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Geological Time Scale

I.

Pre- Cambrian

a. Hadean (4,600- 4,000 mya)

Defining Characteristics:
o formation of Earths crust and main bombardment
Secondary Characteristics:
o continuing erosion and plate tectonics have destroyed Hadean rocks

The name Hadean Eon comes from Hades, the underworld of Greek
mythology. It refers to the hellish conditions of the Earth during the earliest part of
its history, when much of the Earths surface remained molten. The Hadean Eon of
geologic time began with the birth of the solar system, including our planet, Earth,
and ended with the formation of the oldest rocks that are still preserved on the
surface of Earth.
The Hadean is the first period in Earth history, but one for which we have
little record. The Earth began to form about 4.6 billion years ago through the
condensation of material around the sun. As this material collected, further cosmic
material was drawn to it by gravity from all directions, increasing the size of the
Earth. This process created an enormous amount of heat, which melted these
materials and eventually allowed them to separate into different layers. As the
Earth cooled, it acquired the structure we know todayan iron core, silicate mantle,
and thin outer crust.
Formation of the Earth
The Earth initially formed from the same disk of dust and gas from which the
sun itself coalesced. As the mass of the Earth increased, so did the gravitational
force it exerted, and it was bombarded by objects from space ranging in size from
dust particles to small planets (planetismals). The accretion of this material
increased the size of Earth. The impacts of large bodies and the decay of
radioactive elements generated heat that melted the materials of the young Earth,
creating the hellish conditions for which the Hadean Eon was named.
Although heat was generated through decay of radioactive elements and
continued frequent bombardment by asteroids, Earth lost heat to space and slowly
cooled. The different temperatures at which molten iron and silicate minerals
solidify indicate that as it cooled the Earth eventually segregated into an iron core
and silicate mantle. These two features are still part of the fundamental structure of
the Earth today.
Earths Atmosphere
The composition of Earths earliest atmosphere remains unclear and is a
subject of active investigation by geologists. Some geologists believe that the
earliest atmosphere included large amounts of nitrogen. Others believe that it was
composed largely of carbon dioxide and water vapor, along with some other
volcanic gases. These gases would have come from steaming vents and volcanoes
that were common across the Earths surface. Early in the Hadean, Earths
atmosphere was probably too hot to allow liquid water to condense, but fragments
contained within younger rocks suggest that sedimentary rocks formed later in the
Hadean, which implies the existence of surface water.
Earths Surface
Although continents, oceans and an atmosphere must have existed before
the Acasta Gneiss formed, the size of the continents is unclear. While Earth was still
very hot, mantle convection must have been vigorous. It is not known if plate
tectonics operated on the early Earth. One suggestion is that the earliest surface
crust was thin and unstable, and made of minerals with an extremely high content
of iron and magnesium, which are very dense. This early crust may have been

disrupted by upwelling magma at spreading centers and consumed in subduction


zones. Because of the high density of this earliest-formed crust, it would have
descended deep into the melting zone of the mantle and been destroyed. This
process would have led to recycling of the crust almost as soon as it formed. Other
geologists think that early continents formed when plumes of molten mantle rose to
the surface and cooled, but that the Earth was too hot for subduction of plates.

b. Archean (4,000- 2,500 mya)

Defining Characteristics:
o first life appears
Secondary Characteristics:
o plate tectonics established
o oxygen-poor atmosphere

The term Archean means ancient and was originally used to refer to the oldest
known rocks. Rocks of Archean age contain the earliest fossils of life on Earth. Because
these rocks were formed very long agobetween 2.5 to 4.0 billion years agomost
have long since been covered by younger sediments, eroded, or subducted into the
Earth's mantle. Nevertheless, some Archean strata survived in the central parts of
continents. These Archean shields lie in the heart of Canada, Australia, Africa, India,
and Siberia.
By the start of the Archean Eon, the Earth's crust had cooled. The atmosphere was
composed of volcanic gases, including nitrogen, hydrogen, carbon, and possibly low
levels of oxygen. Water vapor was abundant and the first oceans had formed. A
complex set of chemical reactions in these early oceans transformed carbon-containing
molecules into simple, single-celled life forms. Where these chemical reactions
occurred is unclear--hydrothermal vents are one possibility. All life today is descended
from these simple organisms. By the end of the Archean the first photosynthesizing
organisms had evolved and begun to produce oxygen, which was released into the
oceans and atmosphere. This process would dramatically change life on Earth during
the following Proterozoic Eon.
Early Continents and Oceans
Analyses of the approximately 4 billion-year-old Acasta Gneiss suggests that the
first continents and oceans developed before the Archean. The Archean, however, is
the period during which the present continents took shape. Most present continents
have shields at their cores that formed between 3 to 2.5 billion years ago during the
early Archean. Evidence of ancient oceanic crust is often found in today's greenstone
belts. Continental crust gradually grew from selective melting of dark-colored basaltic
igneous rocks within the oceanic crust. Through time, these melts became increasingly
rich in silica, as geologic processes melted more of the lower-temperature, lighterdensity minerals. Much as a helium balloon rises through air, these silica-rich melts
rose from deep within Earth and formed granitic plutons (intrusive bodies) nearer the
surface. Early in the Archean the granitic crust of continents had begun to form from
basaltic crust of the ocean floor. Continental landmasses began forming about 3.7
billion years ago from the horizontal accretion of smaller micro-continents. The Kenoran
orogeny was one such event, which formed what is now the Great Lakes region of
North America during the Late Archean.
The First Life on Earth
Earth was able to support life only after the planet had cooled enough for a rocky
crust to solidify. Once that happened, water vapor from volcanoes condensed in the
atmosphere, fell as rain, and collected on the Earths surface. Besides water vapor,
volcanoes also produced gases rich in the basic ingredients of life: carbon, hydrogen,
oxygen, and nitrogen. Toxic gases such as ammonia and methane were common. At
this point, Earth's early atmosphere consisted entirely of these volcanic gases, and
there was no free oxygen. In the primordial soup of the early seas, organic molecules
concentrated, formed more complex molecules, and became simple cells.

The transition from complex organic molecules to living cells could have occurred in
several environments. Small, warm ponds are one possibility, but recent work has
suggested that deep-sea hydrothermal vents, such as those found along mid-ocean
spreading centers today, may have been the cradle of Earth's life. These environments
contain the chemicals and the source of energy needed to synthesize more complex
organic structures. Although scientists have not succeeded in creating life from organic
molecules in the laboratory, they have reproduced many of the intermediate steps.
Changes in the Atmosphere
During the Archean and early Proterozoic Eons, the deep oceans had large volumes
of dissolved iron. This combined with oxygen, possibly produced by photosynthetic
microorganisms, to produce iron oxides along the continental margins. This rust was
concentrated in sedimentary deposits known as Stromatolites and other microfossils
provide important evidence for the transformation of the oxygen-poor atmosphere of
the Archean, into one that was oxygen-rich in the Proterozoic, like the atmosphere
today. Some microorganisms that build modern stromatolites are capable
of photosynthesis and release free oxygen into the ocean. Chemical traces of
microorganisms, known as biomarkers, show that photosynthetic organisms had
evolved by 2.7 billion years ago, but they were probably not present during the early
Archean.

c. Proterozoic (2,500- 542 mya)

Defining Characteristics:
o first multicellular animals at end of interval
o Map of the Proterozoic World
Secondary Characteristics:
o 4 major mountain-building episodes
o oldest known glaciation

The term Proterozoic comes from the Greek words proteros (meaning first) and
zoon (meaning life). Although early fossils are now thought to occur in older, Archean
rocks, for many years the oldest evidence of life was known from Proterozoic strata.
The Proterozoic represents the last stage of what was once called the Precambrianthe
3.7-billion-year-long period before the Paleozoic Era. Rocks of this age are known from
Australia, Canada, and China.
During the Proterozoic, larger continental landmasses continued to form by the
accretion of smaller ones, often leading to extensive mountain-building. As the
continents began to erode, sediments were washed into the oceans, producing shallowwater marine environments where life could flourish and diversify. Many of these life
forms developed the ability to photosynthesize. As a byproduct of photosynthesis, they
created oxygen, and over billions of years this oxygen transformed the Earths
atmosphere. Because oxygen was toxic to some early life forms, many went extinct.
Others thrived and evolved into the first multicellular organisms, which are preserved
as the Ediacaran fauna. Both the beginning and end of the Proterozoic were marked by
widespread glaciation.
Earths Crust as a Platform for Prokaryotic Life
For much of the history of living things, organisms were exclusively primitive, singlecelled forms such as bacteria and cyanobacteria. Then as now, these simple
prokaryotic cells lacked a nucleus and were relatively small, but they already contained
the same basic hereditary materials found in all life todayDNA and RNA. Many of
these early life forms relied on chemosynthesis for their energy, deriving it from the
chemicals around them instead of from sunlight or other organisms. Some inhabited
environments that seem hostile, such as deep-sea hydrothermal vents.
Eukaryotes and the First Multicellular Life Forms
A fundamental biological change occurred with the appearance of eukaryotes.
Eukaryotes differ from prokaryotes in that their cells contain membranous sacs called

organelles, including mitochondria, chloroplasts, and the nucleus. Many scientists think
these organelles are descended from formerly free-living prokaryotic organisms. Thus,
many important functions of eukaryotic cells, such as photosynthesis, and respiration
(the process by which organisms use oxygen to metabolize organic compounds to
produce energy, giving off carbon dioxide) were acquired through a symbiosis of
independent forms of life. Eukaryotes flourished as the environment became richer in
oxygen, perhaps in part because of their more complex intracellular function.
An important evolutionary innovation was multicellularity. The oldest known
possible multicellular eukaryote is Grypania spiralis, a coiled, ribbon-like fossil two
millimeters wide and over ten centimeters long. It looks very much like a coiled
multicellular alga and has been described from banded iron formations in Michigan 2.1
billion years old. Grypania may not be a eukaryote, but another, unrelated colonial
eukaryote, Horodyskia, is known from sedimentary rocks dated at 1.5 billion years in
western North America and from rocks more than 1 billion years old in Western
Australia.
Changes in the Atmosphere
Earths early atmosphere contained only small amounts of free oxygen, probably
produced entirely by the reaction of sunlight with water vapor from volcanoes. The
oxygen-rich atmosphere that evolved later, and upon which oxygen-breathing life now
depends, was a result of the origin of photosynthesis. During the Precambrian, vast
numbers of single-celled algae and cyanobacteria living in the seas eventually released
enough oxygen to transform the environment. The oldest evidence of cyanobacteria
dates to 2.7 billion years ago, although oxygen did not begin to build up in the
environment until about 2.3 billion years ago. During the transition from oxygen-poor to
oxygen-rich atmosphere, the first banded iron formations may have formed.
Thus, the history of Earths early crust also tells the story of its early atmosphere.
Banded iron formations were precipitated from about 3.1 to about 2 billion years ago
most (92%) during the Proterozoic between 2.5 and 2 billion years ago. Until all the
available iron had been deposited in banded iron formations, oxygen could not build up
in the atmosphere. Red beds appeared only after free oxygen was released into the
atmosphere, beginning about 2.0 to 1.8 billion years ago. They are still being formed
today.
Proterozoic Mountains and Glaciers
In the Proterozoic Eon, four major mountain-building episodes occurred, each of
which was followed by an interval of continental erosion. Mountain-building was caused
by converging plates, just as occurs in present-day plate tectonics. It was accompanied
by intrusions of molten granite that welded an additional belt of younger, igneous rock
around the edges of the original microcontinents. In North America, the Proterozoic
episodes of mountain-building greatly expanded the size of the continent.
Widespread continental glaciations evidently occurred at least twice in the
Proterozoic, once near its beginning and again near its end. Several of these glaciations
extended almost to the equator, much farther south than any recent cooling events.
This unusual situation has led a few geologists to propose that the Earth was almost
entirely covered by glaciers for perhaps several million years during the Proterozoic.
During this Snowball Earth phase, life would have been relegated to hydrothermal
vents and other such refuges until the build-up of carbon dioxide released from
volcanoes warmed Earth from its deep-freeze. The final Marinoan Ice Age marked the
transition to the Cambrian, the first period of the Paleozoic Era.
II.

Phanerozoic
a. Paleozoic
i. Cambrian (542- 488.3 mya)

Defining
Characteristics:
o Cambrian
Explosion
o skeletonized
animals
o early animal
diversification

Secondary
Characteristics:
o diversification
of trilobites
o Burgess Shale
fauna
o warming
climates
o Plate tectonics

The Cambrian was first defined by Adam Sedgwick in England


(Cumbria) and Wales; Cambria is the Roman name for Wales. Since then,
Cambrian-age rocks have been found on every continent; they occur
throughout North America, where the Burgess Shale, in British Columbia,
Canada, is an especially significant deposit of Middle Cambrian rocks that
preserve soft-bodied organisms. Lower Cambrian rocks in Yunnan, China,
preserve the famous soft-bodied Chengjiang biota. In Antarctica, Upper
Cambrian rocks in the Ellsworth Mountains preserve a highly varied shelly
fauna. Cambrian-aged shelly fossils occur widely throughout Australia, Europe,
the Lena and Aldan River areas of Siberia, and in North Africa.
Cambrian rocks were deposited between 543 and 490 million years
ago, and they are now defined as the first period of the Paleozoic Era; it is at
this time that animals having shells (exoskeletons) become widespread in the
fossil record. Before the Cambrian, in the late Neoproterozoic Era (often called
the Vendian), Earth had multicellular life; however, the organisms were soft
bodied. This Vendian soft-bodied biota occurs in the famous Ediacaran beds of
South Australia and now has been found in many other parts of the world.
Before the Vendian, life was largely represented by single-celled organisms
such as Archaea, filamentous microorganisms, algae, and other organisms that
are studied by using microscopes.
The Cambrian Explosion

The fossil record of Proterozoic life is sparse because it was largely soft
bodied or microbial. The advent of the Paleozoic Era brought an incredible
diversification of multicellular animals having many different body plans,
including most of the major groups alive today, such as mollusks, trilobites,
other arthropods, brachiopods, echinoderms, corals, sponges, and chordates
(our ancestors). Most significantly, the first animals having hard outer
skeletons evolved; as a result, many of these shells became fossilized. Most of
the Early Cambrian animals having exoskeletons were small, measuring one to
five millimeters, and are often called the "Small Shelly Fauna". Another unique
feature of the Cambrian fauna is the sponge-like archaeocyaths that formed
reefs during Early Cambrian time. This sudden appearance of many different
kinds of animals having skeletons (shells) in the fossil record is called the
Cambrian Explosion. The Cambrian Fauna is as different from Vendian life as it
is from the Paleozoic Fauna that occurs above Cambrian-age rocks.
The Burgess Shale Fauna

The Burgess Shale was deposited in the ocean near an underwater


algal reef shelf. Occasional undersea landslides buried the animals living there,
and the fine mud prevented decay so that soft parts were preserved. As a
result, this fossil fauna records a host of extinct soft-bodied organisms, many of
which have no living counterparts. Additionally, the Burgess Shale contains
many shelled animals, some of which preserve parts not normally seen, such
as the legs and antennae of trilobites and the setae of brachiopods; the
exceptional preservation allows paleontologists to understand the structure

and biology of the animals better. Some of the soft body plans of Cambrian
animals did not survive beyond that period. To our eyes, some of these body
plans look bizarre and it has taken a long time for paleontologists to
understand them. For example, the first reconstructions of the onycophoran
Hallucigenia were upside-down. The sclerite-bearing Wiwaxia, the arthropodlike predator Anomalocaris, and the five-eyed Opabinia are all strikingly
different from modern animals.
The "Age of Trilobites" and the Cambrian Fauna

The most abundant and diverse animals of Cambrian time were the
trilobites. Trilobites had long antennae, compound eyes, many jointed legs, and
a hard exoskeleton like many of their modern arthropod relatives, such as
lobsters, crabs, and insects. The Cambrian is sometimes called the "Age of
Trilobites" because of their explosive diversification into all marine
environments worldwide. In size, they ranged from a few millimeters (1 mm =
0.25 inches) to 45 centimeters (18 inches).

Following the Cambrian, trilobites remained an abundant and diverse


element of Ordovician marine faunas, but other groups of organisms that had
been more minor elements of Cambrian faunas diversified dramatically. These
include snails, clams, brachiopods, cephalopods, corals, bryozoans, and the
now extinct graptolites. This post-Cambrian radiation, the Paleozoic Fauna,
would dominate marine life until the end of Permian time.
Plate Tectonics at the Start of the Paleozoic

The Cambrian began after the end of the Marinoan ice age, one of
several instances in the past when ice was widespread on Earth's surface. This
marked a shift from late Proterozoic icehouse conditions, during which much of
the world was cold and ice covered. The Cambrian was quite the opposite, with
global warming leading to greenhouse conditions. This climate change was
probably linked to major changes in the positions of the continents.

During the Cambrian, the large Proterozoic landmass (Rodinia) broke


up and plate-tectonic movements pushed North America, Siberia, and
Gondwana, as well as smaller landmasses, toward the equator. Eventually, in
Cambrian time, most of the land was in tropical, subtropical, and temperate
climatic zones; the ice masses melted and oceans flooded the continental
shelves. Marine life flourished in shallow, warm-water continental-shelf
environments and limestone deposition became common; Cambrian
limestones are known from all parts of the world.

ii. Ordovician (488.3- 443.7 mya)

Defining
Characteristics:
o diversification
of marine
invertebrate
Paleozoic
Fauna
o endOrdovician
extinction

Secondary
Characteristics:
o deep-water
faunas,
especially
graptolites
o appearance of
conodonts
o development
of diverse
carbonate
platforms
o first major
Paleozoic ice
age


Charles Lapworth in 1879 named the Ordovician for a set of rock strata
that were particularly well exposed in Wales. He took the name from Ordovices,
the Latin name of a Celtic tribe in the Welsh region during Roman times. By
creating the Ordovician, Lapworth helped settled a dispute, because previously
Adam Sedgwick had assigned these rock layers to his Cambrian, whereas
Roderick Murchison called them Silurian. Lapworth recognized them as distinct
from the (older) Cambrian rocks below and the (younger) Silurian rocks above.
This processdetermining the age of a rock by the fossils contained within it
is called biostratigraphy.

Ordovician rock layers were deposited between 443 and 490 million
years ago. During this time period, marine life saw a major shift from the
Cambrian Fauna, which appeared during the Cambrian Explosion, to the
Paleozoic Fauna, which would dominate the marine world for the next 200
million years. Important new animal forms, such as graptolites and conodonts,
first appeared during the Ordovician, along with the development of complex
shallow-water reef ecosystems. The first land organisms, lichens and
bryophytes, appeared. The end of the period was marked by a major ice age
and the second-largest mass extinction of the Paleozoic Era.
Reef Ecosystems and the Paleozoic Fauna

Life in the Ordovician was quite different from that in the Cambrian.
Although many animal groups survived from the Cambrian, some forms
disappeared and other forms appeared for the first time. These changes
marked the Ordovician as the first example of the Paleozoic Fauna. During this
time, bryozoans, mollusks, echinoderms (particularly the fan-like crinoids),
brachiopods, and trilobites diversified; these groups would dominate marine
faunas throughout the Paleozoic. Many of the unusual animal forms seen in the
Burgess Shale had gone extinct, but graptolites, conodonts, nautiloids, and
jawless fishes had their first known appearances.

These new animals radiated alongside the rugose and tabulate corals
that replaced the Cambrian archaeocyaths. In fact, entirely new types of
ecosystems evolved in the Ordovician, associated with the development of
many types of carbonate (limestone) marine shelves (or platforms). These
warm, shallow-water formations were flooded with sunlight, allowing the
establishment of stable reef ecosystems. These included the first coral/algal
reefs, shallow-water assemblages of mollusks, brachiopods, echinoderms and
trilobites, and more-diverse underwater slope and deeper-water faunas. Many
of these faunas would become even more important in later Paleozoic periods.
New Animals as Biostratigraphic Tools

Another important change in the Ordovician was the evolution of


deeper-water faunas, as many forms of life moved out of their early shallowwater environments. With this change came the appearance of new groups of
animals. One of the most important among these was the graptolites, so
named because their fossils resemble writing on rocks. Graptolitesare
hemichordates, relatives of modern acorn worms and pterobranchs and distant
relatives of chordates and vertebrates. For many years, their exact biological
nature was a mystery to paleontologists, but their rapid rate of evolution made
them useful nonetheless. Because each graptolite species only existed for only
a short time, their fossils could be used to identify the age of a particular rock
quite specifically. This processdetermining the age of a rock by the fossils
contained within itis called biostratigraphy. For more information on the
concept of biostratigraphy, please visit the Foundational Concepts page.
Ordovician Climate and Plate Tectonics


During the Ordovician, tectonic activity occurred in New England and
the British Isles as North America moved toward the equator and began to
collide with Europe. This led to mountain-building along these major plate
margins. As plates continued to shift in position, diverse faunas were brought
into contact with one another. Plate movement may also have contributed to
the formation of the first Paleozoic ice age, as the southern continent of
Gondwana moved over the South Pole. Glaciers began to form in what is now
the Sahara region of Africa. This climate change was possibly associated with
the mass extinction at the end of the Ordovician. These tectonic clues allow
scientists to make sense out of geologic patterns in the Earth, and to map
present-day continents and islands onto these old deposits.

iii. Silurian (443.7- 416 mya)

Defining Characteristics:
o distinct estuarine, freshwater, and terrestrial ecosystems develop
the start of significant life on dry land
Secondary Characteristics:
o first vascular plant fossils
o wide and rapid spread of jawless fishes
o first extensive coral reefs

The Silurian physical world was vastly different in many ways from the
world of today. For example, the equator passed through what is now North
America, and a nearly continuous inland sea extended from New York to
Nevada. Life was also quite different. Following the end-Ordovician extinction
event, Silurian marine faunas rebounded in the warm, shallow continental seas.
Large coral reefs made their first appearance during this time. Although marine
life was diverse and abundant, there were only a few types of small plants and
terrestrial arthropods on land in the Silurian. However, these represent the first
true terrestrial ecosystems on Earth and include the first fossil records of
vascular plants. These terrestrial ecosystems became more complex, especially
during the Late Silurian, as upright plants evolved and populated the wet parts
of lowland landscapes.
Silurian Marine Life

Marine life thrived and diversified during the Silurian; many forms were
similar to those of the earlier Ordovician Period. Although the later Devonian
Period is often called the Age of Fishes, the Silurian also was an important
time in the evolution of this group. In particular, it was marked by the wide and
rapid spread of jawless fishes (agnathans). Only two of their jawless
descendantslampreys and hagfishessurvive today, but many different kinds
populated the Silurian seas.

Among the invertebrates, brachiopods were especially diverse and


abundant. Many survived from the Ordovician, but other new forms evolved as
well, so that brachiopods represented nearly 80% of all Silurian marine
invertebrate species. Although the planktonic graptolites lost many species in
the end-Ordovician mass extinction, they also remained abundant. The
common genus Monograptus included many species that are useful today as
biostratigraphic fossils. Tabulate and rugose corals, calcareous algae,
stromatoporoids, and bryozoans formed shallow-water tropical reefs.
Cephalopods, gastropods, and echinoderms were among the active predators
of the time. Trilobites began to decline in diversity after having reached their
peak in the Cambrian and Ordovician.
The Invasion of Land

Early vascular plants, collectively known as rhyniophytes, first


appeared in the late Early Silurian and diversified considerably through the

Late Silurian. These plants had evolved an internal system of tubular cells
through which water traveled up the body of the plant, allowing them to
maintain cell functions higher above the land surface than would otherwise
have been possible. Simple, small plants such as Cooksonia lacked leaves,
roots, seeds and the capability to grow in diameter, but they were the
dominant early terrestrial plants.

By the Middle Silurian, very early terrestrial communities had


developed around the tiny, vertically growing plants that acted as primary
producers. True herbivores were absent from these early land ecosystems.
Dead plants were recycled by fungi and bacteria. Early arthropodspossibly
related to millipedesalso fed on dead and decaying plant matter rather than
on living plant tissues. These arthropod detritivores may also have digested the
bacteria and fungi that lived in the soil. Other arthropods, possibly related to
centipedes, preyed upon the detritivores. The MidLate Silurian terrestrial biota
was probably confined to relatively wet areas.
Geology and Climate

During the Silurian, Earth underwent considerable changes that had


important repercussions for the environment and life within it. Plate tectonic
activity continued to shift the continents during the Silurian. The great
southern continent of Gondwana drifted farther across the South Pole, while
Siberia, Laurentia and Baltica clustered around the equator. Laurentia and
Baltica collided at the end of the Silurian, forming a new supercontinent,
Euramerica, and raising new mountain ranges. Rising sea level formed a nearly
continuous sea from New York to Nevada, and other shallow seas still covered
parts of other continents.

iv. Devonian (416- 359.2 mya)

Defining Characteristics:
o Age of Fishes
o diverse land invertebrates and first land vertebrates
o diversification of vascular plants
Secondary Characteristics:
o Late Devonian extinction
o first true trees

The recognition of the Devonian Period was the result of great debate
on the part of many nineteenth-century European geologists. The debate
centered on whether the Old Red Sandstonethe rock layers above the
Silurian (and therefore younger)actually represented a distinct system or
merely a later stage of the Silurian. In 1839, Roderick Murchison and Adam
Sedgwick collaborated to name the Devonian, which they based on rock
exposures in Devonshire, England. Devonian-age rocks are also common in
Scotland, central Pennsylvania, western New York, and Greenland, but they
have been found on all continents.
During the Devonian, most of Earths landmasses formed two
neighboring supercontinents, Gondwana and Euramerica. The rest of the
Earths surface was covered by a vast ocean. The Devonian world was
populated by now-extinct, very primitive plants and animals, so it looked much
different from our world today. In the marine realm, many members of the
Paleozoic Fauna continued to diversify. On land, vascular plants and arthropods
formed diverse terrestrial ecosystems, while the earliest tetrapods appeared in
shallow waters nearby.
Life in the Devonian Seas

The first radiation of vertebrates with jaws (gnathostomes) occurred


during the Devonian, which is why it is sometimes known as the Age of

Fishes. Although they first appeared in the Silurian, spiny acanthodians and
armored placoderms reached their peak diversity during the Devonian and
began to dwindle in numbers later in the Paleozoic. Some, like the
giant Dunkleosteus, reached nearly three meters in length and were
menacing marine predators. Placoderms had no teeth, instead relying on selfsharpening bony plates in their jaws that performed the function of teeth.
Chondrichthyan (cartilaginous) fishes, such as sharks, were Ordovician
survivors that thrived during the Devonian. A few shark scales have been
dated to the Late Ordovician, but the earliest shark teeth are Early Devonian,
and the first relatively modern-looking sharks had evolved by the Middle
Devonian. The first truebony fishes (osteichthyans) appear as well, as
both lobe-finned (sarcopterygians) and ray-finned (actinopterygians) forms.
The lobe-finned fish are of particular interest because they gave rise to the
first land vertebrates, around 360 million years ago
Terrestrial Habitats Conquered

The Devonian was the time during which life truly conquered land. At
the start of the period, terrestrial life was restricted to the narrow margin along
the waters edge. The first land plants had already evolved but still required a
moist environment to reproduce. By the middle of the Devonian, the first
shrub- and tree-like plants had appeared, some of which reached heights of
five meters or more. By the Late Devonian, the first true trees and forests had
evolved. This change is marked by the arrival of the genus Archaeopteris,
whose species lived on nearly all Devonian landmasses. Like modern forms,
these trees had extensive root systems, broad leaves, and specialized vascular
systems that facilitated the flow of water and nutrients against the pull of
gravity. All of these features helped plants colonize drier areas, which
expanded lifes terrestrial habitat. Broad leaves provided shade and moderated
temperature and humidity levels, whereas advanced root systems assisted
production of the first soils (pedogenesis) by encouraging weathering of the
land. These early forest habitats had important ecological effects on all other
ecosystems. Gymnosperms appeared at the very end of the Devonian. These
were the first plants to bear seeds, which allowed them to move farther into
drier environments.
The Devonian Extinction

Toward the end of the Devonian, nearly 70% of all invertebrate species
vanished during the Late Devonian extinction. Marine species (especially
tropical ones) suffered the most extinctions, followed by freshwater species,
whereas terrestrial species were hardly affected. In particular, tabulate
coral/stromatoporoid reefs vanished entirely, leading to a great decline in
worldwide coral reef-building until the Triassic Period. Many species of
brachiopods, trilobites, and early fishes went extinct, as did the planktonic
graptolites. These extinctions were not the result of a single major extinction
event, but rather smaller extinction events that occurred over a period of more
than 20 million years.
Climate and Tectonics during the Devonian

During the Devonian, there were two supercontinents, Gondwana and


Euramerica, located relatively close to each other in the Southern Hemisphere.
A vast ocean covered the rest of the globe, and the land was comparatively
arid. Global climate was relatively warm and dry, and there was less of an
equator-to-pole temperature gradient than today. There were no glaciers until
the Late Devonian, when ice began to cover parts of the South Polar region.

In the north, the formation of the continent of Euramerica continued


from the Silurian. The northern branch of the Iapetus Ocean closed and the
Late Caledonian (or Acadian) orogeny continued. The Euramerican continent,

once called the Old Red Continent because of the color of Devonian-age
rocks in Europe and North America, began to drift northward. Gondwana also
drifted north. The formation of Euramerica and the closure of the oceans
between Euramerica and Gondwana initiated the formation of Pangea, which
continued into the early Mesozoic.
v. Carboniferous

1. Mississippian (359.2- 318.1 mya)

Defining
Characteristics:
o first truly
terrestrial
tetrapods

Secondary
Characteristics:
o formation
of
supercon
tinents


In 1869, American geologist Alexander Winchell coined the
name Mississippian in 1869 for rock outcrops along the drainage basin of
the Mississippi River. He distinguished these limestone-rich Lower
Carboniferous rock layers from the coal-bearing beds of the Upper
Carboniferous (or Pennsylvanian). Because these two sets of rocks are
easily distinguished in North America, the terms Mississippian Epoch and
Pennsylvanian Epoch are commonly used by American geologists and
paleontologists. This distinction was less clearly marked in Europe, and
the names Lower and Upper Carboniferious were widely used. The
European subdivision of the Carboniferous has now been formally replaced
by Mississippian and Pennsylvanian by the International Commission on
Stratigraphy.

The Mississippian world was rather uniformly warm, although yet


another ice age began toward the end. As more continents began to
collide, extensive mountain-building occurred. The tropical seas were
home to a great diversity of marine life, including many different kinds of
fishes, brachiopods, bryozoans, mollusks, and echinoderms. On land, a
major habitat division began between seed plants, which preferred drier
habitats and the lycopsids (club mosses), which preferred wetter places.
There were also the first significant radiations of terrestrial tetrapods and
winged insects. In the Pennsylvanian, vast coal swamps would form, the
feature for which the Carboniferous Period was named.

Life in the Seas

Corals, mollusks, arthropods, and echinoderms (particularly


crinoids) flourished in the warm, shallow seas. Fusulinid foraminiferans
made their first appearance in the Mississippian. Brachiopods and
bryozoans also thrived in these marine waters, and both groups were
represented by many different families. Predatory ammonoids and other
cephalopods were abundant, hunting many of these less mobile
organisms. Trilobites were much less common than in previous time
periods. Toward the end of the Mississippian, the continents uplifted to
form more deltas and floodplains that supported gastropods, fresh-water
arthropods, and fishes. Sharks were especially common, along with bony
fishes such as acanthodians, lungfishes, coelacanths, and palaeoniscoids.

Tetrapods and Other Life on Land

Terrestrial arthropods in turn became food for early tetrapods:


the Mississippian was a time of great tetrapod evolutionary radiation.
Many tetrapods were still semiaquatic, but a great variety of early
terrestrial forms had appeared as well. Early forms such as Ichthyostega
and Acanthostega had now been replaced by many new species. Most of
these later Paleozoic amphibians are divided into two major groups
labyrinthodonts and lepospondyls. Labyrinthodonts include
temnospondyls and anthracosaurs (or batrachosaurs), which differ from
each other in skull and vertebral anatomy. Although it has been difficult to
link the modern types of amphibians to these early tetrapods,
paleontologists believe that they were quite similar in basic biology. Like
modern amphibians, early tetrapods were probably tied to the water
throughout their lives. Their eggs, which were not surrounded by any
protective covering (other than a porous membrane), were probably
fertilized in the water.

Mississippian Tectonics and Climate


The uplifting of the continents in the Mississippian reduced the
area covered by inland seas and resulted in less available space for
marine life. As Gondwana continued to move across the South Pole during
the Late Mississippian, temperatures cooled and ice began to form in
great sheets. As more water was frozen into these ice sheets, sea levels
dropped and terrestrial habitats in turn increased. The globally warm
climate became cooler later in the Mississippian.

The supercontinent of Gondwana continued to drift northward


and collide with Euramerica during the Mississippian. The forces in this
collision began to build the Appalachian and Variscan Mountains. The
merged continents formed a nearly continuous landmass that straddled
the equator. In the latest Paleozoic and early Mesozoic the combined
Euramerica and Gondwana would become the western part of the
supercontinent Pangea. Present-day China was still represented by some
isolated subcontinental blocks not yet sutured to Pangea.

2. Pennsylvanian (318.1- 299 mya)

Defining Characteristics:
o Coal Age, very well known for plant fossils that make up
the worlds major coal seams
Secondary Characteristics:
o Age of Amphibians, including the ancestors of modern
terrestrial vertebrates
o diversification of terrestrial and aquatic insects.
o evolution of amniotic egg in reptiles

The Pennsylvanian Subsystem was named for Pennsylvania,


home of some of North Americas richest coal seams. The name was
coined in 1891 by Henry S. Williams for the Upper Carboniferous rock
layers of North America. Like Mississippian, the term Pennsylvanian was
used more commonly by American geologists because the coal-rich Upper
Carboniferous was more easily distinguished from the limestone-rich
Lower Carboniferous in North America than in Europe. The International
Commission on Stratigraphy has now formalized the division of the
Carboniferous into two Subsystems, the Mississippian and Pennsylvanian,
replacing the terms Lower Carboniferous and Upper Carboniferous.
During the Pennsylvanian, most continents were connected in
one landmass and the world was in the throes of an ice age. Nonetheless,
tropical plant growth was very lush. Huge, dense forests of ferns, tree
ferns, club mosses, horsetails, and seed-bearers (newly evolved in the
Late Devonian Period) occupied many terrestrial niches. Lowland swamps
accumulated the dead leaves, trunks, and branches of countless
generations of vegetation. Repeated for tens of millions of years, this
process laid down peat that would later become some of the major coal
seams of the world.

Pennsylvanian Animal Life

The terrestrial animal radiations that had started in the


Mississippian continued into the Pennsylvanian. The great coal forests
housed enormous insects such as dragonflies (Meganeura) with up to 70centimeter wingspans, paleodictyopterans with 55-centimeter wingspans,
and a variety of large protorthopterans and cockroaches. Other giant
terrestrial arthropods included large spider-like animals and a two-meterlong relative of millipedes called Arthropleura. Some of the giant
arthropods were early terrestrial herbivores, such as paleodictyopterans
with mouthparts adapted for piercing and sucking the juices out of plants,
and protorthopterans with chewing mouthparts. These herbivorous insects

became food for predaceous arthropods like the dragonflies and


scorpions, as well as for early tetrapods. These tetrapods were carnivores
(sometimes cannibals) as well as insectivores, but none had yet mastered
herbivory. Early amphibians included reptile-like anthracosaurs, snake-like
astopods, and eel-like semiaquatic forms.

Pennsylvanian Animal Life

The terrestrial animal radiations that had started in the


Mississippian continued into the Pennsylvanian. The great coal forests
housed enormous insects such as dragonflies (Meganeura) with up to 70centimeter wingspans, paleodictyopterans with 55-centimeter wingspans,
and a variety of large protorthopterans and cockroaches. Other giant
terrestrial arthropods included large spider-like animals and a two-meterlong relative of millipedes called Arthropleura. Some of the giant
arthropods were early terrestrial herbivores, such as paleodictyopterans
with mouthparts adapted for piercing and sucking the juices out of plants,
and protorthopterans with chewing mouthparts. These herbivorous insects
became food for predaceous arthropods like the dragonflies and
scorpions, as well as for early tetrapods. These tetrapods were carnivores
(sometimes cannibals) as well as insectivores, but none had yet mastered
herbivory. Early amphibians included reptile-like anthracosaurs, snake-like
astopods, and eel-like semiaquatic forms.

Pennsylvanian Animal Life

The terrestrial animal radiations that had started in the


Mississippian continued into the Pennsylvanian. The great coal forests
housed enormous insects such as dragonflies (Meganeura) with up to 70centimeter wingspans, paleodictyopterans with 55-centimeter wingspans,
and a variety of large protorthopterans and cockroaches. Other giant
terrestrial arthropods included large spider-like animals and a two-meterlong relative of millipedes called Arthropleura. Some of the giant
arthropods were early terrestrial herbivores, such as paleodictyopterans
with mouthparts adapted for piercing and sucking the juices out of plants,
and protorthopterans with chewing mouthparts. These herbivorous insects
became food for predaceous arthropods like the dragonflies and
scorpions, as well as for early tetrapods. These tetrapods were carnivores
(sometimes cannibals) as well as insectivores, but none had yet mastered
herbivory. Early amphibians included reptile-like anthracosaurs, snake-like
astopods, and eel-like semiaquatic forms.

Pennsylvanian Climate and Tectonics

In general the continents continued to collide and merge during


the Pennsylvanian. Pieces once attached to Gondwana drifted north,
approaching Siberia and the other northern continents. These
subcontinent-sized blocks, called North China, South China, and Cimmeria
were probably large islands during part of the late Paleozoic. Ultimately
they collided and merged with Siberia and the other continents to
complete the formation of Pangea during the Early Triassic.

As the continents continued to move around 270 million years


ago, the climate changed correspondingly. Swamps dried out and many
giant plants began to die out as the Pennsylvanian came to an end. The
world of that time was in an ice age, a generally cold global climate
phase. As a result it was climatically zoned, much like today. Ice sheets
were present at high latitudes, but the equator remained relatively wet

and tropical for much of the period. Mid-latitudes were seasonally dry, and
some of the changes in climate and vegetation recorded by fossils and
rocks in North America and Europe probably reflect the movement of
these areas from the warm, wet tropical belt into the seasonally dry
subtropics.

vi. Permian (299- 251 mya)

Defining Characteristics:
o the end of the Permian and the Paleozoic is based on the most
extensive mass extinction in the past 600 million years, leading to
the advent of dinosaurs and modern terrestrial and marine biotas
Secondary Characteristics:
o extensive glaciation at the start gives way to general global
warming
o great diversity of amphibians, diapsids, and synapsids
o the greatest high-level taxonomic diversity of insects of all time
o seed plants become more dominant

The Permian Period marked the end of the Paleozoic Era. The land was
inhabited by a wide diversity of terrestrial insects and vertebrates. Insects
included the earliest representatives of the two most dominant groups of
insectsthe hemipteroids, represented by extant cicadas and lice, and the
highly diverse holometabola, represented by beetles, flies, wasps, and moths.
Also important were the first vertebrate herbivores. Marine faunas were still
dominantly composed of groups belonging to the Paleozoic Fauna. The Earths
continents were coalescing into a single supercontinent, Pangea. The rise of
conifers as dominant elements of tropical vegetation reflects increased
seasonality of rainfall in tropical areas. The Permian is most remarkable for its
conclusion, however: the end-Permian extinction was the largest of the past
600 million years, and it heralded the end of the dominance of the Paleozoic
Fauna. In its aftermath, archosaurs rose to dominate the land, and more
modern groups filled the marine realm.
Terrestrial Animal Life and Evolution of Herbivores

The most important terrestrial herbivores in the Early Permian were the
insects. They represented a mixture of holdovers from the earlier
Pennsylvanian, such as the dragonfly-like palaeodictyopterans and various
forms related to modern grasshoppers and cockroaches, as well as newly
evolved forms such as beetles and scorpionflies that are the earliest known
members of insect groups that undergo true metamorphosis. (Insects that
metamorphose develop from egg to larva to pupa to adult, and generally make
their living in a very different way in the larval and adult phases.) The advent
of insects that could metamorphose had a profound effect on the ecology and
evolution of insects, because immature life stages could use different resources
than adults of the same species. Evidence for these new herbivores is seen in
Permian plant fossils with new types of damage to leaves and seeds.
Sedimentary rocks of Permian age also reveal fossil burrows made by the first
insects to colonize quiet-water aquatic ecosystems.

Permian Terrestrial Floras

Permian floras became much more xerophytic as the global climate


shifted from cold to warm. Tropical regions dried out as rainfall seasonality
increased. Only in what is now China did tropical rain forests and peat-forming
swamps persist into the Late Permian. With this drying a whole new suite of
plants appeared, expanding into many areas of the lowland landscape. Many of
these plants had probably been living in Pennsylvanian-age upland regions,
where soils were better drained than in the lower wetlands and rainfall may

have been more seasonal. Climatic zonation was pronounced, however, and
floras from different climatic belts were markedly distinct.

In addition, competition in these drier areas may have led to


diversification and greater variability among seed-bearing plants. Plants may
also have begun to evolve in response to the fact that they now had to survive
the onslaught of new, abundant vertebrate herbivores. The Permian was rich in
conifers (Walchia, Ernestiodendron), cycad-like plants (Taeniopteris,
Russellites), gigantopterids (Gigantopteridium, Cathaysiopteris, Zeilleropteris,
etc.), and callipterids (Autunia, Rachiphyllum). Most of these were seed
producing. Limited wet spots persisted, identified by the occurrences of tree
fern foliage, calamites (especially along stream banks), and rare tree lycopsids
(Sigillaria).
The Marine Realm and The End-Permian Extinction

The end of the Permian was marked by the greatest mass extinction of
the last 600 million years of Earth history, during which perhaps 90% of marine
animal species disappeared. Major groups such as trilobites, fusulinid
foraminiferans, rugose and tabulate corals, acanthodian and placoderm fishes,
and blastoid echinoderms vanished entirely, Although they did survive,
brachiopods, bryozoans, ammonoids, sharks, bony fishes, crinoids, eurypterids,
ostracods, and many echinoderms lost the majority of their species. Finally,
insects suffered their greatest mass extinction in Earth history.

Several factors have been implicated in this massive extinction. The


formation of Pangea reduced the continental shelves, decreasing the area
available for shallow-water organisms. Rapid warming and glaciation both
occurred during the Permian as well. These events do not seem to have
happened at the same time as the extinction event, however. Indeed, a first
extinction pulse actually occurred during the Middle Permian and may have
been caused by a dramatic drop in sea level. A more likely cause for the endPermian extinction was a series of volcanic eruptions in Siberia, which
produced massive outpourings of lava called flood basalts. This volcanism
covered an area about two-thirds the size of the United States and erupted
very rapidly just at the time of the extinction. It may have caused significant
atmospheric disturbances, global warming, and anoxic (low-oxygen) ocean
waters. The other possible cause is the impact of a large extraterrestrial object,
as occurred with the extinction at the end of the Cretaceous. Direct evidence
for such an impact is sparse, but the available data are consistent with such a
cause.
Tectonics, Climate, and the Formation of Pangea

The southern ice cap melted off permanently during the earliest
Permian. The supercontinent of Pangea was nearly fully assembled by the end
of the period, with only North and South China and Cimmeria remaining
unattached to the east. An open gulf called the Paleo-Tethys Ocean formed
along Pangeas eastern margin. The total length of shorelines continued to be
reduced as the major landmasses coalesced, and shallow seas decreased as
sea level dropped. The salty, inland Zechstein Sea occupied much of Europe.

Throughout these events, the Permian world saw a gradual warming in


climate. There was steep climatic zonation despite temperatures being warmer
than during the Pennsylvanian. Swampy tropical forests gave way to more arid
environments. The dry continental interiors (now very large on the
supercontinent of Pangea) experienced great seasonality, and the polar regions
were barren tundras. Glaciers were also present at these high latitudes.
Nonetheless, many animals were spread across wide regions of the globe in
each climatic area.

b. Mesozoic

i. Triassic (251- 199.6 mya)

Defining Characteristics:
o Early Triassic: the start of lifes remarkable recovery after the endPermian extinction
o the origin and rise of dinosaurs and the first mammals in the
Middle to Late Triassic
Secondary Characteristics:
o the emergence of giant marine reptiles (ichthyosaurs) and the
rapid diversification of cephalopod ammonites in the oceans
o the first truly modern coral reefs in the Middle Triassic

At the start of the Triassic, 252 million years ago, the equatorial
supercontinent of Pangea completed its formation as the final continents
collided with the mainland. Polar ice caps were absent and sea level remained
fairly low and stable. The climate was generally warm and arid, but moist river
and lake environments sustained gymnosperm forests and large amphibians.
Nevertheless, life on Earth was sparse. Only a few types of animals and plants
were left after the great mass extinction at the end of the Permian, and sea life
in particular was severely depleted. The repopulation of Pangea had meager
beginnings, and the rediversification of life took 46 million years. The Early
Triassic is characterized by low diversity in both marine and terrestrial habitats,
but by the Middle or Late Triassic diversity had rebounded in most
environments.
Extinction and Recovery

The first third of the Triassic was a recovery period from the endPermian extinction. This greatest extinction in Earths history, eliminating
approximately 70% of the species of land vertebrates and 90% of marine
animal species, sharply reduced the number of different ways in which plants
and animals made a living. During the Triassic life re-evolved many strategies
for living, and added new ones not seen during the Paleozoic. Newly evolved
scleractinian corals formed small reefs, beginning the recovery of reef
ecosystems. Mollusks such as ammonoids (relatives of the modern chambered
nautilus) were severely reduced in diversity by the extinction but evolved
rapidly afterward to become more diverse than ever before and to dominate
the open-ocean marine invertebrate world.
New types of animals and plants continued to evolve throughout the
Triassic. The first mammals and dinosaurs originated almost simultaneously, in
the late Middle or early Late Triassic. The oldest known fossil of an amniote egg
is from the Early Triassic. In the seas, ichthyosaurs (dolphin-shaped reptiles),
nothosaurs, and placodonts (mollusk-eating reptiles) appeared and thrived.
Some ichthyosaurs reached lengths of 23 meters (75 feet). Turtles,
crocodyliforms, and pterosaurs all made their debuts, along with frogs and
sphenodontians. In fact, by the end of the Triassic, many of the animal groups
we see today had made their first appearances on Earth.
Origin of Mammals

Although the Triassic Period is often remembered as the time when


dinosaurs and other archosaurs rose to dominance, mammals and their
ancestors also played an important role. In fact, these synapsids (once called
mammal-like reptiles) were very abundant in the early Triassic world. They
came in a wide variety of sizes and shapes, from large, cow-like dicynodonts
(such as Lystrosaurus), to fanged, carnivorous gorgonopsians, to tiny
insectivorous tritylodonts. A Triassic landscape in South Africa would have been
filled with great numbers of these early synapsids. By the Middle Triassic
(about the same time as the first dinosaurs) one of these synapsid groups had

evolved into the very first mammals. Although many different types of
mammals evolved later in the Mesozoic, for the most part they remained small,
probably because of the dinosaurs' success. However, later in the Mesozoic
some mammals achieved the size of a 30 lb. (13 kg) dog despite the dominant
success of the dinosaurs. Many types of mammals appeared in the Mesozoic,
and evidence from Early Cretaceous outcrops (130 million-years ago) in China
shows that at least one carnivorous mammal, Repenomamus robustus, feasted
on small dinosaurs.
Origin of the Dinosaurs

The earliest dinosaurs are known from Argentina and include predatory
theropods (Herrerasaurus, Eoraptor) and herbivorous ornithischians
(Pisanosaurus). By the end of the Triassic, dinosaurs were widespread and
dominated most terrestrial ecosystems. The most abundant were small
theropods such as Coelophysis (from North America), and large, herbivorous
prosauropods such as Plateosaurus (from Europe). Rare evidence also exists of
other groups, including the first sauropods and armored dinosaurs. During the
following Jurassic Period, dinosaurs and other archosaurs would become even
more diverse and spectacular, evolving into gigantic sauropods, large
theropods, and birds. Their dominance would continue until the end of the
Mesozoic.
Climate and Plate Tectonics

Pangea was maximally developed at the start of the Triassic, with


Panthalassia (all ocean) occupying the other side of the globe, and the Tethys
Ocean forming an enormous gulf on the eastern margin of Pangea. Even as the
assembly of Pangea was completed, however, it began to rift apart. By the end
of the Triassic, rifting in the center of Pangea had begun. North America began
to pull away from Europe and Africa, and blocks of crust sank to create rift
valleys. During the Late Triassic and Early Jurassic, these valleys were
associated with the initial formation of the Atlantic Ocean. The sediments that
filled these rift valleys are preserved along the eastern margin of North
America and the western edges of Africa and Europe, and they contain
important evidence of Late Triassic organisms and their environments.

Early Triassic climate was quite similar to that at the end of the
Permian. Much of Pangea was warm and dry, and the interior of this
supercontinent was particularly arid. These environments were often
dominated by conifers and other gymnosperms. However, the Triassic also saw
an increase in seasonality, as well as prominent monsoon weather cycles.
Provincial biotas developed as well. In the north (Laurasia), these ecosystems
included ginkgoes, bennettitalians, cycads, and tree ferns. In contrast,
southern (Gondwanan) environments were dominated by seed ferns, most
prominently one called Dicroidium.

ii. Jurassic (199.6- 145.5 mya)

Defining Characteristics:
o The Age of Dinosaurs: dinosaurs become very diverse, evolving
into stegosaurs, theropods, and huge sauropods
Secondary Characteristics:
o origin of birds
o origin of the parasitic feeding guilds in terrestrial ecosystems
possibly replacing predators as top insect carnivores
o Pangea continues breaking apart, and is better separated toward
the end of the Jurassic, with a lush, warm tropical climate

The name Jurassic comes from the Jura Mountains, an extension of the
Swiss Alps into eastern France, where rocks of this age were first studied. They
were first identified as the Jura Kalkstein (Jura Limestone) by Alexander von

Humboldt in 1799, and later termed the Terrains Jurassiques by Alexander


Brongniart. The rocks were officially named the Jurassic System by Leopold von
Buch in 1839. Formed approximately 144206 million years ago, Jurassic rocks
have now been found on every continent.

Pangea was centered on the equator for most of the Jurassic Period,
and Earths climate was decidedly tropical. During the Early Jurassic, some
regions of the world were still arid, but by the Late Jurassic much of the planet
was lush. Great rivers covered North America, and the land was green with
ferns, seed ferns, cycads, ginkgos, and conifers. It was the ideal environment
for the largest animals ever to walk on Earthgiant sauropods such as
Diplodocus, as well as many other types of large dinosaurs. Like today, much of
the land was covered with trees, although flowering plants had not yet evolved.
And, like today, there were many understory plants such as ferns, cycads, and
horsetails, but there was no grass for smaller animals to hide in. Because of the
dinosaurs dominance, mammals were still no larger than rats but had begun
to diversify by the Early Jurassic. The Jurassic seas were filled with many types
of sharks, bony fishes, marine crocodiles, and other marine reptiles of all sizes.
Cephalopods such as ammonites propelled themselves through the oceans.
Birds evolved and began to diversify during the Late Jurassic, but the most
common flying vertebrates were the reptilian pterosaurs. By the end of the
Jurassic, major parts of Europe and North America had become flooded as sea
level rose, and Pangea continued to break apart. As North America and Eurasia
drifted away from Africa and South America, the Atlantic Ocean was born,
creating a barrier for land travel between these regions.
Jurassic Life

Rising sea levels flooded many of the continental interiors during the
Jurassic, creating warm, shallow-water environments where marine animals
and plants could thrive. These regions saw an increase in diversity of
microscopic phytoplankton such as coccolithophores, dinoflagellates, and
foraminiferans. Reef ecosystems continued to flourish, thanks to many species
of corals and sponges. Among these sessile (stationary) organisms lived
gastropods (snails), along with the now-rarer brachiopods and crinoids. In the
waters above swam predatory cephalopods such as ammonoids and
belemnites. Sharks and bony fishes remained common and shared the seas
with ichthyosaurs, plesiosaurs, and other marine reptiles. The first true marine
crocodiles had appeared, alongside the first true teleost fishes (which today are
the most diverse vertebrates on Earth).

Fresh-water environments were home to many invertebrates,


amphibians, turtles, and crocodilians, as in the Triassic. On land, herbivorous
insects diversified, and we see the first examples of many modern forms such
as leaf hoppers, snakeflies, and wasps. One extinct group, the
Kalligrammatidae, a group related to lacewings, possessed large, conspicuous
wings with big eyespots. These insects, with their fluid-feeding mouthparts,
probably were ecologic analogs to modern butterflies. Beetlesthe most
diverse group of organisms on Earth todaybegan their diversification during
the Jurassic as well. Some modern groups of beetles had evolved by this time
and fed on conifers, cycads, and ferns just as their descendents do today. Flies,
beetles and caddisflies diversified in aquatic habitats, feeding in a variety of
ways including filtering, collecting and shredding detritus, eating living plants,
and preying on other aquatic organisms.
Diversity in the Age of Dinosaurs

The Mesozoic, or Age of Dinosaurs, is often illustrated with a scene


from the Late Jurassic Period with its dramatic sauropods. But this was not the
picture throughout the Jurassic. In the Early Jurassic, many dinosaurs were very

similar to their Triassic relatives. These included prosauropods, early theropods


(Dilophosaurus), small ornithischians, and early armored forms (Scelidosaurus).
Sauropods were present but uncommon. By the Late Jurassic, however, these
long-necked, long-tailed herbivores ruled the land. Behemoths such as
Diplodocus, Apatosaurus, and Brachiosaurus were among the largest animals
to ever walk on land. Some of the largest Jurassic dinosaurs may have reached
lengths of 120 feet (35 meters) and weights of 60 tons. Because of their size,
the biology of sauropods is still subject to scientific debates. These revolve
around whether sauropods could lift their heads high in the air or rear up on
their hind legs, how they mated, and what foods they ate.

By the Late Jurassic nearly every major kind of dinosaur had appeared,
and they are found on every continent. Although prosauropods had gone
extinct, the world was now home to armored stegosaurs (Stegosaurus) and
ankylosaurs, herbivorous ornithopods of all sizes (Camptosaurus), and large
(Allosaurus) and small (Ornitholestes) predatory theropods. Many of these
dinosaurs were collected from the Morrison Formation, a thick formation of
mud, silt and sand that was deposited in western North America about 150
million years ago by a large braided river system running across much of the
central part of the continent. These Late Jurassic dinosaurs had close relatives
in Africa and Europe, indicating that these areas were still connected. In the
Cretaceous, however, these connections would finally be severed.
Flight and the Origin of Birds

One of the most important paleontological finds came in 1861, with the
discovery of Archaeopteryx in Late Jurassic (around 146 million years ago)
limestones near Solnhofen, Germany. Although the skeleton of Archaeopteryx
was nearly identical to that of the small theropod dinosaur Compsognathus,
this fossil also bore the unmistakable imprints of feathers. For over 100 years,
Archaeopteryx was the strongest evidence that birds had evolved from
theropod dinosaurs, and it therefore deserved a place on the dinosaur family
tree. Since then, many other feathered dinosaurs have been found in China,
further supporting this hypothesis, and future discoveries will help us
understand exactly how flight evolved in this unique group of theropods.

Although the first bird represents a remarkable evolutionary event, the


Jurassic skies truly belonged to another group of vertebrates, the pterosaurs.
These flying reptiles, relatives of the dinosaurs, had evolved in the Triassic but
by now were very diverse. Pterosaurs lived on nearly every continent and were
far more common than birds throughout the Jurassic. They came in many
shapes and sizes: long-tailed forms such as Rhamphorhynchus, huge-headed
Dimorphodon, and the sparrow-sized and short-tailed Pterodactylus.
Jurassic Climate and Tectonic Activity

Pangea, which had begun forming in the Devonian (400 million years
ago) and lasted through the Triassic, finally began to split apart in the Late
Triassic. By the Middle and Late Jurassic, enough plate movement had occurred
to separate South America from southern Africa. Laurasia (which consisted of
North America and Eurasia) also moved away from Africa and South America,
helping to create the Atlantic Ocean and the Gulf of Mexico. Volcanic activity
was common along these rifting continental margins. At the same time, Eurasia
(Europe and Asia) moved to the south and started to close off the Tethys
Ocean. Sea level gradually rose during the Jurassic, creating epicontinental
seaways in North America and Europe. The end result was a world with many
more separate land-masses, and a great deal more coastline, than the world of
the Triassic.

Much of the Jurassic world was warm and moist, with a greenhouse
climate. Coal deposits formed under forests in Australia and Antarctica.

Although some arid regions remained, much of the rest of Pangea was lush and
green. Northern (Laurasian) and southern (Gondwanan) biotas were still
distinct in many ways, but by the Jurassic, faunas had acquired a more
intercontinental character. Some animals and plants were now found nearly
worldwide, instead of being restricted to particular regions.

iii. Cretaceous (145.5- 65.5 mya)

Defining Characteristics:
o extinction of the dinosaurs
o first appearance and diversification of flowering plants
o extreme global warming
Secondary Characteristics:
o oldest known specimens of termites, ants, and bees
o diversification of birds and many new insect groups
o new kinds of dinosaurs

The name Terrain crtac (chalky terrain) was coined in 1822 by


Belgian geologist D'Omalius d'Halloy to refer to the chalk deposits of the Paris
Basin, France. Later the name Cretaceous (chalk-bearing, from the Latin word
creta, meaning chalk) came to be used for the many chalk deposits around
the world that were formed during this age. One of the most famous examples
is the White Cliffs of Dover, England. Such chalks formed from the bodies of
billions of single-celled marine algae called coccolithophores. The
coccolithophores lived in the sunny surface waters of the ocean, but after
death their shells or scales sank to the bottom where they accumulated, were
buried, and were compressed to form chalk. Coccolithophores are still
important organisms in the oceans today.

The world of the Cretaceous Period (65.5146 million years ago)


brought significant changes to life and to Earth itself. Before this time period,
during the Jurassic, animal life on land was dominated by dinosaurs. Some of
the dominant plants included ferns, cycads, seed ferns, ginkgos, and conifers.
In the seas, marine reptiles (ichthyosaurs and plesiosaurs), sharks, and
ammonites were common. Most of these life forms still dominated the
Cretaceous world, although new types of dinosaurs and plants also appeared.
All of Earths landmasses had been clumped together into one huge
supercontinent called Pangea, but this had begun to break apart during the
Triassic, and seaways had begun to invade Pangea during the Jurassic. By the
Cretaceous this process was well under way, making Earths climate more
equable and greatly affecting both plants and animals.

Terrestrial Life through the Cretaceous

The terrestrial environments in which Early Cretaceous animals lived


were still dominated by ferns, seed ferns, Bennettitaleans, conifers, and
cycads, but angiosperms (flowering plants) also became part of the flora at this
time, about 135 million years ago. Angiosperms diversified rapidly, and by the
end of the Cretaceous were by far the most diverse group of terrestrial plants.
Many explanations have been offered for their rapid diversification, but the
true explanation is probably quite complex. Some Cretaceous flowering plants
may have had their seeds dispersed by animals, but that was also true of other
kinds of plants. Today, some species of flowering plants grow rapidly compared
with living conifers and cycads, so perhaps the success of their Cretaceous
ancestors was related to their rapid growth. Another explanation is that many
angiosperms are pollinated by insects; insect pollination is thought to increase
the rate at which new species evolve. However, several other groups of
Cretaceous plants were also insect-pollinated. Whatever the reasons for the
success of the angiosperms, many new groups of insects evolved during the

Cretaceous, including the oldest known ants and bees as well as newly evolved
groups of pollinating species such as flies, beetles, wasps, and moths. Some
paleontologists think that the coincident evolution of these insect groups and
the diversification of flowering plants is an example of the process of
coevolution, in which two different types of organisms (such as an insect and
plant) become specifically adapted to one another. Insects also evolved more
types of feeding behavior both in quiet-water habitats such as lakes and in
flowing-water habitats.
Life in the Cretaceous Seas

Many groups of marine organisms continued through from the Jurassic


to the Cretaceous. Sharks of all kinds abounded, as well as many species of
bony fishes. Mosasaurs, a new type of aquatic marine lizard, were widely
distributed predators, with some species that reached over 14 meters in
length. Equally dangerous and just as large were plesiosaurs (such as
Kronosaurus) and crocodiles, but these were less common. Ichthyosaurs, which
dominated Triassic and Jurassic oceans, had all but disappeared by the Early
Cretaceous.

Reptiles were not the only marine giants of the Cretaceous. Strangelooking, often gigantic rudistid clams, reminiscent of Paleozoic horn corals,
reached up to one meter in length and formed extensive reefs in shallow
tropical oceans. Inoceramid clams over three meters long occurred in shallow,
warm seas, including environments that were nearly devoid of oxygen.
Ammonite cephalopods continued to diversify into amazing sizes and shapes,
with some coiled forms over two meters across, and other forms that
resembled an extended hook over two meters long.
Extinction of the Dinosaurs

Perhaps the most notable event of the Cretaceous was its conclusion.
About 65 million years ago the second greatest mass extinction in Earth history
occurred, resulting in the loss of the dinosaurs as well as nearly 50% of all the
worlds species. Though not nearly as severe as the end-Permian mass
extinction, the end-Cretaceous extinction is the most famous mass extinction
in Earth history. Other great animals also went extinct at that time, including
flying reptiles (pterosaurs) and the last mosasaurs and plesiosaurs. Many
mollusks, including rudistid and inoceramid clams, ammonites, and belemnites,
also became extinct, as did many species of microscopic marine plankton.
Terrestrial plants also suffered a major extinction at this time; in some regions
up to 60% of latest Cretaceous plant species were absent in the subsequent
Paleocene. Terrestrial insects also suffered a high level of extinction, especially
those that were highly specialized to feed on one or a few types of plants. In
fact, the level of insect herbivoryboth generalized and specializeddid not
recover to latest Cretaceous levels until the Paleocene-Eocene boundary,
approximately 9 million years later. In spite of the severity of extinctions at the
end of the Cretaceous, many types of animals and plants survived and gave
rise to new groups of organisms in the Paleocene.

The causes of the end-Cretaceous extinction are still being debated by


paleontologists. Researchers agree that a major factor was an asteroid about
10 kilometers in diameter that struck what is now the Yucatn peninsula in
Mexico. The effects of the impact were catastrophic, probably including global
forest fires, possibly a period of cold weather due to sunlight-blocking dust and
smoke, and a subsequent period of hot climate caused by the high levels of
CO2 released into the atmosphere by the impact. Evidence for the devastation
of terrestrial vegetation comes in the form of a thin rock layer deposited just
after the impact that is dominated by fossil plants whose present-day relatives
recover well after fires or other disturbances. Some paleontologists argue that

dinosaurs were already in decline before the asteroid impact, so that its
environmental effects merely hastened their extinction. Alternatively, others
point to the high abundance and variety of dinosaur species recorded even in
the sediments deposited just below the asteroid impact layer in the Hell Creek
Formation of western North America.
Shifting Continents and Greenhouse Climates

The Pangean breakup led to an increase in seaways and shorelines,


which may have affected the diversification of terrestrial plants and animals by
creating several isolated continental regions. In addition, rapid seafloor
spreading during the final breakup of Pangea caused a tremendous release of
carbon dioxide gas (CO2). This began in the Early Cretaceous and led to
dramatic global climate warming that culminated about 92 million years ago,
with nearly tropical temperatures extending to very high latitudes (polar
regions). Such extreme warmth resulted in ice-free polar regions that were
populated by dinosaurs, diverse forests and abundant insects. Fossils of
crocodile-like champsosaurs, which could not tolerate extended periods of subfreezing temperatures, have been found in Late Cretaceous sediments well
north of the Arctic Circle. Conditions much warmer than those of today lasted
into the Eocene.
c. Cenozoic
i. Paleogene

1. Paleocene (65.5- 55.8 mya)

Defining Characteristics:
o Age of Mammals begins
o flowering plants and conifers abundant
Secondary Characteristics:
o global climate warming
o abundant plants, fish, crocodiles, and mammals

On the basis of fossil plants, W. P. Schimper separated the lower


part of Lyell's Eocene and named it Paleocene in 1874. The name is
derived from the Greek words palaios, meaning ancient or old, and
kainos, meaning recent. Schimper chose this name to describe the
oldest epoch of the recent Paleogene Period, marking the onset of the
Cenozoic Era. Paleocene rocks are common in the western United States,
South America, western Europe, and eastern Asia.

At the onset of the Paleocene Epoch, Earth was recovering from


the end-Cretaceous asteroid impact. The climate was subtropical almost
to the polar circles, ocean temperatures were high, and polar ice caps
were absent. The oceans invaded many coastal plain areas, as well as
some continental interiors, but mountain-building forced these seas to
retreat. Land bridges existed between North America, Asia, and Europe,
while South America and Antarctica remained connected to each other.
Africa, Australia, and India were island continents or subcontinents.
Mammals began to take advantage of the niches left empty by the
extinction of the dinosaurs, evolving into many new species. Abrupt
warming at the end of the Paleocene followed the release of a large
volume of methane contained in seafloor sediments. This led to a major
extinction of deep-sea foraminiferans and a major reorganization of many
terrestrial and marine communities.

Terrestrial Life through the Paleocene

With the demise of the dinosaurs at the end of the Cretaceous,


the stage was set for mammals and birds to become the dominant land

vertebrates. Although Paleocene bird fossils are rare, mammals are well
represented in Paleocene sediments. Insectivorans, early relatives of true
primates (plesiadapiforms), carnivorans, creodonts, and primitive
herbivores (such as condylarths and early uintatheres) inhabited the
forests. The largest mammal, Pantolambda (a primitive plant eater), was
about the size of a small pony. The multituberculates, small mammals with
chisel-like front teeth that had evolved in the Mesozoic, remained common
in the Paleocene; rodents appeared late in the Paleocene.
Through most of the Paleocene, fossil leaves show low amounts
and few types of damage caused by herbivorous insects. This suggests
that insect herbivores were slow to recover diversity and abundance
following the end-Cretaceous extinction. The diversity of insect feeding
marks on plants did not recover to Cretaceous levels until the late
Paleocene.

Life in the Paleocene Oceans

The marine world of the Paleocene was much more like the
modern marine realm than that of the Cretaceous. In particular, some of
the largest and most common Mesozoic animals had vanished. Although
many Cretaceous species of invertebrates and fishes survived the
Cretaceous extinction event, among aquatic reptiles only turtles,
crocodilians, and champsosaurs (a freshwater, crocodile-like fish eater)
persisted into the Paleocene. Sharks are represented by mackerel sharks,
several genera of sand tiger sharks, and the first small-toothed white
shark. Bony fishes, especially teleosts, became more common as well.
New forms of sea urchins and foraminiferans appeared, alongside more
modern forms of gastropods and bivalves

Climates and Shifting Continents

The continents were mostly separate from one another in the


Paleocene. India was moving north toward southern Asia and would begin
to collide with it during the late Paleocene. Australia had separated from
Antarctica and was moving north. A seaway separated North America from
South America, connecting the Atlantic with the Pacific and Indian Oceans.
The Atlantic Ocean continued widening. South America remained
connected to Antarctica by a narrow peninsula. North America was
connected to Asia by the Bering land bridge, and to Europe by way of
Ellesmere Island and Greenland as well as areas now submerged under
the North Sea. As in the Cretaceous, the Tethys Ocean separated Europe
from Africa. The uplift of the Rocky Mountains signaled the final retreat of
North America's mid-continent seaway, but the ocean still covered parts of
Washington and Oregon, as well as the Atlantic and Gulf coastal plains

Abrupt warming at the end of the Paleocene followed the release


of a large volume of methane contained in seafloor sediments. This led to
the extinction of over 50% of the species of deep-sea foraminiferans. A
subtropical "greenhouse" climate existed nearly worldwide, with
maximum warmth occurring at 55 million years ago.

2. Eocene (55.8- 33.9 mya)

Defining Characteristics:
o first appearances of many modern mammal orders
o maximum extent of warm climate and tropical vegetation
Secondary Characteristics:
o evolution of marine mammals
o cooling climate


In 1833, Charles Lyell derived the name Eocene from the Greek
words Eos (meaning dawn) and Kainos (meaning recent). At the time,
the Paleocene had not yet been named, so the Eocene became the dawn
of the recent (Cenozoic Era). Lyell chose this term because only about
3.5% of fossil mollusks from sediments of this age were recent species.
During the Eocene, volcanoes were active in the Rocky Mountains as the
uplift of this region was completed. The rising mountains were eroded into
sediment that filled the adjacent basins, which (along with nearby large
lakes) became important fossil sites in Wyoming and Colorado.
Spectacular Eocene fossils come from lake deposits at Messel, in
Germany, and from the Green River Formation in southwestern Wyoming.

A dramatic warming event occurred at the onset of the Eocene,


probably due to the release of methane that had been trapped in
sediments on the ocean floor. In fact, the first 5 million years of the
Eocene were warmer than any other time in the Cenozoic. Polar-region
fossils include warm-weather species of plants, alligators, turtles, and
flying lemurs. However, after the middle of the Eocene the climate
became cooler and drier, a trend that continued for the rest of the
Cenozoic Era. Throughout the epoch, mammals continued their rapid postCretaceous diversification. Giant titanothere herbivores, the first whales
and sea cows, numerous hoofed mammals, primates, and rodents
populated the landscapes.

Terrestrial Life during the Eocene

During the warmest part of the early Eocene, palm trees grew as
far north as Alaska and Spitsbergen Island in the North Atlantic.
Crocodilians lived above the Arctic Circle, and forests of dawn redwoods
grew at 80 N latitude. As climate cooled and became more seasonal
during the middle and late Eocene, forests gave way to dry woodlands,
perhaps with open patches of grasses and herbs. The Eocene saw changes
in the distribution of plants, as some species occupied new geographic
regions. Most of the modern bird orders were present by the Eocene, as
well as several unusual, now-extinct species. One, the North American
Diatryma, was a flightless, six-foot-tall predator.

Herbivorous mammals were browsers, feeding on soft


vegetation such as leaves and herbs, or selectively feeding on fruits and
seeds. The first odd-toed hoofed mammals (perissodactyls, such as
horses, tapirs, and rhinos) appeared at the beginning of the Eocene, as did
the first even-toed hoofed mammals (artiodactyls, which included pig-like
omnivores and camels). The largest mammals of the Eocene were the
titanotheres; many species of these browsing perissodactyls lived in North
America and Eurasia, but all were extinct by the end of the epoch. Another
group of large hoofed mammals, the uintatheres, also evolved very large
body size in the Eocene.

Marine Life in the Eocene

One of the most remarkable aspects of the Eocene marine fossil


record is the presence of the first whales. These earliest fossil whales,
from rocks in Pakistan and India, suggest that whales are closely related to
the even-toed ungulates (artiodactyls). Both artiodactyls and early whales
have an unusual feature in that one of the ankle bones (the astragalus) is
shaped like a double pulley. Archaic whales such as Ambulocetus and
Pakicetus demonstrate the transition from a terrestrial to an aquatic way
of life. Later in the Eocene, whales and sea cows (sirenians) adapted more
completely to ocean life and spread worldwide. One type of archaic whale
(Basilosaurus) achieved lengths of 60 feet.


Among the cartilaginous fishes, modern forms such as requiem
sharks increased, while sand tiger sharks decreased. By the Middle
Eocene, the giant sand tiger shark Otodus obliquus was extinct. Mako and
giant-toothed white sharks first appeared at this time as well. However,
bony fishes continued to dominate the seas, as they do today. Marine
invertebrates were also more modern. Paleozoic Fauna animals such as
brachiopods were uncommon, while cephalopods, echinoderms, snails,
and bivalves thrived.

Shifting Continents and Changing Climates

A brief warming event occurred at the start of the Eocene, as


methane was released from ocean-floor sediments. Over a few tens of
thousands of years global temperatures rose as a result of the methane
release, then cooled again in the succeeding 100,000-200,000 years. Even
after this geologically brief interval of very warm climate, subtropical
climates existed up to relatively high northern latitudes. These areas had
much greater rainfall than today, and less seasonal change in
temperature. Carbonate reefs existed in the Bahamas and from Florida to
North Carolina. From the middle Eocene onward the climate cooled and
became drier. At the end of the epoch circumpolar current formed in the
Southern Ocean because of the northward movement of South America.
The circumpolar current helped to isolate Antarctica and probably was an
important factor in the early development of the South Polar Ice Cap. The
cooling of Antarctica in turn meant that cold water began to flow north
along the ocean bottoms from high southern latitudes. These changes in
ocean circulation brought an end to the long greenhouse climate that had
existed since the Mesozoic.

3. Oligocene (33.9- 23.03 mya)

Defining Characteristics:
o transition of woodlands to open grasslands
o appearance of most living families of mammals
o appearance of the two orders (toothed and baleen) of living
whales
o initiation of North Atlantic deep-water formation
Secondary Characteristics:
o development of the Antarctic circumpolar current
o erosion of the Rockies
o decline of browsing mammals

In 1854, Heinrich Ernst von Beyrich defined a new epoch of the


early Paleogene on the basis of fossil-bearing sediments in Belgium and
northern Germany. Sediments of this age were not well represented in
Charles Lyell's British study areas, and he had called these strata the
older Miocene. Von Beyrich derived the name Oligocene from the Greek
words oligos (meaning few or scanty) and kainos (meaning recent).
The name referred to the fact that few modern fossils were found in
Oligocene rocks.

During the Oligocene in central North America, wooded


grasslands gave way to open grasslands with trees only along
watercourses. Volcanic activity continued in the Rocky Mountain region
and along the West Coast. Browsing mammals declined while grazers
increased. Some herbivorous mammals evolved longer limbs, enabling
them to run faster in open country. The seas retreated from most coastal
regions. At the beginning of this epoch, South America continued to
separate from Antarctica. This allowed the development of a circumpolar
current that insulated Antarctica from warm water currents. The ice cap

that had begun to form in the late Eocene expanded, leading to a


significant cooling of global climate.

Terrestrial Life Throughout the Oligocene

Drier climates in the interior of North America led to fewer sites


where plant fossils were preserved, but there is some evidence for more
open forests. Grasslands may have developed in some areas but were
probably not widespread. Along the west coast of North America,
relatively diverse forests of broad-leaved trees and conifers were common.
In higher northern latitudes, forests consisted of broad-leaved deciduous
trees and conifers.

Ungulates were remarkably diverse in the Oligocene. Among the


even-toed forms (artiodactyls), oreodonts (imagine an animal with pig-like
feet but molar teeth more like those of a cow) were very abundant in
North America. Pig-like entelodonts roamed the continents alongside
ruminants such as camels. Odd-toed ungulates (perissodactyls) included
herds of three-toed horses and many forms of rhinoceros. These diverse
Oligocene rhinos included agile running forms, and in Asia, the largest
land mammals of all, the giant indricotheres. Carnivorans included early
members of the dog and cat families, including saber-toothed cats. South
America, which had been isolated from other continents for millions of
years, developed a unique fauna that included edentates (armadillos and
sloths), predatory marsupials (borhyaenids), and giant carnivorous ground
birds (phorusrhacids).

Oligocene Marine Life

Life in the Oligocene oceans looked increasingly familiar. Fossil


pinnipeds (seals and sea lions) first appeared in the late Oligocene seas,
looking much like their modern descendants. Whales had evolved into the
two suborders present today, the toothed whales (odontocetes) and
baleen whales (mysticetes). These marine mammals lived alongside sea
cows and increasingly modern shark species. Bony fishes thrived as well.
Marine invertebrates, too, took on a more familiar appearance.
In the warm tropical seas, carbonate shell reefs extended as far north as
North Carolina. Similarly, coral reefs developed in the Southern
Hemisphere as far south as northern New Zealand. Nummulitid
foraminiferans were common, and the first irregular echinoids (sea
urchins) were present. Other major invertebrate groups included
cephalopods (squid and octopus), bivalves, snails, and crustaceans.

Shifting Continents and Climates

Tectonic activity during the Oligocene caused great changes in


ocean currents. The separation of the Shetland platform from the Faeroe
platform allowed Arctic cold water to flow southward. The separation of
South America from Antarctica that had begun at the end of the Eocene
allowed the formation of a circumpolar current, which insulated Antarctica
from warm-water currents. India continued pushing up the Himalaya
Mountains, causing the first isolation of the Paratethys Ocean, which
occupied central and northern Europe. In turn, this led to low-oxygen
conditions at basin bottoms, reduced salinity, and marine faunas that
were particular to this area. As a result, the mollusks of the Paratethyan
basin began a process of very rapid evolution. At this time the African
plate continued to constrict the Mediterranean basin, but it still had an

eastern opening to the Indian Ocean. The Arabian peninsula and Iran
pushed into Asia Minor. The Atlantic continued to widen.
ii. Neogene

1. Miocene (23.03- 5.332 mya)

Defining Characteristics:
o half of marine invertebrates species are extant forms
o uplift of land separates Tethys Ocean from Indian Ocean
o ocean circulation changes to form major gyres (circles) in N &
S hemispheres
o more seasonal climates in N. Hemisphere
Secondary Characteristics:
o horses and even-toed herbivorous mammals diversify
o whales, seals, sea lions, and walruses diversify

This epoch was named Miocene by Charles Lyell in 1833 for


strata at various European localities. He based this name on his
observation that about one-fifth of the mollusk species found in these
sediments represented living species. The name Miocene is derived from
the Greek words meion (meaning less) and kainos (meaning recent),
referring to the fact that fewer recent species were found in these rocks
than in those of more recent age. (North Carolina).

The Miocene is one of the longest epochs of the Cenozoic Era


and therefore forms a substantial part of the Neogene. It is most noted for
the formation of wide expanses of open grassland across North America
and Eurasia. These open habitats were home to a diversity of new forms
of mammals; in particular, horses, rhinoceros, camels, and antelope-like
mammals were common throughout North America. In the seas, great
changes in ocean circulation encouraged the evolution and spread of
marine vertebrates, including a great diversity of whales, seals, and sea
lions. Distinct continental faunas were characteristic of this period, but so
were migrations between many of the landmasses.

Terrestrial Life Throughout the Miocene

Horses had begun to diversify early in the Neogene, but most of


these species were browsing animals with low-crowned teeth. Their short
legs were well suited to walking through closed, forest-like environments.
However, during the middle Miocene, one group of these horses evolved
durable teeth that allowed them to graze (eat grasses); their high tooth
crowns were better able to withstand the tough, dusty grit on many
grasses. Other lines of browsing horses became extinct before the end of
the Miocene. Grazing horses diversified into several different forms, but
only two of these survived past the Miocene: a three-toed lineage that
became extinct in the ice ages, and a single-toed lineage that gave rise to
modern horses and zebras.

In the Great Plains and Asian steppes, grasses dominated the


landscape, with forests confined to stream courses and wet areas. In
wetter regions such as eastern North America, forests were dominant.
Grazers became more common than browsers among the mammals, with
ruminants enjoying particular success. Elephant-like gomphotheres were
the largest Miocene land mammals in North America. Although horned and
hornless ruminant mammals also diversified, many species became
extinct by the end of the Miocene. Deer and giraffes flourished alongside
early mastodont elephants. The odd chalicotheres, clawed perissodactyl

ungulates, also appeared, as did our own near ancestors in Africa, the first
anthropoid apes.

Miocene Marine Life

In the seas, marine crocodiles were still common and lived as far
north as Maryland. Many species of marine mammals also inhabited the
seas, including baleen and toothed whales, seals, sea lions, walruses, and
sea cows. Both toothed and baleen whales were particularly diverse
during this period. One odd marine mammal that had appeared in the
Oligocene and was common in the North Pacific Ocean during the Miocene
was called Desmostylus. It had four legs that were well adapted both for
living on land and for swimming, and its teeth were shaped like bundles of
cylinders. Distant relatives of elephants and sea cows, Desmostylus and
its kin were entirely extinct by the end of the Miocene.

Some of these marine mammals were undoubtedly preyed upon


by the extinct giant-toothed white shark, Carcharodon megalodon. This
giant shark is well-known from fossils along the east coast of North
America. In fact, by this time most types of living sharks were present in
all the world's seas. Many of the marine invertebrates of the Miocene were
similar to those present today-echinoderms, snails, bivalves, crustaceans,
and cephalopods were common. Marine animals of the ancient Paleozoic
Fauna, such as brachiopods and crinoids, were rare. The first kelp forests
are known from this time.

Shifting Continents and Changing Climates

The Miocene began with a warming of the climate, before the


general Cenozoic cooling trend continued. The Atlantic Ocean continued
widening. The northward movement of Africa, the Arabian peninsula, and
the Indian subcontinent continued pushing up the Alps and the Himalaya
Mountains. Mountain-building cut off the remnants of the Tethys Ocean
from the Indian Ocean, and by the end of the epoch, Gibraltar was
connected to Africa. Thus the Tethys Ocean was separated at both its
eastern and western outlets and confined to the Mediterranean Basin. The
modern Mediterranean Sea is the last remnant of this once-great ocean.

With the complete closure of the Tethys, circumglobal circulation


of warm waters ceased. In the oceans, great gyres or circular currents
brought warm water toward the poles and cold water toward the equator.
Fishes and whales may have used these currents to migrate seasonally
around the ocean basins. Faunal interchange between Asia and North
America continued across the Bering Land Bridge, but by the late Miocene
the bridge had been flooded. Essentially modern patterns of ocean and
atmospheric circulation developed in the Miocene and the major
landmasses came close to their present-day positions.

2. Pliocene (5.332- 2.588 mya)

Defining Characteristics:
o continuation of Age of mammals
o appearance of early bipedal ancestors of humans
o Great American Interchange
Secondary Characteristics:
o expansion of grasslands
o beginning of separation between Atlantic and Pacific marine
organisms

The Pliocene is the final epoch of the Neogene Period. It was


named by Charles Lyell in 1830, after the Greek words pleion (meaning
more) and kainos (meaning recent). The Pliocene represents the

beginning of the familiar modern faunas; many of the plants and animals
from that time have continued to live and evolve to the present day.
Indeed, Lyell devised the name because he noticed that of all the
Cenozoic marine fossils, those from Pliocene strata were the most similar
to modern forms. This characteristic distinguished these layers from
earlier Miocene or Eocene strata.

During this part of the Age of Mammals, the global climate


underwent cyclic variations from cool and dry to warm and wet, then back
to cool conditions again. Pliocene aquatic fossils are numerous, and they
range from tropical marine life to freshwater fish and invertebrates. They
have been found in many places around the world, including Ahl al
Oughlam (Morocco), Bodjong (Indonesia), and the Salada Formation
(Mexico). Dry, open grasslands and savannas were prevalent on land.
Plants very similar to those in modern forests occupied river valleys and
other habitats. In North America, abundant fossils of mastodons,
mammoths, and three-toed horses have been found in Kettleman Hills
(California), Hemphill Beds (Texas), and the Hagerman Fossil Beds National
Monument (Idaho).

Life in the Pliocene

During the Pliocene, the Panamanian land bridge linked North and
South America, allowing terrestrial species to migrate between the two
continents. This event is called the Great American Faunal Interchange, a
time when two long-isolated faunas came into contact. Sixteen native
southern genera moved to the north, including armadillos, giant ground
sloths, flightless predatory birds, marsupials (including opossums), and
porcupines. At the same time, 23 native northern genera moved south,
including cats, dogs, bears, tapirs, camels, and certain rodents. The
exchange was not a balanced one, however. Many more South American
immigrant species became extinct, perhaps as a result of competition and
the inability to adapt to new conditions. As a result, many North American
species now live in South America, but few South American imports still
survive in the north.

Tectonics during the Pliocene

The Pliocene began with the catastrophic origin of the


Mediterranean Sea. The Mediterranean Basin had been dry since the
Miocene, replaced by grasslands. However, at the beginning of the
Pliocene it was reflooded when a tectonic barrier near the Straits of
Gibraltar was breached, allowing water from the Atlantic Ocean to pour
into the basin, probably catastrophically. North and South America were
connected 3.4 million years ago by the formation of the Panamanian
Isthmus, due to the eastward movement of the Caribbean plate. This
closed the Balboa Portal between the Atlantic and Pacific Oceans, creating
significant changes in the marine environment by separating these two
oceans.

The continents were in nearly their present-day positions by the


Pliocene. The Indian plate continued to move northward under the Asian
plate, further elevating the Himalaya Mountains. Other areas of mountainbuilding around the old Tethys Ocean involved the Caucasus region in
Asia, which also generated large orographic changes in climate due to the
rain-shadow effect. In North America, the Cascades, Rockies, and
Appalachian Mountains experienced continued or renewed uplift, along
with the Colorado Plateau. The Sierra Nevada and Alaskan Ranges were
forming as well. Mountain-building was also taking place in Europe,
including the Alps. All these orogenies affected global climate and worked

with astronomically controlled climate cycles to create cooler and more


unstable conditions relative to the Miocene.

Climate Cycles during the Pliocene

Current research on Earth's orbital motions has resulted in


detailed correlation of the sedimentary layers that were deposited in
cycles during this time. These cycles are obvious in some rock strata and
turn out to accurately record climate fluctuations caused by the orbital
variations of the Earth. For example, paleoclimate research has shown
that there was a warming phase at middle and high latitudes during the
middle Pliocene (between 3.15 and 2.85 million years ago), accompanied
by relatively stable tropical temperatures. This information is based on
analysis of microfossils from deep-ocean cores, which give estimates of
ancient sea surface temperatures. These can be used along with pollen
samples from cores taken on land.
iii. Quaternary

1. Pleistocene (2.588 mya- 11,700 yrs)

Defining Characteristics:
o significant human geographic expansion and cultural
development
o marked climatic fluctuations and glacial events
Secondary Characteristics:
o plants and animals very similar to surviving modern forms
o first major human-influenced extinctions

The term Pleistocene is derived from two Greek words, pleistos


(meaning most) and kainos (meaning new or recent). Sir Charles
Lyell introduced this term in 1839 to describe strata in Sicily that had at
least 70% of their molluscan fauna still living today. This distinguished it
from the older Pliocene Epoch, which Lyell had originally thought to be the
youngest fossil rock layer. It represents the first epoch of the Neogene
Period, which is itself the last of the Cenozoic Era.

The Pleistocene is a unique epoch because it is the period during


which our own species, Homo sapiens, evolved. It is also marked by
climatic fluctuations that culminated in widespread continental glaciers.
Many species of vertebrates, especially large mammals, went extinct
during the Pleistocene, but much of the modern flora and fauna are
survivors from this epoch.

Pleistocene Glacial Events

The Pleistocene geological record gives evidence of 20 cycles of


advancing and retreating continental glaciers, though during most of the
Pleistocene glaciers were far more extensive than they are today. Much of
this glaciation occurred at high latitudes and high altitudes, especially in
the Northern Hemisphere. Up to 30% of the Earth's surface was glaciated
periodically during the Pleistocene. Large portions of Europe, North
America (including Greenland), South America, all of Antarctica, and small
sections of Asia were entirely covered by ice. In North America during the
peak of the Wisconsinan glaciation approximately 18,000 years ago, there
were two massive yet independent ice sheets. Both the eastern
Laurentide and the western Cordilleran ice sheets were over 3900 meters
thick. In Europe, ice covered Scandinavia, extended south and east across
Germany and western Russia, and southwest to the British Isles. Another
ice sheet covered most of Siberia. In South America, Patagonia and the
southern Andes mountains were beneath part of the Antarctic ice sheet.

Because so much water was taken up as ice, global sea level dropped
approximately 140 meters.

The causes of the Pleistocene cycle of glacial and interglacial


episodes are still being debated. It appears that continental positions,
oceanic circulation, solar-energy fluctuations, and Earth's orbital cycles
combined to generate these glacial conditions, so perhaps it is
inappropriate to pinpoint any single cause. Some scientists have
calculated that changes in the concentration of greenhouse gases were a
partial reason for large (5-7 C) global temperature swings between the
ice ages and interglacial periods.

Pleistocene Ecosystems and Extinctions

Glacial cycles were not the only geological and climatic


characteristics of the Pleistocene. Volcanic activity was also occurring in
the rift valleys of Africa and in western North and South America. In
southwestern North America, the Colorado River began to carve out the
Grand Canyon.

The Pleistocene was also a time of extinction. By the end of this


epoch, many species of mammals had gone extinct in North America,
including llamas and camels, tapirs and horses, and musk oxen. In
addition, other large mammals such as mammoths and mastodons, sabertooth cats, and ground sloths went completely extinct. Similar extinctions
of large mammals occurred in Australia and South America as well. These
extinctions are a source of active research and controversy. Many
scientists believe that human migrations were an important factor in the
extinctions of large mammals, especially in North America and Australia.
However, the dramatic climate changes that were also occurring may
have been a factor as well.

Human Evolution during the Pleistocene

The evolution of Homo erectus occurred during the Pleistocene,


and fossils of this species have been found in Java, China, Europe, and
throughout Africa. In addition, late Pleistocene fossils discovered in the
Neander valley near Dusseldorf, Germany, represent Neanderthals (Homo
neanderthalensis), a species close to modern humans. Along with
paleoecological information and associated artifact technologies such as
Acheulean and Mousterian cultures (anthropologists call these the
Paleolithic cultures). These fossils provide evidence about Pleistocene
landscapes, climates, and environments. Fossil humans from Dmanisi,
Georgia (west-central Asia), have been dated to the early Pleistocene and
appear to represent the earliest migration of humans out of Africa. There
may have been multiple such migrations. Regardless, by the late
Pleistocene, early modern humans (Homo sapiens) had spread to Australia
and the Americas.

2. Holocene (11,700 yrs- present)

Defining Characteristics:
o impact of Homo sapiens and technology
Secondary Characteristics:
o climate warming following the last ice age
o continents drying out, polar areas contract
o plant communities shifting with the climate


The term Holocene was first proposed at the third International
Geological Congress in 1885. It comes from the Greek words holos
(meaning whole) and kainos (meaning recent), referring to fact that
this epoch is the most recent division of Earth history. However, many
scientists also used the term Recent or Postglacial for this epoch until
1967, when the U.S. Geological Survey formally adopted the term
Holocene and discontinued the use of Recent.

The Holocene is a chronostratigraphic division that follows the


Pleistocene Epoch. By consensus, it covers the last 11,500 years of Earth
history. It is an important time to scientists because during this epoch
most of our modern landscapes and soils evolved. In addition, significant
changes in global climate occurred as the Earth moved into a postglacial
or interglacial regime. In areas that were glaciated, this transition is
marked by a clear stratigraphic boundary due primarily to the scouring
effects and debris that retreating glaciers left on the landscape.

Climate Change and Variability

A series of climatic changes has occurred throughout the


Holocene. In fact, scientists have recorded up to 18 climatic cycles during
this time. Since the Bronze Age (starting about 3500 BC), it has become
increasingly challenging to separate natural climatic trends from humaninduced effects on the environment. Geologists and climatologists
separate the climatic history of the Holocene into three temperate
substages that relate mainly to the Northern Hemisphere (North America
and Eurasia). These include an early cool stage (about 10,000 BC) a
middle climatic optimum (also termed the altithermal, 9000-4000 BC),
and a late stage cooling that contains the little ice ages (starting about
AD 1300). A xerothermic or hypsithermal substage also exists between
these last two stages.

Three other factors must be added to this: complex solar cycles,


Earth's orbital variations (also called Milankovitch cycles, which occur over
intervals of tens to hundreds of thousands of years), and different rates of
change and climatic conditions depending on location. The result is a
complicated picture of the transition from glacial to postglacial conditions.
For example, the retreat of the continental glaciers of the Wisconsinan
stage began in central North America around 10,000 years ago, but it did
not occur in northern Canada and Alaska until nearly 6000 years ago. The
retreat has yet to occur in Greenland. Sea level rise from the melting of
the glaciers affected coastal areas globally, so much so that in the late
1800s some scientists believed that this sea level rise should be the
defining characteristic of the Holocene.

Impact of Human Development

The Holocene has another unique characteristic as well. It is


during this time that humans become an important factor in the processes
of Earth history. Human development has already left a lasting imprint on
the geologic record, and it will likely continue to do so for some time. To
describe these effects, geologists use terms such as anthropogenic
sediment and anthropogenic geomorphology. Examples include
changes in sediment deposition from dam-building and river diversion,
leveling of topographic highs and infilling of lows, as well as attempted
stabilization of variable features such as barrier islands.

Most of these processes continue today, often at an accelerated


pace. As humans increasingly travel between continents, formerly isolated
animals and plants have opportunities to come into contact with one
another. Often the results are devastating, particularly for island

organisms that have few predator defenses. Anthropogenic changes in


Earth's atmosphere are ongoing, with most of the impact yet to be felt.