first
evolutionary tree.
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]

Further development
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]
''On the Origin of Species'' 130

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
Darwin researched how the skulls of different scientists, including Adam Sedgwick, attacked its morality and
pigeon breeds varied, as shown in his Variation scientific errors.[35] Vestiges had significant influence on public
of Plants and Animals Under Domestication of
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]
''On the Origin of Species'' 131


Events leading to publication

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
A photograph of Alfred Russel
thought it worthwhile. The mechanism was similar to Darwin's own theory.[40] Wallace (1823–1913) taken in
Darwin wrote to Lyell that "your words have come true with a vengeance, ... Singapore in 1862
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
⁄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.[2] 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]
''On the Origin of Species'' 132

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]

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.[2] 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 revisions to the sixth edition of the Origin,
using the word "evolution" for the first time,[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]
''On the Origin of Species'' 133

Publication outside Great Britain

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.[2] [65]

The book was widely translated in Darwin's life time, but problems
arose with translating concepts and metaphors, and some translations
American botanist Asa Gray (1810–1888) 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.[2] 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]
''On the Origin of Species'' 134


Title pages and introduction

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 John Gould's illustration of Darwin's Rhea was
published in 1841. The existence of two rhea
on the origin of species—that mystery of mysteries, as it
species with overlapping ranges influenced
has been called by one of our greatest philosophers.[77] Darwin.

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]

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
''On the Origin of Species'' 135

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

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
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
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.
''On the Origin of Species'' 136

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
This tree diagram, used to show the divergence of species, is the only illustration in the
coadaptations between all organic Origin of Species.
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]

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]
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
''On the Origin of Species'' 137

the 1930s and 1940s that a model of heredity became completely integrated with a model of variation.[104]

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
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
''On the Origin of Species'' 138

evolved through natural selection because intermediate stages would not have been adaptive. Darwin proposed
scenarios for the incremental evolution of each feature.[111]

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]

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]
''On the Origin of Species'' 139

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]

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
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.

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]
''On the Origin of Species'' 140

Structure and style

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]

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]
''On the Origin of Species'' 141

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 British cartoonists presented Darwin's theory in
was triumphant.[139] an unthreatening way. In the 1870s iconic
caricatures of Darwin with an ape or monkey
With the exception of a brief hint in the final chapter, Darwin had body emphasised his significance in transforming
ideas, and contributed to widespread
avoided the subject of human evolution. Despite this, the first review
identification of evolutionism with
claimed it made a creed of the "men from monkeys" idea from [137]
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]

Impact on the scientific community

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]
''On the Origin of Species'' 142

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
Huxley used illustrations to show that humans and apes had the same basic skeletal
religious interference, and his article in the
[146] 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]
''On the Origin of Species'' 143

Impact outside Great Britain

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
Haeckel showed a main trunk leading to mankind
evolutionism in a form of neo-Lamarckism involving recapitulation
with minor branches to various animals, unlike
theory.[158] Darwin's branching evolutionary tree.

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]

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
''On the Origin of Species'' 144

rediscovery of Mendelian inheritance invalidated Darwin's views.[160] [161]

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]
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]
The liberal theologian Baden Powell defended
Baden Powell praised "Mr Darwin's masterly volume [supporting] the
evolutionary ideas by arguing that the
grand principle of the self-evolving powers of nature".[172] In America,
introduction of new species should be considered
Asa Gray argued that evolution is the secondary effect, or modus
a natural rather than a miraculous process.
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]
''On the Origin of Species'' 145

Modern influence
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 A modern phylogenetic tree based on genome analysis shows the three-domain
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]

See also
• 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
• History of evolutionary thought
• History of biology
''On the Origin of Species'' 146

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[2] Freeman 1977
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[4] Darwin 1872, p.  xiii (http:/ / darwin-online. org. uk/ content/ frameset?viewtype=text& itemID=F391& pageseq=18)
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[8] Bowler 2003, pp. 27–36, 39–42, 57–62, 67, 70, 77–80
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[11] Bowler 2003, pp. 111–114
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[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
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• Browne, E. Janet (2002), Charles Darwin: Vol. 2 The Power of Place, London: Jonathan Cape,
ISBN 0-7126-6837-3
• Crawford, J. (1859), "(Review of) On the Origin of Species" (
frameset?itemID=CUL-DAR226.1.50&viewtype=text&pageseq=1), Examiner: 722–723. Published
• Darwin, Charles (1845), Journal of Researches Into the Natural History and Geology of the Countries Visited
During the Voyage of H.M.S. Beagle Round the World, Under the Command of Captain Fitz Roy, R.N. (http:// (2nd ed.), London: John
Murray, retrieved 2009-04-22
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Races in the Struggle for Life (
pageseq=1) ( Full image view (
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• Darwin, Charles (1861), On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured
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• Darwin, Charles (1866), On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured
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• Darwin, Charles (1869), On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured
Races in the Struggle for Life (
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• Darwin, Charles (1871), [[The Descent of Man, and Selection in Relation to Sex (
EditorialIntroductions/Freeman_TheDescentofMan.html)]] (1st ed.), London: John Murray, retrieved
• Darwin, Charles (1872), The Origin of Species by Means of Natural Selection, or the Preservation of Favoured
Races in the Struggle for Life (
pageseq=1) (6th ed.), London: John Murray, retrieved 2009-01-09
• Darwin, Charles (1958), Barlow, Nora, ed., [[The Autobiography of Charles Darwin (
uk/EditorialIntroductions/Freeman_LifeandLettersandAutobiography.html)] 1809–1882. With the Original
Omissions Restored. Edited and with Appendix and Notes by his Granddaughter Nora Barlow], London: Collins,
retrieved 2009-01-09
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• Darwin, Charles (2006), "Journal" (

itemID=CUL-DAR158.1-76&pageseq=1), in van Wyhe, John, [Darwin's personal 'Journal' (1809-1881) (http:/
/], Darwin Online,
CUL-DAR158.1-76, retrieved 2010-09-07
• Darwin, Charles; Costa, James T. (2009), The Annotated Origin: A Facsimile of the First Edition of On the Origin
of Species Annotated by James T. Costa, Cambridge, Massachusetts, and London, England: Belknap Press of
Harvard University Press, ISBN 978-0-674-03281-1
• Desmond, Adrian (1989), The Politics of Evolution: Morphology, Medicine, and Reform in Radical London,
Chicago: University of Chicago Press, ISBN 0-226-14374-0
• Desmond, Adrian; Moore, James (1991), Darwin, London: Michael Joseph, Penguin Group, ISBN 0-7181-3430-3
• Dewey, John (1994), "The Influence of Darwinism on Philosophy", in Martin Gardner, Great Essays in Science,
Prometheus Books, ISBN 0-87975-853-8
• Eldredge, Niles (2006), "Confessions of a Darwinist" (
eldredge-confessions-darwinist/), The Virginia Quarterly Review (Spring 2006): 32–53, retrieved 2008-11-04
• Forster, Roger; Marston, Dr Paul (1999), "Genesis Through History" (
aspx?id=6826), Reason Science and Faith (Ivy Cottage: E-Books ed.), Chester, England: Monarch Books,
ISBN 1-85424-441-8, retrieved 2009-03-24
• Freeman, Richard B. (1977), "On the Origin of Species" (
Freeman_OntheOriginofSpecies.html), The Works of Charles Darwin: An Annotated Bibliographical Handlist
(2nd ed.), Folkestone, England: Dawson, ISBN 0712907408, retrieved 2009-02-22
• Herbert, Sandra, ed. (1980), "The Red Notebook of Charles Darwin" (
frameset?viewtype=side&itemID=F1583e&pageseq=1), Bulletin of the British Museum (Natural History)
Historical Series 7: 1–164 Also available here (
• Hodge, Charles (1874), What is Darwinism? (, Scribner
Armstrong, retrieved 2007-01-14
• Huxley, Thomas Henry (1860), "Darwin on the Origin of Species" (
frameset?itemID=A32&viewtype=text&pageseq=1), Westminster Review 17 (April 1860): 541–570. Published
• Huxley, Thomas (1863), Six Lectures to Working Men "On Our Knowledge of the Causes of the Phenomena of
Organic Nature" (Republished in Volume II of his Collected Essays, Darwiniana) (
huxley/CE2/Phen.html), retrieved 2006-12-15
• Keynes, Richard, ed. (2000), Charles Darwin's Zoology Notes & Specimen Lists from H.M.S. Beagle., Cambridge
University Press, ISBN 052167350X
• Kreeft, Peter (2001), Catholic Christianity, San Francisco: Ignatius Press, ISBN 0-89870-798-6
• Larson, Edward J. (2004), Evolution: The Remarkable History of a Scientific Theory, New York: Modern Library,
ISBN 0-8129-6849-2
• Leifchild (1859), "Review of 'Origin'" (
itemID=CUL-DAR226.1.8&pageseq=1), Athenaeum (No. 1673, 19 November 1859), retrieved 2008-11-22
• Lucas, John R. (1979), "Wilberforce and Huxley: A Legendary Encounter" (
legend.html), The Historical Journal 22 (2): 313–330, doi:10.1017/S0018246X00016848, retrieved 2008-11-22
• Malthus, Thomas Robert (1826), An Essay on the Principle of Population: A View of its Past and Present Effects
on Human Happiness; with an Inquiry into Our Prospects Respecting the Future Removal or Mitigation of the
Evils which It Occasions ( (6th ed.), London: John
Murray, retrieved 2006-12-15
• Mayr, Ernst (1982), The Growth of Biological Thought, Harvard University Press, ISBN 0-674-36446-5
• Miles, Sara Joan (2001), "Charles Darwin and Asa Gray Discuss Teleology and Design" (
ASA/PSCF/2001/PSCF9-01Miles.html), Perspectives on Science and Christian Faith 53: 196–201, retrieved
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• Mivart, St. George Jackson (1871), On the Genesis of Species, New York: Appleton
• Moore, James (2006), Evolution and Wonder – Understanding Charles Darwin (http://speakingoffaith., Speaking of Faith (Radio Program), American Public
Media, retrieved 2008-11-22
• Phipps, William E. (1983), "Darwin, the Scientific Creationist" (
asp?title=1681), Christian Century (September 14–21, 1983): 809–811, retrieved 2007-01-11
• Peckham, Morse (ed.) (1959), The Origin of Species: a variorum text (2006 reprint ed.), Philadelphia: University
of Pennsylvania Press., ISBN 978-0-8122-1954-8
• Quammen, David (2006), The Reluctant Mr. Darwin, New York: Atlas Books, ISBN 0-393-05981-2
• Rhodes, Frank H.T. (June, 1987), "Darwinian Gradualism and Its Limits: The development of Darwin's Views on
the Rate and Pattern of Evolutionary Change" (,
Journal of the History of Biology, Humanities, Social Sciences and Law (Springer Netherlands) 20 (2): 139–157,
Saturday, 6 November 2004, doi:10.1007/BF00138435
• Schopf, J. William; Scheibel, Arnold B. (1997), The Origin and Evolution of Intelligence, Boston: Jones and
Bartlett, ISBN 0-7637-0365-6
• Schopf, J. William (2000), "Solution to Darwin's dilemma: Discovery of the missing Precambrian record of life"
(, Proceedings of the National Academy of
Sciences of the USA 97 (13): 6947–6953, doi:10.1073/pnas.97.13.6947, PMID 10860955, PMC 34368, retrieved
• Secord, James A. (2000), Victorian Sensation: The Extraordinary Publication, Reception, and Secret Authorship
of Vestiges of the Natural History of Creation, Chicago: University of Chicago Press, ISBN 0-226-74411-6
• Spencer, Herbert (1864), The Principles of Biology, Vol. 1 (,
London: Williams and Norgate
• van Hoorn, Marijn (2009), Teyler, Winkler, Darwin (Lecture given at the Congress of the European Botanical and
Horticultural Libraries Group, Prague, 23 April 2009) (
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• van Wyhe, John (2007), "Mind the gap: Did Darwin Avoid Publishing his Theory for Many Years?" (http://, Notes and Records of the
Royal Society 61: 177–205, doi:10.1098/rsnr.2006.0171, retrieved 2009-01-05
• van Wyhe, John (2008), Darwin: The Story of the Man and His Theories of Evolution, London: Andre Deutsch,
ISBN 0-233-00251-0
• van Wyhe, John (2009), Charles Darwin: Gentleman Naturalist: A Biographical Sketch (http://darwin-online., The Complete Works of Charles Darwin Online, retrieved 2009-06-06

Further reading
• Janet Browne (2007). Darwin's Origin of Species: A Biography. ISBN 978-0871139535
• David N. Reznick (2009) The Origin Then and Now: An Interpretive Guide to the Origin of Species ISBN

Contemporary reviews
• Carpenter, William Benjamin (1859), "Darwin on the Origin of Species" (
frameset?itemID=A17&viewtype=text&pageseq=1), National Review 10 (December 1859): 188–214. Published
''On the Origin of Species'' 153

• Gray, Asa (1860), "(Review of) The Origin of Species" (

frameset?itemID=A213&viewtype=text&pageseq=1), Athenaeum (1710: 4 August 1860): 161. 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"" (
content/frameset?itemID=A43&viewtype=image&pageseq=1), Macmillan's Magazine 1: 142–148.
• Huxley, Thomas Henry (1859), "Darwin on the Origin of Species" (
frameset?itemID=A166&viewtype=text&pageseq=1), The Times (26 December 1859): 8–9. Published
• Jenkin, Fleeming (1867), "(Review of) The Origin of Species" (
frameset?itemID=A24&viewtype=text&pageseq=1), North British Review 46 (June 1867): 277–318. Published
• Murray, Andrew (1860), "On Mr Darwin's Theory of the Origin of Species" (
content/frameset?itemID=A14&viewtype=text&pageseq=1), Proceedings of the Royal Society of Edinburgh 4:
• Owen, Richard (1860), "Review of Darwin's Origin of Species" (
frameset?itemID=A30&viewtype=text&pageseq=1), Edinburgh Review 3 (April 1860): 487–532. Published
• 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" (
frameset?itemID=A19&viewtype=text&pageseq=1), Quarterly Review 108 (215: July 1860): 225–264.
Published anonymously.
• For further reviews, see Darwin Online: Reviews & Responses to Darwin (
html), Darwin Online, 10 March 2009, retrieved 2009-06-18

External links
• 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
• Origin of Species (, full text with embedded
• Victorian Science Texts (
• Darwin Correspondence Project Home Page (, University Library,
• PDF scans at ( species) AND creator:(darwin))
Darwinism 154

Darwinism is a set of movements and concepts related to ideas of
transmutation of species or evolution, including ideas with no
connection to the work of Charles Darwin.[1] [2] [3] The meaning of
Darwinism has changed over time, and varies depending on who is
using the term.[4] In the United States, Darwinism is often used by
creationists as a pejorative term but in the United Kingdom the term
has no negative connotations, being freely used as a short hand for
evolutionary theory.[5]

The term was coined by Thomas Henry Huxley in April 1860,[6] and
was used to describe evolutionary concepts, including earlier concepts
such as Malthusianism and Spencerism. In the late 19th century it
came to mean the concept that natural selection was the sole
mechanism of evolution, in contrast to Lamarckism, then around 1900
it was eclipsed by Mendelism until the modern evolutionary synthesis
Charles Darwin in 1868
unified Darwin's and Gregor Mendel's ideas. As modern evolutionary
theory has developed, the term has been associated at times with
specific ideas.[4]

While the term has remained in use amongst scientific authors, it is increasingly regarded as an inappropriate
description of modern evolutionary theory.[7] [8] [9] For example, Darwin was unfamiliar with the work of Gregor
Mendel,[10] having as a result only a vague and inaccurate understanding of heredity, and knew nothing of genetic

Conceptions of Darwinism
While the term Darwinism had been used previously to refer to the
work of Erasmus Darwin in the late 18th century, the term as
understood today was introduced when Charles Darwin's 1859 book
On the Origin of Species was reviewed by Thomas Henry Huxley in
the April 1860 issue of the Westminster Review.[13] Having hailed the
book as, "a veritable Whitworth gun in the armoury of liberalism"
promoting scientific naturalism over theology, and praising the
usefulness of Darwin's ideas while expressing professional reservations
about Darwin's gradualism and doubting if it could be proved that
natural selection could form new species,[14] Huxley compared
Darwin's achievement to that of Copernicus in explaining planetary

What if the orbit of Darwinism should be a little too circular?

What if species should offer residual phenomena, here and there,
not explicable by natural selection? Twenty years hence
As "Darwinism" became widely accepted in the
naturalists may be in a position to say whether this is, or is not, 1870s, caricatures of Charles Darwin with an ape
the case; but in either event they will owe the author of "The [12]
or monkey body symbolised evolution.
Darwinism 155

Origin of Species" an immense debt of gratitude...... And viewed as a whole, we do not believe that, since the
publication of Von Baer's "Researches on Development," thirty years ago, any work has appeared calculated to
exert so large an influence, not only on the future of Biology, but in extending the domination of Science over
regions of thought into which she has, as yet, hardly penetrated.[6]
Another important evolutionariy theorist of the same period was Peter Kropotkin who, in his book Mutual Aid: A
Factor of Evolution, advocated a conception of Darwinism counter to that of Huxley. His conception was centred
around what he saw as the widespread use of cooperation as a survival mechanism in human societies and animals.
He used biological and sociological arguments in an attempt to show that the main factor in facilitating evolution is
cooperation between individuals in free-associated societies and groups. This was in order to counteract the
conception of fierce competition as the core of evolution, which provided a rationalisation for the dominant political,
economic and social theories of the time; and the prevalent interpretations of Darwinism, such as those by Huxley,
who is targeted as an opponent by Kropotkin. Kropotkin's conception of Darwinism could be summed up by the
following quote:
In the animal world we have seen that the vast majority of species live in societies, and that they find in
association the best arms for the struggle for life: understood, of course, in its wide Darwinian sense – not as a
struggle for the sheer means of existence, but as a struggle against all natural conditions unfavourable to the
species. The animal species, in which individual struggle has been reduced to its narrowest limits, and the
practice of mutual aid has attained the greatest development, are invariably the most numerous, the most
prosperous, and the most open to further progress. The mutual protection which is obtained in this case, the
possibility of attaining old age and of accumulating experience, the higher intellectual development, and the
further growth of sociable habits, secure the maintenance of the species, its extension, and its further
progressive evolution. The unsociable species, on the contrary, are doomed to decay.
– Peter Kropotkin, Mutual Aid: A Factor of Evolution (1902), Conclusion.

19th Century usage

"Darwinism" soon came to stand for an entire range of evolutionary (and often revolutionary) philosophies about
both biology and society. One of the more prominent approaches was that summed in the phrase "survival of the
fittest" by the philosopher Herbert Spencer, which was later taken to be emblematic of Darwinism even though
Spencer's own understanding of evolution was more similar to that of Jean-Baptiste Lamarck than to that of Darwin,
and predated the publication of Darwin's theory. What is now called "Social Darwinism" was, in its day,
synonymous with "Darwinism" — the application of Darwinian principles of "struggle" to society, usually in support
of anti-philanthropic political agendas. Another interpretation, one notably favoured by Darwin's half-cousin Francis
Galton, was that Darwinism implied that because natural selection was apparently no longer working on "civilised"
people it was possible for "inferior" strains of people (who would normally be filtered out of the gene pool) to
overwhelm the "superior" strains, and voluntary corrective measures would be desirable—the foundation of

“ [Both] a Darwinian 'left' and a Darwinian 'right' were in place before most people had grasped the Darwinian middle, which was where the
maker was.

In Darwin's day there was no rigid definition of the term "Darwinism", and it was used by opponents and proponents
of Darwin's biological theory alike to mean whatever they wanted it to in a larger context. The ideas had
international influence, and Ernst Haeckel developed what was known as Darwinismus in Germany, although, like
Spencer Haeckel's "Darwinism" had only a rough resemblance to the theory of Charles Darwin, and was not centred
on natural selection at all.
Darwinism 156

While the reaction against Darwin's ideas is nowadays often thought to have been widespread immediately, in 1886
Alfred Russel Wallace went on a lecture tour across the United States, starting in New York and going via Boston,
Washington, Kansas, Iowa and Nebraska to California, lecturing on what he called Darwinism without any

Other uses
The term Darwinism is often used in the United States by promoters of creationism, notably by leading members of
the intelligent design movement, as an epithet to attack evolution as though it were an ideology (an "ism") of
philosophical naturalism, or atheism.[17] For example, Phillip E. Johnson makes this accusation of atheism with
reference to Charles Hodge's book What Is Darwinism?.[18] However, unlike Johnson, Hodge confined the term to
exclude those like Asa Gray who combined Christian faith with support for Darwin's natural selection theory, before
answering the question posed in the book's title by concluding: "It is Atheism."[19] [20] [21] Creationists use the term
Darwinism, often pejoratively, to imply that the theory has been held as true only by Darwin and a core group of his
followers, whom they cast as dogmatic and inflexible in their belief.[22] Casting evolution as a doctrine or belief, as
well as a pseudo-religious ideology like Marxism,[23] bolsters religiously motivated political arguments to mandate
equal time for the teaching of creationism in public schools.
However, Darwinism is also used neutrally within the scientific community to distinguish modern evolutionary
theories from those first proposed by Darwin, as well as by historians to differentiate it from other evolutionary
theories from around the same period. For example, Darwinism may be used to refer to Darwin's proposed
mechanism of natural selection, in comparison to more recent mechanisms such as genetic drift and gene flow. It
may also refer specifically to the role of Charles Darwin as opposed to others in the history of evolutionary thought
— particularly contrasting Darwin's results with those of earlier theories such as Lamarckism or later ones such as
the modern synthesis.
In the United Kingdom the term retains its positive sense as a reference to natural selection, and for example Richard
Dawkins wrote in his collection of essays A Devil's Chaplain, published in 2003, that as a scientist he is a

See also
• Darwinism (book)
• Modern evolutionary synthesis
• Neo-Darwinism
• Neural Darwinism
• Social Darwinism
• Darwin Awards
• Pangenesis - Charles Darwin's hypothetical mechanism for heredity
• Universal Darwinism
Darwinism 157

[1] John Wilkins (1998). "How to be Anti-Darwinian" (http:/ / www. talkorigins. org/ faqs/ anti-darwin. html). TalkOrigins Archive. . Retrieved
19 June 2008.
[2] "Expelled Exposed: Why Expelled Flunks » …on what evolution explains" (http:/ / www. expelledexposed. com/ index. php/ contest/
on-what-evolution-explains). National Center for Science Education. . Retrieved 22 December 2008.
[3] based on an European Southern Observatory release (9 December 2006). "Galactic Darwinism :: Astrobiology Magazine - earth science -
evolution distribution Origin of life universe - life beyond :: Astrobiology is study of earth science evolution distribution Origin of life in
universe terrestrial" (http:/ / www. astrobio. net/ news/ index. php?name=News& file=article& sid=2169& theme=Printer). . Retrieved 22
December 2008.
[4] Joel Hanes. "What is Darwinism?" (http:/ / www. talkorigins. org/ faqs/ darwinism. html). TalkOrigins Archive. . Retrieved 19 June 2008.
[5] Scott, Eugenie C.; Branch, Glenn (16 January 2009). "Don’t Call it “Darwinism”" (http:/ / www. springerlink. com/ content/
n47h34357743w4p0/ ?p=e3b030036a4d442a8ce393291fe0688f& pi=9). Evolution: Education and Outreach (New York: Springer) 2 (1): 90.
doi:10.1007/s12052-008-0111-2. ISSN 1936-6434. . Retrieved 17 November 2009.
[6] Huxley, T.H. (April 1860). "ART. VIII.- Darwin on the origin of Species" (http:/ / darwin-online. org. uk/ content/ frameset?viewtype=side&
itemID=A32& pageseq=29). Westminster Review. pp. 541–70. . Retrieved 19 June 2008. "What if the orbit of Darwinism should be a little
too circular?"
[7] John Wilkins (1998). "How to be Anti-Darwinian" (http:/ / www. talkorigins. org/ faqs/ anti-darwin. html). TalkOrigins Archive. . Retrieved
27 June 2008.
[8] Ruse, Michael (2003). Darwin and Design: Does Evolution Have a Purpose? (http:/ / books. google. com/ ?id=SHWaeRiRD-cC&
printsec=frontcover& dq="michael+ ruse"+ darwinism). Cambridge, MA: Harvard University Press. pp. 293. ISBN 0674016319. . Retrieved
18 July 2008.
[9] Olivia Judson (15 July 2008). "Let’s Get Rid of Darwinism" (http:/ / judson. blogs. nytimes. com/ 2008/ 07/ 15/ lets-get-rid-of-darwinism/ ).
New York Times. .
[10] Sclater, Andrew (June 2006). "The extent of Charles Darwin’s knowledge of Mendel" (http:/ / www. springerlink. com/ content/
w112307246x77t37/ ). Journal of Biosciences (Bangalore, India: Springer India / Indian Academy of Sciences) 31 (2): 191–193.
doi:10.1007/BF02703910. PMID 16809850. . Retrieved 3 January 2009.
[11] Laurence Moran (1993). "Random Genetic Drift" (http:/ / www. talkorigins. org/ faqs/ genetic-drift. html). TalkOrigins Archive. . Retrieved
27 June 2008.
[12] Browne 2002, pp. 376–379
[13] "The Huxley File § 4 Darwin's Bulldog" (http:/ / aleph0. clarku. edu/ huxley/ guide4. html). . Retrieved 29 June 2008.
[14] Browne 2002, pp. 105–106
[15] Gopnik 2009, p. 152.
[16] "Evolution and Wonder - Understanding Charles Darwin - Speaking of Faith from American Public Media" (http:/ / speakingoffaith.
publicradio. org/ programs/ darwin/ transcript. shtml). . Retrieved 27 July 2007.
[17] Scott, Eugenie C. (2008). "Creation Science Lite: "Intelligent Design" as the New Anti-Evolutionism" (http:/ / biology. ucf. edu/ ~clp/
Courses/ seminar/ papers/ 07-Scott-scientists_confront-cs_lite. pdf). In Godfrey, Laurie R.; Petto, Andrew J.. Scientists Confront Creationism:
Intelligent Design and Beyond. New York: W. W. Norton. pp. 72. ISBN 0-393-33073-7.
[18] Johnson, Phillip E.. "What is Darwinism?" (http:/ / www. arn. org/ docs/ johnson/ wid. htm). . Retrieved 4 January 2007.
[19] Matthew, Ropp. "Charles Hodge and His Objection to Darwinism" (http:/ / www. theropps. com/ papers/ Winter1997/ CharlesHodge. htm). .
Retrieved 4 January 2007.
[20] Hodge, Charles. "What is Darwinism?" (http:/ / www. gutenberg. org/ files/ 19192/ 19192-8. txt). . Retrieved 4 January 2007.
[21] Hodge, Charles (1874). What is Darwinism?. Scribner, Armstrong, and Company. OCLC 11489956.
[22] Sullivan, M (2005). "From the Beagle to the School Board: God Goes Back to School" (http:/ / www. impactpress. com/ articles/ spring05/
sullivanspring05. html). Impact Press. . Retrieved 18 September 2008.
[23] "Darwinism should be allowed to collapse and end up on the ash heap of history" (http:/ / www. docstoc. com/ docs/ 20835072/
Darwinism-should-be-allowed-to-collapse-and-end-up-on-the-ash-heap-of-history). .
[24] Sheahen, Laura. Religion: For Dummies (http:/ / www. beliefnet. com/ story/ 136/ story_13688_1. html)., interview about
2003 book.
Darwinism 158

• Browne, E. Janet (2002). Charles Darwin: Vol. 2 The Power of Place. London: Jonathan Cape.
ISBN 0712668373.
• Gopnik, Adam (2009). Angels and Ages: A Short Book About Darwin, Lincoln, and Modern Life. London:
Quercus. ISBN 9781847249296.

External links
• Universal Darwinism (
• (Russian) Nikolai Danilevsky. 1885-1889 Darwinism. A Critical Study (
(Дарвинизм. Критическое исследование) at in DjVu format
• Stanford Encyclopedia of Philosophy entry (
• What is Darwinism (
• The Darwinian Revolution (

The Genetical Theory of Natural Selection

The Genetical Theory of Natural Selection is a book by R.A. Fisher first published in 1930 by Clarendon. It is one
of the most important books of the modern evolutionary synthesis[1] and is commonly cited in biology books.

A second, slightly revised edition was republished in 1958. In 1999, a third variorum edition (ISBN 0-19-850440-3),
with the original 1930 text, annotated with the 1958 alterations, notes and alterations accidentally omitted from the
second edition was published, edited by Henry Bennett.

It contains the following chapters:
1. The Nature of Inheritance [2]
2. The Fundamental Theorem of Natural Selection
3. The Evolution of Dominance
4. Variation as determined by Mutation and Selection
5. Variation etc
6. Sexual Reproduction and Sexual Selection [3]
7. Mimicry
8. Man and Society
9. The Inheritance of Human Fertility
10. Reproduction in Relation to Social Class
11. Social Selection of Fertility
12. Conditions of Permanent Civilization
''The Genetical Theory of Natural Selection'' 159

In the preface, Fisher considers some general points, including that there must be an understanding of natural
selection distinct from that of evolution, and that the then-recent advances in the field of genetics (see history of
genetics) now allowed this. In the first chapter, Fisher considers the nature of inheritance, rejecting blending
inheritance in favour of particulate inheritance. The second chapter introduces Fisher's fundamental theorem of
natural selection. The third considers the evolution of dominance, which Fisher believed was strongly influenced by
modifiers. The last five chapters (8-12) include Fisher's more idiosyncratic views on eugenics.

The book is dedicated to Major Leonard Darwin, Fisher's friend, correspondent and son of Charles Darwin, "In
gratitude for the encouragement, given to the author, during the last fifteen years, by discussing many of the
problems dealt with in this book".

Henry Bennett gave an account of the writing and reception of Fisher's Genetical Theory.[4]
Sewall Wright, who had many disagreements with Fisher, reviewed the book and wrote that it was "certain to take
rank as one of the major contributions to the theory of evolution".[5] J.B.S. Haldane described it as "brilliant".[6]
Reginald Punnett was negative, however.[7] '
The Genetical Theory was largely overlooked for 40 years, and in particular the fundamental theorem was
misunderstood. The work had a great effect on W.D. Hamilton, who discovered it as an undergraduate at Oxford[8]
and noted on the rear cover of the 1999 variorum edition:
This is a book which, as a student, I weighed as of equal importance to the entire rest of my undergraduate
Cambridge BA course and, through the time I spent on it, I think it notched down my degree. Most chapters
took me weeks, some months.
...And little modified even by molecular genetics, Fisher's logic and ideas still underpin most of the ever
broadening paths by which Darwinism continues its invasion of human thought.
Unlike in 1958, natural selection has become part of the syllabus of our intellectual life and the topic is
certainly included in every decent course in biology.
For a book that I rate only second in importance in evolution theory to Darwin's Origin (this as joined with its
supplement Of Man), and also rate as undoubtedly one of the greatest books of the twentieth century the
appearance of a variorum edition is a major event...
By the time of my ultimate graduation, will I have understood all that is true in this book and will I get a First?
I doubt it. In some ways some of us have overtaken Fisher; in many, however, this brilliant, daring man is still
far in front.
The publication of the variorum edition in 1999 led to renewed interest in the work and reviews by Laurence Cook
("This is perhaps the most important book on evolutionary genetics ever written"),[9] Brian Charlesworth,[10] Jim
Crow[11] and A.W.F. Edwards[12]
''The Genetical Theory of Natural Selection'' 160

[1] 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.
[2] http:/ / www. blackwellpublishing. com/ ridley/ classictexts/ fisher1. pdf
[3] http:/ / www. blackwellpublishing. com/ ridley/ classictexts/ fisher2. pdf
[4] http:/ / digital. library. adelaide. edu. au/ coll/ special/ fisher/ natsel/ tp_intro. pdf
[5] Wright, S., 1930 The Genetical Theory of Natural Selection: a review (http:/ / jhered. oxfordjournals. org/ cgi/ reprint/ 21/ 8/ 349). J. Hered.
[6] Haldane, J.B.S., 1932 The Causes of Evolution. Longman Green, London.
[7] Punnett, R.C. 1930, A review of The Genetical Theory of Natural Selection, Nature 126: 595-7
[8] Grafen, A. 2004. 'William Donald Hamilton’ (http:/ / users. ox. ac. uk/ ~grafen/ cv/ WDH_memoir. pdf). Biographical Memoirs of Fellows of
the Royal Society, 50, 109-132
[9] Cook, L. 2000 Book reviews. The Genetical Theory of Natural Selection — A Complete Variorum Edition. R. A. Fisher (edited by Henry
Bennett) (http:/ / www. nature. com/ hdy/ journal/ v84/ n3/ full/ 6887132a. html). Heredity 84 (3) , 390–39
[10] Charlesworth, B. 2000 The Genetical Theory of Natural Selection. A Complete Variorum Edition. By R. A. Fisher (edited with foreword
and notes by J. H. Bennett). Oxford University Press. 1999. ISBN 0-19-850440-3. xxi+318 pages. (http:/ / journals. cambridge. org/ action/
displayAbstract?fromPage=online& aid=52675) Genetics Research 75: 369-373
[11] Crow, J.F. 2000 Second only to Darwin - The Genetical Theory of Natural Selection. A Complete Variorum Edition by R.A. Fisher (http:/ /
www. sciencedirect. com/ science?_ob=ArticleURL& _udi=B6VJ1-405BR68-M& _user=10& _rdoc=1& _fmt=& _orig=search& _sort=d&
view=c& _acct=C000050221& _version=1& _urlVersion=0& _userid=10& md5=3a8c3a64ce75187ab109f4b0b9d18ba9) Trends in Ecology
and Evolution, Volume 15, Number 5, 1 May 2000 , pp. 213-214(2)
[12] Edwards, A.W.F. 2000 The Genetical Theory of Natural Selection (http:/ / www. genetics. org/ cgi/ content/ full/ 154/ 4/ 1419) Genetics,
Vol. 154, 1419-1426, April 2000

External links
• Full text of 1930 edition (, Open Library

Neo-Darwinism is a term used to describe the 'modern synthesis' of Darwinian evolution through natural selection
with Mendelian genetics, the latter being a set of primary tenets specifying that evolution involves the transmission
of characteristics from parent to child through the mechanism of genetic transfer, rather than the 'blending process' of
pre-Mendelian evolutionary science. Neo-Darwinism also separates Darwin's ideas of natural selection from his
hypothesis of Pangenesis as a Lamarckian source of variation involving blending inheritance.[1]
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.[2] Weismann and Wallace rejected the
Lamarckian idea of inheritance of acquired characteristics, something that Darwin had not ruled out.[3] The term was
first used in 1895 to explain that evolution occurs solely through natural selection, in other words, without any
mechanism involving the inheritance of acquired characteristics resulting from use or disuse.[4] These two scientists'
complete rejection of Lamarckism came from Weismann's germ plasm theory. Weismann realised that the cells that
produce the germ plasm, or gametes (such as sperm and egg in animals), separate from the somatic cells that go on to
make other body tissues at an early stage in development. Since he could see no obvious means of communication
between the two he asserted that the inheritance of acquired characteristics was therefore impossible; a conclusion
now known as Weismann's barrier.[5]
From the 1880s to the 1930s the term continued to be applied to the panselectionist school of thought, which argued
that natural selection was the main and perhaps sole cause of all evolution.[6] From then until around 1947 the term
was used for the panselectionist followers of R. A. Fisher.
Neo-Darwinism 161

Modern evolutionary synthesis

Following the development, from about 1937 to 1950, of the modern evolutionary synthesis, now generally referred
to as the synthetic view of evolution or the modern synthesis, the term neo-Darwinian is often used to refer to
contemporary evolutionary theory.[7] However, such usage has been described by some as incorrect;[8] [1] [4] with
Ernst Mayr writing in 1984:
"...the term neo-Darwinism for the synthetic theory is wrong, because the term neo-Darwinism was coined by
Romanes in 1895 as a designation of Weismann's theory."'[9]
Despite this, publications such as the Encyclopaedia Britannica,[10] [11] use this term to refer to current evolutionary
theory. This term is also used in the scientific literature, with the academic publishers Blackwell Publishing referring
to "neo-Darwinism as practised today",[12] and some figures in the study of evolution like Richard Dawkins[13] and
Stephen Jay Gould,[14] using the term in their writings and lectures.

See also
• Modern evolutionary synthesis
• The Origin of Species
• Developmental systems theory
• Evolutionary developmental biology
• Neo-Lamarckism

[1] Kutschera U, Niklas KJ (2004). "The modern theory of biological evolution: an expanded synthesis". Naturwissenschaften 91 (6): 255–76.
doi:10.1007/s00114-004-0515-y. PMID 15241603.
[2] Gould The Structure of Evolutionary Theory p. 216
[3] 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 (http:/ / www. springerlink. com/ content/ f6131358k265g3u4/ )
[4] Reif W-E. Junker T. Hoßfeld U. (2000). "The synthetic theory of evolution: general problems and the German contribution to the synthesis".
Theory in Biosciences 119 (1): 41–91(51). doi:10.1078/1431-7613-00004.
[5] Barbieri FD (1989). "The origin of Metazoa and Weismann's germ line theory". Riv. Biol. 82 (1): 61–74. PMID 2665023.
[6] "How to be Anti-Darwinian" (http:/ / www. talkorigins. org/ faqs/ anti-darwin. html). . Retrieved 2007-09-19.
[7] "The Modern Synthesis of Genetics and Evolution" (http:/ / www. talkorigins. org/ faqs/ modern-synthesis. html). . Retrieved 2007-09-19.
[8] Pigliucci, M. (2007). "Do We Need An Extended Evolutionary Synthesis?" (http:/ / www. bioone. org/ perlserv/ ?request=get-abstract).
Evolution 61 (12): 2743–2749. doi:10.1111/j.1558-5646.2007.00246.x. PMID 17924956. .
[9] Mayr E. (1984). "What is Darwinism Today?" (http:/ / links. jstor. org/ sici?sici=0270-8647(1984)1984<145:WIDT>2. 0. CO;2-5).
Proceedings of the Biennial Meeting of the Philosophy of Science Association 2: 145–156. .
[10] "neo-Darwinism" (http:/ / www. britannica. com/ eb/ article-9373225/ neo-Darwinism). . Retrieved 2007-09-19.
[11] "neo-Darwinism" (http:/ / encyclopedia. farlex. com/ neo-Darwinism). Hutchinson Encyclopedia. . Retrieved 2007-09-19.
[12] "A-Z Browser of Evolution" (http:/ / www. blackwellpublishing. com/ ridley/ a-z/ Neo-Darwinism. asp). Blackwell Publishing. . Retrieved
[13] "Lecture on Neo-Darwinism" (http:/ / richarddawkins. net/ article,1345,Lecture-on-Neo-Darwinism,Richard-Dawkins).
: The Official Richard Dawkins Website. . Retrieved 2007-09-19.
[14] "Challenges to Neo-Darwinism and Their Meaning for a Revised View of Human Consciousness" (http:/ / www. tannerlectures. utah. edu/
lectures/ documents/ gould85. pdf). Cambridge University: The Tanner Lectures on Human Values. . Retrieved 2009-03-05.
Neo-Darwinism 162

External links
• The Variation of Animals and Plants Under Domestication (
• Challenges to neo-Darwinism - Stephen Jay Gould ( at the Wayback Machine (archived September 24, 2006).
• Neodarwinism - Richard Dawkins (

Modern evolutionary synthesis

The modern evolutionary synthesis is also referred to as the new synthesis, the modern synthesis, the
evolutionary synthesis and the neo-darwinian synthesis. It is a union of ideas from several biological specialties
which provides a widely accepted account of evolution. 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]
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.
The modern synthesis solved difficulties and confusions caused by the specialisation and poor communication
between biologists in the early years of the 20th century. Discoveries of early geneticists were difficult to reconcile
with gradual evolution and the mechanism of natural selection. The synthesis reconciled the two schools of thought,
while providing evidence that studies of populations in the field were crucial to evolutionary theory. It drew together
ideas from several branches of biology that had become separated, particularly genetics, cytology, systematics,
botany, morphology, ecology and paleontology.

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
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.
Modern evolutionary synthesis 163

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]

Developments leading up to the synthesis

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]

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]
Modern evolutionary synthesis 164

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.[] 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.[] 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.[] [19] [20]

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.[] [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
Modern evolutionary synthesis 165

that were reproductively isolated from all other populations.[] [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.[] [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.[]

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).
Modern evolutionary synthesis 166

After the synthesis

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.

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
The structure of evolutionary biology.
period of time.[29]
The history and causes of evolution (center) are subject to various
• The triumph of Alfred Wegener's idea of continental subdisciplines of evolutionary biology. The areas of segments give
drift came around 1960. The key principle of plate an impression of the contributions of subdisciplines to the literature
of evolutionary biology.
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
• 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]
Modern evolutionary synthesis 167

Symbiotic origin of eukaryotic cell structures

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] a book was published in the U.S.A. in 1927 with the title 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]

Trees of life
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 kingdoms 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
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.

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
Modern evolutionary synthesis 168

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.

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]
A shaft of light has been thrown on the evolution of flatfish (pleuronectiformes), such as plaice, sole, turbot and
halibut, by recent work. 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
Modern evolutionary synthesis 169

of flatfish falls squarely within the evolutionary synthesis.[77]

See also
• Developmental systems theory
• Evolution
• Gene-centered view of evolution
• History of evolutionary thought
• Particulate inheritance theory
• Polymorphism (biology)
• Population genetics
• The Origin of Species

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[2] Mayr, Ernst 2002. What evolution is. Weidenfeld & Nicolson, London. p270
[3] Huxley J.S. 1942. Evolution: the modern synthesis. Allen & Unwin, London. 2nd ed 1963; 3rd ed 1974.
[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] 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
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[11] Bowler pp. 253–256
[12] Mike Ambrose. "Mendel's Peas" (http:/ / www. jic. ac. uk/ germplas/ pisum/ zgs4f. htm). Genetic Resources Unit, John Innes Centre,
Norwich, UK. . 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] Larson Evolution: The Remarkable History of a Scientific Theory pp. 221–243
[19] 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.
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[26] Mayr and Provine 1998 pp. 298–299, 416
[27] Bowler p.361
[28] Pigliucci, Massimo 2007. Do we need an extended evolutionary synthesis? (http:/ / www. blackwell-synergy. com/ doi/ abs/ 10. 1111/ j.
1558-5646. 2007. 00246. x?cookieSet=1& journalCode=evo) 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
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[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.
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[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
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[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.
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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] Walin I.E. 1927. Symbionticism and the origin of species. Williams & Wilkins, Baltimore.
[46] Wells H.G., Huxley J. and Wells G.P. 1930. The science of life. London vol 2, p505. This section (The ABC of genetics) was written by
[47] Sapp, January 1994. Evolution by association: a history of symbiosis. Oxford.
[48] Lederberg J. 1952. Cell genetics and hereditary symbiosis. Physiological Reviews 32, 403–430.
[49] Margulis L and Fester R (eds) 1991. Symbiosis as a source of evolutionary innovation. MIT.
[50] Margulis L. 1993. Symbiosis in cell evolution: microbial communities in the Archaean and Proterozoic eras. Freeman, N.Y.
[51] Maynard Smith J. and Szathmáry E. 1997. The major transitions in evolution. Oxford. The origin of the eukaryote cell is one of the seven
major transitions, according to these authors.
[52] Woese, Carl 1998. The Universal Ancestor. PNAS 95, 6854–6859.
[53] Doolittle, W. Ford 1999. Phylogenetic classification and the Universal Tree. Science 284, 2124–2128.
[54] Doolittle, W. Ford 2000. Uprooting the tree of life. Scientific American 282 (6): 90–95.
[55] Dunn, Casey W. et al 2009. Broad phylogenetic sampling improves resolution of the animal tree of life. Nature 452, 745–749.
[56] Laubichler M. and Maienschein J. 2007. From Embryology to Evo-Devo: a history of developmental evolution. MIT.
[57] de Beer, Gavin 1930. Embryology and evolution. Oxford; 2nd ed 1940 as Embryos and ancestors; 3rd ed 1958, same title.
[58] Bolk, L. 1926. Der Problem der Menschwerdung. Fischer, Jena.
[59] short-list of 25 characters reprinted in Gould, Stephen Jay 1977. Ontogeny and phylogeny. Harvard. p357
[60] Raff R.A. and Kaufman C. 1983. Embryos, genes and evolution: the developmental-genetic basis of evolutionary changes. Macmillan, N.Y.
[61] Carroll, Sean B. 2005. Endless forms most beautiful: the new science of Evo-Devo and the making of the animal kingdom. Norton, N.Y.
[62] Lewis E.B. 1995. The bithorax complex: the first fifty years. Nobel Prize lecture. Repr. in Ringertz N. (ed) 1997. Nobel lectures, Physiology
or Medicine. World Scientific, Singapore.
[63] Lawrence P. 1992. The making of a fly. Blackwell, Oxford.
[64] Duncan I. 1987. The bithorax complex. Ann. Rev. Genetics 21, 285–319.
[65] Lewis E.B. 1992. Clusters of master control genes regulate the development of higher organisms. J. Am. Medical Assoc. 267, 1524–1531.
[66] McGinnis W. et al 1984. A conserved DNA sequence in homeotic genes of the Drosophila antennipedia and bithorax complexes. Nature
308, 428–433.
[67] Scott M.P. and Weiner A.J. 1984. Structural relationships among genes that control developmental sequence homology between the
antennipedia, ultrabithorax and fushi tarazu loci of Drosophila. PNAS USA 81, 4115.
[68] Gehring W. 1999. Master control systems in development and evolution: the homeobox story. Yale.
[69] Shubin N, Tabin C and Carroll S. 1997. Fossils, genes and the evolution of animal limbs. Nature 388, 639–648.
[70] Shubin N, Tabin C and Carroll S. 2009. Deep homology and the origins of evolutionary novelty. Nature 457, p818–823.
[71] Sarich V. 1976. The panda is a bear. Nature 245, 218–220.
[72] Davies D.D. 1964. The giant panda: a morphological study of evolutionary mechanisms. Fieldiana Memoires (Zoology) 3, 1–339.
[73] Stanley Steven M. 1979. Macroevolution: pattern & process. Freeman, San Francisco. p157
[74] Clack, Jenny A. 2002. Gaining Ground: the origin and evolution of tetrapods. Bloomington, Indiana. ISBN 0-253-34054-3
[75] ||cite web |url=[ |title=Jenny Clack homepage}}
[76] Both whale evolution and early insect flight are discussed in Raff R.A. 1996. The shape of life. Chicago. These discussions provide a
welcome synthesis of evo-devo and paleontology.
[77] Janvier, Philip 2008. Squint of the fossil flatfish. Nature 454, 169
[78] Lamark J.B. 1809. Philosophie zoologique. Paris.
[79] Mivart St G. 1871. The genesis of species. Macmillan, London.
[80] Darwin, Charles 1872. The origin of species. 6th ed, Murray, London. p186–188. The whole of Chapter 7 in this edition is taken up with
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Modern evolutionary synthesis 171

[81] Goldschmidt R. Some aspects of evolution. Science 78, 539–547.

[82] Goldschmidt R. 1940. The material basis of evolution. Yale.
[83] Friedman, Matt 2008. The evolutionary origin of flatfish asymmetry. Nature 454, 209–212.

• Allen, Garland. Thomas Hunt Morgan: The Man and His Science, Princeton University Press, 1978 ISBN
• 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
• 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 University Press.
ISBN 0-674-00613-5.
• 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, J. S. Evolution: The Modern Synthesis, Allen and Unwin, 1942 ISBN 0-02-846800-7
• 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, 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
• Smocovitis, V. Betty. Unifying Biology: The Evolutionary Synthesis and Evolutionary Biology, Princeton
University Press, 1996 ISBN 0-691-27226-9
• Wright, S. 1931. "Evolution in Mendelian populations". Genetics 16: 97–159.

External links
• Rose MR, Oakley TH, The new biology: beyond the Modern Synthesis (
content/pdf/1745-6150-2-30.pdf). Biology Direct 2007, 2:30. A review of biology in light of recent innovations
since the initiation of modern synthesis.

Base concepts

Heredity is the passing of traits to offspring (from its parent or ancestors). This is the process by which an offspring
cell or organism acquires or becomes predisposed to the characteristics of its parent cell or organism. Through
heredity, variations exhibited by individuals can accumulate and cause a species to evolve. The study of heredity in
biology is called genetics, which includes the field of epigenetics.

The ancients had a variety of ideas about heredity: Theophrastus proposed that male flowers caused female flowers
to ripen; Hippocrates speculated that "seeds" were produced by various body parts and transmitted to offspring at the
time of conception, and Aristotle thought that male and female semen mixed at conception. Aeschylus, in 458 BC,
proposed the male as the parent, with the female as a "nurse for the young life sown within her."
Various hereditary mechanisms were envisaged without being properly tested or quantified. These included blending
inheritance and the inheritance of acquired traits. Nevertheless, people were able to develop domestic breeds of
animals as well as crops through artificial selection. The inheritance of acquired traits also formed a part of early
Lamarckian ideas on evolution.
In the 9th century AD, the Afro-Arab writer Al-Jahiz considered the effects of the environment on the likelihood of
an animal to survive, and first described the struggle for existence.[1] [2] His ideas on the struggle for existence in the
Book of Animals have been summarized as follows:
"Animals engage in a struggle for existence; for resources, to avoid being eaten and to breed. Environmental
factors influence organisms to develop new characteristics to ensure survival, thus transforming into new
species. Animals that survive to breed can pass on their successful characteristics to offspring."[3]
In 1000 AD, the Arab physician, Abu al-Qasim al-Zahrawi (known as Albucasis in the West), wrote the first clear
description of haemophilia, a hereditary genetic disorder, in his Al-Tasrif. In this work, he wrote of an Andalusian
family whose males died of bleeding after minor injuries.[4]
During the 18th century, Dutch microscopist Antonie van Leeuwenhoek (1632–1723) discovered "animalcules" in
the sperm of humans and other animals. Some scientists speculated they saw a "little man" (homunculus) inside each
sperm. These scientists formed a school of thought known as the "spermists." They contended the only contributions
of the female to the next generation were the womb in which the homunculus grew, and prenatal influences of the
womb. An opposing school of thought, the ovists, believed that the future human was in the egg, and that sperm
merely stimulated the growth of the egg. Ovists thought women carried eggs containing boy and girl children, and
that the gender of the offspring was determined well before conception.
Pangenesis was an idea that males and females formed "pangenes" in every organ. These pangenes subsequently
moved through their blood to the genitals and then to the children. The concept originated with the ancient Greeks,
and influenced biology until as recently as a century ago. The terms "blood relative," "bloodline," "full-blooded,"
and "royal blood" are relics of pangenesis. Francis Galton, Charles Darwin's cousin, experimentally tested and
disproved pangenesis during the 1870s.
Heredity 173

Types of heredity
Dominant and recessive
An allele is said to be dominant if it is always expressed in the appearance of an organism (phenotype). For example,
in peas the allele for green pods, G, is dominant to that for yellow pods, g. Since the allele for green pods is
dominant, pea plants with the pair of alleles GG (homozygote) or Gg (heterozygote) will have green pods. The allele
for yellow pods is recessive. The effects of this allele are only seen when it is present in both chromosomes, gg
The description of a mode of biological inheritance consists of three main categories:
1. Number of involved loci
• Monogenetic (also called "simple") – one locus
• Oligogenetic – few loci
• Polygenetic – many loci
2. Involved chromosomes
• Autosomal – loci are not situated on a sex chromosome
• Gonosomal – loci are situated on a sex chromosome
• X-chromosomal – loci are situated on the X chromosome (the more common case)
• Y-chromosomal – loci are situated on the Y chromosome
• Mitochondrial – loci are situated on the mitochondrial DNA
3. Correlation genotype–phenotype
• Dominant
• Intermediate (also called "codominant")
• Recessive
These three categories are part of every exact description of a mode of inheritance in the above order. Additionally,
more specifications may be added as follows:
4. Coincidental and environmental interactions
• Penetrance
• Complete
• Incomplete (percentual number)
• Expressivity
• Invariable
• Variable
• Heritability (in polygenetic and sometimes also in oligogenetic modes of inheritance)
• Maternal or paternal imprinting phenomena (also see epigenetics)
5. Sex-linked interactions
• Sex-linked inheritance (gonosomal loci)
• Sex-limited phenotype expression (e.g., cryptorchism)
• Inheritance through the maternal line (in case of mitochondrial DNA loci)
• Inheritance through the paternal line (in case of Y-chromosomal loci)
6. Locus–locus interactions
• Epistasis with other loci (e.g., overdominance)
• Gene coupling with other loci (also see crossing over)
• Homozygotous lethal factors
• Semi-lethal factors
Heredity 174

Determination and description of a mode of inheritance is primarily achieved through statistical analysis of pedigree
data. In case the involved loci are known, methods of molecular genetics can also be employed.

Charles Darwin: theory of evolution

When Charles Darwin proposed his theory of evolution in 1859, one of its major problems was the lack of an
underlying mechanism for heredity. Darwin believed in a mix of blending inheritance and the inheritance of acquired
traits (pangenesis). Blending inheritance would lead to uniformity across populations in only a few generations and
thus would remove variation from a population on which natural selection could act. This led to Darwin adopting
some Lamarckian ideas in later editions of On the Origin of Species and his later biological works. Darwin's primary
approach to heredity was to outline how it appeared to work (noticing that traits could be inherited which were not
expressed explicitly in the parent at the time of reproduction, that certain traits could be sex-linked, etc.) rather than
suggesting mechanisms.
Darwin's initial model of heredity was adopted by, and then heavily modified by, his cousin Francis Galton, who laid
the framework for the biometric school of heredity. Galton rejected the aspects of Darwin's pangenesis model which
relied on acquired traits.
The inheritance of acquired traits was shown to have little basis in the 1880s when August Weismann cut the tails off
many generations of mice and found that their offspring continued to develop tails.

Gregor Mendel: father of modern genetics

The idea of particulate inheritance of genes can be attributed to the Moravian[5] monk Gregor Mendel who published
his work on pea plants in 1865. However, his work was not widely known and was rediscovered in 1901. It was
initially assumed the Mendelian inheritance only accounted for large (qualitative) differences, such as those seen by
Mendel in his pea plants—and the idea of additive effect of (quantitative) genes was not realised until R.A. Fisher's
(1918) paper, "The Correlation Between Relatives on the Supposition of Mendelian Inheritance."

Modern development of genetics and heredity

In the 1930s, work by Fisher and others resulted in a combination of Mendelian and biometric schools into the
modern evolutionary synthesis. The modern synthesis bridged the gap between experimental geneticists and
naturalists; and between both and palaeontologists, stating that:[6] [7]
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
3. Selection is overwhelmingly the main mechanism of change; even slight advantages are important when
continued. The object of selection is the phenotype in its surrounding environment. The role of genetic drift is
equivocal; though strongly supported initially by Dobzhansky, it was downgraded later as results from ecological
genetics were obtained.
4. The primacy of population thinking: the genetic diversity carried in natural populations is a key factor in
evolution. The strength of natural selection in the wild was greater than expected; the effect of ecological factors
such as niche occupation and the significance of barriers to gene flow are all important.
5. In palaeontology, the ability to explain historical observations by extrapolation from micro to macro-evolution is
proposed. Historical contingency means explanations at different levels may exist. Gradualism does not mean
constant rate of change.
Heredity 175

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, but an account of Gavin de Beer's work by
Stephen Jay Gould suggests he may be an exception.[8]
Almost all aspects of the synthesis have been challenged at times, with varying degrees of success. There is no
doubt, however, that the synthesis was a great landmark in evolutionary biology. It cleared up many confusions, and
was directly responsible for stimulating a great deal of research in the post-World War II era.
Trofim Lysenko however caused a backlash of what is now called Lysenkoism in the Soviet Union when he
emphasised Lamarckian ideas on the inheritance of acquired traits. This movement affected agricultural research and
led to food shortages in the 1960s and seriously affected the USSR.

See also
• Genetics
• Hard inheritance
• Heritability

Notes and references

[1] Conway Zirkle (1941). Natural Selection before the "Origin of Species", Proceedings of the American Philosophical Society 84 (1), p.
[2] Mehmet Bayrakdar (Third Quarter, 1983). "Al-Jahiz And the Rise of Biological Evolutionism", The Islamic Quarterly. London. (http:/ /
www. salaam. co. uk/ knowledge/ al-jahiz. php)
[3] Gary Dargan, Intelligent Design (http:/ / www. abc. net. au/ rn/ encounter/ stories/ 2006/ 1656361. htm), Encounter, ABC.
[4] Patricia Skinner (2001), Unani-tibbi (http:/ / findarticles. com/ p/ articles/ mi_g2603/ is_0007/ ai_2603000716), Encyclopedia of Alternative
[5] Henig, Robin Marantz (2000). The Monk in the Garden : The Lost and Found Genius of Gregor Mendel, the Father of Genetics. Houghton
Mifflin. ISBN 0-395-97765-7. "The article, written by an obscure Moravian monk named Gregor Mendel"
[6] Mayr & Provine 1998
[7] Mayr E. 1982. The growth of biological thought: diversity, evolution & inheritance. Harvard, Cambs. p567 et seq.
[8] Gould S.J. Ontogeny and phylogeny. Harvard 1977. p221-2

External links
• Stanford Encyclopedia of Philosophy entry on Heredity and Heritability (
Fitness 176

Fitness (often denoted in population genetics models) is a central idea in evolutionary theory. It can be defined
either with respect to a genotype or to a phenotype. In either case, it is equal to the average contribution to the gene
pool of the next generation that is made by an average individual of the specified genotype or phenotype. If
differences between alleles at a given gene affect fitness, then the frequencies of the alleles will change over
generations; the alleles with higher fitness become more common. This process is called natural selection.
An individual's fitness is manifested through its phenotype. As phenotype is affected by both genes and environment,
the fitnesses of different individuals with the same genotype are not necessarily equal, but depend on the
environment in which the individuals live. However, since the fitness of the genotype is an averaged quantity, it will
reflect the reproductive outcomes of all individuals with that genotype.
Inclusive fitness differs from individual fitness by including the ability of an allele in one individual to promote the
survival and/or reproduction of other individuals that share that allele, in preference to individuals with a different
allele. One mechanism of inclusive fitness is kin selection.

Measures of fitness
There are two commonly used measures of fitness; absolute fitness and relative fitness.

Absolute fitness
Absolute fitness ( ) of a genotype is defined as the ratio between the number of individuals with that genotype
after selection to those before selection. It is calculated for a single generation and may be calculated from absolute
numbers or from frequencies. When the fitness is larger than 1.0, the genotype increases in frequency; a ratio smaller
than 1.0 indicates a decrease in frequency.

Absolute fitness for a genotype can also be calculated as the product of the proportion survival times the average

Relative fitness
Relative fitness is quantified as the average number of surviving progeny of a particular genotype compared with
average number of surviving progeny of competing genotypes after a single generation, i.e. one genotype is
normalized at and the fitnesses of other genotypes are measured with respect to that genotype. Relative
fitness can therefore take any nonnegative value, including 0.
While researchers can usually measure relative fitness, absolute fitness is more difficult. It is often difficult to
determine how many individuals of a genotype there were immediately after reproduction.
The two concepts are related, and both of them are equivalent when they are divided by the mean fitness, which is
weighted by genotype frequencies.

Because fitness is a coefficient, and a variable may be multiplied by it several times, biologists may work with "log
fitness" (particularly so before the advent of computers). By taking the logarithm of fitness each term may be added
rather than multiplied.
Fitness 177

Maynard-Smith's Definition
As another example we may mention the definition of fitness given by Maynard Smith in the following way:
"Fitness is a property, not of an individual, but of a class of individuals – for example homozygous for allele A at a
particular locus. Thus the phrase ’expected number of offspring’ means the average number, not the number
produced by some one individual. If the first human infant with a gene for levitation were struck by lightning in its
pram, this would not prove the new genotype to have low fitness, but only that the particular child was unlucky." [1]

Hartl's Definition
Yet another possible measure has been formulated: "The fitness of the individual - having an array x of phenotypes
- is the probability, s(x), that the individual will be included among the group selected as parents of the next
generation." Then, the mean fitness may be determined as a mean over the set of individuals in a large population.

where N is the probability distribution function of phenotypes in the population, and m is its centre of gravity. This
measure is a suitable basis of a model of an evolution selecting individuals. It may in principle take even the stroke
of the lightning into consideration. In the case N is a Gaussian it is fairly easily proved that the average information
(information entropy, disorder, diversity) of a large population may be maximized by Gaussian adaptation - keeping
the mean fitness constant - in accordance with recapitulation, the central limit theorem, the Hardy-Weinberg law and
the second law of thermodynamics. This is in contrast to Fisher's fundamental theorem of natural selection.

The British sociologist Herbert Spencer coined the phrase "survival of the fittest" (though originally, and perhaps
more accurately, "survival of the best fitted") in his 1851 work Social Statics and later used it to characterise what
Charles Darwin had called natural selection.
The British biologist J.B.S. Haldane was the first to quantify fitness, in terms of the modern evolutionary synthesis
of Darwinism and Mendelian genetics starting with his 1924 paper A Mathematical Theory of Natural and Artificial
Selection. The next further advance was the introduction of the concept of inclusive fitness by the British biologist
W.D. Hamilton in 1964 in his paper on The Evolution of Social Behavior.

Fitness landscape
A fitness landscape, first conceptualized by Sewall Wright, is a way of visualising fitness in terms of a
high-dimensional surface, in which height indicates fitness, and each of the other dimensions represents allele
identity for a different gene. Peaks correspond to local fitness maxima; it is often said that natural selection always
progresses uphill but can only do so locally. This can result in suboptimal local maxima becoming stable, because
natural selection cannot return to the less-fit "valleys" of the landscape on the way to reach higher peaks.
Fitness 178

Genetic load
Genetic load measures the average fitness of a population of individuals, relative to a hypothetical population in
which the most fit genotype has become fixed.

See also
• Gene-centered view of evolution
• Inclusive fitness
• Natural selection
• Reproductive success
• Selection coefficient

[1] Maynard-Smith, J. (1989) Evolutionary Genetics ISBN 0198542151
[2] Hartl, D. L. (1981) A Primer of Population Genetics ISBN 0878932712

Further reading
• Sober, E. (2001). The Two Faces of Fitness. In R. Singh, D. Paul, C. Krimbas, and J. Beatty (Eds.), Thinking
about Evolution: Historical, Philosophical, and Political Perspectives. Cambridge University Press, pp.309-321.
Full text (
• Orr HA (August 2009). "Fitness and its role in evolutionary genetics" (
articlerender.fcgi?tool=pmcentrez&artid=2753274). Nat. Rev. Genet. 10 (8): 531–9. doi:10.1038/nrg2603.
PMID 19546856. PMC 2753274.

External links
• Evolution A-Z: Fitness (
• Stanford Encyclopedia of Philosophy entry (
Common descent 179

Common descent
In evolutionary biology, a group of organisms have common descent if they have a common ancestor. "There is
strong quantitative support, by a formal test"[1] for the theory that all living organisms on Earth are descended from a
common ancestor.[2]
Charles Darwin proposed the theory of universal common descent through an evolutionary process in On the Origin
of Species, saying, "There is a grandeur in this view of life, with its several powers, having been originally breathed
into a few forms or into one".[3]
The last universal ancestor (LUA) (or last universal common ancestor, LUCA), that is, the most recent common
ancestor of all currently living organisms,[1] is believed to have appeared about 3.9 billion years ago.[4] [5]
In The Ancestor's Tale, Richard Dawkins coined the word concestor, as a substitute for common ancestor or most
recent common ancestor. This new word is very gradually entering scientific parlance.[6]

In the 1740s, Pierre-Louis Moreau de Maupertuis made the first known suggestion in a series of essays that all
organisms may have had a common ancestor, and that they had diverged through random variation and natural
selection.[7] [8] In Essai de Cosmologie, Maupertuis noted:
Could one not say that, in the fortuitous combinations of the productions of nature, as there must be
some characterized by a certain relation of fitness which are able to subsist, it is not to be wondered at
that this fitness is present in all the species that are currently in existence? Chance, one would say,
produced an innumerable multitude of individuals; a small number found themselves constructed in such
a manner that the parts of the animal were able to satisfy its needs; in another infinitely greater number,
there was neither fitness nor order: all of these latter have perished. Animals lacking a mouth could not
live; others lacking reproductive organs could not perpetuate themselves ... The species we see today are
but the smallest part of what blind destiny has produced ...[9]
In 1790, Immanuel Kant wrote in Kritik der Urtheilskraft (Critique of Judgement) that the analogy of animal forms
implies a common original type, and thus a common parent.[10]
In 1795, Charles Darwin's grandfather, Erasmus Darwin, asked:
[W]ould it be too bold to imagine, that in the great length of time, since the earth began to exist, perhaps
millions of ages before the commencement of the history of mankind, would it be too bold to imagine,
that all warm-blooded animals have arisen from one living filament, which the great First Cause endued
with animality, with the power of acquiring new parts attended with new propensities, directed by
irritations, sensations, volitions, and associations; and thus possessing the faculty of continuing to
improve by its own inherent activity, and of delivering down those improvements by generation to its
posterity, world without end?[11]
In 1859, Charles Darwin's The Origin of Species was published. The views about common descent expressed therein
were that it was possible that there was only one progenitor for all life forms.
"Therefore I should infer from analogy that probably all the organic beings which have ever lived on this
earth have descended from some one primordial form, into which life was first breathed." [3] (p 484)
Darwin's famous closing sentence describes the "grandeur in this view of life, with its several powers, having been
originally breathed into a few forms or into one."[3] (p 490)
Common descent 180

Evidence of universal common descent

Common biochemistry and genetic code

All known forms of life are based on the same fundamental biochemical organisation: genetic information encoded
in DNA, transcribed into RNA, through the effect of protein- and RNA-enzymes, then translated into proteins by
(highly similar) ribosomes, with ATP, NADH and others as energy sources, etc. Furthermore, the genetic code (the
"translation table" according to which DNA information is translated into proteins) is nearly identical for all known
lifeforms, from bacteria to humans. The universality of this code is generally regarded by biologists as definitive
evidence in favor of the theory of universal common descent. Analysis of the small differences in the genetic code
has also provided support for universal common descent.[12] A statistical comparison of various alternative
hypotheses has shown that universal common ancestry is significantly more probable than models involving multiple

Selectively neutral similarities

Similarities which have no adaptive relevance cannot be explained by convergent evolution, and therefore they
provide compelling support for the theory of universal common descent.
Such evidence has come from two areas: amino acid sequences and DNA sequences. Proteins with the same
three-dimensional structure need not have identical amino acid sequences; any irrelevant similarity between the
sequences is evidence for common descent. In certain cases, there are several codons (DNA triplets) that code for the
same amino acid. Thus, if two species use the same codon at the same place to specify an amino acid that can be
represented by more than one codon, that is evidence for a recent common ancestor.

Other similarities
The universality of many aspects of cellular life is often pointed to as supportive evidence to the more compelling
evidence listed above. These similarities include the energy carrier adenosine triphosphate (ATP), and the fact that
all amino acids found in proteins are left-handed. It is possible that these similarities resulted because of the laws of
physics and chemistry, rather than universal common descent and therefore resulted in convergent evolution.

Phylogenetic trees
Another important piece of evidence is
that it is possible to construct detailed
phylogenetic trees (that is, "genealogic
trees" of species) mapping out the
proposed divisions and common
ancestors of all living species. In 2010
an analysis of available genetic data,
mapping them to phylogenetic trees,
gave "firm quantitative support for the
unity of life. ...there is now strong
quantitative support, by a formal
test,[1] for the unity of life.[2]

Traditionally, these trees have been

built using morphological methods, A phylogenetic tree based on rRNA genes.
such as appearance, embryology, etc.
Common descent 181

Recently, it has been possible to construct these trees using molecular data, based on similarities and differences
between genetic and protein sequences. All these methods produce essentially similar results, even though that most
genetic variation has no influence over external morphology. That phylogenetic trees based on different types of
information agree with each other is strong evidence of a real underlying common descent.[14]

Illustrations of common descent

Artificial selection
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.

Dog breeding

The diversity of domesticated dogs is an example of the power of

artificial selection. All breeds share common ancestry, having
descended from wolves. Humans selectively bred them to enhance
specific characteristics, such as color and length or body size. This
created a range of breeds that include the Chihuahua, Great Dane,
Basset Hound, Pug, and Poodle. Wild wolves, which did not undergo
artificial selection, are relatively uniform in comparison.

The Chihuahua mix and Great Dane illustrate the

range of sizes among dog breeds.

Wild cabbage

Early farmers cultivated many popular vegetables from the Brassica

oleracea (wild cabbage) by artificially selecting for certain attributes.
Common vegetables such as cabbage, kale, broccoli, cauliflower,
kohlrabi and Brussels sprouts are all descendants of the wild cabbage
plant.[15] Brussels sprouts were created by artificially selecting for
large bud size. Broccoli was bred by selecting for large flower stalks.
Cabbage was created by selecting for short petioles. Kale was bred by
selecting for large leaves.

Wild Cabbage plant

Common descent 182

Natural selection
Natural selection is the evolutionary process by which heritable traits
that increase an individual's fitness become more common, and
heritable traits that decrease an individual's fitness become less

Darwin's finches

During Charles Darwin's studies on the Galápagos Islands, Darwin

observed 13 species of finches that are closely related and differ most
markedly in the shape of their beaks. The beak of each species is suited
to the food available in its particular environment, suggesting that beak
shapes evolved by natural selection. Large beaks were found on the
Darwin's finches
islands where the primary source of food for the finches is nuts and
therefore the large beaks allowed the birds to be better equipped for
opening the nuts and staying well nourished. Slender beaks were found on the finches which found insects to be the
best source of food on the island they inhabited; their slender beaks allowed the birds to be better equipped for
pulling out the insects from their tiny hiding places. The finch is also found on the main land and it is thought that
they migrated to the islands and began adapting to their environment through natural selection.

[1] Theobald, D.L., Nature 465, 219-222, (2010)
[2] Steel, M. & Penny, D., Nature 465,168 (2010)
[3] Darwin, C., "The Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle For Life", London,
John Murrary, (1859) p. 490
[4] Doolittle, W. Ford (February, 2000). Uprooting the tree of life (http:/ / shiva. msu. montana. edu/ courses/ mb437_537_2004_fall/ docs/
uprooting. pdf). Scientific American 282 (6): 90–95.
[5] 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.
[6] Singham, Mano (October 13, 2006). "Looking for Deep Ancestors" (http:/ / machineslikeus. com/ articles/ LookingForDeep. html). Machines
Like Us. . Retrieved November 19, 2009.
[7] J. S. Bromley, The new Cambridge modern history: The rise of Great Britain and Russia, 1688-1715/25 (http:/ / books. google. com/
books?id=OOgzAAAAIAAJ& pg=PA62& dq=Maupertuis+ "for+ the+ first+ time"& cd=1#v=onepage& q=Maupertuis "for the first time"&
f=false), CUP Archive, 1970, ISBN 9780521075244, pgs. 62-63.
[8] Geoffrey Russell Richards Treasure, The making of modern Europe, 1648-1780 (http:/ / books. google. com/ books?id=C94OAAAAQAAJ&
pg=PA142& dq="blind+ destiny+ has+ produced"& cd=6#v=onepage& q="blind destiny has produced"& f=false), Taylor & Francis, 1985,
ISBN 9780416723700, pg. 142
[9] C. Leon Harris, Evolution, genesis and revelations, with readings from Empedocles to Wilson (http:/ / books. google. com/
books?id=Fkqg3TIkBJwC& pg=PA107& dq=Essai+ de+ Cosmologie+ "blind+ destiny"& cd=1#v=onepage& q=Essai de Cosmologie "blind
destiny"& f=false), SUNY Press, 1981, ISBN 9780873954877, pg. 107
[10] Immanuel Kant and Werner S. Pluhar, Critique of Judgment (http:/ / books. google. com/ books?id=y5wQD-ioaUUC& pg=PA304&
dq=Critique+ of+ Judgment+ "original+ mother"& cd=1#v=onepage& q=& f=false), Hackett Publishing, 1987, ISBN 9780872200258, p. 304:
"Despite all the variety among these forms, they seem to have been produced according to a common archetype, and this analogy among them
reinforces our suspicion that they are actually akin, produced by a common original mother."
[11] Darwin, Erasmus (1818) [1795]. "Generation" (http:/ / books. google. com/ ?id=XRUaAAAAMAAJ& printsec=frontcover& q=).
Zoonomia; or the Laws of Organic Life. 1 (4th American ed.). Philadelphia: Edward Earle. p. 397 [§ 39.4.8]. . Retrieved November 20, 2009.
[12] Knight, Robin, et al. (January 2001). "Rewiring the Keyboard: Evolvability of the Genetic Code". Nature Reviews Genetics 2 (1): 49–58.
doi:10.1038/35047500. PMID 11253070.
[13] Theobald, Douglas L. (13 May 2010). "A formal test of the theory of universal common ancestry.". Nature 465 (7295): 219–222.
doi:10.1038/nature09014. PMID 20463738.
[14] Theobald, Douglas (2004). "Prediction 1.3: Consilience of Independent Phylogenies" (http:/ / www. talkorigins. org/ faqs/ comdesc/
section1. html#independent_convergence). 29+ Evidences for Macroevolution. TalkOrigins Foundation. . Retrieved November 20, 2009.
Common descent 183

[15] Raven, Peter H., et al. (2005). Biology of Plants (7th rev. ed.). New York: W.H. Freeman. ISBN 0716762846. OCLC 183148564. "[These
vegetables were] all produced from a single species of plant (Brassica oleraca), a member of the mustard family."

External links
• 29+ Evidences for Macroevolution: the Scientific Case for Common Descent (
comdesc/). From the TalkOrigins Archive
• The Tree of Life Web Project (

Evidence of common descent

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
Evidence of common descent 184

common descent extends from direct laboratory and 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.

Evidence from comparative physiology and biochemistry

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
While on board HMS Beagle, Charles Darwin collected numerous
species will have a greater fraction of identical specimens, many new to science, which supported his later theory of
sequence and will have shared substitutions when evolution by natural selection.
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 (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.

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. For example, although camera-like eyes are believed to have evolved independently on many
Evidence of common descent 185

separate occasions, 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.

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.

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. 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.[6]

Pseudogenes also known, as Junk DNA, or dead DNA is the extra DNA that is in our genome but does 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 our genome is these (1% working DNA).[7] The way a pseudogene is produced is a gene
that is no longer used will accumulate mutations that make it non-functional but since it is not used; when it
disappears, it does not affect fitness.

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,
Evidence of common descent 186

which subsequently behaves as an endosymbiont, nourishing Hatena, which in turn loses its feeding apparatus and
behaves as an autotroph.[8] [9]
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.

Specific examples

Chromosome 2 in humans

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.[10] [11]

The evidence for this includes:

• The correspondence of chromosome 2 to two ape
chromosomes. The closest human relative, the common Fusion of ancestral chromosomes left distinctive remnants of
telomeres, and a vestigial centromere
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.[12] [13]
• The presence of a vestigial centromere. Normally a chromosome has just one centromere, but in chromosome 2
there are remnants of a second centromere.[14]
• 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.[15]
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."[15]

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.[16]
Evidence of common descent 187

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:[17] the 66th amino acid is isoleucine in the former and threonine in the latter.[16]
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.[18]

Out of Africa hypothesis of human evolution

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.[19]

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.

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.[20] 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.[20] [21] Darwin described how common descent could provide a logical basis for

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

Evidence of common descent 188

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.

Vestigial structures
A strong and direct evidence for common descent comes from vestigial structures.[23] 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

Evolutionary developmental biology and embryonic development

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 stems 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 such as hair growth and loss (lanugo) during human development;[24] 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;[25] 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.
Evidence of common descent 189

An atavism is an evolutionary throwback, such as traits reappearing which had disappeared generations ago.[26]
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.[27] Some examples of this are
hind-legged snakes or whales;[26] [28] [29] the extra toes of ungulates that do not even reach the ground,[30] chicken's
teeth,[31] reemergence of sexual reproduction in Hieracium pilosella and Crotoniidae;[32] and humans with tails,[26]
extra nipples,[28] and large canine teeth.[28]

Specific examples

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.
Evidence of common descent 190

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.

Pentadactyl limb
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
• 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.
Evidence of common descent 191

Pelvic structure of dinosaurs

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.

Insect mouthparts
The basic structures are the same, including a labrum (upper lip), a pair of mandibles, a hypopharynx (floor of
mouth), a pair of maxillae, and a labium. These structures are enlarged and modified; others are reduced and lost.
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.

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.

Recurrent laryngeal nerve in giraffes

The recurrent laryngeal nerve is a

fourth branch of the vagus nerve,
which is a cranial nerve. In mammals
its path is extraordinary: 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 looks not too optimal even

for human, but for giraffes it becomes
quite suboptimal. Due to the lengths of
their necks, the recurrent laryngeal
The path of the recurrent laryngeal nerve in giraffes. The laryngeal nerve is compensated nerve may be up to 4m long (13 ft),
for by subsequent tinkering from natural selection. despite its optimal route being a
distance of just several inches.
Its route is explained by the evolution of mammals from fishes, which had no neck. Therefore the nerve's route was
quite reasonable: it was a relatively short nerve that innervated one gill slit, and passed near the gill arch. Since then
gills have evolved and the gill arch has become a dorsal aorta.[33] [34]
Evidence of common descent 192

Route of the vas deferens

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.[34] [35]

Route of the vas deferens from the testis to the penis

Evidence from paleontology

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.
An insect trapped in amber.
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
Evidence of common descent 193

fossils of which are made in layers that then harden.

Fossil record
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
Fossil trilobite. Trilobites were hard-shelled
arthropods, related to living horseshoe crabs and
structure, whereas younger rocks contain a greater variety of fossils,
spiders, that first appeared in significant often with increasingly complex structures.
numbers around 540 mya, dying out 250 mya.
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.

Extent of the fossil record

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.[36]
Evidence of common descent 194

Despite the relative rarity of suitable conditions for fossilization,

approximately 250,000 fossil species are known.[37] 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.[38]
Many more fossils are still in the ground, in various geological Cynognathus, a Eucynodont, one of a grouping of
formations known to contain a high fossil density, allowing Therapsids ("mammal-like reptiles") that is ancestral to
estimates of the total fossil content of the formation to be made. all modern mammals.

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).[39] It has been estimated that this formation contains 800 billion
vertebrate fossils.[40]

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. F