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Natural selection

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</style><div role="note" class="hatnote navigation-not-searchable">For other uses, see <a href="Natural_selection_(disambiguation)" class="mw-redirect mw-disambig" title="Natural selection (disambiguation)">Natural selection (disambiguation)</a>.</div>

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</style><table class="sidebar sidebar-collapse nomobile nowraplinks" style="border:2px solid #90ee90"><tbody><tr><td class="sidebar-pretitle">Part of a series on</td></tr><tr><th class="sidebar-title-with-pretitle" style="background:#90ee90;padding:0.2em;font-size:175%;font-weight:bold"><a href="Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></th></tr><tr><td class="sidebar-image"><div class="sidebar-caption"><a href="Darwin's_finches" title="Darwin's finches">Darwin's finches</a> by <a href="John_Gould" title="John Gould">John Gould</a></div></td></tr><tr><td class="sidebar-above" style="display:block; margin-bottom:0.55em; background-color: transparent;">

<ul><li><a href="Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index</a></li>

<li><a href="Introduction_to_evolution" title="Introduction to evolution">Introduction</a></li>

<li><a href="Outline_of_evolution" title="Outline of evolution">Outline</a></li></ul>

</div>

<ul><li><a href="Glossary_of_evolutionary_biology" class="mw-redirect" title="Glossary of evolutionary biology">Glossary</a></li>

<li><a href="Evidence_of_common_descent" title="Evidence of common descent">Evidence</a></li>

<li><a href="History_of_life" title="History of life">History</a></li></ul>

</div></td></tr><tr><td class="sidebar-content" style="background:transparent;">

<div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Processes and outcomes</div><div class="sidebar-list-content mw-collapsible-content plainlist"><div class="hlist">

<ul><li><a href="Population_genetics" title="Population genetics">Population genetics</a></li>

<li><a href="Genetic_variation" title="Genetic variation">Variation</a></li>

<li><a href="Genetic_diversity" title="Genetic diversity">Diversity</a></li>

<li><a href="Mutation" title="Mutation">Mutation</a></li>

<li><a href="Adaptation" title="Adaptation">Adaptation</a></li>

<li><a href="Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li>

<li><a href="Genetic_drift" title="Genetic drift">Genetic drift</a></li>

<li><a href="Speciation" title="Speciation">Speciation</a></li>

<li><a href="Adaptive_radiation" title="Adaptive radiation">Adaptive radiation</a></li>

<li><a href="Cooperation_(evolution)" title="Cooperation (evolution)">Co-operation</a></li>

<li><a href="Coevolution" title="Coevolution">Coevolution</a></li>

<li><a href="Coextinction" title="Coextinction">Coextinction</a></li>

<li><a href="Contingency_(evolutionary_biology)" title="Contingency (evolutionary biology)">Contingency</a></li>

<li><a href="Divergent_evolution" title="Divergent evolution">Divergence</a></li>

<li><a href="Convergent_evolution" title="Convergent evolution">Convergence</a></li>

<li><a href="Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li>

<li><a href="Extinction" title="Extinction">Extinction</a></li></ul>

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<div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Natural history</div><div class="sidebar-list-content mw-collapsible-content plainlist"><div class="hlist">

<ul><li><a href="Abiogenesis" title="Abiogenesis">Origin of life</a></li>

<li><a href="Common_descent" title="Common descent">Common descent</a></li>

<li><a href="History_of_life" title="History of life">History of life</a></li>

<li><a href="Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of evolution</a></li>

<li><a href="Human_evolution" title="Human evolution">Human evolution</a>

<ul><li><a href="Recent_human_evolution" title="Recent human evolution">Recent human evolution</a></li></ul></li>

<li><a href="Phylogenetic_tree" title="Phylogenetic tree">Phylogeny</a></li>

<li><a href="Biodiversity" title="Biodiversity">Biodiversity</a></li>

<li><a href="Biogeography" title="Biogeography">Biogeography</a></li>

<li><a href="Taxonomy_(biology)" title="Taxonomy (biology)">Classification</a></li>

<li><a href="Evolutionary_taxonomy" title="Evolutionary taxonomy">Evolutionary taxonomy</a></li>

<li><a href="Cladistics" title="Cladistics">Cladistics</a></li>

<li><a href="Transitional_fossil" title="Transitional fossil">Transitional fossil</a></li>

<li><a href="Extinction_event" title="Extinction event">Extinction event</a></li></ul>

</div></div></div></td>

</tr><tr><td class="sidebar-content" style="background:transparent;">

<div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">History of evolutionary theory</div><div class="sidebar-list-content mw-collapsible-content plainlist"><div class="hlist">

<ul><li><a href="History_of_evolutionary_thought" title="History of evolutionary thought">Overview</a></li>

<li><a href="Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance</a></li>

<li><a href="Transmutation_of_species" title="Transmutation of species">Before Darwin</a></li>

<li><a href="Charles_Darwin" title="Charles Darwin">Darwin</a></li>

<li><a href="On_the_Origin_of_Species" title="On the Origin of Species">Origin of Species</a></li>

<li><a href="The_eclipse_of_Darwinism" title="The eclipse of Darwinism">Before synthesis</a></li>

<li><a href="Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li>

<li><a href="History_of_molecular_evolution" title="History of molecular evolution">Molecular evolution</a></li>

<li><a href="Current_research_in_evolutionary_biology" class="mw-redirect" title="Current research in evolutionary biology">Current research</a></li>

<li><a href="History_of_speciation" title="History of speciation">History of speciation</a></li>

<li><a href="History_of_paleontology" title="History of paleontology">History of paleontology</a> (<a href="Timeline_of_paleontology" title="Timeline of paleontology">timeline</a>)</li></ul>

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<div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Fields and applications</div><div class="sidebar-list-content mw-collapsible-content plainlist">

<ul><li><a href="Applications_of_evolution" title="Applications of evolution">Applications of evolution</a></li>

<li><a href="Biosocial_criminology" title="Biosocial criminology">Biosocial criminology</a></li>

<li><a href="Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li>

<li><a href="Evolutionary_aesthetics" title="Evolutionary aesthetics">Evolutionary aesthetics</a></li>

<li><a href="Evolutionary_anthropology" title="Evolutionary anthropology">Evolutionary anthropology</a></li>

<li><a href="Evolutionary_computation" title="Evolutionary computation">Evolutionary computation</a></li>

<li><a href="Evolutionary_ecology" title="Evolutionary ecology">Evolutionary ecology</a></li>

<li><a href="Evolutionary_economics" title="Evolutionary economics">Evolutionary economics</a></li>

<li><a href="Evolutionary_epistemology" title="Evolutionary epistemology">Evolutionary epistemology</a></li>

<li><a href="Evolutionary_ethics" title="Evolutionary ethics">Evolutionary ethics</a></li>

<li><a href="Evolutionary_game_theory" title="Evolutionary game theory">Evolutionary game theory</a></li>

<li><a href="Evolutionary_linguistics" title="Evolutionary linguistics">Evolutionary linguistics</a></li>

<li><a href="Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li>

<li><a href="Evolutionary_neuroscience" title="Evolutionary neuroscience">Evolutionary neuroscience</a></li>

<li><a href="Evolutionary_physiology" title="Evolutionary physiology">Evolutionary physiology</a></li>

<li><a href="Evolutionary_psychology" title="Evolutionary psychology">Evolutionary psychology</a></li>

<li><a href="Experimental_evolution" title="Experimental evolution">Experimental evolution</a></li>

<li><a href="Invasion_genetics" title="Invasion genetics">Invasion genetics</a></li>

<li><a href="Island_biogeography" class="mw-redirect" title="Island biogeography">Island biogeography</a></li>

<li><a href="Phylogenetics" title="Phylogenetics">Phylogenetics</a></li>

<li><a href="Paleontology" title="Paleontology">Paleontology</a></li>

<li><a href="Selective_breeding" title="Selective breeding">Selective breeding</a></li>

<li><a href="Laboratory_experiments_of_speciation" title="Laboratory experiments of speciation">Speciation experiments</a></li>

<li><a href="Sociobiology" title="Sociobiology">Sociobiology</a></li>

<li><a href="Systematics" title="Systematics">Systematics</a></li>

<li><a href="Universal_Darwinism" title="Universal Darwinism">Universal Darwinism</a></li></ul></div></div></td>

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<div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Social implications</div><div class="sidebar-list-content mw-collapsible-content plainlist">

<ul><li><a href="Eugenics" title="Eugenics">Eugenics</a></li>

<li><a href="Evolution_as_fact_and_theory" title="Evolution as fact and theory">Evolution as fact and theory</a></li>

<li><a href="Dysgenics" title="Dysgenics">Dysgenics</a></li>

<li><a href="Social_effects_of_evolutionary_theory" title="Social effects of evolutionary theory">Social effects</a></li>

<li><a href="Rejection_of_evolution_by_religious_groups" title="Rejection of evolution by religious groups">Creation–evolution controversy</a></li>

<li><a href="Theistic_evolution" title="Theistic evolution">Theistic evolution</a></li>

<li><a href="Objections_to_evolution" title="Objections to evolution">Objections to evolution</a></li>

<li><a href="Level_of_support_for_evolution" title="Level of support for evolution">Level of support</a></li>

<li><a href="Nature_versus_nurture" title="Nature versus nurture">Nature-nurture controversy</a></li></ul></div></div></td>

</tr><tr><td class="sidebar-below hlist">

<ul><li> <a href="Portal%3AEvolutionary_biology" title="Portal:Evolutionary biology">Evolutionary biology portal</a></li>

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Natural selection is the differential survival and reproduction of individuals due to differences in <a href="Phenotype" title="Phenotype">phenotype</a>. It is a key mechanism of <a href="Evolution" title="Evolution">evolution</a>, the change in the <a href="Heredity" title="Heredity">heritable traits</a> characteristic of a <a href="Population" title="Population">population</a> over generations. <a href="Charles_Darwin" title="Charles Darwin">Charles Darwin</a> popularised the term "natural selection", contrasting it with <a href="Selective_breeding" title="Selective breeding">artificial selection</a>, which is intentional, whereas natural selection is not.

<a href="Genetic_diversity" title="Genetic diversity">Variation</a> of traits, both <a href="Genotype" title="Genotype">genotypic</a> and phenotypic, exists within all populations of <a href="Organism" title="Organism">organisms</a>. However, some traits are more likely to facilitate <a href="Survival" title="Survival">survival</a> and <a href="Reproductive_success" title="Reproductive success">reproductive success</a>. Thus, these traits are passed on to the next generation. These traits can also become more <a href="Allele_frequency" title="Allele frequency">common within a population</a> if the environment that favours these traits remains fixed. If new traits become more favoured due to changes in a specific <a href="Ecological_niche" title="Ecological niche">niche</a>, <a href="Microevolution" title="Microevolution">microevolution</a> occurs. If new traits become more favoured due to changes in the broader environment, <a href="Macroevolution" title="Macroevolution">macroevolution</a> occurs. Sometimes, <a href="Speciation" title="Speciation">new species can arise</a> especially if these new traits are radically different from the traits possessed by their predecessors.

The likelihood of these traits being 'selected' and passed down are determined by many factors. Some are likely to be passed down because they <a href="Adaptation" title="Adaptation">adapt</a> well to their environments. Others are passed down because these traits are actively preferred by mating partners, which is known as <a href="Sexual_selection" title="Sexual selection">sexual selection</a>. Female bodies also prefer traits that confer the lowest cost to their reproductive health, which is known as <a href="Fecundity_selection" title="Fecundity selection">fecundity selection</a>.

Natural selection is a cornerstone of modern <a href="Biology" title="Biology">biology</a>. The concept, published by Darwin and <a href="Alfred_Russel_Wallace" title="Alfred Russel Wallace">Alfred Russel Wallace</a> in a <a href="On_the_Tendency_of_Species_to_form_Varieties%3B_and_on_the_Perpetuation_of_Varieties_and_Species_by_Natural_Means_of_Selection" title="On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection">joint presentation of papers in 1858</a>, was elaborated in Darwin's influential 1859 book <a href="On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life</a>. He described natural selection as analogous to artificial selection, a process by which animals and plants with traits considered desirable by human breeders are systematically favoured for reproduction. The concept of natural selection originally developed in the absence of a valid theory of heredity; at the time of Darwin's writing, science had yet to develop modern theories of genetics. The union of traditional <a href="Darwinism" title="Darwinism">Darwinian evolution</a> with subsequent discoveries in <a href="Classical_genetics" title="Classical genetics">classical genetics</a> formed the <a href="Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">modern synthesis of the mid-20th century</a>. The addition of <a href="Molecular_genetics" title="Molecular genetics">molecular genetics</a> has led to <a href="Evolutionary_developmental_biology" title="Evolutionary developmental biology">evolutionary developmental biology</a>, which explains evolution at the molecular level. While genotypes can slowly change by random <a href="Genetic_drift" title="Genetic drift">genetic drift</a>, natural selection remains the primary explanation for <a href="Adaptation" title="Adaptation">adaptive evolution</a>.

<div class="mw-heading mw-heading2"><h2 id="Historical_development">Historical development</h2></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="History_of_evolutionary_thought" title="History of evolutionary thought">History of evolutionary thought</a></div>

<div class="mw-heading mw-heading3"><h3 id="Pre-Darwinian_theories">Pre-Darwinian theories</h3></div>

Several philosophers of the <a href="Classical_era" class="mw-redirect" title="Classical era">classical era</a>, including <a href="Empedocles" title="Empedocles">Empedocles</a><a href="#cite_note-1">[1]</a> and his intellectual successor, the <a href="Roman_Republic" title="Roman Republic">Roman</a> poet <a href="Lucretius" title="Lucretius">Lucretius</a>,<a href="#cite_note-2">[2]</a> expressed the idea that nature produces a huge variety of creatures, randomly, and that only those creatures that manage to provide for themselves and reproduce successfully persist. Empedocles' idea that organisms arose entirely by the incidental workings of causes such as heat and cold was criticised by <a href="Aristotle" title="Aristotle">Aristotle</a> in Book II of <a href="Physics_(Aristotle)" title="Physics (Aristotle)">Physics</a>.<a href="#cite_note-3">[3]</a> He posited natural <a href="Teleology" title="Teleology">teleology</a> in its place, and believed that form was achieved for a purpose, citing the regularity of heredity in species as proof.<a href="#cite_note-4">[4]</a><a href="#cite_note-henry-5">[5]</a> Nevertheless, he accepted <a href="Aristotle's_biology" title="Aristotle's biology">in his biology</a> that new types of animals, <a href="Congenital_disorder" class="mw-redirect" title="Congenital disorder">monstrosities</a> (τερας), can occur in very rare instances (<a href="Generation_of_Animals" title="Generation of Animals">Generation of Animals</a>, Book IV).<a href="#cite_note-6">[6]</a> As quoted in Darwin's 1872 edition of <a href="The_Origin_of_Species" class="mw-redirect" title="The Origin of Species">The Origin of Species</a>, Aristotle considered whether different forms (e.g., of teeth) might have appeared accidentally, but only the useful forms survived:

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</style><blockquote class="templatequote">So what hinders the different parts [of the body] from having this merely accidental relation in nature? as the teeth, for example, grow by necessity, the front ones sharp, adapted for dividing, and the grinders flat, and serviceable for masticating the food; since they were not made for the sake of this, but it was the result of accident. And in like manner as to the other parts in which there appears to exist an adaptation to an end. Wheresoever, therefore, all things together (that is all the parts of one whole) happened like as if they were made for the sake of something, these were preserved, having been appropriately constituted by an internal spontaneity, and whatsoever things were not thus constituted, perished, and still perish.</blockquote><div class="templatequotecite">— Aristotle, Physics, Book II, Chapter 8<a href="#cite_note-7">[7]</a></div>

But Aristotle rejected this possibility in the next paragraph, making clear that he is talking about the <a href="Developmental_biology" title="Developmental biology">development of animals as embryos</a> with the phrase "either invariably or normally come about", not the origin of species:

<blockquote class="templatequote">... Yet it is impossible that this should be the true view. For teeth and all other natural things either invariably or normally come about in a given way; but of not one of the results of chance or spontaneity is this true. We do not ascribe to chance or mere coincidence the frequency of rain in winter, but frequent rain in summer we do; nor heat in the dog-days, but only if we have it in winter. If then, it is agreed that things are either the result of coincidence or for an end, and these cannot be the result of coincidence or spontaneity, it follows that they must be for an end; and that such things are all due to nature even the champions of the theory which is before us would agree. Therefore action for an end is present in things which come to be and are by nature.</blockquote><div class="templatequotecite">— Aristotle, Physics, Book II, Chapter 8<a href="#cite_note-8">[8]</a></div>

The <a href="Struggle_for_existence#Historical_development" title="Struggle for existence">struggle for existence</a> was later described by the <a href="Islam" title="Islam">Islamic</a> writer <a href="Al-Jahiz" title="Al-Jahiz">Al-Jahiz</a> in the 9th century, particularly in the context of top-down population regulation, but not in reference to individual variation or natural selection.<a href="#cite_note-9">[9]</a><a href="#cite_note-10">[10]</a>

At the turn of the 16th century <a href="Leonardo_da_Vinci" title="Leonardo da Vinci">Leonardo da Vinci</a> collected a set of fossils of ammonites as well as other biological material. He extensively reasoned in his writings that the shapes of animals are not given once and forever by the "upper power" but instead are generated in different forms naturally and then selected for reproduction by their compatibility with the environment.<a href="#cite_note-11">[11]</a>

The more recent classical arguments were reintroduced in the 18th century by <a href="Pierre_Louis_Maupertuis" title="Pierre Louis Maupertuis">Pierre Louis Maupertuis</a><a href="#cite_note-12">[12]</a> and others, including Darwin's grandfather, <a href="Erasmus_Darwin" title="Erasmus Darwin">Erasmus Darwin</a>.

Until the early 19th century, the <a href="History_of_creationism#Renaissance_to_Darwin" title="History of creationism">prevailing view</a> in <a href="Western_world" title="Western world">Western societies</a> was that differences between individuals of a species were uninteresting departures from their <a href="Theory_of_Forms" class="mw-redirect" title="Theory of Forms">Platonic ideals</a> (or <a href="https://en.wiktionary.org/wiki/typus" class="extiw external" title="wikt:typus">typus</a>) of <a href="Baraminology" class="mw-redirect" title="Baraminology">created kinds</a>. However, the theory of <a href="Uniformitarianism" title="Uniformitarianism">uniformitarianism</a> in geology promoted the idea that simple, weak forces could act continuously over long periods of time to produce radical changes in the Earth's landscape. The success of this theory raised awareness of the vast scale of <a href="Geologic_time_scale" title="Geologic time scale">geological time</a> and made plausible the idea that tiny, virtually imperceptible changes in successive generations could produce consequences on the scale of differences between species.<a href="#cite_note-Bowler_2003-13">[13]</a>

The early 19th-century zoologist <a href="Jean-Baptiste_Lamarck" title="Jean-Baptiste Lamarck">Jean-Baptiste Lamarck</a> suggested the <a href="Inheritance_of_acquired_characteristics" class="mw-redirect" title="Inheritance of acquired characteristics">inheritance of acquired characteristics</a> as a mechanism for evolutionary change; adaptive traits acquired by an organism during its lifetime could be inherited by that organism's progeny, eventually causing <a href="Transmutation_of_species" title="Transmutation of species">transmutation of species</a>.<a href="#cite_note-14">[14]</a> This theory, <a href="Lamarckism" title="Lamarckism">Lamarckism</a>, was an influence on the Soviet biologist <a href="Trofim_Lysenko" title="Trofim Lysenko">Trofim Lysenko</a>'s ill-fated antagonism to mainstream genetic theory as late as the mid-20th century.<a href="#cite_note-Joravsky-15">[15]</a>

Between 1835 and 1837, the zoologist <a href="Edward_Blyth#On_natural_selection" title="Edward Blyth">Edward Blyth</a> worked on the area of variation, artificial selection, and how a similar process occurs in nature. Darwin acknowledged Blyth's ideas in the first chapter on variation of On the Origin of Species.<a href="#cite_note-16">[16]</a>

<div class="mw-heading mw-heading3"><h3 id="Darwin's_theory">Darwin's theory</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main articles: <a href="Inception_of_Darwin's_theory" title="Inception of Darwin's theory">Inception of Darwin's theory</a> and <a href="Development_of_Darwin's_theory" title="Development of Darwin's theory">Development of Darwin's theory</a></div>

<div role="note" class="hatnote navigation-not-searchable">Further information: <a href="Coloration_evidence_for_natural_selection" title="Coloration evidence for natural selection">Coloration evidence for natural selection</a></div>

In 1859, Charles Darwin set out his theory of evolution by natural selection as an explanation for <a href="Adaptation" title="Adaptation">adaptation</a> and speciation. He defined natural selection as the "principle by which each slight variation [of a trait], if useful, is preserved".<a href="#cite_note-17">[17]</a> The concept was simple but powerful: individuals best adapted to their environments are more likely to survive and reproduce. As long as there is some variation between them and that variation is <a href="Heritability" title="Heritability">heritable</a>, there will be an inevitable selection of individuals with the most advantageous variations. If the variations are heritable, then differential reproductive success leads to the evolution of particular populations of a species, and populations that evolve to be sufficiently different eventually become different species.<a href="#cite_note-18">[18]</a><a href="#cite_note-19">[19]</a>

Darwin's ideas were inspired by the observations that he had made on the <a href="Second_voyage_of_HMS_Beagle" title="Second voyage of HMS Beagle">second voyage of HMS Beagle</a> (1831–1836), and by the work of a political economist, <a href="Thomas_Robert_Malthus" title="Thomas Robert Malthus">Thomas Robert Malthus</a>, who, in <a href="An_Essay_on_the_Principle_of_Population" title="An Essay on the Principle of Population">An Essay on the Principle of Population</a> (1798), noted that population (if unchecked) <a href="Exponential_growth" title="Exponential growth">increases exponentially</a>, whereas the food supply grows only <a href="Linear_function" title="Linear function">arithmetically</a>; thus, inevitable limitations of resources would have demographic implications, leading to a "struggle for existence".<a href="#cite_note-20">[20]</a> When Darwin read Malthus in 1838 he was already primed by his work as a <a href="Natural_history" title="Natural history">naturalist</a> to appreciate the "struggle for existence" in nature. It struck him that as population outgrew resources, "favourable variations would tend to be preserved, and unfavourable ones to be destroyed. The result of this would be the formation of new species."<a href="#cite_note-auto120-21">[21]</a> Darwin wrote:

<blockquote class="templatequote">If during the long course of ages and under varying conditions of life, organic beings vary at all in the several parts of their organisation, and I think this cannot be disputed; if there be, owing to the high geometrical powers of increase of each species, at some age, season, or year, a severe struggle for life, and this certainly cannot be disputed; then, considering the infinite complexity of the relations of all organic beings to each other and to their conditions of existence, causing an infinite diversity in structure, constitution, and habits, to be advantageous to them, I think it would be a most extraordinary fact if no variation ever had occurred useful to each being's own welfare, in the same way as so many variations have occurred useful to man. But if variations useful to any organic being do occur, assuredly individuals thus characterised will have the best chance of being preserved in the struggle for life; and from the strong principle of inheritance they will tend to produce offspring similarly characterised. This principle of preservation, I have called, for the sake of brevity, Natural Selection.</blockquote><div class="templatequotecite">— Darwin summarising natural selection in the fourth chapter of <a href="On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species</a><a href="#cite_note-22">[22]</a></div>

Once he had this <a href="Hypothesis" title="Hypothesis">hypothesis</a>, Darwin was meticulous about gathering and refining evidence of <a href="Consilience" title="Consilience">consilience</a> to meet standards of <a href="Scientific_method" title="Scientific method">methodology</a> before making his <a href="Scientific_theory" title="Scientific theory">scientific theory</a> public.<a href="#cite_note-Bowler_2003-13">[13]</a> He was in the process of writing his "big book" to present his research when the naturalist <a href="Alfred_Russel_Wallace" title="Alfred Russel Wallace">Alfred Russel Wallace</a> independently conceived of the principle and described it in an essay he sent to Darwin to forward to <a href="Charles_Lyell" title="Charles Lyell">Charles Lyell</a>. Lyell and <a href="Joseph_Dalton_Hooker" title="Joseph Dalton Hooker">Joseph Dalton Hooker</a> decided to present his essay together with unpublished writings that Darwin had sent to fellow naturalists, and <a href="On_the_Tendency_of_Species_to_form_Varieties%3B_and_on_the_Perpetuation_of_Varieties_and_Species_by_Natural_Means_of_Selection" title="On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection">On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection</a> was read to the <a href="Linnean_Society_of_London" title="Linnean Society of London">Linnean Society of London</a> announcing co-discovery of the principle in July 1858.<a href="#cite_note-23">[23]</a> Darwin published a detailed account of his evidence and conclusions in <a href="On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species</a> in 1859. In later editions Darwin acknowledged that earlier writers—like <a href="William_Charles_Wells" title="William Charles Wells">William Charles Wells</a> in 1813,<a href="#cite_note-24">[24]</a> and <a href="Patrick_Matthew" title="Patrick Matthew">Patrick Matthew</a> in 1831—had proposed similar basic ideas.<a href="#cite_note-25">[25]</a> However, they had not developed their ideas, or presented evidence to persuade others that the concept was useful.<a href="#cite_note-Bowler_2003-13">[13]</a>

Darwin thought of natural selection by analogy to how farmers select crops or livestock for breeding, which he called "<a href="Artificial_selection" class="mw-redirect" title="Artificial selection">artificial selection</a>"; in his early manuscripts he referred to a "Nature" which would do the selection. At the time, other mechanisms of evolution such as evolution by genetic drift were not yet explicitly formulated, and Darwin believed that selection was likely only part of the story: "I am convinced that Natural Selection has been the main but not exclusive means of modification."<a href="#cite_note-26">[26]</a> In a letter to Charles Lyell in September 1860, Darwin regretted the use of the term "Natural Selection", preferring the term "Natural Preservation".<a href="#cite_note-27">[27]</a>

For Darwin and his contemporaries, natural selection was in essence synonymous with evolution by natural selection. After the publication of On the Origin of Species,<a href="#cite_note-origin-28">[28]</a> educated people generally accepted that evolution had occurred in some form. However, natural selection remained controversial as a mechanism, partly because it was perceived to be too weak to explain the range of observed characteristics of living organisms, and partly because even supporters of evolution balked at its "unguided" and non-<a href="Orthogenesis" title="Orthogenesis">progressive</a> nature,<a href="#cite_note-29">[29]</a> a response that has been characterised as the single most significant impediment to the idea's acceptance.<a href="#cite_note-30">[30]</a> However, some thinkers enthusiastically embraced natural selection; after reading Darwin, <a href="Herbert_Spencer" title="Herbert Spencer">Herbert Spencer</a> introduced the phrase <a href="Survival_of_the_fittest" title="Survival of the fittest">survival of the fittest</a>, which became a popular summary of the theory.<a href="#cite_note-sotf-31">[31]</a><a href="#cite_note-32">[32]</a> The fifth edition of On the Origin of Species published in 1869 included Spencer's phrase as an alternative to natural selection, with credit given: "But the expression often used by Mr. Herbert Spencer of the Survival of the Fittest is more accurate, and is sometimes equally convenient."<a href="#cite_note-33">[33]</a> Although the phrase is still often used by non-biologists, modern biologists avoid it because it is <a href="Tautology_(rhetoric)" class="mw-redirect" title="Tautology (rhetoric)">tautological</a> if "fittest" is read to mean "functionally superior" and is applied to individuals rather than considered as an averaged quantity over populations.<a href="#cite_note-34">[34]</a>

<div class="mw-heading mw-heading3"><h3 id="The_modern_synthesis">The modern synthesis</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis (20th century)</a></div>

Natural selection relies crucially on the idea of heredity, but developed before the basic concepts of <a href="Genetics" title="Genetics">genetics</a>. Although the <a href="Moravia" title="Moravia">Moravian</a> monk <a href="Gregor_Mendel" title="Gregor Mendel">Gregor Mendel</a>, the father of modern genetics, was a contemporary of Darwin's, his work lay in obscurity, only being rediscovered in 1900.<a href="#cite_note-35">[35]</a> With the early 20th-century integration of evolution with <a href="Mendel's_laws" class="mw-redirect" title="Mendel's laws">Mendel's laws</a> of inheritance, the so-called <a href="Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">modern synthesis</a>, scientists generally came to accept natural selection.<a href="#cite_note-Huxley-36">[36]</a><a href="#cite_note-37">[37]</a> The synthesis grew from advances in different fields. Ronald Fisher developed the required mathematical language and wrote <a href="The_Genetical_Theory_of_Natural_Selection" title="The Genetical Theory of Natural Selection">The Genetical Theory of Natural Selection</a> (1930).<a href="#cite_note-fisher-38">[38]</a> <a href="J._B._S._Haldane" title="J. B. S. Haldane">J. B. S. Haldane</a> introduced the concept of the "cost" of natural selection.<a href="#cite_note-39">[39]</a><a href="#cite_note-40">[40]</a>

<a href="Sewall_Wright" title="Sewall Wright">Sewall Wright</a> elucidated the nature of selection and adaptation.<a href="#cite_note-41">[41]</a>

In his book <a href="Genetics_and_the_Origin_of_Species" title="Genetics and the Origin of Species">Genetics and the Origin of Species</a> (1937), <a href="Theodosius_Dobzhansky" title="Theodosius Dobzhansky">Theodosius Dobzhansky</a> established the idea that mutation, <a href="Mutationism" title="Mutationism">once seen as a rival</a> to selection, actually supplied the raw material for natural selection by creating genetic diversity.<a href="#cite_note-42">[42]</a><a href="#cite_note-43">[43]</a>

<div class="mw-heading mw-heading3"><h3 id="A_second_synthesis">A second synthesis</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Evolutionary_developmental_biology#History" title="Evolutionary developmental biology">Evolutionary developmental biology § History</a></div>

<a href="Ernst_Mayr" title="Ernst Mayr">Ernst Mayr</a> recognised the key importance of <a href="Reproductive_isolation" title="Reproductive isolation">reproductive isolation</a> for speciation in his <a href="Systematics_and_the_Origin_of_Species" title="Systematics and the Origin of Species">Systematics and the Origin of Species</a> (1942).<a href="#cite_note-45">[45]</a>

<a href="W._D._Hamilton" title="W. D. Hamilton">W. D. Hamilton</a> conceived of <a href="Kin_selection" title="Kin selection">kin selection</a> in 1964.<a href="#cite_note-Hamilton-46">[46]</a> This synthesis cemented natural selection as the foundation of evolutionary theory, where it remains today. A second synthesis was brought about at the end of the 20th century by advances in <a href="Molecular_genetics" title="Molecular genetics">molecular genetics</a>, creating the field of <a href="Evolutionary_developmental_biology" title="Evolutionary developmental biology">evolutionary developmental biology</a> ("evo-devo"), which seeks to explain the evolution of <a href="Morphology_(biology)" title="Morphology (biology)">form</a> in terms of the <a href="Gene_regulatory_network" title="Gene regulatory network">genetic regulatory programs</a> which control the development of the embryo at molecular level. Natural selection is here understood to act on embryonic development to change the morphology of the adult body.<a href="#cite_note-Gilbert2003-47">[47]</a><a href="#cite_note-Gilbert1996-48">[48]</a><a href="#cite_note-Müller-49">[49]</a><a href="#cite_note-50">[50]</a>

<div class="mw-heading mw-heading2"><h2 id="Terminology">Terminology</h2></div>

The term natural selection is most often defined to operate on heritable traits, because these directly participate in evolution. However, natural selection is "blind" in the sense that changes in phenotype can give a reproductive advantage regardless of whether or not the trait is heritable. Following Darwin's primary usage, the term is used to refer both to the evolutionary consequence of blind selection and to its mechanisms.<a href="#cite_note-origin-28">[28]</a><a href="#cite_note-fisher-38">[38]</a><a href="#cite_note-nomenclature1-51">[51]</a><a href="#cite_note-52">[52]</a> It is sometimes helpful to explicitly distinguish between selection's mechanisms and its effects; when this distinction is important, scientists define "(phenotypic) natural selection" specifically as "those mechanisms that contribute to the selection of individuals that reproduce", without regard to whether the basis of the selection is heritable.<a href="#cite_note-nomenclature2-53">[53]</a><a href="#cite_note-54">[54]</a><a href="#cite_note-55">[55]</a> Traits that cause greater reproductive success of an organism are said to be selected for, while those that reduce success are selected against.<a href="#cite_note-56">[56]</a>

<div class="mw-heading mw-heading2"><h2 id="Mechanism">Mechanism</h2></div>

<div class="mw-heading mw-heading3"><h3 id="Heritable_variation,_differential_reproduction">Heritable variation, differential reproduction</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Genetic_variation" title="Genetic variation">Genetic variation</a></div>

Natural variation occurs among the individuals of any population of organisms. Some differences may improve an individual's chances of surviving and reproducing such that its lifetime reproductive rate is increased, which means that it leaves more offspring. If the traits that give these individuals a reproductive advantage are also <a href="Heritable" class="mw-redirect" title="Heritable">heritable</a>, that is, passed from parent to offspring, then there will be differential reproduction, that is, a slightly higher proportion of fast rabbits or efficient algae in the next generation. Even if the reproductive advantage is very slight, over many generations any advantageous heritable trait becomes dominant in the population. In this way the <a href="Natural_environment" title="Natural environment">natural environment</a> of an organism "selects for" traits that confer a reproductive advantage, causing evolutionary change, as Darwin described.<a href="#cite_note-Michigan-57">[57]</a> This gives the appearance of purpose, but in natural selection there is no intentional choice.<a href="#cite_note-58">[a]</a> Artificial selection is <a href="Teleology" title="Teleology">purposive</a> where natural selection is not, though <a href="Teleology_in_biology" title="Teleology in biology">biologists often use teleological language</a> to describe it.<a href="#cite_note-Stanford-59">[58]</a>

The <a href="Peppered_moth" title="Peppered moth">peppered moth</a> exists in both light and dark colours in Great Britain, but during the <a href="Industrial_Revolution" title="Industrial Revolution">Industrial Revolution</a>, many of the trees on which the moths rested became blackened by <a href="Soot" title="Soot">soot</a>, giving the dark-coloured moths an advantage in hiding from predators. This gave dark-coloured moths a better chance of surviving to produce dark-coloured offspring, and in just fifty years from the first dark moth being caught, nearly all of the moths in industrial <a href="Manchester" title="Manchester">Manchester</a> were dark. The balance was reversed by the effect of the <a href="Clean_Air_Act_1956" title="Clean Air Act 1956">Clean Air Act 1956</a>, and the dark moths became rare again, demonstrating the influence of natural selection on <a href="Peppered_moth_evolution" title="Peppered moth evolution">peppered moth evolution</a>.<a href="#cite_note-60">[59]</a> A recent study, using image analysis and avian vision models, shows that pale individuals more closely match lichen backgrounds than dark morphs and for the first time quantifies the <a href="Camouflage" title="Camouflage">camouflage</a> of moths to <a href="Predation" title="Predation">predation</a> risk.<a href="#cite_note-Walton2018-61">[60]</a> Modern genetic studies show that the switch from light to dark coloration is due to a transposable element insertion into the first intron of the gene cortex.<a href="#cite_note-62">[61]</a>

An example of natural selection in the wild involving a much larger number of genes is given by ash trees in Britain, under selection by an invasive fungus causing <a href="Ash_dieback" class="mw-redirect" title="Ash dieback">ash dieback</a>.<a href="#cite_note-63">[62]</a> This fungus has killed large numbers of ash trees in Europe,<a href="#cite_note-64">[63]</a> and damaged many others, though some trees remain healthy.<a href="#cite_note-65">[64]</a> The genetic basis of health under ash dieback pressure has been shown to be quantitative and highly polygenic.<a href="#cite_note-:0-66">[65]</a> Using genomic prediction models trained on planted trials,<a href="#cite_note-:0-66">[65]</a> geneticists have shown that natural selection is acting on a woodland in Surrey England, causing the new generation of ash trees to be, on average, more genetically resistant to ash dieback than their parents generation.<a href="#cite_note-:1-67">[66]</a> This is due to selection for beneficial gene combinations from among the variation present in the parents.<a href="#cite_note-:1-67">[66]</a>

<div class="mw-heading mw-heading3"><h3 id="Fitness">Fitness</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Fitness_(biology)" title="Fitness (biology)">Fitness (biology)</a></div>

The concept of fitness is central to natural selection. In broad terms, individuals that are more "fit" have better potential for survival, as in the well-known phrase "<a href="Survival_of_the_fittest" title="Survival of the fittest">survival of the fittest</a>", but the precise meaning of the term is much more subtle. Modern evolutionary theory defines fitness not by how long an organism lives, but by how successful it is at reproducing. If an organism lives half as long as others of its species, but has twice as many offspring surviving to adulthood, its genes become more common in the adult population of the next generation. Though natural selection acts on individuals, the effects of chance mean that fitness can only really be defined "on average" for the individuals within a population. The fitness of a particular <a href="Genotype" title="Genotype">genotype</a> corresponds to the average effect on all individuals with that genotype.<a href="#cite_note-Orr2009-68">[67]</a>

A distinction must be made between the concept of "survival of the fittest" and "improvement in fitness". "Survival of the fittest" does not give an "improvement in fitness", it only represents the removal of the less fit variants from a population. A mathematical example of "survival of the fittest" is given by Haldane in his paper "The Cost of Natural Selection".<a href="#cite_note-69">[68]</a> Haldane called this process "substitution" or more commonly in biology, this is called "fixation". This is correctly described by the differential survival and reproduction of individuals due to differences in phenotype. On the other hand, "improvement in fitness" is not dependent on the differential survival and reproduction of individuals due to differences in phenotype, it is dependent on the absolute survival of the particular variant. The probability of a beneficial mutation occurring on some member of a population depends on the total number of replications of that variant. The mathematics of "improvement in fitness was described by Kleinman.<a href="#cite_note-70">[69]</a> An empirical example of "improvement in fitness" is given by the Kishony Mega-plate experiment.<a href="#cite_note-71">[70]</a> In this experiment, "improvement in fitness" depends on the number of replications of the particular variant for a new variant to appear that is capable of growing in the next higher drug concentration region. Fixation or substitution is not required for this "improvement in fitness". On the other hand, "improvement in fitness" can occur in an environment where "survival of the fittest" is also acting. <a href="Richard_Lenski" title="Richard Lenski">Richard Lenski</a>'s classic <a href="E._coli_long-term_evolution_experiment" title="E. coli long-term evolution experiment">E. coli long-term evolution experiment</a> is an example of adaptation in a competitive environment, ("improvement in fitness" during "survival of the fittest").<a href="#cite_note-pmid18524956-72">[71]</a> The probability of a beneficial mutation occurring on some member of the lineage to give improved fitness is slowed by the competition. The variant which is a candidate for a beneficial mutation in this limited carrying capacity environment must first out-compete the "less fit" variants in order to accumulate the requisite number of replications for there to be a reasonable probability of that beneficial mutation occurring.<a href="#cite_note-73">[72]</a>

<div class="mw-heading mw-heading3"><h3 id="Competition">Competition</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Competition_(biology)" title="Competition (biology)">Competition (biology)</a></div>

In biology, competition is an interaction between organisms in which the fitness of one is lowered by the presence of another. This may be because both rely on a <a href="Limiting_factor" title="Limiting factor">limited</a> supply of a resource such as food, water, or <a href="Territory_(animal)" title="Territory (animal)">territory</a>.<a href="#cite_note-74">[73]</a> Competition may be <a href="Intraspecific_competition" title="Intraspecific competition">within</a> or <a href="Interspecific_competition" title="Interspecific competition">between species</a>, and may be direct or indirect.<a href="#cite_note-SahneyBentonFerry2010LinksDiversityVertebrates-75">[74]</a> Species less suited to compete should <a href="Competitive_exclusion_principle" title="Competitive exclusion principle">in theory either adapt or die out</a>, since competition plays a powerful role in natural selection, but according to the "room to roam" theory it may be less important than expansion among larger <a href="Clade" title="Clade">clades</a>.<a href="#cite_note-SahneyBentonFerry2010LinksDiversityVertebrates-75">[74]</a><a href="#cite_note-Jardine2012-76">[75]</a>

Competition is modelled by <a href="R/K_selection_theory" title="R/K selection theory">r/K selection theory</a>, which is based on <a href="Robert_MacArthur" class="mw-redirect" title="Robert MacArthur">Robert MacArthur</a> and <a href="E._O._Wilson" title="E. O. Wilson">E. O. Wilson</a>'s work on <a href="Insular_biogeography" title="Insular biogeography">island biogeography</a>.<a href="#cite_note-77">[76]</a> In this theory, selective pressures drive evolution in one of two stereotyped directions: r- or K-selection.<a href="#cite_note-78">[77]</a> These terms, r and K, can be illustrated in a <a href="Logistic_function#In_ecology:_modeling_population_growth" title="Logistic function">logistic model</a> of <a href="Population_dynamics" title="Population dynamics">population dynamics</a>:<a href="#cite_note-Verhulst-79">[78]</a>

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<mrow>

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</mrow>

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<annotation encoding="application/x-tex">{\displaystyle {\frac {dN}{dt}}=rN\left(1-{\frac {N}{K}}\right)\qquad \!}</annotation>

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</math>

where r is the <a href="Population_growth#Population_growth_rate" title="Population growth">growth rate</a> of the population (N), and K is the <a href="Carrying_capacity" title="Carrying capacity">carrying capacity</a> of its local environmental setting. Typically, r-selected species exploit empty <a href="Ecological_niche" title="Ecological niche">niches</a>, and produce many offspring, each with a relatively low <a href="Probability" title="Probability">probability</a> of surviving to adulthood. In contrast, K-selected species are strong competitors in crowded niches, and <a href="Parental_investment" title="Parental investment">invest</a> more heavily in much fewer offspring, each with a relatively high probability of surviving to adulthood.<a href="#cite_note-Verhulst-79">[78]</a>

<div class="mw-heading mw-heading2"><h2 id="Classification">Classification</h2></div>

Natural selection can act on any heritable <a href="Phenotypic_trait" title="Phenotypic trait">phenotypic trait</a>,<a href="#cite_note-80">[79]</a> and selective pressure can be produced by any aspect of the environment, including sexual selection and <a href="Competition_(biology)" title="Competition (biology)">competition</a> with members of the same or other species.<a href="#cite_note-81">[80]</a><a href="#cite_note-ArnqvistRowe2005-82">[81]</a> However, this does not imply that natural selection is always directional and results in adaptive evolution; natural selection often results in the maintenance of the status quo by eliminating less fit variants.<a href="#cite_note-Michigan-57">[57]</a>

Selection can be classified in several different ways, such as by its effect on a trait, on genetic diversity, by the life cycle stage where it acts, by the unit of selection, or by the resource being competed for.

<div class="mw-heading mw-heading3"><h3 id="By_effect_on_a_trait">By effect on a trait</h3></div>

Selection has different effects on traits. <a href="Stabilizing_selection" title="Stabilizing selection">Stabilizing selection</a> acts to hold a trait at a stable optimum, and in the simplest case all deviations from this optimum are selectively disadvantageous. <a href="Directional_selection" title="Directional selection">Directional selection</a> favours extreme values of a trait. The uncommon <a href="Disruptive_selection" title="Disruptive selection">disruptive selection</a> also acts during transition periods when the current mode is sub-optimal, but alters the trait in more than one direction. In particular, if the trait is quantitative and <a href="Univariate" title="Univariate">univariate</a> then both higher and lower trait levels are favoured. Disruptive selection can be a precursor to <a href="Speciation" title="Speciation">speciation</a>.<a href="#cite_note-Michigan-57">[57]</a>

<div class="mw-heading mw-heading3"><h3 id="By_effect_on_genetic_diversity">By effect on genetic diversity</h3></div>

Alternatively, selection can be divided according to its effect on <a href="Genetic_diversity" title="Genetic diversity">genetic diversity</a>. <a href="Negative_selection_(natural_selection)" title="Negative selection (natural selection)">Purifying or negative selection</a> acts to remove genetic variation from the population (and is opposed by <a href="Mutation#By_inheritance" title="Mutation">de novo mutation</a>, which introduces new variation.<a href="#cite_note-83">[82]</a><a href="#cite_note-84">[83]</a> In contrast, <a href="Balancing_selection" title="Balancing selection">balancing selection</a> acts to maintain genetic variation in a population, even in the absence of de novo mutation, by negative <a href="Frequency-dependent_selection" title="Frequency-dependent selection">frequency-dependent selection</a>. One mechanism for this is <a href="Heterozygote_advantage" title="Heterozygote advantage">heterozygote advantage</a>, where individuals with two different alleles have a selective advantage over individuals with just one allele. The polymorphism at the human <a href="ABO_blood_group" class="mw-redirect" title="ABO blood group">ABO blood group</a> locus has been explained in this way.<a href="#cite_note-85">[84]</a>

<div class="mw-heading mw-heading3"><h3 id="By_life_cycle_stage">By life cycle stage</h3></div>

Another option is to classify selection by the <a href="Biological_life_cycle" title="Biological life cycle">life cycle</a> stage at which it acts. Some biologists recognise just two types: <a href="#Types_of_selection">viability (or survival) selection</a>, which acts to increase an organism's probability of survival, and fecundity (or fertility or reproductive) selection, which acts to increase the rate of reproduction, given survival. Others split the life cycle into further components of selection. Thus viability and survival selection may be defined separately and respectively as acting to improve the probability of survival before and after reproductive age is reached, while fecundity selection may be split into additional sub-components including sexual selection, gametic selection, acting on <a href="Gamete" title="Gamete">gamete</a> survival, and compatibility selection, acting on <a href="Zygote" title="Zygote">zygote</a> formation.<a href="#cite_note-Christiansen1984-86">[85]</a>

<div class="mw-heading mw-heading3"><h3 id="By_unit_of_selection">By unit of selection</h3></div>

Selection can also be classified by the level or <a href="Unit_of_selection" title="Unit of selection">unit of selection</a>. Individual selection acts on the individual, in the sense that adaptations are "for" the benefit of the individual, and result from selection among individuals. <a href="Gene_selection" class="mw-redirect" title="Gene selection">Gene selection</a> acts directly at the level of the gene. In <a href="Kin_selection" title="Kin selection">kin selection</a> and <a href="Intragenomic_conflict" title="Intragenomic conflict">intragenomic conflict</a>, gene-level selection provides a more apt explanation of the underlying process. <a href="Group_selection" title="Group selection">Group selection</a>, if it occurs, acts on groups of organisms, on the assumption that groups replicate and mutate in an analogous way to genes and individuals. There is an ongoing debate over the degree to which group selection occurs in nature.<a href="#cite_note-87">[86]</a>

<div class="mw-heading mw-heading3"><h3 id="By_resource_being_competed_for">By resource being competed for</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Further information: <a href="Sexual_selection" title="Sexual selection">Sexual selection</a></div>

Finally, selection can be classified according to the <a href="Resource_(biology)" title="Resource (biology)">resource</a> being competed for. Sexual selection results from competition for mates. Sexual selection typically proceeds via fecundity selection, sometimes at the expense of viability. <a href="Ecological_selection" class="mw-redirect" title="Ecological selection">Ecological selection</a> is natural selection via any means other than sexual selection, such as kin selection, competition, and <a href="Infanticide_(zoology)" title="Infanticide (zoology)">infanticide</a>. Following Darwin, natural selection is sometimes defined as ecological selection,<a href="#cite_note-Blute_2019-90">[89]</a> in which case sexual selection is considered a separate mechanism.<a href="#cite_note-91">[90]</a>

Sexual selection as first articulated by Darwin (using the example of the <a href="Peafowl" title="Peafowl">peacock</a>'s tail)<a href="#cite_note-DarwinSexualSelection-88">[87]</a> refers specifically to competition for mates,<a href="#cite_note-92">[91]</a> which can be intrasexual, between individuals of the same sex, that is male–male competition, or intersexual, where one gender <a href="Mate_choice" title="Mate choice">chooses mates</a>, most often with males displaying and females choosing.<a href="#cite_note-Hosken2011-93">[92]</a> However, in some species, mate choice is primarily by males, as in some fishes of the family <a href="Syngnathidae" title="Syngnathidae">Syngnathidae</a>.<a href="#cite_note-Eens-94">[93]</a><a href="#cite_note-Barlow-95">[94]</a>

Phenotypic traits can be <a href="Signalling_theory" title="Signalling theory">displayed</a> in one sex and desired in the other sex, causing a <a href="Positive_feedback" title="Positive feedback">positive feedback</a> loop called a <a href="Fisherian_runaway" title="Fisherian runaway">Fisherian runaway</a>, for example, the extravagant plumage of some male birds such as the peacock.<a href="#cite_note-Greenfield-89">[88]</a> An alternate theory proposed by the same <a href="Ronald_Fisher" title="Ronald Fisher">Ronald Fisher</a> in 1930 is the <a href="Sexy_son_hypothesis" title="Sexy son hypothesis">sexy son hypothesis</a>, that mothers want promiscuous sons to give them large numbers of grandchildren and so choose promiscuous fathers for their children. Aggression between members of the same sex is sometimes associated with very distinctive features, such as the antlers of <a href="Deer" title="Deer">stags</a>, which are used in combat with other stags. More generally, intrasexual selection is often associated with <a href="Sexual_dimorphism" title="Sexual dimorphism">sexual dimorphism</a>, including differences in body size between males and females of a species.<a href="#cite_note-Hosken2011-93">[92]</a>

<div class="mw-heading mw-heading2"><h2 id="Arms_races">Arms races</h2></div>

<div role="note" class="hatnote navigation-not-searchable">Further information: <a href="Antimicrobial_resistance" title="Antimicrobial resistance">Antimicrobial resistance</a></div>

Natural selection is seen in action in the development of <a href="Antimicrobial_resistance" title="Antimicrobial resistance">antibiotic resistance</a> in <a href="Microorganism" title="Microorganism">microorganisms</a>. Since the discovery of <a href="Penicillin" title="Penicillin">penicillin</a> in 1928, <a href="Antibiotics" class="mw-redirect" title="Antibiotics">antibiotics</a> have been used to fight bacterial diseases. The widespread misuse of antibiotics has selected for microbial resistance to antibiotics in clinical use, to the point that the <a href="Methicillin-resistant_Staphylococcus_aureus" title="Methicillin-resistant Staphylococcus aureus">methicillin-resistant Staphylococcus aureus</a> (MRSA) has been described as a "superbug" because of the threat it poses to health and its relative invulnerability to existing drugs.<a href="#cite_note-96">[95]</a> Response strategies typically include the use of different, stronger antibiotics; however, new <a href="Strain_(biology)" title="Strain (biology)">strains</a> of MRSA have recently emerged that are resistant even to these drugs.<a href="#cite_note-Schito_2006-97">[96]</a> This is an <a href="Evolutionary_arms_race" title="Evolutionary arms race">evolutionary arms race</a>, in which bacteria develop strains less susceptible to antibiotics, while medical researchers attempt to develop new antibiotics that can kill them. A similar situation occurs with <a href="Pesticide_resistance" title="Pesticide resistance">pesticide resistance</a> in plants and insects. Arms races are not necessarily induced by man; a well-documented example involves the spread of a gene in the butterfly <a href="Hypolimnas_bolina" title="Hypolimnas bolina">Hypolimnas bolina</a> suppressing male-killing activity by <a href="Wolbachia" title="Wolbachia">Wolbachia</a> bacteria parasites on the island of <a href="Samoa" title="Samoa">Samoa</a>, where the spread of the gene is known to have occurred over a period of just five years.<a href="#cite_note-98">[97]</a><a href="#cite_note-99">[98]</a>

<div class="mw-heading mw-heading2"><h2 id="Evolution_by_means_of_natural_selection">Evolution by means of natural selection</h2></div>

<div role="note" class="hatnote navigation-not-searchable">Main articles: <a href="Evolution" title="Evolution">Evolution</a> and <a href="Darwinism" title="Darwinism">Darwinism</a></div>

A prerequisite for natural selection to result in adaptive evolution, novel traits and speciation is the presence of heritable genetic variation that results in fitness differences. Genetic variation is the result of mutations, <a href="Genetic_recombination" title="Genetic recombination">genetic recombinations</a> and alterations in the <a href="Karyotype" title="Karyotype">karyotype</a> (the number, shape, size and internal arrangement of the <a href="Chromosome" title="Chromosome">chromosomes</a>). Any of these changes might have an effect that is highly advantageous or highly disadvantageous, but large effects are rare. In the past, most changes in the genetic material were considered neutral or close to neutral because they occurred in <a href="Noncoding_DNA" class="mw-redirect" title="Noncoding DNA">noncoding DNA</a> or resulted in a <a href="Synonymous_substitution" title="Synonymous substitution">synonymous substitution</a>. However, many mutations in <a href="Non-coding_DNA" title="Non-coding DNA">non-coding DNA</a> have deleterious effects.<a href="#cite_note-NCFitnessEffects-100">[99]</a><a href="#cite_note-NCFitnessEffects2-101">[100]</a> Although both mutation rates and average fitness effects of mutations are dependent on the organism, a majority of mutations in humans are slightly deleterious.<a href="#cite_note-Eyre-Walker-102">[101]</a>

Some mutations occur in <a href="Evo-devo_gene_toolkit" title="Evo-devo gene toolkit">"toolkit" or regulatory genes</a>. Changes in these often have large effects on the phenotype of the individual because they regulate the function of many other genes. Most, but not all, mutations in regulatory genes result in non-viable embryos. Some nonlethal regulatory mutations occur in <a href="Homeobox#Hox_genes" title="Homeobox">HOX genes</a> in humans, which can result in a <a href="Cervical_rib" title="Cervical rib">cervical rib</a><a href="#cite_note-103">[102]</a> or <a href="Polydactyly" title="Polydactyly">polydactyly</a>, an increase in the number of fingers or toes.<a href="#cite_note-104">[103]</a> When such mutations result in a higher fitness, natural selection favours these phenotypes and the novel trait spreads in the population.

Established traits are not immutable; traits that have high fitness in one environmental context may be much less fit if environmental conditions change. In the absence of natural selection to preserve such a trait, it becomes more variable and deteriorate over time, possibly resulting in a <a href="Vestigiality" title="Vestigiality">vestigial</a> manifestation of the trait, also called <a href="Evolutionary_baggage" title="Evolutionary baggage">evolutionary baggage</a>. In many circumstances, the apparently vestigial structure may retain a limited functionality, or may be co-opted for other advantageous traits in a phenomenon known as <a href="Exaptation" title="Exaptation">preadaptation</a>. A famous example of a vestigial structure, the eye of the <a href="Spalax" title="Spalax">blind mole-rat</a>, is believed to retain function in <a href="Photoperiodism" title="Photoperiodism">photoperiod</a> perception.<a href="#cite_note-Sanyal-105">[104]</a>

<div class="mw-heading mw-heading3"><h3 id="Speciation">Speciation</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Speciation" title="Speciation">Speciation</a></div>

Speciation requires a degree of <a href="Reproductive_isolation" title="Reproductive isolation">reproductive isolation</a>—that is, a reduction in gene flow. However, it is intrinsic to the concept of a <a href="Species" title="Species">species</a> that <a href="Hybrid_(biology)" title="Hybrid (biology)">hybrids</a> are selected against, opposing the evolution of reproductive isolation, a problem that was recognised by Darwin. The problem does not occur in <a href="Allopatric" class="mw-redirect" title="Allopatric">allopatric</a> speciation with geographically separated populations, which can diverge with different sets of mutations. <a href="E._B._Poulton" class="mw-redirect" title="E. B. Poulton">E. B. Poulton</a> realized in 1903 that reproductive isolation could evolve through divergence, if each lineage acquired a different, incompatible allele of the same gene. Selection against the heterozygote would then directly create reproductive isolation, leading to the <a href="Bateson%E2%80%93Dobzhansky%E2%80%93Muller_model" title="Bateson–Dobzhansky–Muller model">Bateson–Dobzhansky–Muller model</a>, further elaborated by <a href="H._Allen_Orr" title="H. Allen Orr">H. Allen Orr</a><a href="#cite_note-106">[105]</a> and <a href="Sergey_Gavrilets" title="Sergey Gavrilets">Sergey Gavrilets</a>.<a href="#cite_note-107">[106]</a> With <a href="Reinforcement_(speciation)" title="Reinforcement (speciation)">reinforcement</a>, however, natural selection can favour an increase in pre-zygotic isolation, influencing the process of speciation directly.<a href="#cite_note-Modes_and_Mechanisms_of_Speciation-108">[107]</a>

<div class="mw-heading mw-heading2"><h2 id="Genetic_basis">Genetic basis</h2></div>

<div class="mw-heading mw-heading3"><h3 id="Genotype_and_phenotype">Genotype and phenotype</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Genotype%E2%80%93phenotype_distinction" title="Genotype–phenotype distinction">Genotype–phenotype distinction</a></div>

Natural selection acts on an organism's phenotype, or physical characteristics. Phenotype is determined by an organism's genetic make-up (genotype) and the environment in which the organism lives. When different organisms in a population possess different versions of a gene for a certain trait, each of these versions is known as an <a href="Allele" title="Allele">allele</a>. It is this genetic variation that underlies differences in phenotype. An example is the <a href="ABO" class="mw-redirect" title="ABO">ABO</a> blood type <a href="Antigen" title="Antigen">antigens</a> in humans, where three alleles govern the phenotype.<a href="#cite_note-109">[108]</a>

Some traits are governed by only a single gene, but most traits are influenced by the interactions of many genes. A variation in one of the many genes that contributes to a trait may have only a small effect on the phenotype; together, these genes can produce a continuum of possible phenotypic values.<a href="#cite_note-110">[109]</a>

<div class="mw-heading mw-heading3"><h3 id="Directionality_of_selection">Directionality of selection</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Directional_selection" title="Directional selection">Directional selection</a></div>

When some component of a trait is heritable, selection alters the frequencies of the different alleles, or variants of the gene that produces the variants of the trait. Selection can be divided into three classes, on the basis of its effect on allele frequencies: <a href="Directional_selection" title="Directional selection">directional</a>, <a href="Stabilizing_selection" title="Stabilizing selection">stabilizing</a>, and <a href="Disruptive_selection" title="Disruptive selection">disruptive selection</a>.<a href="#cite_note-Rice-111">[110]</a> Directional selection occurs when an allele has a greater fitness than others, so that it increases in frequency, gaining an increasing share in the population. This process can continue until the allele is <a href="Fixation_(population_genetics)" title="Fixation (population genetics)">fixed</a> and the entire population shares the fitter phenotype.<a href="#cite_note-112">[111]</a> Far more common is stabilizing selection, which lowers the frequency of alleles that have a deleterious effect on the phenotype—that is, produce organisms of lower fitness. This process can continue until the allele is eliminated from the population. Stabilizing selection <a href="Conserved_sequence" title="Conserved sequence">conserves</a> functional genetic features, such as <a href="Protein_biosynthesis" title="Protein biosynthesis">protein-coding genes</a> or <a href="Regulatory_sequence" title="Regulatory sequence">regulatory sequences</a>, over time by selective pressure against deleterious variants.<a href="#cite_note-113">[112]</a> Disruptive (or diversifying) selection is selection favouring extreme trait values over intermediate trait values. Disruptive selection may cause <a href="Sympatric_speciation" title="Sympatric speciation">sympatric speciation</a> through <a href="Niche_partitioning" class="mw-redirect" title="Niche partitioning">niche partitioning</a>.

Some forms of <a href="Balancing_selection" title="Balancing selection">balancing selection</a> do not result in fixation, but maintain an allele at intermediate frequencies in a population. This can occur in <a href="Diploid" class="mw-redirect" title="Diploid">diploid</a> species (with pairs of chromosomes) when <a href="Zygosity#Heterozygous" title="Zygosity">heterozygous</a> individuals (with just one copy of the allele) have a higher fitness than homozygous individuals (with two copies). This is called heterozygote advantage or over-dominance, of which the best-known example is the resistance to malaria in humans heterozygous for <a href="Sickle-cell_anaemia" class="mw-redirect" title="Sickle-cell anaemia">sickle-cell anaemia</a>. Maintenance of allelic variation can also occur through <a href="Disruptive_selection" title="Disruptive selection">disruptive or diversifying selection</a>, which favours genotypes that depart from the average in either direction (that is, the opposite of over-dominance), and can result in a <a href="Multimodal_distribution" title="Multimodal distribution">bimodal distribution</a> of trait values. Finally, balancing selection can occur through frequency-dependent selection, where the fitness of one particular phenotype depends on the distribution of other phenotypes in the population. The principles of <a href="Game_theory" title="Game theory">game theory</a> have been applied to understand the fitness distributions in these situations, particularly in the study of kin selection and the evolution of <a href="Reciprocal_altruism" title="Reciprocal altruism">reciprocal altruism</a>.<a href="#cite_note-Hamilton-46">[46]</a><a href="#cite_note-Trivers-114">[113]</a>

<div class="mw-heading mw-heading3"><h3 id="Selection,_genetic_variation,_and_drift">Selection, genetic variation, and drift</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main articles: <a href="Genetic_variation" title="Genetic variation">Genetic variation</a> and <a href="Genetic_drift" title="Genetic drift">Genetic drift</a></div>

A portion of all genetic variation is functionally neutral, producing no phenotypic effect or significant difference in fitness. <a href="Motoo_Kimura" title="Motoo Kimura">Motoo Kimura</a>'s <a href="Neutral_theory_of_molecular_evolution" title="Neutral theory of molecular evolution">neutral theory of molecular evolution</a> by <a href="Genetic_drift" title="Genetic drift">genetic drift</a> proposes that this variation accounts for a large fraction of observed genetic diversity.<a href="#cite_note-Kimura-115">[114]</a> Neutral events can radically reduce genetic variation through <a href="Population_bottleneck" title="Population bottleneck">population bottlenecks</a>,<a href="#cite_note-116">[115]</a> which among other things can cause the <a href="Founder_effect" title="Founder effect">founder effect</a> in initially small new populations.<a href="#cite_note-Campbell1996-117">[116]</a> When genetic variation does not result in differences in fitness, selection cannot directly affect the frequency of such variation. As a result, the genetic variation at those sites is higher than at sites where variation does influence fitness.<a href="#cite_note-Rice-111">[110]</a> However, after a period with no new mutations, the genetic variation at these sites is eliminated due to genetic drift. Natural selection reduces genetic variation by eliminating maladapted individuals, and consequently the mutations that caused the maladaptation. At the same time, new mutations occur, resulting in a <a href="Mutation%E2%80%93selection_balance" title="Mutation–selection balance">mutation–selection balance</a>. The exact outcome of the two processes depends both on the rate at which new mutations occur and on the strength of the natural selection, which is a function of how unfavourable the mutation proves to be.<a href="#cite_note-Lynch-118">[117]</a>

<a href="Genetic_linkage" title="Genetic linkage">Genetic linkage</a> occurs when the <a href="Locus_(genetics)" title="Locus (genetics)">loci</a> of two alleles are close on a chromosome. During the formation of gametes, recombination reshuffles the alleles. The chance that such a reshuffle occurs between two alleles is inversely related to the distance between them. <a href="Selective_sweep" title="Selective sweep">Selective sweeps</a> occur when an allele becomes more common in a population as a result of positive selection. As the prevalence of one allele increases, closely linked alleles can also become more common by "<a href="Genetic_hitchhiking" title="Genetic hitchhiking">genetic hitchhiking</a>", whether they are neutral or even slightly deleterious. A strong selective sweep results in a region of the genome where the positively selected <a href="Haplotype" title="Haplotype">haplotype</a> (the allele and its neighbours) are in essence the only ones that exist in the population. Selective sweeps can be detected by measuring <a href="Linkage_disequilibrium" title="Linkage disequilibrium">linkage disequilibrium</a>, or whether a given haplotype is overrepresented in the population. Since a selective sweep also results in selection of neighbouring alleles, the presence of a block of strong linkage disequilibrium might indicate a 'recent' selective sweep near the centre of the block.<a href="#cite_note-MaynardSmithHaigh-119">[118]</a>

<a href="Background_selection" title="Background selection">Background selection</a> is the opposite of a selective sweep. If a specific site experiences strong and persistent purifying selection, linked variation tends to be weeded out along with it, producing a region in the genome of low overall variability. Because background selection is a result of deleterious new mutations, which can occur randomly in any haplotype, it does not produce clear blocks of linkage disequilibrium, although with low recombination it can still lead to slightly negative linkage disequilibrium overall.<a href="#cite_note-Keightley_&_Otto_2006-120">[119]</a>

<div class="mw-heading mw-heading2"><h2 id="Impact">Impact</h2></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Universal_Darwinism" title="Universal Darwinism">Universal Darwinism</a></div>

Darwin's ideas, along with those of <a href="Adam_Smith" title="Adam Smith">Adam Smith</a> and <a href="Karl_Marx" title="Karl Marx">Karl Marx</a>, had a profound influence on 19th century thought, including his radical claim that "elaborately constructed forms, so different from each other, and dependent on each other in so complex a manner" evolved from the simplest forms of life by a few simple principles.<a href="#cite_note-121">[120]</a> This inspired some of Darwin's most ardent supporters—and provoked the strongest opposition. Natural selection had the power, according to <a href="Stephen_Jay_Gould" title="Stephen Jay Gould">Stephen Jay Gould</a>, to "dethrone some of the deepest and most traditional comforts of Western thought", such as the belief that humans have a special place in the world.<a href="#cite_note-122">[121]</a>

In the words of the philosopher <a href="Daniel_Dennett" title="Daniel Dennett">Daniel Dennett</a>, "Darwin's dangerous idea" of evolution by natural selection is a "universal acid," which cannot be kept restricted to any vessel or container, as it soon leaks out, working its way into ever-wider surroundings.<a href="#cite_note-123">[122]</a> Thus, in the last decades, the concept of natural selection has spread from <a href="Evolutionary_biology" title="Evolutionary biology">evolutionary biology</a> to other disciplines, including <a href="Evolutionary_computation" title="Evolutionary computation">evolutionary computation</a>, <a href="Quantum_Darwinism" title="Quantum Darwinism">quantum Darwinism</a>, <a href="Evolutionary_economics" title="Evolutionary economics">evolutionary economics</a>, <a href="Evolutionary_epistemology" title="Evolutionary epistemology">evolutionary epistemology</a>, <a href="Evolutionary_psychology" title="Evolutionary psychology">evolutionary psychology</a>, and <a href="Lee_Smolin#Cosmological_natural_selection" title="Lee Smolin">cosmological natural selection</a>. This unlimited applicability has been called <a href="Universal_Darwinism" title="Universal Darwinism">universal Darwinism</a>.<a href="#cite_note-124">[123]</a>

<div class="mw-heading mw-heading3"><h3 id="Origin_of_life">Origin of life</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Abiogenesis" title="Abiogenesis">Abiogenesis</a></div>

How life originated from inorganic matter remains an unresolved problem in biology. One prominent hypothesis is that life first appeared <a href="RNA_world" title="RNA world">in the form of short self-replicating RNA</a> polymers.<a href="#cite_note-125">[124]</a> On this view, life may have come into existence when <a href="RNA" title="RNA">RNA</a> chains first experienced the basic conditions, as conceived by Charles Darwin, for natural selection to operate. These conditions are: heritability, <a href="Genetic_variability" title="Genetic variability">variation of type</a>, and competition for limited resources. The fitness of an early <a href="RNA_world" title="RNA world">RNA replicator</a> would likely have been a function of adaptive capacities that were intrinsic (i.e., determined by the <a href="Nucleic_acid_sequence" title="Nucleic acid sequence">nucleotide sequence</a>) and the availability of resources.<a href="#cite_note-Bernstein-126">[125]</a><a href="#cite_note-Michod-127">[126]</a> The three primary adaptive capacities could logically have been: (1) the capacity to replicate with moderate fidelity (giving rise to both heritability and variation of type), (2) the capacity to avoid decay, and (3) the capacity to acquire and process resources.<a href="#cite_note-Bernstein-126">[125]</a><a href="#cite_note-Michod-127">[126]</a> These capacities would have been determined initially by the folded configurations (including those configurations with <a href="Ribozyme" title="Ribozyme">ribozyme</a> activity) of the RNA replicators that, in turn, would have been encoded in their individual nucleotide sequences.<a href="#cite_note-128">[127]</a>

<div class="mw-heading mw-heading3"><h3 id="Cell_and_molecular_biology">Cell and molecular biology</h3></div>

In 1881, the embryologist <a href="Wilhelm_Roux" title="Wilhelm Roux">Wilhelm Roux</a> published Der Kampf der Theile im Organismus (The Struggle of Parts in the Organism) in which he suggested that the development of an organism results from a Darwinian competition between the parts of the embryo, occurring at all levels, from molecules to organs.<a href="#cite_note-129">[128]</a> In recent years, a modern version of this theory has been proposed by <a href="https://fr.wikipedia.org/wiki/Jean-Jacques_Kupiec" class="extiw external" title="fr:Jean-Jacques Kupiec">Jean-Jacques Kupiec</a>. According to this cellular Darwinism, <a href="Stochastic" title="Stochastic">random variation</a> at the molecular level generates diversity in cell types whereas cell interactions impose a characteristic order on the developing embryo.<a href="#cite_note-130">[129]</a>

<div class="mw-heading mw-heading3"><h3 id="Social_and_psychological_theory">Social and psychological theory</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Evolutionary_psychology" title="Evolutionary psychology">Evolutionary psychology</a></div>

The social implications of the theory of evolution by natural selection also became the source of continuing controversy. <a href="Friedrich_Engels" title="Friedrich Engels">Friedrich Engels</a>, a German <a href="Political_philosophy" title="Political philosophy">political philosopher</a> and co-originator of the ideology of <a href="Communism" title="Communism">communism</a>, wrote in 1872 that "Darwin did not know what a bitter satire he wrote on mankind, and especially on his countrymen, when he showed that free competition, the struggle for existence, which the economists celebrate as the highest historical achievement, is the normal state of the animal kingdom."<a href="#cite_note-131">[130]</a> Herbert Spencer and the eugenics advocate <a href="Francis_Galton" title="Francis Galton">Francis Galton</a>'s interpretation of natural selection as necessarily progressive, leading to supposed advances in intelligence and civilisation, became a justification for <a href="Colonialism" title="Colonialism">colonialism</a>, <a href="Eugenics" title="Eugenics">eugenics</a>, and <a href="Social_Darwinism" title="Social Darwinism">social Darwinism</a>. For example, in 1940, <a href="Konrad_Lorenz" title="Konrad Lorenz">Konrad Lorenz</a>, in writings that he subsequently disowned, used the theory as a justification for policies of the <a href="Nazi" class="mw-redirect" title="Nazi">Nazi</a> state. He wrote "... selection for toughness, heroism, and social utility ... must be accomplished by some human institution, if mankind, in default of selective factors, is not to be ruined by domestication-induced degeneracy. The racial idea as the basis of our state has already accomplished much in this respect."<a href="#cite_note-132">[131]</a> Others have developed ideas that human societies and culture <a href="Sociocultural_evolution" title="Sociocultural evolution">evolve</a> by mechanisms analogous to those that apply to evolution of species.<a href="#cite_note-133">[132]</a>

More recently, work among anthropologists and psychologists has led to the development of <a href="Sociobiology" title="Sociobiology">sociobiology</a> and later of evolutionary psychology, a field that attempts to explain features of <a href="Psychology" title="Psychology">human psychology</a> in terms of adaptation to the ancestral environment. The most prominent example of evolutionary psychology, notably advanced in the early work of <a href="Noam_Chomsky" title="Noam Chomsky">Noam Chomsky</a> and later by <a href="Steven_Pinker" title="Steven Pinker">Steven Pinker</a>, is the hypothesis that the human brain has adapted to <a href="Language_acquisition" title="Language acquisition">acquire</a> the <a href="Grammar" title="Grammar">grammatical</a> rules of <a href="Natural_language" title="Natural language">natural language</a>.<a href="#cite_note-Pinker-134">[133]</a> Other aspects of human behaviour and social structures, from specific cultural norms such as <a href="Westermarck_effect#Westermarck_effect" title="Westermarck effect">incest avoidance</a> to broader patterns such as <a href="Gender_role" title="Gender role">gender roles</a>, have been hypothesised to have similar origins as adaptations to the early environment in which modern humans evolved. By analogy to the action of natural selection on genes, the concept of <a href="Meme" title="Meme">memes</a>—"units of cultural transmission," or culture's equivalents of genes undergoing selection and recombination—has arisen, first described in this form by <a href="Richard_Dawkins" title="Richard Dawkins">Richard Dawkins</a> in 1976<a href="#cite_note-135">[134]</a> and subsequently expanded upon by philosophers such as <a href="Daniel_Dennett" title="Daniel Dennett">Daniel Dennett</a> as explanations for complex cultural activities, including human <a href="Consciousness" title="Consciousness">consciousness</a>.<a href="#cite_note-136">[135]</a>

<div class="mw-heading mw-heading3"><h3 id="Information_and_systems_theory">Information and systems theory</h3></div>

In 1922, <a href="Alfred_J._Lotka" title="Alfred J. Lotka">Alfred J. Lotka</a> proposed that natural selection might be understood as a physical principle that could be described in terms of the use of energy by a system,<a href="#cite_note-137">[136]</a><a href="#cite_note-138">[137]</a> a concept later developed by <a href="Howard_T._Odum" title="Howard T. Odum">Howard T. Odum</a> as the <a href="Maximum_power_principle" title="Maximum power principle">maximum power principle</a> in <a href="Thermodynamics" title="Thermodynamics">thermodynamics</a>, whereby evolutionary systems with selective advantage maximise the rate of useful energy transformation.<a href="#cite_note-139">[138]</a>

The principles of natural selection have inspired a variety of computational techniques, such as "soft" <a href="Artificial_life" title="Artificial life">artificial life</a>, that simulate selective processes and can be highly efficient in 'adapting' entities to an environment defined by a specified <a href="Fitness_function" title="Fitness function">fitness function</a>.<a href="#cite_note-140">[139]</a> For example, a class of heuristic <a href="Mathematical_optimization" title="Mathematical optimization">optimisation</a> algorithms known as <a href="Genetic_algorithm" title="Genetic algorithm">genetic algorithms</a>, pioneered by <a href="John_Henry_Holland" title="John Henry Holland">John Henry Holland</a> in the 1970s and expanded upon by <a href="David_E._Goldberg" title="David E. Goldberg">David E. Goldberg</a>,<a href="#cite_note-141">[140]</a> identify optimal solutions by simulated reproduction and mutation of a population of solutions defined by an initial <a href="Probability_distribution" title="Probability distribution">probability distribution</a>.<a href="#cite_note-142">[141]</a> Such algorithms are particularly useful when applied to problems whose <a href="Energy_landscape" class="mw-redirect" title="Energy landscape">energy landscape</a> is very rough or has many local minima.<a href="#cite_note-143">[142]</a>

<div class="mw-heading mw-heading3"><h3 id="In_fiction">In fiction</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Evolution_in_fiction" title="Evolution in fiction">Evolution in fiction</a></div>

Darwinian evolution by natural selection is pervasive in literature, whether taken optimistically in terms of how humanity may evolve towards perfection, or pessimistically in terms of the dire consequences of the interaction of human nature and the struggle for survival. Among major responses is <a href="Samuel_Butler_(novelist)" title="Samuel Butler (novelist)">Samuel Butler</a>'s 1872 pessimistic <a href="Erewhon" title="Erewhon">Erewhon</a> ("nowhere", written mostly backwards). In 1893 <a href="H._G._Wells" title="H. G. Wells">H. G. Wells</a> imagined "<a href="The_Man_of_the_Year_Million" class="mw-redirect" title="The Man of the Year Million">The Man of the Year Million</a>", transformed by natural selection into a being with a huge head and eyes, and shrunken body.<a href="#cite_note-SFE-144">[143]</a>

<div class="mw-heading mw-heading2"><h2 id="Notes">Notes</h2></div>

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<li id="cite_note-58"><a href="#cite_ref-58">^</a> In <a href="Sexual_selection" title="Sexual selection">sexual selection</a>, a female animal making a choice of mate may be argued to be intending to get the best mate; there is no suggestion that she has any intention to improve the bloodline in the manner of an animal breeder.

</li>

</ol></div></div>

<div class="mw-heading mw-heading2"><h2 id="References">References</h2></div>

<li id="cite_note-1"><a href="#cite_ref-1">^</a> <a href="#CITEREFEmpedocles1898">Empedocles 1898</a>, <a rel="nofollow" class="external text" href="https://history.hanover.edu/texts/presoc/emp.html#book2">On Nature, Book II</a>

</li>

<li id="cite_note-2"><a href="#cite_ref-2">^</a> <a href="#CITEREFLucretius1916">Lucretius 1916</a>, <a rel="nofollow" class="external text" href="http://classics.mit.edu/Carus/nature_things.5.v.html">On the Nature of Things, Book V</a>

</li>

<li id="cite_note-3"><a href="#cite_ref-3">^</a> <a href="#CITEREFAristotle">Aristotle</a>, <a rel="nofollow" class="external text" href="http://classics.mit.edu/Aristotle/physics.2.ii.html">Physics, Book II, Chapters 4 and 8</a>

</li>

</li>

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</style><cite id="CITEREFHenry2006" class="citation journal cs1">Henry, Devin (September 2006). <a rel="nofollow" class="external text" href="http://works.bepress.com/cgi/viewcontent.cgi?article=1010&context=devinhenry">"Aristotle on the Mechanism of Inheritance"</a>. Journal of the History of Biology. 39 (3): 425–455. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs10739-005-3058-y">10.1007/s10739-005-3058-y</a>. <a href="S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:85671523">85671523</a>.</cite>

</li>

<li id="cite_note-6"><a href="#cite_ref-6">^</a> <a href="#CITEREFAriew2002">Ariew 2002</a>

</li>

<li id="cite_note-7"><a href="#cite_ref-7">^</a> <a href="#CITEREFDarwin1872">Darwin 1872</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?itemID=F391&viewtype=text&pageseq=18">xiii</a>

</li>

<li id="cite_note-8"><a href="#cite_ref-8">^</a> <a href="#CITEREFAristotle">Aristotle</a>, <a rel="nofollow" class="external text" href="http://classics.mit.edu/Aristotle/physics.2.ii.html">Physics, Book II, Chapter 8</a>

</li>

<li id="cite_note-9"><a href="#cite_ref-9">^</a> <cite id="CITEREFZirkle1941" class="citation journal cs1"><a href="Conway_Zirkle" title="Conway Zirkle">Zirkle, Conway</a> (25 April 1941). "Natural Selection before the 'Origin of Species'". <a href="Proceedings_of_the_American_Philosophical_Society" title="Proceedings of the American Philosophical Society">Proceedings of the American Philosophical Society</a>. 84 (1): 71–123. <a href="JSTOR_(identifier)" class="mw-redirect" title="JSTOR (identifier)">JSTOR</a> <a rel="nofollow" class="external text" href="https://www.jstor.org/stable/984852">984852</a>.</cite>

</li>

<li id="cite_note-10"><a href="#cite_ref-10">^</a> <a href="#CITEREFAgutterWheatley2008">Agutter & Wheatley 2008</a>, p. 43

</li>

<li id="cite_note-11"><a href="#cite_ref-11">^</a> <cite class="citation book cs1">Leonardo, Codex C. <a href="Institut_de_France" title="Institut de France">Institut de France</a>. Trans. Richter. 2016.</cite>

</li>

<li id="cite_note-12"><a href="#cite_ref-12">^</a> <cite id="CITEREFMaupertuis1746" class="citation journal cs1 cs1-prop-foreign-lang-source"><a href="Pierre_Louis_Maupertuis" title="Pierre Louis Maupertuis">Maupertuis, Pierre Louis</a> (1746). <a class="external text external" href="https://en.wikisource.org/wiki/fr:Les_Loix_du_mouvement_et_du_repos_d%C3%A9duites_d%27un_principe_metaphysique">"Les Loix du mouvement et du repos déduites d'un principe metaphysique" </a> [<a href="https://en.wikisource.org/wiki/Translation:Derivation_of_the_laws_of_motion_and_equilibrium_from_a_metaphysical_principle#I._Assessment_of_the_Proofs_of_God's_Existence_that_are_Based_on_the_Marvels_of_Nature" class="extiw external" title="s:Translation:Derivation of the laws of motion and equilibrium from a metaphysical principle">"Derivation of the laws of motion and equilibrium from a metaphysical principle"</a>]. Histoire de l'Académie Royale des Sciences et des Belles Lettres (in French). Berlin: 267–294.</cite>

</li>

<li id="cite_note-Bowler_2003-13">^ <a href="#cite_ref-Bowler_2003_13-0">a</a> <a href="#cite_ref-Bowler_2003_13-1">b</a> <a href="#cite_ref-Bowler_2003_13-2">c</a> <cite id="CITEREFBowler2003" class="citation book cs1">Bowler, Peter J. (2003). <a rel="nofollow" class="external text" href="https://archive.org/details/evolutionhistory0000bowl_n7y8/page/129">Evolution: The History of an Idea</a> (3rd ed.). Berkeley, CA: <a href="University_of_California_Press" title="University of California Press">University of California Press</a>. pp. <a rel="nofollow" class="external text" href="https://archive.org/details/evolutionhistory0000bowl_n7y8/page/129">129–134</a>, <a rel="nofollow" class="external text" href="https://archive.org/details/evolutionhistory0000bowl_n7y8/page/158">158</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-520-23693-6</bdi>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/43091892">43091892</a>.</cite>

</li>

<li id="cite_note-14"><a href="#cite_ref-14">^</a> <a href="#CITEREFLamarck1809">Lamarck 1809</a>

</li>

<li id="cite_note-Joravsky-15"><a href="#cite_ref-Joravsky_15-0">^</a> <cite id="CITEREFJoravsky1959" class="citation journal cs1">Joravsky, David (January 1959). "Soviet Marxism and Biology before Lysenko". <a href="Journal_of_the_History_of_Ideas" title="Journal of the History of Ideas">Journal of the History of Ideas</a>. 20 (1): 85–104. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.2307%2F2707968">10.2307/2707968</a>. <a href="JSTOR_(identifier)" class="mw-redirect" title="JSTOR (identifier)">JSTOR</a> <a rel="nofollow" class="external text" href="https://www.jstor.org/stable/2707968">2707968</a>.</cite>

</li>

<li id="cite_note-16"><a href="#cite_ref-16">^</a> <a href="#CITEREFDarwin1859">Darwin 1859</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=33&itemID=F373&viewtype=text">18</a>

</li>

<li id="cite_note-17"><a href="#cite_ref-17">^</a> <a href="#CITEREFDarwin1859">Darwin 1859</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?itemID=F373&viewtype=side&pageseq=76">61</a>

</li>

<li id="cite_note-18"><a href="#cite_ref-18">^</a> <a href="#CITEREFDarwin1859">Darwin 1859</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?itemID=F373&viewtype=text&pageseq=20">5</a>

</li>

<li id="cite_note-19"><a href="#cite_ref-19">^</a> <cite id="CITEREFHall,_Brian_K.Hallgrímsson,_Benedikt2008" class="citation book cs1">Hall, Brian K.; Hallgrímsson, Benedikt (2008). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=jrDD3cyA09kC&pg=PA4">Strickberger's Evolution</a> (4th ed.). Jones and Bartlett. pp. 4–6. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-7637-0066-9</bdi>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/796450355">796450355</a>.</cite>

</li>

<li id="cite_note-20"><a href="#cite_ref-20">^</a> <a href="#CITEREFMalthus1798">Malthus 1798</a>

</li>

<li id="cite_note-auto120-21"><a href="#cite_ref-auto120_21-0">^</a> <a href="#CITEREFDarwin1958">Darwin 1958</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F1497&pageseq=124">120</a>

</li>

<li id="cite_note-22"><a href="#cite_ref-22">^</a> <a href="#CITEREFDarwin1859">Darwin 1859</a>, pp. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=144&itemID=F373&viewtype=side">126–127</a>

</li>

<li id="cite_note-23"><a href="#cite_ref-23">^</a> <a href="#CITEREFWallace1871">Wallace 1871</a>

</li>

<li id="cite_note-24"><a href="#cite_ref-24">^</a> <a href="#CITEREFDarwin1866">Darwin 1866</a>, pp. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F385&pageseq=21">xiv–xv</a>

</li>

<li id="cite_note-25"><a href="#cite_ref-25">^</a> <a href="#CITEREFDarwin1861">Darwin 1861</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?itemID=F381&viewtype=text&pageseq=20">xiii</a>

</li>

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<div class="mw-heading mw-heading3"><h3 id="Sources">Sources</h3></div>

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<li><cite id="CITEREFDarwin1872" class="citation book cs1">Darwin, Charles (1872). <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=1&itemID=F391&viewtype=side">The Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life</a> (6th ed.). London: John Murray. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/1185571">1185571</a>.</cite></li>

<li><cite id="CITEREFDarwin1958" class="citation book cs1">Darwin, Charles (1958). Barlow, Nora (ed.). <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?itemID=F1497&viewtype=side&pageseq=1">The Autobiography of Charles Darwin, 1809–1882: With original omissions restored; Edited and with Appendix and Notes by his grand-daughter, Nora Barlow</a>. London: <a href="William_Collins%2C_Sons" title="William Collins, Sons">Collins</a>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/93017940">93017940</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/869541868">869541868</a>.</cite></li>

<li><cite id="CITEREFDawkins1976" class="citation book cs1"><a href="Richard_Dawkins" title="Richard Dawkins">Dawkins, Richard</a> (1976). <a href="The_Selfish_Gene" title="The Selfish Gene">The Selfish Gene</a>. New York: Oxford University Press. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-19-857519-1</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/76029168">76029168</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/2681149">2681149</a>.</cite></li>

<li><cite id="CITEREFDennett1991" class="citation book cs1"><a href="Daniel_Dennett" title="Daniel Dennett">Dennett, Daniel C.</a> (1991). <a href="Consciousness_Explained" title="Consciousness Explained">Consciousness Explained</a> (1st ed.). Boston, MA: <a href="Little%2C_Brown_and_Company" title="Little, Brown and Company">Little, Brown and Company</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-316-18065-8</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/91015614">91015614</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/23648691">23648691</a>.</cite></li>

<li><cite id="CITEREFDennett1995" class="citation book cs1">Dennett, Daniel C. (1995). <a href="Darwin's_Dangerous_Idea" title="Darwin's Dangerous Idea">Darwin's Dangerous Idea: Evolution and the Meanings of Life</a>. New York: <a href="Simon_%26_Schuster" title="Simon & Schuster">Simon & Schuster</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-684-80290-9</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/94049158">94049158</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/31867409">31867409</a>.</cite></li>

<li><cite id="CITEREFDobzhansky1937" class="citation book cs1"><a href="Theodosius_Dobzhansky" title="Theodosius Dobzhansky">Dobzhansky, Theodosius</a> (1937). <a href="Genetics_and_the_Origin_of_Species" title="Genetics and the Origin of Species">Genetics and the Origin of Species</a>. <a href="Columbia_University_Biological_Series" title="Columbia University Biological Series">Columbia University Biological Series</a>. New York: <a href="Columbia_University_Press" title="Columbia University Press">Columbia University Press</a>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/37033383">37033383</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/766405">766405</a>.</cite>

<ul><li><cite id="CITEREFDobzhansky1951" class="citation book cs1">—— (1951). <a rel="nofollow" class="external text" href="https://archive.org/details/in.ernet.dli.2015.547790">Genetics and the Origin of Species</a>. Columbia University Biological Series (3rd revised ed.). New York: Columbia University Press. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/51014816">51014816</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/295774">295774</a>.</cite></li></ul></li>

<li><cite id="CITEREFEisley1958" class="citation book cs1"><a href="Loren_Eiseley" title="Loren Eiseley">Eisley, Loren</a> (1958). Darwin's Century: Evolution and the Men Who Discovered It (1st ed.). Garden City, NY: <a href="Doubleday_(publisher)" title="Doubleday (publisher)">Doubleday</a>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/58006638">58006638</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/168989">168989</a>.</cite></li>

<li><cite id="CITEREFEmpedocles1898" class="citation book cs1"><a href="Empedocles" title="Empedocles">Empedocles</a> (1898). <a rel="nofollow" class="external text" href="https://archive.org/details/cu31924029013162">"Empedokles"</a>. In <a href="Arthur_Fairbanks" title="Arthur Fairbanks">Fairbanks, Arthur</a> (ed.). The First Philosophers of Greece. Translation by Arthur Fairbanks. London: Kegan Paul, Trench, Trübner & Co. Ltd. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/03031810">03031810</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/1376248">1376248</a>.</cite> <a rel="nofollow" class="external text" href="https://archive.org/details/cu31924029013162">The First Philosophers of Greece</a> at the <a href="Internet_Archive" title="Internet Archive">Internet Archive</a>.</li>

<li><cite id="CITEREFEndler1986" class="citation book cs1"><a href="John_Endler" title="John Endler">Endler, John A.</a> (1986). Natural Selection in the Wild. Princeton, NJ: Princeton University Press. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-691-08386-5</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/85042683">85042683</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/12262762">12262762</a>.</cite></li>

<li><cite id="CITEREFEngels1964" class="citation book cs1"><a href="Friedrich_Engels" title="Friedrich Engels">Engels, Friedrich</a> (1964) [1883]. <a href="Dialectics_of_Nature" title="Dialectics of Nature">Dialectics of Nature</a>. 1939 preface by <a href="J.B.S._Haldane" class="mw-redirect" title="J.B.S. Haldane">J.B.S. Haldane</a> (3rd rev. ed.). Moscow, USSR: <a href="Progress_Publishers" title="Progress Publishers">Progress Publishers</a>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/66044448">66044448</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/807047245">807047245</a>.</cite> The book is available from the <a rel="nofollow" class="external text" href="https://www.marxists.org/archive/marx/works/1883/don/index.htm">Marxist Internet Archive</a>.</li>

<li><cite id="CITEREFFalconerMackay1996" class="citation book cs1"><a href="Douglas_Scott_Falconer" title="Douglas Scott Falconer">Falconer, Douglas S.</a>; Mackay, Trudy F.C. (1996). <a rel="nofollow" class="external text" href="https://archive.org/details/introductiontoqu00falc">Introduction to Quantitative Genetics</a> (4th ed.). Harlow, England: <a href="Longman" title="Longman">Longman</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-582-24302-6</bdi>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/824656731">824656731</a>.</cite></li>

<li><cite id="CITEREFFisher1930" class="citation book cs1"><a href="Ronald_Fisher" title="Ronald Fisher">Fisher, Ronald Aylmer</a> (1930). <a href="The_Genetical_Theory_of_Natural_Selection" title="The Genetical Theory of Natural Selection">The Genetical Theory of Natural Selection</a>. Oxford: <a href="Oxford_University_Press#Clarendon_Press" title="Oxford University Press">The Clarendon Press</a>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/30029177">30029177</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/493745635">493745635</a>.</cite></li>

<li><cite id="CITEREFFutuyma2005" class="citation book cs1"><a href="Douglas_J._Futuyma" title="Douglas J. Futuyma">Futuyma, Douglas J.</a> (2005). <a rel="nofollow" class="external text" href="https://archive.org/details/evolution0000futu">Evolution</a>. Sunderland, MA: <a href="Sinauer_Associates" title="Sinauer Associates">Sinauer Associates</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-87893-187-3</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2004029808">2004029808</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/57311264">57311264</a>.</cite></li>

<li><cite id="CITEREFGoldberg1989" class="citation book cs1"><a href="David_E._Goldberg" title="David E. Goldberg">Goldberg, David E.</a> (1989). Genetic Algorithms in Search, Optimization and Machine Learning. Reading, MA: <a href="Addison-Wesley" title="Addison-Wesley">Addison-Wesley Publishing Company</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-201-15767-3</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/88006276">88006276</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/17674450">17674450</a>.</cite></li>

<li><cite id="CITEREFHaldane1932" class="citation book cs1"><a href="J._B._S._Haldane" title="J. B. S. Haldane">Haldane, J B.S.</a> (1932). <a href="The_Causes_of_Evolution" title="The Causes of Evolution">The Causes of Evolution</a>. London; New York: <a href="Longman" title="Longman">Longmans, Green & Co.</a> <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/32033284">32033284</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/5006266">5006266</a>.</cite> "This book is based on a series of lectures delivered in January 1931 at the Prifysgol Cymru, Aberystwyth, and entitled 'A re-examination of Darwinism'."</li>

<li><cite id="CITEREFHaldane1954" class="citation book cs1 cs1-prop-long-vol"><a href="J._B._S._Haldane" title="J. B. S. Haldane">Haldane, J.B.S.</a> (1954). <a rel="nofollow" class="external text" href="http://wellcomelibrary.org/player/b18033878#?asi=0&ai=494&z=-0.6189%2C-0.0449%2C2.8968%2C1.6356">"The Measurement of Natural Selection"</a>. In Montalenti, Giuseppe; Chiarugi, A. (eds.). Atti del IX Congresso Internazionale di Genetica, Bellagio (Como) 24–31 agosto 1953 [Proceedings of the 9th International Congress of Genetics]. Caryologia. Vol. 6 (1953/54) Suppl. Florence, Italy: <a href="University_of_Florence" title="University of Florence">University of Florence</a>. pp. 480–487. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/9069245">9069245</a>.</cite></li>

<li><cite id="CITEREFKauffman1993" class="citation book cs1"><a href="Stuart_Kauffman" title="Stuart Kauffman">Kauffman, Stuart</a> (1993). The Origins of Order: Self-Organisation and Selection in Evolution. New York: Oxford University Press. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-19-507951-7</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/91011148">91011148</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/23253930">23253930</a>.</cite></li>

<li><cite id="CITEREFLamarck1809" class="citation book cs1"><a href="Jean-Baptiste_Lamarck" title="Jean-Baptiste Lamarck">Lamarck, Jean-Baptiste</a> (1809). <a href="Philosophie_Zoologique" class="mw-redirect" title="Philosophie Zoologique">Philosophie Zoologique</a>. Paris: Dentu et L'Auteur. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/2210044">2210044</a>.</cite> <a rel="nofollow" class="external text" href="https://archive.org/details/philosophiezool06unkngoog">Philosophie zoologique (1809)</a> at the <a href="Internet_Archive" title="Internet Archive">Internet Archive</a>.</li>

<li><cite id="CITEREFLear1988" class="citation book cs1"><a href="Jonathan_Lear" title="Jonathan Lear">Lear, Jonathan</a> (1988). <a rel="nofollow" class="external text" href="https://archive.org/details/aristotledesiret0000lear">Aristotle: The Desire to Understand</a>. Cambridge, UK; New York: <a href="Cambridge_University_Press" title="Cambridge University Press">Cambridge University Press</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-521-34762-4</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/87020284">87020284</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/16352317">16352317</a>.</cite></li>

<li><cite id="CITEREFKuhn1996" class="citation book cs1"><a href="Thomas_Kuhn" title="Thomas Kuhn">Kuhn, Thomas S.</a> (1996). The Structure of Scientific Revolutions (3rd ed.). Chicago, IL: University of Chicago Press. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-226-45808-3</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/96013195">96013195</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/34548541">34548541</a>.</cite></li>

<li><cite id="CITEREFLemeySalemiVandamme2009" class="citation book cs1">Lemey, Philippe; Salemi, Marco; Vandamme, Anne-Mieke, eds. (2009). The Phylogenetic Handbook: A Practical Approach to Phylogenetic Analysis and Hypothesis Testing (2nd ed.). Cambridge, UK; New York: Cambridge University Press. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-521-73071-6</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2009464132">2009464132</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/295002266">295002266</a>.</cite></li>

<li><cite id="CITEREFLucretius1916" class="citation book cs1"><a href="Lucretius" title="Lucretius">Lucretius</a> (1916). "Book V". In <a href="William_Ellery_Leonard" title="William Ellery Leonard">Leonard, William Ellery</a> (ed.). <a href="De_rerum_natura" title="De rerum natura">De rerum natura</a>. Translated by William Ellery Leonard. Medford/Somerville, MA: <a href="Tufts_University" title="Tufts University">Tufts University</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/33233743">33233743</a>.</cite></li>

<li><cite id="CITEREFMacArthurWilson2001" class="citation book cs1"><a href="Robert_MacArthur" class="mw-redirect" title="Robert MacArthur">MacArthur, Robert H.</a>; <a href="E._O._Wilson" title="E. O. Wilson">Wilson, Edward O.</a> (2001) [Originally published 1967]. <a href="The_Theory_of_Island_Biogeography" title="The Theory of Island Biogeography">The Theory of Island Biogeography</a>. Princeton Landmarks in Biology. New preface by Edward O. Wilson. Princeton, NJ: Princeton University Press. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-691-08836-5</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/00051495">00051495</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/45202069">45202069</a>.</cite></li>

<li><cite id="CITEREFMalthus1798" class="citation book cs1"><a href="Thomas_Robert_Malthus" title="Thomas Robert Malthus">Malthus, Thomas Robert</a> (1798). <a href="An_Essay_on_the_Principle_of_Population" title="An Essay on the Principle of Population">An Essay on the Principle of Population, As It Affects the Future Improvement of Society: with Remarks on the Speculations of Mr. Godwin, M. Condorcet, and Other Writers</a> (1st ed.). London: J. Johnson. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/46038215">46038215</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/65344349">65344349</a>.</cite> The book is available <a rel="nofollow" class="external text" href="http://www.faculty.rsu.edu/users/f/felwell/www/Theorists/Malthus/Essay.htm#112">here</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20210715132543/http://www.faculty.rsu.edu/users/f/felwell/www/Theorists/Malthus/Essay.htm#112">Archived</a> 15 July 2021 at the <a href="Wayback_Machine" title="Wayback Machine">Wayback Machine</a> from Frank Elwell, <a href="Rogers_State_University" title="Rogers State University">Rogers State University</a>.</li>

<li><cite id="CITEREFMayr1942" class="citation book cs1"><a href="Ernst_Mayr" title="Ernst Mayr">Mayr, Ernst</a> (1942). <a href="Systematics_and_the_Origin_of_Species" title="Systematics and the Origin of Species">Systematics and the Origin of Species from the Viewpoint of a Zoologist</a>. Columbia Biological Series. Vol. 13. New York: Columbia University Press. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/43001098">43001098</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/766053">766053</a>.</cite></li>

<li><cite id="CITEREFMayr2006" class="citation book cs1">Mayr, Ernst (2006) [Originally published 1972; Chicago, IL: Aldine Publishing Co.]. "Sexual Selection and Natural Selection". In Campbell, Bernard G. (ed.). Sexual Selection and the Descent of Man: The Darwinian Pivot. New Brunswick, NJ: <a href="Transaction_Publishers" title="Transaction Publishers">AldineTransaction</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-202-30845-6</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2005046652">2005046652</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/62857839">62857839</a>.</cite></li>

<li><cite id="CITEREFMichod1999" class="citation book cs1">Michod, Richard A. (1999). <a rel="nofollow" class="external text" href="https://archive.org/details/darwiniandynamic00mich">Darwinian Dynamics: Evolutionary Transitions in Fitness and Individuality</a>. Princeton, NJ: Princeton University Press. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-691-02699-2</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/98004166">98004166</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/38948118">38948118</a>.</cite></li>

<li><cite id="CITEREFMiller2000" class="citation book cs1"><a href="Geoffrey_Miller_(psychologist)" title="Geoffrey Miller (psychologist)">Miller, Geoffrey</a> (2000). <a rel="nofollow" class="external text" href="https://archive.org/details/matingmind00geof">The Mating Mind: How Sexual Choice Shaped the Evolution of Human Nature</a> (1st ed.). New York: <a href="Doubleday_(publisher)" title="Doubleday (publisher)">Doubleday</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-385-49516-5</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/00022673">00022673</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/43648482">43648482</a>.</cite></li>

<li><cite id="CITEREFMitchell1996" class="citation book cs1"><a href="Melanie_Mitchell" title="Melanie Mitchell">Mitchell, Melanie</a> (1996). <a rel="nofollow" class="external text" href="https://archive.org/details/introductiontoge00mitc">An Introduction to Genetic Algorithms</a>. Complex Adaptive Systems. Cambridge, MA: <a href="MIT_Press" title="MIT Press">MIT Press</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-262-13316-6</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/95024489">95024489</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/42854439">42854439</a>.</cite></li>

<li><cite id="CITEREFPinker1995" class="citation book cs1"><a href="Steven_Pinker" title="Steven Pinker">Pinker, Steven</a> (1995) [Originally published 1994; New York: <a href="William_Morrow_and_Company" title="William Morrow and Company">William Morrow and Company</a>]. <a href="The_Language_Instinct" title="The Language Instinct">The Language Instinct: How the Mind Creates Language</a> (1st <a href="Harper_Perennial" title="Harper Perennial">Harper Perennial</a> ed.). New York: Harper Perennial. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-06-097651-4</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/94039138">94039138</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/670524593">670524593</a>.</cite></li>

<li><cite id="CITEREFRice2004" class="citation book cs1">Rice, Sean H. (2004). Evolutionary Theory: Mathematical and Conceptual Foundations. Sunderland, MA: Sinauer Associates. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-87893-702-8</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2004008054">2004008054</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/54988554">54988554</a>.</cite></li>

<li><cite id="CITEREFRoux1881" class="citation book cs1"><a href="Wilhelm_Roux" title="Wilhelm Roux">Roux, Wilhelm</a> (1881). <a rel="nofollow" class="external text" href="https://archive.org/details/bub_gb_66lBAAAAYAAJ">Der Kampf der Theile im Organismus</a>. Leipzig: <a href="Wilhelm_Engelmann" title="Wilhelm Engelmann">Wilhelm Engelmann</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/8200805">8200805</a>.</cite> <a rel="nofollow" class="external text" href="https://archive.org/details/derkampfdertheil00roux">Der Kampf der Theile im Organismus</a> at the <a href="Internet_Archive" title="Internet Archive">Internet Archive</a> Retrieved 2015-08-11.</li>

<li><cite id="CITEREFSober1993" class="citation book cs1"><a href="Elliott_Sober" title="Elliott Sober">Sober, Elliott</a> (1993) [Originally published 1984; Cambridge, MA: MIT Press]. The Nature of Selection: Evolutionary Theory in Philosophical Focus. Chicago, IL: <a href="University_of_Chicago_Press" title="University of Chicago Press">University of Chicago Press</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-226-76748-2</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/93010367">93010367</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/896826726">896826726</a>.</cite></li>

<li><cite id="CITEREFWallace1871" class="citation book cs1"><a href="Alfred_Russel_Wallace" title="Alfred Russel Wallace">Wallace, Alfred Russel</a> (1871) [Originally published 1870]. <a rel="nofollow" class="external text" href="http://quod.lib.umich.edu/cgi/t/text/text-idx?c=moa&idno=AJP5195.0001.001&view=toc">Contributions to the Theory of Natural Selection. A Series of Essays</a> (2nd, with corrections and additions ed.). New York: <a href="Macmillan_Publishers" title="Macmillan Publishers">Macmillan & Co.</a> <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/agr04000394">agr04000394</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/809350209">809350209</a>.</cite></li>

<li><cite id="CITEREFWilliams1966" class="citation book cs1"><a href="George_C._Williams_(biologist)" class="mw-redirect" title="George C. Williams (biologist)">Williams, George C.</a> (1966). <a href="Adaptation_and_Natural_Selection" title="Adaptation and Natural Selection">Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought</a>. Princeton Science Library. Princeton, NJ: Princeton University Press. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-691-02615-2</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/65017164">65017164</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/35230452">35230452</a>.</cite></li>

<li><cite id="CITEREFWilson2002" class="citation book cs1"><a href="David_Sloan_Wilson" title="David Sloan Wilson">Wilson, David Sloan</a> (2002). Darwin's Cathedral: Evolution, Religion, and the Nature of Society. Chicago, IL: University of Chicago Press. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-691-02615-2</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2002017375">2002017375</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/48777441">48777441</a>.</cite></li>

<li><cite id="CITEREFZimmerEmlen2013" class="citation book cs1"><a href="Carl_Zimmer" title="Carl Zimmer">Zimmer, Carl</a>; <a href="Douglas_Emlen" title="Douglas Emlen">Emlen, Douglas J.</a> (2013). Evolution: Making Sense of Life (1st ed.). Greenwood Village, CO: Roberts and Company Publishers. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-1-936221-17-2</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2012025118">2012025118</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/767565909">767565909</a>.</cite></li></ul>

</div>

<div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2></div>

<ul><li>For technical audiences

<ul><li><cite class="citation book cs1"><a href="Graham_Bell_(biologist)" title="Graham Bell (biologist)">Bell, Graham</a> (2008). Selection: The Mechanism of Evolution (2nd ed.). Oxford; New York: <a href="Oxford_University_Press" title="Oxford University Press">Oxford University Press</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-19-856972-5</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2007039692">2007039692</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/170034792">170034792</a>.</cite></li>

<li><cite class="citation book cs1">Johnson, Clifford (1976). <a rel="nofollow" class="external text" href="https://archive.org/details/introductiontona00john">Introduction to Natural Selection</a>. Baltimore, MD: University Park Press. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-8391-0936-5</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/76008175">76008175</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/2091640">2091640</a>.</cite></li>

<li><cite class="citation book cs1"><a href="Stephen_Jay_Gould" title="Stephen Jay Gould">Gould, Stephen Jay</a> (2002). <a href="The_Structure_of_Evolutionary_Theory" title="The Structure of Evolutionary Theory">The Structure of Evolutionary Theory</a>. Cambridge, MA: <a href="Harvard_University_Press" title="Harvard University Press">Belknap Press of Harvard University Press</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-674-00613-3</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2001043556">2001043556</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/47869352">47869352</a>.</cite></li>

<li><cite class="citation book cs1"><a href="John_Maynard_Smith" title="John Maynard Smith">Maynard Smith, John</a> (1993) [Originally published 1958; Harmondsworth, England: <a href="Penguin_Books" title="Penguin Books">Penguin Books</a>]. <a href="The_Theory_of_Evolution" title="The Theory of Evolution">The Theory of Evolution</a> (Canto ed.). Cambridge, New York: <a href="Cambridge_University_Press" title="Cambridge University Press">Cambridge University Press</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-521-45128-4</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/93020358">93020358</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/27676642">27676642</a>.</cite></li>

<li><cite class="citation journal cs1"><a href="Karl_Popper" title="Karl Popper">Popper, Karl</a> (December 1978). <a rel="nofollow" class="external text" href="http://www.informationphilosopher.com/solutions/philosophers/popper/natural_selection_and_the_emergence_of_mind.html">"Natural Selection and the Emergence of Mind"</a>. <a href="Dialectica" title="Dialectica">Dialectica</a>. 32 (3–4): 339–355. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1746-8361.1978.tb01321.x">10.1111/j.1746-8361.1978.tb01321.x</a>. <a href="ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0012-2017">0012-2017</a>.</cite></li>

<li><cite class="citation journal cs1">Sammut-Bonnici, Tanya; Wensley, Robin (September 2002). <a rel="nofollow" class="external text" href="http://wrap.warwick.ac.uk/57024/1/WRAP_Sammut-Bonnici_httpwrap%20warwick%20ac%20uk57024%20%282%29.pdf">"Darwinism, probability and complexity: Market-based organizational transformation and change explained through the theories of evolution"</a> (PDF). <a href="International_Journal_of_Management_Reviews" title="International Journal of Management Reviews">International Journal of Management Reviews</a>. 4 (3): 291–315. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2F1468-2370.00088">10.1111/1468-2370.00088</a>. <a href="ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/1460-8545">1460-8545</a>.</cite></li>

<li><cite class="citation book cs1"><a href="Elliott_Sober" title="Elliott Sober">Sober, Elliott</a>, ed. (1994). <a rel="nofollow" class="external text" href="https://archive.org/details/conceptualissues0000unse">Conceptual Issues in Evolutionary Biology</a> (2nd ed.). Cambridge, MA: <a href="MIT_Press" title="MIT Press">MIT Press</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-262-69162-8</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/93008199">93008199</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/28150417">28150417</a>.</cite></li>

<li><cite class="citation book cs1"><a href="George_C._Williams_(biologist)" class="mw-redirect" title="George C. Williams (biologist)">Williams, George C.</a> (1992). Natural Selection: Domains, Levels, and Challenges. Oxford Series in Ecology and Evolution. New York: Oxford University Press. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-19-506933-4</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/91038938">91038938</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/228136567">228136567</a>.</cite></li></ul></li>

<li>For general audiences

<ul><li><cite class="citation book cs1"><a href="Richard_Dawkins" title="Richard Dawkins">Dawkins, Richard</a> (1996). <a rel="nofollow" class="external text" href="https://archive.org/details/climbingmountimp0000dawk">Climbing Mount Improbable</a> (1st American ed.). New York: W.W. Norton & Company. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-393-03930-6</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/34633422">34633422</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/34633422">34633422</a>.</cite></li>

<li><cite class="citation book cs1">Gould, Stephen Jay (1977). <a href="Ever_Since_Darwin" title="Ever Since Darwin">Ever Since Darwin: Reflections in Natural History</a> (1st ed.). New York: W.W. Norton & Company. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-393-06425-4</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/77022504">77022504</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/3090189">3090189</a>.</cite></li>

<li><cite class="citation book cs1"><a href="Steve_Jones_(biologist)" title="Steve Jones (biologist)">Jones, Steve</a> (2000). <a href="Almost_Like_a_Whale" title="Almost Like a Whale">Darwin's Ghost: The Origin of Species Updated</a> (1st ed.). New York: <a href="Random_House" title="Random House">Random House</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-375-50103-6</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/99053246">99053246</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/42690131">42690131</a>.</cite></li>

<li><cite class="citation journal cs1"><a href="Richard_Lewontin" title="Richard Lewontin">Lewontin, Richard C.</a> (September 1978). "Adaptation". <a href="Scientific_American" title="Scientific American">Scientific American</a>. 239 (3): 212–230. <a href="Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1978SciAm.239c.212L">1978SciAm.239c.212L</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fscientificamerican0978-212">10.1038/scientificamerican0978-212</a>. <a href="ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0036-8733">0036-8733</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/705323">705323</a>.</cite></li>

<li><cite class="citation book cs1"><a href="Ernst_Mayr" title="Ernst Mayr">Mayr, Ernst</a> (2002) [Originally published 2001; New York: <a href="Basic_Books" title="Basic Books">Basic Books</a>]. What Evolution Is. Science Masters. London: <a href="Weidenfeld_%26_Nicolson" title="Weidenfeld & Nicolson">Weidenfeld & Nicolson</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-297-60741-0</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2001036562">2001036562</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/248107061">248107061</a>.</cite></li>

<li><cite class="citation book cs1"><a href="Jonathan_Weiner" title="Jonathan Weiner">Weiner, Jonathan</a> (1994). <a href="The_Beak_of_the_Finch" title="The Beak of the Finch">The Beak of the Finch: A Story of Evolution in Our Time</a> (1st ed.). New York: <a href="Alfred_A._Knopf" title="Alfred A. Knopf">Knopf</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-679-40003-5</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/93036755">93036755</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/29029572">29029572</a>.</cite></li></ul></li>

<li>Historical

<ul><li><cite class="citation book cs1"><a href="Marek_Kohn" title="Marek Kohn">Kohn, Marek</a> (2004). A Reason for Everything: Natural Selection and the English Imagination. London: <a href="Faber_and_Faber" class="mw-redirect" title="Faber and Faber">Faber and Faber</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-571-22392-3</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2005360890">2005360890</a>. <a href="OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/57200626">57200626</a>.</cite></li>

<li><cite class="citation journal cs1"><a href="Conway_Zirkle" title="Conway Zirkle">Zirkle, Conway</a> (25 April 1941). "Natural Selection before the 'Origin of Species'". <a href="Proceedings_of_the_American_Philosophical_Society" title="Proceedings of the American Philosophical Society">Proceedings of the American Philosophical Society</a>. 84 (1): 71–123. <a href="ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0003-049X">0003-049X</a>. <a href="JSTOR_(identifier)" class="mw-redirect" title="JSTOR (identifier)">JSTOR</a> <a rel="nofollow" class="external text" href="https://www.jstor.org/stable/984852">984852</a>.</cite></li></ul></li></ul>

<div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2></div>

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<div class="side-box-text plainlist">Wikiquote has quotations related to <a href="https://en.wikiquote.org/wiki/Special:Search/Natural_selection" class="extiw external" title="q:Special:Search/Natural selection">Natural selection</a>.</div></div>

</div>

<ul><li><cite class="citation web cs1"><a href="Charles_Darwin" title="Charles Darwin">Darwin, Charles</a>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20010225025716/http://www.literature.org/authors/darwin-charles/the-origin-of-species/chapter-04.html">"On the Origin of Species"</a>. Archived from <a rel="nofollow" class="external text" href="http://literature.org/authors/darwin-charles/the-origin-of-species/chapter-04.html">the original</a> on 25 February 2001.</cite> – Chapter 4, Natural Selection</li></ul>

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</style></div><div role="navigation" class="navbox" aria-labelledby="Evolutionary_biology753" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Evolutionary_biology753" style="font-size:114%;margin:0 4em"><a href="Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div>

<ul><li><a href="Introduction_to_evolution" title="Introduction to evolution">Introduction</a></li>

<li><a href="Outline_of_evolution" title="Outline of evolution">Outline</a></li>

<li><a href="Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of evolution</a></li>

<li><a href="History_of_life" title="History of life">History of life</a></li>

<li><a href="Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Evolution" title="Evolution">Evolution</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Abiogenesis" title="Abiogenesis">Abiogenesis</a></li>

<li><a href="Adaptation" title="Adaptation">Adaptation</a></li>

<li><a href="Adaptive_radiation" title="Adaptive radiation">Adaptive radiation</a></li>

<li><a href="Altruism_(biology)" title="Altruism (biology)">Altruism</a>

<ul><li><a href="Cheating_(biology)" title="Cheating (biology)">Cheating</a></li>

<li><a href="Reciprocal_altruism" title="Reciprocal altruism">Reciprocal</a></li></ul></li>

<li><a href="Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li>

<li><a href="Cladistics" title="Cladistics">Cladistics</a></li>

<li><a href="Coevolution" title="Coevolution">Coevolution</a>

<ul><li><a href="Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a></li></ul></li>

<li><a href="Common_descent" title="Common descent">Common descent</a></li>

<li><a href="Convergent_evolution" title="Convergent evolution">Convergence</a></li>

<li><a href="Divergent_evolution" title="Divergent evolution">Divergence</a></li>

<li><a href="Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li>

<li><a href="Evidence_of_common_descent" title="Evidence of common descent">Evidence of evolution</a></li>

<li><a href="Evolutionary_arms_race" title="Evolutionary arms race">Evolutionary arms race</a></li>

<li><a href="Evolutionary_pressure" title="Evolutionary pressure">Evolutionary pressure</a></li>

<li><a href="Exaptation" title="Exaptation">Exaptation</a></li>

<li><a href="Extinction" title="Extinction">Extinction</a>

<ul><li><a href="Extinction_event" title="Extinction event">Event</a></li></ul></li>

<li><a href="Homology_(biology)" title="Homology (biology)">Homology</a></li>

<li><a href="Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li>

<li><a href="Macroevolution" title="Macroevolution">Macroevolution</a></li>

<li><a href="Microevolution" title="Microevolution">Microevolution</a></li>

<li><a href="Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li>

<li><a href="Nonadaptive_radiation" title="Nonadaptive radiation">Non-adaptive radiation</a></li>

<li><a href="Abiogenesis" title="Abiogenesis">Origin of life</a></li>

<li><a href="Panspermia" title="Panspermia">Panspermia</a></li>

<li><a href="Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li>

<li><a href="Signalling_theory" title="Signalling theory">Signalling theory</a>

<ul><li><a href="Handicap_principle" title="Handicap principle">Handicap principle</a></li></ul></li>

<li><a href="Speciation" title="Speciation">Speciation</a>

<ul><li><a href="Species" title="Species">Species</a></li>

<li><a href="Species_complex" title="Species complex">Species complex</a></li></ul></li>

<li><a href="Evolutionary_taxonomy" title="Evolutionary taxonomy">Taxonomy</a></li>

<li><a href="Unit_of_selection" title="Unit of selection">Unit of selection</a>

<ul><li><a href="Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene-centered view of evolution</a></li></ul></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Population_genetics" title="Population genetics">Population genetics</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Selective_breeding" title="Selective breeding">Artificial selection</a></li>

<li><a href="Biodiversity" title="Biodiversity">Biodiversity</a></li>

<li><a href="Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li>

<li><a href="Fisher's_principle" title="Fisher's principle">Fisher's principle</a></li>

<li><a href="Fitness_(biology)" title="Fitness (biology)">Fitness</a>

<ul><li><a href="Inclusive_fitness" title="Inclusive fitness">Inclusive</a></li></ul></li>

<li><a href="Genetic_drift" title="Genetic drift">Genetic drift</a></li>

<li><a href="Kin_selection" title="Kin selection">Kin selection</a>

<ul><li><a href="Inbreeding_avoidance" title="Inbreeding avoidance">Inbreeding avoidance</a></li>

<li><a href="Kin_recognition" title="Kin recognition">Kin recognition</a></li>

<li><a href="Parental_investment" title="Parental investment">Parental investment</a></li>

<li><a href="Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li>

<li><a href="Mutation" title="Mutation">Mutation</a></li>

<li><a href="Population" title="Population">Population</a></li>

<li><a href="Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li>

<li><a href="Sexual_selection" title="Sexual selection">Sexual selection</a>

<ul><li><a href="Sexual_selection_in_flowering_plants" title="Sexual selection in flowering plants">Flowering plants</a></li>

<li><a href="Sexual_selection_in_fungi" title="Sexual selection in fungi">Fungi</a></li>

<li><a href="Mate_choice" title="Mate choice">Mate choice</a></li></ul></li>

<li><a href="Social_selection" title="Social selection">Social selection</a></li>

<li><a href="Trivers%E2%80%93Willard_hypothesis" title="Trivers–Willard hypothesis">Trivers–Willard hypothesis</a></li>

<li><a href="Genetic_variation" title="Genetic variation">Variation</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Development</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li>

<li><a href="Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></li>

<li><a href="Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li>

<li><a href="Inversion_(evolutionary_biology)" title="Inversion (evolutionary biology)">Inversion</a></li>

<li><a href="Modularity_(biology)" title="Modularity (biology)">Modularity</a></li>

<li><a href="Phenotypic_plasticity" title="Phenotypic plasticity">Phenotypic plasticity</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="Taxon" title="Taxon">taxa</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Evolution_of_bacteria" title="Evolution of bacteria">Bacteria</a></li>

<li><a href="Evolution_of_birds" title="Evolution of birds">Birds</a>

<ul><li><a href="Origin_of_birds" title="Origin of birds">origin</a></li></ul></li>

<li><a href="Evolution_of_brachiopods" title="Evolution of brachiopods">Brachiopods</a></li>

<li><a href="Evolution_of_molluscs" title="Evolution of molluscs">Molluscs</a>

<ul><li><a href="Evolution_of_cephalopods" title="Evolution of cephalopods">Cephalopods</a></li></ul></li>

<li><a href="Dinosaur#Evolutionary_history" title="Dinosaur">Dinosaurs</a></li>

<li><a href="Evolution_of_fungi" title="Evolution of fungi">Fungi</a></li>

<li><a href="Evolution_of_insects" title="Evolution of insects">Insects</a>

<ul><li><a href="Evolution_of_butterflies" title="Evolution of butterflies">butterflies</a></li></ul></li>

<li><a href="Evolution_of_mammals" title="Evolution of mammals">Mammals</a>

<li><a href="Canidae#Evolution" title="Canidae">canids</a>

<ul><li><a href="Evolution_of_the_wolf" title="Evolution of the wolf">wolves</a></li>

<li><a href="Hyena#Evolution" title="Hyena">hyenas</a></li>

<li><a href="Evolution_of_cetaceans" title="Evolution of cetaceans">dolphins and whales</a></li>

<li><a href="Evolution_of_the_horse" title="Evolution of the horse">horses</a></li>

<li><a href="Evolution_of_Macropodidae" title="Evolution of Macropodidae">Kangaroos</a></li>

<li><a href="Evolution_of_primates" title="Evolution of primates">primates</a>

<ul><li><a href="Human_evolution" title="Human evolution">humans</a></li>

<li><a href="Evolution_of_lemurs" title="Evolution of lemurs">lemurs</a></li></ul></li>

<li><a href="Evolutionary_history_of_plants" title="Evolutionary history of plants">Plants</a>

<ul><li><a href="Pollinator-mediated_selection" title="Pollinator-mediated selection">pollinator-mediated</a></li></ul></li>

<li><a href="Evolution_of_reptiles" title="Evolution of reptiles">Reptiles</a></li>

<li><a href="Evolution_of_spiders" title="Evolution of spiders">Spiders</a></li>

<li><a href="Evolution_of_tetrapods" title="Evolution of tetrapods">Tetrapods</a></li>

<li><a href="Viral_evolution" title="Viral evolution">Viruses</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="Organ_(biology)" title="Organ (biology)">organs</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<li><a href="Evolution_of_flagella" title="Evolution of flagella">Flagella</a></li>

<li><a href="Eukaryote#Origin_of_eukaryotes" title="Eukaryote">Eukaryotes</a>

<ul><li><a href="Symbiogenesis" title="Symbiogenesis">symbiogenesis</a></li>

<li><a href="Chromosome#Eukaryotes" title="Chromosome">chromosome</a></li>

<li><a href="Endomembrane_system#Evolution" title="Endomembrane system">endomembrane system</a></li>

<li><a href="Mitochondrion#Origin_and_evolution" title="Mitochondrion">mitochondria</a></li>

<li><a href="Cell_nucleus#Evolution" title="Cell nucleus">nucleus</a></li>

<li><a href="Plastid#Origin" title="Plastid">plastids</a></li></ul></li>

<li>In animals

<li><a href="Evolution_of_mammalian_auditory_ossicles" title="Evolution of mammalian auditory ossicles">auditory ossicle</a></li>

<li><a href="Evolution_of_nervous_systems" title="Evolution of nervous systems">nervous system</a></li>

<li><a href="Evolution_of_the_brain" title="Evolution of the brain">brain</a></li></ul></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="Biological_process" title="Biological process">processes</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Evolution_of_ageing" title="Evolution of ageing">Aging</a>

<ul><li><a href="Death#Evolution_of_aging_and_mortality" title="Death">Death</a></li>

<li><a href="Programmed_cell_death#Evolutionary_origin_of_mitochondrial_apoptosis" title="Programmed cell death">Programmed cell death</a></li></ul></li>

<li><a href="Origin_of_avian_flight" title="Origin of avian flight">Avian flight</a></li>

<li><a href="Evolution_of_biological_complexity" title="Evolution of biological complexity">Biological complexity</a></li>

<li><a href="Cooperation_(evolution)" title="Cooperation (evolution)">Cooperation</a></li>

<li><a href="Evolution_of_color_vision" title="Evolution of color vision">Color vision</a>

<ul><li><a href="Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">in primates</a></li></ul></li>

<li><a href="Evolution_of_emotion" title="Evolution of emotion">Emotion</a></li>

<li><a href="Empathy#Evolution_across_species" title="Empathy">Empathy</a></li>

<li><a href="Evolutionary_ethics" title="Evolutionary ethics">Ethics</a></li>

<li><a href="Evolution_of_eusociality" title="Evolution of eusociality">Eusociality</a></li>

<li><a href="Immune_system#Evolution_and_other_mechanisms" title="Immune system">Immune system</a></li>

<li><a href="Metabolism#Evolution" title="Metabolism">Metabolism</a></li>

<li><a href="Monogamy_in_animals" title="Monogamy in animals">Monogamy</a></li>

<li><a href="Evolution_of_morality" title="Evolution of morality">Morality</a></li>

<li><a href="Mosaic_evolution" title="Mosaic evolution">Mosaic evolution</a></li>

<li><a href="Multicellular_organism#Evolutionary_history" title="Multicellular organism">Multicellularity</a></li>

<li><a href="Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Sexual reproduction</a>

<ul><li><a href="Anisogamy#Evolution" title="Anisogamy">Gamete differentiation/sexes</a></li>

<li><a href="Biological_life_cycle" title="Biological life cycle">Life cycles/nuclear phases</a></li>

<li><a href="Mating_type" title="Mating type">Mating types</a></li>

<li><a href="Origin_and_function_of_meiosis" title="Origin and function of meiosis">Meiosis</a></li>

<li><a href="Sex-determination_system#Evolution" title="Sex-determination system">Sex-determination</a></li></ul></li>

<li><a href="Evolution_of_snake_venom" title="Evolution of snake venom">Snake venom</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Tempo_and_Mode_in_Evolution" title="Tempo and Mode in Evolution">Tempo and modes</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Phyletic_gradualism" title="Phyletic gradualism">Gradualism</a>/<a href="Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a>/<a href="Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li>

<li><a href="Point_mutation" title="Point mutation">Micromutation</a>/<a href="Mutationism#Later_mutationist_theories" title="Mutationism">Macromutation</a></li>

<li><a href="Uniformitarianism" title="Uniformitarianism">Uniformitarianism</a>/<a href="Catastrophism" title="Catastrophism">Catastrophism</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Speciation" title="Speciation">Speciation</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Allopatric_speciation" title="Allopatric speciation">Allopatric</a></li>

<li><a href="Anagenesis" title="Anagenesis">Anagenesis</a></li>

<li><a href="Catagenesis_(biology)" title="Catagenesis (biology)">Catagenesis</a></li>

<li><a href="Cladogenesis" title="Cladogenesis">Cladogenesis</a></li>

<li><a href="Cospeciation" title="Cospeciation">Cospeciation</a></li>

<li><a href="Ecological_speciation" title="Ecological speciation">Ecological</a></li>

<li><a href="Hybrid_speciation" title="Hybrid speciation">Hybrid</a></li>

<li><a href="Nonecological_speciation" title="Nonecological speciation">Non-ecological</a></li>

<li><a href="Parapatric_speciation" title="Parapatric speciation">Parapatric</a></li>

<li><a href="Peripatric_speciation" title="Peripatric speciation">Peripatric</a></li>

<li><a href="Reinforcement_(speciation)" title="Reinforcement (speciation)">Reinforcement</a></li>

<li><a href="Sympatric_speciation" title="Sympatric speciation">Sympatric</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="History_of_evolutionary_thought" title="History of evolutionary thought">History</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance and Enlightenment</a></li>

<li><a href="Transmutation_of_species" title="Transmutation of species">Transmutation of species</a></li>

<li><a href="David_Hume" title="David Hume">David Hume</a>

<ul><li><a href="Dialogues_Concerning_Natural_Religion" title="Dialogues Concerning Natural Religion">Dialogues Concerning Natural Religion</a></li></ul></li>

<li><a href="Charles_Darwin" title="Charles Darwin">Charles Darwin</a>

<ul><li><a href="On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species</a></li></ul></li>

<li><a href="History_of_paleontology" title="History of paleontology">History of paleontology</a></li>

<li><a href="Transitional_fossil" title="Transitional fossil">Transitional fossil</a></li>

<li><a href="Blending_inheritance" title="Blending inheritance">Blending inheritance</a></li>

<li><a href="Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a></li>

<li><a href="The_eclipse_of_Darwinism" title="The eclipse of Darwinism">The eclipse of Darwinism</a></li>

<li><a href="Neo-Darwinism" title="Neo-Darwinism">Neo-Darwinism</a></li>

<li><a href="Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li>

<li><a href="History_of_molecular_evolution" title="History of molecular evolution">History of molecular evolution</a></li>

<li><a href="Extended_evolutionary_synthesis" title="Extended evolutionary synthesis">Extended evolutionary synthesis</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Philosophy</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Darwinism" title="Darwinism">Darwinism</a></li>

<li><a href="Alternatives_to_Darwinian_evolution" title="Alternatives to Darwinian evolution">Alternatives</a>

<ul><li><a href="Catastrophism" title="Catastrophism">Catastrophism</a></li>

<li><a href="Lamarckism" title="Lamarckism">Lamarckism</a></li>

<li><a href="Orthogenesis" title="Orthogenesis">Orthogenesis</a></li>

<li><a href="Mutationism" title="Mutationism">Mutationism</a></li>

<li><a href="Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li>

<li><a href="Structuralism_(biology)" title="Structuralism (biology)">Structuralism</a>

<ul><li><a href="Spandrel_(biology)" title="Spandrel (biology)">Spandrel</a></li></ul></li>

<li><a href="Theistic_evolution" title="Theistic evolution">Theistic</a></li>

<li><a href="Vitalism" title="Vitalism">Vitalism</a></li></ul></li>

<li><a href="Teleology_in_biology" title="Teleology in biology">Teleology in biology</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Biogeography" title="Biogeography">Biogeography</a></li>

<li><a href="Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li>

<li><a href="Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li>

<li><a href="Group_selection" title="Group selection">Group selection</a>

<ul><li><a href="Cultural_evolution" title="Cultural evolution">Cultural evolution</a></li>

<li><a href="Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li>

<li><a href="Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li></ul></li>

<li><a href="Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory of evolution</a></li>

<li><a href="Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li>

<li><a href="Molecular_evolution" title="Molecular evolution">Molecular evolution</a></li>

<li><a href="Astrobiology" title="Astrobiology">Astrobiology</a></li>

<li><a href="Phylogenetics" title="Phylogenetics">Phylogenetics</a>

<li><a href="Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li>

<li><a href="Protocell" title="Protocell">Protocell</a></li>

<li><a href="Systematics" title="Systematics">Systematics</a></li>

<li><a href="Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></li></ul>

</div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div>

<ul><li> Category</li>

<li> <a href="Portal%3AEvolutionary_biology" title="Portal:Evolutionary biology">Portal</a></li></ul>

</div></td></tr></tbody></table></div>

<div class="navbox-styles"></div><div role="navigation" class="navbox" aria-labelledby="Evolutionary_psychology1033" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Evolutionary_psychology1033" style="font-size:114%;margin:0 4em"><a href="Evolutionary_psychology" title="Evolutionary psychology">Evolutionary psychology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div>

<ul><li><a href="History_of_evolutionary_psychology" title="History of evolutionary psychology">History</a>

<ul><li><a href="History_of_evolutionary_thought" title="History of evolutionary thought">Evolutionary thought</a></li></ul></li>

<li><a href="Theoretical_foundations_of_evolutionary_psychology" title="Theoretical foundations of evolutionary psychology">Theoretical foundations</a>

<ul><li><a href="Adaptationism" title="Adaptationism">Adaptationism</a></li>

<li><a href="Cognitive_revolution" title="Cognitive revolution">Cognitive revolution</a></li>

<li><a href="Cognitivism_(psychology)" title="Cognitivism (psychology)">Cognitivism</a></li>

<li><a href="Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene selection theory</a></li>

<li><a href="Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li></ul></li>

<li><a href="Criticism_of_evolutionary_psychology" title="Criticism of evolutionary psychology">Criticism</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;"><a href="Human_evolution" title="Human evolution">Evolutionary processes</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Psychological_adaptation" title="Psychological adaptation">Adaptations</a></li>

<li><a href="Altruism_(biology)" title="Altruism (biology)">Altruism</a>

<ul><li><a href="Cheating_(biology)" title="Cheating (biology)">Cheating</a></li>

<li><a href="Hamiltonian_spite" title="Hamiltonian spite">Hamiltonian spite</a></li>

<li><a href="Reciprocal_altruism_in_humans" title="Reciprocal altruism in humans">Reciprocal</a></li></ul></li>

<li><a href="Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li>

<li><a href="Spandrel_(biology)" title="Spandrel (biology)">By-products</a></li>

<li><a href="Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li>

<li><a href="Exaptation" title="Exaptation">Exaptation</a></li>

<li><a href="Fitness_(biology)" title="Fitness (biology)">Fitness</a>

<ul><li><a href="Inclusive_fitness_in_humans" title="Inclusive fitness in humans">Inclusive</a></li></ul></li>

<li><a href="Kin_selection" title="Kin selection">Kin selection</a>

<ul><li><a href="Inbreeding_avoidance" title="Inbreeding avoidance">Inbreeding avoidance</a></li>

<li><a href="Kin_recognition" title="Kin recognition">Kin recognition</a></li>

<li><a href="Parental_investment" title="Parental investment">Parental investment</a></li>

<li><a href="Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li>

<li><a href="Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li>

<li><a href="Sexual_selection_in_humans" title="Sexual selection in humans">Sexual selection</a>

<ul><li><a href="Costly_signaling_theory_in_evolutionary_psychology" title="Costly signaling theory in evolutionary psychology">Costly signaling</a></li>

<li><a href="Male_intrasexual_competition" title="Male intrasexual competition">Male</a>/<a href="Female_intrasexual_competition" title="Female intrasexual competition">female intrasexual competition</a></li>

<li><a href="Mate_choice" title="Mate choice">Mate choice</a></li>

<li><a href="Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li></ul></li>

<li><a href="Social_selection" title="Social selection">Social selection</a></li>

<li><a href="Species-typical_behavior" title="Species-typical behavior">Species-typical behavior</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;">Areas</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Evolution_of_cognition" title="Evolution of cognition">Cognition</a> / <a href="Evolution_of_emotion" title="Evolution of emotion">Emotion</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Affect_(psychology)" title="Affect (psychology)">Affect</a>

<ul><li><a href="Affect_display" title="Affect display">Display</a></li>

<li><a href="Display_rules" title="Display rules">Display rules</a></li>

<li><a href="Facial_expression" title="Facial expression">Facial expression</a></li></ul></li>

<li><a href="Behavioral_modernity" title="Behavioral modernity">Behavioral modernity</a></li>

<li><a href="Cognitive_module" title="Cognitive module">Cognitive module</a>/<a href="Modularity_of_mind" title="Modularity of mind">modularity of mind</a>

<ul><li><a href="Adaptive_unconscious" title="Adaptive unconscious">Adaptive unconscious</a>/<a href="Unconscious_cognition" title="Unconscious cognition">Unconscious cognition</a></li>

<li><a href="Automatic_and_controlled_processes" title="Automatic and controlled processes">Automatic and controlled processes</a></li>

<li><a href="Cognitive_specialization" title="Cognitive specialization">Cognitive specialization</a></li>

<li><a href="Computational_theory_of_mind" title="Computational theory of mind">Computational theory of mind</a></li>

<li><a href="Domain-general_learning" title="Domain-general learning">Domain generality</a></li>

<li><a href="Domain_specificity" title="Domain specificity">Domain specificity</a>/<a href="Domain-specific_learning" title="Domain-specific learning">learning</a></li>

<li><a href="Dual_process_theory" title="Dual process theory">Dual process theory</a></li></ul></li>

<li><a href="Cognitive_tradeoff_hypothesis" title="Cognitive tradeoff hypothesis">Cognitive tradeoff hypothesis</a></li>

<li><a href="Evolution_of_the_brain" title="Evolution of the brain">Evolution of the brain</a></li>

<li><a href="Evolution_of_nervous_systems" title="Evolution of nervous systems">Evolution of nervous systems</a></li>

<li><a href="Fight-or-flight_response" title="Fight-or-flight response">Fight-or-flight response</a>

<ul><li><a href="Arachnophobia" title="Arachnophobia">Arachnophobia</a></li>

<li><a href="Fear_of_falling" title="Fear of falling">Basophobia</a></li>

<li><a href="Ophidiophobia" title="Ophidiophobia">Ophidiophobia</a></li></ul></li>

<li><a href="Folk_biology" title="Folk biology">Folk biology</a>/<a href="Folk_taxonomy" title="Folk taxonomy">taxonomy</a></li>

<li><a href="Folk_psychology" title="Folk psychology">Folk psychology</a>/<a href="Theory_of_mind" title="Theory of mind">theory of mind</a></li>

<li><a href="Evolution_of_human_intelligence" title="Evolution of human intelligence">Intelligence</a>

<ul><li><a href="Flynn_effect" title="Flynn effect">Flynn effect</a></li>

<li><a href="Wason_selection_task" title="Wason selection task">Wason selection task</a></li></ul></li>

<li><a href="Motor_control" title="Motor control">Motor control</a>/<a href="Motor_skill" title="Motor skill">skill</a></li>

<li><a href="Human_multitasking" title="Human multitasking">Multitasking</a></li>

<li><a href="Numerical_cognition" title="Numerical cognition">Numerical cognition</a>

<ul><li><a href="Number_sense" title="Number sense">Number sense</a></li>

<li><a href="Approximate_number_system" title="Approximate number system">Approximate number system</a></li>

<li><a href="Parallel_individuation_system" title="Parallel individuation system">Parallel individuation system</a></li></ul></li>

<li><a href="Neuroscience_of_sleep" title="Neuroscience of sleep">Sleep</a></li>

<li><a href="Visual_perception" title="Visual perception">Visual perception</a>

<ul><li><a href="Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">Color vision</a></li>

<li><a href="Na%C3%AFve_physics" title="Naïve physics">Naïve physics</a></li></ul></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Evolutionary_psychology_and_culture" title="Evolutionary psychology and culture">Culture</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Evolutionary_aesthetics" title="Evolutionary aesthetics">Aesthetics</a>

<ul><li><a href="Darwinian_literary_studies" title="Darwinian literary studies">Literary criticism</a></li>

<li><a href="Evolutionary_musicology" title="Evolutionary musicology">Musicology</a></li></ul></li>

<li><a href="Evolutionary_anthropology" title="Evolutionary anthropology">Anthropology</a>

<ul><li><a href="Biological_anthropology" title="Biological anthropology">Biological</a></li></ul></li>

<li><a href="Biosocial_criminology" title="Biosocial criminology">Crime</a></li>

<li><a href="Evolutionary_linguistics" title="Evolutionary linguistics">Language</a>

<ul><li><a href="Origin_of_language" title="Origin of language">Origin</a></li>

<li><a href="Evolutionary_psychology_of_language" title="Evolutionary psychology of language">Psychology</a></li>

<li><a href="Origin_of_speech" title="Origin of speech">Speech</a></li></ul></li>

<li><a href="Evolution_of_morality" title="Evolution of morality">Morality</a>

<ul><li><a href="Moral_foundations_theory" title="Moral foundations theory">Moral foundations</a></li></ul></li>

<li><a href="Evolutionary_psychology_of_religion" title="Evolutionary psychology of religion">Religion</a>

<ul><li><a href="Evolutionary_origin_of_religions" class="mw-redirect" title="Evolutionary origin of religions">Origin</a></li></ul></li>

<li><a href="Cultural_universal" title="Cultural universal">Universals</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Evolutionary_developmental_psychology" title="Evolutionary developmental psychology">Development</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Attachment_theory" title="Attachment theory">Attachment</a></li>

<li><a href="Human_bonding" title="Human bonding">Bonding</a></li>

<li><a href="Affectional_bond" title="Affectional bond">Affectional</a>/<a href="Maternal_bond" title="Maternal bond">maternal</a>/<a href="Paternal_bond" title="Paternal bond">paternal bond</a></li>

<li><a href="Maternal_deprivation" title="Maternal deprivation">Caregiver deprivation</a></li>

<li><a href="Attachment_in_children" title="Attachment in children">Childhood attachment</a></li>

<li><a href="Cinderella_effect" title="Cinderella effect">Cinderella effect</a></li>

<li><a href="Cognitive_development" title="Cognitive development">Cognitive development</a></li>

<li><a href="Evolutionary_educational_psychology" title="Evolutionary educational psychology">Education</a></li>

<li><a href="Language_acquisition" title="Language acquisition">Language acquisition</a></li>

<li><a href="Personality_development" title="Personality development">Personality development</a></li>

<li><a href="Socialization" title="Socialization">Socialization</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Ergonomics" title="Ergonomics">Human factors</a> / <a href="Evolutionary_psychiatry" title="Evolutionary psychiatry">Mental health</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Cognitive_ergonomics" title="Cognitive ergonomics">Cognitive ergonomics</a>

<ul><li><a href="Computer-mediated_communication" title="Computer-mediated communication">Computer-mediated communication</a></li>

<li><a href="Engineering_psychology" title="Engineering psychology">Engineering psychology</a></li>

<li><a href="Human%E2%80%93computer_interaction" title="Human–computer interaction">Human–computer interaction</a></li>

<li><a href="Media_naturalness_theory" title="Media naturalness theory">Media naturalness theory</a></li>

<li><a href="Neuroergonomics" title="Neuroergonomics">Neuroergonomics</a></li></ul></li>

<li><a href="Evolutionary_approaches_to_depression" title="Evolutionary approaches to depression">Depression</a></li>

<li><a href="Digital_media_use_and_mental_health" title="Digital media use and mental health">Digital media use and mental health</a></li>

<li><a href="Accident-proneness" title="Accident-proneness">Hypophobia</a></li>

<li><a href="Imprinted_brain_hypothesis" title="Imprinted brain hypothesis">Imprinted brain hypothesis</a></li>

<li><a href="Mind-blindness" title="Mind-blindness">Mind-blindness</a></li>

<li><a href="Psychological_effects_of_Internet_use" title="Psychological effects of Internet use">Psychological effects of Internet use</a></li>

<li><a href="Rank_theory_of_depression" title="Rank theory of depression">Rank theory of depression</a></li>

<li><a href="Evolution_of_schizophrenia" title="Evolution of schizophrenia">Schizophrenia</a></li>

<li><a href="Screen_time" title="Screen time">Screen time</a></li>

<li><a href="Smartphones_and_pedestrian_safety" title="Smartphones and pedestrian safety">Smartphones and pedestrian safety</a></li>

<li><a href="Social_aspects_of_television" title="Social aspects of television">Social aspects of television</a></li>

<li><a href="Societal_effects_of_cars" title="Societal effects of cars">Societal effects of cars</a>

<ul><li><a href="Distracted_driving" title="Distracted driving">Distracted driving</a></li>

<li><a href="Lead%E2%80%93crime_hypothesis" title="Lead–crime hypothesis">Lead–crime hypothesis</a></li>

<li><a href="Mobile_phones_and_driving_safety" title="Mobile phones and driving safety">Mobile phones and driving safety</a></li>

<li><a href="Texting_while_driving" title="Texting while driving">Texting while driving</a></li></ul></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Sexology" title="Sexology">Sex</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Human_sexual_activity" title="Human sexual activity">Activity</a></li>

<li><a href="Attachment_in_adults" title="Attachment in adults">Adult attachment</a></li>

<li><a href="Age_disparity_in_sexual_relationships" title="Age disparity in sexual relationships">Age disparity</a></li>

<li><a href="Sexual_arousal" title="Sexual arousal">Arousal</a></li>

<li><a href="Concealed_ovulation" title="Concealed ovulation">Concealed ovulation</a></li>

<li><a href="Coolidge_effect" title="Coolidge effect">Coolidge effect</a></li>

<li><a href="Sexual_desire" title="Sexual desire">Desire</a></li>

<li><a href="Sexual_fantasy" title="Sexual fantasy">Fantasy</a></li>

<li><a href="Effects_of_hormones_on_sexual_motivation" title="Effects of hormones on sexual motivation">Hormonal motivation</a></li>

<li><a href="Incest_taboo" title="Incest taboo">Incest taboo</a>

<ul><li><a href="Westermarck_effect" title="Westermarck effect">Westermarck effect</a></li></ul></li>

<li><a href="Sexual_jealousy" title="Sexual jealousy">Jealousy</a></li>

<li><a href="Mate_guarding_in_humans" title="Mate guarding in humans">Mate guarding</a></li>

<li><a href="Mating_preferences" title="Mating preferences">Mating preferences</a></li>

<li><a href="Human_mating_strategies" title="Human mating strategies">Mating strategies</a></li>

<li><a href="Biology_and_sexual_orientation" title="Biology and sexual orientation">Orientation</a></li>

<li><a href="Ovulatory_shift_hypothesis" title="Ovulatory shift hypothesis">Ovulatory shift hypothesis</a></li>

<li><a href="Physical_attractiveness" title="Physical attractiveness">Physical</a>/<a href="Sexual_attraction" title="Sexual attraction">Sexual attraction</a></li>

<li><a href="Human_sexuality" title="Human sexuality">Sexuality</a>/<a href="Human_male_sexuality" title="Human male sexuality">male</a>/<a href="Human_female_sexuality" title="Human female sexuality">female</a></li>

<li><a href="Sexy_son_hypothesis" title="Sexy son hypothesis">Sexy son hypothesis</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Sex_differences_in_psychology" title="Sex differences in psychology">Sex differences</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Aggression" title="Aggression">Aggression</a></li>

<li><a href="Sex_and_gender_differences_in_autism" title="Sex and gender differences in autism">Autism</a></li>

<li><a href="Sex_differences_in_cognition" title="Sex differences in cognition">Cognition</a></li>

<li><a href="Sex_differences_in_crime" title="Sex differences in crime">Crime</a></li>

<li><a href="Sexual_division_of_labour" title="Sexual division of labour">Division of labour</a></li>

<li><a href="Sex_differences_in_emotional_intelligence" title="Sex differences in emotional intelligence">Emotional intelligence</a></li>

<li><a href="Empathising%E2%80%93systemising_theory" title="Empathising–systemising theory">Empathising–systemising theory</a></li>

<li><a href="Gender_role" title="Gender role">Gender role</a></li>

<li><a href="Sex_differences_in_intelligence" title="Sex differences in intelligence">Intelligence</a></li>

<li><a href="Sex_differences_in_memory" title="Sex differences in memory">Memory</a></li>

<li><a href="Mental_disorders_and_gender" title="Mental disorders and gender">Mental health</a></li>

<li><a href="Sex_differences_in_narcissism" title="Sex differences in narcissism">Narcissism</a></li>

<li><a href="Neuroscience_of_sex_differences" title="Neuroscience of sex differences">Neuroscience</a></li>

<li><a href="Sex_differences_in_schizophrenia" title="Sex differences in schizophrenia">Schizophrenia</a></li>

<li><a href="Substance_abuse" title="Substance abuse">Substance abuse</a></li>

<li><a href="Gender_differences_in_suicide" title="Gender differences in suicide">Suicide</a></li>

<li><a href="Variability_hypothesis" title="Variability hypothesis">Variability hypothesis</a></li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;">Related subjects</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Academic disciplines</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Behavioral_economics" title="Behavioral economics">Behavioral</a>/<a href="Evolutionary_economics" title="Evolutionary economics">evolutionary economics</a></li>

<li><a href="Behavioral_epigenetics" title="Behavioral epigenetics">Behavioral epigenetics</a>/<a href="Behavioural_genetics" title="Behavioural genetics">genetics</a></li>

<li><a href="Affective_neuroscience" title="Affective neuroscience">Affective</a>/<a href="Behavioral_neuroscience" title="Behavioral neuroscience">behavioral</a>/<a href="Cognitive_neuroscience" title="Cognitive neuroscience">cognitive</a>/<a href="Evolutionary_neuroscience" title="Evolutionary neuroscience">evolutionary neuroscience</a></li>

<li><a href="Biocultural_anthropology" title="Biocultural anthropology">Biocultural anthropology</a></li>

<li><a href="Biolinguistics" title="Biolinguistics">Biolinguistics</a></li>

<li><a href="Biological_psychiatry" title="Biological psychiatry">Biological psychiatry</a></li>

<li><a href="Cognitive_ethology" title="Cognitive ethology">Cognitive ethology</a></li>

<li><a href="Cognitive_psychology" title="Cognitive psychology">Cognitive psychology</a></li>

<li><a href="Cognitive_science" title="Cognitive science">Cognitive science</a></li>

<li><a href="Comparative_psychology" title="Comparative psychology">Comparative psychology</a></li>

<li><a href="Cross-cultural_psychology" title="Cross-cultural psychology">Cross-cultural psychology</a></li>

<li><a href="Ethology" title="Ethology">Ethology</a></li>

<li><a href="Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></li>

<li><a href="Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li>

<li><a href="Functional_psychology" title="Functional psychology">Functional psychology</a></li>

<li><a href="Neuropsychology" title="Neuropsychology">Neuropsychology</a></li>

<li><a href="Philosophy_of_mind" title="Philosophy of mind">Philosophy of mind</a></li>

<li><a href="Population_genetics" title="Population genetics">Population genetics</a></li>

<li><a href="Primatology" title="Primatology">Primatology</a></li>

<li><a href="Sociobiology" title="Sociobiology">Sociobiology</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Research topics</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Animal_cognition" title="Animal cognition">Animal</a>/<a href="Comparative_cognition" title="Comparative cognition">comparative cognition</a>

<ul><li><a href="Animal_communication" title="Animal communication">Animal communication</a></li>

<li><a href="Animal_consciousness" title="Animal consciousness">Animal consciousness</a></li>

<li><a href="Animal_culture" title="Animal culture">Animal culture</a></li>

<li><a href="Animal_language" title="Animal language">Animal language</a></li>

<li><a href="Animal_navigation" title="Animal navigation">Animal navigation</a></li>

<li><a href="Animal_training" title="Animal training">Animal training</a></li>

<li><a href="Animal-made_art" title="Animal-made art">Animal-made art</a></li>

<li><a href="Cognitive_bias_in_animals" title="Cognitive bias in animals">Cognitive bias</a></li>

<li><a href="Emotion_in_animals" title="Emotion in animals">Emotion</a></li>

<li><a href="G_factor_in_non-humans" title="G factor in non-humans">g factor in non-humans</a></li>

<li><a href="Great_ape_language" title="Great ape language">Great ape language</a></li>

<li><a href="Human%E2%80%93animal_communication" title="Human–animal communication">Human–animal communication</a></li>

<li><a href="Number_sense_in_animals" title="Number sense in animals">Number sense</a></li>

<li><a href="Primate_cognition" title="Primate cognition">Primate cognition</a></li>

<li><a href="Theory_of_mind_in_animals" title="Theory of mind in animals">Theory of mind</a></li>

<li><a href="Cultural_evolution" title="Cultural evolution">Cultural evolution</a></li>

<li><a href="Evolutionary_epistemology" title="Evolutionary epistemology">Evolutionary epistemology</a></li>

<li><a href="Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li>

<li><a href="Unit_of_selection" title="Unit of selection">Unit of selection</a>

<ul><li><a href="Coevolution" title="Coevolution">Coevolution</a></li>

<li><a href="Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li>

<li><a href="Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li>

<li><a href="Fisher's_principle" title="Fisher's principle">Fisher's principle</a></li>

<li><a href="Group_selection" title="Group selection">Group selection</a></li>

<li><a href="Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory</a></li>

<li><a href="Lamarckism" title="Lamarckism">Lamarckism</a>/<a href="Acquired_characteristic" title="Acquired characteristic">acquired characteristics</a></li>

<li><a href="Population" title="Population">Population</a></li>

<li><a href="Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a></li>

<li><a href="Recent_human_evolution" title="Recent human evolution">Recent human evolution</a></li>

<li><a href="Species" title="Species">Species</a></li>

<li><a href="Species_complex" title="Species complex">Species complex</a></li>

<li><a href="Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></li>

<li><a href="Trivers%E2%80%93Willard_hypothesis" title="Trivers–Willard hypothesis">Trivers–Willard hypothesis</a></li></ul></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Theoretical positions</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Cultural_selection_theory" title="Cultural selection theory">Cultural selection theory</a></li>

<li><a href="Determinism" title="Determinism">Determinism</a>/<a href="Indeterminism" title="Indeterminism">indeterminism</a>

<ul><li><a href="Biological_determinism" title="Biological determinism">Biological determinism</a></li>

<li><a href="Connectionism" title="Connectionism">Connectionism</a></li>

<li><a href="Cultural_determinism" title="Cultural determinism">Cultural determinism</a></li>

<li><a href="Environmental_determinism" title="Environmental determinism">Environmental determinism</a></li>

<li><a href="Nature_versus_nurture" title="Nature versus nurture">Nature versus nurture</a></li>

<li><a href="Psychological_nativism" title="Psychological nativism">Psychological nativism</a></li>

<li><a href="Social_constructionism" title="Social constructionism">Social constructionism</a></li>

<li><a href="Social_determinism" title="Social determinism">Social determinism</a></li>

<li><a href="Standard_social_science_model" title="Standard social science model">Standard social science model</a></li></ul></li>

<li><a href="Functionalism_(philosophy_of_mind)" title="Functionalism (philosophy of mind)">Functionalism</a></li>

<li><a href="Memetics" title="Memetics">Memetics</a></li>

<li><a href="Multilineal_evolution" title="Multilineal evolution">Multilineal evolution</a></li>

<li><a href="Neo-Darwinism" title="Neo-Darwinism">Neo-Darwinism</a></li>

<li><a href="Neoevolutionism" title="Neoevolutionism">Neoevolutionism</a></li>

<li><a href="Sociocultural_evolution" title="Sociocultural evolution">Sociocultural evolution</a></li>

<li><a href="Unilineal_evolution" title="Unilineal evolution">Unilineal evolution</a></li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div>

<ul><li><a href="List_of_evolutionary_psychologists" title="List of evolutionary psychologists">Evolutionary psychologists</a></li>

<li> Evolutionary psychology</li>

<li> <a href="Portal%3APsychology" title="Portal:Psychology">Psychology portal</a></li>

<li> <a href="Portal%3AEvolutionary_biology" title="Portal:Evolutionary biology">Evolutionary biology portal</a></li></ul>

</div></td></tr></tbody></table></div>

<div class="navbox-styles"></div><div role="navigation" class="navbox" aria-labelledby="Population_genetics292" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2" style="text-align:center;"><div id="Population_genetics292" style="font-size:114%;margin:0 4em"><a href="Population_genetics" title="Population genetics">Population genetics</a></div></th></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%">Key concepts</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Hardy%E2%80%93Weinberg_principle" title="Hardy–Weinberg principle">Hardy–Weinberg principle</a></li>

<li><a href="Genetic_linkage" title="Genetic linkage">Genetic linkage</a></li>

<li><a href="Identity_by_descent" title="Identity by descent">Identity by descent</a></li>

<li><a href="Linkage_disequilibrium" title="Linkage disequilibrium">Linkage disequilibrium</a></li>

<li><a href="Fisher's_fundamental_theorem_of_natural_selection" title="Fisher's fundamental theorem of natural selection">Fisher's fundamental theorem</a></li>

<li><a href="Neutral_theory_of_molecular_evolution" title="Neutral theory of molecular evolution">Neutral theory</a></li>

<li><a href="Shifting_balance_theory" title="Shifting balance theory">Shifting balance theory</a></li>

<li><a href="Price_equation" title="Price equation">Price equation</a></li>

<li><a href="Coefficient_of_inbreeding" title="Coefficient of inbreeding">Coefficient of inbreeding</a></li>

<li><a href="Coefficient_of_relationship" title="Coefficient of relationship">Coefficient of relationship</a></li>

<li><a href="Selection_coefficient" title="Selection coefficient">Selection coefficient</a></li>

<li><a href="Fitness_(biology)" title="Fitness (biology)">Fitness</a></li>

<li><a href="Heritability" title="Heritability">Heritability</a></li>

<li><a href="Population_structure_(genetics)" title="Population structure (genetics)">Population structure</a></li>

<li><a href="Constructive_neutral_evolution" title="Constructive neutral evolution">Constructive neutral evolution</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%"></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul>

<li><a href="Selective_breeding" title="Selective breeding">Artificial</a></li>

<li><a href="Sexual_selection" title="Sexual selection">Sexual</a></li>

<li><a href="Ecological_selection" class="mw-redirect" title="Ecological selection">Ecological</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%"><div style="display: inline-block; line-height: 1.2em; padding: .1em 0; line-height:1.1em;">Effects of selection on genomic variation</div></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Genetic_hitchhiking" title="Genetic hitchhiking">Genetic hitchhiking</a></li>

<li><a href="Negative_selection_(natural_selection)" title="Negative selection (natural selection)">Background selection</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%"><a href="Genetic_drift" title="Genetic drift">Genetic drift</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Small_population_size" title="Small population size">Small population size</a></li>

<li><a href="Population_bottleneck" title="Population bottleneck">Population bottleneck</a></li>

<li><a href="Founder_effect" title="Founder effect">Founder effect</a></li>

<li><a href="Coalescent_theory" title="Coalescent theory">Coalescence</a></li>

<li><a href="Balding%E2%80%93Nichols_model" title="Balding–Nichols model">Balding–Nichols model</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%">Founders</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Ronald_Fisher" title="Ronald Fisher">R. A. Fisher</a></li>

<li><a href="J._B._S._Haldane" title="J. B. S. Haldane">J. B. S. Haldane</a></li>

<li><a href="Sewall_Wright" title="Sewall Wright">Sewall Wright</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%">Related topics</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Biogeography" title="Biogeography">Biogeography</a></li>

<li><a href="Evolution" title="Evolution">Evolution</a></li>

<li><a href="Evolutionary_game_theory" title="Evolutionary game theory">Evolutionary game theory</a></li>

<li><a href="Fitness_landscape" title="Fitness landscape">Fitness landscape</a></li>

<li><a href="Genetic_genealogy" title="Genetic genealogy">Genetic genealogy</a></li>

<li><a href="Landscape_genetics" title="Landscape genetics">Landscape genetics</a> and <a href="Landscape_genomics" title="Landscape genomics">genomics</a></li>

<li><a href="Microevolution" title="Microevolution">Microevolution</a></li>

<li><a href="Population_genomics" title="Population genomics">Population genomics</a></li>

<li><a href="Phylogeography" title="Phylogeography">Phylogeography</a></li>

<li><a href="Quantitative_genetics" title="Quantitative genetics">Quantitative genetics</a></li></ul>

</div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="text-align:center;"><div>

<ul><li><a href="Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index of evolutionary biology articles</a></li></ul>

</div></td></tr></tbody></table></div>

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This article is issued from <a class="external text" title="Last edited on 2025-08-07" href="https://en.wikipedia.org/wiki/?title=Natural_selection&oldid=1304724438">Wikipedia</a>. The text is available under <a class="external text" href="https://creativecommons.org/licenses/by-sa/4.0/deed.en">Creative Commons Attribution-Share Alike 4.0</a> unless otherwise noted. Additional terms may apply for the media files.

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