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Evolution

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This article is about evolution in biology. For related articles, see Outline of
evolution. For other uses, see Evolution (disambiguation).
For a more accessible and less technical introduction to this topic, see Introdu
ction to evolution.
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Evolutionary biology
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Evolution is change in the heritable characteristics of biological populations o
ver successive generations.[1][2] Evolutionary processes give rise to biodiversi
ty at every level of biological organisation, including the levels of species, i
ndividual organisms, and molecules.[3]
All life on Earth shares a common ancestor known as the last universal common an
cestor (LUCA),[4][5][6] which lived approximately 3.53.8 billion years ago,[7] al
though a study in 2015 found "remains of biotic life" from 4.1 billion years ago
in ancient rocks in Western Australia.[8][9] In July 2016, scientists reported
identifying a set of 355 genes from the LUCA of all organisms living on Earth.[1
0]
Repeated formation of new species (speciation), change within species (anagenesi
s), and loss of species (extinction) throughout the evolutionary history of life
on Earth are demonstrated by shared sets of morphological and biochemical trait
s, including shared DNA sequences.[11] These shared traits are more similar amon
g species that share a more recent common ancestor, and can be used to reconstru
ct a biological "tree of life" based on evolutionary relationships (phylogenetic
s), using both existing species and fossils. The fossil record includes a progre
ssion from early biogenic graphite,[12] to microbial mat fossils,[13][14][15] to
fossilized multicellular organisms. Existing patterns of biodiversity have been
shaped both by speciation and by extinction.[16] More than 99 percent of all sp
ecies that ever lived on Earth are estimated to be extinct.[17][18] Estimates of
Earth's current species range from 10 to 14 million,[19][20] of which about 1.9
million are estimated to have been named[21] and 1.6 million documented in a ce
ntral database to date.[22] More recently, in May 2016, scientists reported that
1 trillion species are estimated to be on Earth currently with only one-thousan
dth of one percent described.[23]
In the mid-19th century, Charles Darwin formulated the scientific theory of evol
ution by natural selection, published in his book On the Origin of Species (1859
). Evolution by natural selection is a process demonstrated by the observation t
hat more offspring are produced than can possibly survive, along with three fact
s about populations: 1) traits vary among individuals with respect to morphology
, physiology, and behaviour (phenotypic variation), 2) different traits confer d
ifferent rates of survival and reproduction (differential fitness), and 3) trait
s can be passed from generation to generation (heritability of fitness).[24] Thu
s, in successive generations members of a population are replaced by progeny of
parents better adapted to survive and reproduce in the biophysical environment i
n which natural selection takes place. This teleonomy is the quality whereby the
process of natural selection creates and preserves traits that are seemingly fi
tted for the functional roles they perform.[25] The processes by which the chang
es occur, from one generation to another, are called evolutionary processes or m
echanisms.[26] The four most widely recognized evolutionary processes are natura
l selection (including sexual selection), genetic drift, mutation and gene migra
tion.[26] Natural selection and genetic drift sort variation; mutation and gene
migration create variation.[26]
Consequences of selection can include meiotic drive[27] (unequal transmission of
certain alleles), nonrandom mating[28] and genetic hitchhiking. In the early 20
th century the modern evolutionary synthesis integrated classical genetics with
Darwin's theory of evolution by natural selection through the discipline of popu
lation genetics. The importance of natural selection as a cause of evolution was
accepted into other branches of biology. Moreover, previously held notions abou
t evolution, such as orthogenesis, evolutionism, and other beliefs about innate
"progress" within the largest-scale trends in evolution, became obsolete.[29] Sc
ientists continue to study various aspects of evolutionary biology by forming an
d testing hypotheses, constructing mathematical models of theoretical biology an
d biological theories, using observational data, and performing experiments in b
oth the field and the laboratory.
In terms of practical application, an understanding of evolution has been instru
mental to developments in numerous scientific and industrial fields, including a
griculture, human and veterinary medicine, and the life sciences in general.[30]
[31][32] Discoveries in evolutionary biology have made a significant impact not
just in the traditional branches of biology but also in other academic disciplin
es, including biological anthropology, and evolutionary psychology.[33][34] Evol
utionary computation, a sub-field of artificial intelligence, involves the appli
cation of Darwinian principles to problems in computer science.
Contents
1 History of evolutionary thought
1.1 Classical times
1.2 Medieval
1.3 Pre-Darwinian
1.4 Darwinian revolution
1.5 Pangenesis
1.6 The 'modern synthesis'
1.7 Further syntheses
2 Heredity
3 Variation
3.1 Mutation
3.2 Sex and recombination
3.3 Gene flow
4 Mechanisms
4.1 Natural selection
4.2 Biased mutation
4.3 Genetic drift
4.4 Genetic hitchhiking
4.5 Gene flow
5 Outcomes
5.1 Adaptation
5.2 Coevolution
5.3 Cooperation
5.4 Speciation
5.5 Extinction
6 Evolutionary history of life
6.1 Origin of life
6.2 Common descent
6.3 Evolution of life
7 Applications
8 Social and cultural responses
9 See also
10 References
11 Bibliography
12 Further reading
13 External links
History of evolutionary thought

Lucretius

Alfred Russel Wallace

Thomas Robert Malthus

Statistician Ronald Fisher

In 1842, Charles Darwin penned his first sketch of On the Origin of Species.[35]
Main article: History of evolutionary thought
Classical times
The proposal that one type of organism could descend from another type goes back
to some of the first pre-Socratic Greek philosophers, such as Anaximander and E
mpedocles.[36] Such proposals survived into Roman times. The poet and philosophe
r Lucretius followed Empedocles in his masterwork De rerum natura (On the Nature
of Things).[37][38]
Medieval
In contrast to these materialistic views, Aristotelianism considered all natural
things as actualisations of fixed natural possibilities, known as forms.[39][40
] This was part of a medieval teleological understanding of nature in which all
things have an intended role to play in a divine cosmic order. Variations of thi
s idea became the standard understanding of the Middle Ages and were integrated
into Christian learning, but Aristotle did not demand that real types of organis
ms always correspond one-for-one with exact metaphysical forms and specifically
gave examples of how new types of living things could come to be.[41]
Pre-Darwinian
In the 17th century, the new method of modern science rejected the Aristotelian
approach. It sought explanations of natural phenomena in terms of physical laws
that were the same for all visible things and that did not require the existence
of any fixed natural categories or divine cosmic order. However, this new appro
ach was slow to take root in the biological sciences, the last bastion of the co
ncept of fixed natural types. John Ray applied one of the previously more genera
l terms for fixed natural types, "species," to plant and animal types, but he st
rictly identified each type of living thing as a species and proposed that each
species could be defined by the features that perpetuated themselves generation
after generation.[42] The biological classification introduced by Carl Linnaeus
in 1735 explicitly recognized the hierarchical nature of species relationships,
but still viewed species as fixed according to a divine plan.[43]
Other naturalists of this time speculated on the evolutionary change of species
over time according to natural laws. In 1751, Pierre Louis Maupertuis wrote of n
atural modifications occurring during reproduction and accumulating over many ge
nerations to produce new species.[44] Georges-Louis Leclerc, Comte de Buffon sug
gested that species could degenerate into different organisms, and Erasmus Darwi
n proposed that all warm-blooded animals could have descended from a single micr
oorganism (or "filament").[45] The first full-fledged evolutionary scheme was Je
an-Baptiste Lamarck's "transmutation" theory of 1809,[46] which envisaged sponta
neous generation continually producing simple forms of life that developed great
er complexity in parallel lineages with an inherent progressive tendency, and po
stulated that on a local level these lineages adapted to the environment by inhe
riting changes caused by their use or disuse in parents.[47][48] (The latter pro
cess was later called Lamarckism.)[47][49][50][51] These ideas were condemned by
established naturalists as speculation lacking empirical support. In particular
, Georges Cuvier insisted that species were unrelated and fixed, their similarit
ies reflecting divine design for functional needs. In the meantime, Ray's ideas
of benevolent design had been developed by William Paley into the Natural Theolo
gy or Evidences of the Existence and Attributes of the Deity (1802), which propo
sed complex adaptations as evidence of divine design and which was admired by Ch
arles Darwin.[52][53][54]
Darwinian revolution
The crucial break from the concept of constant typological classes or types in b
iology came with the theory of evolution through natural selection, which was fo
rmulated by Charles Darwin in terms of variable populations. Partly influenced b
y An Essay on the Principle of Population (1798) by Thomas Robert Malthus, Darwi
n noted that population growth would lead to a "struggle for existence" in which
favorable variations prevailed as others perished. In each generation, many off
spring fail to survive to an age of reproduction because of limited resources. T
his could explain the diversity of plants and animals from a common ancestry thr
ough the working of natural laws in the same way for all types of organism.[55][
56][57][58] Darwin developed his theory of "natural selection" from 1838 onwards
and was writing up his "big book" on the subject when Alfred Russel Wallace sen
t him a version of virtually the same theory in 1858. Their separate papers were
presented together at a 1858 meeting of the Linnean Society of London.[59] At t
he end of 1859, Darwin's publication of his "abstract" as On the Origin of Speci
es explained natural selection in detail and in a way that led to an increasingl
y wide acceptance of concepts of evolution. Thomas Henry Huxley applied Darwin's
ideas to humans, using paleontology and comparative anatomy to provide strong e
vidence that humans and apes shared a common ancestry. Some were disturbed by th
is since it implied that humans did not have a special place in the universe.[60
]
Pangenesis
The mechanisms of reproductive heritability and the origin of new traits remaine
d a mystery. Towards this end, Darwin developed his provisional theory of pangen
esis.[61] In 1865, Gregor Mendel reported that traits were inherited in a predic
table manner through the independent assortment and segregation of elements (lat
er known as genes). Mendel's laws of inheritance eventually supplanted most of D
arwin's pangenesis theory.[62] August Weismann made the important distinction be
tween germ cells that give rise to gametes (such as sperm and egg cells) and the
somatic cells of the body, demonstrating that heredity passes through the germ
line only. Hugo de Vries connected Darwin's pangenesis theory to Weismann's germ
/soma cell distinction and proposed that Darwin's pangenes were concentrated in
the cell nucleus and when expressed they could move into the cytoplasm to change
the cells structure. De Vries was also one of the researchers who made Mendel's
work well-known, believing that Mendelian traits corresponded to the transfer o
f heritable variations along the germline.[63] To explain how new variants origi
nate, de Vries developed a mutation theory that led to a temporary rift between
those who accepted Darwinian evolution and biometricians who allied with de Vrie
s.[48][64][65] In the 1930s, pioneers in the field of population genetics, such
as Ronald Fisher, Sewall Wright and J. B. S. Haldane set the foundations of evol
ution onto a robust statistical philosophy. The false contradiction between Darw
in's theory, genetic mutations, and Mendelian inheritance was thus reconciled.[6
6]
The 'modern synthesis'
In the 1920s and 1930s the so-called modern synthesis connected natural selectio
n, mutation theory, and Mendelian inheritance into a unified theory that applied
generally to any branch of biology. The modern synthesis explained patterns obs
erved across species in populations, through fossil transitions in palaeontology
, and complex cellular mechanisms in developmental biology.[48][67] The publicat
ion of the structure of DNA by James Watson and Francis Crick in 1953 demonstrat
ed a physical mechanism for inheritance.[68] Molecular biology improved our unde
rstanding of the relationship between genotype and phenotype. Advancements were
also made in phylogenetic systematics, mapping the transition of traits into a c
omparative and testable framework through the publication and use of evolutionar
y trees.[69][70] In 1973, evolutionary biologist Theodosius Dobzhansky penned th
at "nothing in biology makes sense except in the light of evolution," because it
has brought to light the relations of what first seemed disjointed facts in nat
ural history into a coherent explanatory body of knowledge that describes and pr
edicts many observable facts about life on this planet.[71]
Further syntheses
Since then, the modern synthesis has been further extended to explain biological
phenomena across the full and integrative scale of the biological hierarchy, fr
om genes to species. This extension, known as evolutionary developmental biology
and informally called "evo-devo," emphasises how changes between generations (e
volution) acts on patterns of change within individual organisms (development).[
72][73][74] Since the beginning of the 21st century and in light of discoveries
made in recent decades, some biologists have argued for an extended evolutionary
synthesis, which would account for the effects of non-genetic inheritance modes
, such as epigenetics, parental effects, ecological and cultural inheritance, an
d evolvability.[75][76]
Heredity
Further information: Introduction to genetics, Genetics, Heredity, and Reaction
norm

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


double helix.
Evolution in organisms occurs through changes in heritable traitsthe inherited ch
aracteristics of an organism. In humans, for example, eye colour is an inherited
characteristic and an individual might inherit the "brown-eye trait" from one o
f their parents.[77] Inherited traits are controlled by genes and the complete s
et of genes within an organism's genome (genetic material) is called its genotyp
e.[78]
The complete set of observable traits that make up the structure and behaviour o
f an organism is called its phenotype. These traits come from the interaction of
its genotype with the environment.[79] As a result, many aspects of an organism
's phenotype are not inherited. For example, suntanned skin comes from the inter
action 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 genotypic variation; a striking example are people with the inher
ited trait of albinism, who do not tan at all and are very sensitive to sunburn.
[80]
Heritable traits are passed from one generation to the next via DNA, a molecule
that encodes genetic information.[78] DNA is a long biopolymer composed of four
types of bases. The sequence of bases along a particular DNA molecule specify th
e 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 resul
ting two cells will inherit the DNA sequence. Portions of a DNA molecule that sp
ecify a single functional unit are called genes; different genes have different
sequences of bases. Within cells, the long strands of DNA form condensed structu
res called chromosomes. The specific location of a DNA sequence within a chromos
ome is known as a locus. If the DNA sequence at a locus varies between individua
ls, the different forms of this sequence are called alleles. DNA sequences can c
hange through mutations, producing new alleles. If a mutation occurs within a ge
ne, the new allele may affect the trait that the gene controls, altering the phe
notype of the organism.[81] However, while this simple correspondence between an
allele and a trait works in some cases, most traits are more complex and are co
ntrolled by quantitative trait loci (multiple interacting genes).[82][83]
Recent findings have confirmed important examples of heritable changes that cann
ot be explained by changes to the sequence of nucleotides in the DNA. These phen
omena are classed as epigenetic inheritance systems.[84] DNA methylation marking
chromatin, self-sustaining metabolic loops, gene silencing by RNA interference
and the three-dimensional conformation of proteins (such as prions) are areas wh
ere epigenetic inheritance systems have been discovered at the organismic level.
[85][86] Developmental biologists suggest that complex interactions in genetic n
etworks and communication among cells can lead to heritable variations that may
underlay some of the mechanics in developmental plasticity and canalisation.[87]
Heritability may also occur at even larger scales. For example, ecological inhe
ritance through the process of niche construction is defined by the regular and
repeated activities of organisms in their environment. This generates a legacy o
f effects that modify and feed back into the selection regime of subsequent gene
rations. Descendants inherit genes plus environmental characteristics generated
by the ecological actions of ancestors.[88] Other examples of heritability in ev
olution that are not under the direct control of genes include the inheritance o
f cultural traits and symbiogenesis.[89][90]
Variation
White peppered moth
Black morph in peppered moth evolution
Further information: Genetic diversity and Population genetics
An individual organism's phenotype results from both its genotype and the influe
nce from the environment it has lived in. A substantial part of the phenotypic v
ariation in a population is caused by genotypic variation.[83] The modern evolut
ionary synthesis defines evolution as the change over time in this genetic varia
tion. The frequency of one particular allele will become more or less prevalent
relative to other forms of that gene. Variation disappears when a new allele rea
ches the point of fixationwhen it either disappears from the population or replac
es the ancestral allele entirely.[91]
Natural selection will only cause evolution if there is enough genetic variation
in a population. Before the discovery of Mendelian genetics, one common hypothe
sis was blending inheritance. But with blending inheritance, genetic variance wo
uld be rapidly lost, making evolution by natural selection implausible. The Hard
yWeinberg principle provides the solution to how variation is maintained in a pop
ulation with Mendelian inheritance. The frequencies of alleles (variations in a
gene) will remain constant in the absence of selection, mutation, migration and
genetic drift.[92]
Variation comes from mutations in the genome, reshuffling of genes through sexua
l reproduction and migration between populations (gene flow). Despite the consta
nt introduction of new variation through mutation and gene flow, most of the gen
ome of a species is identical in all individuals of that species.[93] However, e
ven relatively small differences in genotype can lead to dramatic differences in
phenotype: for example, chimpanzees and humans differ in only about 5% of their
genomes.[94]
Mutation
Main article: Mutation

Duplication of part of a chromosome


Mutations are changes in the DNA sequence of a cell's genome. When mutations occ
ur, they may alter the product of a gene, or prevent the gene from functioning,
or have no effect. Based on studies in the fly Drosophila melanogaster, it has b
een suggested that if a mutation changes a protein produced by a gene, this will
probably be harmful, with about 70% of these mutations having damaging effects,
and the remainder being either neutral or weakly beneficial.[95]
Mutations can involve large sections of a chromosome becoming duplicated (usuall
y by genetic recombination), which can introduce extra copies of a gene into a g
enome.[96] Extra copies of genes are a major source of the raw material needed f
or new genes to evolve.[97] This is important because most new genes evolve with
in gene families from pre-existing genes that share common ancestors.[98] For ex
ample, the human eye uses four genes to make structures that sense light: three
for colour vision and one for night vision; all four are descended from a single
ancestral gene.[99]
New genes can be generated from an ancestral gene when a duplicate copy mutates
and acquires a new function. This process is easier once a gene has been duplica
ted 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 fu
nction.[100][101] Other types of mutations can even generate entirely new genes
from previously noncoding DNA.[102][103]
The generation 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.[104][105] When new genes are assembled from shuffling pre-existin
g parts, domains act as modules with simple independent functions, which can be
mixed together to produce new combinations with new and complex functions.[106]
For example, polyketide synthases are large enzymes that make antibiotics; they
contain up to one hundred independent domains that each catalyse one step in the
overall process, like a step in an assembly line.[107]
Sex and recombination
Further information: Sexual reproduction, Genetic recombination, and Evolution o
f sexual reproduction
In asexual organisms, genes are inherited together, or linked, as they cannot mi
x with genes of other organisms during reproduction. In contrast, the offspring
of sexual organisms contain random mixtures of their parents' chromosomes that a
re produced through independent assortment. In a related process called homologo
us recombination, sexual organisms exchange DNA between two matching chromosomes
.[108] Recombination and reassortment do not alter allele frequencies, but inste
ad change which alleles are associated with each other, producing offspring with
new combinations of alleles.[109] Sex usually increases genetic variation and m
ay increase the rate of evolution.[110][111]

This diagram illustrates the twofold cost of sex. If each individual were to con
tribute to the same number of offspring (two), (a) the sexual population remains
the same size each generation, where the (b) Asexual reproduction population do
ubles in size each generation.
The two-fold cost of sex was first described by John Maynard Smith.[112] The fir
st cost is that in sexually dimorphic species only one of the two sexes can bear
young. (This cost does not apply to hermaphroditic species, like most plants an
d many invertebrates.) The second cost is that any individual who reproduces sex
ually can only pass on 50% of its genes to any individual offspring, with even l
ess passed on as each new generation passes.[113] Yet sexual reproduction is the
more common means of reproduction among eukaryotes and multicellular organisms.
The Red Queen hypothesis has been used to explain the significance of sexual re
production as a means to enable continual evolution and adaptation in response t
o coevolution with other species in an ever-changing environment.[113][114][115]
[116]
Gene flow
Further information: Gene flow
Gene flow is the exchange of genes between populations and between species.[117]
It can therefore be a source of variation that is new to a population or to a s
pecies. Gene flow can be caused by the movement of individuals between separate
populations of organisms, as might be caused by the movement of mice between inl
and and coastal populations, or the movement of pollen between heavy metal toler
ant and heavy metal sensitive populations of grasses.
Gene transfer between species includes the formation of hybrid organisms and hor
izontal gene transfer. Horizontal gene transfer is the transfer of genetic mater
ial from one organism to another organism that is not its offspring; this is mos
t common among bacteria.[118] In medicine, this contributes to the spread of ant
ibiotic resistance, as when one bacteria acquires resistance genes it can rapidl
y transfer them to other species.[119] Horizontal transfer of genes from bacteri
a to eukaryotes such as the yeast Saccharomyces cerevisiae and the adzuki bean w
eevil Callosobruchus chinensis has occurred.[120][121] An example of larger-scal
e transfers are the eukaryotic bdelloid rotifers, which have received a range of
genes from bacteria, fungi and plants.[122] Viruses can also carry DNA between
organisms, allowing transfer of genes even across biological domains.[123]
Large-scale gene transfer has also occurred between the ancestors of eukaryotic
cells and bacteria, during the acquisition of chloroplasts and mitochondria. It
is possible that eukaryotes themselves originated from horizontal gene transfers
between bacteria and archaea.[124]
Mechanisms

Mutation followed by natural selection results in a population with darker colou


ration.
From a Neo-Darwinian perspective, evolution occurs when there are changes in the
frequencies of alleles within a population of interbreeding organisms.[92] For
example, the allele for black colour in a population of moths becoming more comm
on. Mechanisms that can lead to changes in allele frequencies include natural se
lection, genetic drift, genetic hitchhiking, mutation and gene flow.
Natural selection
Further information: Natural selection and Fitness (biology)
Evolution by means of natural selection is the process by which traits that enha
nce survival and reproduction become more common in successive generations of a
population. It has often been called a "self-evident" mechanism because it neces
sarily follows from three simple facts:[24]
Variation exists within populations of organisms with respect to morphol
ogy, physiology, and behaviour (phenotypic variation).
Different traits confer different rates of survival and reproduction (di
fferential fitness).
These traits can be passed from generation to generation (heritability o
f fitness).
More offspring are produced than can possibly survive, and these conditions prod
uce competition between organisms for survival and reproduction. Consequently, o
rganisms with traits that give them an advantage over their competitors are more
likely to pass on their traits to the next generation than those with traits th
at do not confer an advantage.[125]
The central concept of natural selection is the evolutionary fitness of an organ
ism.[126] Fitness is measured by an organism's ability to survive and reproduce,
which determines the size of its genetic contribution to the next generation.[1
26] However, fitness is not the same as the total number of offspring: instead f
itness is indicated by the proportion of subsequent generations that carry an or
ganism's genes.[127] For example, if an organism could survive well and reproduc
e rapidly, but its offspring were all too small and weak to survive, this organi
sm would make little genetic contribution to future generations and would thus h
ave low fitness.[126]
If an allele increases fitness more than the other alleles of that gene, then wi
th each generation this allele will become more common within the population. Th
ese traits are said to be "selected for." Examples of traits that can increase f
itness are enhanced survival and increased fecundity. Conversely, the lower fitn
ess caused by having a less beneficial or deleterious allele results in this all
ele becoming rarerthey are "selected against."[128] Importantly, the fitness of a
n allele is not a fixed characteristic; if the environment changes, previously n
eutral or harmful traits may become beneficial and previously beneficial traits
become harmful.[81] 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 fo
rm (see Dollo's law).[129][130]

These charts depict the different types of genetic selection. On each graph, the
x-axis variable is the type of phenotypic trait and the y-axis variable is the
number of organisms. Group A is the original population and Group B is the popul
ation after selection.
Graph 1 shows directional selection, in which a single extreme phenotype is favo
red.
Graph 2 depicts stabilizing selection, where the intermediate phenotype is favor
ed over the extreme traits.
Graph 3 shows disruptive selection, in which the extreme phenotypes are favored
over the intermediate.
Natural selection within a population for a trait that can vary across a range o
f values, such as height, can be categorised into three different types. The fir
st is directional selection, which is a shift in the average value of a trait ov
er timefor example, organisms slowly getting taller.[131] Secondly, disruptive se
lection is selection for extreme trait values and often results in two different
values becoming most common, with selection against the average value. This wou
ld be when either short or tall organisms had an advantage, but not those of med
ium height. Finally, in stabilising selection there is selection against extreme
trait values on both ends, which causes a decrease in variance around the avera
ge value and less diversity.[125][132] 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 o
rganism to potential mates.[133] Traits that evolved through sexual selection ar
e particularly prominent among males of several animal species. Although sexuall
y favoured, traits such as cumbersome antlers, mating calls, large body size and
bright colours often attract predation, which compromises the survival of indiv
idual males.[134][135] This survival disadvantage is balanced by higher reproduc
tive success in males that show these hard-to-fake, sexually selected traits.[13
6]
Natural selection most generally makes nature the measure against which individu
als 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 inc
ludes all of the organisms...in a given area interacting with the physical envir
onment so that a flow of energy leads to clearly defined trophic structure, biot
ic diversity and material cycles (ie: exchange of materials between living and n
onliving parts) within the system."[137] Each population within an ecosystem occ
upies 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 o
f nature enables scientists to delineate specific forces which, together, compri
se natural selection.
Natural selection can act at different levels of organisation, such as genes, ce
lls, individual organisms, groups of organisms and species.[138][139][140] Selec
tion can act at multiple levels simultaneously.[141] An example of selection occ
urring below the level of the individual organism are genes called transposons,
which can replicate and spread throughout a genome.[142] Selection at a level ab
ove the individual, such as group selection, may allow the evolution of cooperat
ion, as discussed below.[143]
Biased mutation
In addition to being a major source of variation, mutation may also function as
a mechanism of evolution when there are different probabilities at the molecular
level for different mutations to occur, a process known as mutation bias.[144]
If two genotypes, for example one with the nucleotide G and another with the nuc
leotide A in the same position, have the same fitness, but mutation from G to A
happens more often than mutation from A to G, then genotypes with A will tend to
evolve.[145] Different insertion vs. deletion mutation biases in different taxa
can lead to the evolution of different genome sizes.[146][147] Developmental or
mutational biases have also been observed in morphological evolution.[148][149]
For example, according to the phenotype-first theory of evolution, mutations ca
n eventually cause the genetic assimilation of traits that were previously induc
ed by the environment.[150][151][152]
Mutation bias effects are superimposed on other processes. If selection would fa
vor either one out of two mutations, but there is no extra advantage to having b
oth, then the mutation that occurs the most frequently is the one that is most l
ikely to become fixed in a population.[153][154] Mutations leading to the loss o
f function of a gene are much more common than mutations that produce a new, ful
ly functional gene. Most loss of function mutations are selected against. But wh
en selection is weak, mutation bias towards loss of function can affect evolutio
n.[155] For example, pigments are no longer useful when animals live in the dark
ness of caves, and tend to be lost.[156] This kind of loss of function can occur
because of mutation bias, and/or because the function had a cost, and once the
benefit of the function disappeared, natural selection leads to the loss. Loss o
f sporulation ability in Bacillus subtilis during laboratory evolution appears t
o have been caused by mutation bias, rather than natural selection against the c
ost of maintaining sporulation ability.[157] When there is no selection for loss
of function, the speed at which loss evolves depends more on the mutation rate
than it does on the effective population size,[158] indicating that it is driven
more by mutation bias than by genetic drift. In parasitic organisms, mutation b
ias leads to selection pressures as seen in Ehrlichia. Mutations are biased towa
rds antigenic variants in outer-membrane proteins.
Genetic drift
Further information: Genetic drift and Effective population size

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


d 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 are subject to sampling error.[159] As a result, whe
n selective forces are absent or relatively weak, allele frequencies tend to "dr
ift" upward or downward randomly (in a random walk). This drift halts when an al
lele eventually becomes fixed, either by disappearing from the population, or re
placing the other alleles entirely. Genetic drift may therefore eliminate some a
lleles from a population due to chance alone. Even in the absence of selective f
orces, 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.[160]
It is usually difficult to measure the relative importance of selection and neut
ral processes, including drift.[161] The comparative importance of adaptive and
non-adaptive forces in driving evolutionary change is an area of current researc
h.[162]
The neutral theory of molecular evolution proposed that most evolutionary change
s are the result of the fixation of neutral mutations by genetic drift.[163] Hen
ce, in this model, most genetic changes in a population are the result of consta
nt mutation pressure and genetic drift.[164] This form of the neutral theory is
now largely abandoned, since it does not seem to fit the genetic variation seen
in nature.[165][166] However, a more recent and better-supported version of this
model is the nearly neutral theory, where a mutation that would be effectively
neutral in a small population is not necessarily neutral in a large population.[
125] Other alternative theories propose that genetic drift is dwarfed by other s
tochastic forces in evolution, such as genetic hitchhiking, also known as geneti
c draft.[159][167][168]
The time for a neutral allele to become fixed by genetic drift depends on popula
tion size, with fixation occurring more rapidly in smaller populations.[169] The
number of individuals in a population is not critical, but instead a measure kn
own as the effective population size.[170] The effective population is usually s
maller than the total population since it takes into account factors such as the
level of inbreeding and the stage of the lifecycle in which the population is t
he smallest.[170] The effective population size may not be the same for every ge
ne in the same population.[171]
Genetic hitchhiking
Further information: Genetic hitchhiking, HillRobertson effect, and Selective swe
ep
Recombination allows alleles on the same strand of DNA to become separated. Howe
ver, the rate of recombination is low (approximately two events per chromosome p
er 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 i
nherited together, a phenomenon known as linkage.[172] This tendency is measured
by finding how often two alleles occur together on a single chromosome compared
to expectations, which is called their linkage disequilibrium. A set of alleles
that is usually inherited in a group is called a haplotype. This can be importa
nt when one allele in a particular haplotype is strongly beneficial: natural sel
ection 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 geneti
c hitchhiking or genetic draft.[173] Genetic draft caused by the fact that some
neutral genes are genetically linked to others that are under selection can be p
artially captured by an appropriate effective population size.[167]
Gene flow
Further information: Gene flow, Hybrid (biology), and Horizontal gene transfer
Gene flow involves the exchange of genes between populations and between species
.[117] The presence or absence of gene flow fundamentally changes the course of
evolution. Due to the complexity of organisms, any two completely isolated popul
ations will eventually evolve genetic incompatibilities through neutral processe
s, as in the Bateson-Dobzhansky-Muller model, even if both populations remain es
sentially identical in terms of their adaptation to the environment.
If genetic differentiation between populations develops, gene flow between popul
ations can introduce traits or alleles which are disadvantageous in the local po
pulation and this may lead to organisms within these populations evolving mechan
isms that prevent mating with genetically distant populations, eventually result
ing in the appearance of new species. Thus, exchange of genetic information betw
een individuals is fundamentally important for the development of the biological
species concept.
During the development of the modern synthesis, Sewall Wright developed his shif
ting balance theory, which regarded gene flow between partially isolated populat
ions as an important aspect of adaptive evolution.[174] However, recently there
has been substantial criticism of the importance of the shifting balance theory.
[175]
Outcomes
Play media
A visual demonstration of rapid antibiotic resistance evolution by E. coli growi
ng across a plate with increasing concentrations of trimethoprim.[176]
Evolution influences every aspect of the form and behaviour of organisms. Most p
rominent are the specific behavioural and physical adaptations that are the outc
ome of natural selection. These adaptations increase fitness by aiding activitie
s such as finding food, avoiding predators or attracting mates. Organisms can al
so respond to selection by cooperating with each other, usually by aiding their
relatives or engaging in mutually beneficial symbiosis. In the longer term, evol
ution produces new species through splitting ancestral populations of organisms
into new groups that cannot or will not interbreed.
These outcomes of evolution are distinguished based on time scale as macroevolut
ion versus microevolution. Macroevolution refers to evolution that occurs at or
above the level of species, in particular speciation and extinction; whereas mic
roevolution refers to smaller evolutionary changes within a species or populatio
n, in particular shifts in gene frequency and adaptation.[177] In general, macro
evolution is regarded as the outcome of long periods of microevolution.[178] Thu
s, the distinction between micro- and macroevolution is not a fundamental onethe
difference is simply the time involved.[179] However, in macroevolution, the tra
its of the entire species may be important. For instance, a large amount of vari
ation among individuals allows a species to rapidly adapt to new habitats, lesse
ning 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 levelswith microevolution acting on genes and organisms,
versus macroevolutionary processes such as species selection acting on entire sp
ecies and affecting their rates of speciation and extinction.[180][181][182]
A common misconception is that evolution has goals, long-term plans, or an innat
e tendency for "progress," as expressed in beliefs such as orthogenesis and evol
utionism; realistically however, evolution has no long-term goal and does not ne
cessarily produce greater complexity.[183][184][185] Although complex species ha
ve evolved, they occur as a side effect of the overall number of organisms incre
asing and simple forms of life still remain more common in the biosphere.[186] F
or example, the overwhelming majority of species are microscopic prokaryotes, wh
ich form about half the world's biomass despite their small size,[187] and const
itute the vast majority of Earth's biodiversity.[188] Simple organisms have ther
efore been the dominant form of life on Earth throughout its history and continu
e to be the main form of life up to the present day, with complex life only appe
aring more diverse because it is more noticeable.[189] Indeed, the evolution of
microorganisms is particularly important to modern evolutionary research, since
their rapid reproduction allows the study of experimental evolution and the obse
rvation of evolution and adaptation in real time.[190][191]
Adaptation
For more details on this topic, see Adaptation.

Homologous bones in the limbs of tetrapods. The bones of these animals have the
same basic structure, but have been adapted for specific uses.
Adaptation is the process that makes organisms better suited to their habitat.[1
92][193] 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 grindi
ng of grass. By using the term adaptation for the evolutionary process and adapt
ive trait for the product (the bodily part or function), the two senses of the w
ord may be distinguished. Adaptations are produced by natural selection.[194] Th
e following definitions are due to Theodosius Dobzhansky:
1. Adaptation is the evolutionary process whereby an organism becom
es better able to live in its habitat or habitats.[195]
2. Adaptedness is the state of being adapted: the degree to which a
n organism is able to live and reproduce in a given set of habitats.[196]
3. An adaptive trait is an aspect of the developmental pattern of t
he organism which enables or enhances the probability of that organism surviving
and reproducing.[197]
Adaptation may cause either the gain of a new feature, or the loss of an ancestr
al feature. An example that shows both types of change is bacterial adaptation t
o antibiotic selection, with genetic changes causing antibiotic resistance by bo
th modifying the target of the drug, or increasing the activity of transporters
that pump the drug out of the cell.[198] Other striking examples are the bacteri
a Escherichia coli evolving the ability to use citric acid as a nutrient in a lo
ng-term laboratory experiment,[199] Flavobacterium evolving a novel enzyme that
allows these bacteria to grow on the by-products of nylon manufacturing,[200][20
1] and the soil bacterium Sphingobium evolving an entirely new metabolic pathway
that degrades the synthetic pesticide pentachlorophenol.[202][203] An interesti
ng but still controversial idea is that some adaptations might increase the abil
ity of organisms to generate genetic diversity and adapt by natural selection (i
ncreasing organisms' evolvability).[204][205][206][207][208]

A baleen whale skeleton, a and b label flipper bones, which were adapted from fr
ont leg bones: while c indicates vestigial leg bones, suggesting an adaptation f
rom land to sea.[209]
Adaptation occurs through the gradual modification of existing structures. Conse
quently, structures with similar internal organisation may have different functi
ons in related organisms. This is the result of a single ancestral structure bei
ng adapted to function in different ways. The bones within bat wings, for exampl
e, are very similar to those in mice feet and primate hands, due to the descent
of all these structures from a common mammalian ancestor.[210] However, since al
l living organisms are related to some extent,[211] even organs that appear to h
ave 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 comm
on set of homologous genes that control their assembly and function; this is cal
led deep homology.[212][213]
During evolution, some structures may lose their original function and become ve
stigial structures.[214] Such structures may have little or no function in a cur
rent species, yet have a clear function in ancestral species, or other closely r
elated species. Examples include pseudogenes,[215] the non-functional remains of
eyes in blind cave-dwelling fish,[216] wings in flightless birds,[217] the pres
ence of hip bones in whales and snakes,[209] and sexual traits in organisms that
reproduce via asexual reproduction.[218] Examples of vestigial structures in hu
mans include wisdom teeth,[219] the coccyx,[214] the vermiform appendix,[214] an
d other behavioural vestiges such as goose bumps[220][221] and primitive reflexe
s.[222][223][224]
However, many traits that appear to be simple adaptations are in fact exaptation
s: structures originally adapted for one function, but which coincidentally beca
me somewhat useful for some other function in the process.[225] One example is t
he 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, i
n this species, the head has become so flattened that it assists in gliding from
tree to treean exaptation.[225] Within cells, molecular machines such as the bac
terial flagella[226] and protein sorting machinery[227] evolved by the recruitme
nt of several pre-existing proteins that previously had different functions.[177
] Another example is the recruitment of enzymes from glycolysis and xenobiotic m
etabolism to serve as structural proteins called crystallins within the lenses o
f organisms' eyes.[228][229]
An area of current investigation in evolutionary developmental biology is the de
velopmental basis of adaptations and exaptations.[230] This research addresses t
he origin and evolution of embryonic development and how modifications of develo
pment and developmental processes produce novel features.[231] These studies hav
e shown that evolution can alter development to produce new structures, such as
embryonic bone structures that develop into the jaw in other animals instead for
ming part of the middle ear in mammals.[232] It is also possible for structures
that have been lost in evolution to reappear due to changes in developmental gen
es, such as a mutation in chickens causing embryos to grow teeth similar to thos
e of crocodiles.[233] It is now becoming clear that most alterations in the form
of organisms are due to changes in a small set of conserved genes.[234]
Coevolution

Common garter snake (Thamnophis sirtalis sirtalis) has evolved resistance to the
defensive substance tetrodotoxin in its amphibian prey.
Further information: Coevolution
Interactions between organisms can produce both conflict and cooperation. When t
he 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. He
re, the evolution of one species causes adaptations in a second species. These c
hanges in the second species then, in turn, cause new adaptations in the first s
pecies. This cycle of selection and response is called coevolution.[235] An exam
ple is the production of tetrodotoxin in the rough-skinned newt and the evolutio
n of tetrodotoxin resistance in its predator, the common garter snake. In this p
redator-prey pair, an evolutionary arms race has produced high levels of toxin i
n the newt and correspondingly high levels of toxin resistance in the snake.[236
]
Cooperation
Further information: Co-operation (evolution)
Not all co-evolved interactions between species involve conflict.[237] Many case
s of mutually beneficial interactions have evolved. For instance, an extreme coo
peration exists between plants and the mycorrhizal fungi that grow on their root
s and aid the plant in absorbing nutrients from the soil.[238] This is a recipro
cal relationship as the plants provide the fungi with sugars from photosynthesis
. Here, the fungi actually grow inside plant cells, allowing them to exchange nu
trients with their hosts, while sending signals that suppress the plant immune s
ystem.[239]
Coalitions between organisms of the same species have also evolved. An extreme c
ase 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 mak
e up the body of an animal limit their reproduction so they can maintain a stabl
e organism, which then supports a small number of the animal's germ cells to pro
duce offspring. Here, somatic cells respond to specific signals that instruct th
em whether to grow, remain as they are, or die. If cells ignore these signals an
d multiply inappropriately, their uncontrolled growth causes cancer.[240]
Such cooperation within species may have evolved through the process of kin sele
ction, which is where one organism acts to help raise a relative's offspring.[24
1] This activity is selected for because if the helping individual contains alle
les which promote the helping activity, it is likely that its kin will also cont
ain these alleles and thus those alleles will be passed on.[242] Other processes
that may promote cooperation include group selection, where cooperation provide
s benefits to a group of organisms.[243]
Speciation
Main article: Speciation

The four mechanisms of speciation


Speciation is the process where a species diverges into two or more descendant s
pecies.[244]
There are multiple ways to define the concept of "species." The choice of defini
tion is dependent on the particularities of the species concerned.[245] For exam
ple, some species concepts apply more readily toward sexually reproducing organi
sms while others lend themselves better toward asexual organisms. Despite the di
versity of various species concepts, these various concepts can be placed into o
ne of three broad philosophical approaches: interbreeding, ecological and phylog
enetic.[246] The Biological Species Concept (BSC) is a classic example of the in
terbreeding approach. Defined by Ernst Mayr in 1942, the BSC states that "specie
s are groups of actually or potentially interbreeding natural populations, which
are reproductively isolated from other such groups."[247] Despite its wide and
long-term use, the BSC like others is not without controversy, for example becau
se these concepts cannot be applied to prokaryotes,[248] and this is called the
species problem.[245] Some researchers have attempted a unifying monistic defini
tion of species, while others adopt a pluralistic approach and suggest that ther
e may be different ways to logically interpret the definition of a species.[245]
[246]
Barriers to reproduction between two diverging sexual populations are required f
or the populations to become new species. Gene flow may slow this process by spr
eading the new genetic variants also to the other populations. Depending on how
far two species have diverged since their most recent common ancestor, it may st
ill be possible for them to produce offspring, as with horses and donkeys mating
to produce mules.[249] Such hybrids are generally infertile. In this case, clos
ely related species may regularly interbreed, but hybrids will be selected again
st and the species will remain distinct. However, viable hybrids are occasionall
y formed and these new species can either have properties intermediate between t
heir parent species, or possess a totally new phenotype.[250] The importance of
hybridisation in producing new species of animals is unclear, although cases hav
e been seen in many types of animals,[251] with the gray tree frog being a parti
cularly well-studied example.[252]
Speciation has been observed multiple times under both controlled laboratory con
ditions and in nature.[253] In sexually reproducing organisms, speciation result
s from reproductive isolation followed by genealogical divergence. There are fou
r mechanisms for speciation. The most common in animals is allopatric speciation
, which occurs in populations initially isolated geographically, such as by habi
tat fragmentation or migration. Selection under these conditions can produce ver
y rapid changes in the appearance and behaviour of organisms.[254][255] As selec
tion and drift act independently on populations isolated from the rest of their
species, separation may eventually produce organisms that cannot interbreed.[256
]
The second mechanism of speciation is peripatric speciation, which occurs when s
mall populations of organisms become isolated in a new environment. This differs
from allopatric speciation in that the isolated populations are numerically muc
h smaller than the parental population. Here, the founder effect causes rapid sp
eciation after an increase in inbreeding increases selection on homozygotes, lea
ding to rapid genetic change.[257]
The third mechanism of speciation is parapatric speciation. This is similar to p
eripatric speciation in that a small population enters a new habitat, but differ
s in that there is no physical separation between these two populations. Instead
, speciation results from the evolution of mechanisms that reduce gene flow betw
een the two populations.[244] Generally this occurs when there has been a drasti
c change in the environment within the parental species' habitat. One example is
the grass Anthoxanthum odoratum, which can undergo parapatric speciation in res
ponse to localised metal pollution from mines.[258] Here, plants evolve that hav
e resistance to high levels of metals in the soil. Selection against interbreedi
ng with the metal-sensitive parental population produced a gradual change in the
flowering time of the metal-resistant plants, which eventually produced complet
e reproductive isolation. Selection against hybrids between the two populations
may cause reinforcement, which is the evolution of traits that promote mating wi
thin a species, as well as character displacement, which is when two species bec
ome more distinct in appearance.[259]
Geographical isolation of finches on the Galpagos Islands produced over a dozen n
ew 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 ma
y remove genetic differences between parts of a population.[260] Generally, symp
atric speciation in animals requires the evolution of both genetic differences a
nd non-random mating, to allow reproductive isolation to evolve.[261]
One type of sympatric speciation involves crossbreeding of two related species t
o 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 f
rom each parent are from different species and cannot successfully pair. However
, it is more common in plants because plants often double their number of chromo
somes, to form polyploids.[262] This allows the chromosomes from each parental s
pecies to form matching pairs during meiosis, since each parent's chromosomes ar
e represented by a pair already.[263] An example of such a speciation event is w
hen the plant species Arabidopsis thaliana and Arabidopsis arenosa crossbred to
give the new species Arabidopsis suecica.[264] This happened about 20,000 years
ago,[265] and the speciation process has been repeated in the laboratory, which
allows the study of the genetic mechanisms involved in this process.[266] Indeed
, chromosome doubling within a species may be a common cause of reproductive iso
lation, as half the doubled chromosomes will be unmatched when breeding with und
oubled organisms.[267]
Speciation events are important in the theory of punctuated equilibrium, which a
ccounts for the pattern in the fossil record of short "bursts" of evolution inte
rspersed with relatively long periods of stasis, where species remain relatively
unchanged.[268] In this theory, speciation and rapid evolution are linked, with
natural selection and genetic drift acting most strongly on organisms undergoin
g 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 organ
isms undergoing speciation and rapid evolution are found in small populations or
geographically restricted habitats and therefore rarely being preserved as foss
ils.[181]
Extinction
Further information: Extinction

Tyrannosaurus rex. Non-avian dinosaurs died out in the CretaceousPaleogene extinc


tion event at the end of the Cretaceous period.
Extinction is the disappearance of an entire species. Extinction is not an unusu
al event, as species regularly appear through speciation and disappear through e
xtinction.[269] Nearly all animal and plant species that have lived on Earth are
now extinct,[270] and extinction appears to be the ultimate fate of all species
.[271] These extinctions have happened continuously throughout the history of li
fe, although the rate of extinction spikes in occasional mass extinction events.
[272] The CretaceousPaleogene extinction event, during which the non-avian dinosa
urs became extinct, is the most well-known, but the earlier PermianTriassic extin
ction event was even more severe, with approximately 96% of all marine species d
riven to extinction.[272] The Holocene extinction event is an ongoing mass extin
ction associated with humanity's expansion across the globe over the past few th
ousand years. Present-day extinction rates are 1001000 times greater than the bac
kground rate and up to 30% of current species may be extinct by the mid 21st cen
tury.[273] Human activities are now the primary cause of the ongoing extinction
event;[274] global warming may further accelerate it in the future.[275]
The role of extinction in evolution is not very well understood and may depend o
n which type of extinction is considered.[272] The causes of the continuous "low
-level" extinction events, which form the majority of extinctions, may be the re
sult of competition between species for limited resources (the competitive exclu
sion principle).[72] 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.[139] 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.[276]
Evolutionary history of life
Human timeline
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Human-like
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Nakalipithecus
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Orrorin
Ardipithecus
Australopithecus
Homo habilis
Homo erectus
Neanderthal
Homo sapiens

Earlier apes

Possibly bipedal

Earliest bipedal

Earliest stone tools

Earliest exit
from Africa

Earliest fire use

Earliest cooking

Earliest clothes

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Also see: Life timeline and Nature timeline
Life timeline
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water
Single-celled
life
photosynthesis
Eukaryotes
Multicellular
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Land life
Dinosaurs
Mammals
Flowers

Earliest Earth (4540)

Earliest water

Earliest life

LHB meteorites

Earliest oxygen

Atmospheric oxygen

Oxygen crisis

Earliest sexual reproduction

Ediacara biota

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Earliest humans
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Axis scale: millions of years.


Orange labels: known ice ages.
Also see: Human timeline and Nature timeline
Main article: Evolutionary history of life
See also: Timeline of evolutionary history of life and Timeline of human evoluti
on
Origin of life
Further information: Abiogenesis, Panspermia, and RNA world hypothesis
The Earth is about 4.54 billion years old.[277][278][279] The earliest undispute
d evidence of life on Earth dates from at least 3.5 billion years ago,[7][280] d
uring the Eoarchean Era after a geological crust started to solidify following t
he earlier molten Hadean Eon. Microbial mat fossils have been found in 3.48 bill
ionyearold sandstone in Western Australia.[13][14][15] Other early physical ev
idence of a biogenic substance is graphite in 3.7 billionyearold metasedimenta
ry rocks discovered in Western Greenland[12] as well as "remains of biotic life"
found in 4.1 billionyearold rocks in Western Australia.[8][9] According to on
e of the researchers, "If life arose relatively quickly on Earth then it could b
e common in the universe."[8]
More than 99 percent of all species, amounting to over five billion species,[281
] that ever lived on Earth are estimated to be extinct.[17][18] Estimates on the
number of Earth's current species range from 10 million to 14 million,[19][20]
of which about 1.9 million are estimated to have been named[21] and 1.6 million
documented in a central database to date,[22] leaving at least 80 percent not ye
t described.
Highly energetic chemistry is thought to have produced a selfreplicating molecu
le around 4 billion years ago, and half a billion years later the last common an
cestor of all life existed.[5] The current scientific consensus is that the comp
lex biochemistry that makes up life came from simpler chemical reactions.[282] T
he beginning of life may have included selfreplicating molecules such as RNA[28
3] and the assembly of simple cells.[284]
Common descent
Further information: Common descent and Evidence of common descent
All organisms on Earth are descended from a common ancestor or ancestral gene po
ol.[211][285] Current species are a stage in the process of evolution, with thei
r diversity the product of a long series of speciation and extinction events.[28
6] The common descent of organisms was first deduced from four simple facts abou
t organisms: First, they have geographic distributions that cannot be explained
by local adaptation. Second, the diversity of life is not a set of completely un
ique organisms, but organisms that share morphological similarities. Third, vest
igial traits with no clear purpose resemble functional ancestral traits and fina
lly, that organisms can be classified using these similarities into a hierarchy
of nested groupssimilar to a family tree.[287] However, modern research has sugge
sted that, due to horizontal gene transfer, this "tree of life" may be more comp
licated than a simple branching tree since some genes have spread independently
between distantly related species.[288][289]

The hominoids are descendants of a common ancestor.


Past species have also left records of their evolutionary history. Fossils, alon
g with the comparative anatomy of presentday organisms, constitute the morpholo
gical, or anatomical, record.[290] By comparing the anatomies of both modern and
extinct species, paleontologists can infer the lineages of those species. Howev
er, this approach is most successful for organisms that had hard body parts, suc
h as shells, bones or teeth. Further, as prokaryotes such as bacteria and archae
a share a limited set of common morphologies, their fossils do not provide infor
mation on their ancestry.
More recently, evidence for common descent has come from the study of biochemica
l similarities between organisms. For example, all living cells use the same bas
ic set of nucleotides and amino acids.[291] The development of molecular genetic
s has revealed the record of evolution left in organisms' genomes: dating when s
pecies diverged through the molecular clock produced by mutations.[292] For exam
ple, these DNA sequence comparisons have revealed that humans and chimpanzees sh
are 98% of their genomes and analysing the few areas where they differ helps she
d light on when the common ancestor of these species existed.[293]
Evolution of life
Main articles: Evolutionary history of life and Timeline of evolutionary history
of life

Evolutionary tree showing the divergence of modern species from their common anc
estor in the centre.[294] The three domains are coloured, with bacteria blue, ar
chaea green and eukaryotes red.
Prokaryotes inhabited the Earth from approximately 34 billion years ago.[295][296
] No obvious changes in morphology or cellular organisation occurred in these or
ganisms over the next few billion years.[297] The eukaryotic cells emerged betwe
en 1.62.7 billion years ago. The next major change in cell structure came when ba
cteria were engulfed by eukaryotic cells, in a cooperative association called en
dosymbiosis.[298][299] The engulfed bacteria and the host cell then underwent co
evolution, with the bacteria evolving into either mitochondria or hydrogenosomes
.[300] Another engulfment of cyanobacteriallike organisms led to the formation
of chloroplasts in algae and plants.[301]
The history of life was that of the unicellular eukaryotes, prokaryotes and arch
aea until about 610 million years ago when multicellular organisms began to appe
ar in the oceans in the Ediacaran period.[295][302] The evolution of multicellul
arity occurred in multiple independent events, in organisms as diverse as sponge
s, brown algae, cyanobacteria, slime moulds and myxobacteria.[303] In January 20
16, scientists reported that, about 800 million years ago, a minor genetic chang
e in a single molecule called GKPID may have allowed organisms to go from a sin
gle cell organism to one of many cells.[304]
Soon after the emergence of these first multicellular organisms, a remarkable am
ount of biological diversity appeared over approximately 10 million years, in an
event called the Cambrian explosion. Here, the majority of types of modern anim
als appeared in the fossil record, as well as unique lineages that subsequently
became extinct.[305] Various triggers for the Cambrian explosion have been propo
sed, including the accumulation of oxygen in the atmosphere from photosynthesis.
[306]
About 500 million years ago, plants and fungi colonised the land and were soon f
ollowed by arthropods and other animals.[307] Insects were particularly successf
ul and even today make up the majority of animal species.[308] Amphibians first
appeared around 364 million years ago, followed by early amniotes and birds arou
nd 155 million years ago (both from "reptile"like lineages), mammals around 129
million years ago, homininae around 10 million years ago and modern humans arou
nd 250,000 years ago.[309][310][311] However, despite the evolution of these lar
ge animals, smaller organisms similar to the types that evolved early in this pr
ocess continue to be highly successful and dominate the Earth, with the majority
of both biomass and species being prokaryotes.[188]
Applications
Main articles: Applications of evolution, Selective breeding, and Evolutionary c
omputation
Concepts and models used in evolutionary biology, such as natural selection, hav
e many applications.[312]
Artificial selection is the intentional selection of traits in a population of o
rganisms. This has been used for thousands of years in the domestication of plan
ts and animals.[313] More recently, such selection has become a vital part of ge
netic engineering, with selectable markers such as antibiotic resistance genes b
eing used to manipulate DNA. Proteins with valuable properties have evolved by r
epeated rounds of mutation and selection (for example modified enzymes and new a
ntibodies) in a process called directed evolution.[314]
Understanding the changes that have occurred during an organism's evolution can
reveal the genes needed to construct parts of the body, genes which may be invol
ved in human genetic disorders.[315] For example, the Mexican tetra is an albino
cavefish that lost its eyesight during evolution. Breeding together different p
opulations of this blind fish produced some offspring with functional eyes, sinc
e different mutations had occurred in the isolated populations that had evolved
in different caves.[316] This helped identify genes required for vision and pigm
entation.[317]
Many human diseases are not static phenomena, but capable of evolution. Viruses,
bacteria, fungi and cancers evolve to be resistant to host immune defences, as
well as pharmaceutical drugs.[318][319][320] These same problems occur in agricu
lture with pesticide[321] and herbicide[322] resistance. It is possible that we
are facing the end of the effective life of most of available antibiotics[323] a
nd predicting the evolution and evolvability[324] of our pathogens and devising
strategies to slow or circumvent it is requiring deeper knowledge of the complex
forces driving evolution at the molecular level.[325]
In computer science, simulations of evolution using evolutionary algorithms and
artificial life started in the 1960s and were extended with simulation of artifi
cial selection.[326] Artificial evolution became a widely recognised optimisatio
n method as a result of the work of Ingo Rechenberg in the 1960s. He used evolut
ion strategies to solve complex engineering problems.[327] Genetic algorithms in
particular became popular through the writing of John Henry Holland.[328] Pract
ical applications also include automatic evolution of computer programmes.[329]
Evolutionary algorithms are now used to solve multidimensional problems more ef
ficiently than software produced by human designers and also to optimise the des
ign of systems.[330]
Social and cultural responses
Further information: Social effects of evolutionary theory, 1860 Oxford evolutio
n debate, Creationevolution controversy, and Objections to evolution

As evolution became widely accepted in the 1870s, caricatures of Charles Darwin


with an ape or monkey body symbolised evolution.[331]
In the 19th century, particularly after the publication of On the Origin of Spec
ies in 1859, the idea that life had evolved was an active source of academic deb
ate centred on the philosophical, social and religious implications of evolution
. Today, the modern evolutionary synthesis is accepted by a vast majority of sci
entists.[72] However, evolution remains a contentious concept for some theists.[
332]
While various religions and denominations have reconciled their beliefs with evo
lution through concepts such as theistic evolution, there are creationists who b
elieve that evolution is contradicted by the creation myths found in their relig
ions and who raise various objections to evolution.[177][333][334] As had been d
emonstrated by responses to the publication of Vestiges of the Natural History o
f Creation in 1844, the most controversial aspect of evolutionary biology is the
implication of human evolution that humans share common ancestry with apes and
that the mental and moral faculties of humanity have the same types of natural c
auses as other inherited traits in animals.[335] In some countries, notably the
United States, these tensions between science and religion have fuelled the curr
ent creationevolution controversy, a religious conflict focusing on politics and
public education.[336] While other scientific fields such as cosmology[337] and
Earth science[338] also conflict with literal interpretations of many religious
texts, evolutionary biology experiences significantly more opposition from relig
ious literalists.
The teaching of evolution in American secondary school biology classes was uncom
mon 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 textbo
oks for a generation, but it was gradually reintroduced later and became legall
y protected with the 1968 Epperson v. Arkansas decision. Since then, the competi
ng religious belief of creationism was legally disallowed in secondary school cu
rricula in various decisions in the 1970s and 1980s, but it returned in pseudosc
ientific form as intelligent design (ID), to be excluded once again in the 2005
Kitzmiller v. Dover Area School District case.[339]
See also
Book: Evolution
Argument from poor design
Biocultural evolution
Biological classification
Evidence of common descent
Evolutionary anthropology
Evolutionary ecology
Evolutionary epistemology
Evolutionary neuroscience
Evolution of biological complexity
Evolution of plants
Timeline of the evolutionary history of life
Unintelligent design
Universal Darwinism
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Further reading
Further information: Bibliography of biology
Library resources about
Evolution
Online books
Resources in your library
Resources in other libraries
Introductory reading
Barrett, Paul H.; Weinshank, Donald J.; Gottleber, Timothy T., eds. (198
1). A Concordance to Darwin's Origin of Species, First Edition. Ithaca, NY: Corn
ell University Press. ISBN0-8014-1319-2. LCCN80066893. OCLC610057960.
Carroll, Sean B. (2005). Endless Forms Most Beautiful: The New Science o
f Evo Devo and the Making of the Animal Kingdom. illustrations by Jamie W. Carro
ll, Josh P. Klaiss, Leanne M. Olds (1st ed.). New York: W. W. Norton & Company.
ISBN0-393-06016-0. LCCN2004029388. OCLC57316841.
Charlesworth, Brian; Charlesworth, Deborah (2003). Evolution: A Very Sho
rt Introduction. Very Short Introductions. Oxford; New York: Oxford University P
ress. ISBN0-19-280251-8. LCCN2003272247. OCLC51668497.
Gould, Stephen Jay (1989). Wonderful Life: The Burgess Shale and the Nat
ure of History (1st ed.). New York: W. W. Norton & Company. ISBN0-393-02705-8. LC
CN88037469. OCLC18983518.
Jones, Steve (1999). Almost Like a Whale: The Origin of Species Updated.
London; New York: Doubleday. ISBN0-385-40985-0. LCCN2002391059. OCLC41420544.
(2000). Darwin's Ghost: The Origin of Species Updated (1st ed.). New York
: Random House. ISBN0-375-50103-7. LCCN99053246. OCLC42690131. American version.
Mader, Sylvia S. (2007). Biology. Significant contributions by Murray P.
Pendarvis (9th ed.). Boston, MA: McGraw-Hill Higher Education. ISBN978-0-07-2464
63-4. LCCN2005027781. OCLC61748307.
Maynard Smith, John (1993). The Theory of Evolution (Canto ed.). Cambrid
ge; New York: Cambridge University Press. ISBN0-521-45128-0. LCCN93020358. OCLC2767
6642.
Pallen, Mark J. (2009). The Rough Guide to Evolution. Rough Guides Refer
ence Guides. London; New York: Rough Guides. ISBN978-1-85828-946-5. LCCN2009288090
. OCLC233547316.
Advanced reading
Barton, Nicholas H.; Briggs, Derek E. G.; Eisen, Jonathan A.; et al. (20
07). Evolution. Cold Spring Harbor, NY: Cold Spring Harbor Laboratory Press. ISB
N978-0-87969-684-9. LCCN2007010767. OCLC86090399.
Coyne, Jerry A.; Orr, H. Allen (2004). Speciation. Sunderland, MA: Sinau
er Associates. ISBN0-87893-089-2. LCCN2004009505. OCLC55078441.
Bergstrom, Carl T.; Dugatkin, Lee Alan (2012). Evolution (1st ed.). New
York: W. W. Norton & Company. ISBN978-0-393-91341-5. LCCN2011036572. OCLC729341924.
Gould, Stephen Jay (2002). The Structure of Evolutionary Theory. Cambrid
ge, MA: Belknap Press of Harvard University Press. ISBN0-674-00613-5. LCCN20010435
56. OCLC47869352.
Hall, Brian K.; Olson, Wendy, eds. (2003). Keywords and Concepts in Evol
utionary Developmental Biology. Cambridge, MA: Harvard University Press. ISBN0-67
4-00904-5. LCCN2002192201. OCLC50761342.
Kauffman, Stuart A. (1993). The Origins of Order: Self-organization and
Selection in Evolution. New York, NY: Oxford University Press.
Maynard Smith, John; Szathmry, Ers (1995). The Major Transitions in Evolut
ion. Oxford; New York: W.H. Freeman Spektrum. ISBN0-7167-4525-9. LCCN94026965. OCL
C30894392.
Mayr, Ernst (2001). What Evolution Is. New York: Basic Books. ISBN0-465-0
4426-3. LCCN2001036562. OCLC47443814.
Minelli, Alessandro (2009). Forms of Becoming: The Evolutionary Biology
of Development. Translation by Mark Epstein. Princeton, NJ; Oxford: Princeton Un
iversity Press. ISBN978-0-691-13568-7. LCCN2008028825. OCLC233030259.
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Evolution on In Our Time at the BBC. (listen now)
"Evolution". New Scientist. Retrieved 2011-05-30.
"Evolution Resources from the National Academies". Washington, D.C.: Nat
ional Academy of Sciences. Retrieved 2011-05-30.
"Understanding Evolution: your one-stop resource for information on evol
ution". University of California, Berkeley. Retrieved 2011-05-30.
"Evolution of Evolution 150 Years of Darwin's 'On the Origin of Species'
". Arlington County, VA: National Science Foundation. Retrieved 2011-05-30.
Human Timeline (Interactive) Smithsonian, National Museum of Natural His
tory (August 2016).
Experiments concerning the process of biological evolution
Lenski, Richard E. "Experimental Evolution". Michigan State University.
Retrieved 2013-07-31.
Chastain, Erick; Livnat, Adi; Papadimitriou, Christos; Vazirani, Umesh (
July 22, 2014). "Algorithms, games, and evolution". Proc. Natl. Acad. Sci. U.S.A
. Washington, D.C.: National Academy of Sciences. 111 (29): 1062010623. Bibcode:2
014PNAS..11110620C. doi:10.1073/pnas.1406556111. ISSN0027-8424. Retrieved 2015-01
-03.
Online lectures
Carroll, Sean B. "The Making of the Fittest". Retrieved 2011-05-30.
Stearns, Stephen C. "Principles of Evolution, Ecology and Behavior". Ret
rieved 2011-08-30.
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