History of the Earth

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For the history of modern humans, see History of the world.

Geological time put in a diagram called a geological clock, showing the relative lengths
of the eons of the Earth's history.
The history of the Earth describes the most important events and fundamental stages in
the development of the planet Earth from its formation 4.6 billion years ago to the present
day.[1] Nearly all branches of natural science have contributed to the understanding of the
main events of the Earth's past. The age of Earth is approximately one-third of the age of
the universe.[2] Immense geological and biological changes have occurred during that
time span.

Contents
[hide]

1 Hadean and Archaean

o 1.1 Origin of the solar system
o 1.2 Origin of the Earth's core and first atmosphere
o 1.3 The giant impact hypothesis
o 1.4 Origin of the oceans and atmosphere
o 1.5 The first continents
o 1.6 Origin of life
2 Proterozoic eon
o 2.1 The oxygen revolution

2.2 Snowball Earth and the origin of the ozone layer

2.3 Proterozoic development of life
2.4 Rodinia and other supercontinents
2.5 Late Proterozoic climate and life
3 Paleozoic era
o 3.1 Cambrian explosion
o 3.2 Paleozoic tectonics, paleogeography and climate
o 3.3 Colonization of land
4 Mesozoic
5 Cenozoic era (Recent life)
o 5.1 Human evolution
o 5.2 Civilization
o 5.3 Recent events
6 See also
7 Notes
o 7.1 References
o 7.2 Literature
o
o
o
o

7.3 External links

Hadean and Archaean

Main articles: Hadean and Archaean
Starting with the Earth's formation by accretion from the solar nebula 4.54 billion years
ago (4.54 Ga),[1] the first eon in the Earth's history is called the Hadean.[3] It lasted until
the Archaean eon, which began 3.8 Ga. The oldest rocks found on Earth date to about 4.0
Ga, and the oldest detrital zircon crystals in some rocks have been dated to about 4.4 Ga,
[4]
close to the formation of the Earth's crust and the Earth itself. Because not much
material from this time is preserved, little is known about Hadean times, but scientists
hypothesize at an estimated 4.53 Ga,[nb 1] shortly after formation of an initial crust, the
proto-Earth was impacted by a smaller protoplanet, which ejected part of the mantle and
crust into space and created the Moon.[6][7][8]
During the Hadean, the Earth's surface was under a continuous bombardment by
meteorites, and volcanism must have been severe due to the large heat flow and
geothermal gradient. The detrital zircon crystals dated to 4.4 Ga show evidence of having
undergone contact with liquid water, considered as proof that the planet already had
oceans or seas at that time.[4] From crater counts on other celestial bodies it is inferred
that a period of intense meteorite impacts, called the "Late Heavy Bombardment", began
about 4.1 Ga, and concluded around 3.8 Ga, at the end of the Hadean.[9]
By the beginning of the Archaean, the Earth had cooled significantly. It would have been
impossible for most present day life forms to exist due to the composition of the
Archaean atmosphere, which lacked oxygen and an ozone layer. Nevertheless it is

believed that primordial life began to evolve by the early Archaean, with some possible
fossil finds dated to around 3.5 Ga.[10] Some researchers, however, speculate that life
could have begun during the early Hadean, as far back as 4.4 Ga, surviving the possible
Late Heavy Bombardment period in hydrothermal vents below the Earth's surface.[11]

Origin of the solar system

An artist's impression of protoplanetary disk.

Main article: Formation and evolution of the Solar System
The Solar System (including the Earth) formed from a large, rotating cloud of interstellar
dust and gas called the solar nebula, orbiting the Milky Way's galactic center. It was
composed of hydrogen and helium created shortly after the Big Bang 13.7 Ga and heavier
elements ejected by supernovas.[12] About 4.6 Ga, the solar nebula began to contract,
possibly due to the shock wave of a nearby supernova. Such a shock wave would have
also caused the nebula to rotate and gain angular momentum. As the cloud began to
accelerate its rotation, gravity and inertia flattened it into a protoplanetary disk oriented
perpendicularly to its axis of rotation. Most of the mass concentrated in the middle and
began to heat up, but small perturbations due to collisions and the angular momentum of
other large debris created the means by which protoplanets up to several kilometres in
length began to form, orbiting the nebular center.
The infall of material, increase in rotational speed and the crush of gravity created an
enormous amount of kinetic heat at the center. Its inability to transfer that energy away
through any other process at a rate capable of relieving the build-up resulted in the disk's
center heating up. Ultimately, nuclear fusion of hydrogen into helium began, and
eventually, after contraction, a T Tauri star ignited to create the Sun. Meanwhile, as
gravity caused matter to condense around the previously perturbed objects outside the
gravitational grasp of the new sun, dust particles and the rest of the protoplanetary disk
began separating into rings. Successively larger fragments collided with one another and
became larger objects, ultimately becoming protoplanets.[13] These included one
collection about 150 million kilometers from the center: Earth. The planet formed about
4.54 billion years ago (within an uncertainty of 1%)[1] and was largely completed within
1020 million years.[14] The solar wind of the newly formed T Tauri star cleared out most
of the material in the disk that had not already condensed into larger bodies.
Computer simulations have shown that planets with distances equal to the terrestrial
planets in our solar system can be created from a protoplanetary disk.[15] The now widely
accepted nebular hypothesis suggests that the same process, which gave rise to the solar

system's planets, produces accretion disks around virtually all newly forming stars in the
universe, some of which yield planets.[16]

Origin of the Earth's core and first atmosphere

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

The giant impact hypothesis

Main articles: Origin and evolution of the Moon and Giant impact hypothesis
The Earth's relatively large natural satellite, the Moon, is unique.[nb 2] During the Apollo
program, rocks from the Moon's surface were brought to Earth. Radiometric dating of
these rocks has shown the Moon to be 4527 10 million years old,[17] about 30 to 55
million years younger than other bodies in the solar system.[18] (New evidence suggests
the Moon formed even later, 4.480.02Ga,

or 70110Ma

after the start of the Solar

System.[5]) Another notable feature is the relatively low density of the Moon, which must
mean it does not have a large metallic core, like all other terrestrial bodies in the solar
system. The Moon has a bulk composition closely resembling the Earth's mantle and
crust together, without the Earth's core. This has led to the giant impact hypothesis, the
idea that the Moon was formed during a giant impact of the proto-Earth with another
protoplanet by accretion of the material blown off the mantles of the proto-Earth and
impactor.[19][8]
The impactor, sometimes named Theia, is thought to have been a little smaller than the
current planet Mars. It could have formed by accretion of matter about 150 million
kilometres from the Sun and Earth, at their fourth or fifth Lagrangian point. Its orbit may
have been stable at first, but destabilized as Theia's mass increased due to the accretion of
matter. Theia oscillated in larger and larger orbits around the Lagrangian point until it
finally collided with Earth about 4.533 Ga.[7][nb 1] Models reveal that when an impactor

this size struck the proto-Earth at a low angle and relatively low speed (820 km/sec),
much material from the mantles (and proto-crusts) of the proto-Earth and the impactor
was ejected into space, where much of it stayed in orbit around the Earth. This material
would eventually form the Moon. However, the metallic cores of the impactor would
have sunk through the Earth's mantle to fuse with the Earth's core, depleting the Moon of
metallic material.[20] The giant impact hypothesis thus explains the Moon's abnormal
composition.[21] The ejecta in orbit around the Earth could have condensed into a single
body within a couple of weeks. Under the influence of its own gravity, the ejected
material became a more spherical body: the Moon.[22]
The radiometric ages show the Earth existed already for at least 10 million years before
the impact, enough time to allow for differentiation of the Earth's primitive mantle and
core. Then, when the impact occurred, only material from the mantle was ejected, leaving
the Earth's core of heavy siderophile elements untouched.
The impact had some important consequences for the young Earth. It released a
enormous amount of energy, causing both the Earth and Moon to be completely molten.
Immediately after the impact, the Earth's mantle was vigorously convecting, the surface
was a large magma ocean. The planet's first atmosphere must have been completely
blown away by the enormous amount of energy released.[23] The impact is also thought to
have changed Earths axis to produce the large 23.5 axial tilt that is responsible for
Earths seasons (a simple, ideal model of the planets origins would have axial tilts of 0
with no recognizable seasons). It may also have sped up Earths rotation.

Origin of the oceans and atmosphere

Because the Earth lacked an atmosphere immediately after the giant impact, cooling must
have occurred quickly. Within 150 million years, a solid crust with a basaltic composition
must have formed. The felsic continental crust of today did not yet exist. Within the
Earth, further differentiation could only begin when the mantle had at least partly
solidified again. Nevertheless, during the early Archaean (about 3.0 Ga) the mantle was
still much hotter than today, probably around 1600C. This means the fraction of partially
molten material was still much larger than today.
Steam escaped from the crust, and more gases were released by volcanoes, completing
the second atmosphere. Additional water was imported by bolide collisions, probably
from asteroids ejected from the outer asteroid belt under the influence of Jupiter's gravity.
The large amount of water on Earth can never have been produced by volcanism and
degassing alone. It is assumed the water was derived from impacting comets that
contained ice.[24]:130-132 Though most comets are today in orbits farther away from the Sun
than Neptune, computer simulations show they were originally far more common in the
inner parts of the solar system. However, most of the water on Earth was probably
derived from small impacting protoplanets, objects comparable with today's small icy
moons of the outer planets.[25] Impacts of these objects can have enriched the terrestrial
planets (Mercury, Venus, the Earth and Mars) with water, carbon dioxide, methane,

ammonia, nitrogen and other volatiles. If all water on Earth was derived from comets
alone, millions of comet impacts would be required to support this theory. Computer
simulations illustrate that this is not an unreasonable number.[24]:131
As the planet cooled, clouds formed. Rain created the oceans. Recent evidence suggests
the oceans may have begun forming by 4.2 Ga,[26] or as early as 4.4 Ga.[4] In any event, by
the start of the Archaean eon the Earth was already covered with oceans. The new
atmosphere probably contained water vapor, carbon dioxide, nitrogen, and smaller
amounts of other gases.[27] As the output of the Sun was only 70% of the current amount,
significant amounts of greenhouse gas in the atmosphere most likely prevented the
surface water from freezing.[28] Free oxygen would have been bound by hydrogen or
minerals on the surface. Volcanic activity was intense and, without an ozone layer to
hinder its entry, ultraviolet radiation flooded the surface.

Lithified stromatolites on the shores of Lake Thetis (Western Australia). Stromatolites are
formed by colonies of single celled organisms like cyanobacteria or chlorophyta. These
colonies of algae entrap sedimentary grains, thus forming the draped sedimentary layers
of a stromatolite. Archaean stromatolites are the first direct fossil traces of life on Earth,
even though little preserved fossilized cells have been found inside them. The Archaean
and Proterozoic oceans could have been full of algal mats like these.

The first continents

Mantle convection, the process that drives plate tectonics today, is a result of heat flow
from the core to the Earth's surface. It involves the creation of rigid tectonic plates at
mid-oceanic ridges. These plates are destroyed by subduction into the mantle at
subduction zones. The inner Earth was warmer during the Hadean and Archaean eons, so
convection in the mantle must have been faster. When a process similar to present day
plate tectonics did occur, this would have gone faster too. Most geologists believe that
during the Hadean and Archaean, subduction zones were more common, and therefore
tectonic plates were smaller.
The initial crust, formed when the Earth's surface first solidified, totally disappeared from
a combination of this fast Hadean plate tectonics and the intense impacts of the Late
Heavy Bombardment. It is, however, assumed that this crust must have been basaltic in
composition, like today's oceanic crust, because little crustal differentiation had yet taken
place. The first larger pieces of continental crust, which is a product of differentiation of

lighter elements during partial melting in the lower crust, appeared at the end of the
Hadean, about 4.0 Ga. What is left of these first small continents are called cratons. These
pieces of late Hadean and early Archaean crust form the cores around which today's
continents grew.
The oldest rocks on Earth are found in the North American craton of Canada. They are
tonalites from about 4.0 Ga. They show traces of metamorphism by high temperature, but
also sedimentary grains that have been rounded by erosion during transport by water,
showing rivers and seas existed then.[24]
Cratons consist primarily of two alternating types of terranes. The first are so called
greenstone belts, consisting of low grade metamorphosed sedimentary rocks. These
"greenstones" are similar to the sediments today found in oceanic trenches, above
subduction zones. For this reason, greenstones are sometimes seen as evidence for
subduction during the Archaean. The second type is a complex of felsic magmatic rocks.
These rocks are mostly tonalite, trondhjemite or granodiorite, types of rock similar in
composition to granite (hence such terranes are called TTG-terranes). TTG-complexes
are seen as the relicts of the first continental crust, formed by partial melting in basalt.
The alternation between greenstone belts and TTG-complexes is interpreted as a tectonic
situation in which small proto-continents were separated by a thorough network of
subduction zones.

Origin of life

The replicator in virtually all known life is deoxyribonucleic acid. DNA is far more
complex than the original replicator and its replication systems are highly elaborate.
Main article: Abiogenesis
The details of the origin of life are unknown, but the basic principles have been
established. There are two schools of thought about the origin of life. One suggests that
organic components arrived on Earth from space (see Panspermia), while the other
argues that they originated on Earth. Nevertheless, both schools suggest similar
mechanisms by which life initially arose.[29]
If life arose on Earth, the timing of this event is highly speculativeperhaps it arose
around 4 Ga.[30] It is possible that, as a result of repeated formation and destruction of
oceans during that time period caused by high energy asteroid bombardment, life may
have arisen and extinguished more than once.[4]
In the energetic chemistry of early Earth, a molecule gained the ability to make copies of
itselfa replicator. (More accurately, it promoted the chemical reactions which produced
a copy of itself.) The replication was not always accurate: some copies were slightly
different from their parent.

If the change destroyed the copying ability of the molecule, the molecule did not produce
any copies, and the line died out. On the other hand, a few rare changes might make the
molecule replicate faster or better: those strains would become more numerous and
successful. This is an early example of evolution on abiotic material. The variations
present in matter and molecules combined with the universal tendency for systems to
move towards a lower energy state allowed for an early method of natural selection. As
choice raw materials (food) became depleted, strains which could utilize different
materials, or perhaps halt the development of other strains and steal their resources,
became more numerous.[31]:563-578
The nature of the first replicator is unknown because its function was long since
superseded by lifes current replicator, DNA. Several models have been proposed
explaining how a replicator might have developed. Different replicators have been
posited, including organic chemicals such as modern proteins, nucleic acids,
phospholipids, crystals,[32] or even quantum systems.[33] There is currently no way to
determine whether any of these models closely fits the origin of life on Earth.
One of the older theories, one which has been worked out in some detail, will serve as an
example of how this might occur. The high energy from volcanoes, lightning, and
ultraviolet radiation could help drive chemical reactions producing more complex
molecules from simple compounds such as methane and ammonia.[34]:38 Among these
were many of the simpler organic compounds, including nucleobases and amino acids,
which are the building blocks of life. As the amount and concentration of this organic
soup increased, different molecules reacted with one another. Sometimes more complex
molecules would resultperhaps clay provided a framework to collect and concentrate
organic material.[34]:39
Certain molecules could speed up a chemical reaction. All this continued for a long time,
with reactions occurring at random, until by chance it produced a replicator molecule. In
any case, at some point, the function of the replicator was superseded by DNA; all known
life (except some viruses and prions) use DNA as their replicator, in an almost identical
manner (see Genetic code).

A small section of a cell membrane. This modern cell membrane is far more sophisticated
than the original simple phospholipid bilayer (the small blue spheres with two tails).
Proteins and carbohydrates serve various functions in regulating the passage of material
through the membrane and in reacting to the environment.

Modern life has its replicating material packaged inside a cellular membrane. It is easier
to understand the origin of the cell membrane than the origin of the replicator, because a
cell membrane is made of phospholipid molecules, which often form a bilayer
spontaneously when placed in water. Under certain conditions, many such spheres can be
formed (see The bubble theory).[34]:40
The prevailing theory is that the membrane formed after the replicator, which perhaps by
then was RNA (the RNA world hypothesis), along with its replicating apparatus and other
biomolecules. Initial protocells may have simply burst when they grew too large; the
scattered contents may then have recolonized other bubbles. Proteins that stabilized the
membrane, or that later assisted in an orderly division, would have promoted the
proliferation of those cell lines.
RNA is a likely candidate for an early replicator, because it can both store genetic
information and catalyze reactions. At some point DNA took over the genetic storage role
from RNA, and proteins known as enzymes took over the catalysis role, leaving RNA to
transfer information, synthesize proteins and regulate the process. There is increasing
belief that these early cells evolved in association with undersea volcanic vents known as
black smokers[34]:42 or even hot, deep rocks.[31]:580
It is believed that of this multiplicity of protocells, only one line survived. Current
phylogentic evidence suggests that the last universal common ancestor lived during the
early Archean eon, perhaps roughly 3.5 Ga or earlier.[35][36] This LUCA cell is the
ancestor of all life on Earth today. It was probably a prokaryote, possessing a cell
membrane and probably ribosomes, but lacking a nucleus or membrane-bound organelles
such as mitochondria or chloroplasts.
Like all modern cells, it used DNA as its genetic code, RNA for information transfer and
protein synthesis, and enzymes to catalyze reactions. Some scientists believe that instead
of a single organism being the last universal common ancestor, there were populations of
organisms exchanging genes in lateral gene transfer.[35]

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

The oxygen revolution

Main article: Oxygen catastrophe

The harnessing of the suns energy led to several major changes in life on Earth.

Graph showing range of estimated partial pressure of atmospheric oxygen through

geologic time [37]

A banded iron formation from the 3.15 Ga Moories Group, Barberton Greenstone Belt,
South Africa. Red layers represent the times when oxygen was available, gray layers
were formed in anoxic circumstances.
The first cells were likely heterotrophs, using surrounding organic molecules (including
those from other cells) as raw material and an energy source.[31]:564-566 As the food supply
diminished, a new strategy evolved in some cells. Instead of relying on the diminishing
amounts of free-existing organic molecules, these cells adopted sunlight as an energy
source. Estimates vary, but by about 3 Ga, something similar to modern oxygenic
photosynthesis had probably developed, which made the suns energy available not only
to autotrophs but also to the heterotrophs that consumed them.[38][39] This type of
photosynthesis, which became by far the most common, used the abundant carbon
dioxide and water as raw materials and, with the energy of sunlight, produced energy-rich
organic molecules (carbohydrates).
Moreover, oxygen was released as a waste product of the photosynthesis.[37] At first, it
became bound up with limestone, iron, and other minerals. There is substantial proof of
this in iron-oxide rich layers in geological strata that correspond with this period. The
reaction of the minerals with oxygen would have turned the oceans green. When most of

the exposed readily-reacting minerals were oxidized, oxygen finally began to accumulate
in the atmosphere. Though each cell only produced a minute amount of oxygen, the
combined metabolism of many cells over a vast time transformed Earths atmosphere to
its current state.[34]:50-51 Among the oldest examples of oxygen-producing lifeforms are
fossil stromatolites. This was Earths third atmosphere.
Some of the oxygen was stimulated by incoming ultraviolet radiation to form ozone,
which collected in a layer near the upper part of the atmosphere. The ozone layer
absorbed, and still absorbs, a significant amount of the ultraviolet radiation that once had
passed through the atmosphere. It allowed cells to colonize the surface of the ocean and
eventually the land:[40] without the ozone layer, ultraviolet radiation bombarding the
surface would have caused unsustainable levels of mutation in exposed cells.
Photosynthesis had another, major, and world-changing impact. Oxygen was toxic;
probably much life on Earth died out as its levels rose in what is known as the "oxygen
catastrophe".[40] Resistant forms survived and thrived, and some developed the ability to
use oxygen to increase their metabolism and obtain more energy from the same food.

Snowball Earth and the origin of the ozone layer

An oxygen-rich atmosphere had two principal advantages for life. Organisms not using
oxygen for their metabolism, such as anaerobe bacteria, base their metabolism on
fermentation. The abundance of oxygen makes respiration possible, a much more
effective energy source for life than fermentation. The second advantage of an oxygenrich atmosphere is that oxygen forms ozone in the higher atmosphere, causing the
emergence of the Earth's ozone layer. The ozone layer protects the Earth's surface from
ultraviolet radiation, which is harmful for life. Without the ozone layer, the development
of more complex life later on would probably have been impossible.[24]:219-220
The natural evolution of the Sun made it progressively more luminous during the
Archaean and Proterozoic eons; the Sun's luminosity increases 6% every billion years.
[24]:165
As a result, the Earth began to receive more heat from the Sun in the Proterozoic
eon. However, the Earth did not get warmer. Instead, the geological record seems to
suggest it cooled dramatically during the early Proterozoic. Glacial deposits found in all
cratons show that about 2.3 Ga, the Earth underwent its first big ice age (the Makganyene
ice age).[41] Some scientists suggest this and following Proterozoic ice ages were so
severe that the planet was totally frozen over from the poles to the equator, a hypothesis
called Snowball Earth. Not all geologists agree with this scenario and older, Archaean ice
ages have been postulated, but the ice age 2.3 Ga is the first such event for which the
evidence is widely accepted.
The ice age around 2.3 Ga could have been directly caused by the increased oxygen
concentration in the atmosphere, which caused the decrease of methane (CH4) in the
atmosphere. Methane is a strong greenhouse gas, but with oxygen it reacts to form CO2, a
less effective greenhouse gas.[24]:172 When free oxygen became available in the

atmosphere, the concentration of methane could have decreased dramatically, enough to

counter the effect of the increasing heat flow from the Sun.

Proterozoic development of life

Main article: Endosymbiotic theory

Some of the pathways by which the various endosymbionts might have arisen.
Modern taxonomy classifies life into three domains. The time of the origin of these
domains is uncertain. The Bacteria domain probably first split off from the other forms of
life (sometimes called Neomura), but this supposition is controversial. Soon after this, by
2 Ga,[42] the Neomura split into the Archaea and the Eukarya. Eukaryotic cells (Eukarya)
are larger and more complex than prokaryotic cells (Bacteria and Archaea), and the origin
of that complexity is only now becoming known.
Around this time, the first proto-mitochondrion was formed. A bacterial cell related to
todays Rickettsia,[43] which had learned how to metabolize oxygen, entered a larger
prokaryotic cell, which lacked that capability. Perhaps the large cell attempted to ingest
the smaller one but failed (possibly due to the evolution of prey defenses). The smaller
cell may have tried to parasitize the larger one. In any case, the smaller cell survived
inside the larger cell. Using oxygen, it metabolized the larger cells waste products and
derived more energy. Some of this excess energy was returned to the host. The smaller
cell replicated inside the larger one. Soon, a stable symbiosis developed between the large
cell and the smaller cells inside it. Over time, the host cell acquired some of the genes of
the smaller cells, and the two kinds became dependent on each other: the larger cell could
not survive without the energy produced by the smaller ones, and these in turn could not
survive without the raw materials provided by the larger cell. The whole cell is now
considered a single organism, and the smaller cells are classified as organelles called
mitochondria.
A similar event occurred with photosynthetic cyanobacteria[44] entering large
heterotrophic cells and becoming chloroplasts.[34]:60-61[31]:536-539 Probably as a result of these
changes, a line of cells capable of photosynthesis split off from the other eukaryotes more

than 1 billion years ago. There were probably several such inclusion events, as the figure
at right suggests. Besides the well-established endosymbiotic theory of the cellular origin
of mitochondria and chloroplasts, it has been suggested that cells led to peroxisomes,
spirochetes led to cilia and flagella, and that perhaps a DNA virus led to the cell nucleus,
[45],[46]
though none of these theories is widely accepted.[47]

Green algae of the genus Volvox are believed to be similar to the first multicellular plants.
Archaeans, bacteria, and eukaryotes continued to diversify and to become more complex
and better adapted to their environments. Each domain repeatedly split into multiple
lineages, although little is known about the history of the archaea and bacteria. Around
1.1 Ga, the supercontinent Rodinia was assembling.[48] The plant, animal, and fungi lines
had all split, though they still existed as solitary cells. Some of these lived in colonies,
and gradually some division of labor began to take place; for instance, cells on the
periphery might have started to assume different roles from those in the interior. Although
the division between a colony with specialized cells and a multicellular organism is not
always clear, around 1 billion years ago[49] the first multicellular plants emerged, probably
green algae.[50] Possibly by around 900 Ma[31]:488 true multicellularity had also evolved in
animals.
At first it probably resembled todays sponges, which have totipotent cells that allow a
disrupted organism to reassemble itself.[31]:483-487 As the division of labor was completed in
all lines of multicellular organisms, cells became more specialized and more dependent
on each other; isolated cells would die.

Rodinia and other supercontinents

Wilson cycle timeline from 1 Ga, depicting Rodinia and Pangaea supercontinent
formation and separation
When the theory of plate tectonics was developed around 1960, geologists began to
reconstruct the movements and positions of the continents in the past. This appeared

relatively easy until about 250 million years ago, when all continents were united in what
is called the "supercontinent" Pangaea. Before that time, reconstructions cannot rely on
apparent similarities in coastlines or ages of oceanic crust, but only on geologic
observations and paleomagnetic data.[24]:95
Throughout the history of the Earth, there have been times when the continental mass
came together to form a supercontinent, followed by the break-up of the supercontinent
and new continents moving apart again. This repetition of tectonic events is called a
Wilson cycle. The further back in time, the scarcer and harder to interpret the data get. It
is at least clear that, about 1000 to 830 Ma, most continental mass was united in the
supercontinent Rodinia.[51] It is highly probable Rodinia was not the first supercontinent,
and many earlier supercontinents have been proposed. This means plate tectonic
processes similar to today's must have been active during the Proterozoic.
After the break-up of Rodinia about 800 Ma, it is possible the continents joined again
around 550 Ma. The hypothetical supercontinent is sometimes referred to as Pannotia or
Vendia. The evidence for it is a phase of continental collision known as the Pan-African
orogeny, which joined the continental masses of current-day Africa, South-America,
Antarctica and Australia. It is extremely likely, however, that the aggregation of
continental masses was not completed, since a continent called Laurentia (roughly
equivalent to current-day North America) had already started breaking off around 610
Ma. It is at least certain that by the end of the Proterozoic eon, most of the continental
mass lay united in a position around the south pole.[52]

Late Proterozoic climate and life

A 580 million year old fossil of Spriggina floundensi, an animal from the Ediacaran
period. Such life forms could have been ancestors to the many new forms that origined in
the Cambrian Explosion.
The end of the Proterozoic saw at least two Snowball Earths, so severe that the surface of
the oceans may have been completely frozen. This happened about 710 and 640 Ma, in
the Cryogenian period. These severe glaciations are less easy to explain than the early
Proterozoic Snowball Earth. Most paleoclimatologists think the cold episodes had
something to do with the formation of the supercontinent Rodinia. Because Rodinia was
centered on the equator, rates of chemical weathering increased and carbon dioxide (CO2)

was taken from the atmosphere. Because CO2 is an important greenhouse gas, climates
cooled globally.
In the same way, during the Snowball Earths most of the continental surface was in
permafrost, which decreased chemical weathering again, leading to the end of the
glaciations. An alternative hypothesis is that enough carbon dioxide escaped through
volcanic outgassing that the resulting greenhouse effect raised global temperatures.[53]
Increased volcanic activity resulted from the break-up of Rodinia at about the same time.
The Cryogenian period was followed by the Ediacaran period, which was characterized
by a rapid development of new multicellular lifeforms. If there is a connection between
the end of the severe ice ages and the increase in diversity of life, is not clear, but it does
not seem coincidental. The new forms of life, called Ediacara biota, were larger and more
diverse than ever. Most scientists think some of them may have been the precursors of the
new life forms of the following Cambrian period. Though the taxonomy of most
Ediacaran life forms is unclear, some are proposed to have been ancestors of groups of
modern life.[54] Important developments were the origin of muscular and neural cells.
None of the Ediacaran fossils had hard body parts like skeletons. These first appear after
the boundary between the Proterozoic and Phanerozoic eons or Ediacaran and Cambrian
periods.

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

Cambrian explosion
Main article: Cambrian explosion
Apparently, the rate of the evolution of life accelerated in the Cambrian period (542-488
Ma). The sudden emergence of many new species, phyla, and forms in this period is
called the Cambrian Explosion. The biological formenting in the Cambrian Explosion
was unpreceded before and since that time.[24]:229 Whereas the Ediacaran life forms appear
yet primitive and not easy to put in any modern group, at the end of the Cambrian most
modern phyla were already present. The development of hard body parts such as shells,

skeletons or exoskeletons in animals like molluscs, echinoderms, crinoids and arthropods

(a well-known group of arthropods from the lower Paleozoic are the trilobites) made the
preservation and fossilisation of such life forms easier than those of their Proterozoic
ancestors.[55] For this reason, much more is known about life in and after the Cambrian
than about that of older periods. The boundary between the Cambrian and Ordovician
(the following period, 488-444 Ma) is characterized by a large mass-extinction, in which
some of the new groups disappeared altogether.[56] Some of these Cambrian groups appear
complex but are quite different from modern life; examples are Anomalocaris and
Haikouichthys.
During the Cambrian, the first vertebrate animals, among them the first fishes, had
appeared.[57] A creature that could have been the ancestor of the fishes, or was probably
closely related to it, was Pikaia. It had a primitive notochord, a structure that could have
developed into a vertebral column later. The first fishes with jaws (Gnathostomata)
appeared during the Ordovician. The colonisation of new niches resulted in massive body
sizes. In this way, fishes with increasing sizes evolved during the early Paleozoic, such as
the titanic placoderm Dunkleosteus, which could grow 7 meters long.

Paleozoic tectonics, paleogeography and climate

At the end of the Proterozoic, the supercontinent Pannotia had broken apart in the smaller
continents Laurentia, Baltica, Siberia and Gondwana. During periods when continents
move apart, more oceanic crust is formed by volcanic activity. Because young volcanic
crust is relatively hotter and less dense than old oceanic crust, the ocean floors will rise
during such periods. This causes the sea level to rise. Therefore, in the first half of the
Paleozoic, large areas of the continents were below sea level.
Early Paleozoic climates were warmer than today, but the end of the Ordovician saw a
short ice age during which glaciers covered the south pole, where the huge continent
Gondwana was situated. Traces of glaciation from this period are only found on former
Gondwana. During the Late Ordovician ice age, a number of mass extinctions took place,
in which many brachiopods, trilobites, Bryozoa and corals disappeared. These marine
species could probably not contend with the decreasing temperature of the sea water.[58]
After the extinctions new species evolved, more diverse and better adapted. They would
fill the niches left by the extinct species.
The continents Laurentia and Baltica collided between 450 and 400 Ma, during the
Caledonian Orogeny, to form Laurussia. Traces of the mountain belt which resulted from
this collision can be found in Scandinavia, Scotland and the northern Appalachians. In
the Devonian period (416-359 Ma) Gondwana and Siberia began to move towards
Laurussia. The collision of Siberia with Laurussia caused the Uralian Orogeny, the
collision of Gondwana with Laurussia is called the Variscan or Hercynian Orogeny in
Europe or the Alleghenian Orogeny in North America. The latter phase took place during
the Carboniferous period (359-299 Ma) and resulted in the formation of the last
supercontinent, Pangaea.

Colonization of land

For most of Earths history, there were no multicellular organisms on land. Parts of the
surface may have vaguely resembled this view of Mars.[citation needed]
Oxygen accumulation from photosynthesis resulted in the formation of an ozone layer
that absorbed much of Suns ultraviolet radiation, meaning unicellular organisms that
reached land were less likely to die, and prokaryotes began to multiply and become better
adapted to survival out of the water. Prokaryotes had probably colonized the land as early
as 2.6 Ga[59] even before the origin of the eukaryotes. For a long time, the land remained
barren of multicellular organisms. The supercontinent Pannotia formed around 600 Ma
and then broke apart a short 50 million years later.[60] Fish, the earliest vertebrates,
evolved in the oceans around 530 Ma.[31]:354 A major extinction event occurred near the
end of the Cambrian period,[61] which ended 488 Ma.[62]
Several hundred million years ago, plants (probably resembling algae) and fungi started
growing at the edges of the water, and then out of it.[63]:138-140 The oldest fossils of land
fungi and plants date to 480460 Ma, though molecular evidence suggests the fungi may
have colonized the land as early as 1000 Ma and the plants 700 Ma.[64] Initially remaining
close to the waters edge, mutations and variations resulted in further colonization of this
new environment. The timing of the first animals to leave the oceans is not precisely
known: the oldest clear evidence is of arthropods on land around 450 Ma,[65] perhaps
thriving and becoming better adapted due to the vast food source provided by the
terrestrial plants. There is also some unconfirmed evidence that arthropods may have
appeared on land as early as 530 Ma.[66]
At the end of the Ordovician period, 440 Ma, additional extinction events occurred,
perhaps due to a concurrent ice age.[58] Around 380 to 375 Ma, the first tetrapods evolved
from fish.[67] It is thought that perhaps fins evolved to become limbs which allowed the
first tetrapods to lift their heads out of the water to breathe air. This would allow them to
live in oxygen-poor water or pursue small prey in shallow water.[67] They may have later
ventured on land for brief periods. Eventually, some of them became so well adapted to
terrestrial life that they spent their adult lives on land, although they hatched in the water
and returned to lay their eggs. This was the origin of the amphibians. About 365 Ma,
another period of extinction occurred, perhaps as a result of global cooling.[68] Plants
evolved seeds, which dramatically accelerated their spread on land, around this time (by
approximately 360 Ma).[69][70]

Pangaea, the most recent supercontinent, existed from 300 to 180 Ma. The outlines of the
modern continents and other land masses are indicated on this map.
Some 20 million years later (340 Ma[31]:293-296), the amniotic egg evolved, which could be
laid on land, giving a survival advantage to tetrapod embryos. This resulted in the
divergence of amniotes from amphibians. Another 30 million years (310 Ma[31]:254-256) saw
the divergence of the synapsids (including mammals) from the sauropsids (including
birds and reptiles). Other groups of organisms continued to evolve, and lines diverged
in fish, insects, bacteria, and so onbut less is known of the details. The most recent
hypothesized supercontinent, called Pangaea, formed 300 Ma.

Mesozoic
The most severe extinction event to date took place 250 Ma, at the boundary of the
Permian and Triassic periods; 95% of life on Earth died out. That started the Mesozoic
era (meaning middle life) that spanned 187 million years,[71] possibly due to the Siberian
Traps volcanic event. The discovery of the Wilkes Land crater in Antarctica may indicate
a connection with the Permian-Triassic extinction, but the age of that crater is not known.
[72]
Among other speculative theories, it has been suggested that what is now the Gulf of
Mexico was created by a large bolide impact event at that time.[73] Life persevered, and
around 230 Ma,[74] dinosaurs split off from their reptilian ancestors. An extinction event
between the Triassic and Jurassic periods 200 Ma spared many of the dinosaurs,[75] and
they soon became dominant among the vertebrates. Though some of the mammalian lines
began to separate during this period, existing mammals were probably all small animals
resembling shrews.[31]:169
By 180 Ma, Pangaea broke up into Laurasia and Gondwana. The boundary between avian
and non-avian dinosaurs is not clear, but Archaeopteryx, traditionally considered one of
the first birds, lived around 150 Ma.[76] The earliest evidence for the angiosperms
evolving flowers is during the Cretaceous period, some 20 million years later (132 Ma).
[77]
Competition with birds drove many pterosaurs to extinction and the dinosaurs were

probably already in decline[78] when, 65 Ma, a 10-kilometre (6.2 mi) meteorite probably
struck Earth just off the Yucatn Peninsula where the Chicxulub crater is today. This
ejected vast quantities of particulate matter and vapor into the air that occluded sunlight,
inhibiting photosynthesis. Most large animals, including the non-avian dinosaurs, became
extinct,[79] marking the end of the Cretaceous period and Mesozoic era. Thereafter, in the
Paleocene epoch, mammals rapidly diversified, grew larger, and became the dominant
vertebrates. Perhaps a couple of million years later (around 63 Ma), the last common
ancestor of primates lived.[31]:160 By the late Eocene epoch, 34 Ma, some terrestrial
mammals had returned to the oceans to become animals such as Basilosaurus which
eventually led to dolphins and baleen whales.[80]

Cenozoic era (Recent life)

Human evolution

Australopithecus africanus, an early hominid.

Main article: Human evolution
A small African ape living around six Ma was the last animal whose descendants would
include both modern humans and their closest relatives, the bonobo and chimpanzees.
[31]:100-101
Only two branches of its family tree have surviving descendants. Very soon after
the split, for reasons that are still debated, apes in one branch developed the ability to
walk upright.[31]:95-99 Brain size increased rapidly, and by 2 Ma, the first animals classified
in the genus Homo had appeared.[63]:300 Of course, the line between different species or
even genera is somewhat arbitrary as organisms continuously change over generations.
Around the same time, the other branch split into the ancestors of the common
chimpanzee and the ancestors of the bonobo as evolution continued simultaneously in all
life forms.[31]:100-101
The ability to control fire probably began in Homo erectus (or Homo ergaster), probably
at least 790,000 years ago[81] but perhaps as early as 1.5 Ma.[31]:67 In addition, it has
sometimes suggested that the use and discovery of controlled fire may even predate
Homo erectus. Fire was possibly used by the early Lower Paleolithic (Oldowan) hominid
Homo habilis or strong australopithecines such as Paranthropus.[82]

It is more difficult to establish the origin of language; it is unclear whether Homo erectus
could speak or if that capability had not begun until Homo sapiens.[31]:67 As brain size
increased, babies were born earlier, before their heads grew too large to pass through the
pelvis. As a result, they exhibited more plasticity, and thus possessed an increased
capacity to learn and required a longer period of dependence. Social skills became more
complex, language became more sophisticated, and tools became more elaborate. This
contributed to further cooperation and intellectual development.[83]:7 Modern humans
(Homo sapiens) are believed to have originated somewhere around 200,000 years ago or
earlier in Africa; the oldest fossils date back to around 160,000 years ago.[84]
The first humans to show signs of spirituality are the Neanderthals (usually classified as a
separate species with no surviving descendants); they buried their dead, often apparently
with food or tools.[85]:17 However, evidence of more sophisticated beliefs, such as the early
Cro-Magnon cave paintings (probably with magical or religious significance)[85]:17-19 did
not appear until some 32,000 years ago.[86] Cro-Magnons also left behind stone figurines
such as Venus of Willendorf, probably also signifying religious belief.[85]:17-19 By 11,000
years ago, Homo sapiens had reached the southern tip of South America, the last of the
uninhabited continents (except for Antarctica, which remained undiscovered until 1820
AD).[87] Tool use and communication continued to improve, and interpersonal
relationships became more intricate.

Civilization
Main article: History of the world
Further information: History of Africa, History of the Americas, History of
Antarctica, and History of Eurasia

Vitruvian Man by Leonardo da Vinci epitomizes the advances in art and science seen
during the Renaissance.

Throughout more than 90% of its history, Homo sapiens lived in small bands as nomadic
hunter-gatherers.[83]:8 As language became more complex, the ability to remember and
communicate information resulted in a new replicator: the meme.[88] Ideas could be
exchanged quickly and passed down the generations.
Cultural evolution quickly outpaced biological evolution, and history proper began.
Somewhere between 8500 and 7000 BC, humans in the Fertile Crescent in Middle East
began the systematic husbandry of plants and animals: agriculture.[89] This spread to
neighboring regions, and developed independently elsewhere, until most Homo sapiens
lived sedentary lives in permanent settlements as farmers.
Not all societies abandoned nomadism, especially those in isolated areas of the globe
poor in domesticable plant species, such as Australia.[90] However, among those
civilizations that did adopt agriculture, the relative stability and increased productivity
provided by farming allowed the population to expand.
Agriculture had a major impact; humans began to affect the environment as never before.
Surplus food allowed a priestly or governing class to arise, followed by increasing
division of labor. This led to Earths first civilization at Sumer in the Middle East,
between 4000 and 3000 BC.[83]:15 Additional civilizations quickly arose in ancient Egypt,
at the Indus River valley and in China.
Starting around 3000 BC, Hinduism, one of the oldest religions still practiced today,
began to take form.[91] Others soon followed. The invention of writing enabled complex
societies to arise: record-keeping and libraries served as a storehouse of knowledge and
increased the cultural transmission of information. Humans no longer had to spend all
their time working for survivalcuriosity and education drove the pursuit of knowledge
and wisdom.
Various disciplines, including science (in a primitive form), arose. New civilizations
sprang up, traded with one another, and fought for territory and resources. Empires soon
began to develop. By around 500 BC, there were empires in the Middle East, Iran, India,
China, and Greece, on nearly equal footing; at times one empire expanded, only to
decline or be driven back later.[83]:3
In the fourteenth century, the Renaissance began in Italy with advances in religion, art,
and science.[83]:317-319 European civilization began to change beginning in 1500, leading to
the scientific and industrial revolutions. That continent began to exert political and
cultural dominance over human societies around the planet.[83]:295-299 From 1914 to 1918
and 1939 to 1945, nations around the world were embroiled in world wars.
Established following World War I, the League of Nations was a first step in establishing
international institutions to settle disputes peacefully. After failing to prevent World War
II, it was replaced by the United Nations. In 1992, several European nations joined in the
European Union. As transportation and communication improved, the economies and

political affairs of nations around the world have become increasingly intertwined. This
globalization has often produced both conflict and collaboration.

Recent events
Main article: Modern era
See also: Modernity and Future

Four and a half billion years after the planet's formation, Earths life broke free of the
biosphere. For the first time in history, Earth was viewed from space.
Change has continued at a rapid pace from the mid-1940s to today. Technological
developments include nuclear weapons, computers, genetic engineering, and
nanotechnology. Economic globalization spurred by advances in communication and
transportation technology has influenced everyday life in many parts of the world.
Cultural and institutional forms such as democracy, capitalism, and environmentalism
have increased influence. Major concerns and problems such as disease, war, poverty,
violent radicalism, and recently, human-caused climate change have risen as the world
population increases.[92]
In 1957, the Soviet Union launched the first artificial satellite into orbit and, soon
afterward, Yuri Gagarin became the first human in space. Neil Armstrong, an American,
was the first to set foot on another astronomical object, the Moon. Unmanned probes
have been sent to all the known planets in the solar system, with some (such as Voyager)
having left the solar system. The Soviet Union and the United States were the earliest
leaders in space exploration in the 20th Century. Five space agencies, representing over
fifteen countries,[93] have worked together to build the International Space Station. Aboard
it, there has been a continuous human presence in space since 2000.[

Timeline of evolution
From Wikipedia, the free encyclopedia

Jump to: navigation, search

For the history of evolutionary biology, see History of evolutionary thought.
This article is about the timeline of evolution in a general aspect. See also the timeline of
human evolution.
This timeline of the evolution of life outlines the major events in the development of life
on the planet Earth (See Organism). For a thorough explanatory context, see the history
of Earth, and geologic time scale. The dates given in this article are estimates based on
scientific evidence.
In biology, evolution is the process by which populations of organisms acquire and pass
on novel traits from generation to generation. Its occurrence over large stretches of time
explains the origin of new species and ultimately the vast diversity of the biological
world. Contemporary species are related to each other through common descent, products
of evolution and speciation over billions of years.

Contents
[hide]

1 Basic timeline
2 Detailed timeline
o 2.1 Hadean Eon
o 2.2 Archean Eon
o 2.3 Proterozoic Eon
o 2.4 Phanerozoic Eon
2.4.1 Paleozoic Era
2.4.2 Mesozoic Era
2.4.3 Cenozoic Era
3 See also
4 Further reading
5 References

6 External links

[edit] Basic timeline

Life on Earth
view discuss edit
-4500

-4000

-3500

-3000

-2500

-2000

-1500

-1000

-500

Life?
Photosynthesis?
Eukaryotes
Complex multicellular life
Animals
Land plants
"Dinosaurs"
Mammals
Flowers

Formation
of Earth

Meteorite bombardment

Atmospheric oxygen

Ediacara biota

Cambrian explosion

Modern-looking
humans

Axis scale: millions of years ago.

Dates prior to 1 billion years ago are speculative.

The basic timeline is a 4.6 billion year old Earth, with (very approximate) dates:

3.8 billion years of simple cells (prokaryotes),

3 billion years of photosynthesis,

2 billion years of complex cells (eukaryotes),

1 billion years of multicellular life,
600 million years of simple animals,
570 million years of arthropods (ancestors of insects, arachnids and crustaceans),
550 million years of complex animals,
500 million years of fish and proto-amphibians,
475 million years of land plants,
400 million years of insects and seeds,
360 million years of amphibians,
300 million years of reptiles,
200 million years of mammals,
150 million years of birds,
130 million years of flowers,
65 million years since the non-avian dinosaurs died out,
2.5 million years since the appearance of the genus Homo,
200,000 years since humans started looking like they do today,
25,000 years since Neanderthals died out.

[edit] Detailed timeline

Note that Ma, megaannum, means "million years ago".

[edit] Hadean Eon

3800 Ma and earlier.
Date

Event

4600 Ma The planet Earth forms from the accretion disc

revolving around the young Sun.

4533 Ma According to one plausible theory, the planet Earth

and the planet Theia collide, sending countless
moonlets into orbit around the young Earth. These
moonlets eventually coalesce to form the Moon.[1]
The gravitational pull of the new Moon stabilises
the Earth's fluctuating axis of rotation and sets up
the conditions in which life formed[citation needed].

4100 Ma The surface of the Earth cools enough for the crust
to solidify. The atmosphere and the oceans form.[2]

PAH infall,[3] and Iron-Sulfide synthesis along deep

ocean platelet boundaries, may have led to the RNA
world of competing organic compounds.

Between 4500 and 3500 Ma The earliest life appears, possibly derived from selfreproducing RNA molecules.[4][5] The replication of
these organisms requires resources like energy,
space, and smaller building blocks, which soon
become limited, resulting in competition, with
natural selection favouring those molecules which
are more efficient at replication. DNA molecules
then take over as the main replicators and these
archaic genomes soon develop inside enclosing
membranes which provide a stable physical and
chemical environment conducive to their
replication: proto-cells.[6][7][8]

3900 Ma Late Heavy Bombardment: peak rate of impact

events upon the inner planets by meteors. This
constant disturbance may have obliterated any life
that had evolved to that point, or possibly not, as
some early microbes could have survived in
hydrothermal vents below the Earth's surface;[9] or
life might have been transported to Earth by a
meteor.[10]
Somewhere between 3900 - 2500 Ma Cells resembling prokaryotes appear.[11] These first
organisms are chemoautotrophs: they use carbon
dioxide as a carbon source and oxidize inorganic
materials to extract energy. Later, prokaryotes
evolve glycolysis, a set of chemical reactions that
free the energy of organic molecules such as
glucose and store it in the chemical bonds of ATP.
Glycolysis (and ATP) continue to be used in almost
all organisms, unchanged, to this day.[12][13]

[edit] Archean Eon

3800 Ma 2500 Ma

Date

Event

3500 Ma Lifetime of the last universal ancestor;[14][15] the split between bacteria and
archaea occurs.[16]
Bacteria develop primitive forms of photosynthesis which at first do not produce
oxygen.[17] These organisms generate ATP by exploiting a proton gradient, a
mechanism still used in virtually all organisms.
3000 Ma Photosynthesizing cyanobacteria evolve; they use water as a reducing agent,
thereby producing oxygen as waste product.[18] The oxygen initially oxidizes
dissolved iron in the oceans, creating iron ore. The oxygen concentration in the
atmosphere subsequently rises, acting as a poison for many bacteria. The moon
is still very close to the earth and causes tides 1000 feet high. The earth is
continually wracked by hurricane force winds. These extreme mixing influences
are thought to stimulate evolutionary processes. (See Oxygen catastrophe)

[edit] Proterozoic Eon

2500 Ma 542 Ma
Date

Event

By 2100 Ma Eukaryotic cells appear.[19] Eukaryotes contain membrane-bound organelles

with diverse functions, probably derived from prokaryotes engulfing each
other via phagocytosis. (See Endosymbiosis)
By 1200 Ma Sexual reproduction first appears, increasing the rate of evolution.[20]

1200 Ma Simple multicellular organisms evolve, mostly consisting of cell colonies of

limited complexity.
850630 Ma A global glaciation may have occurred.[21][22] Opinion is divided on whether
it increased or decreased biodiversity or the rate of evolution.[23][24][25]

580542 Ma The Ediacaran biota represent the first large, complex multicellular
organisms - although their affinities remain a subject of debate.[26]

580500 Ma Most modern phyla of animals begin to appear in the fossil record during the
Cambrian explosion.[27][28]

580540 Ma The accumulation of atmospheric oxygen allows the formation of an ozone

layer.[29] This blocks ultraviolet radiation, permitting the colonisation of the
land.[29]

[edit] Phanerozoic Eon

542 Ma present
The Phanerozoic Eon, literally the "period of well-displayed life", marks the appearance
in the fossil record of abundant, shell-forming and/or trace-making organisms. It is
subdivided into three eras, the Paleozoic, Mesozoic and Cenozoic, which are divided by
major mass extinctions.
[edit] Paleozoic Era
542 Ma 251.0 Ma
Date

Event

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

530 Ma The first known footprints on land date to 530 Ma, indicating that early animal
explorations may have predated the development of terrestrial plants.[30]

525 Ma Earliest graptolites.

510 Ma First cephalopods (Nautiloids) and chitons.

505 Ma Fossilization of the Burgess Shale.

485 Ma First vertebrates with true bones (jawless fishes).

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

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

434 Ma The first primitive plants move onto land,[31][citation needed] having evolved from
green algae living along the edges of lakes.[32] They are accompanied by fungi,
which may have aided the colonisation of land through symbiosis.

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

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

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

and ammonoids.

363 Ma By the start of the Carboniferous Period, the Earth begins to be recognisable.
Insects roamed the land and would soon take to the skies; sharks swam the
oceans as top predators,[33] and vegetation covered the land, with seed-bearing
plants and forests soon to flourish.
Four-limbed tetrapods gradually gain adaptations which will help them occupy
a terrestrial life-habit.
360 Ma First crabs and ferns. Land flora dominated by seed ferns.
350 Ma First large sharks, ratfishes, and hagfish.
340 Ma Diversification of amphibians.
330 Ma First amniote vertebrates (Paleothyris).
305 Ma Earliest diapsid reptiles (e.g. Petrolacosaurus).
280 Ma Earliest beetles, seed plants and conifers diversify while lepidodendrids and
sphenopsids decrease. Terrestrial temnospondyl amphibians and pelycosaurs
(e.g. Dimetrodon) diversify in species.

251.4 Ma The Permian-Triassic extinction event eliminates over 90-95% of marine

species. Terrestrial organisms were not as seriously affected as the marine biota.
This "clearing of the slate" may have led to an ensuing diversification, however
life on land took 30M years to completely recover.[34]
[edit] Mesozoic Era
Date

Event

From 251.4 Ma The Mesozoic Marine Revolution begins: increasingly well-adapted and
diverse predators pressurise sessile marine groups; the "balance of power"
in the oceans shifts dramatically as some groups of prey adapt more
rapidly and effectively than others.

245 Ma Earliest ichthyosaurs.

240 Ma Increase in diversity of gomphodont cynodonts and rhynchosaurs.

225 Ma Earliest dinosaurs (prosauropods), first cardiid bivalves, diversity in

cycads, bennettitaleans, and conifers. First teleost fishes.

215 Ma First mammals (e.g. Eozostrodon), minor vertebrate extinctions occur

220 Ma

Eoraptor, among the earliest dinosaurs, appeared in the fossil record 230
million years ago.
Gymnosperm forests dominate the land; herbivores grow to huge sizes in
order to accommodate the large guts necessary to digest the nutrient-poor
plants.[citation needed], first flies and turtles (Odontochelys).
200 Ma The first accepted evidence for viruses (at least, the group Geminiviridae)
exists.[35] Viruses are still poorly understood and may have arisen before

"life" itself, or may be a more recent phenomenon.

Major extinctions in terrestrial vertebrates and large amphibians
195 Ma First pterosaurs with specialized feeding (Dorygnathus). First sauropod
dinosaurs. Diversification in small, ornithischian dinosaurs:
heterodontosaurids, fabrosaurids, and scelidosaurids.
190 Ma Pliosaurs appear in the fossil record. First lepidopteran insects
(Archaeolepis), hermit crabs, modern starfish, irregular echinoids,
corbulid bivalves, and tubulipore bryozoans. Extensive development of
sponge reefs.
170 Ma Earliest salamanders, newts, cryptoclidid & elasmosaurid plesiosaurs, and
cladotherian mammals. Cynodonts become extinct while sauropod
dinosaurs diversify.
165 Ma First rays and glycymeridid bivalves.
161 Ma Ceratopsian dinosaurs appear in the fossil record (Yinlong)
155 Ma First blood-sucking insects (ceratopogonids), rudist bivalves, and
cheilosome bryozoans. Archaeopteryx, a possible ancestor to the birds,
appears in the fossil record, along with triconodontid and symmetrodont
mammals. Diversity in stegosaurian and theropod dinosaurs.
130 Ma The rise of the Angiosperms: These flowering plants boast structures that
attract insects and other animals to spread pollen. This innovation causes
a major burst of animal evolution through co-evolution. First freshwater
pelomedusid turtles.
115 Ma First monotreme mammals.
110 Ma First hesperornithes, toothed diving birds. Earliest limopsid, verticordiid,
and thyasirid bivalves.
106 Ma Spinosaurus, the largest theropod dinosaur, appears in the fossil record.
100 Ma Earliest bees.
90 Ma Extinction of ichthyosaurs. Earliest snakes and nuculanid bivalves. Large
diversification in angiosperms: magnoliids, rosids, hamamelidids,
monocots, and ginger. Earliest examples of ticks.
80 Ma First ants and termites.
70 Ma Multituberculate mammals increase in diversity. First yoldiid bivalves.
68 Ma Tyrannosaurus, the largest terrestrial predator of North America appears
in the fossil record. First species of Triceratops.
[edit] Cenozoic Era
65.5 Ma present

Date

Event

65.5 Ma File:KT Impact2.jpg

The CretaceousTertiary extinction event eradicates about half of all animal
species, including mosasaurs, pterosaurs, plesiosaurs, ammonites,
belemnites, rudist and inoceramid bivalves, most planktic foraminifers, and
all of the dinosaurs excluding their descendants the birds [36]
From 65 Ma Rapid dominance of conifers and ginkgos in high latitudes, along with
mammals becoming the dominant species. First psammobiid bivalves. Rapid
diversification in ants.
63 Ma Evolution of the creodonts, an important group of carnivorous mammals.
60 Ma Diversification of large, flightless birds. Earliest true primates, along with
the first semelid bivalves, edentates, carnivorous and lipotyphlan mammals,
and owls. The ancestors of the carnivorous mammals (miacids) were alive.
56 Ma Gastornis, a large, flightless bird appears in the fossil record, becoming an
apex predator at the time.
55 Ma Modern bird groups diversify (first song birds, parrots, loons, swifts,
woodpeckers), first whale (Himalayacetus), earliest rodents, lagomorphs,
armadillos, appearance of sirenians, proboscideans, perissodactyl and
artiodactyl mammals in the fossil record. Angiosperms diversify. The
ancestor (according to theory) of the species in Carcharodon, the early mako
shark Isurus hastalis, is alive.
52 Ma First bats appear (Onychonycteris).
50 Ma Peak diversity of dinoflagellates and nanofossils, increase in diversity of
anomalodesmatan and heteroconch bivalves, brontotheres, tapirs,
rhinoceroses, and camels appear in the fossil record, diversification of
primates.
40 Ma Modern type butterflies and moths appear. Extinction of Gastornis.
Basilosaurus, one of the first of the giant whales, appeared in the fossil
record.
35 Ma Grasses evolve from among the angiosperms; grassland begin to expand.
Slight increase in diversity of cold-tolerant ostracods and foraminifers, along
with major extinctions of gastropods, reptiles, and amphibians. Many
modern mammal groups begin to appear: first glyptodonts, ground sloths,
dogs, peccaries, and the first eagles and hawks. Diversity in toothed and
baleen whales.
33 Ma Evolution of the thylacinid marsupials (Badjcinus).
30 Ma First balanids and eucalypts, extinction of embrithopod and brontothere
mammals, earliest pigs and cats.

28 Ma Paraceratherium appears in the fossil record, the largest terrestrial mammal

that ever lived.
25 Ma First deer.
20 Ma First giraffes and giant anteaters, increase in bird diversity.
15 Ma Mammut appears in the fossil record, first bovids and kangaroos, diversity in
Australian megafauna.
10 Ma Grasslands and savannas are established, diversity in insects, especially ants
and termites, horses increase in body size and develop high-crowned teeth,
major diversification in grassland mammals and snakes.
6.5 Ma First hominin (Sahelanthropus).
6 Ma Australopithecines diversify (Orrorin, Ardipithecus)
5 Ma First tree sloths and hippopotami, diversification of grazing herbivores, large
carnivorous mammals, burrowing rodents, kangaroos, birds, and small
carnivores, vultures increase in size, decrease in the number of perissodactyl
mammals
4.8 Ma Mammoths appear in the fossil record.
4 Ma Evolution of Australopithecus, Stupendemys appears in the fossil record as
the largest freshwater turtle.
3 Ma The Great American Interchange, where various land and freshwater faunas
migrated between North and South America. Armadillos, opossums,
hummingbirds, and vampire bats traveled to North America while horses,
tapirs, saber-toothed cats, and deer entered South America.
2 Ma Extinction of australopithecines. First members of the genus Homo appear in
the fossil record. Diversification of conifers in high latitudes.
1.2 Ma Evolution of Homo antecessor.
200 ka Anatomically modern humans appear in Africa.[37][38][39] Around 50,000 years
before present they start colonising the other continents, replacing the
Neanderthals in Europe and other hominins in Asia.
10 ka The Holocene Epoch starts 10,000[40] years ago after the Late Glacial
Maximum. The last mammoth species, the woolly mammoth, becomes
extinct.
Present day The impact of humanity is felt in all corners of the globe and contributing to
a dramatically rising extinction rate, estimated in 1995 to be 100 times the
background rate.[41][42]

[edit] See also

Book:Evolution

Books are collections of articles which can be downloaded or ordered in print.

Evolutionary history of life

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

History of evolutionary thought

From Wikipedia, the free encyclopedia

Jump to: navigation, search

This article is about the history of evolutionary thought in biology. For the history of
evolutionary thought in the social sciences, see Sociocultural evolution. For the history
of religious discussions, see History of the creation-evolution controversy.

The Tree of Life as depicted by Ernst Haeckel in The Evolution of Man (1879) illustrates
the 19th-century view that evolution was a progressive process leading towards man.
Evolutionary thought, the conception that species change over time, has roots in
antiquity, in the ideas of the ancient Greeks, Romans, and Chinese as well as in medieval
Islamic science. However, until the 18th century, Western biological thinking was
dominated by essentialism, the belief that every species has essential characteristics that
are unalterable. This began to change during the Enlightenment when evolutionary
cosmology and the mechanical philosophy spread from the physical sciences to natural
history. Naturalists began to focus on the variability of species; the emergence of
paleontology with the concept of extinction further undermined the static view of nature.
In the early 19th century, Jean-Baptiste Lamarck proposed his theory of the transmutation
of species, the first fully formed scientific theory of evolution.
In 1858, Charles Darwin and Alfred Russel Wallace published a new evolutionary theory
that was explained in detail in Darwin's On the Origin of Species (1859). Unlike
Lamarck, Darwin proposed common descent and a branching tree of life. The theory was
based on the idea of natural selection, and it synthesized a broad range of evidence from
animal husbandry, biogeography, geology, morphology, and embryology.
The debate over Darwin's work led to the rapid acceptance of the general concept of
evolution, but the specific mechanism he proposed, natural selection, was not widely
accepted until it was revived by developments in biology that occurred during 1920s
through the 1940s. Before that time most biologists argued that other factors were
responsible for evolution. Alternatives to natural selection suggested during the eclipse of
Darwinism included inheritance of acquired characteristics (neo-Lamarckism), an innate
drive for change (orthogenesis), and sudden large mutations (saltationism). The synthesis
of natural selection with Mendelian genetics during the 1920s and 1930s founded the new
discipline of population genetics. Throughout the 1930s and 1940s, population genetics
became integrated with other biological fields, resulting in a widely applicable theory of
evolution that encompassed much of biologythe modern evolutionary synthesis.
Following the establishment of evolutionary biology, studies of mutation and variation in
natural populations, combined with biogeography and systematics, led to sophisticated
mathematical and causal models of evolution. Paleontology and comparative anatomy
allowed more detailed reconstructions of the history of life. After the rise of molecular
genetics in the 1950s, the field of molecular evolution developed, based on protein
sequences and immunological tests, and later incorporating RNA and DNA studies. The
gene-centered view of evolution rose to prominence in the 1960s, followed by the neutral
theory of molecular evolution, sparking debates over adaptationism, the units of
selection, and the relative importance of genetic drift versus natural selection. In the late
20th century, DNA sequencing led to molecular phylogenetics and the reorganization of
the tree of life into the three-domain system. In addition, the newly recognized factors of
symbiogenesis and horizontal gene transfer introduced yet more complexity into
evolutionary history.

Part of the Biology series on

Evolution

Mechanisms and processes

Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation
Research and history
Introduction
Evidence
Evolutionary history of life
History
Modern synthesis
Social effect
Theory and fact
Objections / Controversy
Evolutionary biology fields
Cladistics
Ecological genetics
Evolutionary development
Evolutionary psychology
Human evolution
Molecular evolution
Phylogenetics
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Biology portal v d e

Contents

[hide]

1 Antiquity
o 1.1 Greeks
o 1.2 Chinese
o 1.3 Romans
o 1.4 Augustine of Hippo
2 Middle Ages
o 2.1 Islamic philosophy and the struggle for existence
o 2.2 Christian philosophy and the great chain of being
o 2.3 Thomas Aquinas on creation and natural processes
3 Renaissance and Enlightenment
4 Early 19th century
o 4.1 Paleontology and geology
o 4.2 Transmutation of species
o 4.3 Anticipations of natural selection
o 4.4 Natural selection
5 18591930s: Darwin and his legacy
o 5.1 Application to humans
o 5.2 Alternatives to natural selection
o 5.3 Mendelian genetics, biometrics, and mutation
6 1920s1940s
o 6.1 Population genetics
o 6.2 Modern evolutionary synthesis
7 1940s1960s: Molecular biology and evolution
8 Late 20th century
o 8.1 Gene-centered view
o 8.2 Sociobiology
o 8.3 Evolutionary paths and processes
o 8.4 Microbiology, horizontal gene transfer, and endosymbiosis
o 8.5 Evolutionary developmental biology
9 21st century
o 9.1 Epigenetic inheritance
10 Unconventional evolutionary theory
o 10.1 Omega point
o 10.2 Gaia hypothesis
o 10.3 Transhumanism
11 See also
12 Notes
13 References

14 External links

[edit] Antiquity
[edit] Greeks

Several ancient Greek philosophers discussed ideas that involved change in living
organisms over time. Anaximander (c.610546 BC) proposed that the first animals lived
in water and animals that live on land were generated from them.[1] Empedocles (c. 490
430 BC) wrote of a non-supernatural origin for living things.[2] Empedocles suggested
that adaptation did not require an organizer or final cause. Aristotle summarized his idea:
"Wherever then all the parts came about just what they would have been if they had come
to be for an end, such things survived, being organized spontaneously in a fitting way;
whereas those which grew otherwise perished and continue to perish ..." although
Aristotle himself rejected this view.[3]

Plato (left) and Aristotle (right), a detail of The School of Athens

Plato (c. 428348 BC) was, in the words of biologist and historian Ernst Mayr, "the great
antihero of evolutionism",[4] as he established the philosophy of essentialism, which he
called the Theory of Forms. This theory holds that objects observed in the real world are
only reflections of a limited number of essences (eide). Variation is merely the result of
an imperfect reflection of these constant essences. In his Timaeus, Plato set forth the idea
that the Demiurge had created the cosmos and everything in it because, being good, and
hence, "... free from jealousy, He desired that all things should be as like Himself as they
could be". The creator created all conceivable forms of life, since "... without them the
universe will be incomplete, for it will not contain every kind of animal which it ought to
contain, if it is to be perfect". This idea, that all potential forms of life are essential to a
perfect creation, is called the plenitude principle, and greatly influenced Christian
thought.[5]
Aristotle (384322 BC), one of the most influential of the Greek philosophers, is the
earliest natural historian whose work has been preserved in any real detail. His writings
on biology were the result of his research into natural history on and around the isle of
Lesbos, and have survived in the form of four books, usually known by their Latin
names, De anima (on the essence of life), Historia animalium (inquiries about animals),
De generatione animalium (reproduction), and De partibus animalium (anatomy).
Aristotle's works contain some remarkably astute observations and interpretationsalong
with sundry myths and mistakesreflecting the uneven state of knowledge during his

time.[6] However, for Charles Singer, "Nothing is more remarkable than [Aristotle's]
efforts to [exhibit] the relationships of living things as a scala natur."[6] This scala
natur, described in Historia animalium, classified organisms in relation to a hierarchical
"Ladder of Life" or "Chain of Being", placing them according to their complexity of
structure and function, with organisms that showed greater vitality and ability to move
described as "higher organisms".[5]

[edit] Chinese
Ideas on evolution were expressed by ancient Chinese thinkers such as Zhuangzi (Chuang
Tzu), a Taoist philosopher who lived around the 4th century BC. According to Joseph
Needham, Taoism explicitly denies the fixity of biological species and Taoist
philosophers speculated that species had developed differing attributes in response to
differing environments.[7] Humans, nature and the heavens are seen as existing in a state
of "constant transformation" known as the Tao, in contrast with the more static view of
nature typical of Western thought.[8]

[edit] Romans
Titus Lucretius Carus (d. 50 BC), the Roman philosopher and atomist, wrote the poem
On the Nature of Things (De rerum natura), which provides the best surviving
explanation of the ideas of the Greek Epicurean philosophers. It describes the
development of the cosmos, the Earth, living things, and human society through purely
naturalistic mechanisms, without any reference to supernatural involvement. On the
Nature of things would influence the cosmological and evolutionary speculations of
philosophers and scientists during and after the Renaissance.[9][10]

[edit] Augustine of Hippo

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

[edit] Middle Ages

[edit] Islamic philosophy and the struggle for existence
See also: Early Islamic philosophy and Science in medieval Islam

Although Greek and Roman evolutionary ideas died out in Europe after the fall of the
Roman Empire, they were not lost to Islamic philosophers and scientists. In the Islamic
Golden Age, early evolutionary ideas were explored by Islamic philosophers. These ideas
included not only transmutation of one kind of living thing into another but also
transmutation from non living to living: "from mineral to plant, from plant to animal, and
from animal to man".[15][16]
The first Muslim biologist and philosopher to speculate in detail about evolution was the
Afro-Arab writer al-Jahiz in the 9th century. In the Book of Animals, he considered the
effects of the environment on an animal's chances for survival, and described the struggle
for existence.[17][18] Al-Jahiz was also wrote vivid descriptions of food chains.[19] Al-Jahiz
speculated on the influence of the environment on animals and developed an early theory
of evolution. Al-Jahiz considered the effects of the environment on the likelihood of an
animal to survive, and first described the struggle for existence.[17] He described the
struggle for existence in terms that anticipate natural selection. Al-Jahiz's ideas on the
struggle for existence in the Book of Animals have been summarized as follows:
Animals engage in a struggle for existence; for resources, to avoid being eaten and to breed.
Environmental factors influence organisms to develop new characteristics to ensure survival, thus
transforming into new species. Animals that survive to breed can pass on their successful
characteristics to offspring.[20][unreliable source?]

Ibn Miskawayh's al-Fawz al-Asghar and the Brethren of Purity's Encyclopedia of the
Brethren of Purity (The Epistles of Ikhwan al-Safa) set forth ideas on how species
developed: from matter into vapor and thence to water, then minerals into plants and then
animals, leading to apes and, finally, humans.[21][22] Abu Rayhan Biruni explained the idea
of artificial selection and argued that nature works in much the same way.[23]

[edit] Christian philosophy and the great chain of being

Drawing of the great chain of being from Rhetorica Christiana (1579) by Diego Valades
During the Early Middle Ages, Greek classical learning was all but lost to the West.
However, contact with the Islamic world, where Greek manuscripts were preserved and
expanded, soon led to a massive spate of Latin translations in the 12th century. Europeans
were re-introduced to the works of Plato and Aristotle, as well as to Islamic thought.
Christian thinkers of the scholastic school, in particular Abelard and Thomas Aquinas,
combined Aristotelian classification with Plato's ideas of the goodness of God, and of all
potential life forms being present in a perfect creation, to organize all inanimate, animate,
and spiritual beings into a huge interconnected system: the scala natur, or great chain of
being.[5][24]
Within this system, everything that existed could be placed in order, from "lowest" to
"highest", with Hell at the bottom and God at the topbelow God, an angelic hierarchy
marked by the orbits of the planets, mankind in an intermediate position, and worms the
lowest of the animals. As the universe was ultimately perfect, the great chain was also
perfect. There were no empty links in the chain, and no link was represented by more
than one species. Therefore no species could ever move from one position to another.
Thus, in this Christianized version of Plato's perfect universe, species could never
change, but remained forever fixed, in accordance with the text of Genesis. For humans
to forget their position was seen as sinful, whether they behaved like lower animals or
aspired to a higher station than was given them by their Creator.[5]
Creatures on adjacent steps were expected to closely resemble each other, an idea
expressed in the saying: natura non facit saltum ("nature does not make leaps").[5] This
basic concept of the great chain of being greatly influenced the thinking of Western
civilization for centuries (and still has an influence today). It formed a part of the
argument from design presented by natural theology. As a classification system, it

became the major organizing principle and foundation of the emerging science of biology
in the 17th and 18th centuries.[5]

[edit] Thomas Aquinas on creation and natural processes

While the development of the great chain of being and the argument from design by
Christian theologians contributed to the view that the natural world fit into an unchanging
designed hierarchy, some theologians were more open to the possibility that the world
might have developed through natural processes. Thomas Aquinas went even farther than
Augustine of Hippo in arguing that scriptural texts like Genesis should not be interpreted
in a literal way that conflicted with or constrained what natural philosophers learned
about the workings of the natural world. He felt that the autonomy of nature was a sign of
God's goodness and that there was no conflict between the concept of a divinely created
universe, and the idea that the universe may have evolved over time through natural
mechanisms.[25] However, Aquinas disputed the views of those like the ancient Greek
philosopher Empodocles who held that such natural processes showed that the universe
could have developed without an underlying purpose. Rather holding that:
Hence, it is clear that nature is nothing but a certain kind of art, i.e., the divine art, impressed
upon things, by which these things are moved to a determinate end. It is as if the shipbuilder were
able to give to timbers that by which they would move themselves to take the form of a ship. [26]

[edit] Renaissance and Enlightenment

Main article: Evolutionary ideas of the Renaissance and Enlightenment

Pierre Belon compared the skeletons of birds and humans in his Book of Birds (1555).
In the first half of the 17th century, Ren Descartes's mechanical philosophy encouraged
the use of the metaphor of the universe as a machine, a concept that would come to
characterise the scientific revolution.[27] Between 1650 and 1800, some evolutionist
theories supported the view that the universe, including life on Earth, had developed
mechanically, entirely without divine guidance. In contrast, most contemporary theories
of evolution, such of those of Gottfried Leibniz and J. G. Herder, held that evolution was
a fundamentally spiritual process.[28] In 1751, Pierre Louis Maupertuis veered toward
more materialist ground. He wrote of natural modifications occurring during reproduction

and accumulating over the course of many generations, producing races and even new
species, a description that anticipated in general terms the concept of natural selection.[29]
Later in the 18th century, the French philosopher G. L. L. Buffon, one of the leading 18th
century naturalists, suggested that what most people referred to as species were really just
well-marked varieties, modified from an original form by environmental factors. For
example, he believed that lions, tigers, leopards and house cats might all have a common
ancestor. He further speculated that the 200 or so species of mammals then known might
have descended from as few as 38 original animal forms. Buffon's evolutionary ideas
were limited; he believed each of the original forms had arisen through spontaneous
generation and that each was shaped by "internal moulds" that limited the amount of
change. Buffon's works, Natural History and The Epochs of Nature, containing well
developed theories about a completely materialistic origin for the Earth and his ideas
questioning the fixity of species, were extremely influential.[30][31]
Between 1767 and 1792, James Burnett, Lord Monboddo included in his writings not
only the concept that man had descended from primates, but also that, in response to the
environment, creatures had found methods of transforming their characteristics over long
time intervals.[32] Charles Darwin's grandfather, Erasmus Darwin, published Zonomia in
1796, which suggested that "all warm-blooded animals have arisen from one living
filament".[33] In his 1802 poem Temple of Nature, he described the rise of life from minute
organisms living in mud to all of its modern diversity.[34]

[edit] Early 19th century

Diagram of the geologic timescale from an 1861 book by Richard Owen showing the
appearance of major animal types

[edit] Paleontology and geology

See also: History of paleontology
In 1796, Georges Cuvier published his findings on the differences between living
elephants and those found in the fossil record. His analysis demonstrated that mammoths
and mastodons were distinct species different from any living animal, effectively ending
a long-running debate over whether the extinction of a species was possible.[35] In 1788,
James Hutton described gradual geological processes operating continuously over deep
time.[36] In the 1790s William Smith began the process of ordering rock strata by
examining fossils in the layers while he worked on his geologic map of England.
Independently, in 1811, Georges Cuvier and Alexandre Brongniart published an
influential study of the geologic history of the region around Paris, based on the
stratigraphic succession of rock layers. These works helped establish the antiquity of the
Earth.[37] Cuvier advocated catastrophism to explain the patterns of extinction and faunal
succession revealed by the fossil record.
Knowledge of the fossil record continued to advance rapidly during the first few decades
of the 19th century. By the 1840s, the outlines of the geologic timescale were becoming
clear, and in 1841 John Phillips named three major eras, based on the predominant fauna
of each: the Paleozoic, dominated by marine invertebrates and fish, the Mesozoic, the age
of reptiles, and the current Cenozoic age of mammals. This progressive picture of the
history of life was accepted even by conservative English geologists like Adam Sedgwick
and William Buckland; however, like Cuvier, they attributed the progression to repeated
catastrophic episodes of extinction followed by new episodes of creation.[38] Unlike
Cuvier, Buckland and some other advocates of natural theology among British geologists
made efforts to explicitly link the last catastrophic episode proposed by Cuvier to the
biblical flood.[39][40]
From 1830 to 1833, Charles Lyell published his multi-volume work Principles of
Geology, which, building on Hutton's ideas, advocated a uniformitarian alternative to the
catastrophic theory of geology. Lyell claimed that, rather than being the products of
cataclysmic (and possibly supernatural) events, the geologic features of the Earth are
better explained as the result of the same gradual geologic forces observable in the
present daybut acting over immensely long periods of time. Although Lyell opposed
evolutionary ideas (even questioning the consensus that the fossil record demonstrates a
true progression), his concept that the Earth was shaped by forces working gradually over
an extended period, and the immense age of the Earth assumed by his theories, would
strongly influence future evolutionary thinkers such as Charles Darwin.[41]

[edit] Transmutation of species

Main article: Transmutation of species

Diagram from Vestiges of the Natural History of Creation (1844) by Robert Chambers
shows a model of development where fish (F), reptiles (R), and birds (B) represent
branches from a path leading to mammals (M).
Jean-Baptiste Lamarck proposed, in his Philosophie Zoologique of 1809, a theory of the
transmutation of species. Lamarck did not believe that all living things shared a common
ancestor but rather that simple forms of life were created continuously by spontaneous
generation. He also believed that an innate life force drove species to become more
complex over time, advancing up a linear ladder of complexity that was related to the
great chain of being. Lamarck recognized that species were adapted to their environment.
He explained this by saying that the same innate force driving increasing complexity
caused the organs of an animal (or a plant) to change based on the use or disuse of those
organs, just as muscles are affected by exercise. He argued that these changes would be
inherited by the next generation and produce slow adaptation to the environment. It was
this secondary mechanism of adaptation through the inheritance of acquired
characteristics that would become known as Lamarckism and would influence
discussions of evolution into the 20th century.[42][43]
A radical British school of comparative anatomy that included the anatomist Robert Grant
was closely in touch with Lamarck's French school of Transformationism. One of the
French scientists who influenced Grant was the anatomist tienne Geoffroy Saint-Hilaire,
whose ideas on the unity of various animal body plans and the homology of certain
anatomical structures would be widely influential and lead to intense debate with his
colleague Georges Cuvier. Grant became an authority on the anatomy and reproduction
of marine invertebrates. He developed Lamarck's and Erasmus Darwin's ideas of

transmutation and evolutionism, and investigated homology, even proposing that plants
and animals had a common evolutionary starting point. As a young student Charles
Darwin joined Grant in investigations of the life cycle of marine animals. In 1826 an
anonymous paper, probably written by Robert Jameson, praised Lamarck for explaining
how higher animals had "evolved" from the simplest worms; this was the first use of the
word "evolved" in a modern sense.[44][45]
In 1844, the Scottish publisher Robert Chambers anonymously published an extremely
controversial but widely read book entitled Vestiges of the Natural History of Creation.
This book proposed an evolutionary scenario for the origins of the Solar System and life
on Earth. It claimed that the fossil record showed a progressive ascent of animals with
current animals being branches off a main line that leads progressively to humanity. It
implied that the transmutations lead to the unfolding of a preordained plan that had been
woven into the laws that governed the universe. In this sense it was less completely
materialistic than the ideas of radicals like Robert Grant, but its implication that humans
were only the last step in the ascent of animal life incensed many conservative thinkers.
The high profile of the public debate over Vestiges, with its depiction of evolution as a
progressive process, would greatly influence the perception of Darwin's theory a decade
later.[46][47]
Ideas about the transmutation of species were associated with the radical materialism of
the Enlightenment and were attacked by more conservative thinkers. Georges Cuvier
attacked the ideas of Lamarck and Geoffroy Saint-Hilaire, agreeing with Aristotle that
species were immutable. Cuvier believed that the individual parts of an animal were too
closely correlated with one another to allow for one part of the anatomy to change in
isolation from the others, and argued that the fossil record showed patterns of
catastrophic extinctions followed by re-population, rather than gradual change over time.
He also noted that drawings of animals and animal mummies from Egypt, which were
thousands of years old, showed no signs of change when compared with modern animals.
The strength of Cuvier's arguments and his scientific reputation helped keep
transmutational ideas out of the mainstream for decades.[48]

This 1847 diagram by Richard Owen shows his conceptual archetype for all vertebrates.
In Britain the philosophy of natural theology remained influential. William Paley's 1802
book Natural Theology with its famous watchmaker analogy had been written at least in
part as a response to the transmutational ideas of Erasmus Darwin.[49] Geologists
influenced by natural theology, such as Buckland and Sedgwick, made a regular practice
of attacking the evolutionary ideas of Lamarck, Grant, and The Vestiges of the Natural
History of Creation.[50][51] Although the geologist Charles Lyell opposed scriptural

geology, he also believed in the immutability of species, and in his Principles of Geology
(18301833), he criticized Lamarck's theories of development.[41] Idealists such as Louis
Agassiz and Richard Owen believed that each species was fixed and unchangeable
because it represented an idea in the mind of the creator. They believed that relationships
between species could be discerned from developmental patterns in embryology, as well
as in the fossil record, but that these relationships represented an underlying pattern of
divine thought, with progressive creation leading to increasing complexity and
culminating in humanity. Owen developed the idea of "archetypes" in the Divine mind
that would produce a sequence of species related by anatomical homologies, such as
vertebrate limbs. Owen led a public campaign that successfully marginalized Robert
Grant in the scientific community. Darwin would make good use of the homologies
analyzed by Owen in his own theory, but the harsh treatment of Grant, and the
controversy surrounding Vestiges, showed him the need to ensure that his own ideas were
scientifically sound.[45][52][53]

[edit] Anticipations of natural selection

Several writers anticipated aspects of Darwin's theory, and in the third edition of On the
Origin of Species published in 1861 Darwin named those he knew about in an
introductory appendix, An Historical Sketch of the Recent Progress of Opinion on the
Origin of Species, which he expanded in later editions.[54]
In 1813, William Charles Wells read before the Royal Society essays assuming that there
had been evolution of humans, and recognising the principle of natural selection. Charles
Darwin and Alfred Russel Wallace were unaware of this work when they jointly
published the theory in 1858, but Darwin later acknowledged that Wells had recognised
the principle before them, writing that the paper "An Account of a White Female, part of
whose Skin resembles that of a Negro" was published in 1818, and "he distinctly
recognises the principle of natural selection, and this is the first recognition which has
been indicated; but he applies it only to the races of man, and to certain characters
alone."[55] When Darwin was developing his theory, he was influenced by Augustin de
Candolle's natural system of classification, which laid emphasis on the war between
competing species.[56][57]
Patrick Matthew wrote in the obscure book Naval Timber & Arboriculture (1831) of
"continual balancing of life to circumstance. ... [The] progeny of the same parents, under
great differences of circumstance, might, in several generations, even become distinct
species, incapable of co-reproduction."[58] Charles Darwin discovered this work after the
initial publication of the Origin. In the brief historical sketch that Darwin included in the
3rd edition he says "Unfortunately the view was given by Mr. Matthew very briefly in an
Appendix to a work on a different subject ... He clearly saw, however, the full force of the
principle of natural selection."[59]
It is possible to look through the history of biology from the ancient Greeks onwards and
discover anticipations of almost all of Darwin's key ideas. However, as historian of
science Peter J. Bowler says, "Through a combination of bold theorizing and

comprehensive evaluation, Darwin came up with a concept of evolution that was unique
for the time." Bowler goes on to say that simple priority alone is not enough to secure a
place in the history of science; someone has to develop an idea and convince others of its
importance to have a real impact.[60]
T. H. Huxley said in his essay on the reception of the Origin of Species:
The suggestion that new species may result from the selective action of external conditions upon
the variations from their specific type which individuals present and which we call spontaneous
because we are ignorant of their causation is as wholly unknown to the historian of scientific
ideas as it was to biological specialists before 1858. But that suggestion is the central idea of the
Origin of Species, and contains the quintessence of Darwinism.[61]

Darwin's first sketch of an evolutionary tree from his First Notebook on Transmutation of
Species (1837)

[edit] Natural selection

Main articles: Inception of Darwin's theory, Development of Darwin's theory, and
Publication of Darwin's theory
The biogeographical patterns Charles Darwin observed in places such as the Galapagos
islands during the voyage of the Beagle caused him to doubt the fixity of species, and in
1837 Darwin started the first of a series of secret notebooks on transmutation. Darwin's
observations led him to view transmutation as a process of divergence and branching,
rather than the ladder-like progression envisioned by Lamarck and others. In 1838 he
read the new 6th edition of An Essay on the Principle of Population, written in the late
1700s by Thomas Malthus. Malthus' idea of population growth leading to a struggle for
survival combined with Darwin's knowledge on how breeders selected traits, led to the
inception of Darwin's theory of natural selection. Darwin did not publish his ideas on
evolution for 20 years. However he did share them with certain other naturalists and
friends, starting with Joseph Hooker, with whom he discussed his unpublished 1844

essay on natural selection. During this period he used the time he could spare from his
other scientific work to slowly refine his ideas and, aware of the intense controversy
around transmutation, amass evidence to support them. In September 1854 he began full
time work on writing his book on natural selection.[53][62][63]
Unlike Darwin, Alfred Russel Wallace, influenced by the book Vestiges of the Natural
History of Creation, already suspected that transmutation of species occurred when he
began his career as a naturalist. By 1855 his biogeographical observations during his field
work in South America and the Malay Archipelago made him confident enough in a
branching pattern of evolution to publish a paper stating that every species originated in
close proximity to an already existing closely allied species. Like Darwin, it was
Wallace's consideration of how the ideas of Malthus might apply to animal populations
that led him to conclusions very similar to those reached by Darwin about the role of
natural selection. In February 1858 Wallace, unaware of Darwin's unpublished ideas,
composed his thoughts into an essay and mailed them to Darwin, asking for his opinion.
The result was the joint publication in July of an extract from Darwin's 1844 essay along
with Wallace's letter. Darwin also began work on a short abstract summarising his theory,
which he would publish in 1859 as On the Origin of Species.[64]

Diagram by O.C. Marsh of the evolution of horse feet and teeth over time as reproduced
in T.H Huxley's 1876 book Professor Huxley in America

[edit] 18591930s: Darwin and his legacy

See also: Reaction to Darwin's theory
By the 1850s whether or not species evolved was a subject of intense debate, with
prominent scientists arguing both sides of the issue.[65] However, it was the publication of

Charles Darwin's On the Origin of Species (1859) that fundamentally transformed the
discussion over biological origins.[66] Darwin argued that his branching version of
evolution explained a wealth of facts in biogeography, anatomy, embryology, and other
fields of biology. He also provided the first cogent mechanism by which evolutionary
change could persist: his theory of natural selection.[67]
One of the first and most important naturalists to be convinced by Origin of the reality of
evolution was the British anatomist Thomas Henry Huxley. Huxley recognized that
unlike the earlier transmutational ideas of Lamarck and Vestiges, Darwin's theory
provided a mechanism for evolution without supernatural involvement, even if Huxley
himself was not completely convinced that natural selection was the key evolutionary
mechanism. Huxley would make advocacy of evolution a cornerstone of the program of
the X Club to reform and professionalise science by displacing natural theology with
naturalism and to end the domination of British natural science by the clergy. By the early
1870s in English-speaking countries, thanks partly to these efforts, evolution had become
the mainstream scientific explanation for the origin of species.[67] In his campaign for
public and scientific acceptance of Darwin's theory, Huxley made extensive use of new
evidence for evolution from paleontology. This included evidence that birds had evolved
from reptiles, including the discovery of Archaeopteryx in Europe, and a number of
fossils of primitive birds with teeth found in North America. Another important line of
evidence was the finding of fossils that helped trace the evolution of the horse from its
small five-toed ancestors.[68] However, acceptance of evolution among scientists in nonEnglish speaking nations such as France, and the countries of southern Europe and Latin
America was slower. An exception to this was Germany, where both August Weismann
and Ernst Haeckel championed this idea: Haeckel used evolution to challenge the
established tradition of metaphysical idealism in German biology, much as Huxley used it
to challenge natural theology in Britain.[69] Haeckel and other German scientists would
take the lead in launching an ambitious programme to reconstruct the evolutionary
history of life based on morphology and embryology.[70]
Darwin's theory succeeded in profoundly altering scientific opinion regarding the
development of life and in producing a small philosophical revolution.[71] However, this
theory could not explain several critical components of the evolutionary process.
Specifically, Darwin was unable to explain the source of variation in traits within a
species, and could not identify a mechanism that could pass traits faithfully from one
generation to the next. Darwin's hypothesis of pangenesis, while relying in part on the
inheritance of acquired characteristics, proved to be useful for statistical models of
evolution that were developed by his cousin Francis Galton and the "biometric" school of
evolutionary thought. However, this idea proved to be of little use to other biologists.[72]

[edit] Application to humans

This illustration was the frontispiece of Thomas Henry Huxley's book Evidence as to
Man's Place in Nature (1863).
Charles Darwin was aware of the severe reaction in some parts of the scientific
community against the suggestion made in Vestiges of the Natural History of Creation
that humans had arisen from animals by a process of transmutation. Therefore he almost
completely ignored the topic of human evolution in The Origin of Species. Despite this
precaution, the issue featured prominently in the debate that followed the book's
publication. For most of the first half of the 19th century, the scientific community
believed that, although geology had shown that the Earth and life were very old, human
beings had appeared suddenly just a few thousand years before the present. However, a
series of archaeological discoveries in the 1840s and 1850s showed stone tools associated
with the remains of extinct animals. By the early 1860s, as summarized in Charles Lyell's
1863 book Geological Evidences of the Antiquity of Man, it had become widely accepted
that humans had existed during a prehistoric period which stretched many thousands of
years before the start of written history. This view of human history was more compatible
with an evolutionary origin for humanity than was the older view. On the other hand, at
that time there was no fossil evidence to demonstrate human evolution. The only human
fossils found before the discovery of Java man in the 1890s were either of anatomically
modern humans or of Neanderthals that were too close, especially in the critical
characteristic of cranial capacity, to modern humans for them to be convincing
intermediates between humans and other primates.[73]
Therefore the debate that immediately followed the publication of The Origin of Species
centered on the similarities and differences between humans and modern apes. Carolus
Linnaeus had been criticised in the 18th century for grouping humans and apes together
as primates in his ground breaking classification system.[74] Richard Owen vigorously
defended the classification suggested by Cuvier and Johann Friedrich Blumenbach that
placed humans in a separate order from any of the other mammals, which by the early
19th century had become the orthodox view. On the other hand, Thomas Henry Huxley
sought to demonstrate a close anatomical relationship between humans and apes. In one
famous incident, Huxley showed that Owen was mistaken in claiming that the brains of
gorillas lacked a structure present in human brains. Huxley summarized his argument in
his highly influential 1863 book Evidence as to Man's Place in Nature. Another
viewpoint was advocated by Charles Lyell and Alfred Russel Wallace. They agreed that
humans shared a common ancestor with apes, but questioned whether any purely

materialistic mechanism could account for all the differences between humans and apes,
especially some aspects of the human mind.[73]
In 1871, Darwin published The Descent of Man, and Selection in Relation to Sex, which
contained his views on human evolution. Darwin argued that the differences between the
human mind and the minds of the higher animals were a matter of degree rather than of
kind. For example, he viewed morality as a natural outgrowth of instincts that were
beneficial to animals living in social groups. He argued that all the differences between
humans and apes were explained by a combination of the selective pressures that came
from our ancestors moving from the trees to the plains, and sexual selection. The debate
over human origins, and over the degree of human uniqueness continued well into the
20th century.[73]

[edit] Alternatives to natural selection

Main article: The eclipse of Darwinism

This photo from Henry Fairfield Osborn's 1918 book Origin and Evolution of Life shows
models depicting the evolution of Titanothere horns over time, which Osborn claimed
was an example of an orthogenic trend in evolution.
The concept of evolution was widely accepted in scientific circles within a few years of
the publication of Origin, but the acceptance of natural selection as its driving mechanism
was much less widespread. The four major alternatives to natural selection in the late
19th century were theistic evolution, neo-Lamarckism, orthogenesis, and saltationism.
Theistic evolution (a term promoted by Darwin's greatest American advocate Asa Gray)
was the idea that God intervened in the process of evolution to guide it in such a way that
the living world could still be considered to be designed. However, this idea gradually
fell out of favor among scientists, as they became more and more committed to the idea
of methodological naturalism and came to believe that direct appeals to supernatural
involvement were scientifically unproductive. By 1900, theistic evolution had largely
disappeared from professional scientific discussions, although it retained a strong popular
following.[75][76]

In the late 19th century, the term neo-Lamarckism came to be associated with the position
of naturalists who viewed the inheritance of acquired characteristics as the most
important evolutionary mechanism. Advocates of this position included the British writer
and Darwin critic Samuel Butler, the German biologist Ernst Haeckel, and the American
paleontologist Edward Drinker Cope. They considered Lamarckism to be philosophically
superior to Darwin's idea of selection acting on random variation. Cope looked for, and
thought he found, patterns of linear progression in the fossil record. Inheritance of
acquired characteristics was part of Haeckel's recapitulation theory of evolution, which
held that the embryological development of an organism repeats its evolutionary history.
[75][76]
Critics of neo-Lamarckism, such as the German biologist August Weismann and
Alfred Russel Wallace, pointed out that no one had ever produced solid evidence for the
inheritance of acquired characteristics. Despite these criticisms, neo-Lamarckism
remained the most popular alternative to natural selection at the end of the 19th century,
and would remain the position of some naturalists well into the 20th century.[75][76]
Orthogenesis was the hypothesis that life has an innate tendency to change, in a unilinear
fashion, towards ever-greater perfection. It had a significant following in the 19th
century, and its proponents included the Russian biologist Leo Berg and the American
paleontologist Henry Fairfield Osborn. Orthogenesis was popular among some
paleontologists, who believed that the fossil record showed a gradual and constant
unidirectional change. Saltationism was the idea that new species arise as a result of large
mutations. It was seen as a much faster alternative to the Darwinian concept of a gradual
process of small random variations being acted on by natural selection, and was popular
with early geneticists such as Hugo de Vries, William Bateson, and early in his career, T.
H. Morgan. It became the basis of the mutation theory of evolution.[75][76]

Diagram from T.H. Morgan's 1919 book The Physical Basis of Heredity, showing the
sex-linked inheritance of the white-eyed mutation in Drosophila melanogaster

[edit] Mendelian genetics, biometrics, and mutation

The so-called rediscovery of Gregor Mendel's laws of inheritance in 1900 ignited a fierce
debate between two camps of biologists. In one camp were the Mendelians, who were
focused on discrete variations and the laws of inheritance. They were led by William
Bateson (who coined the word genetics) and Hugo de Vries (who coined the word

mutation). Their opponents were the biometricians, who were interested in the continuous
variation of characteristics within populations. Their leaders, Karl Pearson and Walter
Frank Raphael Weldon, followed in the tradition of Francis Galton, who had focused on
measurement and statistical analysis of variation within a population. The biometricians
rejected Mendelian genetics on the basis that discrete units of heredity, such as genes,
could not explain the continuous range of variation seen in real populations. Weldon's
work with crabs and snails provided evidence that selection pressure from the
environment could shift the range of variation in wild populations, but the Mendelians
maintained that the variations measured by biometricians were too insignificant to
account for the evolution of new species.[77][78]
When T. H. Morgan began experimenting with breeding the fruit fly Drosophila
melanogaster, he was a saltationist who hoped to demonstrate that a new species could be
created in the lab by mutation alone. Instead, the work at his lab between 1910 and 1915
reconfirmed Mendelian genetics and provided solid experimental evidence linking it to
chromosomal inheritance. His work also demonstrated that most mutations had relatively
small effects, such as a change in eye color, and that rather than creating a new species in
a single step, mutations served to increase variation within the existing population.[77][78]

[edit] 1920s1940s
See also: Modern evolutionary synthesis

Biston betularia f. typica is the white-bodied form of the peppered moth.

Biston betularia f. carbonaria is the black-bodied form of the peppered moth.

[edit] Population genetics

The Mendelian and biometrician models were eventually reconciled with the
development of population genetics. A key step was the work of the British biologist and
statistician R.A. Fisher. In a series of papers starting in 1918 and culminating in his 1930
book The Genetical Theory of Natural Selection, Fisher showed that the continuous

variation measured by the biometricians could be produced by the combined action of

many discrete genes, and that natural selection could change gene frequencies in a
population, resulting in evolution. In a series of papers beginning in 1924, another British
geneticist, J.B.S. Haldane, applied statistical analysis to real-world examples of natural
selection, such as the evolution of industrial melanism in peppered moths, and showed
that natural selection worked at an even faster rate than Fisher assumed.[79][80]
The American biologist Sewall Wright, who had a background in animal breeding
experiments, focused on combinations of interacting genes, and the effects of inbreeding
on small, relatively isolated populations that exhibited genetic drift. In 1932, Wright
introduced the concept of an adaptive landscape and argued that genetic drift and
inbreeding could drive a small, isolated sub-population away from an adaptive peak,
allowing natural selection to drive it towards different adaptive peaks. The work of
Fisher, Haldane and Wright founded the discipline of population genetics. This integrated
natural selection with Mendelian genetics, which was the critical first step in developing
a unified theory of how evolution worked.[79][80]

[edit] Modern evolutionary synthesis

In the first few decades of the 20th century, most field naturalists continued to believe
that Lamarckian and orthogenic mechanisms of evolution provided the best explanation
for the complexity they observed in the living world. But as the field of genetics
continued to develop, those views became less tenable.[81] Theodosius Dobzhansky, a
postdoctoral worker in T. H. Morgan's lab, had been influenced by the work on genetic
diversity by Russian geneticists such as Sergei Chetverikov. He helped to bridge the
divide between the foundations of microevolution developed by the population
geneticists and the patterns of macroevolution observed by field biologists, with his 1937
book Genetics and the Origin of Species. Dobzhansky examined the genetic diversity of
wild populations and showed that, contrary to the assumptions of the population
geneticists, these populations had large amounts of genetic diversity, with marked
differences between sub-populations. The book also took the highly mathematical work
of the population geneticists and put it into a more accessible form. In Great Britain E.B.
Ford, the pioneer of ecological genetics, continued throughout the 1930s and 1940s to
demonstrate the power of selection due to ecological factors including the ability to
maintain genetic diversity through genetic polymorphisms such as human blood types.
Ford's work would contribute to a shift in emphasis during the course of the modern
synthesis towards natural selection over genetic drift.[79][80][82][83]
Evolutionary biologist Ernst Mayr was influenced by the work of the German biologist
Bernhard Rensch showing the influence of local environmental factors on the geographic
distribution of sub-species and closely related species. Mayr followed up on
Dobzhansky's work with the 1942 book Systematics and the Origin of Species, which
emphasized the importance of allopatric speciation in the formation of new species. This
form of speciation occurs when the geographical isolation of a sub-population is followed
by the development of mechanisms for reproductive isolation. Mayr also formulated the
biological species concept that defined a species as a group of interbreeding or potentially

interbreeding populations that were reproductively isolated from all other populations.[79]
[80][84]

In the 1944 book Tempo and Mode in Evolution, George Gaylord Simpson showed that
the fossil record was consistent with the irregular non-directional pattern predicted by the
developing evolutionary synthesis, and that the linear trends that earlier paleontologists
had claimed supported orthogenesis and neo-Lamarckism did not hold up to closer
examination. In 1950, G. Ledyard Stebbins published Variation and Evolution in Plants,
which helped to integrate botany into the synthesis. The emerging cross-disciplinary
consensus on the workings of evolution would be known as the modern evolutionary
synthesis. It received its name from the book Evolution: The Modern Synthesis by Julian
Huxley.[79][80]
The evolutionary synthesis provided a conceptual corein particular, natural selection
and Mendelian population geneticsthat tied together many, but not all, biological
disciplines. It helped establish the legitimacy of evolutionary biology, a primarily
historical science, in a scientific climate that favored experimental methods over
historical ones.[85] The synthesis also resulted in a considerable narrowing of the range of
mainstream evolutionary thought (what Stephen Jay Gould called the "hardening of the
synthesis"): by the 1950s, natural selection acting on genetic variation was virtually the
only acceptable mechanism of evolutionary change (panselectionism), and
macroevolution was simply considered the result of extensive microevolution.[86][87]

[edit] 1940s1960s: Molecular biology and evolution

Main article: History of molecular evolution
The middle decades of the 20th century saw the rise of molecular biology, and with it an
understanding of the chemical nature of genes as sequences of DNA and their
relationship, through the genetic code, to protein sequences. At the same time,
increasingly powerful techniques for analyzing proteins, such as protein electrophoresis
and sequencing, brought biochemical phenomena into realm of the synthetic theory of
evolution. In the early 1960s, biochemists Linus Pauling and Emile Zuckerkandl
proposed the molecular clock hypothesis: that sequence differences between homologous
proteins could be used to calculate the time since two species diverged. By 1969, Motoo
Kimura and others provided a theoretical basis for the molecular clock, arguing thatat
the molecular level at leastmost genetic mutations are neither harmful nor helpful and
that genetic drift, rather than natural selection, is responsible for a large portion of genetic
change: the neutral theory of molecular evolution.[88] Studies of protein differences within
species also brought molecular data to bear on population genetics by providing estimates
of the level of heterozygosity in natural populations.[89]
From the early 1960s, molecular biology was increasingly seen as a threat to the
traditional core of evolutionary biology. Established evolutionary biologistsparticularly
Ernst Mayr, Theodosius Dobzhansky and G. G. Simpson, three of the architects of the
modern synthesiswere extremely skeptical of molecular approaches, especially when it

came to the connection (or lack thereof) to natural selection. The molecular clock
hypothesis and the neutral theory were particularly controversial, spawning the neutralistselectionist debate over the relative importance of drift and selection, which continued
into the 1980s without a clear resolution.[90][91]

[edit] Late 20th century

[edit] Gene-centered view
In the mid-1960s, George C. Williams strongly critiqued explanations of adaptations
worded in terms of "survival of the species" (group selection arguments). Such
explanations were largely replaced by a gene-centered view of evolution, epitomized by
the kin selection arguments of W. D. Hamilton, George R. Price and John Maynard
Smith.[92] This viewpoint would be summarized and popularized in the influential 1976
book The Selfish Gene by Richard Dawkins.[93] Models of the period showed that group
selection was severely limited in its strength; though newer models do admit the
possibility of significant multi-level selection.[94]
In 1973, Leigh Van Valen proposed the term "Red Queen", which he took from Through
the Looking-Glass by Lewis Carroll, to describe a scenario where a species involved in
one or more evolutionary arms races would have to constantly change just to keep pace
with the species with which it was co-evolving. Hamilton, Williams and others suggested
that this idea might explain the evolution of sexual reproduction: the increased genetic
diversity caused by sexual reproduction would help maintain resistance against rapidly
evolving parasites, thus making sexual reproduction common, despite the tremendous
cost from the gene-centric point of view of a system where only half of an organism's
genome is passed on during reproduction.[95][96] The gene-centric view has also led to an
increased interest in Darwin's old idea of sexual selection,[97] and more recently in topics
such as sexual conflict and intragenomic conflict.

[edit] Sociobiology
W. D. Hamilton's work on kin selection contributed to the emergence of the discipline of
sociobiology. The existence of altruistic behaviors has been a difficult problem for
evolutionary theorists from the beginning.[98] Significant progress was made in 1964
when Hamilton formulated the inequality in kin selection known as Hamilton's rule,
which showed how eusociality in insects (the existence of sterile worker classes) and
many other examples of altruistic behavior could have evolved through kin selection.
Other theories followed, some derived from game theory, such as reciprocal altruism.[99]
In 1975, E.O. Wilson published the influential and highly controversial book
Sociobiology: The New Synthesis which claimed evolutionary theory could help explain
many aspects of animal, including human, behavior. Critics of sociobiology, including
Stephen Jay Gould and Richard Lewontin, claimed that sociobiology greatly overstated
the degree to which complex human behaviors could be determined by genetic factors.
They also claimed that the theories of sociobiologists often reflected their own
ideological biases. Despite these criticisms, work has continued in sociobiology and the

related discipline of evolutionary psychology, including work on other aspects of the

altruism problem.[100][101]

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

[edit] Evolutionary paths and processes

One of the most prominent debates arising during the 1970s was over the theory of
punctuated equilibrium. Niles Eldredge and Stephen Jay Gould proposed that there was a
pattern of fossil species that remained largely unchanged for long periods (what they
termed stasis), interspersed with relatively brief periods of rapid change during
speciation.[102][103] Improvements in sequencing methods resulted in a large increase of
sequenced genomes, allowing the testing and refining of evolutionary theories using this
huge amount of genome data.[104] Comparisons between these genomes provide insights
into the molecular mechanisms of speciation and adaptation.[105][106] These genomic
analyses have produced fundamental changes in the understanding of the evolutionary
history of life, such as the proposal of the three-domain system by Carl Woese.[107]
Advances in computational hardware and software allow the testing and extrapolation of
increasingly advanced evolutionary models and the development of the field of systems
biology.[108] One of the results has been an exchange of ideas between theories of
biological evolution and the field of computer science known as evolutionary
computation, which attempts to mimic biological evolution for the purpose of developing
new computer algorithms. Discoveries in biotechnology now allow the modification of
entire genomes, advancing evolutionary studies to the level where future experiments
may involve the creation of entirely synthetic organisms.[109]

[edit] Microbiology, horizontal gene transfer, and endosymbiosis

Main article: Horizontal gene transfer
Microbiology was largely ignored by early evolutionary theory. This was due to the
paucity of morphological traits and the lack of a species concept in microbiology,
particularly amongst prokaryotes.[110] Now, evolutionary researchers are taking advantage
of their improved understanding of microbial physiology and ecology, produced by the
comparative ease of microbial genomics, to explore the taxonomy and evolution of these

organisms.[111] These studies are revealing unanticipated levels of diversity amongst

microbes.[112][113]
One particularly important outcome from studies on microbial evolution was the
discovery in Japan of horizontal gene transfer in 1959.[114] This transfer of genetic
material between different species of bacteria was first recognized because it played a
major role in the spread of antibiotic resistance.[115] More recently, as knowledge of
genomes has continued to expand, it has been suggested that lateral transfer of genetic
material has played an important role in the evolution of all organisms.[116] These high
levels of horizontal gene transfer have led to suggestions that the family tree of today's
organisms, the so-called "tree of life", is more similar to an interconnected web or net.[117]
[118]

Indeed, as part of the endosymbiotic theory for the origin of organelles, a form of
horizontal gene transfer has been a critical step in the evolution of eukaryotes such as
fungi, plants, and animals.[119][120] The endosymbiotic theory holds that organelles within
the cells of eukorytes such as mitochondria and chloroplasts, had descended from
independent bacteria that came to live symbiotically within other cells. It had been
suggested in the late 19th century when similarities between mitochondria and bacteria
were noted, but largely dismissed until it was revived and championed by Lynn Margulis
in the 1960s and 70s; Margulis was able to make use of new evidence that such
organelles had their own DNA that was inherited independently from that in the cell's
nucleus.[121]

[edit] Evolutionary developmental biology

Main article: Evolutionary developmental biology
In the 1980s and 1990s the tenets of the modern evolutionary synthesis came under
increasing scrutiny. There was a renewal of structuralist themes in evolutionary biology
in the work of biologists such as Brian Goodwin and Stuart Kauffman, which
incorporated ideas from cybernetics and systems theory, and emphasized the selforganizing processes of development as factors directing the course of evolution. The
evolutionary biologist Stephen Jay Gould revived earlier ideas of heterochrony,
alterations in the relative rates of developmental processes over the course of evolution,
to account for the generation of novel forms, and, with the evolutionary biologist Richard
Lewontin, wrote an influential paper in 1979 suggesting that a change in one biological
structure, or even a structural novelty, could arise incidentally as an accidental result of
selection on another structure, rather than through direct selection for that particular
adaptation. They called such incidental structural changes "spandrels" after an
architectural feature.[122] Later, Gould and Vrba discussed the acquisition of new functions
by novel structures arising in this fashion, calling them "exaptations".[123]
Molecular data regarding the mechanisms underlying development accumulated rapidly
during the 1980s and '90s. It became clear that the diversity of animal morphology was
not the result of different sets of proteins regulating the development of different animals,

but from changes in the deployment of a small set of proteins that were common to all
animals.[124] These proteins became known as the "developmental toolkit".[125] Such
perspectives influenced the disciplines of phylogenetics, paleontology and comparative
developmental biology, and spawned the new discipline of evolutionary developmental
biology.[126]
More recent work in this field by Mary Jane West-Eberhard has emphasized phenotypic
and developmental plasticity.[127] It has been suggested, for example, that the rapid
emergence of basic animal body plans in the Cambrian explosion was due in part to
changes in the environment acting on inherent material properties of cell aggregates, such
as differential cell adhesion and biochemical oscillation. The resulting forms were later
stabilized by natural selection.[128] Experimental and theoretical research on these and
related ideas have been presented in the multi-authored volume Origination of
Organismal Form.

[edit] 21st century

[edit] Epigenetic inheritance
Main article: Epigenetics
Yet another area where developmental biology has led to the questioning of some tenets
of the modern evolutionary synthesis is in the field of epigenetics, the study of the effect
of environmental factors on the way genes express themselves during development. By
the first decade of the 21st century it had become accepted that in some cases such
environmental factors could affect the expression of genes in subsequent generations
even though the offspring were not exposed to the same environmental factors, and there
had been no genetic changes. This shows that in some cases non genetic changes to an
organism can be inherited and it has been suggested that such inheritance can help with
adaptation to local conditions and affect evolution.[129][130] Some have suggested that in
some cases a form of Lamarckian evolution may occur.[131]

[edit] Unconventional evolutionary theory

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

[edit] Gaia hypothesis

Teilhard de Chardin's ideas have been seen as being connected to the more specific Gaia
theory by James Lovelock, who proposed that the living and nonliving parts of Earth can
be viewed as a complex interacting system with similarities to a single organism.[132] The

Gaia hypothesis has also been viewed by Lynn Margulis[133] and others as an extension of
endosymbiosis and exosymbiosis.[134] This modified hypothesis postulates that all living
things have a regulatory effect on the Earth's environment that promotes life overall.

[edit] Transhumanism
Futurists have often viewed scientific and technological progress as a continuation of
biological evolution. Among these, transhumanists often view such technological
evolution itself as a goal in their philosophy, possibly in the form of a technological
singularity.

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Evolutionary history of life

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Adaptation
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The evolutionary history of life on Earth traces the processes by which living and fossil
organisms evolved. It stretches from the origin of life on Earth, thought to be over 3,500
million years ago, to the present day. The similarities between all present day organisms
indicate the presence of a common ancestor from which all known species have diverged
through the process of evolution.[1]
Microbial mats of coexisting bacteria and archaea were the dominant form of life in the
early Archean and many of the major steps in early evolution are thought to have taken

place within them.[2] The evolution of oxygenic photosynthesis, around 3,500 million
years ago, eventually led to the oxygenation of the atmosphere, beginning around 2,400
million years ago.[3] While eukaryotic cells may have been present earlier, their evolution
accelerated when they began to use oxygen in their metabolism. The earliest evidence of
complex eukaryotes with organelles, dates from 1,850 million years ago. Later, around
1,700 million years ago, multicellular organisms began to appear, with differentiated cells
performing specialised functions.[4]
The earliest land plants date back to around 450 million years ago,[5] though evidence
suggests that algal scum formed on the land as early as 1,200 million years ago. Land
plants were so successful that they are thought to have contributed to the late Devonian
extinction event.[6] Invertebrate animals appear during the Vendian period,[7] while
vertebrates originated about 525 million years ago during the Cambrian explosion.[8]
During the Permian period, synapsids, including the ancestors of mammals, dominated
the land,[9] but the PermianTriassic extinction event 251 million years ago came close to
wiping out all complex life.[10] During the recovery from this catastrophe, archosaurs
became the most abundant land vertebrates, displacing therapsids in the mid-Triassic.[11]
One archosaur group, the dinosaurs, dominated the Jurassic and Cretaceous periods,[12]
while the ancestors of mammals survived only as small insectivores.[13] After the
CretaceousTertiary extinction event 65 million years ago killed off the non-avian
dinosaurs[14] mammals increased rapidly in size and diversity.[15] Such mass extinctions
may have accelerated evolution by providing opportunities for new groups of organisms
to diversify.[16]
Fossil evidence indicates that flowering plants appeared and rapidly diversified in the
Early Cretaceous, between 130 million years ago and 90 million years ago, probably
helped by coevolution with pollinating insects. Flowering plants and marine
phytoplankton are still the dominant producers of organic matter. Social insects appeared
around the same time as flowering plants. Although they occupy only small parts of the
insect "family tree", they now form over half the total mass of insects. Humans evolved
from a lineage of upright-walking apes whose earliest fossils date from over 6 million
years ago. Although early members of this lineage had chimp-sized brains, there are signs
of a steady increase in brain size after about 3 million years ago.

Contents
[hide]

1 Earliest history of Earth

2 Earliest evidence for life on Earth
3 Origins of life on Earth
o 3.1 Life "seeded" from elsewhere
o 3.2 Independent emergence on Earth
3.2.1 Replication first: RNA world
3.2.2 Metabolism first: Iron-sulfur world
3.2.3 Membranes first: Lipid world
3.2.4 The clay theory

4 Environmental and evolutionary impact of microbial mats

5 Diversification of eukaryotes
6 Multicellular organisms and sexual reproduction
o 6.1 Multicellularity
o 6.2 Evolution of sexual reproduction
o 6.3 Fossil evidence for multicellularity and sexual reproduction
7 Emergence of animals
8 Colonization of land
o 8.1 Evolution of soil
o 8.2 Plants and the Late Devonian wood crisis
o 8.3 Land invertebrates
o 8.4 Land vertebrates
9 Dinosaurs, birds and mammals
10 Flowering plants
11 Social insects
12 Humans
13 Mass extinctions
14 The present
15 See also
16 Footnotes
17 References
18 Further reading

19 External links

[edit] Earliest history of Earth

History of Earth and its life
-4500

-4000

-3500

-3000

-2500

-2000

-1500

-1000

-500

Hadean

Archean
Protero
-zoic
Phanero
-zoic
Eo
Paleo
Meso
Neo
Paleo
Meso
Neo
Paleo
Meso
Ceno

Solar system formed

Impact formed Moon

? Cool surface, oceans, atmosphere

Late Heavy Bombardment

? Earliest evidence of life

Oxygenation of atmosphere

Earliest multicellular organism

Earliest known fungi

Earliest known cnidarians

? Cambrian explosion

Earliest land invertebrates and plants

Earliest land vertebrates

Earliest known dinosaur

Extinction of non-avian dinosaurs

Scale:
Millions of years

Main article: History of the Earth

The oldest meteorite fragments found on Earth are about 4,540 million years old, and this
has convinced scientists that the whole Solar system, including Earth, formed around that
time.[17] About 40 million years later a planetoid struck the Earth, throwing into orbit the
material that formed the Moon.[18]
Until recently the oldest rocks found on Earth were about 3,800 million years old,[17] and
this led scientists to believe for decades that Earth's surface was molten until then. Hence
they named this part of Earth's history the Hadean eon, whose name means "hellish".[19]
However analysis of zircons formed 4,400 to 4,000 million years ago indicates that
Earth's crust solidified about 100 million years after the planet's formation and that Earth
quickly acquired oceans and an atmosphere, which may have been capable of supporting
life.[20]
Evidence from the Moon indicates that from 4,000 to 3,800 million years ago it suffered a
Late Heavy Bombardment by debris that was left over from the formation of the Solar
system, and Earth, having stronger gravity, should have experienced an even heavier
bombardment.[19][21] While there is no direct evidence of conditions on Earth
4,000 to 3,800 million years ago, there is no reason to think that the Earth was not also
affected by this late heavy bombardment.[22] This event may well have stripped away any
previous atmosphere and oceans; in this case gases and water from comet impacts may
have contributed to their replacement, although volcanic outgassing on Earth would have
contributed at least half.[23]

[edit] Earliest evidence for life on Earth

The earliest identified organisms were minute and relatively featureless, so their fossils
look like small rods, which are very difficult to tell apart from structures which form
through physical processes. The oldest undisputed evidence of life on Earth, interpreted
as fossilized bacteria, dates to 3,000 million years ago.[24] Other finds in rocks dated to
about 3,500 million years ago have been interpreted as bacteria,[25] and geochemical
evidence seemed to show the presence of life 3,800 million years ago.[26] However these
analyses were closely scrutinized, and non-biological processes were found which could
produce all of the "signatures of life" that had been reported.[27][28] While this does not
prove that the structures found had a non-biological origin, they cannot be taken as clear
evidence for the presence of life. Currently, the oldest unchallenged evidence for life is
geochemical signatures from rocks deposited 3,400 million years ago,[24][29] although there
has been little time for these recent reports (2006) to be examined by critics.

[edit] Origins of life on Earth

Evolutionary tree showing the divergence of modern species from their common ancestor
in the center.[30] The three domains are colored, with bacteria blue, archaea green, and
eukaryotes red.
Further information: Evidence of common descent, Common descent, and Homology
(biology)
Biochemists reason that all living organisms on Earth must share a single last universal
ancestor, because it would be virtually impossible that two or more separate lineages
could have independently developed the many complex biochemical mechanisms shared
by all living organisms.[31][32] However the earliest organisms for which fossil evidence is
available are bacteria, which are far too complex to have arisen directly from non-living
materials.[33] The lack of fossil or geochemical evidence for earlier types of organism has
left plenty of scope for hypotheses, which fall into two main groups: that life arose
spontaneously on Earth, and that it was "seeded" from elsewhere in the universe.[34]

[edit] Life "seeded" from elsewhere

Main articles: Panspermia, Life on Mars, Fermi paradox, and Rare Earth hypothesis
The idea that life Earth was "seeded" from elsewhere in the universe dates back at least to
the fifth century BC.[35] In the twentieth century it was proposed by the physical chemist
Svante Arrhenius,[36] by the astronomers Fred Hoyle and Chandra Wickramasinghe,[37] and
by molecular biologist Francis Crick and chemist Leslie Orgel.[38] There are three main
versions of the "seeded from elsewhere" hypothesis: from elsewhere in our Solar system
via fragments knocked into space by a large meteor impact, in which case the only
credible source is Mars;[39] by alien visitors, possibly as a result of accidental
contamination by micro-organisms that they brought with them;[38] and from outside the
Solar system but by natural means.[36][39] Experiments suggest that some micro-organisms
can survive the shock of being catapulted into space and some can survive exposure to
radiation for several days, but there is no proof that they can survive in space for much

longer periods.[39] Scientists are divided over the likelihood of life arising independently
on Mars,[40] or on other planets in our galaxy.[39]

[edit] Independent emergence on Earth

Main article: Abiogenesis
Life on earth is based on carbon and water. Carbon provides stable frameworks for
complex chemicals and can be easily extracted from the environment, especially from
carbon dioxide. The only other element with similar chemical properties, silicon, forms
much less stable structures and, because most of its compounds are solids, would be more
difficult for organisms to extract. Water is an excellent solvent and has two other useful
properties: the fact that ice floats enables aquatic organisms to survive beneath it in
winter; and its molecules have electrically negative and positive ends, which enables it to
form a wider range of compounds than other solvents can. Other good solvents, such as
ammonia, are liquid only at such low temperatures that chemical reactions may be too
slow to sustain life, and lack water's other advantages.[41] Organisms based on alternative
biochemistry may however be possible on other planets.[42]
Research on how life might have emerged unaided from non-living chemicals focuses on
three possible starting points: self-replication, an organism's ability to produce offspring
that are very similar to itself; metabolism, its ability to feed and repair itself; and external
cell membranes, which allow food to enter and waste products to leave, but exclude
unwanted substances.[43] Research on abiogenesis still has a long way to go, since
theoretical and empirical approaches are only beginning to make contact with each other.
[44][45]

[edit] Replication first: RNA world

Main articles: Last universal ancestor and RNA world

The replicator in virtually all known life is deoxyribonucleic acid. DNA's structure and
replication systems are far more complex than those of the original replicator.[33]

Even the simplest members of the three modern domains of life use DNA to record their
"recipes" and a complex array of RNA and protein molecules to "read" these instructions
and use them for growth, maintenance and self-replication. This system is far too
complex to have emerged directly from non-living materials.[33] The discovery that some
RNA molecules can catalyze both their own replication and the construction of proteins
led to the hypothesis of earlier life-forms based entirely on RNA.[46] These ribozymes
could have formed an RNA world in which there were individuals but no species, as
mutations and horizontal gene transfers would have meant that the offspring in each
generation were quite likely to have different genomes from those that their parents
started with.[47] RNA would later have been replaced by DNA, which is more stable and
therefore can build longer genomes, expanding the range of capabilities a single organism
can have.[47][48][49] Ribozymes remain as the main components of ribosomes, modern cells'
"protein factories".[50]
Although short self-replicating RNA molecules have been artificially produced in
laboratories,[51] doubts have been raised about where natural non-biological synthesis of
RNA is possible.[52] The earliest "ribozymes" may have been formed of simpler nucleic
acids such as PNA, TNA or GNA, which would have been replaced later by RNA.[53][54]
In 2003 it was proposed that porous metal sulfide precipitates would assist RNA
synthesis at about 100 C (212 F) and ocean-bottom pressures near hydrothermal vents.
In this hypothesis lipid membranes would be the last major cell components to appear
and until then the proto-cells would be confined to the pores.[55]
[edit] Metabolism first: Iron-sulfur world
Main article: Iron-sulfur world theory

A series of experiments starting in 1997 showed that early stages in the formation of
proteins from inorganic materials including carbon monoxide and hydrogen sulfide could
be achieved by using iron sulfide and nickel sulfide as catalysts. Most of the steps
required temperatures of about 100 C (212 F) and moderate pressures, although one
stage required 250 C (482 F) and a pressure equivalent to that found under 7 kilometres
(4.3 mi) of rock. Hence it was suggested that self-sustaining synthesis of proteins could
have occurred near hydrothermal vents.[56]
[edit] Membranes first: Lipid world

= water-attracting heads of lipid molecules

= water-repellent tails

Cross-section through a liposome.

It has been suggested that double-walled "bubbles" of lipids like those that form the
external membranes of cells may have been an essential first step.[57] Experiments that
simulated the conditions of the early Earth have reported the formation of lipids, and
these can spontaneously form liposomes, double-walled "bubbles", and then reproduce
themselves. Although they are not intrinsically information-carriers as nucleic acids are,
they would be subject to natural selection for longevity and reproduction. Nucleic acids
such as RNA might then have formed more easily within the liposomes than they would
have outside.[58]
[edit] The clay theory
Main articles: Graham Cairns-Smith#Clay Theory and RNA world

RNA is complex and there are doubts about whether it can be produced non-biologically
in the wild.[52] Some clays, notably montmorillonite, have properties that make them
plausible accelerators for the emergence of an RNA world: they grow by self-replication
of their crystalline pattern; they are subject to an analog of natural selection, as the clay
"species" that grows fastest in a particular environment rapidly becomes dominant; and
they can catalyze the formation of RNA molecules.[59] Although this idea has not become
the scientific consensus, it still has active supporters.[60]
Research in 2003 reported that montmorillonite could also accelerate the conversion of
fatty acids into "bubbles", and that the "bubbles" could encapsulate RNA attached to the
clay. These "bubbles" can then grow by absorbing additional lipids and then divide. The
formation of the earliest cells may have been aided by similar processes.[61]
A similar hypothesis presents self-replicating iron-rich clays as the progenitors of
nucleotides, lipids and amino acids.[62]

[edit] Environmental and evolutionary impact of

microbial mats
Main articles: Microbial mat and Oxygen catastrophe

Modern stromatolites in Shark Bay, Western Australia.

Microbial mats are multi-layered, multi-species colonies of bacteria and other organisms
that are generally only a few millimeters thick, but still contain a wide range of chemical
environments, each of which favors a different set of micro-organisms.[63] To some extent
each mat forms its own food chain, as the by-products of each group of micro-organisms
generally serve as "food" for adjacent groups.[64]
Stromatolites are stubby pillars built as microbes in mats slowly migrate upwards to
avoid being smothered by sediment deposited on them by water.[63] There has been
vigorous debate about the validity of alleged fossils from before 3,000 million years ago,
[65]
with critics arguing that so-called stromatolites could have been formed by nonbiological processes.[27] In 2006 another find of stromatolites was reported from the same
part of Australia as previous ones, in rocks dated to 3,500 million years ago.[66]

In modern underwater mats the top layer often consists of photosynthesizing

cyanobacteria which create an oxygen-rich environment, while the bottom layer is
oxygen-free and often dominated by hydrogen sulfide emitted by the organisms living
there.[64] It is estimated that the appearance of oxygenic photosynthesis by bacteria in
mats increased biological productivity by a factor of between 100 and 1,000. The
reducing agent used by oxygenic photosynthesis is water, which is much more plentiful
than the geologically-produced reducing agents required by the earlier non-oxygenic
photosynthesis.[67] From this point onwards life itself produced significantly more of the
resources it needed than did geochemical processes.[68] Oxygen is toxic to organisms that
are not adapted to it, but greatly increases the metabolic efficiency of oxygen-adapted
organisms.[69][70]
Oxygen became a significant component of Earth's atmosphere about 2,400 million years
ago.[71] Although eukaryotes may have been present much earlier,[72][73] the oxygenation of
the atmosphere was a prerequisite for the evolution of the most complex eukaryotic cells,
from which all multicellular organisms are built.[74] The boundary between oxygen-rich
and oxygen-free layers in microbial mats would have moved upwards when
photosynthesis shut down overnight, and then downwards as it resumed on the next day.
This would have created selection pressure for organisms in this intermediate zone to
acquire the ability to tolerate and then to use oxygen, possibly via endosymbiosis, where
one organism lives inside another and both of them benefit from their association.[2]
Cyanobacteria have the most complete biochemical "toolkits" of all the mat-forming
organisms. Hence they are the most self-sufficient of the mat organisms and were welladapted to strike out on their own both as floating mats and as the first of the
phytoplankton, providing the basis of most marine food chains.[2]

[edit] Diversification of eukaryotes

Eukaryotes Bikonta

Apusozoa
Archaeplastida (Land plants, green algae, red algae, and
glaucophytes)
Chromalveolata
Rhizaria
Excavata

Unikonta

Amoebozoa
Opisthokonta

Metazoa (Animals)
Choanozoa
Eumycota (Fungi)

One possible family tree of eukaryotes

[75][76]

Main article: Eukaryote

Eukaryotes may have been present long before the oxygenation of the atmosphere,[72] but
most modern eukaryotes require oxygen, which their mitochondria use to fuel the
production of ATP, the internal energy supply of all known cells.[74] In the 1970s it was
proposed and, after much debate, widely accepted that eukaryotes emerged as a result of
a sequence of endosymbioses between "procaryotes". For example: a predatory microorganism invaded a large procaryote, probably an archaean, but the attack was
neutralized, and the attacker took up residence and evolved into the first of the
mitochondria; one of these chimeras later tried to swallow a photosynthesizing
cyanobacterium, but the victim survived inside the attacker and the new combination
became the ancestor of plants; and so on. After each endosymbiosis began, the partners
would have eliminated unproductive duplication of genetic functions by re-arranging
their genomes, a process which sometimes involved transfer of genes between them.[77][78]
[79]
Another hypothesis proposes that mitochondria were originally sulfur- or hydrogenmetabolising endosymbionts, and became oxygen-consumers later.[80] On the other hand
mitochondria might have been part of eukaryotes' original equipment.[81]
There is a debate about when eukaryotes first appeared: the presence of steranes in
Australian shales may indicate that eukaryotes were present 2,700 million years ago;[73]
however an analysis in 2008 concluded that these chemicals infiltrated the rocks less than
2,200 million years ago and prove nothing about the origins of eukaryotes.[82] Fossils of
the alga Grypania have been reported in 1,850 million-year-old rocks (originally dated to
2,100 million years ago but later revised[83]), and indicates that eukaryotes with organelles
had already evolved.[84] A diverse collection of fossil algae were found in rocks dated
between 1,500 million years ago and 1,400 million years ago.[85] The earliest known
fossils of fungi date from 1,430 million years ago.[86]

[edit] Multicellular organisms and sexual reproduction

[edit] Multicellularity
Main articles: Multicellular organism , Evolution of multicellularity , and Sexual
reproduction

A slime mold solves a maze. The mold (yellow) explored and filled the maze (left). When
the researchers placed sugar (red) at two separate points, the mold concentrated most of
its mass there and left only the most efficient connection between the two points (right).
[87]

The simplest definitions of "multicellular", for example "having multiple cells", could
include colonial cyanobacteria like Nostoc. Even a professional biologist's definition such
as "having the same genome but different types of cell" would still include some genera
of the green alga Volvox, which have cells that specialize in reproduction.[88]
Multicellularity evolved independently in organisms as diverse as sponges and other
animals, fungi, plants, brown algae, cyanobacteria, slime moulds and myxobacteria.[83][89]
For the sake of brevity this article focuses on the organisms that show the greatest
specialization of cells and variety of cell types, although this approach to the evolution of
complexity could be regarded as "rather anthropocentric".[90]
The initial advantages of multicellularity may have included: increased resistance to
predators, many of which attacked by engulfing; the ability to resist currents by attaching
to a firm surface; the ability to reach upwards to filter-feed or to obtain sunlight for
photosynthesis;[91] the ability to create an internal environment that gives protection
against the external one;[90] and even the opportunity for a group of cells to behave
"intelligently" by sharing information.[87] These features would also have provided
opportunities for other organisms to diversify, by creating more varied environments than
flat microbial mats could.[91]
Multicellularity with differentiated cells is beneficial to the organism as a whole but
disadvantageous from the point of view of individual cells, most of which lose the
opportunity to reproduce themselves. In an asexual multicellular organism, rogue cells
which retain the ability to reproduce may take over and reduce the organism to a mass of
undifferentiated cells. Sexual reproduction eliminates such rogue cells from the next
generation and therefore appears to be a prerequisite for complex multicellularity.[91]
The available evidence indicates that eukaryotes evolved much earlier but remained
inconspicuous until a rapid diversification around 1,000 million years ago. The only
respect in which eukaryotes clearly surpass bacteria and archaea is their capacity for
variety of forms, and sexual reproduction enabled eukaryotes to exploit that advantage by
producing organisms with multiple cells that differed in form and function.[91]

[edit] Evolution of sexual reproduction

Main article: Evolution of sexual reproduction
The defining characteristic of sexual reproduction is recombination, in which each of the
offspring receives 50% of its genetic inheritance from each of the parents.[92] Bacteria also
exchange DNA by bacterial conjugation, the benefits of which include resistance to
antibiotics and other toxins, and the ability to utilize new metabolites.[93] However
conjugation is not a means of reproduction, and is not limited to members of the same
species there are cases where bacteria transfer DNA to plants and animals.[94]
The disadvantages of sexual reproduction are well-known: the genetic reshuffle of
recombination may break up favorable combinations of genes; and since males do not
directly increase the number of offspring in the next generation, an asexual population

can out-breed and displace in as little as 50 generations a sexual population that is equal
in every other respect.[92] Nevertheless the great majority of animals, plants, fungi and
protists reproduce sexually. There is strong evidence that sexual reproduction arose early
in the history of eukaryotes and that the genes controlling it have changed very little since
then.[95] How sexual reproduction evolved and survived is an unsolved puzzle.[96]
The Red Queen Hypothesis suggests that sexual reproduction provides protection against
parasites, because it is easier for parasites to evolve means of overcoming the defenses of
genetically identical clones than those of sexual species that present moving targets, and
there is some experimental evidence for this. However there is still doubt about whether
it would explain the survival of sexual species if multiple similar clone species were
present, as one of the clones may survive the attacks of parasites for long enough to outbreed the sexual species.[92]
The Mutation Deterministic Hypothesis assumes that each organism has more than one
harmful mutation and the combined effects of these mutations are more harmful than the
sum of the harm done by each individual mutation. If so, sexual recombination of genes
will reduce the harm done that bad mutations do to offspring and at the same time
eliminate some bad mutations from the gene pool by isolating them in individuals that
perish quickly because they have an above-average number of bad mutations. However
the evidence suggests that the MDH's assumptions are shaky, because many species have
on average less than one harmful mutation per individual and no species that has been
investigated shows evidence of synergy between harmful mutations.[92]
The random nature of recombination causes the relative abundance of alternative traits to
vary from one generation to another. This genetic drift is insufficient on its own to make
sexual reproduction advantageous, but a combination of genetic drift and natural
selection may be sufficient. When chance produces combinations of good traits, natural
selection gives a large advantage to lineages in which these traits become genetically
linked. On the other hand the benefits of good traits are neutralized if they appear along
with bad traits. Sexual recombination gives good traits the opportunities to become linked
with other good traits, and mathematical models suggest this may be more than enough to
offset the disadvantages of sexual reproduction.[96] Other combinations of hypotheses that
are inadequate on their own are also being examined.[92]

[edit] Fossil evidence for multicellularity and sexual reproduction

Horodyskia may have been an early metazoan,[83] or a colonial foraminiferan[97]

The earliest known fossil organism that is clearly multicellular, Qingshania,[note 1] dated to
1,700 million years ago, appears to consist of virtually identical cells. A red alga called
Bangiomorpha, dated at 1,200 million years ago, is the earliest known organism which
has differentiated, specialized cells, and is also the oldest known sexually-reproducing
organism.[91] The 1,430 million-year-old fossils interpreted as fungi appear to have been
multicellular with differentiated cells.[86] The "string of beads" organism Horodyskia,
found in rocks dated from 1,500 million years ago to 900 million years ago, may have
been an early metazoan;[83] however it has also been interpreted as a colonial
foraminiferan.[97]

[edit] Emergence of animals

Main articles: Animal, Ediacara biota, Cambrian Explosion, Burgess shale type fauna,
and Stem group
Bilaterians
Deuterostomes (chordates, hemichordates,
echinoderms)
Protostomes
Ecdysozoa (anthropods, nematodes,
tardigrades, etc.)
Lophotrochozoa (molluscs, annelids,
brachiopods, etc.)
Acoelomorpha
Cnidaria (jellyfish, sea anemones, hydras)
Ctenophora (comb jellies)
Placozoa
Porifera (sponges): Calcarea
Porifera: Hexactinellida & Demospongiae
Choanoflagellata
Mesomycetozoea
A family tree of the animals.[98]

Animals are multicellular eukaryotes,[note 2] and are distinguished from plants, algae, and
fungi by lacking cell walls.[99] All animals are motile,[100] if only at certain life stages. All
animals except sponges have bodies differentiated into separate tissues, including
muscles, which move parts of the animal by contracting, and nerve tissue, which
transmits and processes signals.[101]
The earliest widely-accepted animal fossils are rather modern-looking cnidarians (the
group that includes jellyfish, sea anemones and hydras), possibly from around 580
million years ago, although fossils from the Doushantuo Formation can only be dated
approximately. Their presence implies that the cnidarian and bilaterian lineages had
already diverged.[102]
The Ediacara biota, which flourished for the last 40 million years before the start of the
Cambrian,[103] were the first animals more than a very few centimeters long. Many were
flat and had a "quilted" appearance, and seemed so strange that there was a proposal to
classify them as a separate kingdom, Vendozoa.[104] Others, however, been interpreted as
early molluscs (Kimberella[105][106]), echinoderms (Arkarua[107]), and arthropods
(Spriggina,[108] Parvancorina[109]). There is still debate about the classification of these
specimens, mainly because the diagnostic features which allow taxonomists to classify
more recent organisms, such as similarities to living organisms, are generally absent in
the Ediacarans. However there seems little doubt that Kimberella was at least a
triploblastic bilaterian animal, in other words significantly more complex than cnidarians.
[110]

The small shelly fauna are a very mixed collection of fossils found between the Late
Ediacaran and Mid Cambrian periods. The earliest, Cloudina, shows signs of successful
defense against predation and may indicate the start of an evolutionary arms race. Some
tiny Early Cambrian shells almost certainly belonged to molluscs, while the owners of
some "armor plates", Halkieria and Microdictyon, were eventually identified when more
complete specimens were found in Cambrian lagersttten that preserved soft-bodied
animals.[111]

Opabinia made the largest single contribution to modern interest in the Cambrian
explosion.[112]
In the 1970s there was already a debate about whether the emergence of the modern
phyla was "explosive" or gradual but hidden by the shortage of Pre-Cambrian animal
fossils.[111] A re-analysis of fossils from the Burgess Shale lagersttte increased interest in
the issue when it revealed animals, such as Opabinia, which did not fit into any known

phylum. At the time these were interpreted as evidence that the modern phyla had
evolved very rapidly in the "Cambrian explosion" and that the Burgess Shale's "weird
wonders" showed that the Early Cambrian was a uniquely experimental period of animal
evolution.[113] Later discoveries of similar animals and the development of new theoretical
approaches led to the conclusion that many of the "weird wonders" were evolutionary
"aunts" or "cousins" of modern groups[114] for example that Opabinia was a member of
the lobopods, a group which includes the ancestors of the arthropods, and that it may
have been closely related to the modern tardigrades.[115] Nevertheless there is still much
debate about whether the Cambrian explosion was really explosive and, if so, how and
why it happened and why it appears unique in the history of animals.[116]

Acanthodians were among the earliest vertebrates with jaws[117]

Most of the animals at the heart of the Cambrian explosion debate are protostomes, one
of the two main groups of complex animals. One deuterostome group, the echinoderms,
many of which have hard calcite "shells", are fairly common from the Early Cambrian
small shelly fauna onwards.[111] Other deuterostome groups are soft-bodied, and most of
the significant Cambrian deuterostome fossils come from the Chengjiang fauna, a
lagersttte in China.[118] The Chengjiang fossils Haikouichthys and Myllokunmingia
appear to be true vertebrates,[119] and Haikouichthys had distinct vertebrae, which may
have been slightly mineralized.[120] Vertebrates with jaws, such as the Acanthodians, first
appeared in the Late Ordovician.[121]

[edit] Colonization of land

Adaptation to life on land is a major challenge: all land organisms need to avoid dryingout and all those above microscopic size have to resist gravity; respiration and gas
exchange systems have to change; reproductive systems cannot depend on water to carry
eggs and sperm towards each other.[122][123] Although the earliest good evidence of land
plants and animals dates back to the Ordovician period (488 to 444 million years ago),
modern land ecosystems only appeared in the late Devonian, about 385 to 359 million
years ago.[124]

[edit] Evolution of soil

Before the colonization of land, soil, a combination of mineral particles and decomposed
organic matter, did not exist. Land surfaces would have been either bare rock or unstable
sand produced by weathering. Water and any nutrients in it would have drained away
very quickly.[124]

Lichens growing on concrete

Films of cyanobacteria, which are not plants but use the same photosynthesis
mechanisms, have been found in modern deserts, and only in areas that are unsuitable for
vascular plants. This suggests that microbial mats may have been the first organisms to
colonize dry land, possibly in the Precambrian. Mat-forming cyanobacteria could have
gradually evolved resistance to desiccation as they spread from the seas to tidal zones and
then to land.[124] Lichens, which are symbiotic combinations of a fungus (almost always
an ascomycete) and one or more photosynthesizers (green algae or cyanobacteria),[125] are
also important colonizers of lifeless environments,[124] and their ability to break down
rocks contributes to soil formation in situations where plants cannot survive.[125] The
earliest known ascomycete fossils date from 423 to 419 million years ago in the Silurian.
[124]

Soil formation would have been very slow until the appearance of burrowing animals,
which mix the mineral and organic components of soil and whose feces are a major
source of the organic components.[124] Burrows have been found in Ordovician sediments,
and are attributed to annelids ("worms") or arthropods.[124][126]

[edit] Plants and the Late Devonian wood crisis

Main article: Evolutionary history of plants

Reconstruction of Cooksonia, a vascular plant from the Silurian.

Fossilized trees from the Mid-Devonian Gilboa fossil forest.

In aquatic algae, almost all cells are capable of photosynthesies and are nearly
independent. Life on land required plants to become internally more complex and
specialized: photosynthesis was most efficient at the top; roots were required in order to
extract water from the ground; the parts in between became supports and transport
systems for water and nutrients.[122][127]
Spores of land plants, possibly rather like liverworts, have been found in Mid Ordovician
rocks dated to about 476 million years ago. In Mid Silurian rocks 430 million years ago
there are fossils of actual plants including clubmosses such as Baragwanathia; most were
under 10 centimetres (3.9 in) high, and some appear closely related to vascular plants, the
group that includes trees.[127]
By the Late Devonian 370 million years ago, trees such as Archaeopteris were so
abundant that they changed river systems from mostly braided to mostly meandering,
because their roots bound the soil firmly.[128] In fact they caused a "Late Devonian wood
crisis",[129] because:

They removed more carbon dioxide from the atmosphere, reducing the
greenhouse effect and thus causing an ice age in the Carboniferous period.[130] In
later ecosystems the carbon dioxide "locked up" in wood is returned to the
atmosphere by decomposition of dead wood. However the earliest fossil evidence
of fungi that can decompose wood also comes from the Late Devonian.[131]
The increasing depth of plants' roots led to more washing of nutrients into rivers
and seas by rain. This caused algal blooms whose high consumption of oxygen
caused anoxic events in deeper waters, increasing the extinction rate among deepwater animals.[130]

[edit] Land invertebrates

Animals had to change their feeding and excretory systems, and most land animals
developed internal fertilization of their eggs. The difference in refractive index between
water and air required changes in their eyes. On the other hand in some ways movement
and breathing became easier, and the better transmission of high-frequency sounds in air
encouraged the development of hearing.[123]

Some trace fossils from the Cambrian-Ordovician boundary about 490 million years ago
are interpreted as the tracks of large amphibious arthropods on coastal sand dunes, and
may have been made by euthycarcinoids,[132] which are thought to be evolutionary "aunts"
of myriapods.[133] Other trace fossils from the Late Ordovician a little over 445 million
years ago probably represent land invertebrates, and there is clear evidence of numerous
arthropods on coasts and alluvial plains shortly before the Silurian-Devonian boundary,
about 415 million years ago, including signs that some arthropods ate plants.[134]
Arthropods were well pre-adapted to colonise land, because their existing jointed
exoskeletons provided protection against desiccation, support against gravity and a means
of locomotion that was not dependent on water.[135]
The fossil record of other major invertebrate groups on land is poor: none at all for nonparasitic flatworms, nematodes or nemerteans; some parasitic nematodes have been
fossilized in amber; annelid worm fossils are known from the Carboniferous, but they
may still have been aquatic animals; the earliest fossils of gastropods on land date from
the Late Carboniferous, and this group may have had to wait until leaf litter became
abundant enough to provide the moist conditions they need.[123]
The earliest confirmed fossils of flying insects date from the Late Carboniferous, but it is
thought that insects developed the ability to fly in the Early Carboniferous or even Late
Devonian. This gave them a wider range of ecological niches for feeding and breeding,
and a means of escape from predators and from unfavorable changes in the environment.
[136]
About 99% of modern insect species fly or are descendants of flying species.[137]

[edit] Land vertebrates

Main article: Tetrapod

Acanthostega changed views about the early evolution of tetrapods[138]

"Fish"
Osteolepiformes ("fish")
Panderichthyidae
Obruchevichthidae
Acanthostega
Ichthyostega
Tulerpeton
Early amphibians

Family tree of tetrapods[139]

Tetrapods, vertebrates with four limbs, evolved from other rhipidistians over a relatively
short timespan during the Late Devonian, between 370 million years ago and 360 million
years ago.[140] From the 1950s to the early 1980s it was thought that tetrapods evolved
from fish that had already acquired the ability to crawl on land, possibly in order to go
from a pool that was drying out to one that was deeper. However in 1987 nearly-complete
fossils of Acanthostega from about 363 million years ago showed that this Late Devonian
transitional animal had legs and both lungs and gills, but could never have survived on
land: its limbs and its wrist and ankle joints were too weak to bear its weight; its ribs
were too short to prevent its lungs from being squeezed flat by its weight; its fish-like tail
fin would have been damaged by dragging on the ground. The current hypothesis is that
Acanthostega, which was about 1 metre (3.3 ft) long, was a wholly aquatic predator that
hunted in shallow water. Its skeleton differed from that of most fish, in ways that enabled
it to raise its head to breathe air while its body remained submerged, including: its jaws
show modifications that would have enabled it to gulp air; the bones at the back of its
skull are locked together, providing strong attachment points for muscles that raised its
head; the head is not joined to the shoulder girdle and it has a distinct neck.[138]
The Devonian proliferation of land plants may help to explain why air-breathing would
have been an advantage: leaves falling into streams and rivers would have encouraged the
growth of aquatic vegetation; this would have attracted grazing invertebrates and small
fish that preyed on them; they would have been attractive prey but the environment was
unsuitable for the big marine predatory fish; air-breathing would have been necessary
because these waters would have been short of oxygen, since warm water holds less
dissolved oxygen than cooler marine water and since the decomposition of vegetation
would have used some of the oxygen.[138]
Later discoveries revealed earlier transitional forms between Acanthostega and
completely fish-like animals.[141] Unfortunately there is then a gap of about 30 million
years between the fossils of ancestral tetrapods and Mid Carboniferous fossils of
vertebrates that look well-adapted for life on land. Some of these look like early relatives
of modern amphibians, most of which need to keep their skins moist and to lay their eggs
in water, while others are accepted as early relatives of the amniotes, whose water-proof
skins and eggs enable them to live and breed far from water.[139]

[edit] Dinosaurs, birds and mammals

Main articles: Dinosaur evolution, Origin of Birds, and Evolution of mammals
Amniote Synapsids
Early synapsids (extinct)
s
Pelycosaurs
Extinct pelycosaurs

Therapsids
Sauropsid
s

Extinct therapsids

Mammaliformes
Extinct
Anapsids; whether turtles belong here is debated[142]
Captorhinidae and Protorothyrididae
Diapsid
s

Araeoscelidia (extinct)
Squamata (lizards and snakes)

Possible family tree of dinosaurs, birdsArchosaur

and mammals[143][144]
Amniotes, whose eggs can survive in dry environments, probably evolved in the Late
Carboniferous period, between 330 million years ago and 314 million years ago. The
earliest fossils of the two surviving amniote groups, synapsids and sauropsids, date from
around 313 million years ago.[143][144] The synapsid pelycosaurs and their descendants the
therapsids are the most common land vertebrates in the best-known Permian fossil beds,
between 229 million years ago and 251 million years ago. However at the time these
were all in temperate zones at middle latitudes, and there is evidence that hotter, drier
environments nearer the Equator were dominated by sauropsids and amphibians.[145]
The Permian-Triassic extinction wiped out almost all land vertebrates,[146] as well as the
great majority of other life.[147] During the slow recovery from this catastrophe, estimated
to be 30M years,[148] a previously obscure sauropsid group became the most abundant and
diverse terrestrial vertebrates: a few fossils of archosauriformes ("shaped like
archosaurs") have been found in Late Permian rocks,[149] but by the Mid Triassic
archosaurs were the dominant land vertebrates. Dinosaurs distinguished themselves from
other archosaurs in the Late Triassic, and became the dominant land vertebrates of the

Jurassic and Cretaceous periods, between 199 million years ago and 65 million years ago.
[150]

During the Late Jurassic, birds evolved from small, predatory theropod dinosaurs.[151] The
first birds inherited teeth and long, bony tails from their dinosaur ancestors,[151] but some
developed horny, toothless beaks by the very Late Jurassic[152] and short pygostyle tails by
the Early Cretaceous.[153]
While the archosaurs and dinosaurs were becoming more dominant in the Triassic, the
mammaliform successors of the therapsids could only survive as small, mainly nocturnal
insectivores. This apparent set-back may actually have promoted the evolution of
mammals, for example nocturnal life may have accelerated the development of
endothermy ("warm-bloodedness") and hair or fur.[154] By 195 million years ago in the
Early Jurassic there were animals that were very nearly mammals.[155] Unfortunately there
is a gap in the fossil record throughout the Mid Jurassic.[156] However fossil teeth
discovered in Madagascar indicate that true mammals existed at least 167 million years
ago.[157] After dominating land vertebrate niches for about 150 million years, the
dinosaurs perished 65 million years ago in the CretaceousTertiary extinction along with
many other groups of organisms.[158] Mammals throughout the time of the dinosaurs had
been restricted to a narrow range of taxa, sizes and shapes, but increased rapidly in size
and diversity after the extinction,[159][160] with bats taking to the air within 13 million
years,[161] and cetaceans to the sea within 15 million years.[162]

[edit] Flowering plants

Main articles: Flowering plant and Gymnosperm
Gymnosperms
Gnetales
Gymnosperms
(gymnosperm)
Welwitschia
(gymnosperm)
Ephedra
(gymnosperm)
Bennettitales
Angiosperms
(flowering plants)
One possible family tree of flowering plants.
[163]

Angiosperms
(flowering plants)
Cycads
(gymnosperm)
Bennettitales
Gingko

Gnetales
(gymnosperm)
Conifers
(gymnosperm)
Another possible family tree.[164]

The 250,000 to 400,000 species of flowering plants outnumber all other ground plants
combined, and are the dominant vegetation in most terrestrial ecosystems. There is fossil
evidence that flowering plants diversified rapidly in the Early Cretaceous, between 130
million years ago and 90 million years ago,[163][164] and that their rise was associated with
that of pollinating insects.[164] Among modern flowering plants Magnolias are thought to

be close to the common ancestor of the group.[163] However paleontologists have not
succeeded in identifying the earliest stages in the evolution of flowering plants.[163][164]

[edit] Social insects

Main article: Social insects
The social insects are remarkable because the great majority of individuals in each colony
are sterile. This appears contrary to basic concepts of evolution such as natural selection
and the selfish gene. In fact there are very few eusocial insect species: only 15 out of
approximately 2,600 living families of insects contain eusocial species, and it seems that
eusociality has evolved independently only 12 times among arthropods, although some
eusocial lineages have diversified into several families. Nevertheless social insects have
been spectacularly successful; for example although ants and termites account for only
about 2% of known insect species, they form over 50% of the total mass of insects. Their
ability to control a territory appears to be the foundation of their success.[165]

These termite mounds have survived a bush fire.

The sacrifice of breeding opportunities by most individuals has long been explained as a
consequence of these species' unusual haplodiploid method of sex determination, which
has the paradoxical consequence that two sterile worker daughters of the same queen
share more genes with each other than they would with their offspring if they could
breed.[166] However Wilson and Hlldobler argue that this explanation is faulty: for
example, it is based on kin selection, but there is no evidence of nepotism in colonies that
have multiple queens. Instead, they write, eusociality evolves only in species that are
under strong pressure from predators and competitors, but in environments where it is
possible to build "fortresses"; after colonies have established this security, they gain other

advantages though co-operative foraging. In support of this explanation they cite the
appearance of eusociality in bathyergid mole rats,[165] which are not haplodiploid.[167]
The earliest fossils of insects have been found in Early Devonian rocks from about 400
million years ago, which preserve only a few varieties of flightless insect. The Mazon
Creek lagersttten from the Late Carboniferous, about 300 million years ago, include
about 200 species, some gigantic by modern standards, and indicate that insects had
occupied their main modern ecological niches as herbivores, detritivores and insectivores.
Social termites and ants first appear in the Early Cretaceous, and advanced social bees
have been found in Late Cretaceous rocks but did not become abundant until the Mid
Cenozoic.[168]

[edit] Humans
Main article: Human evolution
Modern humans evolved from a lineage of upright-walking apes that has been traced
back over 6 million years ago to Sahelanthropus.[169] The first known stone tools were
made about 2.5 million years ago, apparently by Australopithecus garhi, and were found
near animal bones that bear scratches made by these tools.[170] The earliest hominines had
chimp-sized brains, but there has been a fourfold increase in the last 3 million years; a
statistical analysis suggests that hominine brain sizes depend almost completely on the
date of the fossils, while the species to which they are assigned has only slight influence.
[171]
There is a long-running debate about whether modern humans evolved all over the
world simultaneously from existing advanced hominines or are descendants of a single
small population in Africa, which then migrated all over the world less than
200,000 years ago and replaced previous hominine species.[172] There is also debate about
whether anatomically-modern humans had an intellectual, cultural and technological
"Great Leap Forward" under 100,000 years ago and, if so, whether this was due to
neurological changes that are not visible in fossils.[173]

[edit] Mass extinctions

Main article: Mass extinction

KT

TrJ
PTr
Late D
OS
Millions of years ago

Apparent extinction intensity, i.e. the fraction of genera going extinct at any given time, as
reconstructed from the fossil record. (Graph not meant to include recent epoch of Holocene
extinction event)

Life on earth has suffered occasional mass extinctions at least since 542 million years
ago. Although they are disasters at the time, mass extinctions have sometimes accelerated
the evolution of life on earth. When dominance of particular ecological niches passes
from one group of organisms to another, it is rarely because the new dominant group is
"superior" to the old and usually because an extinction event eliminates the old dominant
group and makes way for the new one.[174][175]
The fossil record appears to show that the gaps between mass extinctions are becoming
longer and the average and background rates of extinction are decreasing. Both of these
phenomena could be explained in one or more ways:[176]

The oceans may have become more hospitable to life over the last 500 million
years and less vulnerable to mass extinctions: dissolved oxygen became more
widespread and penetrated to greater depths; the development of life on land
reduced the run-off of nutrients and hence the risk of eutrophication and anoxic
events; and marine ecosystems became more diversified so that food chains were
less likely to be disrupted.[177][178]
Reasonably complete fossils are very rare, most extinct organisms are represented
only by partial fossils, and complete fossils are rarest in the oldest rocks. So
paleontologists have mistakenly assigned parts of the same organism to different
genera which were often defined solely to accommodate these finds the story of
Anomalocaris is an example of this. The risk of this mistake is higher for older
fossils because these are often unlike parts of any living organism. Many of the
"superfluous" genera are represented by fragments which are not found again and
the "superfluous" genera appear to become extinct very quickly.[176]

All genera
"Well-defined" genera
Trend line
"Big Five" mass extinctions
Other mass extinctions
Million years ago
Thousands of genera

Phanerozoic biodiversity as shown by the fossil record

Biodiversity in the fossil record, which is

"the number of distinct genera alive at any given time; that is, those whose first
occurrence predates and whose last occurrence postdates that time"[179]
shows a different trend: a fairly swift rise from 542 to 400 million years ago; a slight
decline from 400 to 200 million years ago, in which the devastating PermianTriassic
extinction event is an important factor; and a swift rise from 200 million years ago to the
present.[179]

[edit] The present

Oxygenic photosynthesis accounts for virtually all of the production of organic matter
from non-organic ingredients. Production is split about evenly between land and marine
plants, and phytoplankton are the dominant marine producers.[180]

The processes that drive evolution are still operating. Well-known examples include the
changes in coloration of the peppered moth over the last 200 years and the more recent
appearance of pathogens that are resistant to antibiotics.[181][182] There is even evidence
that humans are still evolving, and possibly at an accelerating rate over the last 40,000
years.[183]

Evolution
From Wikipedia, the free encyclopedia

Jump to: navigation, search

This article is about evolution in biology. For other uses, see Evolution (disambiguation).
For a generally accessible and less technical introduction to the topic, see Introduction
to evolution.
Part of the Biology series on

Evolution

Mechanisms and processes

Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation
Research and history
Introduction
Evidence
Evolutionary history of life

History
Modern synthesis
Social effect
Theory and fact
Objections / Controversy
Evolutionary biology fields
Cladistics
Ecological genetics
Evolutionary development
Evolutionary psychology
Human evolution
Molecular evolution
Phylogenetics
Population genetics

Biology portal v d e

Evolution is the change in the inherited traits of a population of organisms through

successive generations.[1] After a population splits into smaller groups, these groups
evolve independently and may eventually diversify into new species. A nested hierarchy
of anatomical and genetic similarities, geographical distribution of similar species and the
fossil record indicate that all organisms are descended from a common ancestor through a
long series of these divergent events, stretching back in a tree of life that has grown over
the 3,500 million years of life on Earth.[2]
Evolution is the product of two opposing forces: processes that constantly introduce
variation in traits, and processes that make particular variants become more common or
rare. A trait is a particular characteristic, such as eye color, height, or a behavior, that is
expressed when an organism's genes interact with its environment, translating its
genotypic predispositions into phenotypic phenomena. Genes vary within populations, so
organisms show heritable differences (variation) in their traits.
The main cause of variation is mutation, which changes the sequence of a gene. Altered
genes, or alleles, are then inherited by offspring. There can sometimes also be transfer of
genes between species. Two main processes cause variants to become more common or
rare in a population. One is natural selection, which causes traits that aid survival and
reproduction to become more common, and traits that hinder survival and reproduction to
become more rare.[1][3] Natural selection occurs because only a few individuals in each
generation will survive, since resources are limited and organisms produce many more
offspring than their environment can support. Over many generations, mutations produce
successive, small, random changes in traits, which are then filtered by natural selection
and the beneficial changes retained. This adjusts traits so they become suited to an
organism's environment: these adjustments are called adaptations.[4] Not every trait,
however, is an adaptation. Another cause of evolution is genetic drift, an independent

process that produces entirely random changes in how common traits are in a population.
Genetic drift comes from the role that chance plays in whether a trait will be passed on to
the next generation.
Evolutionary biologists document the fact that evolution occurs, and also develop and test
theories that explain its causes. The study of evolutionary biology began in the midnineteenth century, when research into the fossil record and the diversity of living
organisms convinced most scientists that species changed over time.[5][6] The mechanism
driving these changes remained unclear until the theories of natural selection were
independently proposed by Charles Darwin and Alfred Wallace. In 1859, Darwin's
seminal work On the Origin of Species brought the new theories of evolution by natural
selection to a wide audience,[7] leading to the overwhelming acceptance of evolution
among scientists.[8][9][10][11] In the 1930s, Darwinian natural selection became understood in
combination with Mendelian inheritance, forming the modern evolutionary synthesis,[12]
which connected the units of evolution (genes) and the mechanism of evolution (natural
selection). This powerful explanatory and predictive theory has become the central
organizing principle of modern biology, directing research and providing a unifying
explanation for the history and diversity of life on Earth.[9][10][13] Evolution is therefore
applied and studied in fields as diverse as ecology, anthropology, conservation biology,
paleontology, agriculture, medicine, psychology, philosophy and others.

Contents
[hide]

1 History of evolutionary thought

2 Heredity
3 Variation
o 3.1 Mutation
o 3.2 Sex and recombination
o 3.3 Population genetics
o 3.4 Gene flow
4 Mechanisms
o 4.1 Natural selection
o 4.2 Genetic drift
5 Outcomes
o 5.1 Adaptation
o 5.2 Co-evolution
o 5.3 Co-operation
o 5.4 Speciation
o 5.5 Extinction
6 Evolutionary history of life
o 6.1 Origin of life
o 6.2 Common descent
o 6.3 Evolution of life
7 Social and cultural responses

8 Applications
9 See also
10 References
11 Further reading

12 External links

History of evolutionary thought

For more details on this topic, see History of evolutionary thought.

Around 1854 Charles Darwin began writing out what became On the Origin of Species.
The scientific inquiry into the origin of species can be dated to at least the 6th century
BCE, with the Greek philosopher Anaximander.[14] Others who considered evolutionary
ideas included the Greek philosopher Empedocles, the Roman philosopher-poet
Lucretius, the Afro-Arab biologist Al-Jahiz,[15] the Persian philosopher Ibn Miskawayh,
the Brethren of Purity,[16] and the Chinese philosopher Zhuangzi.[17] As biological
knowledge grew in the 18th century, evolutionary ideas were set out by a few natural
philosophers including Pierre Maupertuis in 1745 and Erasmus Darwin in 1796.[18] The
ideas of the biologist Jean-Baptiste Lamarck about transmutation of species influenced
radicals, but were rejected by mainstream scientists. Charles Darwin formulated his idea
of natural selection in 1838 and was still developing his theory in 1858 when Alfred
Russel Wallace sent him a similar theory, and both were presented to the Linnean Society
of London in separate papers.[19] At the end of 1859, Darwin's publication of On the
Origin of Species explained natural selection in detail and presented evidence leading to
increasingly wide acceptance of the occurrence of evolution.
Debate about the mechanisms of evolution continued, and Darwin could not explain the
source of the heritable variations which would be acted on by natural selection. Like
Lamarck, he thought that parents passed on adaptations acquired during their lifetimes,[20]
a theory which was subsequently dubbed Lamarckism.[21] In the 1880s, August
Weismann's experiments indicated that changes from use and disuse were not heritable,
and Lamarckism gradually fell from favour.[22][23] More significantly, Darwin could not
account for how traits were passed down from generation to generation. In 1865 Gregor
Mendel found that traits were inherited in a predictable manner.[24] When Mendel's work
was rediscovered in 1900s, disagreements over the rate of evolution predicted by early

geneticists and biometricians led to a rift between the Mendelian and Darwinian models
of evolution.
Yet it was the rediscovery of Gregor Mendels pioneering work on the fundamentals of
genetics (of which Darwin and Wallace were unaware) by Hugo de Vries and others in
the early 1900s that provided the impetus for a better understanding of how variation
occurs in plant and animal traits. That variation is the main fuel used by natural selection
to shape the wide variety of adaptive traits observed in organic life. Even though Hugo de
Vries and other early geneticists rejected gradual natural selection, their rediscovery of
and subsequent work on genetics eventually provided a solid basis on which the theory of
evolution stood even more convincingly than when it was originally proposed.[25]
The apparent contradiction between Darwins theory of evolution by natural selection and
Mendels work was reconciled in the 1920s and 1930s by evolutionary biologists such as
J.B.S. Haldane, Sewall Wright, and particularly Ronald Fisher, who set the foundations
for the establishment of the field of population genetics. The end result was a
combination of evolution by natural selection and Mendelian inheritance, the modern
evolutionary synthesis.[26] In the 1940s, the identification of DNA as the genetic material
by Oswald Avery and colleagues and the subsequent publication of the structure of DNA
by James Watson and Francis Crick in 1953, demonstrated the physical basis for
inheritance. Since then, genetics and molecular biology have become core parts of
evolutionary biology and have revolutionized the field of phylogenetics.[12]
In its early history, evolutionary biology primarily drew in scientists from traditional
taxonomically oriented disciplines, whose specialist training in particular organisms
addressed general questions in evolution. As evolutionary biology expanded as an
academic discipline, particularly after the development of the modern evolutionary
synthesis, it began to draw more widely from the biological sciences.[12] Currently the
study of evolutionary biology involves scientists from fields as diverse as biochemistry,
ecology, genetics and physiology, and evolutionary concepts are used in even more
distant disciplines such as psychology, medicine, philosophy and computer science. In
the 21st century, current research in evolutionary biology deals with several areas where
the modern evolutionary synthesis may need modification or extension, such as assessing
the relative importance of various ideas on the unit of selection and evolvability and how
to fully incorporate the findings of evolutionary developmental biology.[27][28]

Heredity
Further information: Introduction to genetics, Genetics, and Heredity

DNA structure. Bases are in the center, surrounded by phosphatesugar chains in a

double helix.
Evolution in organisms occurs through changes in heritable traits particular
characteristics of an organism. In humans, for example, eye color is an inherited
characteristic and an individual might inherit the "brown-eye trait" from one of their
parents.[29] Inherited traits are controlled by genes and the complete set of genes within an
organism's genome is called its genotype.[30]
The complete set of observable traits that make up the structure and behavior of an
organism is called its phenotype. These traits come from the interaction of its genotype
with the environment.[31] As a result, many aspects of an organism's phenotype are not
inherited. For example, suntanned skin comes from the interaction between a person's
genotype and sunlight; thus, suntans are not passed on to people's children. However,
some people tan more easily than others, due to differences in their genotype; a striking
example are people with the inherited trait of albinism, who do not tan at all and are very
sensitive to sunburn.[32]
Heritable traits are passed from one generation to the next via DNA, a molecule that
encodes genetic information.[30] DNA is a long polymer composed of four types of bases.
The sequence of bases along a particular DNA molecule specify the genetic information,
in a manner similar to a sequence of letters spelling out a sentence. Before a cell divides,
the DNA is copied, so that each of the resulting two cells will inherit the DNA sequence.
Portions of a DNA molecule that specify a single functional unit are called genes;
different genes have different sequences of bases. Within cells, the long strands of DNA
form condensed structures called chromosomes. A specific location within a chromosome
is known as a locus. If the DNA sequence at a locus varies between individuals, the
different forms of this sequence are called alleles. DNA sequences can change through

mutations, producing new alleles. If a mutation occurs within a gene, the new allele may
affect the trait that the gene controls, altering the phenotype of the organism.
However, while this simple correspondence between an allele and a trait works in some
cases, most traits are more complex and are controlled by multiple interacting genes.[33][34]
The study of such complex traits is a major area of current genetic research. Another
unsolved question in genetics under active research asks whether or not epigenetics is
important in evolution; that is, whether certain heritable traits, aspects of the
environmental context of genome, are implicated in the evolution of organisms without
there being any necessary change in the gene sequence.[35]

Variation
Further information: Genetic diversity and Population genetics
An individual organism's phenotype results from both its genotype and the influence from
the environment it has lived in. A substantial part of the variation in phenotypes in a
population is caused by the differences between their genotypes.[34] The modern
evolutionary synthesis defines evolution as the change over time in this genetic variation.
The frequency of one particular allele will fluctuate, becoming more or less prevalent
relative to other forms of that gene. Evolutionary forces act by driving these changes in
allele frequency in one direction or another. Variation disappears when a new allele
reaches the point of fixation when it either disappears from the population or replaces
the ancestral allele entirely.[36]
Variation comes from mutations in genetic material, migration between populations (gene
flow), and the reshuffling of genes through sexual reproduction. Variation also comes
from exchanges of genes between different species; for example, through horizontal gene
transfer in bacteria, and hybridization in plants.[37] Despite the constant introduction of
variation through these processes, most of the genome of a species is identical in all
individuals of that species.[38] However, even relatively small changes in genotype can
lead to dramatic changes in phenotype: for example, chimpanzees and humans differ in
only about 5% of their genomes.[39]

Mutation
Further information: Mutation and Molecular evolution

Duplication of part of a chromosome

Random mutations constantly occur in the genomes of organisms; these mutations create
genetic variation. Mutations are changes in the DNA sequence of a cell's genome and are
caused by radiation, viruses, transposons and mutagenic chemicals, as well as errors that
occur during meiosis or DNA replication.[40][41][42] These mutations involve several
different types of change in DNA sequences; these can either have no effect, alter the
product of a gene, or prevent the gene from functioning. Studies in the fly Drosophila
melanogaster suggest that if a mutation changes a protein produced by a gene, this will
probably be harmful, with about 70 percent of these mutations having damaging effects,
and the remainder being either neutral or weakly beneficial.[43] Due to the damaging
effects that mutations can have on cells, organisms have evolved mechanisms such as
DNA repair to remove mutations.[40] Therefore, the optimal mutation rate for a species is a
trade-off between costs of a high mutation rate, such as deleterious mutations, and the
metabolic costs of maintaining systems to reduce the mutation rate, such as DNA repair
enzymes.[44] Viruses that use RNA as their genetic material have rapid mutation rates,[45]
which can be an advantage since these viruses will evolve constantly and rapidly, and
thus evade the defensive responses of e.g. the human immune system.[46]
Mutations can involve large sections of a chromosome becoming duplicated (usually by
genetic recombination), which can introduce extra copies of a gene into a genome.[47]
Extra copies of genes are a major source of the raw material needed for new genes to
evolve.[48] This is important because most new genes evolve within gene families from
pre-existing genes that share common ancestors.[49] For example, the human eye uses four
genes to make structures that sense light: three for color vision and one for night vision;
all four are descended from a single ancestral gene.[50] New genes can be created from an
ancestral gene when a duplicate copy mutates and acquires a new function. This process
is easier once a gene has been duplicated because it increases the redundancy of the
system; one gene in the pair can acquire a new function while the other copy continues to
perform its original function.[51][52] Other types of mutation can even create entirely new
genes from previously noncoding DNA.[53][54] The creation of new genes can also involve

small parts of several genes being duplicated, with these fragments then recombining to
form new combinations with new functions.[55][56] When new genes are assembled from
shuffling pre-existing parts, domains act as modules with simple independent functions,
which can be mixed together creating new combinations with new and complex
functions.[57] For example, polyketide synthases are large enzymes that make antibiotics;
they contain up to one hundred independent domains that each catalyze one step in the
overall process, like a step in an assembly line.[58]
Changes in chromosome number may involve even larger mutations, where segments of
the DNA within chromosomes break and then rearrange. For example, two chromosomes
in the Homo genus fused to produce human chromosome 2; this fusion did not occur in
the lineage of the other apes, and they retain these separate chromosomes.[59] In evolution,
the most important role of such chromosomal rearrangements may be to accelerate the
divergence of a population into new species by making populations less likely to
interbreed, and thereby preserving genetic differences between these populations.[60]
Sequences of DNA that can move about the genome, such as transposons, make up a
major fraction of the genetic material of plants and animals, and may have been
important in the evolution of genomes.[61] For example, more than a million copies of the
Alu sequence are present in the human genome, and these sequences have now been
recruited to perform functions such as regulating gene expression.[62] Another effect of
these mobile DNA sequences is that when they move within a genome, they can mutate
or delete existing genes and thereby produce genetic diversity.[41]

Sex and recombination

Further information: Genetic recombination and Sexual reproduction
In asexual organisms, genes are inherited together, or linked, as they cannot mix with
genes of other organisms during reproduction. In contrast, the offspring of sexual
organisms contain random mixtures of their parents' chromosomes that are produced
through independent assortment. In a related process called homologous recombination,
sexual organisms exchange DNA between two matching chromosomes.[63] Recombination
and reassortment do not alter allele frequencies, but instead change which alleles are
associated with each other, producing offspring with new combinations of alleles.[64] Sex
usually increases genetic variation and may increase the rate of evolution.[65][66] However,
asexuality is advantageous in some environments as it can evolve in previously-sexual
animals.[67] Here, asexuality might allow the two sets of alleles in their genome to diverge
and gain different functions.[68]
Recombination allows even alleles that are close together in a strand of DNA to be
inherited independently. However, the rate of recombination is low (approximately two
events per chromosome per generation). As a result, genes close together on a
chromosome may not always be shuffled away from each other, and genes that are close
together tend to be inherited together, a phenomenon known as linkage.[69] This tendency
is measured by finding how often two alleles occur together on a single chromosome,

which is called their linkage disequilibrium. A set of alleles that is usually inherited in a
group is called a haplotype. This can be important when one allele in a particular
haplotype is strongly beneficial: natural selection can drive a selective sweep that will
also cause the other alleles in the haplotype to become more common in the population;
this effect is called genetic hitchhiking.[70]
When alleles cannot be separated by recombination such as in mammalian Y
chromosomes, which pass intact from fathers to sons harmful mutations accumulate.[71]
[72]
By breaking up allele combinations, sexual reproduction allows the removal of
harmful mutations and the retention of beneficial mutations.[73] In addition, recombination
and reassortment can produce individuals with new and advantageous gene combinations.
These positive effects are balanced by the fact that sex reduces an organism's
reproductive rate, can cause mutations and may separate beneficial combinations of
genes.[73] The reasons for the evolution of sexual reproduction are therefore unclear and
this question is still an active area of research in evolutionary biology,[74][75] that has
prompted ideas such as the Red Queen hypothesis.[76]

Population genetics

White peppered moth

Black morph in peppered moth evolution

Further information: Population genetics
From a genetic viewpoint, evolution is a generation-to-generation change in the
frequencies of alleles within a population that shares a common gene pool.[77] A
population is a localized group of individuals belonging to the same species. For
example, all of the moths of the same species living in an isolated forest represent a
population. A single gene in this population may have several alternate forms, which
account for variations between the phenotypes of the organisms. An example might be a
gene for coloration in moths that has two alleles: black and white. A gene pool is the
complete set of alleles for a gene in a single population; the allele frequency measures the
fraction of the gene pool composed of a single allele (for example, what fraction of moth

coloration genes are the black allele). Evolution occurs when there are changes in the
frequencies of alleles within a population of interbreeding organisms; for example, the
allele for black color in a population of moths becoming more common.
To understand the mechanisms that cause a population to evolve, it is useful to consider
what conditions are required for a population not to evolve. The Hardy-Weinberg
principle states that the frequencies of alleles (variations in a gene) in a sufficiently large
population will remain constant if the only forces acting on that population are the
random reshuffling of alleles during the formation of the sperm or egg, and the random
combination of the alleles in these sex cells during fertilization.[78] Such a population is
said to be in Hardy-Weinberg equilibrium; it is not evolving.[79]

Gene flow
Further information: Gene flow, Hybrid (biology), and Horizontal gene transfer

When they mature, male lions leave the pride where they were born and take over a new
pride to mate, causing gene flow between prides.[80]
Gene flow is the exchange of genes between populations, which are usually of the same
species.[81] Examples of gene flow within a species include the migration and then
breeding of organisms, or the exchange of pollen. Gene transfer between species includes
the formation of hybrid organisms and horizontal gene transfer.
Migration into or out of a population can change allele frequencies, as well as introducing
genetic variation into a population. Immigration may add new genetic material to the
established gene pool of a population. Conversely, emigration may remove genetic
material. As barriers to reproduction between two diverging populations are required for
the populations to become new species, gene flow may slow this process by spreading
genetic differences between the populations. Gene flow is hindered by mountain ranges,
oceans and deserts or even man-made structures such as the Great Wall of China, which
has hindered the flow of plant genes.[82]
Depending on how far two species have diverged since their most recent common
ancestor, it may still be possible for them to produce offspring, as with horses and
donkeys mating to produce mules.[83] Such hybrids are generally infertile, due to the two

different sets of chromosomes being unable to pair up during meiosis. In this case,
closely related species may regularly interbreed, but hybrids will be selected against and
the species will remain distinct. However, viable hybrids are occasionally formed and
these new species can either have properties intermediate between their parent species, or
possess a totally new phenotype.[84] The importance of hybridization in creating new
species of animals is unclear, although cases have been seen in many types of animals,[85]
with the gray tree frog being a particularly well-studied example.[86]
Hybridization is, however, an important means of speciation in plants, since polyploidy
(having more than two copies of each chromosome) is tolerated in plants more readily
than in animals.[87][88] Polyploidy is important in hybrids as it allows reproduction, with
the two different sets of chromosomes each being able to pair with an identical partner
during meiosis.[89] Polyploids also have more genetic diversity, which allows them to
avoid inbreeding depression in small populations.[90]
Horizontal gene transfer is the transfer of genetic material from one organism to another
organism that is not its offspring; this is most common among bacteria.[91] In medicine,
this contributes to the spread of antibiotic resistance, as when one bacteria acquires
resistance genes it can rapidly transfer them to other species.[92] Horizontal transfer of
genes from bacteria to eukaryotes such as the yeast Saccharomyces cerevisiae and the
adzuki bean beetle Callosobruchus chinensis may also have occurred.[93][94] An example
of larger-scale transfers are the eukaryotic bdelloid rotifers, which appear to have
received a range of genes from bacteria, fungi, and plants.[95] Viruses can also carry DNA
between organisms, allowing transfer of genes even across biological domains.[96] Largescale gene transfer has also occurred between the ancestors of eukaryotic cells and
prokaryotes, during the acquisition of chloroplasts and mitochondria.[97]

Mechanisms
The two main mechanisms that produce evolution are natural selection and genetic drift.
Natural selection is the process which favors genes that aid survival and reproduction.
Genetic drift is the random change in the frequency of alleles, caused by the random
sampling of a generation's genes during reproduction. The relative importance of natural
selection and genetic drift in a population varies depending on the strength of the
selection and the effective population size, which is the number of individuals capable of
breeding.[98] Natural selection usually predominates in large populations, whereas genetic
drift dominates in small populations. The dominance of genetic drift in small populations
can even lead to the fixation of slightly deleterious mutations.[99] As a result, changing
population size can dramatically influence the course of evolution. Population
bottlenecks, where the population shrinks temporarily and therefore loses genetic
variation, result in a more uniform population.[36]

Natural selection
Further information: Natural selection and Fitness (biology)

Natural selection of a population for dark coloration.

Natural selection is the process by which genetic mutations that enhance reproduction
become, and remain, more common in successive generations of a population. It has
often been called a "self-evident" mechanism because it necessarily follows from three
simple facts:

Heritable variation exists within populations of organisms.

Organisms produce more offspring than can survive.
These offspring vary in their ability to survive and reproduce.

These conditions produce competition between organisms for survival and reproduction.
Consequently, organisms with traits that give them an advantage over their competitors
pass these advantageous traits on, while traits that do not confer an advantage are not
passed on to the next generation.[100]
The central concept of natural selection is the evolutionary fitness of an organism.[101]
Fitness is measured by an organism's ability to survive and reproduce, which determines
the size of its genetic contribution to the next generation.[101] However, fitness is not the
same as the total number of offspring: instead fitness is indicated by the proportion of
subsequent generations that carry an organism's genes.[102] For example, if an organism
could survive well and reproduce rapidly, but its offspring were all too small and weak to
survive, this organism would make little genetic contribution to future generations and
would thus have low fitness.[101]
If an allele increases fitness more than the other alleles of that gene, then with each
generation this allele will become more common within the population. These traits are
said to be "selected for". Examples of traits that can increase fitness are enhanced
survival, and increased fecundity. Conversely, the lower fitness caused by having a less
beneficial or deleterious allele results in this allele becoming rarer they are "selected
against".[3] Importantly, the fitness of an allele is not a fixed characteristic; if the

environment changes, previously neutral or harmful traits may become beneficial and
previously beneficial traits become harmful.[1] However, even if the direction of selection
does reverse in this way, traits that were lost in the past may not re-evolve in an identical
form (see Dollo's law).[103][104]
Natural selection within a population for a trait that can vary across a range of values,
such as height, can be categorized into three different types. The first is directional
selection, which is a shift in the average value of a trait over time for example
organisms slowly getting taller.[105] Secondly, disruptive selection is selection for extreme
trait values and often results in two different values becoming most common, with
selection against the average value. This would be when either short or tall organisms had
an advantage, but not those of medium height. Finally, in stabilizing selection there is
selection against extreme trait values on both ends, which causes a decrease in variance
around the average value and less diversity.[100][106] This would, for example, cause
organisms to slowly become all the same height.
A special case of natural selection is sexual selection, which is selection for any trait that
increases mating success by increasing the attractiveness of an organism to potential
mates.[107] Traits that evolved through sexual selection are particularly prominent in males
of some animal species, despite traits such as cumbersome antlers, mating calls or bright
colors that attract predators, decreasing the survival of individual males.[108] This survival
disadvantage is balanced by higher reproductive success in males that show these hard to
fake, sexually selected traits.[109]
Natural selection most generally makes nature the measure against which individuals, and
individual traits, are more or less likely to survive. "Nature" in this sense refers to an
ecosystem, that is, a system in which organisms interact with every other element,
physical as well as biological, in their local environment. Eugene Odum, a founder of
ecology, defined an ecosystem as: "Any unit that includes all of the organisms...in a given
area interacting with the physical environment so that a flow of energy leads to clearly
defined trophic structure, biotic diversity, and material cycles (ie: exchange of materials
between living and nonliving parts) within the system."[110] Each population within an
ecosystem occupies a distinct niche, or position, with distinct relationships to other parts
of the system. These relationships involve the life history of the organism, its position in
the food chain, and its geographic range. This broad understanding of nature enables
scientists to delineate specific forces which, together, comprise natural selection.
An active area of research is the unit of selection, with natural selection being proposed
to work at the level of genes, cells, individual organisms, groups of organisms and
species.[111][112] None of these are mutually exclusive and selection can act on multiple
levels simultaneously.[113] An example of selection occurring below the level of the
individual organism are genes called transposons, which can replicate and spread
throughout a genome.[114] Selection at a level above the individual, such as group
selection, may allow the evolution of co-operation, as discussed below.[115]

Genetic drift

Further information: Genetic drift and Effective population size

Simulation of genetic drift of 20 unlinked alleles in populations of 10 (top) and 100

(bottom). Drift to fixation is more rapid in the smaller population.
Genetic drift is the change in allele frequency from one generation to the next that occurs
because alleles in offspring are a random sample of those in the parents, as well as from
the role that chance plays in determining whether a given individual will survive and
reproduce. In mathematical terms, alleles are subject to sampling error. As a result, when
selective forces are absent or relatively weak, allele frequencies tend to "drift" upward or
downward randomly (in a random walk). This drift halts when an allele eventually
becomes fixed, either by disappearing from the population, or replacing the other alleles
entirely. Genetic drift may therefore eliminate some alleles from a population due to
chance alone. Even in the absence of selective forces, genetic drift can cause two separate
populations that began with the same genetic structure to drift apart into two divergent
populations with different sets of alleles.[116]
The time for an allele to become fixed by genetic drift depends on population size, with
fixation occurring more rapidly in smaller populations.[117] The precise measure of
population that is important is called the effective population size. The effective
population is always smaller than the total population since it takes into account factors
such as the level of inbreeding, the number of animals that are too old or young to breed,
and the lower probability of animals that live far apart managing to mate with each other.
[118]

An example when genetic drift is probably of central importance in determining a trait is

the loss of pigments from animals that live in caves, a change that produces no obvious
advantage or disadvantage in complete darkness.[119] However, it is usually difficult to
measure the relative importance of selection and drift,[120] so the comparative importance
of these two forces in driving evolutionary change is an area of current research.[121]
These investigations were prompted by the neutral theory of molecular evolution, which
proposed that most evolutionary changes are the result of the fixation of neutral

mutations that do not have any immediate effects on the fitness of an organism.[122]
Hence, in this model, most genetic changes in a population are the result of constant
mutation pressure and genetic drift.[123] This form of the neutral theory is now largely
abandoned, since it does not seem to fit the genetic variation seen in nature.[124][125]
However, a more recent and better-supported version of this model is the nearly neutral
theory, where most mutations only have small effects on fitness.[100]

Outcomes
Evolution influences every aspect of the form and behavior of organisms. Most
prominent are the specific behavioral and physical adaptations that are the outcome of
natural selection. These adaptations increase fitness by aiding activities such as finding
food, avoiding predators or attracting mates. Organisms can also respond to selection by
co-operating with each other, usually by aiding their relatives or engaging in mutually
beneficial symbiosis. In the longer term, evolution produces new species through splitting
ancestral populations of organisms into new groups that cannot or will not interbreed.
These outcomes of evolution are sometimes divided into macroevolution, which is
evolution that occurs at or above the level of species, such as extinction and speciation,
and microevolution, which is smaller evolutionary changes, such as adaptations, within a
species or population.[126] In general, macroevolution is regarded as the outcome of long
periods of microevolution.[127] Thus, the distinction between micro- and macroevolution is
not a fundamental one the difference is simply the time involved.[128] However, in
macroevolution, the traits of the entire species may be important. For instance, a large
amount of variation among individuals allows a species to rapidly adapt to new habitats,
lessening the chance of it going extinct, while a wide geographic range increases the
chance of speciation, by making it more likely that part of the population will become
isolated. In this sense, microevolution and macroevolution might involve selection at
different levels with microevolution acting on genes and organisms, versus
macroevolutionary processes such as species selection acting on entire species and
affecting their rates of speciation and extinction.[129][130][131]
A common misconception is that evolution has goals or long-term plans; realistically
however, evolution has no long-term goal and does not necessarily produce greater
complexity.[132][133] Although complex species have evolved, they occur as a side effect of
the overall number of organisms increasing, and simple forms of life still remain more
common in the biosphere.[134] For example, the overwhelming majority of species are
microscopic prokaryotes, which form about half the world's biomass despite their small
size,[135] and constitute the vast majority of Earth's biodiversity.[136] Simple organisms have
therefore been the dominant form of life on Earth throughout its history and continue to
be the main form of life up to the present day, with complex life only appearing more
diverse because it is more noticeable.[137] Indeed, the evolution of microorganisms is
particularly important to modern evolutionary research, since their rapid reproduction
allows the study of experimental evolution and the observation of evolution and
adaptation in real time.[138][139]

Adaptation
For more details on this topic, see Adaptation.
Adaptation is one of the basic phenomena of biology,[140] and is the process whereby an
organism becomes better suited to its habitat.[141][142] Also, the term adaptation may refer
to a trait that is important for an organism's survival. For example, the adaptation of
horses' teeth to the grinding of grass, or the ability of horses to run fast and escape
predators. By using the term adaptation for the evolutionary process, and adaptive trait
for the product (the bodily part or function), the two senses of the word may be
distinguished. Adaptations are produced by natural selection.[143] The following
definitions are due to Theodosius Dobzhansky.
1. Adaptation is the evolutionary process whereby an organism becomes better
able to live in its habitat or habitats.[144]
2. Adaptedness is the state of being adapted: the degree to which an organism is
able to live and reproduce in a given set of habitats.[145]
3. An adaptive trait is an aspect of the developmental pattern of the organism
which enables or enhances the probability of that organism surviving and
reproducing.[146]
Adaptation may cause either the gain of a new feature, or the loss of an ancestral feature.
An example that shows both types of change is bacterial adaptation to antibiotic
selection, with genetic changes causing antibiotic resistance by both modifying the target
of the drug, or increasing the activity of transporters that pump the drug out of the cell.[147]
Other striking examples are the bacteria Escherichia coli evolving the ability to use citric
acid as a nutrient in a long-term laboratory experiment,[148] Flavobacterium evolving a
novel enzyme that allows these bacteria to grow on the by-products of nylon
manufacturing,[149][150] and the soil bacterium Sphingobium evolving an entirely new
metabolic pathway that degrades the synthetic pesticide pentachlorophenol.[151][152] An
interesting but still controversial idea is that some adaptations might increase the ability
of organisms to generate genetic diversity and adapt by natural selection (increasing
organisms' evolvability).[153][154]

A baleen whale skeleton, a and b label flipper bones, which were adapted from front leg
bones: while c indicates vestigial leg bones, suggesting an adaptation from land to sea.[155]
Adaptation occurs through the gradual modification of existing structures. Consequently,
structures with similar internal organization may have different functions in related

organisms. This is the result of a single ancestral structure being adapted to function in
different ways. The bones within bat wings, for example, are very similar to those in mice
feet and primate hands, due to the descent of all these structures from a common
mammalian ancestor.[156] However, since all living organisms are related to some extent,
[157]
even organs that appear to have little or no structural similarity, such as arthropod,
squid and vertebrate eyes, or the limbs and wings of arthropods and vertebrates, can
depend on a common set of homologous genes that control their assembly and function;
this is called deep homology.[158][159]
During adaptation, some structures may lose their original function and become vestigial
structures.[160] Such structures may have little or no function in a current species, yet have
a clear function in ancestral species, or other closely related species. Examples include
pseudogenes,[161] the non-functional remains of eyes in blind cave-dwelling fish,[162] wings
in flightless birds,[163] and the presence of hip bones in whales and snakes.[155] Examples
of vestigial structures in humans include wisdom teeth,[164] the coccyx,[160] and the
vermiform appendix.[160]
However, many traits that appear to be simple adaptations are in fact exaptations:
structures originally adapted for one function, but which coincidentally became
somewhat useful for some other function in the process.[165] One example is the African
lizard Holaspis guentheri, which developed an extremely flat head for hiding in crevices,
as can be seen by looking at its near relatives. However, in this species, the head has
become so flattened that it assists in gliding from tree to treean exaptation.[165] Within
cells, molecular machines such as the bacterial flagella[166] and protein sorting
machinery[167] evolved by the recruitment of several pre-existing proteins that previously
had different functions.[126] Another example is the recruitment of enzymes from
glycolysis and xenobiotic metabolism to serve as structural proteins called crystallins
within the lenses of organisms' eyes.[168][169]
A critical principle of ecology is that of competitive exclusion: no two species can occupy
the same niche in the same environment for a long time.[170] Consequently, natural
selection will tend to force species to adapt to different ecological niches. This may mean
that, for example, two species of cichlid fish adapt to live in different habitats, which will
minimize the competition between them for food.[171]
An area of current investigation in evolutionary developmental biology is the
developmental basis of adaptations and exaptations.[172] This research addresses the origin
and evolution of embryonic development and how modifications of development and
developmental processes produce novel features.[173] These studies have shown that
evolution can alter development to create new structures, such as embryonic bone
structures that develop into the jaw in other animals instead forming part of the middle
ear in mammals.[174] It is also possible for structures that have been lost in evolution to
reappear due to changes in developmental genes, such as a mutation in chickens causing
embryos to grow teeth similar to those of crocodiles.[175] It is now becoming clear that
most alterations in the form of organisms are due to changes in a small set of conserved
genes.[176]

Co-evolution
Further information: Co-evolution
Interactions between organisms can produce both conflict and co-operation. When the
interaction is between pairs of species, such as a pathogen and a host, or a predator and
its prey, these species can develop matched sets of adaptations. Here, the evolution of one
species causes adaptations in a second species. These changes in the second species then,
in turn, cause new adaptations in the first species. This cycle of selection and response is
called co-evolution.[177] An example is the production of tetrodotoxin in the rough-skinned
newt and the evolution of tetrodotoxin resistance in its predator, the common garter
snake. In this predator-prey pair, an evolutionary arms race has produced high levels of
toxin in the newt and correspondingly high levels of toxin resistance in the snake.[178]

Co-operation
Further information: Co-operation (evolution)
However, not all interactions between species involve conflict.[179] Many cases of
mutually beneficial interactions have evolved. For instance, an extreme cooperation
exists between plants and the mycorrhizal fungi that grow on their roots and aid the plant
in absorbing nutrients from the soil.[180] This is a reciprocal relationship as the plants
provide the fungi with sugars from photosynthesis. Here, the fungi actually grow inside
plant cells, allowing them to exchange nutrients with their hosts, while sending signals
that suppress the plant immune system.[181]
Coalitions between organisms of the same species have also evolved. An extreme case is
the eusociality found in social insects, such as bees, termites and ants, where sterile
insects feed and guard the small number of organisms in a colony that are able to
reproduce. On an even smaller scale, the somatic cells that make up the body of an
animal limit their reproduction so they can maintain a stable organism, which then
supports a small number of the animal's germ cells to produce offspring. Here, somatic
cells respond to specific signals that instruct them whether to grow, remain as they are, or
die. If cells ignore these signals and multiply inappropriately, their uncontrolled growth
causes cancer.[40]
Such cooperation within species may have evolved through the process of kin selection,
which is where one organism acts to help raise a relative's offspring.[182] This activity is
selected for because if the helping individual contains alleles which promote the helping
activity, it is likely that its kin will also contain these alleles and thus those alleles will be
passed on.[183] Other processes that may promote cooperation include group selection,
where cooperation provides benefits to a group of organisms.[184]

Speciation
Further information: Speciation

The four mechanisms of speciation.

Speciation is the process where a species diverges into two or more descendant species.
[185]
Evolutionary biologists view species as statistical phenomena and not categories or
types. This view is counterintuitive since the classical idea of species is still widely held,
with a species seen as a class of organisms exemplified by a "type specimen" that bears
all the traits common to this species. Instead, a species is now defined as a separately
evolving lineage that forms a single gene pool. Although properties such as genetics and
morphology are used to help separate closely related lineages, this definition has fuzzy
boundaries.[186] Indeed, the exact definition of the term "species" is still controversial,
particularly in prokaryotes,[187] and this is called the species problem.[188] Biologists have
proposed a range of more precise definitions, but the definition used is a pragmatic
choice that depends on the particularities of the species concerned.[188] Typically the
actual focus on biological study is the population, an observable interacting group of
organisms, rather than a species, an observable similar group of individuals.
Speciation has been observed multiple times under both controlled laboratory conditions
and in nature.[189] In sexually reproducing organisms, speciation results from reproductive
isolation followed by genealogical divergence. There are four mechanisms for speciation.
The most common in animals is allopatric speciation, which occurs in populations
initially isolated geographically, such as by habitat fragmentation or migration. Selection
under these conditions can produce very rapid changes in the appearance and behaviour
of organisms.[190][191] As selection and drift act independently on populations isolated from
the rest of their species, separation may eventually produce organisms that cannot
interbreed.[192]
The second mechanism of speciation is peripatric speciation, which occurs when small
populations of organisms become isolated in a new environment. This differs from

allopatric speciation in that the isolated populations are numerically much smaller than
the parental population. Here, the founder effect causes rapid speciation through both
rapid genetic drift and selection on a small gene pool.[193]
The third mechanism of speciation is parapatric speciation. This is similar to peripatric
speciation in that a small population enters a new habitat, but differs in that there is no
physical separation between these two populations. Instead, speciation results from the
evolution of mechanisms that reduce gene flow between the two populations.[185]
Generally this occurs when there has been a drastic change in the environment within the
parental species' habitat. One example is the grass Anthoxanthum odoratum, which can
undergo parapatric speciation in response to localized metal pollution from mines.[194]
Here, plants evolve that have resistance to high levels of metals in the soil. Selection
against interbreeding with the metal-sensitive parental population produced a gradual
change in the flowering time of the metal-resistant plants, which eventually produced
complete reproductive isolation. Selection against hybrids between the two populations
may cause reinforcement, which is the evolution of traits that promote mating within a
species, as well as character displacement, which is when two species become more
distinct in appearance.[195]

Geographical isolation of finches on the Galpagos Islands produced over a dozen new
species.
Finally, in sympatric speciation species diverge without geographic isolation or changes
in habitat. This form is rare since even a small amount of gene flow may remove genetic
differences between parts of a population.[196] Generally, sympatric speciation in animals
requires the evolution of both genetic differences and non-random mating, to allow
reproductive isolation to evolve.[197]
One type of sympatric speciation involves cross-breeding of two related species to
produce a new hybrid species. This is not common in animals as animal hybrids are
usually sterile. This is because during meiosis the homologous chromosomes from each
parent are from different species and cannot successfully pair. However, it is more
common in plants because plants often double their number of chromosomes, to form
polyploids.[198] This allows the chromosomes from each parental species to form a
matching pair during meiosis, since as each parent's chromosomes is represented by a

pair already.[199] An example of such a speciation event is when the plant species
Arabidopsis thaliana and Arabidopsis arenosa cross-bred to give the new species
Arabidopsis suecica.[200] This happened about 20,000 years ago,[201] and the speciation
process has been repeated in the laboratory, which allows the study of the genetic
mechanisms involved in this process.[202] Indeed, chromosome doubling within a species
may be a common cause of reproductive isolation, as half the doubled chromosomes will
be unmatched when breeding with undoubled organisms.[88]
Speciation events are important in the theory of punctuated equilibrium, which accounts
for the pattern in the fossil record of short "bursts" of evolution interspersed with
relatively long periods of stasis, where species remain relatively unchanged.[203] In this
theory, speciation and rapid evolution are linked, with natural selection and genetic drift
acting most strongly on organisms undergoing speciation in novel habitats or small
populations. As a result, the periods of stasis in the fossil record correspond to the
parental population, and the organisms undergoing speciation and rapid evolution are
found in small populations or geographically restricted habitats, and therefore rarely
being preserved as fossils.[204]

Extinction
Further information: Extinction

Tyrannosaurus rex. Non-avian dinosaurs died out in the CretaceousTertiary extinction

event at the end of the Cretaceous period.
Extinction is the disappearance of an entire species. Extinction is not an unusual event, as
species regularly appear through speciation, and disappear through extinction.[205] Nearly
all animal and plant species that have lived on earth are now extinct,[206] and extinction
appears to be the ultimate fate of all species.[207] These extinctions have happened
continuously throughout the history of life, although the rate of extinction spikes in
occasional mass extinction events.[208] The CretaceousTertiary extinction event, during
which the non-avian dinosaurs went extinct, is the most well-known, but the earlier
PermianTriassic extinction event was even more severe, with approximately 96 percent
of species driven to extinction.[208] The Holocene extinction event is an ongoing mass
extinction associated with humanity's expansion across the globe over the past few
thousand years. Present-day extinction rates are 1001000 times greater than the
background rate, and up to 30 percent of species may be extinct by the mid 21st century.

[209]

Human activities are now the primary cause of the ongoing extinction event;[210]
global warming may further accelerate it in the future.[211]
The role of extinction in evolution is not very well understood and may depend on which
type of extinction is considered.[208] The causes of the continuous "low-level" extinction
events, which form the majority of extinctions, may be the result of competition between
species for limited resources (competitive exclusion).[12] If one species can out-compete
another, this could produce species selection, with the fitter species surviving and the
other species being driven to extinction.[111] The intermittent mass extinctions are also
important, but instead of acting as a selective force, they drastically reduce diversity in a
nonspecific manner and promote bursts of rapid evolution and speciation in survivors.[212]

Evolutionary history of life

Main article: Evolutionary history of life
See also: Timeline of evolution and Timeline of human evolution

Origin of life
Further information: Abiogenesis and RNA world hypothesis
The origin of life is a necessary precursor for biological evolution, but understanding that
evolution occurred once organisms appeared and investigating how this happens does not
depend on understanding exactly how life began.[213] The current scientific consensus is
that the complex biochemistry that makes up life came from simpler chemical reactions,
but it is unclear how this occurred.[214] Not much is certain about the earliest
developments in life, the structure of the first living things, or the identity and nature of
any last universal common ancestor or ancestral gene pool.[215][216] Consequently, there is
no scientific consensus on how life began, but proposals include self-replicating
molecules such as RNA,[217] and the assembly of simple cells.[218]

Common descent
Further information: Evidence of common descent, Common descent, and Homology
(biology)

The hominoids are descendants of a common ancestor.

All organisms on Earth are descended from a common ancestor or ancestral gene pool.[157]
Current species are a stage in the process of evolution, with their diversity the product of
a long series of speciation and extinction events.[219] The common descent of organisms
was first deduced from four simple facts about organisms: First, they have geographic
distributions that cannot be explained by local adaptation. Second, the diversity of life is
not a set of completely unique organisms, but organisms that share morphological
similarities. Third, vestigial traits with no clear purpose resemble functional ancestral
traits, and finally, that organisms can be classified using these similarities into a hierarchy
of nested groups similar to a family tree.[7] However, modern research has suggested
that, due to horizontal gene transfer, this "tree of life" may be more complicated than a
simple branching tree since some genes have spread independently between distantly
related species.[220][221]
Past species have also left records of their evolutionary history. Fossils, along with the
comparative anatomy of present-day organisms, constitute the morphological, or
anatomical, record.[222] By comparing the anatomies of both modern and extinct species,
paleontologists can infer the lineages of those species. However, this approach is most
successful for organisms that had hard body parts, such as shells, bones or teeth. Further,
as prokaryotes such as bacteria and archaea share a limited set of common morphologies,
their fossils do not provide information on their ancestry.
More recently, evidence for common descent has come from the study of biochemical
similarities between organisms. For example, all living cells use the same basic set of
nucleotides and amino acids.[223] The development of molecular genetics has revealed the
record of evolution left in organisms' genomes: dating when species diverged through the
molecular clock produced by mutations.[224] For example, these DNA sequence
comparisons have revealed that humans and chimpanzees share 96% of their genomes
and analyzing the few areas where they differ helps shed light on when the common
ancestor of these species existed.[225]

Evolution of life
For more details on this topic, see Timeline of evolution.

Evolutionary tree showing the divergence of modern species from their common ancestor
in the center.[226] The three domains are colored, with bacteria blue, archaea green, and
eukaryotes red.
Despite the uncertainty on how life began, it is generally accepted that prokaryotes
inhabited the Earth from approximately 34 billion years ago.[2][227] No obvious changes
in morphology or cellular organization occurred in these organisms over the next few
billion years.[228]
The eukaryotes were the next major change in cell structure. These came from ancient
bacteria being engulfed by the ancestors of eukaryotic cells, in a cooperative association
called endosymbiosis.[97][229] The engulfed bacteria and the host cell then underwent coevolution, with the bacteria evolving into either mitochondria or hydrogenosomes.[230] An
independent second engulfment of cyanobacterial-like organisms led to the formation of
chloroplasts in algae and plants.[231] It is unknown when the first eukaryotic cells appeared
though they first emerged between 1.6 2.7 billion years ago.
The history of life was that of the unicellular eukaryotes, prokaryotes, and archaea until
about 610 million years ago when multicellular organisms began to appear in the oceans
in the Ediacaran period.[2][232] The evolution of multicellularity occurred in multiple
independent events, in organisms as diverse as sponges, brown algae, cyanobacteria,
slime moulds and myxobacteria.[233]
Soon after the emergence of these first multicellular organisms, a remarkable amount of
biological diversity appeared over approximately 10 million years, in an event called the
Cambrian explosion. Here, the majority of types of modern animals appeared in the fossil
record, as well as unique lineages that subsequently became extinct.[234] Various triggers
for the Cambrian explosion have been proposed, including the accumulation of oxygen in
the atmosphere from photosynthesis.[235] About 500 million years ago, plants and fungi
colonized the land, and were soon followed by arthropods and other animals.[236] Insects
were particularly successful and even today make up the majority of animal species.[237]

Amphibians first appeared around 300 million years ago, followed by early amniotes,
then mammals around 200 million years ago and birds around 100 million years ago
(both from "reptile"-like lineages). However, despite the evolution of these large animals,
smaller organisms similar to the types that evolved early in this process continue to be
highly successful and dominate the Earth, with the majority of both biomass and species
being prokaryotes.[136]

Social and cultural responses

Further information: Social effect of evolutionary theory and Objections to evolution

As Darwinism became widely accepted in the 1870s, caricatures of Charles Darwin with
an ape or monkey body symbolised evolution.[238]
In the 19th century, particularly after the publication of On the Origin of Species in 1859,
the idea that life had evolved was an active source of academic debate centered on the
philosophical, social and religious implications of evolution. Nowadays, the fact that
organisms evolve is uncontested in the scientific literature and the modern evolutionary
synthesis is widely accepted by scientists.[12] However, evolution remains a contentious
concept for some theists.[239]
While various religions and denominations have reconciled their beliefs with evolution
through concepts such as theistic evolution, there are creationists who believe that
evolution is contradicted by the creation myths found in their respective religions and
who raise various objections to evolution.[126][240][241] As had been demonstrated by
responses to the publication of Vestiges of the Natural History of Creation in 1844, the
most controversial aspect of evolutionary biology is the implication of human evolution
that human mental and moral faculties, which had been thought purely spiritual, are not
distinctly separated from those of other animals.[6] In some countriesnotably the United
Statesthese tensions between science and religion have fueled the current creationevolution controversy, a religious conflict focusing on politics and public education.[242]
While other scientific fields such as cosmology[243] and earth science[244] also conflict with
literal interpretations of many religious texts, evolutionary biology experiences
significantly more opposition from religious literalists.

The teaching of evolution in American secondary school biology classes was uncommon
in most of the first half of the 20th century. The Scopes Trial decision of 1925 caused the
subject to become very rare in American secondary biology textbooks for a generation,
but it was gradually re-introduced about a generation later and legally protected with the
1968 Epperson v. Arkansas decision. Since then, the competing religious belief of
creationism was legally disallowed in secondary school curricula in various decisions in
the 1970s and 1980s, but it returned in the form of intelligent design, to be excluded once
again in the 2005 Kitzmiller v. Dover Area School District case.[245]
Another example somewhat associated with evolutionary theory that is now widely
regarded as unwarranted is "Social Darwinism", a derogatory term associated with the
19th century Malthusian theory developed by Whig philosopher Herbert Spencer. It was
later expanded by others into ideas about "survival of the fittest" in commerce and human
societies as a whole, and led to claims that social inequality, sexism, racism, and
imperialism were justified.[246] However, these ideas contradict Darwin's own views, and
contemporary scientists and philosophers consider these ideas to be neither mandated by
evolutionary theory nor supported by data.[247][248][249]

Applications
Further information: Artificial selection and Evolutionary computation
Evolutionary biology, and in particular the understanding of how organisms evolve
through natural selection, is an area of science with many practical applications.[250] A
major technological application of evolution is artificial selection, which is the intentional
selection of certain traits in a population of organisms. Humans have used artificial
selection for thousands of years in the domestication of plants and animals.[251] More
recently, such selection has become a vital part of genetic engineering, with selectable
markers such as antibiotic resistance genes being used to manipulate DNA in molecular
biology. It is also possible to use repeated rounds of mutation and selection to evolve
proteins with particular properties, such as modified enzymes or new antibodies, in a
process called directed evolution.[252]
Understanding the changes that have occurred during organism's evolution can reveal the
genes needed to construct parts of the body, genes which may be involved in human
genetic disorders.[253] For example, the Mexican tetra is an albino cavefish that lost its
eyesight during evolution. Breeding together different populations of this blind fish
produced some offspring with functional eyes, since different mutations had occurred in
the isolated populations that had evolved in different caves.[254] This helped identify genes
required for vision and pigmentation, such as crystallins and the melanocortin 1 receptor.
[255]
Similarly, comparing the genome of the Antarctic icefish, which lacks red blood cells,
to close relatives such as the zebrafish revealed genes needed to make these blood cells.
[256]

As evolution can produce highly optimized processes and networks, it has many
applications in computer science. Here, simulations of evolution using evolutionary

algorithms and artificial life started with the work of Nils Aall Barricelli in the 1960s, and
was extended by Alex Fraser, who published a series of papers on simulation of artificial
selection.[257] Artificial evolution became a widely recognized optimization method as a
result of the work of Ingo Rechenberg in the 1960s and early 1970s, who used evolution
strategies to solve complex engineering problems.[258] Genetic algorithms in particular
became popular through the writing of John Holland.[259] As academic interest grew,
dramatic increases in the power of computers allowed practical applications, including
the automatic evolution of computer programs.[260] Evolutionary algorithms are now used
to solve multi-dimensional problems more efficiently than software produced by human
designers, and also to optimize the design of systems.[261]

See also
Book:Evolution
Books are collections of articles which can be downloaded or ordered in print.

Geologic time scale

From Wikipedia, the free encyclopedia

Jump to: navigation, search

This clock representation shows some of the major units of geological time and definitive
events of Earth history. The Hadean eon represents the time before fossil record of life on
Earth; its upper boundary is now regarded as 4.0 Ga [1]. Other subdivisions reflect the
evolution of life; the Archean and Proterozoic are both eons, the Palaeozoic, Mesozoic
and Cenozoic are eras of the Phanerozoic eon. The two million year Quaternary period,
the time of recognizable humans, is too small to be visible at this scale.
The geologic time scale provides a system of chronologic measurement relating
stratigraphy to time that is used by geologists, paleontologists and other earth scientists to
describe the timing and relationships between events that have occurred during the
history of the Earth. The table of geologic time spans presented here agrees with the dates
and nomenclature proposed by the International Commission on Stratigraphy, and uses
the standard color codes of the United States Geological Survey.
Evidence from radiometric dating indicates that the Earth is about 4.570 billion years old.
The geological or deep time of Earth's past has been organized into various units
according to events which took place in each period. Different spans of time on the time
scale are usually delimited by major geological or paleontological events, such as mass
extinctions. For example, the boundary between the Cretaceous period and the Paleogene
period is defined by the CretaceousTertiary extinction event, which marked the demise

of the dinosaurs and of many marine species. Older periods which predate the reliable
fossil record are defined by absolute age.
Each era on the scale is separated from the next by a major event or change.

Contents
[hide]

1 Graphical timelines
2 Terminology
3 History of the time scale and names
4 Table of geologic time
5 See also
6 References and footnotes

7 External links

[edit] Graphical timelines

The second and third timelines are each subsections of their preceding timeline as
indicated by asterisks.

Millions of Years

The Holocene (the latest epoch) is too short to be shown clearly on this timeline.

[edit] Terminology
e h

Units in geochronology and stratigraphy[2]

Segments of rock
(strata) in
chronostratigraphy

Periods of time in
geochronology

Notes

Eonothem

Eon

4 total, half a
billion years or
more

Erathem

Era

12 total, several
hundred million
years

System

Period

Series

Epoch

tens of millions of
years

Stage

Age

millions of years

Chronozone

Chron

smaller than an
age/stage, not used
by the ICS
timescale

The largest defined unit of time is the supereon, composed of eons. Eons are divided into
eras, which are in turn divided into periods, epochs and ages. The terms eonothem,
erathem, system, series, and stage are used to refer to the layers of rock that correspond to
these periods of geologic time.

Geologists qualify these units as Upper, Middle, and Lower. Examples are "Upper
Jurassic" and "Middle Cambrian". The adjectives are capitalized when the subdivision is
formally recognized, and lower case when not; thus "early Miocene" but "Early Jurassic."
The names of subdivisions can vary when discussing either position in the geologic
record or corresponding time period, by use of either upper/lower (for series) or late/early
(for epochs). For example, the Lower Jurassic Series in geochronology corresponds to the
Early Jurassic Epoch in chronostratigraphy.[3]
Geologic units from the same time but different parts of the world often look different
and contain different fossils, so the same period was historically given different names in
different locales. For example, in North America the Lower Cambrian is called the
Waucoban series that is then subdivided into zones based on succession of trilobites. In
East Asia and Siberia, the same unit is split into Tommotian, Atdabanian, and Botomian
stages. A key aspect of the work of the International Commission on Stratigraphy is to
reconcile this conflicting terminology and define universal horizons that can be used
around the world.[4]

[edit] History of the time scale and names

Main articles: History of geology and History of paleontology

Animation showing Earth's palaeogeographic reconstruction beginning from early

Cambrian period.

Diagram of geological time scale, where the past is toward the bottom of the spiral
In classical antiquity, Aristotle saw that fossil seashells from rocks were similar to those
found on the beach and deduced that the fossils were once part of living animals. He
reasoned that the positions of land and sea had changed over long periods of time.
Leonardo da Vinci concurred with Aristotle's view that fossils were the remains of
ancient life.[5]

The 11th-century Persian geologist Avicenna (Ibn Sina) examined various fossils and
deduced that they originated from the petrifaction of plants and animals.[6] He also first
proposed one of the principles underlying geologic time scales: the law of superposition
of strata. While discussing the origins of mountains in The Book of Healing in 1027, he
outlined the principle as follows:[7][8]
It is also possible that the sea may have happened to flow little by little over the land consisting of
both plain and mountain, and then have ebbed away from it. ... It is possible that each time the
land was exposed by the ebbing of the sea a layer was left, since we see that some mountains
appear to have been piled up layer by layer, and it is therefore likely that the clay from which they
were formed was itself at one time arranged in layers. One layer was formed first, then at a
different period, a further was formed and piled, upon the first, and so on. Over each layer there
spread a substance of different material, which formed a partition between it and the next layer;
but when petrification took place something occurred to the partition which caused it to break up
and disintegrate from between the layers (possibly referring to unconformity). ... As to the
beginning of the sea, its clay is either sedimentary or primeval, the latter not being sedimentary. It
is probable that the sedimentary clay was formed by the disintegration of the strata of mountains.
Such is the formation of mountains.

His contemporary, Abu Rayhan Biruni (973-1048), discovered the existence of shells and
fossils in regions that once were seas and later became dry land, such as the Indian
subcontinent. Based on this evidence, he realized that the Earth is constantly evolving
and proposed that the Earth had an age, but that its origin was too distant to measure.[9]
Later in the 11th century, the Chinese naturalist, Shen Kuo (10311095), also recognized
the concept of 'deep time'.[10]
The principles underlying geologic (geological) time scales were later laid down by
Nicholas Steno in the late 17th century. Steno argued that rock layers (or strata) are laid
down in succession, and that each represents a "slice" of time. He also formulated the law
of superposition, which states that any given stratum is probably older than those above it
and younger than those below it. While Steno's principles were simple, applying them to
real rocks proved complex. Over the course of the 18th century geologists realized that:
1. Sequences of strata were often eroded, distorted, tilted, or even inverted after
deposition;
2. Strata laid down at the same time in different areas could have entirely different
appearances;
3. The strata of any given area represented only part of the Earth's long history.

A comparative geological timescale

The first serious attempts to formulate a geological time scale that could be applied
anywhere on Earth were made in the late 18th century. The most influential of those early
attempts (championed by Abraham Werner, among others) divided the rocks of the
Earth's crust into four types: Primary, Secondary, Tertiary, and Quaternary. Each type of
rock, according to the theory, formed during a specific period in Earth history. It was thus
possible to speak of a "Tertiary Period" as well as of "Tertiary Rocks." Indeed, "Tertiary"
(now Paleocene-Pliocene) and "Quaternary" (now Pleistocene-Holocene) remained in use
as names of geological periods well into the 20th century.
The Neptunist theories popular at this time (expounded by Werner) proposed that all
rocks had precipitated out of a single enormous flood. A major shift in thinking came
when James Hutton presented his Theory of the Earth; or, an Investigation of the Laws
Observable in the Composition, Dissolution, and Restoration of Land Upon the Globe
before the Royal Society of Edinburgh in March and April 1785. It has been said that "as
things appear from the perspective of the twentieth century, James Hutton in those
reading became the founder of modern geology"[11] Hutton proposed that the interior of
the Earth was hot, and that this heat was the engine which drove the creation of new rock:
land was eroded by air and water and deposited as layers in the sea; heat then
consolidated the sediment into stone, and uplifted it into new lands. This theory was
dubbed "Plutonist" in contrast to the flood-oriented theory.
The identification of strata by the fossils they contained, pioneered by William Smith,
Georges Cuvier, Jean d'Omalius d'Halloy, and Alexandre Brogniart in the early 19th
century, enabled geologists to divide Earth history more precisely. It also enabled them to
correlate strata across national (or even continental) boundaries. If two strata (however
distant in space or different in composition) contained the same fossils, chances were
good that they had been laid down at the same time. Detailed studies between 1820 and
1850 of the strata and fossils of Europe produced the sequence of geological periods still
used today.
The process was dominated by British geologists, and the names of the periods reflect
that dominance. The "Cambrian," (the Roman name for Wales) and the "Ordovician," and
"Silurian", named after ancient Welsh tribes, were periods defined using stratigraphic
sequences from Wales.[12] The "Devonian" was named for the English county of Devon,
and the name "Carboniferous" was simply an adaptation of "the Coal Measures," the old
British geologists' term for the same set of strata. The "Permian" was named after Perm,
Russia, because it was defined using strata in that region by Scottish geologist Roderick
Murchison. However, some periods were defined by geologists from other countries. The
"Triassic" was named in 1834 by a German geologist Friedrich Von Alberti from the three
distinct layers (Latin trias meaning triad) red beds, capped by chalk, followed by black
shales that are found throughout Germany and Northwest Europe, called the 'Trias'.
The "Jurassic" was named by a French geologist Alexandre Brogniart for the extensive
marine limestone exposures of the Jura Mountains. The "Cretaceous" (from Latin creta
meaning 'chalk') as a separate period was first defined by Belgian geologist Jean
d'Omalius d'Halloy in 1822, using strata in the Paris basin[13] and named for the extensive
beds of chalk (calcium carbonate deposited by the shells of marine invertebrates).

British geologists were also responsible for the grouping of periods into Eras and the
subdivision of the Tertiary and Quaternary periods into epochs.
When William Smith and Sir Charles Lyell first recognized that rock strata represented
successive time periods, time scales could be estimated only very imprecisely since
various kinds of rates of change used in estimation were highly variable. While
creationists had been proposing dates of around six or seven thousand years for the age of
the Earth based on the Bible, early geologists were suggesting millions of years for
geologic periods with some even suggesting a virtually infinite age for the Earth.
Geologists and paleontologists constructed the geologic table based on the relative
positions of different strata and fossils, and estimated the time scales based on studying
rates of various kinds of weathering, erosion, sedimentation, and lithification. Until the
discovery of radioactivity in 1896 and the development of its geological applications
through radiometric dating during the first half of the 20th century (pioneered by such
geologists as Arthur Holmes) which allowed for more precise absolute dating of rocks,
the ages of various rock strata and the age of the Earth were the subject of considerable
debate.
The first geologic time scale was eventually published in 1913 by the British geologist
Arthur Holmes.[14] He greatly furthered the newly created discipline of geochronology
and published the world renowned book The Age of the Earth in 1913 in which he
estimated the Earth's age to be at least 1.6 billion years.[15]
In 1977, the Global Commission on Stratigraphy (now the International Commission on
Stratigraphy) started an effort to define global references (Global Boundary Stratotype
Sections and Points) for geologic periods and faunal stages. The commission's most
recent work is described in the 2004 geologic time scale of Gradstein et al.[16]. A UML
model for how the timescale is structured, relating it to the GSSP, is also available[17].

[edit] Table of geologic time

The following table summarizes the major events and characteristics of the periods of
time making up the geologic time scale. As above, this time scale is based on the
International Commission on Stratigraphy. (See lunar geologic timescale for a discussion
of the geologic subdivisions of Earth's moon.) This table is arranged with the most recent
geologic periods at the top, and the most ancient at the bottom. The height of each table
entry does not correspond to the duration of each subdivision of time.
The content of the table is based on the current official geologic time scale of the
International Commission on Stratigraphy,[18] with the epoch names altered to the
early/late format from lower/upper as recommended by the ICS when dealing with
chronostratigraphy.[3]
vde

Geologic time scale[hide]

Super

Eon

Era

Period[1

Epoch

Age[20]

Major events

Start,

eon

million
years
ago[20]

9]

Phanero Cenozoic[2 Quatern Holocene

1]
zoic
ary

Atlantic
Boreal

The last
0.011430
glacial period 0.00013[
ends; rise of 21][23]
human
civilization.
Quaternary
Ice Age
recedes, and
the current
interglacial
begins.
Younger
Dryas cold
spell occurs,
Sahara forms
from
savannah, and
agriculture
begins,
allowing
humans to
build cities.
Paleolithic/N
eolithic
(Stone Age)
cultures begin
around 10000
BC, giving
way to
Copper Age
(3500 BC)
and Bronze
Age (2500
BC). Cultures
continue to
grow in
complexity
and technical
advancement
through the
Iron Age
(1200 BC),
giving rise to
many prehistoric

cultures
throughout
the world,
eventually
leading into
Classical
Antiquity,
such as the
Roman
Empire and
even to the
Middle Ages
and present
day. Little Ice
Age (stadial)
causes brief
cooling in
Northern
Hemisphere
from 1400 to
1850. Also
refer to the
List of
archaeologica
l periods for
clarification
on early
cultures and
ages. Mount
Tambora
erupts in
1815, causing
the Year
Without a
Summer
(1816) in
Europe and
North
America from
a volcanic
winter.
Atmospheric
CO2 levels
start creeping
from 100
parts per
million
volume

(ppmv) at the
end of the last
glaciation to
the current
level of 385
ppmv,
causing
global
warming and
Pleistocene Late/Tyrrhenian Flourishing
0.126
Stage/Eemian/San and then
0.005*
gamonian
extinction of
many large
Middle
0.500?
mammals
(Pleistocene 1.806
Early
*
megafauna). 0.005
Evolution of 2.588
Gelasian
anatomically 0.005*
modern
humans.
Quaternary
Ice Age
continues
with
glaciations
and
interstadials
(and the
accompanyin
g fluctuations
from 100 to
300 ppmv in
atmospheric
CO2
levels[22]),
further
intensificatio
n of Icehouse
Earth
conditions,

roughly 1.6
Ma. Last
glacial
maximum
(30000 years
ago), last
glacial period
(18000
15000 years
ago). Dawn
of human
stone-age
cultures, with
increasing
technical
complexity
relative to
previous ice
age cultures,
such as
engravings
and clay
statues (e.g.
Venus of
Lespugue),
particularly in
the
Mediterranea
n and Europe.
Lake Toba
supervolcano
erupts 75000
years before
present,
causing a
volcanic
winter that
pushes
humanity to
the brink of
extinction.
Pleistocene
ends with
Oldest Dryas,
Older
Dryas/Aller
d and
Younger

Neoge
ne

Pliocene

Miocene

Dryas climate
events, with
Younger
Dryas
Piacenzian/Blanca Intensificatio 3.600
n
n of present 0.005*
Icehouse
conditions,
present
(Quaternary)
ice age begins
roughly 2.58
Ma; cool and
dry climate.
Australopithe
cines, many 5.332 *
Zanclean
0.005
of the
existing
genera of
mammals,
and recent
mollusks
appear. Homo
habilis
appears.
Messinian
Tortonian
Serravallian
Langhian
Burdigalian
Aquitanian

Moderate
Icehouse
climate,
puncuated by
ice ages;
Orogeny in
northern
hemisphere.
Modern
mammal and
bird families
became
recognizable.
Horses and
mastodons
diverse.
Grasses
become

7.246
0.05*
11.608
0.05*
13.65
0.05*
15.97
0.05*
20.43
0.05*
23.03
0.05*

ubiquitous.
First apes
appear (for
reference see
the article:
"Sahelanthrop
us
tchadensis").
Kaikoura
Orogeny
forms
Southern Alps
in New
Zealand,
continues
today.
Orogeny of
the Alps in
Europe slows,
but continues
to this day.
Carpathean
orogeny
forms
Carpathian
Mountains in
Central and
Eastern
Europe.
Hellenic
orogeny in
Greece and
Aegean Sea
slows, but
continues to
this day.
Middle
Miocene
Disruption
occurs.
Widespread
forests slowly
draw in
massive
amounts of
CO2,
gradually
lowering the

Chattian

Oligocene

Eocene
Paleog
ene

Rupelian

Priabonian
Bartonian
Lutetian
Ypresian

level of
atmospheric
CO2 from 650
Warm but
28.4 0.1*
cooling
climate,
moving
towards
Icehouse;
Rapid
evolution and
diversificatio
33.9
n of fauna,
0.1*
especially
mammals.
Major
evolution and
dispersal of
modern types
of flowering
plants
Moderate,
cooling
climate.
Archaic
mammals
(e.g.
Creodonts,
Condylarths,
Uintatheres,
etc) flourish
and continue
to develop
during the
epoch.
Appearance
of several
"modern"
mammal
families.
Primitive
whales
diversify.
First grasses.

37.2 0.1*
40.4
0.2*
48.6
0.2*
55.8
0.2*

Paleocene Thanetian
Selandian
Danian

Reglaciation
of Antarctica
and formation
of its ice cap;
Azolla event
triggers ice
age, and the
Icehouse
Earth climate
that would
follow it to
this day, from
the settlement
and decay of
seafloor algae
drawing in
massive
amounts of
atmospheric
carbon
dioxide[22],
Climate
58.7 0.2*
tropical.
61.7
Modern
0.3*
plants appear;
65.5
Mammals
*
diversify into 0.3

Mesozoic Cretace
ous

Maastrichtian
Campanian
Santonian
Late

Coniacian
Turonian
Cenomanian

Early

Albian
Aptian
Barremian
Hauterivian
Valanginian

a number of
primitive
lineages
following the
extinction of
the dinosaurs.
First large
mammals (up
to bear or
small hippo
size). Alpine
orogeny in
Flowering
70.6 0.6*
plants
83.5
proliferate,
0.7*
along with
new types of 85.8
*
insects. More 0.7
modern
89.3
teleost fish
1.0*
begin to
93.5
appear.
*
Ammonites, 0.8
belemnites, 99.6
rudist
0.9*
bivalves,
112.0
echinoids and
1.0*
sponges all
125.0
common.
1.0*
Many new
types of
130.0
dinosaurs
1.5*
(e.g.
Tyrannosaurs, 136.4*
Titanosaurs, 2.0
duck bills,
140.2

 3.0*

Berriasian

Jurassi
c

Tithonian
Late

Kimmeridgian
Oxfordian

Middle

Callovian
Bathonian
Bajocian

and horned
dinosaurs)
evolve on
land, as do
Eusuchia
(modern
crocodilians);
and
mosasaurs
and modern
sharks appear
in the sea.
Primitive
birds
gradually
replace
pterosaurs.
Gymnosperm
s (especially
conifers,
Bennettitales
and cycads)
and ferns
common.
Many types
of dinosaurs,
such as
sauropods,
carnosaurs,
and

145.5
4.0*

150.8
4.0*
155.7
4.0*
161.2
4.0*
164.7
4.0
167.7
3.5*
171.6

 3.0*
Aalenian

175.6
2.0*

Toarcian

183.0
1.5*

Pliensbachian

189.6
1.5*

Sinemurian

196.5
1.0*

Early

Hettangian

Triassi Late
c

Rhaetian

stegosaurs.
Mammals
common but
small. First
birds and
199.6
lizards.
0.6*
Ichthyosaurs
and
plesiosaurs
diverse.
Bivalves,
Ammonites
and
belemnites
abundant. Sea
urchins very
common,
along with
crinoids,
starfish,
sponges, and
terebratulid
and
Archosaurs 203.6
dominant on 1.5*

Norian
Carnian
Ladinian
Middle
Anisian
Early

Olenekian
Induan

land as
dinosaurs, in
the oceans as
Ichthyosaurs
and
nothosaurs,
and in the air
as pterosaurs.
Cynodonts
become
smaller and
more
mammal-like,
while first
mammals and
crocodilia
appear.
Dicrodium
flora common
on land.
Many large
aquatic
temnospondyl
amphibians.
Ceratitic
ammonoids
extremely
common.
Modern
corals and
teleost fish
appear, as do
many modern
insect clades.
Andean
Orogeny in
South
America.
Cimmerian
Orogeny in
Asia.
Rangitata
Orogeny
begins in
New Zealand.
HunterBowen
Orogeny in

216.5
2.0*
228.0
2.0*
237.0
2.0*
245.0
1.5*
249.7
1.5*
251.0
0.7*

Northern
Australia,
Queensland
and New
Paleozoic Permian
Landmasses 253.8
Changhsingian
unite into
0.7*
Lopingian
supercontinen 260.4
Wuchiapingian
t Pangaea,
0.7*
creating the
Appalachians. 265.8
Capitanian
0.7*
End of
Guadalupi Wordian/Kazania Permo268.4
Carboniferou 0.7*
an
n
s glaciation.
270.6
Roadian/Ufimian Synapsid
0.7*
reptiles
(pelycosaurs 275.6
Cisuralian
Kungurian
and
0.7*
therapsids)
284.4
become
Artinskian
0.7*
plentiful,
294.6
while
Sakmarian
parareptiles 0.8*
and
Asselian
299.0
temnospondyl 0.8*
amphibians
remain
common. In
the midPermian,
coal-age flora
are replaced
by conebearing
gymnosperms
(the first true
seed plants)
and by the
first true
mosses.
Beetles and
flies evolve.
Marine life
flourishes in
warm shallow

Carbon
iferous[ Late
24]
/
Pennsy
Middle

Gzhelian
Kasimovian
Moscovian

reefs;
productid and
spiriferid
brachiopods,
bivalves,
forams, and
ammonoids
all abundant.
PermianTriassic
extinction
event occurs
251 Ma: 95%
of life on
Earth
becomes
extinct,
including all
trilobites,
graptolites,
and blastoids.
Winged
303.9
insects radiate 0.9*
suddenly;
306.5
some (esp.
1.0*
Protodonata
311.7

 1.1*

lEarly
vanian

Carbon
Late
iferous[
24]
Middle
/
Mississippian Early

Bashkirian

Serpukhovian
Visan
Tournaisian

318.1
and
1.3*
Palaeodictyop
tera) are quite
large.
Amphibians
common and
diverse. First
reptiles and
coal forests
(scale trees,
ferns, club
trees, giant
horsetails,
Cordaites,
etc.). Highestever
atmospheric
oxygen
levels.
Goniatites,
brachiopods,
Large
326.4
primitive
1.6*
trees, first
345.3
land
2.1*
vertebrates,
359.2
and

Devoni
an

Famennian
Late
Frasnian
Middle

Givetian
Eifelian

2.5*
amphibious
sea-scorpions
live amid
coal-forming
coastal
swamps.
Lobe-finned
rhizodonts are
dominant big
fresh-water
predators. In
the oceans,
early sharks
are common
and quite
diverse;
echinoderms
(especially
crinoids and
blastoids)
First
374.5
clubmosses, 2.6*
horsetails and 385.3
ferns appear, 2.6*
as do the first
seed-bearing 391.8
2.7*
plants
(progymnosp 397.5

Early

Emsian
Pragian
Lochkovian

erms), first
trees (the
progymnospe
rm
Archaeopteri
s), and first
(wingless)
insects.
Strophomenid
and atrypid
brachiopods,
rugose and
tabulate
corals, and
crinoids are
all abundant
in the oceans.
Goniatite
ammonoids
are plentiful,
while squidlike coleoids
arise.
Trilobites and
armoured
agnaths
decline, while
jawed fishes
(placoderms,
lobe-finned
and rayfinned fish,
and early
sharks) rule
the seas. First
amphibians
still aquatic.
"Old Red
Continent" of
Euramerica.
Beginning of
Acadian
Orogeny for
Anti-Atlas
Mountains of
North Africa,
and
Appalachian

2.7*
407.0
2.8*
411.2
2.8*
416.0
2.8*

Mountains of
North
America, also
the Antler,
Siluria
no faunal stages First Vascular 418.7
Pridoli
n
defined
plants (the
2.7*
rhyniophytes 421.3
Ludfordian
and their
2.6*
Ludlow/Ca
relatives),
yugan
422.9
first
Gorstian
2.5*
millipedes
Homerian/Lockpor and
426.2
arthropleurids 2.4*
tian
Wenlock
on land. First
Sheinwoodian/To jawed fishes, 428.2
nawandan
2.3*
as well as
Llandover Telychian/Ontaria many
436.0
armoured
y/
n
1.9*
jawless fish,
Alexandria
439.0
populate the
Aeronian
n
1.8*
seas. SeaRhuddanian
443.7
scorpions
1.5*
reach large
size. Tabulate
and rugose
corals,
brachiopods
(Pentamerida
,
Rhynchonelli
da, etc.), and
crinoids all
abundant.
Trilobites and
mollusks
diverse;
graptolites
not as varied.
Beginning of
Caledonian
Orogeny for
hills in
England,
Ireland,

Ordovi
cian

Hirnantian
Late

other faunal
stages
Darriwilian

Middle

Early

other faunal
stages
Arenig
Tremadocian

Wales,
Scotland, and
the
Scandinavian
Mountains.
Also
continued
into Devonian
period as the
Acadian
Invertebrates 445.6
diversify into 1.5*
many new
460.9
types (e.g.,
1.6*
long straight468.1
shelled
*
cephalopods). 1.6
Early corals, 471.8
articulate
1.6*
brachiopods
478.6
(Orthida,
*
Strophomenid 1.7
a, etc.),
488.3
bivalves,
1.7*
nautiloids,
trilobites,
ostracods,
bryozoa,
many types of
echinoderms
(crinoids,
cystoids,
starfish, etc.),
branched
graptolites,
and other taxa
all common.
Conodonts

(early
planktonic
vertebrates)
appear. First
green plants
Cambri
Major
496.0
other faunal stages
an
diversificatio 2.0*
n of life in the
Furongian Paibian/Ibexian/ Cambrian
Ayusokkanian/Sa
501.0
Explosion.
kian/
2.0*
Many
fossils;
Aksayan
most modern
other faunal
animal phyla 513.0
Middle
stages/Albertan
appear. First 2.0
chordates
Early
other faunal
542.0
appear, along 1.0*
stages/
Waucoban/Tommo with a
number of
tian/
Atdabanian/Botom extinct,
problematic
ian
phyla. Reefbuilding
Archaeocyath
a abundant;
then vanish.
Trilobites,
priapulid
worms,
sponges,
inarticulate
brachiopods
(unhinged
lampshells),
and many
other animals
numerous.
Anomalocari
ds are giant
predators,
while many
Ediacaran
fauna die out.
Prokaryotes,
protists (e.g.,

forams),
fungi and
algae
continue to
present day.
Gondwana
emerges.
Petermann
Orogeny on
the Australian
Continent
tapers off
(550-535
Ma). Ross
Orogeny in
Antarctica.
Adelaide
Geosyncline
(Delamerian
Orogeny),
Preca Proter- NeoEdiacar Good fossils of the first multi-celled animals. 630 +5/mozoic[26] proterozoi an
Ediacaran biota flourish worldwide in seas. 30*
[2
[26]
brian
c
Simple trace fossils of possible worm-like
5]
Trichophycus, etc. First sponges and
trilobitomorphs. Enigmatic forms include
many soft-jellied creatures shaped like bags,
disks, or quilts (like Dickinsonia). Taconic
Orogeny in North America. Aravalli Range

orogeny in Indian Subcontinent. Beginning of

Petermann Orogeny on Australian Continent.
Beardmore Orogeny in Antarctica, 633-620
Ma.
Possible "Snowball Earth" period. Fossils still
Cryoge rare. Rodinia landmass begins to break up.
850[27]
nian
Late Ruker / Nimrod Orogeny in Antarctica
tapers off.
Rodinia supercontinent persists. Trace fossils
of simple multi-celled eukaryotes. First
radiation of dinoflagellate-like acritarchs.
Grenville Orogeny tapers off in North
America. Pan-African orogeny in Africa.
Lake Ruker / Nimrod Orogeny in Antarctica,
Tonian
1000[27]
1000 150 Ma. Edmundian Orogeny (c. 920
- 850 Ma), Gascoyne Complex, Western
Australia. Adelaide Geosyncline laid down on
Australian Continent, beginning of Adelaide
Geosyncline (Delamerian Orogeny) in that
continent.
Narrow highly metamorphic belts due to
orogeny as Rodinia formed. Late Ruker /
Stenian Nimrod Orogeny in Antarctica possibly
begins. Musgrave Orogeny (c. 1080 Ma),
Musgrave Block, Central Australia.

1200[27]

Platform covers expand. Barramundi

Calym Orogeny, MacArthur Basin, Northern
mian Australia, and Isan Orogeny, c. 1600 Ma,
Mount Isa Block, Queensland

1600[27]

MesoPlatform covers continue to expand. Green

proterozoi Ectasia
algae colonies in the seas. Grenville Orogeny 1400[27]
c[26]
n
in North America.

Paleoproterozoi
c[26]

First complex single-celled life: protists with

nuclei. Columbia is the primordial
supercontinent. Kimban Orogeny in
Australian Continent ends. Yapungku
Statheri
Orogeny on North Yilgarn craton, in Western 1800[27]
an
Australia. Mangaroon Orogeny, 1680-1620
Ma, on the Gascoyne Complex in Western
Australia. Kararan Orogeny (1650- Ma),
Gawler Craton, South Australia.
Orosiri The atmosphere became oxygenic. Vredefort 2050[27]
an
and Sudbury Basin asteroid impacts. Much
orogeny. Penokean and Trans-Hudsonian
Orogenies in North America. Early Ruker

Orogeny in Antarctica, 2000 - 1700 Ma.

Glenburgh Orogeny, Glenburgh Terrane,
Australian Continent c. 2005 - 1920 Ma.
Kimban Orogeny, Gawler craton in Australian
Continent begins.
Rhyaci Bushveld Formation formed. Huronian
an
glaciation.

2300[27]

Oxygen catastrophe: banded iron formations

Sideria
formed. Sleaford Orogeny on Australian
2500[27]
n
Continent, Gawler Craton 2440-2420 Ma.
Stabilization of most modern cratons; possible mantle
Neoarchea overturn event. Insell Orogeny, 2650 150 Ma.
2800[27]
n[26]
Abitibi greenstone belt in present-day Ontario and
Quebec begins to form, stablizes by 2600 Ma.
First stromatolites (probably colonial cyanobacteria).
Oldest macrofossils. Humboldt Orogeny in Antarctica.
Mesoarch
Blake River Megacaldera Complex begins to form in 3200[27]
ean[26]
present-day Ontario and Quebec, ends by roughly
Archea
[26]
2696 Ma.
n
First known oxygen-producing bacteria. Oldest
definitive microfossils. Oldest cratons on earth (such
Paleoarch
as the Canadian Shield and the Pilbara Craton) may 3600[27]
ean[26]
have formed during this period[28]. Rayner Orogeny in
Antarctica.
Eoarchea Simple single-celled life (probably bacteria and
n[26]
perhaps archaea). Oldest probable microfossils.

3800

Early
This era overlaps the end of the Late Heavy
Imbrian[26]
Bombardment of the inner solar system.
[30]

c.3850

Nectarian[
26][30]

Hadean
[26][29]

c.3920

Oldest known rock (4030 Ma)[31]. The first Lifeforms

Basin
and self-replicating RNA molecules may have evolved
Groups[26]
c.4150
on earth around 4000 Ma during this era. Napier
[30]
Orogeny in Antarctica, 4000 200 Ma.
Cryptic[26]
[30]

[edit] See also

This unit gets its name from the lunar geologic

timescale when the Nectaris Basin and other major
lunar basins were formed by large impact events.

Age of the Earth

Oldest known mineral (Zircon, 44068 Ma[32]).

Formation of Moon (4533 Ma), probably from giant
impact. Formation of Earth (4567.17 to 4570 Ma)

c.4570

Anthropocene/Homogenocene
Deep time
Geological history of Earth
Geology of Mars/areology
Graphical timeline of the universe

History of the Earth

List of fossil

Timeline of human evolution

From Wikipedia, the free encyclopedia

Jump to: navigation, search

This article is about the timeline of human evolution. For a timeline of general evolution
see Timeline of evolution.
See also Human evolution for more details on this topic.

Evolutionary tree
The timeline of human evolution outlines the major events in the development of
human species, and the evolution of humans' ancestors. It includes a brief explanation of
some animals, species or genera, which are possible ancestors of Homo sapiens sapiens.
It does not address the origin of life, which is addressed by abiogenesis, but presents a
possible line of descendants that led to humans. This timeline is based on studies from

paleontology, developmental biology, morphology and from anatomical and genetic data.
The study of human evolution is a major component of anthropology.

Contents
[hide]

1 Homo sapiens taxonomy

2 Timeline
o 2.1 First living beings
o 2.2 Chordates
o 2.3 Tetrapodes
o 2.4 Mammals
o 2.5 Primates
o 2.6 Hominidae
o 2.7 Homo
3 See also
4 References

5 External links

[edit] Homo sapiens taxonomy

The cladistic line of descent (taxonomic rank) of homo sapiens sapiens (modern humans)
is as follows:
domain: eukaryotes (2.100.000.000 years ago)
kingdom: animalia (590.000.000 years ago)
phylum: chordata (530.000.000 years ago)
subphylum: vertebrata (505.000.000 years ago)
class: mammalia (220.000.000 years ago)
subclass: theriiformes
infraclass: eutheria (125.000.000 years ago)
magnorder: boreoeutheria
superorder: euarchontoglires (supraprimates) (100.000.000 years ago)
order: primates (75.000.000 years ago)
suborder: haplorrhini (tarsiers, monkeys, apes, "dry-nosed" primates) (40.000.000 years ago)
infraorder: simiiformes (simians, "higher" primates)
parvorder: catarrhini ("narrow nosed" primates) (30.000.000 years ago)
superfamily: hominoidea (25.000.000 years ago)
family: hominidae (great apes) (15.000.000 years ago)
subfamily: homininae (4.500.000 years ago)
tribe: hominini
subtribe: hominina (3.000.000 years ago)
genus: homo (2.500.000 years ago)
species: homo sapiens (500.000 years ago)
sub-species: homo sapiens sapiens (200.000 years ago)

[edit] Timeline
[edit] First living beings
Date

Event

4000 Ma The earliest life appears.

(million
years ago) Further information: Origin of life

3900 Ma Cells resembling prokaryotes appear.

Further information: Cell (biology)#Origins of cells

2500 Ma First organisms to utilize oxygen.

2100 Ma More complex cells appear: the eukaryotes.

Further information: Eukaryote#Origin and evolution
1200 Ma Sexual reproduction evolves, leading to faster evolution.[1]

900 Ma

Choanoflagellate
The choanoflagellates may look similar to the ancestors of the entire animal
kingdom, and in particular they may be the direct ancestors of Sponges.[2]
Proterospongia (members of the Choanoflagellata) are the best living examples
of what the ancestor of all animals may have looked like.
They live in colonies, and show a primitive level of cellular specialization for
different tasks.

600 Ma It is thought that the earliest multicellular animal was a sponge-like creature.
Sponges are among the simplest of animals, with partially differentiated
tissues.
Sponges (Porifera) are the phylogenetically oldest animal phylum extant today.
580 Ma The movement of all animals may have started with cnidarians. Almost all
cnidarians possess nerves and muscles and, because they are the simplest
animals to possess it, their direct ancestors were very likely the first animals to
use nerves and muscles together. Cnidarians are also the first animals with an
actual body of definite form and shape. They have radial symmetry. The first
eyes evolved at this time.
550 Ma

Flatworm
Flatworms are the earliest animals to have a brain, and the simplest animals
alive to have bilateral symmetry. They are also the simplest animals with
organs that form from three germ layers.
540 Ma Acorn worms are considered more highly specialised and advanced than other
similarly shaped worm-like creatures. They have a circulatory system with a
heart that also functions as a kidney. Acorn worms have a gill-like structure
used for breathing, a structure similar to that of primitive fish. Acorn worms
are thus sometimes said to be a link between vertebrates and invertebrates[citation
needed]
.

[edit] Chordates
Date

Event

530 Ma

Pikaia
One of the earliest known ancestor of the chordates is Pikaia.[3] It is the first

known animal with a notochord. Pikaia is believed to be the ancestor of all

chordates and vertebrates.[4]
The Lancelet, still living today, retains some characteristics of the primitive
chordates. It resembles Pikaia
Other earliest known chordate-like fossils is from a conodonts an "eel-shaped
animal of 4-20 cm (1-8 in) long" with a pair of huge eyes at the head end and a
complex basket of teeth.
505 Ma

Agnatha
The first vertebrates appear: the ostracoderms, jawless fish related to present-day
lampreys and hagfishes. Haikouichthys and Myllokunmingia are examples of
these jawless fish, or Agnatha. (See also prehistoric fish). They were jawless and
their internal skeletons were cartilaginous. They lacked the paired (pectoral and
pelvic) fins of more advanced fish. They were the Precursors to the Osteichthyes
(bony fish). [5]
480 Ma

A Placoderm
The Placodermi were prehistoric fishes. Placoderms were the first of the jawed
fishes, their jaws evolving from the first of their gill arches [6]. Their head and
thorax were covered by articulated armoured plates and the rest of the body was
scaled or naked.
400 Ma First Coelacanth appears; this order of animals had been thought to have no
extant members until living specimens were discovered in 1938. It is often
referred to as a living fossil.
375 Ma Tiktaalik is a genus of sarcopterygian (lobe-finned) fishes from the late Devonian
with many tetrapod-like features. It shows a clear link between Panderichthys and
Acanthostega.

[edit] Tetrapodes
Date

Event

365 Ma

Panderichthys
Some fresh water lobe-finned fish (Sarcopterygii) develop legs and give rise to
the Tetrapoda.
The first tetrapods evolved in shallow and swampy freshwater habitats.
Primitive tetrapods developed from a lobe-finned fish (an "osteolepid
Sarcopterygian"), with a two-lobed brain in a flattened skull, a wide mouth and a
short snout, whose upward-facing eyes show that it was a bottom-dweller, and
which had already developed adaptations of fins with fleshy bases and bones. The
"living fossil" coelacanth is a related lobe-finned fish without these shallow-water
adaptations. These fishes used their fins as paddles in shallow-water habitats
choked with plants and detritus. The universal tetrapod characteristics of front
limbs that bend backward at the elbow and hind limbs that bend forward at the
knee can plausibly be traced to early tetrapods living in shallow water.[7]
Panderichthys is a 90-130 cm (35-50 in) long fish from the Late Devonian period.
It has a large tetrapod-like head. Panderichthys exhibits features transitional
between lobe-finned fishes and early tetrapods.
Lungfishes retain some characteristics of the early Tetrapodas. One example is
the Queensland Lungfish.
315 Ma

Acanthostega

Ichthyostega
Acanthostega is an extinct amphibian, among the first animals to have
recognizable limbs. It is a candidate for being one of the first vertebrates to be
capable of coming onto land. It lacked wrists, and was generally poorly adapted
for life on land. The limbs could not support the animal's weight. Acanthostega
had both lungs and gills, also indicating it was a link between lobe-finned fish

and terrestrial vertebrates.

Ichthyostega is an early tetrapod. Being one of the first animals with legs, arms,
and finger bones, Ichthyostega is seen as a hybrid between a fish and an
amphibian. Ichthyostega' had legs but its limbs probably weren't used for
walking, they may have spent very brief periods out of water and would have
used their legs to paw their way through the mud.[8]
Amphibia were the first four-legged animals to develop lungs.
Amphibians living today still retain many characteristics of the early tetrapods.
300 Ma

Hylonomus
From amphibians came the first reptiles: Hylonomus is the earliest known reptile.
It was 20 cm (8 in) long (including the tail) and probably would have looked
rather similar to modern lizards. It had small sharp teeth and probably ate
millipedes and early insects. It is a precursor of later Amniotes and mammal-like
reptiles.
Evolution of the amniotic egg gives rise to the Amniota, reptiles that can
reproduce on land and lay eggs on dry land. They did not need to return to water
for reproduction. This adaptation gave them the capability to colonize the uplands
for the first time.
Reptiles have advanced nervous system, compared to amphibians. They have
twelve pairs of cranial nerves.

[edit] Mammals
Date

256 Ma

Event

Phthinosuchus, an early Therapsid

Shortly after the appearance of the first reptiles, two branches split off. One
branch is the Diapsids, from which come the modern reptiles. The other branch is
Synapsida, which had temporal fenestra, a pair of holes in their skulls behind the
eyes, which were used to increase the space for jaw muscles.
The earliest mammal-like reptiles are the pelycosaurs. The pelycosaurs were the
first animals to have temporal fenestra. Pelycosaurs are not Therapsids but soon
they gave rise to them. The Therapsida were the direct ancestor of mammals.
The therapsids have temporal fenestrae larger and more mammal-like than
pelycosaurs, their teeth show more serial differentiation; and later forms had
evolved a secondary palate. A secondary palate enables the animal to eat and
breathe at the same time and is a sign of a more active, perhaps warm-blooded,
way of life. [9]
220 Ma One sub-group of therapsids, the cynodonts evolved more mammal-like
characteristics.
The jaws of cynodonts resemble modern mammal jaws. It is very likely this
group of animals contains a species which is the direct ancestor of all modern
mammals.[10]
220 Ma

Repenomamus
From Eucynodontia (cynodonts) came the first mammals. Most early mammals
were small and shrew-like animals that fed on insects. Although there is no
evidence in the fossil record, it is likely that these animals had a constant body
temperature, milk glands for their young. The neocortex region of the brain first
evolved in mammals and thus is unique to them.
125 Ma

Eomaia scansoria

Eomaia scansoria, a eutherian mammal, leads to the formation of modern

placental mammals. It looks like modern dormouse, climbing small shrubs in
Liaoning, China.
100 Ma Common genetic ancestor of mice and humans (base of the clade
Euarchontoglires).

[edit] Primates
Date

Event

6585 Ma

Carpolestes simpsoni

A Plesiadapis without fur.

A group of small, nocturnal and arboreal, insect-eating mammals called the
Euarchonta begins a speciation that will lead to the primate, treeshrew and
flying lemur orders. The Primatomorpha is a subdivision of Euarchonta that
includes the primates and the proto-primate Plesiadapiformes. One of the early
proto-primates is Plesiadapis. Plesiadapis still had claws and the eyes located
on each side of the head. Because of this they were faster on the ground than
on the top of the trees, but they began to spend long times on lower branches of
trees, feeding on fruits and leaves. The Plesiadapiformes very likely contain the
species which is the ancestor of all primates.[11]
One of the last Plesiadapiformes is Carpolestes simpsoni. It had grasping digits

but no forward facing eyes.

47 Ma Darwinius masillae, a transitional form between the prosimians (lemurs and
other primitive primates) and the simians (monkeys, apes). It looked much like
a lemur but had opposable thumbs.
40 Ma Primates diverge into suborders Strepsirrhini (wet-nosed primates) and
Haplorrhini (dry nosed primates). Strepsirrhini contains most of the
prosimians; modern examples include the lemurs and lorises. The haplorrhines
include the three living groups the prosimian tarsiers, the simian monkeys, and
apes. One of the earliest haplorrhines is Teilhardina asiatica, a mouse-sized,
diurnal creature with small eyes. The Haplorrhini metabolism lost the ability to
make its own Vitamin C. This means that it and all its descendants had to
include fruit in its diet, where Vitamin C could be obtained externally.
30 Ma

Aegyptopithecus
Haplorrhini splits into infraorders Platyrrhini and Catarrhini. Platyrrhines, New
World monkeys, have prehensile tails and males are color blind. They may
have migrated to South America on a raft of vegetation across the Atlantic
ocean (circa 4,500 km, 2,800 mi). Catarrhines mostly stayed in Africa as the
two continents drifted apart. One ancestor of catarrhines might be
Aegyptopithecus.
25 Ma

Proconsul
Catarrhini splits into 2 superfamilies, Old World monkeys (Cercopithecoidea)
and apes (Hominoidea). Our trichromatic color vision had its genetic origins in
this period.
Proconsul was an early genus of catarrhine primates. They had a mixture of
Old World monkey and ape characteristics. Proconsul's monkey-like features

include thin tooth enamel, a light build with a narrow chest and short
forelimbs, and an arboreal quadrupedal lifestyle. Its ape-like features are its
lack of a tail, ape-like elbows, and a slightly larger brain relative to body size.
Proconsul africanus is a possible ancestor of both great and lesser apes, and
humans.

[edit] Hominidae
Date

Event

15 Ma Hominidae (great apes) speciate from the ancestors of the gibbon (lesser apes).
13 Ma Homininae ancestors speciate from the ancestors of the orangutan[12].
Pierolapithecus catalaunicus is believed to be a common ancestor of humans and
the great apes or at least a species that brings us closer to a common ancestor than
any previous fossil discovery.
Pierolapithecus had special adaptations for tree climbing, just as humans and
other great apes do: a wide, flat ribcage, a stiff lower spine, flexible wrists, and
shoulder blades that lie along its back.
10 Ma Hominini speciate from the ancestors of the gorillas.
7 Ma

Sahelanthropus tchadensis
Hominina speciate from the ancestors of the chimpanzees. The latest common
ancestor lived around the time of Sahelanthropus tchadensis, ca. 7 Ma [2]; S.
tchadensis is sometimes claimed to be the last common ancestor of humans and
chimpanzees, but this is disputed. The earliest known human ancestor post-dating

the separation of the human and the chimpanzee lines is Orrorin tugenensis
(Millennium Man, Kenya; ca. 6 Ma). Both chimpanzees and humans have a
larynx that repositions during the first two years of life to a spot between the
pharynx and the lungs, indicating that the common ancestors have this feature, a
precursor of speech.
4.4 Ma Ardipithecus is a very early hominin genus (subfamily Homininae). Two species
are described in the literature: A. ramidus, which lived about 4.4 million years
ago[13] during the early Pliocene, and A. kadabba, dated to approximately 5.6
million years ago[14] (late Miocene). A. ramidus had a small brain, measuring
between 300 and 350 cm3. This is about the same size as modern bonobo and
female common chimpanzee brain, but much smaller than the brain of
australopithecines like Lucy (~400 to 550 cm3) and slightly over a fifth the size of
the modern Homo sapiens brain. Ardipithecus was aboreal, meaning it lived
largely in the forest where it competed with other forest animals for food,
including the contemporary ancestor for the chimpanzees. Ardipithecus was likely
bipedal as evidenced by its bowl shaped pelvis and centered foramen magnum,
though its feet were still adapted for grasping rather than walking for long
distances.
3.6 Ma

Australopithecus afarensis
Some Australopithecus afarensis left human-like footprints on volcanic ash in
Laetoli, Kenya (Northern Tanzania) which provides strong evidence of full-time
bipedalism. Australopithecus afarensis lived between 3.9 and 2.9 million years
ago. It is thought that A. afarensis was ancestral to both the genus
Australopithecus and the genus Homo. Compared to the modern and extinct great
apes, A. afarensis has reduced canines and molars, although they are still
relatively larger than in modern humans. A. afarensis also has a relatively small
brain size (~380-430cm) and a prognathic (i.e. projecting anteriorly) face.
Australopithecines have been found in Savannah environments and likely
increased its diet to include meat from scavenging opportunities. An analysis of
Australopithecus africanus lower vertebrae suggests that females had changes to
support bipedalism even while pregnant.

3.5 Ma Kenyanthropus platyops, a possible ancestor of Homo, emerges from the

Australopithecus genus.
3 Ma The bipedal australopithecines (a genus of the Hominina subtribe) evolve in the
savannas of Africa being hunted by Dinofelis. Loss of body hair takes place in the
period 3-2 Ma, in parallel with the development of full bipedalism.

[edit] Homo
Date

Event

2.5 Ma

Homo habilis
Appearance of Homo. Homo habilis is thought to be the ancestor of the lankier
and more sophisticated Homo ergaster. Lived side by side with Homo erectus
until at least 1.44 Ma, making it highly unlikely that Homo erectus directly
evolved out of Homo habilis. First stone tools, beginning of the Lower Paleolithic.
Further information: Homo rudolfensis
1.8 Ma

A reconstruction of Homo erectus.

Homo erectus evolves in Africa. Homo erectus would bear a striking resemblance
to modern humans, but had a brain about 74 percent of the size of modern man. Its
forehead is less sloping and the teeth are smaller. Other hominid designations such
as Homo georgicus, Homo ergaster, Homo pekinensis, Homo heidelbergensis are
often put under the umbrella species name of Homo erectus[15]. Starting with
Homo georgicus found in what is now the Republic of Georgia dated at 1.8 Ma,
the pelvis and backbone grew more human-like and gave georgicus the ability
cover very long distances in order to follow herds of other animals. This is the
oldest fossil of a hominid found (so far) outside of Africa. Control of fire by early
humans is achieved 1.5 Ma by Homo ergaster. Homo ergaster reaches a height of
around 1.9 metres (6.2 ft). Evolution of dark skin, which is linked to the loss of
body hair in human ancestors, is complete by 1.2 Ma. Homo pekinensis first
appears in Asia around 700 Ka but according to the theory of a recent African
origin of modern humans, they could not be human ancestors, but rather, were just
a cousin offshoot species from Homo ergaster. Homo heidelbergensis was a very
large hominid that had a more advanced complement of cutting tools and may
have hunted big game such as horses.
516 ka Homo antecessor is the common genetic ancestor of humans and Neanderthal.[16]
At present estimate, humans have approximately 20,00025,000 genes and share
99% of their DNA with the now extinct Neanderthal [17] and 95-99% of their DNA
with their closest living evolutionary relative, the chimpanzees[18][19]. The human
variant of the FOXP2 gene (linked to the control of speech) has been found to be
identical in Neanderthal[20]. It can therefore be deduced that Homo antecessor
would also have had the human FOXP2 gene.
355 ka

A reconstruction of Homo heidelbergensis

Three 1.5 m (5 ft) tall Homo heidelbergensis left footprints in powdery volcanic
ash solidified in Italy. Homo heidelbergensis is the common ancestor of both
Homo neanderthalensis and Homo sapiens. It is morphologically very similar to
Homo erectus but Homo heidelbergensis had a larger brain-case, about 93% the
size of that of Homo sapiens. The species was tall, 1.8 m (6 ft) on average, and
more muscular than modern humans. Beginning of the Middle Paleolithic.

195 ka

Homo sapiens sapiens (Pioneer plaque)

Omo1, Omo2 (Ethiopia, Omo river) are the earliest fossil evidence for archaic
Homo sapiens, evolved from Homo heidelbergensis.
160 ka Homo sapiens (Homo sapiens idaltu) in Ethiopia, Awash River, Herto village,
practice mortuary rituals and butcher hippos. Potential earliest evidence of
behavioral modernity consistent with the continuity hypothesis including use of
red ochre and fishing[21].
150 ka Mitochondrial Eve is a woman that lived in East Africa. She is the statistically
expected most recent female ancestor common to all mitochondrial lineages in
humans alive today. Note that there is no evidence of any characteristic or genetic
drift that significantly differentiated her from the contemporary social group she
lived with at the time. Her ancestors were homo sapiens and her mother had the
same mtDNA.
70 ka Appearance of mitochondrial haplogroup L2. Behavioral modernity according to
the "great leap forward" theory[22].
60 ka Y-chromosomal Adam lives in Africa. He is the most recent common ancestor
from whom all male human Y chromosomes are descended. Appearance of
mitochondrial haplogroups M and N, which participate in the migration out of
Africa.
50 ka Migration to South Asia. M168 mutation (carried by all non-African males).
Beginning of the Upper Paleolithic. mt-haplogroups U, K.
40 ka Migration to Australia and Europe (Cro-Magnon).
25 ka Neanderthals die out. Y-Haplogroup R2; mt-haplogroups J, X.
12 ka Beginning of the Mesolithic / Holocene. Y-Haplogroup R1a; mt-haplogroups V, T.
Evolution of light skin in Europeans (SLC24A5). Homo floresiensis dies out,
leaving Homo sapiens as the only living species of the genus Homo.

History of the world

From Wikipedia, the free encyclopedia

Jump to: navigation, search

For the history of Earth which includes the time before human existence, see History of
the Earth. For the field of historical study that examines history from a global
perspective, see World History. For other uses, see History of the world
(disambiguation).
Not to be confused with recorded history.
The history of the world is the recorded memory of the experience of Homo sapiens.
Ancient human history[1] begins with the invention, independently at several sites on
Earth, of writing, which created the infrastructure for lasting, accurately transmitted
memories and thus for the diffusion and growth of knowledge.[2][3] Nevertheless, an
appreciation of the roots of civilization requires at least cursory consideration to
humanity's prehistory.
During the Agricultural Revolution between 8,500 and 7,000 BCE in the Fertile Crescent
humans began the systematic husbandry of plants and animals agriculture.[4][5][6] It
spread to neighboring regions, and also developed independently elsewhere, until most
humnas lived sedentary lives as farmers in permanent settlements[7] centered about lifesustaining bodies of water. The relative security and increased productivity provided by
farming allowed these communities to expand.. They grew over time into increasingly
larger units in parallel with the evolution of ever more efficient means of transport.
Surplus food made possible an increasing division of labor, the rise of a leisured upper
class, and the development of cities and thus of civilization. The growing complexity of
human societies necessitated systems of accounting. Beginning in the Bronze Age this led
to writing.[8]
Civilizations developed on the banks of rivers. By 3,000 BCE they had arisen in the
Middle East's Mesopotamia (the "land between the Rivers" Euphrates and Tigris),[9] on
the banks of Egypt's River Nile,[10][11][12] in India's Indus River valley,[13][14][15] and along the
great rivers of China.
The history of the Old World is commonly divided into Antiquity (in the ancient Near
East,[16][17][18] the Mediterranean basin of classical antiquity, ancient China,[19] and ancient
India, up to about the 6th century); the Middle Ages,[20][21] from the 6th through the 15th
centuries; the Early Modern period,[22] including the European Renaissance, from the 16th
century to about 1750; and the Modern period, from the Age of Enlightenment and the
Industrial Revolution, beginning about 1750, to the present. In Europe, the fall of the
Western Roman Empire (476 CE) is commonly taken as signaling the end of antiquity
and the beginning of the Middle Ages.

A thousand years later, in the mid-15th century, Johannes Gutenberg's invention of

modern printing,[23] employing movable type, revolutionized communication, helping end
the Middle Ages and usher in modern times, the European Renaissance[24][25] and the
Scientific Revolution.[26]
By the 18th century, the accumulation of knowledge and technology, especially in
Europe, had reached a critical mass that brought about the Industrial Revolution.[27] Over
the quarter-millennium since, the growth of knowledge, technology, commerce, and of
the potential destructiveness of war has accelerated, creating the opportunities and perils
that now confront the human communities that inhabit the planet.[28][29]

Contents
[hide]

1 Prehistory
2 Ancient history
o 2.1 Origin of civilization
3 Ancient empires
o 3.1 Religion and philosophy
o 3.2 Regional empires
o 3.3 Declines and falls
4 Middle Ages
5 Modern history
o 5.1 Early Modern period
5.1.1 Rise of Europe
5.1.2 Age of Discovery
o 5.2 19th century
o 5.3 20th century to present
5.3.1 Early 20th century
5.3.2 Late 20th century
5.3.3 21st century
6 See also
o 6.1 History topics
o 6.2 History by period
o 6.3 History by region
7 Notes
8 References

9 Further reading

[edit] Prehistory
Main article: Paleolithic

Homo sapiens first arose in the Earth between 400 and 250 thousand years ago during the
Palaeolithic period. This occurred after a long period of evolution. Ancestors of humans,
such as Homo erectus, had been using simple tools for many millennia, but as time
progressed, tools became far more refined and complex. At some point, humans had
begun using fire for heat and for cooking. Humans also developed language sometime
during the Paleolithic, as well as a conceptual repertoire that included systematic burial of
the dead and adornment of the living. During this period, all humans lived as huntergatherers, who were generally nomadic.
Modern humans spread rapidly from Africa and the frost-free zones of Europe and Asia.
The rapid expansion of humankind to North America and Oceania took place at the
climax of the most recent Ice Age, when temperate regions of today were extremely
inhospitable. Yet, humans had colonised nearly all the ice-free parts of the globe by the
end of the Ice Age, some 12,000 years ago.
The Agricultural Revolution, beginning about 10,000 BCE, saw the development of
agriculture. Farming permitted far denser populations, which in time organised into
states. Agriculture also created food surpluses that could support people not directly
engaged in food production. The development of agriculture permitted the creation of the
first cities. These were centres of trade, manufacture and political power with nearly no
agricultural production of their own. Cities established a symbiosis with their surrounding
countrysides, absorbing agricultural products and providing, in return, manufactures and
varying degrees of military control and protection.[30][31][32]

Cuneiform script, the earliest known writing system

The development of cities equated, both etymologically and in fact, with the rise of
civilization itself: first Sumerian civilization, in lower Mesopotamia (3500 BCE),[33][34]
followed by Egyptian civilization along the Nile (3300 BCE)[12] and Harappan civilization
in the Indus Valley (3300 BCE).[35][36] Elaborate cities grew up, with high levels of social
and economic complexity. Each of these civilizations was so different from the others
that they almost certainly originated independently. It was at this time, and due to the
needs of cities, that writing and extensive trade were introduced.

This period also saw the origins of complex religion.[37][38][39] Religious belief in this
period commonly consisted in the worship of a Mother Goddess, a Sky Father, and of the
Sun and Moon as deities.[40] (See also: Sun worship.) Shrines developed, which over time
evolved into temple establishments, complete with a complex hierarchy of priests and
priestesses and other functionaries. Typical of the Neolithic was a tendency to worship
anthropomorphic deities. Some of the earliest surviving written religious scriptures are
the Pyramid Texts, produced by the Egyptians, the oldest of which date to between 2400
and 2300 BCE.[41] Some archeologists suggest, based on ongoing excavations of a temple
complex at Gbekli Tepe ("Potbelly Hill") in southern Turkey, dating from ca. 11,500
years ago, that religion predated the Agricultural Revolution rather than following in its
wake, as had generally been assumed.[42]

[edit] Ancient history

Main article: Ancient history

[edit] Origin of civilization

Main article: Bronze Age

Ancient Egyptians built the Great Pyramids of Giza.

The Bronze Age forms part of a three-age system. In this system, in some areas of the
world, the Bronze Age follows the Stone Age. During this era the most fertile areas of the
world saw city states and the first civilizations develop. These were concentrated on four
fertile river valleys: The Tigris and Euphrates in Mesopotamia, the Nile in Egypt, the
Indus in South Asia, and the Yangtze and Yellow River in China.
Mesopotamia saw the rise of the Sumerian city states. It was in these cities that the
earliest known form of writing, cuneiform script, appeared ca. 3000 BCE. Cuneiform
writing began as a system of pictographs. Over time, the pictorial representations became
simplified and more abstract. Cuneiforms were written on clay tablets, on which symbols
were drawn with a blunt reed for a stylus. Writing made the administration of a large state
far easier. This era also saw new military technologies, such as chariots, that allowed
armies to move faster.
These developments lead to the development of empires. The first empire, controlling a
large territory and many cities, developed in Egypt that formed with the unification of

Lower and Upper Egypt c. 3100 BCE . Over the next millennia the other river valleys
would also see monarchical empires rise to power. In the 24th century BCE, the
Akkadian Empire arose in Mesopotamia.[43] and in China the Xia Dynasty arose c. 2200
BCE.
Over the next millennia civilizations would develop across the world. Trade would
increasingly become a source of power as states with access to important resources or
controlling important trade routes would rise to dominance. In c.2,500 BCE the Kingdom
of Kerma developed in Sudan south of Egpyt. In modern Turkey the Hittites controlled a
large empire and by 1600 BCE, Mycenaean Greece begins to develop.[44][45] In India this
era was the Vedic period, which laid the foundations of Hinduism and other cultural
aspects of early Indian society, and ended in the 500s BCE. From around 550 BCE, many
independent kingdoms and republics known as the Mahajanapadas were established
across the country. In the Americas, civilizations such as the Maya, Zapotec, Moche, and
Nazca emerged in Mesoamerica and Peru at the end of the 1st millennium BCE.

[edit] Ancient empires

[edit] Religion and philosophy
Main article: Axial age
Main articles: History of philosophy, Timeline of religion, and History of religion

Angkor Wat temple, Cambodia, early 12th century

Beginning in the sixth century BCE a set of transformative religious and philosophical
ideas developed. During this century Chinese Confucianism, Indian Buddhism and
Jainism, Jewish Monotheism, and Persian Zoroastrianism all developed. In the fifth
century Socrates and Plato would lay the foundations of Ancient Greek philosophy.
In the east, three schools of thought were to dominate Chinese thinking until the modern
day. These were Taoism,[46] Legalism[47] and Confucianism.[48] The Confucian tradition,
which would attain dominance, looked for political morality not to the force of law but to
the power and example of tradition. Confucianism would later spread into the Korean
peninsula and toward Japan.

In the west, the Greek philosophical tradition, represented by Socrates,[49] Plato,[50] and
Aristotle,[51][52] was diffused throughout Europe and the Middle East in the 4th century
BCE by the conquests of Alexander III of Macedon, more commonly known as
Alexander the Great.[53][54][55]

[edit] Regional empires

Main articles: Civilization and Global empire

The Parthenon epitomizes the sophisticated culture of the ancient Greeks.

The millennia from 500 BCE to 500 CE saw a series of empires of unprecedented size
developed. Well trained professional armies, unifying ideologies, and advanced
bureaucracies allowed a single emperor to rule over tens of millions of subjects.
This period in the history of the world was marked by slow but steady technological
advances, with important developments such as the stirrup and moldboard plow arriving
every few centuries. There were, however, in some regions, periods of rapid
technological progress. Most important, perhaps, was the Mediterranean area during the
Hellenistic period, when hundreds of technologies were invented.[56][57][58] Such periods
were followed by periods of technological decay, as during the Roman Empire's decline
and fall and the ensuing early medieval period.
The great empires depended on military annexation of territory and on the formation of
defended settlements to become agricultural centres.[59] The relative peace that the
empires brought encouraged international trade, most notably the massive trade routes in
the Mediterranean that had been developed by the time of the Hellenistic Age, and the
Silk Road.

Ptolemy's world map, reconstituted from his Geographia (ca. 150)

The empires faced common problems associated with maintaining huge armies and
supporting a central bureaucracy. These costs fell most heavily on the peasantry, while
land-owning magnates were increasingly able to evade centralised control and its costs.
The pressure of barbarians on the frontiers hastened the process of internal dissolution.
China's Han Empire fell into civil war in 220 CE, while its Roman counterpart became
increasingly decentralised and divided about the same time.
In the west, the Greeks established a civilization that is the foundational culture of
modern western civilization. Some centuries later, in the 3rd century BCE, the Romans
began expanding their territory through conquest and colonization. By the reign of
Emperor Augustus (late 1st century BCE), Rome controlled all the lands surrounding the
Mediterranean. By the reign of Emperor Trajan (early 2nd century CE), Rome controlled
much of the land from England to Mesopotamia.
In the third century BCE, most of South Asia was united into the Maurya Empire by
Chandragupta Maurya and flourished under Ashoka the Great. From the third century
CE, the Gupta dynasty oversaw the period referred to as ancient India's Golden Age.
Empires in Southern India included those of the Chalukyas,the Rashtrakutas, the
Hoysalas, the Cholas and the Vijayanagara Empire. Science, engineering, art, literature,
astronomy, and philosophy flourished under the patronage of these kings.

[edit] Declines and falls

The great empires of Eurasia were all located on the temperate coastal plains. From the
Central Asian steppe horse based nomads dominated a large part of the continent. The
development of the stirrup and the breeding of horses strong enough to carry a fully
armed archer made the nomads a constant threat to the more settled civilizations.
The gradual break-up of the Roman Empire,[60][61] spanning several centuries after the 2nd
century CE, coincided with the spread of Christianity westward from the Middle East.
The western Roman Empire fell[62] under the domination of Germanic tribes in the 5th
century, and these polities gradually developed into a number of warring states, all
associated in one way or another with the Roman Catholic Church. The remaining part of
the Roman Empire, in the eastern Mediterranean, would henceforth be the Byzantine
Empire.[63] Centuries later, a limited unity would be restored to western Europe through
the establishment of the Holy Roman Empire[64] in 962, which comprised a number of
states in what is now Germany, Switzerland, Belgium, Italy, and France.
In China, dynasties would similarly rise and fall.[65][66] After the fall of the Eastern Han
Dynasty[67] and the demise of the Three Kingdoms, Nomadic tribes from the north began
to invade in the 4th century CE, eventually conquering areas of Northern China and
setting up many small kingdoms. The Sui Dynasty reunified China in 581, and under the
succeeding Tang Dynasty (618-907) China entered a second golden age. The Tang

Dynasty also splintered, however, and after half a century of turmoil the Northern Song
Dynasty reunified China in 982. Yet pressure from nomadic empires to the north became
increasingly urgent. North China was lost to the Jurchens in 1141, and the Mongol
Empire[68][69] conquered all of China in 1279, as well as almost all of Eurasia's landmass,
missing only central and western Europe, and most of Southeast Asia and Japan.
In these times, northern India was ruled by the Guptas. In southern India, three prominent
Dravidian kingdoms emerged: Cheras, Cholas and Pandyas. The ensuing stability
contributed to heralding in the golden age of Hindu culture in the 4th and 5th centuries
CE.

Machu Picchu, "the Lost City of the Incas"the most recognizable symbol of Inca
civilization
Also at this time in Central America,[70] vast societies began to be built, the most notable
being the Maya and Aztecs of Mesoamerica. As the mother culture of the Olmecs[71]
gradually declined, the great Mayan city-states slowly rose in number and prominence,
and Maya culture spread throughout Yucatn and surrounding areas. The later empire of
the Aztecs was built on neighboring cultures and was influenced by conquered peoples
such as the Toltecs.
In South America, the 14th and 15th centuries saw the rise of the Inca. The Inca Empire
of Tawantinsuyu, with its capital at Cusco, spanned the entire Andes Mountain Range.[72]
[73]
The Inca were prosperous and advanced, known for an excellent road system and
unrivaled masonry.

[edit] Middle Ages

Main article: Middle Ages
The Middle Ages are commonly dated from the fall of the Western Roman Empire in the
5th century.

The period corresponds to the Islamic conquests[74], subsequent Islamic golden age[75][76],
and commencement and expansion of the Islamic/Arab Slave Trade followed by the
Mongol invasions in the Middle East and Central Asia. South Asia saw a series of middle
kingdoms of India followed by the establishment of Islamic empires in India. The
Chinese Empire saw the succession of the Sui, Tang, Liao, Yuan and Ming Dynasties.
The Black Death was one of the deadliest pandemics in human history. Starting in Asia,
the disease reached Mediterranean and western Europe during the late 1340s,[77] and
killed tens of millions of Europeans in six years; between a third and a half of the total
population.[78]
The Middle Ages[79] witnessed the first sustained urbanization of northern and western
Europe. Many modern European states owe their origins to events unfolding in the
Middle Ages; present European political boundaries are, in many regards, the result of the
military and dynastic achievements during this tumultuous period.[80]
The Middle Ages lasted until the beginning of the Early Modern Period[22] in the 16th
century, marked by the rise of nation-states, the division of Western Christianity in the
Reformation,[81] the rise of humanism in the Italian Renaissance,[82] and the beginnings of
European overseas expansion which allowed for the Columbian Exchange.[83]

[edit] Modern history

Main article: Modern history
Modern history (the "modern period," the "modern era," "modern times") is history of the
period following the Middle Ages. "Contemporary history" encompasses historic events
that are immediately relevant to the present time; its intentionally loose ambit includes
major events such as World War II, but not those whose immediate effects have
dissipated.

[edit] Early Modern period

Main article: Early Modern period
"Early modern period"[84] is a term used by historians to refer to the period in Western
Europe and its first colonies that spans the centuries between the Middle Ages and the
Industrial Revolution. The early modern period is characterized by the rise to importance
of science and by increasingly rapid technological progress, secularized civic politics,
and the nation-state. Capitalist economies began their rise, initially in northern Italian
republics such as Genoa. The early modern period also saw the rise and dominance of the
mercantilist economic theory. As such, the early modern period represents the decline and
eventual disappearance, in much of the European sphere, of feudalism, serfdom and the
power of the Catholic Church. The period includes the Protestant Reformation, the
disastrous Thirty Years' War, the European colonization of the Americas, and the peak of
European witch-hunting.

[edit] Rise of Europe

Further information: History of Europe

The movable-type printing press arose in the mid-15th century. Less than 50 years later,
nine million books were in print.
Nearly all the agricultural civilizations have been heavily constrained by their
environments. Productivity remained low, and climatic changes easily instigated boom
and bust cycles that brought about civilizations' rise and fall. By about 1500, however,
there was a qualitative change in world history. Technological advance and the wealth
generated by trade gradually brought about a widening of possibilities.[85][86][87][88][89][90][91]
[92][93][94][95]

Outwardly, Europe's Renaissance, beginning in the 14th century,[96] consisted in the

rediscovery of the classical world's scientific contributions, and in the economic and
social rise of Europe. But the Renaissance also engendered a culture of inquisitiveness
which ultimately led to Humanism,[97] the Scientific Revolution,[98] and finally the great
transformation of the Industrial Revolution. The Scientific Revolution in the 17th century,
however, had no immediate impact on technology; only in the second half of the 18th
century did scientific advances begin to be applied to practical invention.
The advantages that Europe had developed by the mid-18th century were two: an
entrepreneurial culture,[citation needed] and the wealth generated by the Atlantic trade
(including the African slave trade). By the late 16th century, American silver accounted
for one-fifth of Spain's total budget.[citation needed] The profits of the slave trade and of West
Indian plantations amounted to 5% of the British economy at the time of the Industrial
Revolution.[99] While some historians conclude that, in 1750, labour productivity in the
most developed regions of China was still on a par with that of Europe's Atlantic
economy (see the NBER Publications by Carol H. Shiue and Wolfgang Keller[100]), other
historians like Angus Maddison hold that the per-capita productivity of western Europe
had by the late Middle Ages surpassed that of all other regions.[101]

A number of explanations are proffered as to why, from the late Middle Ages on, Europe
rose to surpass other civilizations, become the home of the Industrial Revolution,[102] and
dominate the world. Max Weber argued that it was due to a Protestant work ethic that
encouraged Europeans to work harder and longer than others. Another socioeconomic
explanation looks to demographics: Europe, with its celibate clergy, colonial emigration,
high-mortality urban centers, periodic famines and outbreaks of the Black Death,
continual warfare, and late age of marriage had far more restrained population growth,
compared to Asian cultures. A relative shortage of labour meant that surpluses could be
invested in labour-saving technological advances such as water-wheels and mills,
spinners and looms, steam engines and shipping, rather than fueling population growth.
Many have also argued that Europe's institutions were superior,[103][104] that property rights
and free-market economics were stronger than elsewhere due to an ideal of freedom
peculiar to Europe. In recent years, however, scholars such as Kenneth Pomeranz have
challenged this view, although the revisionist approach to world history has also met with
criticism for systematically "downplaying" European achievements.[105]

Vasco da Gama reached India by sea in 1498.

Europe's geography may also have played an important role. The Middle East, India and
China are all ringed by mountains but, once past these outer barriers, are relatively flat.
By contrast, the Pyrenees, Alps, Apennines, Carpathians and other mountain ranges run

through Europe, and the continent is also divided by several seas. This gave Europe some
degree of protection from the peril of Central Asian invaders. Before the era of firearms,
these nomads were militarily superior to the agricultural states on the periphery of the
Eurasian continent and, if they broke out into the plains of northern India or the valleys of
China, were all but unstoppable. These invasions were often devastating. The Golden Age
of Islam[106] was ended by the Mongol sack of Baghdad in 1258. India and China were
subject to periodic invasions, and Russia spent a couple of centuries under the MongolTatar Yoke. Central and western Europe, logistically more distant from the Central Asian
heartland, proved less vulnerable to these threats.
Geography also contributed to important geopolitical differences. For most of their
histories, China, India and the Middle East were each unified under a single dominant
power that expanded until it reached the surrounding mountains and deserts. In 1600 the
Ottoman Empire[107] controlled almost all the Middle East, the Ming Dynasty ruled China,
[108][109]
and the Mughal Empire held sway over India. By contrast, Europe was almost
always divided into a number of warring states. Pan-European empires, with the notable
exception of the Roman Empire, tended to collapse soon after they arose. Another
doubtless important geographic factor in the rise of Europe was the Mediterranean Sea,
which, for millennia, had functioned as a maritime superhighway fostering the exchange
of goods, people, ideas and inventions.
[edit] Age of Discovery
Main article: Age of Discovery

Columbus sought India aboard the Santa Maria in 1492.

In the fourteenth century, the Renaissance began in Europe.[110][111] Some modern
scholars[who?] have questioned whether this flowering of art and Humanism was a benefit
to science. The era did see an important fusion of Arab and European knowledge.[112][113]
One of the most important developments was the caravel, which combined the
Mediterranean lateen sail with European square rigging to create the first vessels that

could safely sail the Atlantic Ocean.[114] Along with important developments in
navigation, this technology allowed the Italian Christopher Columbus in 1492 to journey
across the Atlantic Ocean and bridge the gap between Afro-Eurasia and the Americas.
This had dramatic effects on both continents. The Europeans brought with them viral
diseases that American natives had never encountered,[115] and uncertain numbers of
natives died in a series of devastating epidemics. The Europeans also had the
technological advantage of horses, steel and guns that helped them overpower the Aztec
and Incan empires as well as North American cultures.[116]
Gold and resources from the Americas began to be stripped from the land and people and
shipped to Europe, while at the same time large numbers of European colonists began to
emigrate to the Americas.[117][118] To meet the great demand for labor in the new colonies,
the mass import of Africans as slaves began.[119] Soon much of the Americas had a large
racial underclass of slaves. In West Africa, a series of thriving states developed along the
coast, becoming prosperous from the exploitation of suffering interior African peoples.
Europe's maritime expansion unsurprisingly given that continent's geography was
largely the work of its Atlantic states: Portugal, Spain, England, France, and the
Netherlands. The Portuguese and Spanish Empires were the predominant conquerors and
source of influence, and their union resulted in the Iberian Union,[120] the first global
empire, on which the "sun never set". Soon the more northern English, French and Dutch
began to dominate the Atlantic. In a series of wars fought in the 17th and 18th centuries,
culminating with the Napoleonic Wars, Britain emerged as the new world power.
Meanwhile the voyages of Admiral Zheng He were halted by China's Ming Dynasty
(13681644), established after the expulsion of the Mongols. A Chinese commercial
revolution, sometimes described as "incipient capitalism", was also abortive. The Ming
Dynasty would eventually fall to the Manchus, whose Qing Dynasty at first oversaw a
period of calm and prosperity but would increasingly fall prey to Western encroachment.

[edit] 19th century

Main article: 19th century
After Europeans had achieved dominance over the Americas, their imperial appetites
turned to the countries of Asia. In the 19th century the European states had a distinct
technological advantage over Asian states and peoples.[citation needed] Britain gained control
of the Indian subcontinent, Egypt and the Malay Peninsula;[citation needed] the French took
Indochina;[citation needed] while the Dutch cemented their control over the Dutch East Indies.
[citation needed]
In addition, Russia colonised large pre-agricultural areas of Siberia.[citation needed]
The British also colonised places inhabited by Neolithic peoples, including Australia,
New Zealand and South Africa. large numbers of British colonists emigrated to these
colonies.[citation needed] In the late 19th century, the European powers divided the remaining
areas of Africa.[citation needed] Within Europe, economic and military challenges created a
system of nation states, and ethno-linguistic groupings began to identify themselves as

distinctive nations with aspirations for cultural and political autonomy.[citation needed] This
nationalism would become important to peoples across the world in the twentieth century.
[citation needed]

This era in European culture saw the Age of Reason lead to the Scientific Revolution.
The Scientific Revolution changed humanity's understanding of the world and happened
simultaneously with the Industrial Revolution, a major transformation of the world's
economies.[citation needed] The Industrial Revolution began in Great Britain[citation needed] and used
new modes of production the factory, mass production, and mechanisation to
manufacture a wide array of goods faster and using less labour than previously.[citation needed]
The Age of Reason also led to the beginnings of modern democracy in the late-18th
century American and French Revolutions.[citation needed] Democracy would grow to have a
profound effect on world events and on quality of life.[dubious discuss][neutrality is disputed]
During the Industrial Revolution, the world economy became reliant on coal as a
fuel[citation needed], as new methods of transport, such as railways and steamships, effectively
shrank the world.[citation needed] Meanwhile, industrial pollution and environmental damage,
present since the discovery of fire and the beginning of civilization, accelerated
drastically.

[edit] 20th century to present

Main articles: 20th century and 21st century
[edit] Early 20th century

World War I, fought by the Allies (green) and Central Powers (orange), ended the
German, Austro-Hungarian, Russian and Ottoman Empires.
The 20th century[121][122][123] opened with Europe at an apex of wealth and power, and with
much of the world under its direct colonial control or its indirect domination.[124] Much of
the rest of the world was influenced by heavily Europeanized nations: the United States
and Japan.[125] As the century unfolded, however, the global system dominated by rival
powers was subjected to severe strains, and ultimately yielded to a more fluid structure of
independent nations organized on Western models.
This transformation was catalysed by wars of unparalleled scope and devastation. World
War I[126] destroyed many of Europe's empires and monarchies, and weakened Britain and
France.[127] In its aftermath, powerful ideologies arose. The Russian Revolution[128][129][130]

of 1917 created the first communist state, while the 1920s and 1930s saw militaristic
fascist dictatorships gain control in Italy, Germany, Spain and elsewhere.[131]
Ongoing national rivalries, exacerbated by the economic turmoil of the Great Depression,
helped precipitate World War II.[132][133] The militaristic dictatorships of Europe and Japan
pursued an ultimately doomed course of imperialist expansionism. Their defeat opened
the way for the advance of communism into Central Europe, Yugoslavia, Bulgaria,
Romania, Albania, China, North Vietnam and North Korea.

Nuclear bombs, dropped on Japan in 1945, ended World War II and opened the Cold War.
Following World War II, in 1945, the United Nations was founded in the hope of allaying
conflicts among nations and preventing future wars.[134][135] The war had, however, left
two nations, the United States[136] and the Soviet Union, with principal power to guide
international affairs.[137] Each was suspicious of the other and feared a global spread of the
other's political-economic model. This led to the Cold War, a forty-year stand-off
between the United States, the Soviet Union, and their respective allies. With the
development of nuclear weapons[138] and the subsequent arms race, all of humanity were
put at risk of nuclear war between the two superpowers.[139] Such war being viewed as
impractical, proxy wars were instead waged, at the expense of non-nuclear-armed Third
World countries.
[edit] Late 20th century
The Cold War lasted through to the ninth decade of the twentieth century, when the
Soviet Union's communist system began to collapse, unable to compete economically
with the United States and western Europe; the Soviets' Central European "satellites"
reasserted their national sovereignty, and in 1991 the Soviet Union itself disintegrated.[140]
[141][142]
The United States for the time being was left as the "sole remaining superpower".
[143][144][145]

In the early postwar decades, the African and Asian colonies of the Belgian, British,
Dutch, French and other west European empires won their formal independence.[146][147]
These nations faced challenges in the form of neocolonialism, poverty, illiteracy and
endemic tropical diseases.[148][149] Many of the Western and Central European nations
gradually formed a political and economic community, the European Union, which
subsequently expanded eastward to include former Soviet satellites.[150][151][152][153]

Last Moon landing Apollo 17 (1972)

The twentieth century saw exponential progress in science and technology, and increased
life expectancy and standard of living for much of humanity. As the developed world
shifted from a coal-based to a petroleum-based economy, new transport technologies,
along with the dawn of the Information Age,[154] led to increased globalization.[155][156][157]
Space exploration reached throughout the solar system. The structure of DNA, the very
template of life, was discovered,[158][159][160] and the human genome was sequenced, a
major milestone in the understanding of human biology and the treatment of disease.[161]
[162][163][164][165]
Global literacy rates continued to rise, and the percentage of the world's
labor pool needed to produce humankind's food supply continued to drop.
The century saw the development of new global threats, such as nuclear proliferation,
worldwide epidemics of diseases, global climate change,[166][167] massive deforestation,
overpopulation, and the dwindling of global resources (particularly fossil fuels).[168]
[edit] 21st century
Main article: Contemporary history
As the 20th century yielded to the 21st, worldwide demand and competition for resources
rose due to growing populations and industrialization, with resulting increased levels of
environmental degradation.[169] This led to development of alternate sources of energy
such as solar and other renewable energy varieties, to proposals for cleaner fossil-fuel
technologies, and to consideration of expanded use of nuclear energy.[170][171][172]

[edit] See also