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Human evolution

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</style><div role="note" class="hatnote navigation-not-searchable">"Anthropogeny" redirects here. For the study of humans, see <a href="Anthropology" title="Anthropology">Anthropology</a>.</div>

<a href="Homo_sapiens" class="mw-redirect" title="Homo sapiens">Homo sapiens</a> is a distinct species of the <a href="Hominid" class="mw-redirect" title="Hominid">hominid</a> family of <a href="Primates" class="mw-redirect" title="Primates">primates</a>, which also includes all the <a href="Great_ape" class="mw-redirect" title="Great ape">great apes</a>.<a href="#cite_note-1">[1]</a> Over their <a href="Evolution" title="Evolution">evolutionary</a> history, humans gradually developed traits such as <a href="Human_skeletal_changes_due_to_bipedalism" title="Human skeletal changes due to bipedalism">bipedalism</a>, <a href="Dexterity" class="mw-redirect" title="Dexterity">dexterity</a>, and <a href="Complex_language" class="mw-redirect" title="Complex language">complex language</a>,<a href="#cite_note-HallHallgrímsson2011c-2">[2]</a> as well as interbreeding with other <a href="Hominins" class="mw-redirect" title="Hominins">hominins</a> (a tribe of the <a href="Homininae" title="Homininae">African hominid</a> subfamily),<a href="#cite_note-:3-3">[3]</a> indicating that human evolution was not linear but weblike.<a href="#cite_note-Ackermann_2015-4">[4]</a><a href="#cite_note-Denisovans_&_Neandertals-5">[5]</a><a href="#cite_note-Human_Hybrids-6">[6]</a><a href="#cite_note-Mosaic_humans,_the_hybrid_species-7">[7]</a> The study of the origins of humans involves <a href="Interdisciplinary" class="mw-redirect" title="Interdisciplinary">several</a> scientific disciplines, including <a href="Biological_anthropology" title="Biological anthropology">physical</a> and <a href="Evolutionary_anthropology" title="Evolutionary anthropology">evolutionary anthropology</a>, <a href="Paleontology" title="Paleontology">paleontology</a>, and <a href="Genetics" title="Genetics">genetics</a>; the field is also known by the terms anthropogeny, anthropogenesis, and anthropogony<a href="#cite_note-8">[8]</a><a href="#cite_note-Marlowe_54–67-9">[9]</a>—with the latter two sometimes used to refer to the related subject of <a href="Hominization" title="Hominization">hominization</a>.

<a href="Primate_evolution" class="mw-redirect" title="Primate evolution">Primates diverged</a> from other <a href="Mammal" title="Mammal">mammals</a> about <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=85">85</a> million years ago (<a href="Myr" class="mw-redirect" title="Myr">mya</a>), in the <a href="Late_Cretaceous" title="Late Cretaceous">Late Cretaceous</a> period, with their earliest fossils appearing over 55 mya, during the <a href="Paleocene" title="Paleocene">Paleocene</a>.<a href="#cite_note-10">[10]</a> Primates produced successive clades leading to the <a href="Ape" title="Ape">ape</a> superfamily, which gave rise to the hominid and the <a href="Gibbon" title="Gibbon">gibbon</a> families; these diverged some 15–20 mya. African and <a href="Ponginae" title="Ponginae">Asian hominids</a> (including <a href="Orangutan" title="Orangutan">orangutans</a>) diverged about 14 mya. <a href="Hominini" title="Hominini">Hominins</a> (including the <a href="Australopithecine" title="Australopithecine">Australopithecine</a> and <a href="Panina" title="Panina">Panina</a> subtribes) parted from the <a href="Gorillini" title="Gorillini">Gorillini</a> tribe between 8 and 9 mya; Australopithecine (including the extinct biped ancestors of humans) separated from the Pan genus (containing <a href="Chimpanzee" title="Chimpanzee">chimpanzees</a> and <a href="Bonobo" title="Bonobo">bonobos</a>) 4–7 mya.<a href="#cite_note-:4-11">[11]</a> The <a href="Homo" title="Homo">Homo</a> genus is evidenced by the appearance of <a href="H._habilis" class="mw-redirect" title="H. habilis">H. habilis</a> over 2 mya,<a href="#cite_note-habilis-12">[a]</a> while <a href="Anatomically_modern_human" class="mw-redirect" title="Anatomically modern human">anatomically modern humans</a> emerged in Africa approximately 300,000 years ago.

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<div class="mw-heading mw-heading2"><h2 id="Before_Homo">Before Homo</h2></div>

<div role="note" class="hatnote navigation-not-searchable">For evolutionary history before primates, see <a href="Evolution_of_mammals" title="Evolution of mammals">Evolution of mammals</a>, <a href="History_of_life" title="History of life">History of life</a>, and <a href="Timeline_of_human_evolution" title="Timeline of human evolution">Timeline of human evolution</a>.</div>

<div class="mw-heading mw-heading3"><h3 id="Early_evolution_of_primates">Early evolution of primates</h3></div>

<div role="note" class="hatnote navigation-not-searchable">See also: <a href="Evolution_of_primates" title="Evolution of primates">Evolution of primates</a></div>

The evolutionary history of primates can be traced back 65 million years.<a href="#cite_note-FOOTNOTEMaxwell1984296-13">[12]</a><a href="#cite_note-14">[13]</a><a href="#cite_note-15">[14]</a><a href="#cite_note-FOOTNOTEMartin200112032–12038-16">[15]</a><a href="#cite_note-17">[16]</a> One of the oldest known primate-like mammal species, the <a href="Plesiadapis" title="Plesiadapis">Plesiadapis</a>, came from North America;<a href="#cite_note-18">[17]</a><a href="#cite_note-19">[18]</a><a href="#cite_note-20">[19]</a><a href="#cite_note-21">[20]</a><a href="#cite_note-22">[21]</a><a href="#cite_note-23">[22]</a> another, <a href="Archicebus" title="Archicebus">Archicebus</a>, came from <a href="China" title="China">China</a>.<a href="#cite_note-NYT-20130605-24">[23]</a> Other similar basal primates were widespread in Eurasia and Africa during the tropical conditions of the Paleocene and <a href="Eocene" title="Eocene">Eocene</a>.

David R. Begun<a href="#cite_note-Kordos-p17-25">[24]</a> concluded that early primates flourished in Eurasia and that a lineage leading to the African apes and humans, including to <a href="Dryopithecus" title="Dryopithecus">Dryopithecus</a>, migrated south from Europe or Western Asia into Africa. The surviving tropical population of primates—which is seen most completely in the Upper Eocene and lowermost <a href="Oligocene" title="Oligocene">Oligocene</a> fossil beds of the <a href="Faiyum" title="Faiyum">Faiyum</a> depression southwest of <a href="Cairo" title="Cairo">Cairo</a>—gave rise to all extant primate species, including the <a href="Lemur" title="Lemur">lemurs</a> of <a href="Madagascar" title="Madagascar">Madagascar</a>, <a href="Loris" title="Loris">lorises</a> of Southeast Asia, <a href="Galago" title="Galago">galagos</a> or "bush babies" of Africa, and to the <a href="Simian" title="Simian">anthropoids</a>, which are the <a href="New_World_monkey" title="New World monkey">Platyrrhines</a> or New World monkeys, the <a href="Catarrhines" class="mw-redirect" title="Catarrhines">Catarrhines</a> or Old World monkeys, and the great apes, including humans and other hominids.

The earliest known catarrhine is <a href="Kamoyapithecus" title="Kamoyapithecus">Kamoyapithecus</a> from the uppermost Oligocene at Eragaleit in the northern <a href="Great_Rift_Valley%2C_Kenya" title="Great Rift Valley, Kenya">Great Rift Valley</a> in Kenya, dated to 24 million years ago.<a href="#cite_note-FOOTNOTECameron2004[httpsbooksgooglecombooksidfnqzb4_UVfkCpgPA76_76]-26">[25]</a> Its ancestry is thought to be species related to <a href="Aegyptopithecus" title="Aegyptopithecus">Aegyptopithecus</a>, <a href="Propliopithecus" title="Propliopithecus">Propliopithecus</a>, and <a href="Parapithecus" title="Parapithecus">Parapithecus</a> from the Faiyum, at around 35 mya.<a href="#cite_note-FOOTNOTEWallace2004[httpsbooksgooglecombooksidYoyQQEOKGVQCpgPA240_240]-27">[26]</a> In 2010, <a href="Saadanius" title="Saadanius">Saadanius</a> was described as a close relative of the last common ancestor of the <a href="Crown_group" title="Crown group">crown</a> catarrhines, and tentatively dated to 29–28 mya, helping to fill an 11-million-year gap in the fossil record.<a href="#cite_note-2010Zalmout-28">[27]</a>

In the <a href="Early_Miocene" title="Early Miocene">Early Miocene</a>, about 22 million years ago, the many kinds of <a href="Arboreal_locomotion" title="Arboreal locomotion">arboreally</a>-adapted (tree-dwelling) primitive catarrhines from East Africa suggest a long history of prior diversification. <a href="Fossil" title="Fossil">Fossils</a> at 20 million years ago include fragments attributed to <a href="Victoriapithecus" title="Victoriapithecus">Victoriapithecus</a>, the earliest Old World monkey. Among the genera thought to be in the <a href="Ape" title="Ape">ape</a> lineage leading up to 13 million years ago are <a href="Proconsul_(mammal)" title="Proconsul (mammal)">Proconsul</a>, <a href="Rangwapithecus" title="Rangwapithecus">Rangwapithecus</a>, <a href="Dendropithecus" title="Dendropithecus">Dendropithecus</a>, Limnopithecus, <a href="Nacholapithecus" title="Nacholapithecus">Nacholapithecus</a>, <a href="Equatorius" title="Equatorius">Equatorius</a>, <a href="Nyanzapithecus_pickfordi" class="mw-redirect" title="Nyanzapithecus pickfordi">Nyanzapithecus</a>, <a href="Afropithecus" title="Afropithecus">Afropithecus</a>, Heliopithecus, and <a href="Kenyapithecus" title="Kenyapithecus">Kenyapithecus</a>, all from East Africa.

The presence of other generalized non-cercopithecids of <a href="Middle_Miocene" title="Middle Miocene">Middle Miocene</a> from sites far distant, such as <a href="Otavipithecus" title="Otavipithecus">Otavipithecus</a> from cave deposits in Namibia, and <a href="Pierolapithecus" title="Pierolapithecus">Pierolapithecus</a> and <a href="Dryopithecus" title="Dryopithecus">Dryopithecus</a> from France, Spain and Austria, is evidence of a wide diversity of forms across Africa and the Mediterranean basin during the relatively warm and equable climatic regimes of the Early and Middle Miocene. The youngest of the <a href="Miocene" title="Miocene">Miocene</a> hominoids, <a href="Oreopithecus" title="Oreopithecus">Oreopithecus</a>, is from coal beds in Italy that have been dated to 9 million years ago.

Molecular evidence indicates that the lineage of gibbons diverged from the line of great apes some 18–12 mya, and that of orangutans (subfamily <a href="Ponginae" title="Ponginae">Ponginae</a>)<a href="#cite_note-29">[b]</a> diverged from the other great apes at about 12 million years; there are no fossils that clearly document the ancestry of gibbons, which may have originated in a so-far-unknown Southeast Asian hominoid population, but fossil proto-orangutans may be represented by <a href="Sivapithecus" title="Sivapithecus">Sivapithecus</a> from India and <a href="Griphopithecus" title="Griphopithecus">Griphopithecus</a> from Turkey, dated to around 10 mya.<a href="#cite_note-FOOTNOTESrivastava2009[httpsbooksgooglecombooksidkCerOsM8XMwCpgPA87_87]-30">[28]</a>

Hominidae subfamily <a href="Homininae" title="Homininae">Homininae</a> (African hominids) diverged from Ponginae (orangutans) about 14 mya. Hominins (including humans and the Australopithecine and <a href="Panina" title="Panina">Panina</a> subtribes) parted from the <a href="Gorillini" title="Gorillini">Gorillini</a> tribe (gorillas) between 8 and 9 mya; Australopithecine (including the extinct biped ancestors of humans) separated from the Pan genus (containing chimpanzees and bonobos) 4–7 mya.<a href="#cite_note-:4-11">[11]</a> The Homo genus is evidenced by the appearance of H. habilis over 2 mya,<a href="#cite_note-habilis-12">[a]</a> while anatomically modern humans emerged in Africa approximately 300,000 years ago.

<div class="mw-heading mw-heading3"><h3 id="Divergence_of_the_human_clade_from_other_great_apes">Divergence of the human clade from other great apes</h3></div>

Species close to the last common ancestor of gorillas, chimpanzees and humans may be represented by <a href="Nakalipithecus" title="Nakalipithecus">Nakalipithecus</a> fossils found in Kenya and <a href="Ouranopithecus" title="Ouranopithecus">Ouranopithecus</a> found in Greece. Molecular evidence suggests that between 8 and 4 million years ago, first the gorillas, and then the chimpanzees (genus Pan) split off from the line leading to the humans. Human DNA is approximately 98.4% identical to that of chimpanzees when comparing single nucleotide polymorphisms (see <a href="Human_evolutionary_genetics" title="Human evolutionary genetics">human evolutionary genetics</a>). The fossil record, however, of gorillas and chimpanzees is limited; both poor preservation – rain forest soils tend to be acidic and dissolve bone – and <a href="Sampling_bias" title="Sampling bias">sampling bias</a> probably contribute to this problem.

Other hominins probably adapted to the drier environments outside the equatorial belt; and there they encountered antelope, hyenas, dogs, pigs, elephants, horses, and others. The equatorial belt contracted after about 8 million years ago, and there is very little fossil evidence for the split—thought to have occurred around that time—of the hominin lineage from the lineages of gorillas and chimpanzees. The earliest fossils argued by some to belong to the human lineage are Sahelanthropus tchadensis (7 Ma) and Orrorin tugenensis (6 Ma), followed by Ardipithecus (5.5–4.4 Ma), with species Ar. kadabba and <a href="Ar._ramidus" class="mw-redirect" title="Ar. ramidus">Ar. ramidus</a>.

It has been argued in a study of the life history of Ar. ramidus that the species provides evidence for a suite of anatomical and behavioral adaptations in very early hominins unlike any species of extant great ape.<a href="#cite_note-aramidus-32">[30]</a> This study demonstrated affinities between the skull morphology of Ar. ramidus and that of infant and juvenile chimpanzees, suggesting the species evolved a juvenalised or <a href="Paedomorphic" class="mw-redirect" title="Paedomorphic">paedomorphic</a> craniofacial morphology via <a href="Heterochrony" title="Heterochrony">heterochronic</a> dissociation of growth trajectories. It was also argued that the species provides support for the notion that very early hominins, akin to bonobos (Pan paniscus) the less aggressive species of the genus Pan, may have evolved via the process of <a href="Self-domestication" title="Self-domestication">self-domestication</a>. Consequently, arguing against the so-called "chimpanzee referential model"<a href="#cite_note-sayers-33">[31]</a> the authors suggest it is no longer tenable to use chimpanzee (Pan troglodytes) social and mating behaviors in models of early hominin social evolution. When commenting on the absence of aggressive canine morphology in Ar. ramidus and the implications this has for the evolution of hominin social psychology, they wrote:

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</style><blockquote class="templatequote">Of course Ar. ramidus differs significantly from bonobos, bonobos having retained a functional canine honing complex. However, the fact that Ar. ramidus shares with bonobos reduced sexual dimorphism, and a more paedomorphic form relative to chimpanzees, suggests that the developmental and social adaptations evident in bonobos may be of assistance in future reconstructions of early hominin social and sexual psychology. In fact the trend towards increased maternal care, female mate selection and self-domestication may have been stronger and more refined in Ar. ramidus than what we see in bonobos.<a href="#cite_note-aramidus-32">[30]</a>: 128 </blockquote>

The authors argue that many of the basic human adaptations evolved in the ancient forest and woodland ecosystems of late <a href="Miocene" title="Miocene">Miocene</a> and early <a href="Pliocene" title="Pliocene">Pliocene</a> Africa. Consequently, they argue that humans may not represent evolution from a chimpanzee-like ancestor as has traditionally been supposed. This suggests many modern human adaptations represent <a href="Phylogeny" class="mw-redirect" title="Phylogeny">phylogenetically</a> deep traits and that the behavior and morphology of chimpanzees may have evolved subsequent to the split with the common ancestor they share with humans.

<div class="mw-heading mw-heading3"><h3 id="Genus_Australopithecus">Genus Australopithecus</h3></div>

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

The genus <a href="Australopithecus" title="Australopithecus">Australopithecus</a> evolved in eastern Africa around 4 million years ago before spreading throughout the continent and eventually becoming extinct 2 million years ago. During this time period various forms of australopiths existed, including <a href="Australopithecus_anamensis" title="Australopithecus anamensis">Australopithecus anamensis</a>, <a href="Australopithecus_afarensis" title="Australopithecus afarensis">A. afarensis</a>, <a href="Australopithecus_sediba" title="Australopithecus sediba">A. sediba</a>, and <a href="Australopithecus_africanus" title="Australopithecus africanus">A. africanus</a>. There is still some debate among academics whether certain African hominid species of this time, such as <a href="Australopithecus_robustus" class="mw-redirect" title="Australopithecus robustus">A. robustus</a> and <a href="Australopithecus_boisei" class="mw-redirect" title="Australopithecus boisei">A. boisei</a>, constitute members of the same genus; if so, they would be considered to be "robust australopiths" while the others would be considered "gracile australopiths". However, if these species do indeed constitute their own genus, then they may be given their own name, Paranthropus.

<ul><li><a href="Australopithecus" title="Australopithecus">Australopithecus</a> (4–1.8 Ma), with species <a href="Australopithecus_anamensis" title="Australopithecus anamensis">A. anamensis</a>, <a href="Australopithecus_afarensis" title="Australopithecus afarensis">A. afarensis</a>, <a href="Australopithecus_africanus" title="Australopithecus africanus">A. africanus</a>, <a href="Australopithecus_bahrelghazali" title="Australopithecus bahrelghazali">A. bahrelghazali</a>, <a href="Australopithecus_garhi" title="Australopithecus garhi">A. garhi</a>, and <a href="Australopithecus_sediba" title="Australopithecus sediba">A. sediba</a>;</li>

<li><a href="Kenyanthropus" title="Kenyanthropus">Kenyanthropus</a> (3–2.7 Ma), with species <a href="Kenyanthropus" title="Kenyanthropus">K. platyops</a>;</li>

<li><a href="Paranthropus" title="Paranthropus">Paranthropus</a> (3–1.2 Ma), with species <a href="Paranthropus_aethiopicus" title="Paranthropus aethiopicus">P. aethiopicus</a>, <a href="Paranthropus_boisei" title="Paranthropus boisei">P. boisei</a>, and <a href="Paranthropus_robustus" title="Paranthropus robustus">P. robustus</a></li></ul>

A new proposed species <a href="Australopithecus_deyiremeda" title="Australopithecus deyiremeda">Australopithecus deyiremeda</a> is claimed to have been discovered living at the same time period of <a href="Australopithecus_afarensis" title="Australopithecus afarensis">A. afarensis</a>. There is debate whether <a href="Australopithecus_deyiremeda" title="Australopithecus deyiremeda">A. deyiremeda</a> is a new species or is <a href="Australopithecus_afarensis" title="Australopithecus afarensis">A. afarensis</a>.<a href="#cite_note-34">[32]</a> Australopithecus prometheus, otherwise known as <a href="Little_Foot" title="Little Foot">Little Foot</a>, has recently been dated at 3.67 million years old through a new dating technique, making the genus Australopithecus as old as afarensis.<a href="#cite_note-35">[33]</a> Given the opposable big toe found on Little Foot, it seems that the specimen was a good climber. It is thought given the night predators of the region that he built a nesting platform at night in the trees in a similar fashion to chimpanzees and gorillas.

<div class="mw-heading mw-heading2"><h2 id="Evolution_of_genus_Homo">Evolution of genus Homo</h2></div>

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

<table id="Container" role="presentation" class="nomobile toccolours searchaux" style="text-align:left;padding:0 0.5em;border-style:solid;float:right;clear:right;margin:0.3em 0 0.8em 1.4em;overflow:hidden;min-width:29.760em"><tbody><tr><td colspan="3" id="Title" style="padding:1em 1em 0 1em"><div style="background-color:#77bb77;padding:0 0.2em 0 0.2em;font-weight:bold;text-align:center"><a href="Timeline_of_human_evolution" title="Timeline of human evolution">Hominin timeline</a></div></td></tr><tr><td colspan="3" id="Navbox" style="padding:0;margin:0 1em 0 0;text-align:right"><style data-mw-deduplicate="TemplateStyles:r1129693374">

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</style></td></tr><tr style="vertical-align:top"><td id="Scale" style="padding:0;margin:0.7em 0 0.7em 0;width:4.2em;position:relative;float:left;font-size:100%;height:50em;border-right:1px solid #242020"><div style="float:right;position:absolute;right:-1px;top:50.00em;transform:translateY(-50%)">−10 —</div><div style="float:right;position:absolute;right:-1px;top:47.50em;transform:translateY(-50%)">–</div><div style="float:right;position:absolute;right:-1px;top:45.00em;transform:translateY(-50%)">−9 —</div><div style="float:right;position:absolute;right:-1px;top:42.50em;transform:translateY(-50%)">–</div><div style="float:right;position:absolute;right:-1px;top:40.00em;transform:translateY(-50%)">−8 —</div><div style="float:right;position:absolute;right:-1px;top:37.50em;transform:translateY(-50%)">–</div><div style="float:right;position:absolute;right:-1px;top:35.00em;transform:translateY(-50%)">−7 —</div><div style="float:right;position:absolute;right:-1px;top:32.50em;transform:translateY(-50%)">–</div><div style="float:right;position:absolute;right:-1px;top:30.00em;transform:translateY(-50%)">−6 —</div><div style="float:right;position:absolute;right:-1px;top:27.50em;transform:translateY(-50%)">–</div><div style="float:right;position:absolute;right:-1px;top:25.00em;transform:translateY(-50%)">−5 —</div><div style="float:right;position:absolute;right:-1px;top:22.50em;transform:translateY(-50%)">–</div><div style="float:right;position:absolute;right:-1px;top:20.00em;transform:translateY(-50%)">−4 —</div><div style="float:right;position:absolute;right:-1px;top:17.50em;transform:translateY(-50%)">–</div><div style="float:right;position:absolute;right:-1px;top:15.00em;transform:translateY(-50%)">−3 —</div><div style="float:right;position:absolute;right:-1px;top:12.50em;transform:translateY(-50%)">–</div><div style="float:right;position:absolute;right:-1px;top:10.00em;transform:translateY(-50%)">−2 —</div><div style="float:right;position:absolute;right:-1px;top:7.50em;transform:translateY(-50%)">–</div><div style="float:right;position:absolute;right:-1px;top:5.00em;transform:translateY(-50%)">−1 —</div><div style="float:right;position:absolute;right:-1px;top:2.50em;transform:translateY(-50%)">–</div><div style="float:right;position:absolute;right:-1px;top:-0.00em;transform:translateY(-50%)">0 —</div></td><td id="Timeline" class="toccolours notheme" style="padding:0px;margin:0.7em 0 0.7em 0;position:relative;font-size:100%;width:13em;height:50em;float:left;border:none;background-color:#FFE4F1"><div class="notheme" style="font-size:100%;background-color:#936a00;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:26.665em;height:23.335em;left:0.000em;width:1.300em"><div style="font-size:100%;position:relative;top:0em;left:-0.1em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-11.667em;transform:translateY(-50%);z-index:5;pointer-events:initial"><style data-mw-deduplicate="TemplateStyles:r1231500821">

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</style><a href="Miocene" title="Miocene">Miocene</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#f1c309;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:12.900em;height:13.765em;left:0.000em;width:1.300em"><div style="font-size:100%;position:relative;top:0.5em;left:-0.1em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-6.883em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Pliocene" title="Pliocene">Pliocene</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#c1e0c1;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.058em;height:12.841em;left:0.000em;width:1.300em"><div style="font-size:100%;position:relative;top:0.75em;left:-0.1em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-6.421em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Pleistocene" title="Pleistocene">Pleistocene</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:14.000em;height:36.000em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:-6.7em;left:-0.3em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-18.000em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Hominini" title="Hominini">Hominini</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:48.500em;height:0.500em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:0em;left:0.1em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.250em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Nakalipithecus" title="Nakalipithecus">Nakalipithecus</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:46.500em;height:1.000em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:0.5em;left:0.6em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.500em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Samburupithecus" title="Samburupithecus">Samburupithecus</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:44.500em;height:0.500em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:0em;left:0.2em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.250em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Ouranopithecus" title="Ouranopithecus">Ouranopithecus</a> (<a href="Ouranopithecus_turkae" title="Ouranopithecus turkae">Ou. turkae</a>) (<a href="Ouranopithecus_macedoniensis" title="Ouranopithecus macedoniensis">Ou. macedoniensis</a>)</div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:39.500em;height:0.500em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:0.1em;left:0.4em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.250em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Chororapithecus" title="Chororapithecus">Chororapithecus</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:35.000em;height:10.000em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:-1.75em;left:-0.2em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-5.000em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Oreopithecus" title="Oreopithecus">Oreopithecus</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:39.500em;height:0.500em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:1.75em;left:-0.4em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.250em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Sivapithecus" title="Sivapithecus">Sivapithecus</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:34.500em;height:0.500em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:0.5em;left:0.3em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.250em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Sahelanthropus" title="Sahelanthropus">Sahelanthropus</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:34.500em;height:0.500em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:2em;left:0.3em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.250em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Graecopithecus" title="Graecopithecus">Graecopithecus</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:29.500em;height:0.500em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:0.5em;left:-1.5em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.250em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Orrorin" title="Orrorin">Orrorin</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:29.500em;height:0.500em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:2.5em;left:0.15em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.250em;transform:translateY(-50%);z-index:5;pointer-events:initial">(<a href="Orrorin" title="Orrorin">O. praegens</a>) (<a href="Orrorin" title="Orrorin">O. tugenensis</a>)</div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:22.000em;height:6.850em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:-3.75em;left:-0.3em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-3.425em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Ardipithecus" title="Ardipithecus">Ardipithecus</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:27.700em;height:1.150em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:0.75em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.575em;transform:translateY(-50%);z-index:5;pointer-events:initial">(<a href="Ardipithecus_kadabba" title="Ardipithecus kadabba">Ar. kadabba</a>)</div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:21.500em;height:1.000em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:1em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.500em;transform:translateY(-50%);z-index:5;pointer-events:initial">(<a href="Ardipithecus_ramidus" title="Ardipithecus ramidus">Ar. ramidus</a>)</div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:6.000em;height:16.500em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:3em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-8.250em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Australopithecus" title="Australopithecus">Australopithecus</a> (<a href="Australopithecus_africanus" title="Australopithecus africanus">Au. africanus</a>) (<a href="Australopithecus_afarensis" title="Australopithecus afarensis">Au. afarensis</a>) (<a href="Australopithecus_anamensis" title="Australopithecus anamensis">Au. anamensis</a>)</div></div></div><div class="notheme" style="font-size:100%;background-color:#DCEEFF;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:8.250em;height:5.750em;left:2.600em;width:10.400em"><div style="font-size:100%;position:relative;top:0.9em;left:-0.5em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-2.875em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Homo_habilis" title="Homo habilis">H. habilis</a> (<a href="Homo_rudolfensis" title="Homo rudolfensis">H. rudolfensis</a>) (<a href="Australopithecus_garhi" title="Australopithecus garhi">Au. garhi</a>)</div></div></div><div class="notheme" style="font-size:100%;background-color:#FFE4F1;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.570em;height:9.430em;left:3.900em;width:9.100em"><div style="font-size:100%;position:relative;top:1.5em;left:-1em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-4.715em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Homo_erectus" title="Homo erectus">H. erectus</a> (<a href="Homo_antecessor" title="Homo antecessor">H. antecessor</a>) (<a href="Homo_ergaster" title="Homo ergaster">H. ergaster</a>) (<a href="Australopithecus_sediba" title="Australopithecus sediba">Au. sediba</a>)</div></div></div><div class="notheme" style="font-size:100%;background-color:#FFE4F1;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:6.000em;height:2.500em;left:2.600em;width:10.400em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-1.250em;transform:translateY(-50%);z-index:5;pointer-events:initial"></div></div></div><div class="notheme" style="font-size:100%;background-color:#7DFFFF;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:1.000em;height:2.500em;left:2.600em;width:10.400em"><div style="font-size:100%;position:relative;top:1em;left:-0.2em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-1.250em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Homo_heidelbergensis" title="Homo heidelbergensis">H. heidelbergensis</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#F3F300;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.000em;height:1.500em;left:3.900em;width:9.100em"><div style="font-size:100%;position:relative;top:0em;left:-0.7em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.750em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Anatomically_modern_human" class="mw-redirect" title="Anatomically modern human">Homo sapiens</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#F3F300;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.000em;height:0.200em;left:1.300em;width:3.900em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.100em;transform:translateY(-50%);z-index:5;pointer-events:initial"></div></div></div><div class="notheme" style="font-size:100%;background-color:#7DFFFF;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.200em;height:1.050em;left:2.600em;width:1.300em"><div style="font-size:100%;position:relative;top:1.8em;left:2.6em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.525em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Neanderthal" title="Neanderthal">Neanderthals</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#F3F300;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.200em;height:0.370em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.185em;transform:translateY(-50%);z-index:5;pointer-events:initial"></div></div></div><div class="notheme" style="font-size:100%;background-color:#7DFFFF;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.200em;height:0.370em;left:2.600em;width:1.300em"><div style="font-size:100%;position:relative;top:1.4em;left:2.8em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.185em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="Denisovan" title="Denisovan">Denisovans</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:6.000em;height:8.500em;left:11.700em;width:1.300em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-4.250em;transform:translateY(-50%);z-index:5;pointer-events:initial"> </div></div></div></td><td id="Annotations" style="padding:0;margin:0.7em 0 0.7em 0;float:left;position:relative;width:10.860em;height:50em;pointer-events:none"><table role="presentation" style="position:absolute;z-index:15;top:50.000em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle">←</td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="Timeline_of_human_evolution#Hominidae" title="Timeline of human evolution">Earlier apes</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:45.000em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle">←</td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="Gorilla%E2%80%93human_last_common_ancestor" title="Gorilla–human last common ancestor">Gorilla split</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:42.500em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle">←</td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="Chimpanzee%E2%80%93human_last_common_ancestor" title="Chimpanzee–human last common ancestor">Chimpanzee split</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:35.000em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle">←</td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="Human_skeletal_changes_due_to_bipedalism" title="Human skeletal changes due to bipedalism">Earliest bipedal</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:29.000em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle">←</td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="Ardipithecus_kadabba" title="Ardipithecus kadabba">Earliest sign of Ardipithecus</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:22.500em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle">←</td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="Australopithecus_anamensis" title="Australopithecus anamensis">Earliest sign of Australopithecus</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:17.000em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle">←</td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="Stone_Age#Beginning_of_the_Stone_Age" title="Stone Age">Earliest stone tools</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:14.000em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle">←</td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.25em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="LD_350-1" title="LD 350-1">Earliest sign of Homo</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:10.600em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle">←</td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.25em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="Early_expansions_of_hominins_out_of_Africa" title="Early expansions of hominins out of Africa">Dispersal beyond Africa</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:8.500em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle">←</td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="Control_of_fire_by_early_humans" title="Control of fire by early humans">Earliest fire</a> / <a href="Cooking#History" title="Cooking">cooking</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:1.000em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle">←</td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.6em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="Rock_art#East_Asia" title="Rock art">Earliest rock art</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:0.625em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle">←</td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.1em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="Clothing#Origin_and_history" title="Clothing">Earliest clothes</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:0.250em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle">←</td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:-0.4em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="Behavioral_modernity" title="Behavioral modernity">Modern humans</a></div></div></td></tr></tbody></table><div class="annot-bar" style="width:auto;font-size:100%;position:absolute;text-align:center;pointer-events:none;margin-top:21.000em"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;text-align:left;pointer-events:none;top:0.25em;left:-5em;width:7em"><div style="position:relative;width:auto;z-index:10;font-size:90%;color:var( --color-base, #000);vertical-align:middle;line-height:105%;bottom:0;pointer-events:initial"><a href="Hominidae" title="Hominidae">H o m i n i d s</a></div></div></div><div class="annot-bar" style="width:auto;font-size:100%;position:absolute;text-align:center;pointer-events:none;margin-top:3.000em"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;text-align:left;pointer-events:none;top:3.25em;left:-5em;width:7em"><div style="position:relative;width:auto;z-index:10;font-size:90%;color:var( --color-base, #000);vertical-align:middle;line-height:105%;bottom:0;pointer-events:initial"><a href="Paranthropus" title="Paranthropus">P a r a n t h r o p u s</a></div></div></div></td></tr><tr><td id="Caption" colspan="3" style="padding:0;margin:0 0.2em 0.7em 0.2em"><div id="Caption" class="toccolours" style="margin:0 0.5em;border-style:none;clear:both;text-align:center;width:26.000em"><div style="float:left;font-size:85%;">(<a href="Myr" class="mw-redirect" title="Myr">million years ago</a>)</div></div></td></tr></tbody></table>

The earliest documented representative of the genus Homo is <a href="Homo_habilis" title="Homo habilis">Homo habilis</a>, which evolved around <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=2.8">2.8</a> million years ago,<a href="#cite_note-autogenerated1-36">[34]</a> and is arguably the earliest species for which there is positive evidence of the use of stone tools. The brains of these early hominins were about the same size as that of a chimpanzee, although it has been suggested that this was the time in which the human <a href="SRGAP2" title="SRGAP2">SRGAP2</a> <a href="Gene" title="Gene">gene</a> doubled, producing a more rapid wiring of the frontal cortex. During the next million years a process of rapid <a href="Encephalization" class="mw-redirect" title="Encephalization">encephalization</a> occurred, and with the arrival of <a href="Homo_erectus" title="Homo erectus">Homo erectus</a> and <a href="Homo_ergaster" title="Homo ergaster">Homo ergaster</a> in the <a href="Fossil#Dating" title="Fossil">fossil record</a>, cranial capacity had doubled to 850 cm3.<a href="#cite_note-FOOTNOTESwisherCurtisLewin2001-37">[35]</a> (Such an increase in human brain size is equivalent to each generation having 125,000 more <a href="Neuron" title="Neuron">neurons</a> than their parents.) It is believed that H. erectus and H. ergaster were the first to use fire and complex tools, and were the first of the hominin line to leave Africa, spreading throughout Africa, Asia, and Europe between <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=1.3–1.8">1.3 to 1.8</a> million years ago.

According to the recent African origin theory, modern humans evolved in Africa possibly from <a href="H._heidelbergensis" class="mw-redirect" title="H. heidelbergensis">H. heidelbergensis</a>, <a href="H._rhodesiensis" class="mw-redirect" title="H. rhodesiensis">H. rhodesiensis</a> or <a href="H._antecessor" class="mw-redirect" title="H. antecessor">H. antecessor</a> and migrated out of the continent some 50,000 to 100,000 years ago, gradually replacing local populations of H. erectus, <a href="Denisova_hominin" class="mw-redirect" title="Denisova hominin">Denisova hominins</a>, <a href="H._floresiensis" class="mw-redirect" title="H. floresiensis">H. floresiensis</a>, <a href="H._luzonensis" class="mw-redirect" title="H. luzonensis">H. luzonensis</a> and <a href="H._neanderthalensis" class="mw-redirect" title="H. neanderthalensis">H. neanderthalensis</a>, whose ancestors had left Africa in earlier migrations.<a href="#cite_note-FOOTNOTEStringer1994242-38">[36]</a><a href="#cite_note-FOOTNOTEMcHenry2009265-39">[37]</a><a href="#cite_note-40">[38]</a><a href="#cite_note-41">[39]</a><a href="#cite_note-42">[40]</a> <a href="Archaic_humans" class="mw-redirect" title="Archaic humans">Archaic Homo sapiens</a>, the forerunner of <a href="Anatomically_modern_humans" class="mw-redirect" title="Anatomically modern humans">anatomically modern humans</a>, evolved in the <a href="Middle_Paleolithic" title="Middle Paleolithic">Middle Paleolithic</a> between 400,000 and 250,000 years ago.<a href="#cite_note-43">[41]</a><a href="#cite_note-44">[42]</a><a href="#cite_note-45">[43]</a> Recent <a href="DNA" title="DNA">DNA</a> evidence suggests that several <a href="Haplotype" title="Haplotype">haplotypes</a> of <a href="Neanderthal" title="Neanderthal">Neanderthal</a> origin are present among all non-African populations, and Neanderthals and other hominins, such as Denisovans, may have contributed up to 6% of their <a href="Genome" title="Genome">genome</a> to present-day humans, suggestive of a <a href="Interbreeding_between_archaic_and_modern_humans" title="Interbreeding between archaic and modern humans">limited interbreeding between these species</a>.<a href="#cite_note-pmid21179161-46">[44]</a><a href="#cite_note-pmid20439435-47">[45]</a><a href="#cite_note-10.1126/science.1209202-48">[46]</a> According to some anthropologists, the transition to <a href="Behavioral_modernity" title="Behavioral modernity">behavioral modernity</a> with the development of symbolic culture, language, and specialized <a href="Lithic_technology" title="Lithic technology">lithic technology</a> happened around 50,000 years ago (beginning of the <a href="Upper_Paleolithic" title="Upper Paleolithic">Upper Paleolithic</a>),<a href="#cite_note-49">[47]</a> although others point to evidence of a gradual change over a longer time span during the Middle Paleolithic.<a href="#cite_note-Mcbrearty_Brooks-50">[48]</a>

Homo sapiens is the only <a href="Extant_species" class="mw-redirect" title="Extant species">extant species</a> of its genus, Homo. While some (extinct) Homo species might have been ancestors of Homo sapiens, many, perhaps most, were likely "cousins", having <a href="Speciate" class="mw-redirect" title="Speciate">speciated</a> away from the ancestral hominin line.<a href="#cite_note-52">[50]</a><a href="#cite_note-FOOTNOTEBryson2004522–543-53">[51]</a> There is yet no consensus as to which of these groups should be considered a separate species and which should be subspecies; this may be due to the dearth of fossils or to the slight differences used to classify species in the genus Homo.<a href="#cite_note-FOOTNOTEBryson2004522–543-53">[51]</a> The <a href="Sahara_pump_theory" title="Sahara pump theory">Sahara pump theory</a> (describing an occasionally passable <a href="Wet_Sahara" class="mw-redirect" title="Wet Sahara">"wet" Sahara</a> desert) provides one possible explanation of the intermittent migration and speciation in the genus Homo.

Based on archaeological and paleontological evidence, it has been possible to infer, to some extent, the ancient dietary practices<a href="#cite_note-NYT-20150813-54">[52]</a> of various Homo species and to study the role of diet in physical and behavioral evolution within Homo.<a href="#cite_note-Leonard_2007-55">[53]</a><a href="#cite_note-FOOTNOTEWalker2007[httpsbooksgooglecombooksid6mxZ1hNBHgkCpgPA3_3–10]-56">[54]</a><a href="#cite_note-FOOTNOTEUngarTeaford2002-57">[55]</a><a href="#cite_note-FOOTNOTEBogin1997[httpswebarchiveorgweb20031203003838httpcitdscarutorontocaANTAO1ProjectsBoginhtml_96–142]-58">[56]</a><a href="#cite_note-59">[57]</a>

Some anthropologists and archaeologists subscribe to the <a href="Toba_catastrophe_theory" class="mw-redirect" title="Toba catastrophe theory">Toba catastrophe theory</a>, which posits that the <a href="Supereruption" class="mw-redirect" title="Supereruption">supereruption</a> of <a href="Lake_Toba" title="Lake Toba">Lake Toba</a> on Sumatra in Indonesia some 70,000 years ago caused global starvation,<a href="#cite_note-60">[58]</a> killing the majority of humans and creating a <a href="Population_bottleneck" title="Population bottleneck">population bottleneck</a> that affected the genetic inheritance of all humans today.<a href="#cite_note-61">[59]</a> The genetic and archaeological evidence for this remains in question however.<a href="#cite_note-62">[60]</a> A 2023 genetic study suggests that a similar human <a href="Population_bottleneck" title="Population bottleneck">population bottleneck</a> of between 1,000 and 100,000 survivors occurred "around 930,000 and 813,000 years ago ... lasted for about 117,000 years and brought human ancestors close to extinction."<a href="#cite_note-NYT-20230831-63">[61]</a><a href="#cite_note-SCI-20230831-64">[62]</a>

<div class="mw-heading mw-heading3"><h3 id="H._habilis_and_H._gautengensis">H. habilis and H. gautengensis</h3></div>

Homo habilis lived from about 2.8<a href="#cite_note-autogenerated1-36">[34]</a> to 1.4 Ma. The species evolved in South and East Africa in the <a href="Piacenzian" title="Piacenzian">Late Pliocene</a> or <a href="Early_Pleistocene" title="Early Pleistocene">Early Pleistocene</a>, 2.5–2 Ma, when it diverged from the australopithecines with the development of smaller molars and larger brains. One of the first known hominins, it made <a href="Stone_tool" title="Stone tool">tools from stone</a> and perhaps animal bones, leading to its name homo habilis (Latin 'handy man') bestowed by discoverer <a href="Louis_Leakey" title="Louis Leakey">Louis Leakey</a>. Some scientists have proposed moving this species from Homo into Australopithecus due to the morphology of its skeleton being more adapted to <a href="Arboreal_locomotion" title="Arboreal locomotion">living in trees</a> rather than <a href="Bipedalism" title="Bipedalism">walking on two legs</a> like later hominins.<a href="#cite_note-65">[63]</a>

In May 2010, a new species, <a href="Homo_gautengensis" title="Homo gautengensis">Homo gautengensis</a>, was discovered in South Africa.<a href="#cite_note-toothy-66">[64]</a>

<div class="mw-heading mw-heading3"><h3 id="H._rudolfensis_and_H._georgicus">H. rudolfensis and H. georgicus</h3></div>

These are proposed species names for fossils from about 1.9–1.6 Ma, whose relation to Homo habilis is not yet clear.

<ul><li>Homo rudolfensis refers to a single, incomplete skull from Kenya. Scientists have suggested that this was a specimen of Homo habilis, but this has not been confirmed.<a href="#cite_note-67">[65]</a></li>

<li><a href="Homo_georgicus" class="mw-redirect" title="Homo georgicus">Homo georgicus</a>, from <a href="Georgia_(country)" title="Georgia (country)">Georgia</a>, may be an intermediate form between Homo habilis and Homo erectus,<a href="#cite_note-68">[66]</a> or a subspecies of Homo erectus.<a href="#cite_note-69">[67]</a></li></ul>

<div class="mw-heading mw-heading3"><h3 id="H._ergaster_and_H._erectus">H. ergaster and H. erectus</h3></div>

The first fossils of Homo erectus were discovered by Dutch physician <a href="Eugene_Dubois" class="mw-redirect" title="Eugene Dubois">Eugene Dubois</a> in 1891 on the Indonesian island of Java. He originally named the material <a href="Anthropopithecus" title="Anthropopithecus">Anthropopithecus</a> erectus (1892–1893, considered at this point as a chimpanzee-like fossil primate) and <a href="Pithecanthropus" class="mw-redirect" title="Pithecanthropus">Pithecanthropus</a> erectus (1893–1894, changing his mind as of based on its morphology, which he considered to be intermediate between that of humans and apes).<a href="#cite_note-70">[68]</a> Years later, in the 20th century, the German physician and <a href="Paleoanthropologist" class="mw-redirect" title="Paleoanthropologist">paleoanthropologist</a> <a href="Franz_Weidenreich" title="Franz Weidenreich">Franz Weidenreich</a> (1873–1948) compared in detail the characters of Dubois' <a href="Java_Man" title="Java Man">Java Man</a>, then named Pithecanthropus erectus, with the characters of the <a href="Peking_Man" title="Peking Man">Peking Man</a>, then named Sinanthropus pekinensis. Weidenreich concluded in 1940 that because of their anatomical similarity with modern humans it was necessary to gather all these specimens of Java and China in a single species of the genus Homo, the species H. erectus.<a href="#cite_note-71">[69]</a><a href="#cite_note-72">[70]</a>

Homo erectus lived from about 1.8 Ma to about 108,000 years ago.<a href="#cite_note-Rizal2020-73">[71]</a> This population appears to have died out when the savannah corridors closed, and tropical jungle took over.;<a href="#cite_note-74">[72]</a> however, nearby <a href="H._floresiensis" class="mw-redirect" title="H. floresiensis">H. floresiensis</a> survived it. The early phase of H. erectus, from 1.8 to 1.25 Ma, is considered by some to be a separate species, H. ergaster, or as H. erectus ergaster, a subspecies of H. erectus. Many paleoanthropologists now use the term Homo ergaster for the non-Asian forms of this group, and reserve H. erectus only for those fossils that are found in Asia and meet certain skeletal and dental requirements which differ slightly from H. ergaster.

In Africa in the Early Pleistocene, 1.5–1 Ma, some populations of Homo habilis are thought to have evolved larger brains and to have made more elaborate stone tools; these differences and others are sufficient for anthropologists to classify them as a new species, Homo erectus—in Africa.<a href="#cite_note-75">[73]</a> This species also may have used fire to cook meat. <a href="Catching_Fire%3A_How_Cooking_Made_Us_Human" title="Catching Fire: How Cooking Made Us Human">Richard Wrangham</a> notes that Homo seems to have been ground dwelling, with reduced intestinal length, smaller dentition, and "brains [swollen] to their current, horrendously fuel-inefficient size",<a href="#cite_note-76">[74]</a> and hypothesizes that control of fire and cooking, which released increased nutritional value, was the key adaptation that separated Homo from tree-sleeping Australopithecines.<a href="#cite_note-77">[75]</a>

<div role="note" class="hatnote navigation-not-searchable">See also: <a href="Control_of_fire_by_early_humans" title="Control of fire by early humans">Control of fire by early humans</a></div>

<div class="mw-heading mw-heading3"><h3 id="H._cepranensis_and_H._antecessor">H. cepranensis and H. antecessor</h3></div>

These are proposed as species intermediate between H. erectus and H. heidelbergensis.

<ul><li>H. antecessor is known from fossils from Spain and England that are dated 1.2 Ma–500 <a href="Year#SI_prefix_multipliers" title="Year">ka</a>.<a href="#cite_note-78">[76]</a><a href="#cite_note-79">[77]</a></li>

<li><a href="Homo_cepranensis" class="mw-redirect" title="Homo cepranensis">H. cepranensis</a> refers to a single skull cap from Italy, estimated to be about 800,000 years old.<a href="#cite_note-80">[78]</a></li></ul>

<div class="mw-heading mw-heading3"><h3 id="H._heidelbergensis">H. heidelbergensis</h3></div>

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

H. heidelbergensis ("Heidelberg Man") lived from about 800,000 to about 300,000 years ago. It is also proposed as Homo sapiens heidelbergensis or Homo sapiens paleohungaricus.<a href="#cite_note-81">[79]</a>

<div class="mw-heading mw-heading3"><h3 id="H._rhodesiensis,_and_the_Gawis_cranium">H. rhodesiensis, and the Gawis cranium</h3></div>

<ul><li>H. rhodesiensis, estimated to be 300,000–125,000 years old. Most current researchers place Rhodesian Man within the group of Homo heidelbergensis, though other designations such as archaic Homo sapiens and Homo sapiens rhodesiensis have been proposed.</li>

<li>In February 2006 a fossil, the <a href="Gawis_cranium" title="Gawis cranium">Gawis cranium</a>, was found which might possibly be a species intermediate between H. erectus and H. sapiens or one of many evolutionary dead ends. The skull from Gawis, Ethiopia, is believed to be 500,000–250,000 years old. Only summary details are known, and the finders have not yet released a peer-reviewed study. Gawis man's facial features suggest that it is either an intermediate species or an example of a "Bodo man" female.<a href="#cite_note-82">[80]</a></li></ul>

<div class="mw-heading mw-heading3"><h3 id="Neanderthal_and_Denisovan">Neanderthal and Denisovan</h3></div>

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

Homo neanderthalensis, alternatively designated as Homo sapiens neanderthalensis,<a href="#cite_note-83">[81]</a> lived in Europe and Asia from 400,000<a href="#cite_note-84">[82]</a> to about 28,000 years ago.<a href="#cite_note-85">[83]</a>

There are a number of clear anatomical differences between <a href="Anatomically_modern_humans" class="mw-redirect" title="Anatomically modern humans">anatomically modern humans</a> (AMH) and Neanderthal specimens, many relating to the superior Neanderthal adaptation to cold environments. Neanderthal <a href="Surface_to_volume_ratio" class="mw-redirect" title="Surface to volume ratio">surface to volume ratio</a> was even lower than that among modern <a href="Inuit" title="Inuit">Inuit</a> populations, indicating superior retention of body heat.

Neanderthals also had significantly larger brains, as shown from brain endocasts, casting doubt on their intellectual inferiority to modern humans. However, the higher body mass of Neanderthals may have required larger brain mass for body control.<a href="#cite_note-dunbar-86">[84]</a> Also, recent research by Pearce, <a href="Chris_Stringer" title="Chris Stringer">Stringer</a>, and Dunbar has shown important differences in brain architecture. The larger size of the Neanderthal orbital chamber and <a href="Occipital_lobe" title="Occipital lobe">occipital lobe</a> suggests that they had a better visual acuity than modern humans, useful in the dimmer light of glacial Europe.

Neanderthals may have had less <a href="Dunbar's_number" title="Dunbar's number">brain capacity available for social functions</a>. Inferring social group size from endocranial volume (minus occipital lobe size) suggests that Neanderthal groups may have been limited to 120 individuals, compared to 144<a href="#cite_note-87">[85]</a> possible relationships for modern humans. Larger social groups could imply that modern humans had less risk of inbreeding within their clan, trade over larger areas (confirmed in the distribution of stone tools), and faster spread of social and technological innovations. All these may have all contributed to modern Homo sapiens replacing Neanderthal populations by 28,000 BP.<a href="#cite_note-dunbar-86">[84]</a>

Earlier evidence from sequencing mitochondrial DNA suggested that no significant gene flow occurred between H. neanderthalensis and H. sapiens, and that the two were separate species that shared a common ancestor about 660,000 years ago.<a href="#cite_note-88">[86]</a><a href="#cite_note-89">[87]</a><a href="#cite_note-90">[88]</a> However, a sequencing of the Neanderthal genome in 2010 indicated that Neanderthals did indeed interbreed with anatomically modern humans c. 45,000-80,000 years ago, around the time modern humans migrated out from Africa, but before they dispersed throughout Europe, Asia and elsewhere.<a href="#cite_note-neandersequence-91">[89]</a> The genetic sequencing of a 40,000-year-old <a href="Pe%C8%99tera_cu_Oase" title="Peștera cu Oase">human skeleton from Romania</a> showed that 11% of its genome was Neanderthal, implying the individual had a Neanderthal ancestor 4–6 generations previously,<a href="#cite_note-92">[90]</a> in addition to a contribution from earlier interbreeding in the Middle East. Though this interbred Romanian population seems not to have been ancestral to modern humans, the finding indicates that interbreeding happened repeatedly.<a href="#cite_note-93">[91]</a>

All modern non-African humans have about 1% to 4% (or 1.5% to 2.6% by more recent data) of their DNA derived from Neanderthals.<a href="#cite_note-greenetal-94">[92]</a><a href="#cite_note-neandersequence-91">[89]</a><a href="#cite_note-A_high-coverage_Neandertal_genome_f-95">[93]</a> This finding is consistent with recent studies indicating that the divergence of some human alleles dates to one Ma, although this interpretation has been questioned.<a href="#cite_note-96">[94]</a><a href="#cite_note-97">[95]</a> Neanderthals and AMH Homo sapiens could have co-existed in Europe for as long as 10,000 years, during which AMH populations exploded, vastly outnumbering Neanderthals, possibly outcompeting them by sheer numbers.<a href="#cite_note-98">[96]</a>

In 2008, archaeologists working at the site of <a href="Denisova_Cave" title="Denisova Cave">Denisova Cave</a> in the <a href="Altai_Mountains" title="Altai Mountains">Altai Mountains</a> of <a href="Siberia" title="Siberia">Siberia</a> uncovered a small bone fragment from the fifth finger of a juvenile member of another human species, the Denisovans.<a href="#cite_note-99">[97]</a> Artifacts, including a bracelet, excavated in the cave at the same level were <a href="Radiocarbon_dating" title="Radiocarbon dating">carbon dated</a> to around 40,000 BP. As DNA had survived in the fossil fragment due to the cool climate of the Denisova Cave, both mtDNA and nuclear DNA were sequenced.<a href="#cite_note-pmid21179161-46">[44]</a><a href="#cite_note-Pääbo_et_al.-100">[98]</a>

While the divergence point of the mtDNA was unexpectedly deep in time,<a href="#cite_note-The_Scientist-101">[99]</a> the full genomic sequence suggested the Denisovans belonged to the same lineage as Neanderthals, with the two diverging shortly after their line split from the lineage that gave rise to modern humans.<a href="#cite_note-pmid21179161-46">[44]</a> Modern humans are known to have overlapped with Neanderthals in Europe and the Near East for possibly more than 40,000 years,<a href="#cite_note-102">[100]</a> and the discovery raises the possibility that Neanderthals, Denisovans, and modern humans may have co-existed and interbred. The existence of this distant branch creates a much more complex picture of humankind during the <a href="Late_Pleistocene" title="Late Pleistocene">Late Pleistocene</a> than previously thought.<a href="#cite_note-Pääbo_et_al.-100">[98]</a><a href="#cite_note-103">[101]</a> Evidence has also been found that as much as 6% of the DNA of some modern <a href="Melanesians" title="Melanesians">Melanesians</a> derive from Denisovans, indicating limited interbreeding in Southeast Asia.<a href="#cite_note-Reich_2011-104">[102]</a><a href="#cite_note-105">[103]</a>

Alleles thought to have originated in Neanderthals and Denisovans have been identified at several genetic loci in the genomes of modern humans outside Africa. <a href="Human_leukocyte_antigen" title="Human leukocyte antigen">Human leukocyte antigen</a> (HLA) haplotypes from Denisovans and Neanderthal represent more than half the HLA alleles of modern Eurasians,<a href="#cite_note-10.1126/science.1209202-48">[46]</a> indicating strong positive selection for these <a href="Introgressed" class="mw-redirect" title="Introgressed">introgressed</a> alleles. Corinne Simoneti at Vanderbilt University, in Nashville and her team have found from medical records of 28,000 people of European descent that the presence of Neanderthal DNA segments may be associated with a higher rate of depression.<a href="#cite_note-106">[104]</a>

The flow of genes from Neanderthal populations to modern humans was not all one way. Sergi Castellano of the Max Planck Institute for <a href="Evolutionary_Anthropology" class="mw-redirect" title="Evolutionary Anthropology">Evolutionary Anthropology</a> reported in 2016 that while Denisovan and Neanderthal genomes are more related to each other than they are to us, Siberian Neanderthal genomes show more similarity to modern human genes than do European Neanderthal populations. This suggests Neanderthal populations interbred with modern humans around 100,000 years ago, probably somewhere in the Near East.<a href="#cite_note-107">[105]</a>

Studies of a Neanderthal child at Gibraltar show from brain development and tooth eruption that Neanderthal children may have matured more rapidly than Homo sapiens.<a href="#cite_note-108">[106]</a>

<div class="mw-heading mw-heading3"><h3 id="H._floresiensis">H. floresiensis</h3></div>

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

H. floresiensis, which lived from approximately 190,000 to 50,000 years <a href="Before_present" class="mw-redirect" title="Before present">before present</a> (BP), has been nicknamed the <a href="Hobbit" title="Hobbit">hobbit</a> for its small size, possibly a result of <a href="Insular_dwarfism" title="Insular dwarfism">insular dwarfism</a>.<a href="#cite_note-:0-109">[107]</a> H. floresiensis is intriguing both for its size and its age, being an example of a recent species of the genus Homo that exhibits derived traits not shared with modern humans. In other words, H. floresiensis shares a common ancestor with modern humans, but split from the modern human lineage and followed a distinct evolutionary path. The main find was a skeleton believed to be a woman of about 30 years of age. Found in 2003, it has been dated to approximately 18,000 years old. The living woman was estimated to be one meter in height, with a brain volume of just 380 cm3 (considered small for a chimpanzee and less than a third of the H. sapiens average of 1400 cm3).<a href="#cite_note-:0-109">[107]</a>

However, there is an ongoing debate over whether H. floresiensis is indeed a separate species.<a href="#cite_note-:1-110">[108]</a> Some scientists hold that H. floresiensis was a modern H. sapiens with pathological dwarfism.<a href="#cite_note-Martin-111">[109]</a> This hypothesis is supported in part, because some modern humans who live on <a href="Flores" title="Flores">Flores</a>, the Indonesian island where the skeleton was found, are <a href="Pygmy_peoples" title="Pygmy peoples">pygmies</a>. This, coupled with pathological dwarfism, could have resulted in a significantly diminutive human. The other major attack on H. floresiensis as a separate species is that it was found with tools only associated with H. sapiens.<a href="#cite_note-Martin-111">[109]</a>

The hypothesis of pathological dwarfism, however, fails to explain additional <a href="Homo_floresiensis#Anatomy" title="Homo floresiensis">anatomical features</a> that are unlike those of modern humans (diseased or not) but much like those of ancient members of our genus. Aside from cranial features, these features include the form of bones in the wrist, forearm, shoulder, knees, and feet. Additionally, this hypothesis fails to explain the find of multiple examples of individuals with these same characteristics, indicating they were common to a large population, and not limited to one individual.<a href="#cite_note-:1-110">[108]</a>

In 2016, fossil teeth and a partial jaw from hominins assumed to be ancestral to H. floresiensis were discovered<a href="#cite_note-Callaway2016b-112">[110]</a> at <a href="Mata_Menge" title="Mata Menge">Mata Menge</a>, about 74 km (46 mi) from Liang Bua. They date to about 700,000 years ago<a href="#cite_note-Brumm2016-113">[111]</a> and are noted by Australian archaeologist Gerrit van den Bergh for being even smaller than the later fossils.<a href="#cite_note-114">[112]</a>

<div class="mw-heading mw-heading3"><h3 id="H._luzonensis">H. luzonensis</h3></div>

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

A small number of specimens from the island of <a href="Luzon" title="Luzon">Luzon</a>, dated 50,000 to 67,000 years ago, have recently been assigned by their discoverers, based on dental characteristics, to a novel human species, H. luzonensis.<a href="#cite_note-115">[113]</a>

<div class="mw-heading mw-heading3"><h3 id="H._sapiens">H. sapiens</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main articles: <a href="Archaic_humans" class="mw-redirect" title="Archaic humans">Archaic humans</a>, <a href="Early_modern_human" title="Early modern human">Early modern human</a>, <a href="Interbreeding_between_archaic_and_modern_humans" title="Interbreeding between archaic and modern humans">Interbreeding between archaic and modern humans</a>, and <a href="Human#Evolution" title="Human">Human § Evolution</a></div>

H. sapiens (the adjective <a href="https://en.wiktionary.org/wiki/sapiens" class="extiw external" title="wikt:sapiens">sapiens</a> is Latin for "wise" or "intelligent") emerged in Africa around 300,000 years ago, likely derived from <a href="H._heidelbergensis" class="mw-redirect" title="H. heidelbergensis">H. heidelbergensis</a> or a related lineage.<a href="#cite_note-Schlebusch2017-116">[114]</a><a href="#cite_note-Guardian-117">[115]</a> In September 2019, scientists reported the computerized determination, based on 260 <a href="CT_scan" title="CT scan">CT scans</a>, of a virtual <a href="Human_skull" class="mw-redirect" title="Human skull">skull shape</a> of the last common human ancestor to <a href="Modern_human" class="mw-redirect" title="Modern human">modern humans</a> (H. sapiens), representative of the earliest modern humans, and suggested that modern humans arose between 260,000 and 350,000 years ago through a merging of populations in <a href="East_Africa" title="East Africa">East</a> and South Africa.<a href="#cite_note-NYT-20190910-118">[116]</a><a href="#cite_note-NAT-20190910-119">[117]</a>

Between 400,000 years ago and the second interglacial period in the <a href="Middle_Pleistocene" title="Middle Pleistocene">Middle Pleistocene</a>, around 250,000 years ago, the trend in <a href="Brain_size#Cranial_capacity" title="Brain size">intra-cranial volume expansion</a> and the elaboration of stone tool technologies developed, providing evidence for a transition from H. erectus to H. sapiens. The direct evidence suggests there was a migration of H. erectus <a href="Recent_African_origin_of_modern_humans" title="Recent African origin of modern humans">out of Africa</a>, then a further speciation of H. sapiens from H. erectus in Africa. A subsequent migration (both within and out of Africa) eventually replaced the earlier dispersed H. erectus. This migration and origin theory is usually referred to as the "recent single-origin hypothesis" or "out of Africa" theory. H. sapiens <a href="Archaic_human_admixture_with_modern_humans" class="mw-redirect" title="Archaic human admixture with modern humans">interbred with archaic humans</a> both in Africa and in Eurasia, in Eurasia notably with Neanderthals and Denisovans.<a href="#cite_note-pmid21179161-46">[44]</a><a href="#cite_note-Reich_2011-104">[102]</a>

The <a href="Toba_catastrophe_theory" class="mw-redirect" title="Toba catastrophe theory">Toba catastrophe theory</a>, which postulates a <a href="Population_bottleneck" title="Population bottleneck">population bottleneck</a> for H. sapiens about 70,000 years ago,<a href="#cite_note-ambrose1998-120">[118]</a> was controversial from its first proposal in the 1990s and by the 2010s had very little support.<a href="#cite_note-121">[119]</a> Distinctive <a href="Human_genetic_variability" class="mw-redirect" title="Human genetic variability">human genetic variability</a> has arisen as the result of the <a href="Founder_effect" title="Founder effect">founder effect</a>, by <a href="Archaic_human_admixture_with_modern_humans" class="mw-redirect" title="Archaic human admixture with modern humans">archaic admixture</a> and by <a href="Recent_human_evolution" title="Recent human evolution">recent evolutionary pressures</a>.

<div class="mw-heading mw-heading2"><h2 id="Anatomical_changes">Anatomical changes</h2></div>

Since <a href="Homo_sapiens" class="mw-redirect" title="Homo sapiens">Homo sapiens</a> separated from its <a href="Chimpanzee%E2%80%93human_last_common_ancestor" title="Chimpanzee–human last common ancestor">last common ancestor</a> shared with <a href="Chimpanzee" title="Chimpanzee">chimpanzees</a>, human evolution is characterized by a number of <a href="Morphology_(biology)" title="Morphology (biology)">morphological</a>, <a href="Human_development_(biology)" class="mw-redirect" title="Human development (biology)">developmental</a>, <a href="Human_physiology" class="mw-redirect" title="Human physiology">physiological</a>, <a href="Human_behavior" title="Human behavior">behavioral</a>, and environmental changes.<a href="#cite_note-Marlowe_54–67-9">[9]</a> Environmental (cultural) evolution discovered much later during the <a href="Pleistocene" title="Pleistocene">Pleistocene</a> played a significant role in human evolution observed via human transitions between subsistence systems.<a href="#cite_note-122">[120]</a><a href="#cite_note-Marlowe_54–67-9">[9]</a> The most significant of these adaptations are bipedalism, increased brain size, lengthened <a href="Ontogeny" title="Ontogeny">ontogeny</a> (gestation and infancy), and decreased <a href="Sexual_dimorphism" title="Sexual dimorphism">sexual dimorphism</a>. The relationship between these changes is the subject of ongoing debate.&#91;[[Wikipedia:Citing_sources|<span_title="This_citation_requires_a_reference_to_the_specific_page_or_range_of_pages_in_which_the_material_appears.&#32;(February_2015)">page&nbsp;needed]]&#93;_123-0" class="reference"><a href="#cite_note-FOOTNOTEBoydSilk2003[[Category:Wikipedia_articles_needing_page_number_citations_from_February_2015]]<sup_class="noprint_Inline-Template_"_style="white-space:nowrap;">&#91;[[Wikipedia:Citing_sources|<span_title="This_citation_requires_a_reference_to_the_specific_page_or_range_of_pages_in_which_the_material_appears.&#32;(February_2015)">page&nbsp;needed]]&#93;-123">[121]</a> Other significant morphological changes included the evolution of a <a href="Thumb#Grips" title="Thumb">power and precision grip</a>, a change first occurring in <a href="H._erectus" class="mw-redirect" title="H. erectus">H. erectus</a>.<a href="#cite_note-FOOTNOTEBruesSnow1965[httpsbooksgooglecombooksid9WemAAAAIAAJpgPA1_1–39]-124">[122]</a>

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

<a href="Bipedalism" title="Bipedalism">Bipedalism</a> (walking on two legs) is the basic adaptation of the hominid and is considered the main cause behind a suite of skeletal changes shared by all bipedal hominids. The earliest hominin, of presumably primitive bipedalism, is considered to be either <a href="Sahelanthropus" title="Sahelanthropus">Sahelanthropus</a><a href="#cite_note-Brunet2002-125">[123]</a> or <a href="Orrorin" title="Orrorin">Orrorin</a>, both of which arose some 6 to 7 million years ago. The non-bipedal knuckle-walkers, the <a href="Gorilla" title="Gorilla">gorillas</a> and chimpanzees, diverged from the hominin line over a period covering the same time, so either Sahelanthropus or Orrorin may be our last shared ancestor. <a href="Ardipithecus" title="Ardipithecus">Ardipithecus</a>, a full biped, arose approximately 5.6 million years ago.<a href="#cite_note-126">[124]</a>

The early bipeds eventually evolved into the australopithecines and still later into the genus <a href="Homo" title="Homo">Homo</a>. There are several theories of the adaptation value of bipedalism. It is possible that bipedalism was favored because it freed the hands for reaching and carrying food, saved energy during locomotion,<a href="#cite_note-Kwang_Hyun_2015_929–934-127">[125]</a> enabled long-distance running and hunting, provided an enhanced field of vision, and helped avoid hyperthermia by reducing the surface area exposed to direct sun; features all advantageous for thriving in the new savanna and woodland environment created as a result of the East African Rift Valley uplift versus the previous closed forest habitat.<a href="#cite_note-Kwang_Hyun_2015_929–934-127">[125]</a><a href="#cite_note-FOOTNOTEDeSalleTattersall2008[httpsbooksgooglecombooksidBf4Sitw7YaICpgPA146_146]-128">[126]</a><a href="#cite_note-FOOTNOTECurry2008106–109-129">[127]</a> A 2007 study provides support for the hypothesis that bipedalism evolved because it used less energy than quadrupedal knuckle-walking.<a href="#cite_note-130">[128]</a><a href="#cite_note-131">[129]</a> However, recent studies suggest that bipedality without the <a href="Control_of_fire_by_early_humans" title="Control of fire by early humans">ability to use fire</a> would not have allowed global dispersal.<a href="#cite_note-132">[130]</a> This change in gait saw a lengthening of the legs proportionately when compared to the length of the arms, which were shortened through the removal of the need for <a href="Brachiation" title="Brachiation">brachiation</a>. Another change is the shape of the big toe. Recent studies suggest that australopithecines still lived part of the time in trees as a result of maintaining a grasping big toe. This was progressively lost in habilines.

Anatomically, the evolution of bipedalism has been accompanied by a large number of skeletal changes, not just to the legs and pelvis, but also to the <a href="Human_vertebral_column" class="mw-redirect" title="Human vertebral column">vertebral column</a>, feet and ankles, and skull.<a href="#cite_note-FOOTNOTEAielloDean1990-133">[131]</a> The <a href="Femur" title="Femur">femur</a> evolved into a slightly more angular position to move the center of gravity toward the geometric center of the body. The knee and ankle joints became increasingly robust to better support increased weight. To support the increased weight on each vertebra in the upright position, the human vertebral column became S-shaped and the <a href="Lumbar_vertebrae" title="Lumbar vertebrae">lumbar vertebrae</a> became shorter and wider. In the feet the big toe moved into alignment with the other toes to help in forward locomotion. The arms and forearms shortened relative to the legs making it easier to run. The <a href="Foramen_magnum" title="Foramen magnum">foramen magnum</a> migrated under the skull and more anterior.<a href="#cite_note-FOOTNOTEKondo1985-134">[132]</a>

The most significant changes occurred in the pelvic region, where the long downward facing <a href="Ilium_(bone)" title="Ilium (bone)">iliac blade</a> was shortened and widened as a requirement for keeping the center of gravity stable while walking;<a href="#cite_note-FOOTNOTESrivastava2009[httpsbooksgooglecombooksidkCerOsM8XMwCpgPA87_87]-30">[28]</a> bipedal hominids have a shorter but broader, bowl-like pelvis due to this. A drawback is that the birth canal of bipedal apes is smaller than in knuckle-walking apes, though there has been a widening of it in comparison to that of australopithecine and modern humans, thus permitting the passage of newborns due to the increase in cranial size. This is limited to the upper portion, since further increase can hinder normal bipedal movement.<a href="#cite_note-FOOTNOTEStrickberger2000[httpsbooksgooglecombooksidfeMItLo5gwgCpgPA476_475–476]-135">[133]</a>

The shortening of the pelvis and smaller birth canal evolved as a requirement for bipedalism and had significant effects on the process of human birth, which is much more difficult in modern humans than in other primates. During human birth, because of the variation in size of the pelvic region, the fetal head must be in a transverse position (compared to the mother) during entry into the birth canal and rotate about 90 degrees upon exit.<a href="#cite_note-FOOTNOTETrevathan2011[httpsbooksgooglecombooksidPotlqpl-jxgCpgPA20_20]-136">[134]</a> The smaller birth canal became a limiting factor to brain size increases in early humans and prompted a shorter gestation period leading to the relative immaturity of human offspring, who are unable to walk much before 12 months and have greater <a href="Neoteny" title="Neoteny">neoteny</a>, compared to other primates, who are mobile at a much earlier age.<a href="#cite_note-FOOTNOTECurry2008106–109-129">[127]</a> The increased brain growth after birth and the increased dependency of children on mothers had a major effect upon the female reproductive cycle,<a href="#cite_note-137">[135]</a> and the more frequent appearance of <a href="Alloparenting" title="Alloparenting">alloparenting</a> in humans when compared with other hominids.<a href="#cite_note-138">[136]</a> Delayed human sexual maturity also led to the evolution of <a href="Menopause" title="Menopause">menopause</a> with one explanation, the <a href="Grandmother_hypothesis" title="Grandmother hypothesis">grandmother hypothesis</a>, providing that elderly women could better pass on their genes by taking care of their daughter's offspring, as compared to having more children of their own.<a href="#cite_note-139">[137]</a><a href="#cite_note-140">[138]</a>

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

The human species eventually developed a much larger brain than that of other primates—typically 1,330 cm3 (81 cu in) in modern humans, nearly three times the size of a chimpanzee or gorilla brain.<a href="#cite_note-Schoeneman-144">[141]</a> After a period of stasis with Australopithecus anamensis and Ardipithecus, species which had smaller brains as a result of their bipedal locomotion,<a href="#cite_note-145">[142]</a> the pattern of <a href="Encephalization" class="mw-redirect" title="Encephalization">encephalization</a> started with Homo habilis, whose 600 cm3 (37 cu in) brain was slightly larger than that of chimpanzees. This evolution continued in Homo erectus with 800–1,100 cm3 (49–67 cu in), and reached a maximum in Neanderthals with 1,200–1,900 cm3 (73–116 cu in), larger even than modern Homo sapiens. This brain increase manifested during postnatal <a href="Neural_development" class="mw-redirect" title="Neural development">brain growth</a>, far exceeding that of other apes (<a href="Heterochrony" title="Heterochrony">heterochrony</a>). It also allowed for extended periods of <a href="Observational_learning" title="Observational learning">social learning</a> and <a href="Language_acquisition" title="Language acquisition">language acquisition</a> in juvenile humans, beginning as much as 2 million years ago. Encephalization may be due to a dependency on calorie-dense, difficult-to-acquire food.<a href="#cite_note-146">[143]</a>

Furthermore, the changes in the structure of <a href="Human_brain" title="Human brain">human brains</a> may be even more significant than the increase in size.<a href="#cite_note-Park2007-147">[144]</a><a href="#cite_note-Bruner2007-148">[145]</a><a href="#cite_note-149">[146]</a><a href="#cite_note-Leonard_2007-55">[53]</a> Fossilized skulls shows the brain size in early humans fell within the range of modern humans 300,000 years ago, but only got its present-day brain shape between 100,000 and 35,000 years ago.<a href="#cite_note-150">[147]</a> The <a href="Temporal_lobe" title="Temporal lobe">temporal lobes</a>, which contain centers for language processing, have increased disproportionately, as has the <a href="Prefrontal_cortex" title="Prefrontal cortex">prefrontal cortex</a>, which has been related to complex decision-making and moderating social behavior.<a href="#cite_note-Schoeneman-144">[141]</a> Encephalization has been tied to increased starches<a href="#cite_note-NYT-20150813-54">[52]</a> and meat<a href="#cite_note-151">[148]</a><a href="#cite_note-152">[149]</a> in the diet, however a 2022 meta study called into question the role of meat.<a href="#cite_note-153">[150]</a> Other factors are the development of cooking,<a href="#cite_note-Organ_2011-154">[151]</a> and it has been proposed that intelligence increased as a response to an increased necessity for <a href="Dunbar's_number" title="Dunbar's number">solving social problems</a> as human society became more complex.<a href="#cite_note-David-Barrett-155">[152]</a> Changes in skull morphology, such as smaller mandibles and mandible muscle attachments, allowed more room for the brain to grow.<a href="#cite_note-FOOTNOTEBownRose1987-156">[153]</a>

The increase in volume of the <a href="Neocortex" title="Neocortex">neocortex</a> also included a rapid increase in size of the <a href="Cerebellum" title="Cerebellum">cerebellum</a>. Its function has traditionally been associated with balance and fine motor control, but more recently with <a href="Speech" title="Speech">speech</a> and <a href="Cognition" title="Cognition">cognition</a>. The great apes, including hominids, had a more pronounced cerebellum relative to the neocortex than other primates. It has been suggested that because of its function of sensory-motor control and learning complex muscular actions, the cerebellum may have underpinned human technological adaptations, including the preconditions of speech.<a href="#cite_note-157">[154]</a><a href="#cite_note-158">[155]</a><a href="#cite_note-159">[156]</a><a href="#cite_note-160">[157]</a>

The immediate survival advantage of encephalization is difficult to discern, as the major brain changes from Homo erectus to Homo heidelbergensis were not accompanied by major changes in technology. It has been suggested that the changes were mainly social and behavioural, including increased empathic abilities,<a href="#cite_note-161">[158]</a><a href="#cite_note-162">[159]</a> increases in size of social groups,<a href="#cite_note-David-Barrett-155">[152]</a><a href="#cite_note-163">[160]</a><a href="#cite_note-164">[161]</a> and increased behavioral plasticity.<a href="#cite_note-165">[162]</a> Humans are unique in the ability to acquire information through social transmission and adapt that information.<a href="#cite_note-166">[163]</a> The emerging field of <a href="Cultural_evolution" title="Cultural evolution">cultural evolution</a> studies human sociocultural change from an evolutionary perspective.<a href="#cite_note-167">[164]</a>

<div class="mw-heading mw-heading3"><h3 id="Sexual_dimorphism">Sexual dimorphism</h3></div>

The reduced degree of <a href="Sexual_dimorphism_in_humans" class="mw-redirect" title="Sexual dimorphism in humans">sexual dimorphism in humans</a> is visible primarily in the reduction of the male <a href="Canine_tooth" title="Canine tooth">canine tooth</a> relative to other ape species (except gibbons) and reduced brow ridges and general robustness of males. Another important physiological change related to <a href="Sexuality" class="mw-redirect" title="Sexuality">sexuality</a> in humans was the evolution of <a href="Concealed_ovulation" title="Concealed ovulation">hidden estrus</a>. Humans are the only hominoids in which the female is fertile year round and in which no special signals of fertility are produced by the body (such as genital swelling or overt changes in proceptivity during estrus).<a href="#cite_note-181">[177]</a>

Nonetheless, humans retain a degree of sexual dimorphism in the distribution of body hair and subcutaneous fat, and in the overall size, males being around 15% larger than females.<a href="#cite_note-182">[178]</a> These changes taken together have been interpreted as a result of an increased emphasis on <a href="Pair_bond" title="Pair bond">pair bonding</a> as a possible solution to the requirement for increased parental investment due to the prolonged infancy of offspring.<a href="#cite_note-183">[179]</a>

<div class="mw-heading mw-heading3"><h3 id="Ulnar_opposition">Ulnar opposition</h3></div>

The ulnar opposition—the contact between the thumb and the tip of the <a href="Little_finger" title="Little finger">little finger</a> of the same hand—is unique to the <a href="Homo" title="Homo">genus Homo</a>,<a href="#cite_note-184">[180]</a> including Neanderthals, the <a href="Atapuerca_Mountains" title="Atapuerca Mountains">Sima de los Huesos</a> <a href="Hominini" title="Hominini">hominins</a> and <a href="Homo_sapiens" class="mw-redirect" title="Homo sapiens">anatomically modern humans</a>.<a href="#cite_note-Miriam-185">[181]</a><a href="#cite_note-186">[182]</a> In other primates, the thumb is short and unable to touch the little finger.<a href="#cite_note-Miriam-185">[181]</a> The ulnar opposition facilitates the precision grip and power grip of the human hand, underlying all the skilled manipulations.

<div class="mw-heading mw-heading3"><h3 id="Other_changes">Other changes</h3></div>

A number of other changes have also characterized the evolution of humans, among them an increased reliance on vision rather than smell (highly reduced <a href="Olfactory_bulb" title="Olfactory bulb">olfactory bulb</a>); a longer juvenile developmental period and higher infant dependency;<a href="#cite_note-Kuzawa2014-187">[183]</a> a smaller gut and small, misaligned teeth; faster basal metabolism;<a href="#cite_note-Pontzer2016-188">[184]</a> loss of body hair;<a href="#cite_note-MarkusJRantala2007-189">[185]</a> an increase in

<a href="Eccrine_sweat_gland" title="Eccrine sweat gland">eccrine sweat gland</a> density that is ten times higher than any other <a href="Catarrhini" title="Catarrhini">catarrhinian</a> primates,<a href="#cite_note-190">[186]</a> yet humans use 30% to 50% less water per day compared to chimps and gorillas;<a href="#cite_note-191">[187]</a> more <a href="Rapid_eye_movement_sleep" title="Rapid eye movement sleep">REM sleep</a> but less sleep in total;<a href="#cite_note-192">[188]</a> a change in the shape of the dental arcade from u-shaped to parabolic; development of a <a href="Chin" title="Chin">chin</a> (found in Homo sapiens alone); <a href="Temporal_styloid_process" title="Temporal styloid process">styloid processes</a>; and a descended <a href="Larynx" title="Larynx">larynx</a>. As the human hand and arms adapted to the making of tools and were used less for climbing, the shoulder blades changed too. As a side effect, it allowed human ancestors to throw objects with greater force, speed and accuracy.<a href="#cite_note-193">[189]</a>

<div class="mw-heading mw-heading2"><h2 id="Use_of_tools">Use of tools</h2></div>

<div role="note" class="hatnote navigation-not-searchable">See also: <a href="Hunting_hypothesis" title="Hunting hypothesis">Hunting hypothesis</a></div>

The use of tools has been interpreted as a sign of intelligence, and it has been theorized that tool use may have stimulated certain aspects of human evolution, especially the continued expansion of the human brain.<a href="#cite_note-auto-195">[191]</a> Paleontology has yet to explain the expansion of this organ over millions of years despite being extremely demanding in terms of energy consumption. The brain of a modern human consumes, on average, about 13 watts (260 kilocalories per day), a fifth of the body's resting power consumption.<a href="#cite_note-196">[192]</a> Increased tool use would allow hunting for energy-rich meat products, and would enable processing more energy-rich plant products. Researchers have suggested that early hominins were thus under evolutionary pressure to increase their capacity to create and use tools.<a href="#cite_note-197">[193]</a>

Precisely when early humans started to use tools is difficult to determine, because the more primitive these tools are (for example, sharp-edged stones) the more difficult it is to decide whether they are natural objects or human artifacts.<a href="#cite_note-auto-195">[191]</a> There is some evidence that the australopithecines (4 Ma) <a href="Osteodontokeratic_culture" title="Osteodontokeratic culture">may have used broken bones as tools</a>, but this is debated.<a href="#cite_note-FOOTNOTERobinson2008[httpsbooksgooglecombooksid-kOvkb-xbU8CpgPA398_398]-198">[194]</a>

<a href="Tool_use_by_animals" class="mw-redirect" title="Tool use by animals">Many species make and use tools</a>, but it is the human genus that dominates the areas of making and using more complex tools. The oldest known tools are flakes from West Turkana, Kenya, which date to 3.3 million years ago.<a href="#cite_note-Harmand_310–315-199">[195]</a> The next oldest stone tools are from <a href="Gona%2C_Ethiopia" title="Gona, Ethiopia">Gona, Ethiopia</a>, and are considered the beginning of the Oldowan technology. These tools date to about 2.6 million years ago.<a href="#cite_note-200">[196]</a> A Homo fossil was found near some <a href="Oldowan" title="Oldowan">Oldowan</a> tools, and its age was noted at 2.3 million years old, suggesting that maybe the Homo species did indeed create and use these tools. It is a possibility but does not yet represent solid evidence.<a href="#cite_note-FOOTNOTEFreemanHerron2007786–788-201">[197]</a> The <a href="Third_metacarpal_styloid_process" title="Third metacarpal styloid process">third metacarpal styloid process</a> enables the hand bone to lock into the wrist bones, allowing for greater amounts of pressure to be applied to the wrist and hand from a grasping thumb and fingers. It allows humans the dexterity and strength to make and use complex tools. This unique anatomical feature separates humans from other apes and other nonhuman primates, and is not seen in human fossils older than 1.8 million years.<a href="#cite_note-202">[198]</a>

Bernard Wood noted that Paranthropus co-existed with the early Homo species in the area of the "Oldowan Industrial Complex" over roughly the same span of time. Although there is no direct evidence which identifies Paranthropus as the tool makers, their anatomy lends to indirect evidence of their capabilities in this area. Most paleoanthropologists agree that the early Homo species were indeed responsible for most of the Oldowan tools found. They argue that when most of the Oldowan tools were found in association with human fossils, Homo was always present, but Paranthropus was not.<a href="#cite_note-FOOTNOTEFreemanHerron2007786–788-201">[197]</a>

In 1994, Randall Susman used the anatomy of opposable thumbs as the basis for his argument that both the Homo and Paranthropus species were toolmakers. He compared bones and muscles of human and chimpanzee thumbs, finding that humans have 3 muscles which are lacking in chimpanzees. Humans also have thicker metacarpals with broader heads, allowing more precise grasping than the chimpanzee hand can perform. Susman posited that modern anatomy of the human opposable thumb is an evolutionary response to the requirements associated with making and handling tools and that both species were indeed toolmakers.<a href="#cite_note-FOOTNOTEFreemanHerron2007786–788-201">[197]</a>

<div class="mw-heading mw-heading2"><h2 id="Transition_to_behavioral_modernity">Transition to behavioral modernity</h2></div>

<div role="note" class="hatnote navigation-not-searchable">Further information: <a href="Behavioral_modernity" title="Behavioral modernity">Behavioral modernity</a></div>

Anthropologists describe <a href="Modern_human_behavior" class="mw-redirect" title="Modern human behavior">modern human behavior</a> to include cultural and behavioral traits such as specialization of tools, use of jewellery and images (such as cave drawings), organization of living space, rituals (such as grave gifts), specialized hunting techniques, exploration of less hospitable geographical areas, and <a href="Barter" title="Barter">barter</a> trade networks, as well as more general traits such as language and complex symbolic thinking. Debate continues as to whether a "revolution" led to modern humans ("big bang of human consciousness"), or whether the evolution was more gradual.<a href="#cite_note-Mcbrearty_Brooks-50">[48]</a>

Until about 50,000–40,000 years ago, the use of stone tools seems to have progressed stepwise. Each phase (H. habilis, H. ergaster, H. neanderthalensis) marked a new technology, followed by very slow development until the next phase. Currently paleoanthropologists are debating whether these Homo species possessed some or many modern human behaviors. They seem to have been culturally conservative, maintaining the same technologies and foraging patterns over very long periods.

Around 50,000 <a href="Before_Present" title="Before Present">BP</a>, human culture started to evolve more rapidly. The transition to behavioral modernity has been characterized by some as a "Great Leap Forward",<a href="#cite_note-FOOTNOTEDiamond199939-203">[199]</a> or as the "Upper Palaeolithic Revolution",<a href="#cite_note-204">[200]</a> due to the sudden appearance in the archaeological record of distinctive signs of modern behavior and <a href="Big_game_hunting" class="mw-redirect" title="Big game hunting">big game hunting</a>.<a href="#cite_note-oppenheimer-205">[201]</a> Evidence of behavioral modernity significantly earlier also exists from Africa, with older evidence of abstract imagery, widened subsistence strategies, more sophisticated tools and weapons, and other "modern" behaviors, and many scholars have recently argued that the transition to modernity occurred sooner than previously believed.<a href="#cite_note-Mcbrearty_Brooks-50">[48]</a><a href="#cite_note-Henshilwood_etal_2002-206">[202]</a><a href="#cite_note-Henshilwood_Marean_2003-207">[203]</a><a href="#cite_note-Backwell-208">[204]</a>

Other scholars consider the transition to have been more gradual, noting that some features had already appeared among archaic African Homo sapiens 300,000–200,000 years ago.<a href="#cite_note-209">[205]</a><a href="#cite_note-210">[206]</a><a href="#cite_note-211">[207]</a><a href="#cite_note-The_Atlantic-555674-212">[208]</a><a href="#cite_note-Brooks-213">[209]</a> Recent evidence suggests that the Australian Aboriginal population separated from the African population 75,000 years ago, and that they made a 160 km (99 mi) sea journey 60,000 years ago, which may diminish the significance of the Upper Paleolithic Revolution.<a href="#cite_note-214">[210]</a>

Modern humans started burying their dead, making clothing from animal hides, hunting with more sophisticated techniques (such as using <a href="Trapping_pit" title="Trapping pit">pit traps</a> or driving animals off cliffs), and <a href="Cave_painting" title="Cave painting">cave painting</a>.<a href="#cite_note-215">[211]</a> As human culture advanced, different populations innovated existing technologies: artifacts such as fish hooks, buttons, and bone needles show signs of cultural variation, which had not been seen prior to 50,000 BP. Typically, the older H. neanderthalensis populations did not vary in their technologies, although the <a href="Chatelperronian" class="mw-redirect" title="Chatelperronian">Chatelperronian</a> assemblages have been found to be Neanderthal imitations of H. sapiens <a href="Aurignacian" title="Aurignacian">Aurignacian</a> technologies.<a href="#cite_note-216">[212]</a>

<div class="mw-heading mw-heading2"><h2 id="Recent_and_ongoing_human_evolution">Recent and ongoing human evolution</h2></div>

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

<a href="Anatomically_modern_human" class="mw-redirect" title="Anatomically modern human">Anatomically modern human</a> populations continue to evolve, as they are affected by both natural selection and <a href="Genetic_drift" title="Genetic drift">genetic drift</a>. Although <a href="Selection_pressure" class="mw-redirect" title="Selection pressure">selection pressure</a> on some traits, such as resistance to smallpox, has decreased in the modern age, humans are still undergoing natural selection for many other traits. Some of these are due to specific environmental pressures, while others are related to lifestyle changes since the development of agriculture (10,000 years ago), urbanization (5,000), and industrialization (250 years ago). It has been argued that human evolution has accelerated since the development of agriculture 10,000 years ago and civilization some 5,000 years ago, resulting, it is claimed, in substantial genetic differences between different current human populations,<a href="#cite_note-FOOTNOTECochranHarpending2009-217">[213]</a> and more recent research indicates that for some traits, the developments and innovations of human culture have driven a new form of selection that coexists with, and in some cases has largely replaced, natural selection.<a href="#cite_note-218">[214]</a>

Particularly conspicuous is variation in superficial characteristics, such as <a href="Afro-textured_hair" class="mw-redirect" title="Afro-textured hair">Afro-textured hair</a>, or the recent evolution of <a href="Light_skin" title="Light skin">light skin</a> and blond hair in some populations, which are attributed to differences in climate. Particularly strong selective pressures have resulted in <a href="High-altitude_adaptation_in_humans" title="High-altitude adaptation in humans">high-altitude adaptation in humans</a>, with different ones in different isolated populations. Studies of the <a href="High-altitude_adaptation_in_humans#Genetic_basis" title="High-altitude adaptation in humans">genetic basis</a> show that some developed very recently, with Tibetans evolving over 3,000 years to have high proportions of an allele of <a href="EPAS1" title="EPAS1">EPAS1</a> that is adaptive to high altitudes.

Other evolution is related to <a href="Endemic_disease" class="mw-redirect" title="Endemic disease">endemic diseases</a>: the presence of <a href="Malaria" title="Malaria">malaria</a> selects for <a href="Sickle_cell_trait" title="Sickle cell trait">sickle cell trait</a> (the <a href="Heterozygous" class="mw-redirect" title="Heterozygous">heterozygous</a> form of sickle cell gene), while in the absence of malaria, the health effects of <a href="Sickle-cell_anemia" class="mw-redirect" title="Sickle-cell anemia">sickle-cell anemia</a> select against this trait. For another example, the population at risk of the severe debilitating disease <a href="Kuru_(disease)" title="Kuru (disease)">kuru</a> has significant over-representation of an immune variant of the <a href="PRNP" class="mw-redirect" title="PRNP">prion protein</a> gene G127V versus non-immune alleles. The frequency of this <a href="Mutation" title="Mutation">genetic variant</a> is due to the survival of immune persons.<a href="#cite_note-220">[216]</a><a href="#cite_note-221">[217]</a> Some reported trends remain unexplained and the subject of ongoing research in the novel field of evolutionary medicine: <a href="Polycystic_ovary_syndrome" title="Polycystic ovary syndrome">polycystic ovary syndrome</a> (PCOS) reduces fertility and thus is expected to be subject to extremely strong negative selection, but its relative commonality in human populations suggests a counteracting selection pressure. The identity of that pressure remains the subject of some debate.<a href="#cite_note-222">[218]</a>

Recent human evolution related to agriculture includes genetic resistance to infectious disease that has appeared in human populations by crossing the species barrier from domesticated animals,<a href="#cite_note-FOOTNOTEDiamond1999-223">[219]</a> as well as changes in metabolism due to changes in diet, such as <a href="Lactase_persistence" title="Lactase persistence">lactase persistence</a>.

Culturally-driven evolution can defy the expectations of natural selection: while human populations experience some pressure that drives a selection for producing children at younger ages, the advent of effective contraception, higher education, and changing social norms have driven the observed selection in the opposite direction.<a href="#cite_note-224">[220]</a> However, culturally-driven selection need not necessarily work counter or in opposition to natural selection: some proposals to explain the high rate of recent human brain expansion indicate a kind of feedback whereupon the brain's increased social learning efficiency encourages cultural developments that in turn encourage more efficiency, which drive more complex cultural developments that demand still-greater efficiency, and so forth.<a href="#cite_note-225">[221]</a> Culturally-driven evolution has an advantage in that in addition to the genetic effects, it can be observed also in the archaeological record: the development of stone tools across the Palaeolithic period connects to culturally-driven cognitive development in the form of skill acquisition supported by the culture and the development of increasingly complex technologies and the cognitive ability to elaborate them.<a href="#cite_note-226">[222]</a>

In contemporary times, since industrialization, some trends have been observed: for instance, menopause is evolving to occur later.<a href="#cite_note-doi10.1073/pnas.0906199106-227">[223]</a> Other reported trends appear to include lengthening of the human reproductive period and reduction in cholesterol levels, blood glucose and blood pressure in some populations.<a href="#cite_note-doi10.1073/pnas.0906199106-227">[223]</a>

<div class="mw-heading mw-heading2"><h2 id="History_of_study">History of study</h2></div>

<div role="note" class="hatnote navigation-not-searchable">For a chronological guide, see <a href="Timeline_of_human_evolution" title="Timeline of human evolution">Timeline of human evolution</a>.</div>

<div class="mw-heading mw-heading3"><h3 id="Before_Darwin">Before Darwin</h3></div>

The name Homo of the biological genus to which humans belong is <a href="Latin" title="Latin">Latin</a> for 'human'.<a href="#cite_note-228">[e]</a> It was chosen originally by <a href="Carl_Linnaeus" title="Carl Linnaeus">Carl Linnaeus</a> in his classification system.<a href="#cite_note-229">[f]</a> The English word human is from the Latin humanus, the adjectival form of homo. The Latin homo derives from the <a href="Proto-Indo-European_language" title="Proto-Indo-European language">Indo-European</a> root *dhghem, or 'earth'.<a href="#cite_note-230">[224]</a> Linnaeus and other scientists of his time also considered the great apes to be the closest relatives of humans based on <a href="Morphology_(biology)" title="Morphology (biology)">morphological</a> and <a href="Anatomy" title="Anatomy">anatomical</a> similarities.<a href="#cite_note-231">[225]</a>

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

The possibility of linking humans with earlier apes by descent became clear only after 1859 with the publication of <a href="Charles_Darwin" title="Charles Darwin">Charles Darwin</a>'s <a href="On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species</a>, in which he argued for the idea of the evolution of new species from earlier ones. Darwin's book did not address the question of human evolution, saying only that "Light will be thrown on the origin of man and his history."<a href="#cite_note-FOOTNOTEDarwin1859[httpdarwin-onlineorgukcontentframesetitemIDF373viewtypetextpageseq506_367–404]-232">[226]</a>

The first debates about the nature of human evolution arose between <a href="Thomas_Henry_Huxley" title="Thomas Henry Huxley">Thomas Henry Huxley</a> and <a href="Richard_Owen" title="Richard Owen">Richard Owen</a>. Huxley argued for human evolution from apes by illustrating many of the similarities and differences between humans and other apes, and did so particularly in his 1863 book <a href="Evidence_as_to_Man's_Place_in_Nature" class="mw-redirect" title="Evidence as to Man's Place in Nature">Evidence as to Man's Place in Nature</a>. Many of Darwin's early supporters (such as <a href="Alfred_Russel_Wallace" title="Alfred Russel Wallace">Alfred Russel Wallace</a> and <a href="Charles_Lyell" title="Charles Lyell">Charles Lyell</a>) did not initially agree that the origin of the mental capacities and the moral sensibilities of humans could be explained by <a href="Natural_selection" title="Natural selection">natural selection</a>, though this later changed. Darwin applied the theory of evolution and <a href="Sexual_selection" title="Sexual selection">sexual selection</a> to humans in his 1871 book <a href="The_Descent_of_Man%2C_and_Selection_in_Relation_to_Sex" title="The Descent of Man, and Selection in Relation to Sex">The Descent of Man, and Selection in Relation to Sex</a>.<a href="#cite_note-FOOTNOTEDarwin1981-233">[227]</a>

<div class="mw-heading mw-heading3"><h3 id="First_fossils">First fossils</h3></div>

A major problem in the 19th century was the lack of <a href="Transitional_fossil" title="Transitional fossil">fossil intermediaries</a>. <a href="Neanderthal" title="Neanderthal">Neanderthal</a> remains were discovered in a limestone quarry in 1856, three years before the publication of On the Origin of Species, and Neanderthal fossils had been discovered in Gibraltar even earlier, but it was originally claimed that these were the remains of a modern human who had suffered some kind of illness.<a href="#cite_note-FOOTNOTEMontgomery198895–96-234">[228]</a> Despite the 1891 discovery by <a href="Eug%C3%A8ne_Dubois" title="Eugène Dubois">Eugène Dubois</a> of what is now called Homo erectus at <a href="Trinil" title="Trinil">Trinil</a>, <a href="Java" title="Java">Java</a>, it was only in the 1920s when such fossils were discovered in Africa, that intermediate species began to accumulate.<a href="#cite_note-235">[229]</a> In 1925, <a href="Raymond_Dart" title="Raymond Dart">Raymond Dart</a> described <a href="Australopithecus_africanus" title="Australopithecus africanus">Australopithecus africanus</a>.<a href="#cite_note-236">[230]</a> The <a href="Type_(biology)" title="Type (biology)">type specimen</a> was the <a href="Taung_Child" title="Taung Child">Taung Child</a>, an australopithecine infant which was discovered in a cave. The child's remains were a remarkably well-preserved tiny skull and an <a href="Endocast" title="Endocast">endocast</a> of the brain.

Although the brain was small (410 cm3), its shape was rounded, unlike that of chimpanzees and gorillas, and more like a modern human brain. Also, the specimen showed short <a href="Canine_tooth" title="Canine tooth">canine teeth</a>, and the position of the <a href="Foramen_magnum" title="Foramen magnum">foramen magnum</a> (the hole in the skull where the spine enters) was evidence of <a href="Bipedal" class="mw-redirect" title="Bipedal">bipedal</a> locomotion. All of these traits convinced Dart that the Taung Child was a bipedal human ancestor, a transitional form between apes and humans.

<div class="mw-heading mw-heading3"><h3 id="The_East_African_fossils">The East African fossils</h3></div>

During the 1960s and 1970s, hundreds of fossils were found in East Africa in the regions of the <a href="Olduvai_Gorge" title="Olduvai Gorge">Olduvai Gorge</a> and <a href="Lake_Turkana" title="Lake Turkana">Lake Turkana</a>. These searches were carried out by the Leakey family, with <a href="Louis_Leakey" title="Louis Leakey">Louis Leakey</a> and his wife <a href="Mary_Leakey" title="Mary Leakey">Mary Leakey</a>, and later their son <a href="Richard_Leakey" title="Richard Leakey">Richard</a> and daughter-in-law <a href="Meave_Leakey" title="Meave Leakey">Meave</a>, fossil hunters and paleoanthropologists. From the fossil beds of Olduvai and Lake Turkana they amassed specimens of the early hominins: the australopithecines and Homo species, and even H. erectus.

These finds cemented Africa as the cradle of humankind. In the late 1970s and the 1980s, <a href="Ethiopia" title="Ethiopia">Ethiopia</a> emerged as the new hot spot of paleoanthropology after <a href="Lucy_(Australopithecus)" title="Lucy (Australopithecus)">"Lucy"</a>, the most complete fossil member of the species <a href="Australopithecus_afarensis" title="Australopithecus afarensis">Australopithecus afarensis</a>, was found in 1974 by <a href="Donald_Johanson" title="Donald Johanson">Donald Johanson</a> near <a href="Hadar%2C_Ethiopia" title="Hadar, Ethiopia">Hadar</a> in the desertic <a href="Afar_Triangle" title="Afar Triangle">Afar Triangle</a> region of northern Ethiopia. Although the specimen had a small brain, the pelvis and leg bones were almost identical in function to those of modern humans, showing with certainty that these hominins had walked erect.<a href="#cite_note-FOOTNOTEJohansonEdey198120–22,_184–185-237">[231]</a> Lucy was classified as a new species, Australopithecus afarensis, which is thought to be more closely related to the genus Homo as a direct ancestor, or as a close relative of an unknown ancestor, than any other known hominid or hominin from this early time range.<a href="#cite_note-238">[232]</a> (The specimen was nicknamed "Lucy" after <a href="The_Beatles" title="The Beatles">the Beatles</a>' song "<a href="Lucy_in_the_Sky_with_Diamonds" title="Lucy in the Sky with Diamonds">Lucy in the Sky with Diamonds</a>", which was played loudly and repeatedly in the camp during the excavations.)<a href="#cite_note-FOOTNOTEJohansonEdey198122-239">[233]</a> The <a href="Afar_Triangle" title="Afar Triangle">Afar Triangle</a> area would later yield discovery of many more hominin fossils, particularly those uncovered or described by teams headed by <a href="Tim_D._White" title="Tim D. White">Tim D. White</a> in the 1990s, including <a href="Ardipithecus_ramidus" title="Ardipithecus ramidus">Ardipithecus ramidus</a> and <a href="Ardipithecus_kadabba" title="Ardipithecus kadabba">A. kadabba</a>.<a href="#cite_note-240">[234]</a>

In 2013, fossil skeletons of <a href="Homo_naledi" title="Homo naledi">Homo naledi</a>, an <a href="Extinct_species" class="mw-redirect" title="Extinct species">extinct species</a> of <a href="Hominin" class="mw-redirect" title="Hominin">hominin</a> assigned (provisionally) to the genus Homo, were found in the <a href="Rising_Star_Cave" title="Rising Star Cave">Rising Star Cave</a> system, a site in South Africa's <a href="Cradle_of_Humankind" title="Cradle of Humankind">Cradle of Humankind</a> region in <a href="Gauteng" title="Gauteng">Gauteng</a> province near <a href="Johannesburg" title="Johannesburg">Johannesburg</a>.<a href="#cite_note-241">[235]</a><a href="#cite_note-NGS-20150910-242">[236]</a> As of September 2015, fossils of at least fifteen individuals, amounting to 1,550 specimens, have been excavated from the cave.<a href="#cite_note-NGS-20150910-242">[236]</a> The species is characterized by a body mass and stature similar to small-bodied human populations, a smaller <a href="Endocranium" title="Endocranium">endocranial</a> volume similar to <a href="Australopithecus" title="Australopithecus">Australopithecus</a>, and a <a href="Skull" title="Skull">cranial</a> <a href="Morphology_(biology)" title="Morphology (biology)">morphology</a> (skull shape) similar to early Homo species. The skeletal anatomy combines primitive features known from <a href="Australopithecine" title="Australopithecine">australopithecines</a> with features known from early hominins. The individuals show signs of having been deliberately disposed of within the cave near the time of death. The fossils were dated close to 250,000 years ago,<a href="#cite_note-243">[237]</a> and thus are not ancestral but contemporary with the first appearance of larger-brained <a href="Anatomically_modern_humans" class="mw-redirect" title="Anatomically modern humans">anatomically modern humans</a>.<a href="#cite_note-eLIFE-2017a-244">[238]</a>

<div class="mw-heading mw-heading3"><h3 id="The_genetic_revolution">The genetic revolution</h3></div>

The genetic revolution in studies of human evolution started when <a href="Vincent_Sarich" title="Vincent Sarich">Vincent Sarich</a> and <a href="Allan_Wilson_(biologist)" title="Allan Wilson (biologist)">Allan Wilson</a> measured the strength of immunological cross-reactions of <a href="Serum_(blood)" title="Serum (blood)">blood serum</a> <a href="Albumin" title="Albumin">albumin</a> between pairs of creatures, including humans and African apes (chimpanzees and gorillas).<a href="#cite_note-MolClock-245">[239]</a> The strength of the reaction could be expressed numerically as an immunological distance, which was in turn proportional to the number of <a href="Amino_acid" title="Amino acid">amino acid</a> differences between homologous proteins in different species. By constructing a calibration curve of the ID of species' pairs with known divergence times in the fossil record, the data could be used as a <a href="Molecular_clock" title="Molecular clock">molecular clock</a> to estimate the times of divergence of pairs with poorer or unknown fossil records.

In their seminal 1967 paper in <a href="Science_(journal)" title="Science (journal)">Science</a>, Sarich and Wilson estimated the divergence time of humans and apes as four to five million years ago,<a href="#cite_note-MolClock-245">[239]</a> at a time when standard interpretations of the fossil record gave this divergence as at least 10 to as much as 30 million years. Subsequent fossil discoveries, notably "Lucy", and reinterpretation of older fossil materials, notably <a href="Sivapithecus" title="Sivapithecus">Ramapithecus</a>, showed the younger estimates to be correct and validated the albumin method.

Progress in <a href="DNA_sequencing" title="DNA sequencing">DNA sequencing</a>, specifically <a href="Mitochondrial_DNA" title="Mitochondrial DNA">mitochondrial DNA</a> (mtDNA) and then <a href="Human_Y-chromosome_DNA_haplogroup" title="Human Y-chromosome DNA haplogroup">Y-chromosome DNA</a> (Y-DNA) advanced the understanding of human origins.<a href="#cite_note-FOOTNOTEDeSalleTattersall2008[httpsbooksgooglecombooksidBf4Sitw7YaICpgPA146_146]-128">[126]</a><a href="#cite_note-FOOTNOTEM'charek2005[httpsbooksgooglecombooksidn-K436zeUY8CpgPA96_96]-246">[240]</a><a href="#cite_note-FOOTNOTETrent2005[httpsbooksgooglecombooksid772BEp8ZyMYCpgPA6_6]-247">[241]</a> Application of the <a href="Molecular_clock" title="Molecular clock">molecular clock</a> principle revolutionized the study of <a href="Molecular_evolution" title="Molecular evolution">molecular evolution</a>.

On the basis of a separation from the <a href="Orangutan" title="Orangutan">orangutan</a> between 10 and 20 million years ago, earlier studies of the molecular clock suggested that there were about 76 mutations per generation that were not inherited by human children from their parents; this evidence supported the divergence time between hominins and chimpanzees noted above. However, a 2012 study in Iceland of 78 children and their parents suggests a mutation rate of only 36 mutations per generation; this datum extends the separation between humans and chimpanzees to an earlier period greater than 7 million years ago (<a href="Year#SI_prefix_multipliers" title="Year">Ma</a>). Additional research with 226 offspring of wild chimpanzee populations in eight locations suggests that chimpanzees reproduce at age 26.5 years on average; which suggests the human divergence from chimpanzees occurred between 7 and 13 mya. And these data suggest that Ardipithecus (4.5 Ma), Orrorin (6 Ma) and Sahelanthropus (7 Ma) all may be on the hominid <a href="Lineage_(evolution)" title="Lineage (evolution)">lineage</a>, and even that the separation may have occurred outside the <a href="East_African_Rift" title="East African Rift">East African Rift</a> region.

Furthermore, analysis of the two species' genes in 2006 provides evidence that after human ancestors had started to diverge from chimpanzees, interspecies mating between "proto-human" and "proto-chimpanzees" nonetheless occurred regularly enough to change certain genes in the new <a href="Gene_pool" title="Gene pool">gene pool</a>:

<dl><dd>A new comparison of the human and chimpanzee genomes suggests that after the two lineages separated, they may have begun interbreeding... A principal finding is that the <a href="X_chromosome" title="X chromosome">X chromosomes</a> of humans and chimpanzees appear to have <a href="Genetic_divergence" title="Genetic divergence">diverged</a> about 1.2 million years more recently than the other chromosomes.</dd></dl>

The research suggests:

<dl><dd>There were in fact two splits between the human and chimpanzee lineages, with the first being followed by interbreeding between the two populations and then a second split. The suggestion of a hybridization has startled paleoanthropologists, who nonetheless are treating the new genetic data seriously.<a href="#cite_note-248">[242]</a></dd></dl>

<div class="mw-heading mw-heading3"><h3 id="The_quest_for_the_earliest_hominin">The quest for the earliest hominin</h3></div>

In the 1990s, several teams of paleoanthropologists were working throughout Africa looking for evidence of the earliest divergence of the hominin lineage from the great apes. In 1994, Meave Leakey discovered <a href="Australopithecus_anamensis" title="Australopithecus anamensis">Australopithecus anamensis</a>. The find was overshadowed by Tim D. White's 1995 discovery of Ardipithecus ramidus, which pushed back the fossil record to <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=4.2">4.2</a> million years ago.

In 2000, <a href="Martin_Pickford" title="Martin Pickford">Martin Pickford</a> and <a href="Brigitte_Senut" title="Brigitte Senut">Brigitte Senut</a> discovered, in the <a href="Tugen_Hills" title="Tugen Hills">Tugen Hills</a> of <a href="Kenya" title="Kenya">Kenya</a>, a 6-million-year-old bipedal hominin which they named <a href="Orrorin" title="Orrorin">Orrorin tugenensis</a>. And in 2001, a team led by <a href="Michel_Brunet_(paleontologist)" title="Michel Brunet (paleontologist)">Michel Brunet</a> discovered the skull of <a href="Sahelanthropus" title="Sahelanthropus">Sahelanthropus tchadensis</a> which was dated as <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=7.2">7.2</a> million years ago, and which Brunet argued was a bipedal, and therefore a hominid—that is, a hominin (<abbr title="confer">cf</abbr> <a href="Hominidae" title="Hominidae">Hominidae</a>; terms "hominids" and hominins).

<div class="mw-heading mw-heading3"><h3 id="Human_dispersal">Human dispersal</h3></div>

<div role="note" class="hatnote navigation-not-searchable">See also: <a href="Early_human_migrations" title="Early human migrations">Early human migrations</a>, <a href="Recent_African_origin_of_modern_humans" title="Recent African origin of modern humans">Recent African origin of modern humans</a>, <a href="Multiregional_origin_of_modern_humans" title="Multiregional origin of modern humans">Multiregional origin of modern humans</a>, and <a href="Archaic_humans_in_Southeast_Asia" title="Archaic humans in Southeast Asia">Archaic humans in Southeast Asia</a></div>

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</style><div class="thumb tmulti tright"><div class="thumbinner multiimageinner" style="width:562px;max-width:562px"><div class="trow"><div class="tsingle" style="width:324px;max-width:324px"><div class="thumbimage" style="height:229px;overflow:hidden"></div><div class="thumbcaption">A global mapping model of human migration, based from divergence of the <a href="Mitochondrial_DNA" title="Mitochondrial DNA">mitochondrial DNA</a> (which indicates the <a href="Lineage_(genetic)" title="Lineage (genetic)">matrilineage</a>).<a href="#cite_note-Behar_2008-249">[243]</a><a href="#cite_note-Gonder_2007-250">[244]</a><a href="#cite_note-Tishkoff_2009-251">[245]</a> Timescale (<a href="Year#SI_prefix_multipliers" title="Year">ka</a>) indicated by colours.</div></div><div class="tsingle" style="width:234px;max-width:234px"><div class="thumbimage" style="height:229px;overflow:hidden"></div><div class="thumbcaption">A "trellis" (as <a href="Milford_H._Wolpoff" title="Milford H. Wolpoff">Milford H. Wolpoff</a> called it) that emphasizes back-and-forth gene flow among geographic regions<a href="#cite_note-252">[246]</a></div></div></div><div class="trow" style="display:flex"><div class="thumbcaption">Different models for the beginning of the present human species</div></div></div></div>

Anthropologists in the 1980s were divided regarding some details of reproductive barriers and migratory dispersals of the genus Homo. Subsequently, genetics has been used to investigate and resolve these issues. According to the <a href="Sahara_pump_theory" title="Sahara pump theory">Sahara pump theory</a> evidence suggests that the genus Homo have migrated out of Africa at least three and possibly four times (e.g. Homo erectus, Homo heidelbergensis and two or three times for Homo sapiens). Recent evidence suggests these dispersals are closely related to fluctuating periods of climate change.<a href="#cite_note-253">[247]</a>

Recent evidence suggests that humans may have left Africa half a million years earlier than previously thought. A joint Franco-Indian team has found human artifacts in the Siwalk Hills north of New Delhi dating back at least 2.6 million years. This is earlier than the previous earliest finding of genus Homo at <a href="Dmanisi" title="Dmanisi">Dmanisi</a>, in <a href="Georgia_(country)" title="Georgia (country)">Georgia</a>, dating to 1.85 million years. Although controversial, tools found at a Chinese cave strengthen the case that humans used tools as far back as 2.48 million years ago.<a href="#cite_note-254">[248]</a> This suggests that the Asian "Chopper" tool tradition, found in Java and northern China may have left Africa before the appearance of the <a href="Acheulian" class="mw-redirect" title="Acheulian">Acheulian</a> hand axe.

<div class="mw-heading mw-heading4"><h4 id="Dispersal_of_modern_Homo_sapiens">Dispersal of modern Homo sapiens</h4></div>

Up until the genetic evidence became available, there were two dominant models for the dispersal of modern humans. The <a href="Multiregional_evolution" class="mw-redirect" title="Multiregional evolution">multiregional hypothesis</a> proposed that the genus Homo contained only a single interconnected population as it does today (not separate species), and that its evolution took place worldwide continuously over the last couple of million years. This model was proposed in 1988 by <a href="Milford_H._Wolpoff" title="Milford H. Wolpoff">Milford H. Wolpoff</a>.<a href="#cite_note-255">[249]</a><a href="#cite_note-256">[250]</a> In contrast, the "out of Africa" model proposed that modern H. sapiens speciated in Africa recently (that is, approximately 200,000 years ago) and the subsequent migration through <a href="Eurasia" title="Eurasia">Eurasia</a> resulted in the nearly complete replacement of other Homo species. This model has been developed by <a href="Chris_Stringer" title="Chris Stringer">Chris Stringer</a> and Peter Andrews.<a href="#cite_note-257">[251]</a><a href="#cite_note-258">[252]</a>

Sequencing mtDNA and Y-DNA sampled from a wide range of indigenous populations revealed ancestral information relating to both male and female genetic heritage, and strengthened the "out of Africa" theory and weakened the views of multiregional evolutionism.<a href="#cite_note-FOOTNOTEWebster2010[httpsbooksgooglecombooksidcdiy1EhJVW0CpgPA53_53]-259">[253]</a> Aligned in genetic tree differences were interpreted as supportive of a recent single origin.<a href="#cite_note-FOOTNOTERamachandranTangGutenkunstBustamante2010[httpsbooksgooglecombooksidFlfPSpBvKLgCpgPA606_606]-260">[254]</a>

"Out of Africa" has thus gained much support from research using female mitochondrial DNA and the male <a href="Y_chromosome" title="Y chromosome">Y chromosome</a>. After analysing genealogy trees constructed using 133 types of mtDNA, researchers concluded that all were descended from a female African progenitor, dubbed <a href="Mitochondrial_Eve" title="Mitochondrial Eve">Mitochondrial Eve</a>. "Out of Africa" is also supported by the fact that mitochondrial genetic diversity is highest among African populations.<a href="#cite_note-Cann-261">[255]</a>

A broad study of African genetic diversity, headed by <a href="Sarah_Tishkoff" title="Sarah Tishkoff">Sarah Tishkoff</a>, found the <a href="San_people" title="San people">San people</a> had the greatest genetic diversity among the 113 distinct populations sampled, making them one of 14 "ancestral population clusters". The research also located a possible origin of modern human migration in southwestern Africa, near the coastal border of <a href="Namibia" title="Namibia">Namibia</a> and <a href="Angola" title="Angola">Angola</a>.<a href="#cite_note-262">[256]</a> The fossil evidence was insufficient for archaeologist <a href="Richard_Leakey" title="Richard Leakey">Richard Leakey</a> to resolve the debate about exactly where in Africa modern humans first appeared.<a href="#cite_note-FOOTNOTELeakey199487–89-263">[257]</a> Studies of <a href="Haplogroup" title="Haplogroup">haplogroups</a> in <a href="Human_Y-chromosome_DNA_haplogroup" title="Human Y-chromosome DNA haplogroup">Y-chromosomal DNA</a> and <a href="Human_mitochondrial_DNA_haplogroup" title="Human mitochondrial DNA haplogroup">mitochondrial DNA</a> have largely supported a recent African origin.<a href="#cite_note-264">[258]</a> All the evidence from autosomal DNA also predominantly supports a Recent African origin. However, evidence for <a href="Archaic_human_admixture_with_modern_humans" class="mw-redirect" title="Archaic human admixture with modern humans">archaic admixture in modern humans</a>, both in Africa and later, throughout Eurasia has recently been suggested by a number of studies.<a href="#cite_note-265">[259]</a>

Recent sequencing of Neanderthal<a href="#cite_note-greenetal-94">[92]</a> and Denisovan<a href="#cite_note-pmid21179161-46">[44]</a> genomes shows that some admixture with these populations has occurred. All modern human groups outside Africa have 1–4% or (according to more recent research) about 1.5–2.6% Neanderthal <a href="Allele" title="Allele">alleles</a> in their genome,<a href="#cite_note-A_high-coverage_Neandertal_genome_f-95">[93]</a> and some <a href="Melanesians" title="Melanesians">Melanesians</a> have an additional 4–6% of Denisovan alleles. These new results do not contradict the "out of Africa" model, except in its strictest interpretation, although they make the situation more complex. After recovery from a <a href="Genetic_bottleneck" class="mw-redirect" title="Genetic bottleneck">genetic bottleneck</a> that some researchers speculate might be linked to the <a href="Toba_catastrophe_theory" class="mw-redirect" title="Toba catastrophe theory">Toba supervolcano catastrophe</a>, a fairly small group left Africa and interbred with Neanderthals, probably in the Middle East, on the Eurasian steppe or even in North Africa before their departure. Their still predominantly African descendants spread to populate the world. A fraction in turn interbred with Denisovans, probably in southeastern Asia, before populating Melanesia.<a href="#cite_note-Reich_2011-104">[102]</a> <a href="Human_leukocyte_antigen" title="Human leukocyte antigen">HLA</a> haplotypes of Neanderthal and Denisova origin have been identified in modern Eurasian and Oceanian populations.<a href="#cite_note-10.1126/science.1209202-48">[46]</a> The Denisovan <a href="EPAS1" title="EPAS1">EPAS1</a> gene has also been found in Tibetan populations.<a href="#cite_note-266">[260]</a> Studies of the human genome using machine learning have identified additional genetic contributions in Eurasians from an "unknown" ancestral population potentially related to the Neanderthal-Denisovan lineage.<a href="#cite_note-267">[261]</a>

There are still differing theories on whether there was a single exodus from Africa or several. A multiple dispersal model involves the Southern Dispersal theory,<a href="#cite_note-lahr-268">[262]</a><a href="#cite_note-269">[263]</a><a href="#cite_note-270">[264]</a> which has gained support in recent years from genetic, linguistic and archaeological evidence. In this theory, there was a coastal dispersal of modern humans from the <a href="Horn_of_Africa" title="Horn of Africa">Horn of Africa</a> crossing the Bab el Mandib to Yemen at a lower sea level around 70,000 years ago. This group helped to populate Southeast Asia and Oceania, explaining the discovery of early human sites in these areas much earlier than those in the <a href="Levant" title="Levant">Levant</a>.<a href="#cite_note-lahr-268">[262]</a> This group seems to have been dependent upon marine resources for their survival.

<a href="Stephen_Oppenheimer" title="Stephen Oppenheimer">Stephen Oppenheimer</a> has proposed a second wave of humans may have later dispersed through the Persian Gulf oases, and the Zagros mountains into the Middle East. Alternatively it may have come across the <a href="Sinai_Peninsula" title="Sinai Peninsula">Sinai Peninsula</a> into Asia, from shortly after 50,000 yrs BP, resulting in the bulk of the human populations of Eurasia. It has been suggested that this second group possibly possessed a more sophisticated "big game hunting" tool technology and was less dependent on coastal food sources than the original group. Much of the evidence for the first group's expansion would have been destroyed by the rising sea levels at the end of each <a href="Glacial_maximum" class="mw-redirect" title="Glacial maximum">glacial maximum</a>.<a href="#cite_note-lahr-268">[262]</a> The multiple dispersal model is contradicted by studies indicating that the populations of Eurasia and the populations of Southeast Asia and Oceania are all descended from the same mitochondrial DNA L3 <a href="Lineage_(genetic)" title="Lineage (genetic)">lineages</a>, which support a single migration out of Africa that gave rise to all non-African populations.<a href="#cite_note-271">[265]</a>

On the basis of the early date of Badoshan Iranian Aurignacian, Oppenheimer suggests that this second dispersal may have occurred with a pluvial period about 50,000 years before the present, with modern human big-game hunting cultures spreading up the Zagros Mountains, carrying modern human genomes from Oman, throughout the Persian Gulf, northward into Armenia and Anatolia, with a variant travelling south into Israel and to Cyrenicia.<a href="#cite_note-oppenheimer-205">[201]</a>

Recent genetic evidence suggests that all modern non-African populations, including those of Eurasia and Oceania, are descended from a single wave that left Africa between 65,000 and 50,000 years ago.<a href="#cite_note-Posth-272">[266]</a><a href="#cite_note-273">[267]</a><a href="#cite_note-ReferenceC-274">[268]</a>

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

The evidence on which scientific accounts of human evolution are based comes from many fields of <a href="Natural_science" title="Natural science">natural science</a>. The main source of knowledge about the evolutionary process has traditionally been the fossil record, but since the development of genetics beginning in the 1970s, DNA analysis has come to occupy a place of comparable importance. The studies of ontogeny, <a href="Phylogenetics" title="Phylogenetics">phylogeny</a> and especially <a href="Evolutionary_developmental_biology" title="Evolutionary developmental biology">evolutionary developmental biology</a> of both vertebrates and invertebrates offer considerable insight into the evolution of all life, including how humans evolved. The specific study of the origin and life of humans is <a href="Anthropology" title="Anthropology">anthropology</a>, particularly paleoanthropology which focuses on the study of human <a href="Prehistory" title="Prehistory">prehistory</a>.<a href="#cite_note-FOOTNOTEStanfordAllenAntón2009-275">[269]</a>

<div class="mw-heading mw-heading3"><h3 id="Evidence_from_genetics">Evidence from genetics</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main articles: <a href="Human_evolutionary_genetics" title="Human evolutionary genetics">Human evolutionary genetics</a> and <a href="Human_genetic_variation" title="Human genetic variation">Human genetic variation</a></div>

The closest living relatives of humans are bonobos and chimpanzees (both genus Pan) and gorillas (genus Gorilla).<a href="#cite_note-Wood-276">[270]</a> With the sequencing of both the human and chimpanzee genome, as of 2012 estimates of the similarity between their DNA sequences range between 95% and 99%.<a href="#cite_note-Wood-276">[270]</a><a href="#cite_note-277">[271]</a><a href="#cite_note-sayers-33">[31]</a> It is also noteworthy that mice share around 97.5% of their working DNA with humans.<a href="#cite_note-278">[272]</a> By using the technique called the <a href="Molecular_clock" title="Molecular clock">molecular clock</a> which estimates the time required for the number of divergent mutations to accumulate between two lineages, the approximate date for the split between lineages can be calculated.

The gibbons (family Hylobatidae) and then the orangutans (genus Pongo) were the first groups to split from the line leading to the hominins, including humans—followed by gorillas (genus Gorilla), and, ultimately, by the chimpanzees (genus Pan). The splitting date between hominin and chimpanzee lineages is placed by some between <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=4–8">4 to 8</a> million years ago, that is, during the <a href="Late_Miocene" title="Late Miocene">Late Miocene</a>.<a href="#cite_note-FOOTNOTEDawkins2004-279">[273]</a><a href="#cite_note-280">[274]</a><a href="#cite_note-281">[275]</a><a href="#cite_note-Ruvolo1997-282">[276]</a> Speciation, however, appears to have been unusually drawn out. Initial divergence occurred sometime between <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=7–13">7 to 13</a> million years ago, but ongoing hybridization blurred the separation and delayed complete separation during several millions of years. Patterson (2006) dated the final divergence at <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=5–6">5 to 6</a> million years ago.<a href="#cite_note-283">[277]</a>

Genetic evidence has also been employed to compare species within the genus Homo, investigating <a href="Archaic_human_admixture_with_modern_humans" class="mw-redirect" title="Archaic human admixture with modern humans">gene flow between early modern humans and Neanderthals</a>, and to enhance the understanding of the early human migration patterns and splitting dates. By comparing the parts of the genome that are <a href="Neutral_theory_of_molecular_evolution" title="Neutral theory of molecular evolution">not under natural selection</a> and which therefore accumulate mutations at a fairly steady rate, it is possible to reconstruct a genetic tree incorporating the entire human species since the last shared ancestor.

Each time a certain mutation (<a href="Single-nucleotide_polymorphism" title="Single-nucleotide polymorphism">single-nucleotide polymorphism</a>) appears in an individual and is passed on to his or her descendants, a haplogroup is formed including all of the descendants of the individual who will also carry that mutation. By comparing mitochondrial <a href="DNA" title="DNA">DNA</a> which is inherited only from the mother, geneticists have concluded that the last female common ancestor whose <a href="Genetic_marker" title="Genetic marker">genetic marker</a> is found in all modern humans, the so-called <a href="Mitochondrial_Eve" title="Mitochondrial Eve">mitochondrial Eve</a>, must have lived around 200,000 years ago.

Human evolutionary genetics studies how <a href="Human_genome" title="Human genome">human genomes</a> differ among individuals, the evolutionary past that gave rise to them, and their current effects. Differences between genomes have <a href="Anthropology" title="Anthropology">anthropological</a>, medical and <a href="Forensic_science" title="Forensic science">forensic</a> implications and applications. Genetic data can provide important insight into human evolution.

In May 2023, scientists reported a more complicated pathway of human evolution than previously understood. According to the studies, humans evolved from different places and times in Africa, instead of from a single location and period of time.<a href="#cite_note-NYT-20230517-284">[278]</a><a href="#cite_note-NAT-20230517-285">[279]</a>

<div class="mw-heading mw-heading3"><h3 id="Evidence_from_the_fossil_record">Evidence from the fossil record</h3></div>

There is little fossil evidence for the divergence of the gorilla, chimpanzee and hominin lineages.<a href="#cite_note-286">[280]</a> The earliest fossils that have been proposed as members of the hominin lineage are Sahelanthropus tchadensis dating from <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=7">7</a> million years ago, Orrorin tugenensis dating from <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=5.7">5.7</a> million years ago, and Ardipithecus kadabba dating to <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=5.6">5.6</a> million years ago. Each of these have been argued to be a <a href="Bipedal" class="mw-redirect" title="Bipedal">bipedal</a> ancestor of later hominins but, in each case, the claims have been contested. It is also possible that one or more of these species are ancestors of another branch of African apes, or that they represent a shared ancestor between hominins and other apes.

The question then of the relationship between these early fossil species and the hominin lineage is still to be resolved. From these early species, the australopithecines arose around <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=4">4</a> million years ago and diverged into <a href="Paranthropus" title="Paranthropus">robust</a> (also called <a href="Paranthropus" title="Paranthropus">Paranthropus</a>) and <a href="Australopithecus" title="Australopithecus">gracile</a> branches, one of which (possibly <a href="Australopithecus_garhi" title="Australopithecus garhi">A. garhi</a>) probably went on to become ancestors of the genus Homo. The australopithecine species that is best represented in the fossil record is Australopithecus afarensis with more than 100 fossil individuals represented, found from Northern Ethiopia (such as the famous "Lucy"), to Kenya, and South Africa. Fossils of robust australopithecines such as A. robustus (or alternatively <a href="Paranthropus_robustus" title="Paranthropus robustus">Paranthropus robustus</a>) and A./P. boisei are particularly abundant in South Africa at sites such as <a href="Kromdraai" class="mw-redirect" title="Kromdraai">Kromdraai</a> and <a href="Swartkrans" title="Swartkrans">Swartkrans</a>, and around <a href="Lake_Turkana" title="Lake Turkana">Lake Turkana</a> in Kenya.

The earliest member of the genus Homo is Homo habilis which evolved around <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=2.8">2.8</a> million years ago.<a href="#cite_note-autogenerated1-36">[34]</a> H. habilis is the first species for which we have positive evidence of the use of stone tools. They developed the <a href="Oldowan" title="Oldowan">Oldowan</a> lithic technology, named after the Olduvai Gorge in which the first specimens were found. Some scientists consider <a href="Homo_rudolfensis" title="Homo rudolfensis">Homo rudolfensis</a>, a larger bodied group of fossils with similar morphology to the original H. habilis fossils, to be a separate species, while others consider them to be part of H. habilis—simply representing intraspecies variation, or perhaps even <a href="Sexual_dimorphism" title="Sexual dimorphism">sexual dimorphism</a>. The brains of these early hominins were about the same size as that of a chimpanzee, and their main adaptation was bipedalism as an adaptation to terrestrial living.

During the next million years, a process of encephalization began and, by the arrival (about <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=1.9">1.9</a> million years ago) of H. erectus in the fossil record, cranial capacity had doubled. H. erectus were the first of the hominins to emigrate from Africa, and, from <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=1.8–1.3">1.8 to 1.3</a> million years ago, this species spread through Africa, Asia, and Europe. One population of H. erectus, also sometimes classified as separate species H. ergaster, remained in Africa and evolved into H. sapiens. It is believed that H. erectus and H. ergaster were the first to use fire and complex tools. In Eurasia, H. erectus evolved into species such as <a href="H._antecessor" class="mw-redirect" title="H. antecessor">H. antecessor</a>, <a href="H._heidelbergensis" class="mw-redirect" title="H. heidelbergensis">H. heidelbergensis</a> and <a href="H._neanderthalensis" class="mw-redirect" title="H. neanderthalensis">H. neanderthalensis</a>. The earliest fossils of anatomically modern humans are from the Middle Paleolithic, about 300–200,000 years ago such as the Herto and <a href="Omo_remains" title="Omo remains">Omo remains</a> of Ethiopia, <a href="Jebel_Irhoud" title="Jebel Irhoud">Jebel Irhoud</a> remains of Morocco, and Florisbad remains of South Africa; <a href="Skhul_and_Qafzeh_hominins" title="Skhul and Qafzeh hominins">later fossils</a> from the <a href="Skhul_Cave" title="Skhul Cave">Skhul Cave</a> in Israel and Southern Europe begin around 90,000 years ago (<a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=0.09">0.09</a> million years ago).

As modern humans spread out from Africa, they encountered other hominins such as H. neanderthalensis and the Denisovans, who may have evolved from populations of H. erectus that had left Africa around <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=2">2</a> million years ago. The nature of interaction between early humans and these sister species has been a long-standing source of controversy, the question being whether humans replaced these earlier species or whether they were in fact similar enough to interbreed, in which case these earlier populations may have contributed genetic material to modern humans.<a href="#cite_note-FOOTNOTEWood2009[httpslinkspringercomchapter1010072F978-1-4020-9980-9_3page-1_17–27]-287">[281]</a><a href="#cite_note-NYT-01302012-288">[282]</a>

This migration out of Africa is estimated to have begun about 70–50,000 years <a href="Before_Present" title="Before Present">BP</a> and modern humans subsequently spread globally, replacing earlier hominins either through competition or hybridization. They inhabited Eurasia and Oceania by 40,000 years BP, and the Americas by at least 14,500 years BP.<a href="#cite_note-289">[283]</a>

<div class="mw-heading mw-heading3"><h3 id="Inter-species_breeding">Inter-species breeding</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Further information: <a href="Interbreeding_between_archaic_and_modern_humans" title="Interbreeding between archaic and modern humans">Interbreeding between archaic and modern humans</a></div>

The hypothesis of interbreeding, also known as hybridization, admixture or hybrid-origin theory, has been discussed ever since the discovery of Neanderthal remains in the 19th century.<a href="#cite_note-290">[284]</a> The linear view of human evolution began to be abandoned in the 1970s as different species of humans were discovered that made the linear concept increasingly unlikely. In the 21st century with the advent of molecular biology techniques and computerization, <a href="Whole-genome_sequencing" class="mw-redirect" title="Whole-genome sequencing">whole-genome sequencing</a> of Neanderthal and human <a href="Genome" title="Genome">genome</a> were performed, confirming recent admixture between different human species.<a href="#cite_note-greenetal-94">[92]</a> In 2010, evidence based on molecular biology was published, revealing unambiguous examples of interbreeding between archaic and modern humans during the <a href="Middle_Paleolithic" title="Middle Paleolithic">Middle Paleolithic</a> and early <a href="Upper_Paleolithic" title="Upper Paleolithic">Upper Paleolithic</a>. It has been demonstrated that interbreeding happened in several independent events that included Neanderthals and Denisovans, as well as several unidentified hominins.<a href="#cite_note-291">[285]</a> Today, approximately 2% of DNA from all non-African populations (including Europeans, Asians, and <a href="Oceanians" class="mw-redirect" title="Oceanians">Oceanians</a>) is Neanderthal,<a href="#cite_note-greenetal-94">[92]</a> with traces of Denisovan heritage.<a href="#cite_note-NG-20180822-292">[286]</a> Also, 4–6% of modern <a href="Melanesians" title="Melanesians">Melanesian</a> genetics are Denisovan.<a href="#cite_note-NG-20180822-292">[286]</a> Comparisons of the human genome to the genomes of Neandertals, Denisovans and apes can help identify features that set modern humans apart from other hominin species. In a 2016 <a href="Comparative_genomics" title="Comparative genomics">comparative genomics</a> study, a Harvard Medical School/UCLA research team made a world map on the distribution and made some predictions about where Denisovan and Neanderthal genes may be impacting modern human biology.<a href="#cite_note-293">[287]</a><a href="#cite_note-294">[288]</a>

For example, comparative studies in the mid-2010s found several <a href="Phenotypic_trait" title="Phenotypic trait">traits</a> related to neurological, immunological,<a href="#cite_note-295">[289]</a> developmental, and metabolic phenotypes, that were developed by archaic humans to European and Asian environments and inherited to modern humans through admixture with local hominins.<a href="#cite_note-296">[290]</a><a href="#cite_note-297">[291]</a>

Although the narratives of human evolution are often contentious, several discoveries since 2010 show that human evolution should not be seen as a simple linear or branched progression, but a mix of related species.<a href="#cite_note-pmid21179161-46">[44]</a><a href="#cite_note-Denisovans_&_Neandertals-5">[5]</a><a href="#cite_note-Human_Hybrids-6">[6]</a><a href="#cite_note-Mosaic_humans,_the_hybrid_species-7">[7]</a> In fact, genomic research has shown that hybridization between substantially diverged lineages is the rule, not the exception, in human evolution.<a href="#cite_note-Ackermann_2015-4">[4]</a> Furthermore, it is argued that hybridization was an essential creative force in the emergence of modern humans.<a href="#cite_note-Ackermann_2015-4">[4]</a>

<div class="mw-heading mw-heading3"><h3 id="Stone_tools">Stone tools</h3></div>

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

Stone tools are first attested around 2.6 million years ago, when hominins in Eastern Africa used so-called core <a href="Oldowan" title="Oldowan">tools</a>, <a href="Chopper_(archaeology)" title="Chopper (archaeology)">choppers</a> made out of round cores that had been split by simple strikes.<a href="#cite_note-Plummer-298">[292]</a> This marks the beginning of the <a href="Paleolithic" title="Paleolithic">Paleolithic</a>, or Old <a href="Stone_Age" title="Stone Age">Stone Age</a>; its end is taken to be the end of the last <a href="Last_glacial_period" class="mw-redirect" title="Last glacial period">Ice Age</a>, around 10,000 years ago. The Paleolithic is subdivided into the <a href="Lower_Paleolithic" title="Lower Paleolithic">Lower Paleolithic</a> (Early Stone Age), ending around 350,000–300,000 years ago, the <a href="Middle_Paleolithic" title="Middle Paleolithic">Middle Paleolithic</a> (Middle Stone Age), until 50,000–30,000 years ago, and the <a href="Upper_Paleolithic" title="Upper Paleolithic">Upper Paleolithic</a>, (Late Stone Age), 50,000–10,000 years ago.

Archaeologists working in the Great Rift Valley in Kenya have discovered the oldest known stone tools in the world. Dated to around 3.3 million years ago, the implements are some 700,000 years older than stone tools from Ethiopia that previously held this distinction.<a href="#cite_note-Harmand_310–315-199">[195]</a><a href="#cite_note-299">[293]</a><a href="#cite_note-300">[294]</a><a href="#cite_note-301">[295]</a>

The period from 700,000 to 300,000 years ago is also known as the <a href="Acheulean" title="Acheulean">Acheulean</a>, when H. ergaster (or erectus) made large stone <a href="Hand_axe" title="Hand axe">hand axes</a> out of <a href="Flint" title="Flint">flint</a> and <a href="Quartzite" title="Quartzite">quartzite</a>, at first quite rough (Early Acheulian), later "<a href="Retouch_(lithics)" title="Retouch (lithics)">retouched</a>" by additional, more-subtle strikes at the sides of the <a href="Lithic_flake" title="Lithic flake">flakes</a>. After 350,000 BP the more refined so-called <a href="Levallois_technique" title="Levallois technique">Levallois technique</a> was developed, a series of consecutive strikes, by which scrapers, slicers ("racloirs"), needles, and flattened needles were made.<a href="#cite_note-Plummer-298">[292]</a> Finally, after about 50,000 BP, ever more refined and specialized flint tools were made by the Neanderthals and the immigrant <a href="Cro-Magnon" title="Cro-Magnon">Cro-Magnons</a> (knives, blades, skimmers). Bone tools were also made by H. sapiens in Africa by 90,000–70,000 years ago<a href="#cite_note-Henshilwood_etal_2002-206">[202]</a><a href="#cite_note-302">[296]</a> and are also known from early H. sapiens sites in Eurasia by about 50,000 years ago.

<div class="mw-heading mw-heading2"><h2 id="Species_list">Species list</h2></div>

This list is in <a href="Chronological" class="mw-redirect" title="Chronological">chronological</a> order across the table by <a href="Genus" title="Genus">genus</a>. Some species/subspecies names are well-established, and some are less established – especially in genus Homo. Please see articles for more information.

<th><a href="Sahelanthropus" title="Sahelanthropus">Sahelanthropus</a>

</th>

<th><a href="Homo" title="Homo">Homo</a> (humans)

</th></tr>

<tr>

<td>S. tchadensis

</td>

<td><a href="Homo_gautengensis" title="Homo gautengensis">H. gautengensis</a>

</td></tr>

<tr>

<th><a href="Orrorin" title="Orrorin">Orrorin</a>

</th>

<td><a href="Homo_habilis" title="Homo habilis">H. habilis</a>

</td></tr>

<tr>

<td>O. tugenensis

</td>

<td><a href="Homo_rudolfensis" title="Homo rudolfensis">H. rudolfensis</a>

</td></tr>

<tr>

<th><a href="Ardipithecus" title="Ardipithecus">Ardipithecus</a>

</th>

<td><a href="Homo_floresiensis" title="Homo floresiensis">H. floresiensis</a>

</td></tr>

<tr>

<td><a href="Ardipithecus#Ardipithecus_kadabba" title="Ardipithecus">A. kadabba</a>

</td>

<td><a href="Homo_ergaster" title="Homo ergaster">H. ergaster</a>

</td></tr>

<tr>

<td><a href="Ardipithecus#Ardipithecus_ramidus" title="Ardipithecus">A. ramidus</a>

</td>

<td><a href="Homo_erectus" title="Homo erectus">H. erectus</a>

</td></tr>

<tr>

<th><a href="Australopithecus" title="Australopithecus">Australopithecus</a>

</th>

<td>• <a href="H._e._georgicus" class="mw-redirect" title="H. e. georgicus">H. e. georgicus</a>

</td></tr>

<tr>

<td><a href="Australopithecus_anamensis" title="Australopithecus anamensis">A. anamensis</a>

</td>

<td><a href="Homo_cepranensis" class="mw-redirect" title="Homo cepranensis">H. cepranensis</a>

</td></tr>

<tr>

<td><a href="Australopithecus_afarensis" title="Australopithecus afarensis">A. afarensis</a>

</td>

<td><a href="Homo_antecessor" title="Homo antecessor">H. antecessor</a>

</td></tr>

<tr>

<td><a href="Australopithecus_bahrelghazali" title="Australopithecus bahrelghazali">A. bahrelghazali</a>

</td>

<td><a href="Homo_heidelbergensis" title="Homo heidelbergensis">H. heidelbergensis</a>

</td></tr>

<tr>

<td><a href="Australopithecus_africanus" title="Australopithecus africanus">A. africanus</a>

</td>

<td><a href="Homo_rhodesiensis" title="Homo rhodesiensis">H. rhodesiensis</a>

</td></tr>

<tr>

<td><a href="Australopithecus_garhi" title="Australopithecus garhi">A. garhi</a>

</td>

<td><a href="Homo_naledi" title="Homo naledi">H. naledi</a>

</td></tr>

<tr>

<td><a href="Australopithecus_sediba" title="Australopithecus sediba">A. sediba</a>

</td>

<td><a href="Florisbad_Skull" title="Florisbad Skull">H. helmei</a>

</td></tr>

<tr>

<th><a href="Kenyanthropus" title="Kenyanthropus">Kenyanthropus</a>

</th>

<td><a href="Neanderthal" title="Neanderthal">H. neanderthalensis</a>

</td></tr>

<tr>

<td>K. platyops

</td>

<td><a href="H._sapiens" class="mw-redirect" title="H. sapiens">H. sapiens</a>

</td></tr>

<tr>

<th><a href="Paranthropus" title="Paranthropus">Paranthropus</a>

</th>

<td>• <a href="Homo_sapiens_idaltu" class="mw-redirect" title="Homo sapiens idaltu">H. s. idaltu</a>

</td></tr>

<tr>

<td><a href="Paranthropus_aethiopicus" title="Paranthropus aethiopicus">P. aethiopicus</a>

</td>

<td>• <a href="Cro-Magnon" title="Cro-Magnon">H. s. sapiens</a> (early)

</td></tr>

<tr>

<td><a href="Paranthropus_boisei" title="Paranthropus boisei">P. boisei</a>

</td>

<td>• <a href="H._s._sapiens" class="mw-redirect" title="H. s. sapiens">H. s. sapiens</a> (modern)

</td></tr>

<tr>

<td><a href="Paranthropus_robustus" title="Paranthropus robustus">P. robustus</a>

</td>

<td>

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

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<ul><li><a href="Adaptive_evolution_in_the_human_genome" title="Adaptive evolution in the human genome">Adaptive evolution in the human genome</a></li>

<li><a href="Amity%E2%80%93enmity_complex" title="Amity–enmity complex">Amity–enmity complex</a></li>

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

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

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

<li><a href="Evolution_of_morality" title="Evolution of morality">Evolution of morality</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_origin_of_religion" title="Evolutionary origin of religion">Evolutionary origin of religion</a></li>

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

<li><a href="Human_behavioral_ecology" title="Human behavioral ecology">Human behavioral ecology</a></li>

<li><a href="Human_origins_(disambiguation)" class="mw-redirect mw-disambig" title="Human origins (disambiguation)">Human origins</a></li>

<li><a href="Human_vestigiality" title="Human vestigiality">Human vestigiality</a></li>

<li><a href="List_of_human_evolution_fossils" title="List of human evolution fossils">List of human evolution fossils</a></li>

<li><a href="Molecular_paleontology" title="Molecular paleontology">Molecular paleontology</a></li>

<li><a href="Obstetrical_dilemma" title="Obstetrical dilemma">Obstetrical dilemma</a></li>

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

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

<li><a href="Prehistory_of_nakedness_and_clothing" title="Prehistory of nakedness and clothing">Prehistory of nakedness and clothing</a></li>

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

<li><a href="Transgenerational_trauma" title="Transgenerational trauma">Transgenerational trauma</a></li>

<li><a href="Timeline_of_human_evolution" title="Timeline of human evolution">Timeline of human evolution</a></li></ul>

</div>

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

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<li id="cite_note-habilis-12">^ <a href="#cite_ref-habilis_12-0">a</a> <a href="#cite_ref-habilis_12-1">b</a> <a href="#cite_ref-habilis_12-2">c</a> The conventional estimate on the age of H. habilis is at roughly 2.1 to 2.3 million years.<a href="#cite_note-FOOTNOTEStringer1994242-38">[36]</a><a href="#cite_note-170">[167]</a> Suggestions for pushing back the age to 2.8 Mya were made in 2015 based on the discovery of <a href="LD_350-1" title="LD 350-1">a jawbone</a>.<a href="#cite_note-171">[168]</a>

</li>

<li id="cite_note-29"><a href="#cite_ref-29">^</a> Not to be confused with <a href="Pongidae" title="Pongidae">Pongidae</a>, an obsolete family which grouped together <a href="Orangutan" title="Orangutan">orangutans</a>, <a href="Gorilla" title="Gorilla">gorillas</a> and <a href="Chimpanzee" title="Chimpanzee">chimpanzees</a> to separate them from humans

</li>

<li id="cite_note-143"><a href="#cite_ref-143">^</a> There is no general agreement on the line of <a href="Species" title="Species">special</a> descent of H. sapiens from H. erectus. Some of the species depicted in the image may not actually represent a direct evolutionary ancestor to H. sapiens, and may not directly derive from one another, namely:

<ul><li>H. heidelbergensis likely did not descend from H. antecessor.<a href="#cite_note-Welker2020-141">[139]</a></li>

<li>H. heidelbergensis is likely not an ancestor to H. sapiens, nor is H. antecessor.<a href="#cite_note-Welker2020-141">[139]</a></li>

<li>H. ergaster is often considered the next evolutionary ancestor to H. sapiens following H. erectus, however, there is considerable uncertainty as to the accuracy of classifying it as a separate species from H. erectus at all.<a href="#cite_note-142">[140]</a></li></ul>

</li>

<li id="cite_note-174"><a href="#cite_ref-174">^</a> H. erectus in the narrow sense (the Asian species) was extinct by 140,000 years ago, <a href="Homo_erectus_soloensis" class="mw-redirect" title="Homo erectus soloensis">Homo erectus soloensis</a>, found in <a href="Java_(island)" class="mw-redirect" title="Java (island)">Java</a>, is considered the latest known survival of H. erectus. Formerly dated to as late as 50,000 to 40,000 years ago, a 2011 study pushed back the date of its extinction of H. e. soloensis to 143,000 years ago at the latest, more likely before 550,000 years ago.<a href="#cite_note-173">[170]</a>

</li>

<li id="cite_note-228"><a href="#cite_ref-228">^</a> The Latin word which refers to adult

males only is vir

</li>

<li id="cite_note-229"><a href="#cite_ref-229">^</a> See the <a href="Binomial_nomenclature" title="Binomial nomenclature">Binomial nomenclature</a> and <a href="Systema_Naturae" title="Systema Naturae">Systema Naturae</a> articles.

</li>

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

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

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<div style="line-height: 1.6em; font-weight: bold;"><div>Full list of authors</div></div>

<ul class="mw-collapsible-content" style="margin-top: 0; margin-bottom: 0; line-height: inherit;"><li style="line-height: inherit; margin: 0">Clive Finlayson </li><li style="line-height: inherit; margin: 0">Francisco Giles Pacheco </li><li style="line-height: inherit; margin: 0">Joaquín Rodríguez-Vidal </li><li style="line-height: inherit; margin: 0">Darren A. Fa </li><li style="line-height: inherit; margin: 0">José María Gutierrez López </li><li style="line-height: inherit; margin: 0">Antonio Santiago Pérez </li><li style="line-height: inherit; margin: 0">Geraldine Finlayson </li><li style="line-height: inherit; margin: 0">Ethel Allue </li><li style="line-height: inherit; margin: 0">Javier Baena Preysler </li><li style="line-height: inherit; margin: 0">Isabel Cáceres </li><li style="line-height: inherit; margin: 0">José S. Carrión </li><li style="line-height: inherit; margin: 0">Yolanda Fernández-Jalvo </li><li style="line-height: inherit; margin: 0">Christopher P. Gleed-Owen </li><li style="line-height: inherit; margin: 0">Francisco J. Jimenez-Espejo </li><li style="line-height: inherit; margin: 0">Pilar López Martínez </li><li style="line-height: inherit; margin: 0">José Antonio López Sáez </li><li style="line-height: inherit; margin: 0">José Antonio Riquelme Cantal </li><li style="line-height: inherit; margin: 0">Antonio Sánchez Marco </li><li style="line-height: inherit; margin: 0">Francisco Giles Guzman </li><li style="line-height: inherit; margin: 0">Kimberly Brown </li><li style="line-height: inherit; margin: 0">Noemí Fuentes </li><li style="line-height: inherit; margin: 0">Claire A. Valarino </li><li style="line-height: inherit; margin: 0">Antonio Villalpando </li><li style="line-height: inherit; margin: 0">Christopher B. Stringer </li><li style="line-height: inherit; margin: 0">Francisca Martinez Ruiz </li><li style="line-height: inherit; margin: 0">Tatsuhiko Sakamoto</li></ul>

</div>

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<div style="line-height: 1.6em; font-weight: bold;"><div>Full list of authors</div></div>

<ul class="mw-collapsible-content" style="margin-top: 0; margin-bottom: 0; line-height: inherit;"><li style="line-height: inherit; margin: 0">Richard E. Green </li><li style="line-height: inherit; margin: 0">Johannes Krause </li><li style="line-height: inherit; margin: 0">Adrian W. Briggs </li><li style="line-height: inherit; margin: 0">Tomislav Maricic </li><li style="line-height: inherit; margin: 0">Udo Stenzel </li><li style="line-height: inherit; margin: 0">Martin Kircher </li><li style="line-height: inherit; margin: 0">Nick Patterson </li><li style="line-height: inherit; margin: 0">Heng Li </li><li style="line-height: inherit; margin: 0">Weiwei Zhai </li><li style="line-height: inherit; margin: 0">Markus Hsi-Yang Fritz </li><li style="line-height: inherit; margin: 0">Nancy F. Hansen </li><li style="line-height: inherit; margin: 0">Eric Y. Durand </li><li style="line-height: inherit; margin: 0">Anna-Sapfo Malaspinas </li><li style="line-height: inherit; margin: 0">Jeffrey D. Jensen </li><li style="line-height: inherit; margin: 0">Tomas Marques-Bonet </li><li style="line-height: inherit; margin: 0">Can Alkan </li><li style="line-height: inherit; margin: 0">Kay Prüfer </li><li style="line-height: inherit; margin: 0">Matthias Meyer </li><li style="line-height: inherit; margin: 0">Hernán A. Burbano </li><li style="line-height: inherit; margin: 0">Jeffrey M. Good </li><li style="line-height: inherit; margin: 0">Rigo Schultz </li><li style="line-height: inherit; margin: 0">Ayinuer Aximu-Petri </li><li style="line-height: inherit; margin: 0">Anne Butthof </li><li style="line-height: inherit; margin: 0">Barbara Höber </li><li style="line-height: inherit; margin: 0">Barbara Höffner </li><li style="line-height: inherit; margin: 0">Madlen Siegemund </li><li style="line-height: inherit; margin: 0">Antje Weihmann </li><li style="line-height: inherit; margin: 0">Chad Nusbaum </li><li style="line-height: inherit; margin: 0">Eric S. Lander </li><li style="line-height: inherit; margin: 0">Carsten Russ </li><li style="line-height: inherit; margin: 0">Nathaniel Novod </li><li style="line-height: inherit; margin: 0">Jason Affourtit </li><li style="line-height: inherit; margin: 0">Michael Egholm </li><li style="line-height: inherit; margin: 0">Christine Verna </li><li style="line-height: inherit; margin: 0">Pavao Rudan </li><li style="line-height: inherit; margin: 0">Dejana Brajkovic </li><li style="line-height: inherit; margin: 0">Željko Kucan </li><li style="line-height: inherit; margin: 0">Ivan Gušic </li><li style="line-height: inherit; margin: 0">Vladimir B. Doronichev </li><li style="line-height: inherit; margin: 0">Liubov V. Golovanova </li><li style="line-height: inherit; margin: 0">Carles Lalueza-Fox </li><li style="line-height: inherit; margin: 0">Marco de la Rasilla </li><li style="line-height: inherit; margin: 0">Javier Fortea </li><li style="line-height: inherit; margin: 0">Antonio Rosas </li><li style="line-height: inherit; margin: 0">Ralf W. Schmitz </li><li style="line-height: inherit; margin: 0">Philip L. F. Johnson </li><li style="line-height: inherit; margin: 0">Evan E. Eichler </li><li style="line-height: inherit; margin: 0">Daniel Falush </li><li style="line-height: inherit; margin: 0">Ewan Birney </li><li style="line-height: inherit; margin: 0">James C. Mullikin </li><li style="line-height: inherit; margin: 0">Montgomery Slatkin </li><li style="line-height: inherit; margin: 0">Rasmus Nielsen </li><li style="line-height: inherit; margin: 0">Janet Kelso </li><li style="line-height: inherit; margin: 0">Michael Lachmann </li><li style="line-height: inherit; margin: 0">David Reich </li><li style="line-height: inherit; margin: 0"><a href="Svante_P%C3%A4%C3%A4bo" title="Svante Pääbo">Svante Pääbo</a></li></ul>

</div>

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<div style="line-height: 1.6em; font-weight: bold;"><div>Full list of authors</div></div>

<ul class="mw-collapsible-content" style="margin-top: 0; margin-bottom: 0; line-height: inherit;"><li style="line-height: inherit; margin: 0">Adam Brumm </li><li style="line-height: inherit; margin: 0">Gerrit D. van den Bergh </li><li style="line-height: inherit; margin: 0">Michael Storey </li><li style="line-height: inherit; margin: 0">Iwan Kurniawan </li><li style="line-height: inherit; margin: 0">Brent V. Alloway </li><li style="line-height: inherit; margin: 0">Ruly Setiawan </li><li style="line-height: inherit; margin: 0">Erick Setiyabudi </li><li style="line-height: inherit; margin: 0">Rainer Grün </li><li style="line-height: inherit; margin: 0">Mark W. Moore </li><li style="line-height: inherit; margin: 0">Dida Yurnaldi </li><li style="line-height: inherit; margin: 0">Mika R. Puspaningrum </li><li style="line-height: inherit; margin: 0">Unggul P. Wibowo </li><li style="line-height: inherit; margin: 0">Halmi Insani </li><li style="line-height: inherit; margin: 0">Indra Sutisna </li><li style="line-height: inherit; margin: 0">John A. Westgate </li><li style="line-height: inherit; margin: 0">Nick J. G. Pearce </li><li style="line-height: inherit; margin: 0">Mathieu Duval </li><li style="line-height: inherit; margin: 0">Hanneke J. M. Meijer </li><li style="line-height: inherit; margin: 0">Fachroel Aziz </li><li style="line-height: inherit; margin: 0">Thomas Sutikna </li><li style="line-height: inherit; margin: 0">Sander van der Kaars </li><li style="line-height: inherit; margin: 0">Stephanie Flude </li><li style="line-height: inherit; margin: 0">Michael J. Morwood</li></ul>

</div>

</li>

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<li id="cite_note-FOOTNOTEBoydSilk2003[[Category:Wikipedia_articles_needing_page_number_citations_from_February_2015]]<sup_class="noprint_Inline-Template_"_style="white-space:nowrap;">&#91;[[Wikipedia:Citing_sources|<span_title="This_citation_requires_a_reference_to_the_specific_page_or_range_of_pages_in_which_the_material_appears.&#32;(February_2015)">page&nbsp;needed]]&#93;-123"><a href="#cite_ref-FOOTNOTEBoydSilk2003[[Category:Wikipedia_articles_needing_page_number_citations_from_February_2015]]<sup_class="noprint_Inline-Template_"_style="white-space:nowrap;">&#91;[[Wikipedia:Citing_sources|<span_title="This_citation_requires_a_reference_to_the_specific_page_or_range_of_pages_in_which_the_material_appears.&#32;(February_2015)">page&nbsp;needed]]&#93;_123-0">^</a> <a href="#CITEREFBoydSilk2003">Boyd & Silk 2003</a>, p. .

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<div style="line-height: 1.6em; font-weight: bold;"><div>Full list of authors</div></div>

<ul class="mw-collapsible-content" style="margin-top: 0; margin-bottom: 0; line-height: inherit;"><li style="line-height: inherit; margin: 0">Michel Brunet </li><li style="line-height: inherit; margin: 0">Franck Guy </li><li style="line-height: inherit; margin: 0">David Pilbeam </li><li style="line-height: inherit; margin: 0">Hassane Taisso Mackaye </li><li style="line-height: inherit; margin: 0">Andossa Likius </li><li style="line-height: inherit; margin: 0">Djimdoumalbaye Ahounta </li><li style="line-height: inherit; margin: 0">Alain Beauvilain </li><li style="line-height: inherit; margin: 0">Cécile Blondel </li><li style="line-height: inherit; margin: 0">Hervé Bocherens </li><li style="line-height: inherit; margin: 0">Jean-Renaud Boisserie </li><li style="line-height: inherit; margin: 0">Louis De Bonis </li><li style="line-height: inherit; margin: 0">Yves Coppens </li><li style="line-height: inherit; margin: 0">Jean Dejax </li><li style="line-height: inherit; margin: 0">Christiane Denys </li><li style="line-height: inherit; margin: 0">Philippe Duringer </li><li style="line-height: inherit; margin: 0">Véra Eisenmann </li><li style="line-height: inherit; margin: 0">Gongdibé Fanone </li><li style="line-height: inherit; margin: 0">Pierre Fronty </li><li style="line-height: inherit; margin: 0">Denis Geraads </li><li style="line-height: inherit; margin: 0">Thomas Lehmann </li><li style="line-height: inherit; margin: 0">Fabrice Lihoreau </li><li style="line-height: inherit; margin: 0">Antoine Louchart </li><li style="line-height: inherit; margin: 0">Adoum Mahamat </li><li style="line-height: inherit; margin: 0">Gildas Merceron </li><li style="line-height: inherit; margin: 0">Guy Mouchelin </li><li style="line-height: inherit; margin: 0">Olga Otero </li><li style="line-height: inherit; margin: 0">Pablo Pelaez Campomanes </li><li style="line-height: inherit; margin: 0">Marcia Ponce De Leon </li><li style="line-height: inherit; margin: 0">Jean-Claude Rage </li><li style="line-height: inherit; margin: 0">Michel Sapanet </li><li style="line-height: inherit; margin: 0">Mathieu Schuster </li><li style="line-height: inherit; margin: 0">Jean Sudre </li><li style="line-height: inherit; margin: 0">Pascal Tassy </li><li style="line-height: inherit; margin: 0">Xavier Valentin </li><li style="line-height: inherit; margin: 0">Patrick Vignaud </li><li style="line-height: inherit; margin: 0">Laurent Viriot </li><li style="line-height: inherit; margin: 0">Antoine Zazzo </li><li style="line-height: inherit; margin: 0">Christoph Zollikofer</li></ul>

</div>

</li>

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<li id="cite_note-Kwang_Hyun_2015_929–934-127">^ <a href="#cite_ref-Kwang_Hyun_2015_929–934_127-0">a</a> <a href="#cite_ref-Kwang_Hyun_2015_929–934_127-1">b</a> <cite id="CITEREFKwang_Hyun2015" class="citation journal cs1">Kwang Hyun, Ko (2015). "Origins of Bipedalism". Brazilian Archives of Biology and Technology. 58 (6): 929–934. <a href="ArXiv_(identifier)" class="mw-redirect" title="ArXiv (identifier)">arXiv</a>:<a rel="nofollow" class="external text" href="https://arxiv.org/abs/1508.02739">1508.02739</a>. <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/2015arXiv150802739K">2015arXiv150802739K</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1590%2FS1516-89132015060399">10.1590/S1516-89132015060399</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:761213">761213</a>.</cite>

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<li id="cite_note-FOOTNOTEDeSalleTattersall2008[httpsbooksgooglecombooksidBf4Sitw7YaICpgPA146_146]-128">^ <a href="#cite_ref-FOOTNOTEDeSalleTattersall2008[httpsbooksgooglecombooksidBf4Sitw7YaICpgPA146_146]_128-0">a</a> <a href="#cite_ref-FOOTNOTEDeSalleTattersall2008[httpsbooksgooglecombooksidBf4Sitw7YaICpgPA146_146]_128-1">b</a> <a href="#CITEREFDeSalleTattersall2008">DeSalle & Tattersall 2008</a>, p. <a rel="nofollow" class="external text" href="https://books.google.com/books?id=Bf4Sitw7YaIC&pg=PA146">146</a>.

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<li id="cite_note-FOOTNOTECurry2008106–109-129">^ <a href="#cite_ref-FOOTNOTECurry2008106–109_129-0">a</a> <a href="#cite_ref-FOOTNOTECurry2008106–109_129-1">b</a> <a href="#CITEREFCurry2008">Curry 2008</a>, pp. 106–109.

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<li id="cite_note-132"><a href="#cite_ref-132">^</a> <cite id="CITEREFDavid-BarrettDunbar2016" class="citation journal cs1">David-Barrett, T.; Dunbar, R. I. M. (2016). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4874949">"Bipedality and Hair-loss Revisited: The Impact of Altitude and Activity Scheduling"</a>. Journal of Human Evolution. 94: 72–82. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.jhevol.2016.02.006">10.1016/j.jhevol.2016.02.006</a>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4874949">4874949</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/27178459">27178459</a>.</cite>

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<li id="cite_note-FOOTNOTEAielloDean1990-133"><a href="#cite_ref-FOOTNOTEAielloDean1990_133-0">^</a> <a href="#CITEREFAielloDean1990">Aiello & Dean 1990</a>.

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<li id="cite_note-FOOTNOTEKondo1985-134"><a href="#cite_ref-FOOTNOTEKondo1985_134-0">^</a> <a href="#CITEREFKondo1985">Kondo 1985</a>.

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<li id="cite_note-FOOTNOTEStrickberger2000[httpsbooksgooglecombooksidfeMItLo5gwgCpgPA476_475–476]-135"><a href="#cite_ref-FOOTNOTEStrickberger2000[httpsbooksgooglecombooksidfeMItLo5gwgCpgPA476_475–476]_135-0">^</a> <a href="#CITEREFStrickberger2000">Strickberger 2000</a>, pp. <a rel="nofollow" class="external text" href="https://books.google.com/books?id=feMItLo5gwgC&pg=PA476">475–476</a>.

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<li id="cite_note-FOOTNOTETrevathan2011[httpsbooksgooglecombooksidPotlqpl-jxgCpgPA20_20]-136"><a href="#cite_ref-FOOTNOTETrevathan2011[httpsbooksgooglecombooksidPotlqpl-jxgCpgPA20_20]_136-0">^</a> <a href="#CITEREFTrevathan2011">Trevathan 2011</a>, p. <a rel="nofollow" class="external text" href="https://books.google.com/books?id=Potlqpl-jxgC&pg=PA20">20</a>.

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<li id="cite_note-Welker2020-141">^ <a href="#cite_ref-Welker2020_141-0">a</a> <a href="#cite_ref-Welker2020_141-1">b</a> <cite id="CITEREFWelkerRamos-MadrigalGutenbrunner2020" class="citation journal cs1">Welker, F.; Ramos-Madrigal, J.; Gutenbrunner, P.; et al. (April 1, 2020). <a rel="nofollow" class="external text" href="http://eprints.whiterose.ac.uk/159068/1/Welker_etal_Hominin1_AAM.docx">"The dental proteome of Homo antecessor"</a>. <a href="Nature_(journal)" title="Nature (journal)">Nature</a>. 580 (7802): 235–238. <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/2020Natur.580..235W">2020Natur.580..235W</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%2Fs41586-020-2153-8">10.1038/s41586-020-2153-8</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/1476-4687">1476-4687</a>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7582224">7582224</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/32269345">32269345</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:214736611">214736611</a>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20211110200345/https://eprints.whiterose.ac.uk/159068/1/Welker_etal_Hominin1_AAM.docx">Archived</a> from the original on November 10, 2021. Retrieved November 10, 2021.</cite><div class="collapsible-list mw-collapsible mw-collapsed" style="text-align: left;">

<div style="line-height: 1.6em; font-weight: bold;"><div>Full list of authors</div></div>

<ul class="mw-collapsible-content" style="margin-top: 0; margin-bottom: 0; line-height: inherit;"><li style="line-height: inherit; margin: 0">Frido Welker </li><li style="line-height: inherit; margin: 0">Jazmín Ramos-Madrigal </li><li style="line-height: inherit; margin: 0">Petra Gutenbrunner </li><li style="line-height: inherit; margin: 0">Meaghan Mackie </li><li style="line-height: inherit; margin: 0">Shivani Tiwary </li><li style="line-height: inherit; margin: 0">Rosa Rakownikow Jersie-Christensen </li><li style="line-height: inherit; margin: 0">Cristina Chiva </li><li style="line-height: inherit; margin: 0">Marc R. Dickinson </li><li style="line-height: inherit; margin: 0">Martin Kuhlwilm </li><li style="line-height: inherit; margin: 0">Marc de Manuel </li><li style="line-height: inherit; margin: 0">Pere Gelabert </li><li style="line-height: inherit; margin: 0">María Martinón-Torres </li><li style="line-height: inherit; margin: 0">Ann Margvelashvili </li><li style="line-height: inherit; margin: 0">Juan Luis Arsuaga </li><li style="line-height: inherit; margin: 0">Eudald Carbonell </li><li style="line-height: inherit; margin: 0">Tomas Marques-Bonet </li><li style="line-height: inherit; margin: 0">Kirsty Penkman </li><li style="line-height: inherit; margin: 0">Eduard Sabidó </li><li style="line-height: inherit; margin: 0">Jürgen Cox </li><li style="line-height: inherit; margin: 0">Jesper V. Olsen </li><li style="line-height: inherit; margin: 0">David Lordkipanidze </li><li style="line-height: inherit; margin: 0">Fernando Racimo </li><li style="line-height: inherit; margin: 0">Carles Lalueza-Fox </li><li style="line-height: inherit; margin: 0">José María Bermúdez de Castro </li><li style="line-height: inherit; margin: 0">Eske Willerslev </li><li style="line-height: inherit; margin: 0">Enrico Cappellini</li></ul>

</div>

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<li id="cite_note-173"><a href="#cite_ref-173">^</a> <cite id="CITEREFIndriatiSwisherLepreQuinn2011" class="citation journal cs1">Indriati, E.; Swisher, C. C.; Lepre, C.; Quinn, R. L.; Suriyanto, R. A.; et al. (2011). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3126814">"The age of the 20 meter Solo River terrace, Java, Indonesia and the survival of Homo erectus in Asia"</a>. <a href="PLoS_One" class="mw-redirect" title="PLoS One">PLoS One</a>. 6 (6): e21562. <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/2011PLoSO...621562I">2011PLoSO...621562I</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1371%2Fjournal.pone.0021562">10.1371/journal.pone.0021562</a>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3126814">3126814</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/21738710">21738710</a>.</cite><div class="collapsible-list mw-collapsible mw-collapsed" style="text-align: left;">

<div style="line-height: 1.6em; font-weight: bold;"><div>Full list of authors</div></div>

<ul class="mw-collapsible-content" style="margin-top: 0; margin-bottom: 0; line-height: inherit;"><li style="line-height: inherit; margin: 0">Etty Indriati </li><li style="line-height: inherit; margin: 0">Carl C. Swisher III </li><li style="line-height: inherit; margin: 0">Christopher Lepre </li><li style="line-height: inherit; margin: 0">Rhonda L. Quinn </li><li style="line-height: inherit; margin: 0">Rusyad A. Suriyanto </li><li style="line-height: inherit; margin: 0">Agus T. Hascaryo </li><li style="line-height: inherit; margin: 0">Rainer Grün </li><li style="line-height: inherit; margin: 0">Craig S. Feibel </li><li style="line-height: inherit; margin: 0">Briana L. Pobiner </li><li style="line-height: inherit; margin: 0">Maxime Aubert </li><li style="line-height: inherit; margin: 0">Wendy Lees </li><li style="line-height: inherit; margin: 0">Susan C. Antón</li></ul>

</div>

</li>

<li id="cite_note-Zeitoun_2003_148–156-175"><a href="#cite_ref-Zeitoun_2003_148–156_175-0">^</a> <cite id="CITEREFZeitoun2003" class="citation journal cs1">Zeitoun, V. (September 2003). <a rel="nofollow" class="external text" href="https://doi.org/10.1002%2Far.b.10028">"High occurrence of a basicranial feature in Homo erectus: Anatomical description of the preglenoid tubercle"</a>. The Anatomical Record Part B: The New Anatomist. 274 (1): 148–56. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1002%2Far.b.10028">10.1002/ar.b.10028</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/12964205">12964205</a>.</cite>

</li>

<li id="cite_note-Proceedings_2015-176"><a href="#cite_ref-Proceedings_2015_176-0">^</a> <cite id="CITEREFDemboMatzkeMooersCollard2015" class="citation journal cs1">Dembo, M.; Matzke, N. J.; Mooers, A. Ø.; Collard, M. (August 2015). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4528516">"Bayesian analysis of a morphological supermatrix sheds light on controversial fossil hominin relationships"</a>. Proceedings. Biological Sciences. 282 (1812): 20150943. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1098%2Frspb.2015.0943">10.1098/rspb.2015.0943</a>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4528516">4528516</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/26202999">26202999</a>.</cite>

</li>

<li id="cite_note-:2-177"><a href="#cite_ref-:2_177-0">^</a> <cite id="CITEREFDemboRadovčićGarvinLaird2016" class="citation journal cs1">Dembo, M.; Radovčić, D.; et al. (August 2016). "The evolutionary relationships and age of Homo naledi: An assessment using dated Bayesian phylogenetic methods". Journal of Human Evolution. 97: 17–26. <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/2016JHumE..97...17D">2016JHumE..97...17D</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.jhevol.2016.04.008">10.1016/j.jhevol.2016.04.008</a>. <a href="Hdl_(identifier)" class="mw-redirect" title="Hdl (identifier)">hdl</a>:<a rel="nofollow" class="external text" href="https://hdl.handle.net/2164%2F8796">2164/8796</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/27457542">27457542</a>.</cite><div class="collapsible-list mw-collapsible mw-collapsed" style="text-align: left;">

<div style="line-height: 1.6em; font-weight: bold;"><div>Full list of authors</div></div>

<ul class="mw-collapsible-content" style="margin-top: 0; margin-bottom: 0; line-height: inherit;"><li style="line-height: inherit; margin: 0">Mana Dembo </li><li style="line-height: inherit; margin: 0">Davorka Radovčić </li><li style="line-height: inherit; margin: 0">Heather M. Garvinc </li><li style="line-height: inherit; margin: 0">Myra F. Laird </li><li style="line-height: inherit; margin: 0">Lauren Schroeder </li><li style="line-height: inherit; margin: 0">Jill E. Scott </li><li style="line-height: inherit; margin: 0">Juliet Brophy </li><li style="line-height: inherit; margin: 0">Rebecca R. Ackermann </li><li style="line-height: inherit; margin: 0">Chares M. Musiba </li><li style="line-height: inherit; margin: 0">Darryl J. de Ruiter </li><li style="line-height: inherit; margin: 0">Arne Ø. Mooers </li><li style="line-height: inherit; margin: 0">Mark Collard</li></ul>

</div>

</li>

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<div style="line-height: 1.6em; font-weight: bold;"><div>Full list of authors</div></div>

<ul class="mw-collapsible-content" style="margin-top: 0; margin-bottom: 0; line-height: inherit;"><li style="line-height: inherit; margin: 0">Simon Mead </li><li style="line-height: inherit; margin: 0">Jerome Whitfield </li><li style="line-height: inherit; margin: 0">Mark Poulter </li><li style="line-height: inherit; margin: 0">Paresh Shah </li><li style="line-height: inherit; margin: 0">James Uphill </li><li style="line-height: inherit; margin: 0">Tracy Campbell </li><li style="line-height: inherit; margin: 0">Huda Al-Dujaily </li><li style="line-height: inherit; margin: 0">Holger Hummerich </li><li style="line-height: inherit; margin: 0">Jon Beck </li><li style="line-height: inherit; margin: 0">Charles A. Mein </li><li style="line-height: inherit; margin: 0">Claudio Verzilli </li><li style="line-height: inherit; margin: 0">John Whittaker </li><li style="line-height: inherit; margin: 0">Michael P. Alpers </li><li style="line-height: inherit; margin: 0">John Collinge</li></ul>

</div>

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<li id="cite_note-FOOTNOTEDarwin1981-233"><a href="#cite_ref-FOOTNOTEDarwin1981_233-0">^</a> <a href="#CITEREFDarwin1981">Darwin 1981</a>.

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<li id="cite_note-FOOTNOTEMontgomery198895–96-234"><a href="#cite_ref-FOOTNOTEMontgomery198895–96_234-0">^</a> <a href="#CITEREFMontgomery1988">Montgomery 1988</a>, pp. 95–96.

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<li id="cite_note-FOOTNOTEJohansonEdey198120–22,_184–185-237"><a href="#cite_ref-FOOTNOTEJohansonEdey198120–22,_184–185_237-0">^</a> <a href="#CITEREFJohansonEdey1981">Johanson & Edey 1981</a>, pp. 20–22, 184–185.

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<li id="cite_note-FOOTNOTEJohansonEdey198122-239"><a href="#cite_ref-FOOTNOTEJohansonEdey198122_239-0">^</a> <a href="#CITEREFJohansonEdey1981">Johanson & Edey 1981</a>, p. 22.

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<li id="cite_note-240"><a href="#cite_ref-240">^</a> <cite id="CITEREFShreeve2010" class="citation journal cs1">Shreeve, Jamie (July 2010). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20160126024510/http://ngm.nationalgeographic.com/2010/07/middle-awash/shreeve-text">"The Evolutionary Road"</a>. National Geographic. <a href="ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0027-9358">0027-9358</a>. Archived from <a rel="nofollow" class="external text" href="http://ngm.nationalgeographic.com/2010/07/middle-awash/shreeve-text">the original</a> on January 26, 2016. Retrieved May 28, 2015.</cite>

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<li id="cite_note-241"><a href="#cite_ref-241">^</a> <cite id="CITEREFBergerHawksde_RuiterChurchill2015" class="citation journal cs1">Berger, L. R.; Hawks, J.; de Ruiter, D. J.; Churchill, S. E.; et al. (September 10, 2015). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4559886">"Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa"</a>. eLife. 4. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.7554%2FeLife.09560">10.7554/eLife.09560</a>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4559886">4559886</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/26354291">26354291</a>.</cite><div class="collapsible-list mw-collapsible mw-collapsed" style="text-align: left;">

<div style="line-height: 1.6em; font-weight: bold;"><div>Full list of authors</div></div>

<ul class="mw-collapsible-content" style="margin-top: 0; margin-bottom: 0; line-height: inherit;"><li style="line-height: inherit; margin: 0">Lee R Berger </li><li style="line-height: inherit; margin: 0">John Hawks </li><li style="line-height: inherit; margin: 0">Darryl J de Ruiter </li><li style="line-height: inherit; margin: 0">Steven E Churchill </li><li style="line-height: inherit; margin: 0">Peter Schmid </li><li style="line-height: inherit; margin: 0">Lucas K Delezene </li><li style="line-height: inherit; margin: 0">Tracy L Kivell </li><li style="line-height: inherit; margin: 0">Heather M Garvin </li><li style="line-height: inherit; margin: 0">Scott A Williams </li><li style="line-height: inherit; margin: 0">Jeremy M DeSilva </li><li style="line-height: inherit; margin: 0">Matthew M Skinner </li><li style="line-height: inherit; margin: 0">Charles M Musiba </li><li style="line-height: inherit; margin: 0">Noel Cameron </li><li style="line-height: inherit; margin: 0">Trenton W Holliday </li><li style="line-height: inherit; margin: 0">William Harcourt-Smith </li><li style="line-height: inherit; margin: 0">Rebecca R Ackermann </li><li style="line-height: inherit; margin: 0">Markus Bastir </li><li style="line-height: inherit; margin: 0">Barry Bogin </li><li style="line-height: inherit; margin: 0">Debra Bolter </li><li style="line-height: inherit; margin: 0">Juliet Brophy </li><li style="line-height: inherit; margin: 0">Zachary D Cofran </li><li style="line-height: inherit; margin: 0">Kimberly A Congdon </li><li style="line-height: inherit; margin: 0">Andrew S Deane </li><li style="line-height: inherit; margin: 0">Mana Dembo </li><li style="line-height: inherit; margin: 0">Michelle Drapeau </li><li style="line-height: inherit; margin: 0">Marina C Elliott </li><li style="line-height: inherit; margin: 0">Elen M Feuerriegel </li><li style="line-height: inherit; margin: 0">Daniel Garcia-Martinez </li><li style="line-height: inherit; margin: 0">David J Green </li><li style="line-height: inherit; margin: 0">Alia Gurtov </li><li style="line-height: inherit; margin: 0">Joel D Irish </li><li style="line-height: inherit; margin: 0">Ashley Kruger </li><li style="line-height: inherit; margin: 0">Myra F Laird </li><li style="line-height: inherit; margin: 0">Damiano Marchi </li><li style="line-height: inherit; margin: 0">Marc R Meyer </li><li style="line-height: inherit; margin: 0">Shahed Nalla </li><li style="line-height: inherit; margin: 0">Enquye W Negash </li><li style="line-height: inherit; margin: 0">Caley M Orr </li><li style="line-height: inherit; margin: 0">Davorka Radovcic </li><li style="line-height: inherit; margin: 0">Lauren Schroeder </li><li style="line-height: inherit; margin: 0">Jill E Scott </li><li style="line-height: inherit; margin: 0">Zachary Throckmorton </li><li style="line-height: inherit; margin: 0">Matthew W Tocheri </li><li style="line-height: inherit; margin: 0">Caroline VanSickle </li><li style="line-height: inherit; margin: 0">Christopher S Walker </li><li style="line-height: inherit; margin: 0">Pianpian Wei </li><li style="line-height: inherit; margin: 0">Bernhard Zipfel</li></ul>

</div>

</li>

<li id="cite_note-NGS-20150910-242">^ <a href="#cite_ref-NGS-20150910_242-0">a</a> <a href="#cite_ref-NGS-20150910_242-1">b</a> <cite id="CITEREFShreeve2015" class="citation web cs1">Shreeve, Jamie (September 10, 2015). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20180414054102/https://news.nationalgeographic.com/2015/09/150910-human-evolution-change/">"This Face Changes the Human Story. But How?"</a>. <a href="National_Geographic_News" class="mw-redirect" title="National Geographic News">National Geographic News</a>. Archived from <a rel="nofollow" class="external text" href="http://news.nationalgeographic.com/2015/09/150910-human-evolution-change/">the original</a> on April 14, 2018. Retrieved September 10, 2015.</cite>

</li>

<li id="cite_note-243"><a href="#cite_ref-243">^</a> <a href="#CITEREFDirksRobertsHilbert-WolfKramers2017">Dirks et al. (2017)</a>: between 335 and 236 ka. The lower limit of 236 ka is due to optically stimulated luminescence dating of sediments with U-Th and palaeomagnetic analyses of flowstones; the upper limit of 335 ka is due to U-series and electron spin resonance (US-ESR) dating of two H. naledi teeth, to 253+82 −70 ka, for an estimated age of the fossils of 253+82 −17 ka.

</li>

<li id="cite_note-eLIFE-2017a-244"><a href="#cite_ref-eLIFE-2017a_244-0">^</a> <cite id="CITEREFDirksRobertsHilbert-WolfKramers2017" class="citation journal cs1">Dirks PH, Roberts EM, Hilbert-Wolf H, Kramers JD, et al. (May 9, 2017). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5423772">"The age of Homo naledi and associated sediments in the Rising Star Cave, South Africa"</a>. eLife. 6: e24231. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.7554%2FeLife.24231">10.7554/eLife.24231</a>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5423772">5423772</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/28483040">28483040</a>.</cite><div class="collapsible-list mw-collapsible mw-collapsed" style="text-align: left;">

<div style="line-height: 1.6em; font-weight: bold;"><div>Full list of authors</div></div>

<ul class="mw-collapsible-content" style="margin-top: 0; margin-bottom: 0; line-height: inherit;"><li style="line-height: inherit; margin: 0"> Paul H.G.M. Dirks</li><li style="line-height: inherit; margin: 0"> Eric M. Roberts</li><li style="line-height: inherit; margin: 0"> Hannah Hilbert-Wolf</li><li style="line-height: inherit; margin: 0"> Jan D. Kramers</li><li style="line-height: inherit; margin: 0"> John Hawks</li><li style="line-height: inherit; margin: 0"> Anthony Dosseto</li><li style="line-height: inherit; margin: 0"> Mathieu Duval</li><li style="line-height: inherit; margin: 0"> Marina Elliott</li><li style="line-height: inherit; margin: 0"> Mary Evans</li><li style="line-height: inherit; margin: 0"> Rainer Grün</li><li style="line-height: inherit; margin: 0"> John Hellstrom</li><li style="line-height: inherit; margin: 0"> Andy I.R. Herries</li><li style="line-height: inherit; margin: 0"> Renaud Joannes-Boyau</li><li style="line-height: inherit; margin: 0"> Tebogo V. Makhubela</li><li style="line-height: inherit; margin: 0"> Christa J. Placzek</li><li style="line-height: inherit; margin: 0"> Jessie Robbins</li><li style="line-height: inherit; margin: 0"> Carl Spandler</li><li style="line-height: inherit; margin: 0"> Jelle Wiersma</li><li style="line-height: inherit; margin: 0"> Jon Woodhead</li><li style="line-height: inherit; margin: 0"> Lee R. Berger</li></ul>

</div>

</li>

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<li id="cite_note-FOOTNOTETrent2005[httpsbooksgooglecombooksid772BEp8ZyMYCpgPA6_6]-247"><a href="#cite_ref-FOOTNOTETrent2005[httpsbooksgooglecombooksid772BEp8ZyMYCpgPA6_6]_247-0">^</a> <a href="#CITEREFTrent2005">Trent 2005</a>, p. <a rel="nofollow" class="external text" href="https://books.google.com/books?id=772BEp8ZyMYC&pg=PA6">6</a>.

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<div style="line-height: 1.6em; font-weight: bold;"><div>Full list of authors</div></div>

<ul class="mw-collapsible-content" style="margin-top: 0; margin-bottom: 0; line-height: inherit;"><li style="line-height: inherit; margin: 0">Doron M. Behar </li><li style="line-height: inherit; margin: 0">Richard Villems </li><li style="line-height: inherit; margin: 0">Himla Soodyall </li><li style="line-height: inherit; margin: 0">Jason Blue-Smith </li><li style="line-height: inherit; margin: 0">Luisa Pereira </li><li style="line-height: inherit; margin: 0">Ene Metspalu </li><li style="line-height: inherit; margin: 0">Rosaria Scozzari </li><li style="line-height: inherit; margin: 0">Heeran Makkan </li><li style="line-height: inherit; margin: 0">Shay Tzur </li><li style="line-height: inherit; margin: 0">David Comas </li><li style="line-height: inherit; margin: 0">Jaume Bertranpetit </li><li style="line-height: inherit; margin: 0">Lluis Quintana-Murci </li><li style="line-height: inherit; margin: 0">Chris Tyler-Smith </li><li style="line-height: inherit; margin: 0">R. Spencer Wells </li><li style="line-height: inherit; margin: 0">Saharon Rosset </li><li style="line-height: inherit; margin: 0">The Genographic Consortium</li></ul>

</div>

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<div style="line-height: 1.6em; font-weight: bold;"><div>Full list of authors</div></div>

<ul class="mw-collapsible-content" style="margin-top: 0; margin-bottom: 0; line-height: inherit;"><li style="line-height: inherit; margin: 0">Sarah A. Tishkoff </li><li style="line-height: inherit; margin: 0">Floyd A. Reed </li><li style="line-height: inherit; margin: 0">Françoise R. Friedlaender </li><li style="line-height: inherit; margin: 0">Christopher Ehret </li><li style="line-height: inherit; margin: 0">Alessia Ranciaro </li><li style="line-height: inherit; margin: 0">Alain Froment </li><li style="line-height: inherit; margin: 0">Jibril B. Hirbo </li><li style="line-height: inherit; margin: 0">Agnes A. Awomoyi </li><li style="line-height: inherit; margin: 0">Jean-Marie Bodo </li><li style="line-height: inherit; margin: 0">Ogobara Doumbo </li><li style="line-height: inherit; margin: 0">Muntaser Ibrahim </li><li style="line-height: inherit; margin: 0">Abdalla T. Juma </li><li style="line-height: inherit; margin: 0">Maritha J. Kotze </li><li style="line-height: inherit; margin: 0">Godfrey Lema </li><li style="line-height: inherit; margin: 0">Jason H. Moore </li><li style="line-height: inherit; margin: 0">Holly Mortensen </li><li style="line-height: inherit; margin: 0">Thomas B. Nyambo </li><li style="line-height: inherit; margin: 0">Sabah A. Omar </li><li style="line-height: inherit; margin: 0">Kweli Powell </li><li style="line-height: inherit; margin: 0">Gideon S. Pretorius </li><li style="line-height: inherit; margin: 0">Michael W. Smith </li><li style="line-height: inherit; margin: 0">Mahamadou A. Thera </li><li style="line-height: inherit; margin: 0">Charles Wambebe </li><li style="line-height: inherit; margin: 0">James L. Weber </li><li style="line-height: inherit; margin: 0">Scott M. Williams</li></ul>

</div>

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

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<li><cite id="CITEREFDarwin1981" class="citation book cs1"><a href="Charles_Darwin" title="Charles Darwin">Darwin, Charles</a> (1981) [Originally published 1871; London: <a href="John_Murray_(publishing_house)" title="John Murray (publishing house)">John Murray</a>]. <a href="The_Descent_of_Man%2C_and_Selection_in_Relation_to_Sex" title="The Descent of Man, and Selection in Relation to Sex">The Descent of Man, and Selection in Relation to Sex</a>. Introduction by <a href="John_Tyler_Bonner" title="John Tyler Bonner">John Tyler Bonner</a> and <a href="Robert_May%2C_Baron_May_of_Oxford" title="Robert May, Baron May of Oxford">Robert M. May</a> (Reprint ed.). <a href="Princeton_University_Press" title="Princeton University Press">Princeton University Press</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-691-02369-4</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/80008679">80008679</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/7197127">7197127</a>.</cite></li>

<li><cite id="CITEREFDawkins2004" class="citation book cs1"><a href="Richard_Dawkins" title="Richard Dawkins">Dawkins, Richard</a> (2004). <a href="The_Ancestor's_Tale" title="The Ancestor's Tale">The Ancestor's Tale: A Pilgrimage to the Dawn of Evolution</a>. Boston: <a href="Houghton_Mifflin" class="mw-redirect" title="Houghton Mifflin">Houghton Mifflin</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-618-00583-3</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2004059864">2004059864</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/56617123">56617123</a>.</cite></li>

<li><cite id="CITEREFDeSalleTattersall2008" class="citation book cs1">DeSalle, Rob; <a href="Ian_Tattersall" title="Ian Tattersall">Tattersall, Ian</a> (2008). <a rel="nofollow" class="external text" href="https://archive.org/details/humanoriginswhat0000desa">Human Origins: What Bones and Genomes Tell Us About Ourselves</a>. Texas A&M University Anthropology Series. Vol. 13. College Station: <a href="Texas_A%26M_University_Press" title="Texas A&M University Press">Texas A&M University Press</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-1-58544-567-7</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/144520427">144520427</a>.</cite></li>

<li><cite id="CITEREFDiamond1999" class="citation book cs1"><a href="Jared_Diamond" title="Jared Diamond">Diamond, Jared</a> (1999). <a href="Guns%2C_Germs%2C_and_Steel" title="Guns, Germs, and Steel">Guns, Germs, and Steel: The Fates of Human Societies</a>. New York: <a href="W._W._Norton_%26_Co." class="mw-redirect" title="W. W. Norton & Co.">W. W. Norton & Co.</a> <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-393-31755-8</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2005284124">2005284124</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/35792200">35792200</a>.</cite></li>

<li><cite id="CITEREFFreemanHerron2007" class="citation book cs1">Freeman, Scott; Herron, Jon C. (2007). Evolutionary Analysis (4th ed.). Upper Saddle River, New Jersey: <a href="Prentice_Hall" title="Prentice Hall">Pearson Prentice Hall</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-13-227584-2</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2006034384">2006034384</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/73502978">73502978</a>.</cite></li>

<li><cite id="CITEREFJohansonEdey1981" class="citation book cs1"><a href="Donald_C._Johanson" class="mw-redirect" title="Donald C. Johanson">Johanson, Donald</a>; Edey, Maitland (1981). Lucy, the Beginnings of Humankind. St Albans: Granada. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-586-08437-3</bdi>.</cite></li>

<li><cite id="CITEREFMontgomery1988" class="citation book cs1">Montgomery, William M. (1988) [Originally published 1974]. "Germany". In Glick, Thomas F. (ed.). The Comparative Reception of Darwinisms. <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>978-0-226-29977-8</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/87035814">87035814</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/17328115">17328115</a>.</cite> "The Conference on the Comparative Reception of Darwinism was held in Austin, Texas, on April 22 and 23, 1972, under the joint sponsorship of the <a href="American_Council_of_Learned_Societies" title="American Council of Learned Societies">American Council of Learned Societies</a> and the <a href="University_of_Texas_at_Austin" title="University of Texas at Austin">University of Texas at Austin</a>"</li>

<li><cite id="CITEREFKondo1985" class="citation journal cs1">Kondo, Shiro, ed. (1985). "Primate Morphophysiology, Locomotor Analyses, and Human Bipedalism". American Journal of Physical Anthropology. 70 (2). <a href="University_of_Tokyo_Press" title="University of Tokyo Press">University of Tokyo Press</a>: 278–279. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1002%2Fajpa.1330700214">10.1002/ajpa.1330700214</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-4-13-066093-8</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/85173489">85173489</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/12352830">12352830</a>.</cite></li>

<li><cite id="CITEREFLeakey1994" class="citation book cs1"><a href="Richard_Leakey" title="Richard Leakey">Leakey, Richard E.</a> (1994). <a rel="nofollow" class="external text" href="https://archive.org/details/originofhumankin00leak">The Origin of Humankind</a>. Science Masters Series. New York: <a href="Basic_Books" title="Basic Books">Basic Books</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-465-03135-1</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/94003617">94003617</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/30739453">30739453</a>.</cite></li>

<li><cite id="CITEREFM'charek2005" class="citation book cs1">M'charek, Amade (2005). The Human Genome Diversity Project: An Ethnography of Scientific Practice. Cambridge Studies in Society and the Life Sciences. <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>978-0-521-83222-9</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2004052648">2004052648</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/55600894">55600894</a>.</cite></li>

<li><cite id="CITEREFMartin2001" class="citation encyclopaedia cs1"><a href="Robert_D._Martin" title="Robert D. Martin">Martin, Robert D.</a> (2001). "Primates, Evolution of". In <a href="Neil_Smelser" title="Neil Smelser">Smelser, Neil J.</a>; <a href="Paul_Baltes" title="Paul Baltes">Baltes, Paul B.</a> (eds.). <a href="International_Encyclopedia_of_the_Social_%26_Behavioral_Sciences" title="International Encyclopedia of the Social & Behavioral Sciences">International Encyclopedia of the Social & Behavioral Sciences</a>. Amsterdam / New York: <a href="Elsevier" title="Elsevier">Elsevier</a>. pp. 12032–12038. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2FB0-08-043076-7%2F03083-7">10.1016/B0-08-043076-7/03083-7</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-08-043076-8</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2001044791">2001044791</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/47869490">47869490</a>.</cite></li>

<li><cite id="CITEREFMaxwell1984" class="citation book cs1">Maxwell, Mary (1984). Human Evolution: A Philosophical Anthropology. New York: <a href="Columbia_University_Press" title="Columbia University Press">Columbia University Press</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-231-05946-6</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/83024005">83024005</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/10163036">10163036</a>.</cite></li>

<li><cite id="CITEREFMcHenry2009" class="citation book cs1"><a href="Henry_McHenry_(anthropologist)" title="Henry McHenry (anthropologist)">McHenry, Henry M.</a> (2009). <a rel="nofollow" class="external text" href="https://archive.org/details/evolutionfirstfo00mich">"Human Evolution"</a>. In <a href="Michael_Ruse" title="Michael Ruse">Ruse, Michael</a>; <a href="Joseph_Travis" title="Joseph Travis">Travis, Joseph</a> (eds.). Evolution: The First Four Billion Years. Foreword by <a href="Edward_O._Wilson" class="mw-redirect" title="Edward O. Wilson">Edward O. Wilson</a>. Cambridge, Massachusetts: <a href="Harvard_University_Press" title="Harvard University Press">Belknap Press of Harvard University</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-674-03175-3</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2008030270">2008030270</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/225874308">225874308</a>.</cite></li>

<li><cite id="CITEREFRamachandranTangGutenkunstBustamante2010" class="citation book cs1"><a href="Sohini_Ramachandran" title="Sohini Ramachandran">Ramachandran, Sohini</a>; Tang, Hua; Gutenkunst, Ryan N.; <a href="Carlos_D._Bustamante" title="Carlos D. Bustamante">Bustamante, Carlos D.</a> (2010). "Genetics and Genomics of Human Population Structure". In Speicher, Michael R.; <a href="Stylianos_Antonarakis" title="Stylianos Antonarakis">Antonarakis, Stylianos E.</a>; Motulsky, Arno G. (eds.). Vogel and Motulsky's Human Genetics: Problems and Approaches (4th ed.). Heidelberg / London / New York: <a href="Springer_Science%2BBusiness_Media" title="Springer Science+Business Media">Springer</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2F978-3-540-37654-5">10.1007/978-3-540-37654-5</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-3-540-37653-8</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2009931325">2009931325</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/549541244">549541244</a>.</cite></li>

<li><cite id="CITEREFRobinson2008" class="citation book cs1"><a href="John_T._Robinson" class="mw-redirect" title="John T. Robinson">Robinson, J. T.</a> (2008) [Originally published 1963; Chicago, Illinois: Aldine Transaction]. "Adaptive Radiation in the Australopithecines and the Origin of Man". In <a href="Francis_Clark_Howell" title="Francis Clark Howell">Howell, F. Clark</a>; Bourlière, François (eds.). African Ecology and Human Evolution. New Brunswick, New Jersey: <a href="Transaction_Publishers" title="Transaction Publishers">Transaction Publishers</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-202-36136-9</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2007024716">2007024716</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/144770218">144770218</a>.</cite></li>

<li><cite id="CITEREFSrivastava2009" class="citation book cs1">Srivastava, R. P. (2009). Morphology of the Primates and Human Evolution. New Delhi: PHI Learning. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-81-203-3656-8</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/423293609">423293609</a>.</cite></li>

<li><cite id="CITEREFStanfordAllenAntón2009" class="citation book cs1"><a href="Craig_Stanford" title="Craig Stanford">Stanford, Craig</a>; Allen, John S.; Antón, Susan C. (2009). <a rel="nofollow" class="external text" href="https://archive.org/details/biologicalanthro00stan">Biological Anthropology: The Natural History of Humankind</a> (2nd ed.). Upper Saddle River, New Jersey: <a href="Pearson_Prentice_Hall" class="mw-redirect" title="Pearson Prentice Hall">Pearson Prentice Hall</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-13-601160-6</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2007052429">2007052429</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/187548835">187548835</a>.</cite></li>

<li><cite id="CITEREFStrickberger2000" class="citation book cs1">Strickberger, Monroe W. (2000). Evolution (3rd ed.). Sudbury, Massachusetts: <a href="Jones_%26_Bartlett_Publishers" class="mw-redirect" title="Jones & Bartlett Publishers">Jones & Bartlett Publishers</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-7637-1066-8</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/99032072">99032072</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/41431683">41431683</a>.</cite></li>

<li><cite id="CITEREFStringer1994" class="citation book cs1"><a href="Chris_Stringer" title="Chris Stringer">Stringer, Chris B.</a> (1994) [1992]. "Evolution of Early Humans". In <a href="Steve_Jones_(biologist)" title="Steve Jones (biologist)">Jones, Steve</a>; Martin, Robert D.; <a href="David_Pilbeam" title="David Pilbeam">Pilbeam, David</a> (eds.). <a rel="nofollow" class="external text" href="https://archive.org/details/cambridgeencyclo00step">The Cambridge Encyclopedia of Human Evolution</a>. <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>978-0-521-32370-3</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/92018037">92018037</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/444512451">444512451</a>.</cite></li>

<li><cite id="CITEREFSwisherCurtisLewin2001" class="citation book cs1">Swisher, Carl C. III; <a href="Garniss_Curtis" title="Garniss Curtis">Curtis, Garniss H.</a>; <a href="Roger_Lewin" title="Roger Lewin">Lewin, Roger</a> (2001) [Originally published 2000]. Java Man: How Two Geologists Changed Our Understanding of Human Evolution. <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>978-0-226-78734-3</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2001037337">2001037337</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/48066180">48066180</a>.</cite></li>

<li><cite id="CITEREFTrent2005" class="citation book cs1">Trent, Ronald J. (2005). Molecular Medicine: An Introductory Text (3rd ed.). Burlington, Massachusetts: <a href="Elsevier_Academic_Press" class="mw-redirect" title="Elsevier Academic Press">Elsevier Academic Press</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-12-699057-7</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2004028087">2004028087</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/162577235">162577235</a>.</cite></li>

<li><cite id="CITEREFTrevathan2011" class="citation book cs1">Trevathan, Wenda R. (2011) [Originally published 1987; New York: Aldine De Gruyter]. Human Birth: An Evolutionary Perspective. New Brunswick, New Jersey: <a href="Transaction_Publishers" title="Transaction Publishers">Transaction Publishers</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-1-4128-1502-4</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2010038249">2010038249</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/669122326">669122326</a>.</cite></li>

<li><cite id="CITEREFUngarTeaford2002" class="citation book cs1"><a href="Peter_Ungar" title="Peter Ungar">Ungar, Peter S.</a>; Teaford, Mark F., eds. (2002). Human Diet: Its Origin and Evolution. Westport, Connecticut: Bergin & Garvey. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-89789-736-5</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2001043790">2001043790</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/537239907">537239907</a>.</cite></li>

<li><cite id="CITEREFWalker2007" class="citation book cs1"><a href="Alan_Walker_(anthropologist)" title="Alan Walker (anthropologist)">Walker, Alan</a> (2007). "Early Hominin Diets: Overview and Historical Perspectives". In Ungar, Peter (ed.). Evolution of the Human Diet: The Known, the Unknown, and the Unknowable. Human Evolution Series. <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>978-0-19-518346-7</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2005036120">2005036120</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/132816551">132816551</a>.</cite></li>

<li><cite id="CITEREFWallace2004" class="citation book cs1"><a href="David_Rains_Wallace" title="David Rains Wallace">Wallace, David Rains</a> (2004). Beasts of Eden: Walking Whales, Dawn Horses, and Other Enigmas of Mammal Evolution. Berkeley: <a href="University_of_California_Press" title="University of California Press">University of California Press</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-520-24684-3</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2003022857">2003022857</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/53254011">53254011</a>.</cite></li>

<li><cite id="CITEREFWebster2010" class="citation book cs1"><a href="Donovan_Webster" title="Donovan Webster">Webster, Donovan</a> (2010). Meeting the Family: One Man's Journey Through His Human Ancestry. Foreword by <a href="Spencer_Wells" title="Spencer Wells">Spencer Wells</a>. Washington, DC: <a href="National_Geographic_Society" title="National Geographic Society">National Geographic Society</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-1-4262-0573-6</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2009050471">2009050471</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/429022321">429022321</a>.</cite></li>

<li><cite id="CITEREFWood2009" class="citation book cs1">Wood, Bernard A. (2009). "Where Does the Genus Homo Begin, and How Would We Know?". In <a href="Frederick_E._Grine" title="Frederick E. Grine">Grine, Frederick E.</a>; <a href="John_G._Fleagle" title="John G. Fleagle">Fleagle, John G.</a>; Leakey, Richard E. (eds.). <a rel="nofollow" class="external text" href="https://archive.org/details/firsthumansorigi00grin">The First Humans – Origin and Early Evolution of the Genus Homo</a>. Vertebrate Paleobiology and Paleoanthropology. Dordrecht: <a href="Springer_Science%2BBusiness_Media" title="Springer Science+Business Media">Springer Netherlands</a>. pp. <a rel="nofollow" class="external text" href="https://archive.org/details/firsthumansorigi00grin/page/n26">17</a>–28. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2F978-1-4020-9980-9_3">10.1007/978-1-4020-9980-9_3</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-1-4020-9979-3</bdi>. <a href="ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/1877-9077">1877-9077</a>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2009927083">2009927083</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/310400980">310400980</a>.</cite> "Contributions from the Third Stony Brook Human Evolution Symposium and Workshop October 3–7, 2006."</li></ul>

</div>

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

<ul><li><cite id="CITEREFAlexander1990" class="citation book cs1"><a href="Richard_D._Alexander" title="Richard D. Alexander">Alexander, Richard D.</a> (1990). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20160304185713/http://qcpages.qc.edu/Biology/LahtiSites/RDAlexander/Pubs/Alexander90.pdf">How Did Humans Evolve? Reflections on the Uniquely Unique Species</a> (PDF). Special Publication. Ann Arbor: <a href="List_of_museums_and_collections_at_the_University_of_Michigan#Museum_of_Zoology" title="List of museums and collections at the University of Michigan">Museum of Zoology, University of Michigan</a>. pp. 1–38. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/90623893">90623893</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/22860997">22860997</a>. Archived from <a rel="nofollow" class="external text" href="http://qcpages.qc.edu/Biology/LahtiSites/RDAlexander/Pubs/Alexander90.pdf">the original</a> (PDF) on March 4, 2016. Retrieved May 6, 2015.</cite></li>

<li><cite id="CITEREFBartonBriggsEisenGoldstein2007" class="citation book cs1"><a href="Nick_Barton" title="Nick Barton">Barton, Nicholas H.</a>; <a href="Derek_Briggs" title="Derek Briggs">Briggs, Derek E. G.</a>; <a href="Jonathan_Eisen" title="Jonathan Eisen">Eisen, Jonathan A.</a>; et al. (2007). Evolution. <a href="Cold_Spring_Harbor_Laboratory" title="Cold Spring Harbor Laboratory">Cold Spring Harbor Laboratory</a> Press. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-87969-684-9</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2007010767">2007010767</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/86090399">86090399</a>.</cite></li>

<li><cite id="CITEREFBellwood2022" class="citation book cs1"><a href="Peter_Bellwood" title="Peter Bellwood">Bellwood, Peter</a> (2022). The Five-Million-Year Odyssey: The Human Journey from Ape to Agriculture. Princeton: Princeton University Press. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-691-19757-9</bdi>.</cite></li>

<li><cite id="CITEREFEnardPrzeworskiFisherLai2002" class="citation journal cs1">Enard, Wolfgang; Przeworski, Molly; <a href="Simon_Fisher" title="Simon Fisher">Fisher, Simon E.</a>; et al. (August 22, 2002). "Molecular evolution of FOXP2, a gene involved in speech and language". <a href="Nature_(journal)" title="Nature (journal)">Nature</a>. 418 (6900): 869–872. <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/2002Natur.418..869E">2002Natur.418..869E</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%2Fnature01025">10.1038/nature01025</a>. <a href="Hdl_(identifier)" class="mw-redirect" title="Hdl (identifier)">hdl</a>:<a rel="nofollow" class="external text" href="https://hdl.handle.net/11858%2F00-001M-0000-0012-CB89-A">11858/00-001M-0000-0012-CB89-A</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/0028-0836">0028-0836</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/12192408">12192408</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:4416233">4416233</a>.</cite></li>

<li><cite id="CITEREFFlinnGearyWard2005" class="citation journal cs1">Flinn, Mark V.; <a href="David_C._Geary" title="David C. Geary">Geary, David C.</a>; Ward, Carol V. (2005). <a rel="nofollow" class="external text" href="http://web.missouri.edu/~gearyd/Flinnetal2005.pdf">"Ecological dominance, social competition, and coalitionary arms races: Why humans evolved extraordinary intelligence"</a> (PDF). Evolution and Human Behavior. 26 (1): 10–46. <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/2005EHumB..26...10F">2005EHumB..26...10F</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.evolhumbehav.2004.08.005">10.1016/j.evolhumbehav.2004.08.005</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/1090-5138">1090-5138</a>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20070604224126/http://web.missouri.edu/~gearyd/Flinnetal2005.pdf">Archived</a> (PDF) from the original on June 4, 2007. Retrieved May 5, 2015.</cite></li>

<li><cite id="CITEREFGalinon-Melenec2015" class="citation book cs1">Galinon-Melenec, Béatrice (2015). "From "Traces" and "Human Trace" to "Human–Trace Paradigm"". In Parrend, Pierre; Bourgine, Paul; Collet, Pierre (eds.). First Complex systems Digital Campus World E-Conference. Tempe, Arizona: <a href="Springer_Nature" title="Springer Nature">Springer</a>.</cite></li>

<li><cite id="CITEREFGibbons2006" class="citation book cs1">Gibbons, Ann (2006). <a rel="nofollow" class="external text" href="https://archive.org/details/firsthumanraceto00gibb">The First Human: The Race to Discover our Earliest Ancestors</a>. New York: <a href="Doubleday_(publisher)" title="Doubleday (publisher)">Doubleday</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-385-51226-8</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2005053780">2005053780</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/61652817">61652817</a>.</cite></li>

<li><cite id="CITEREFHartwig2002" class="citation book cs1"><a href="Walter_Hartwig" title="Walter Hartwig">Hartwig, Walter C.</a>, ed. (2002). The Primate Fossil Record. Cambridge Studies in Biological and Evolutionary Anthropology. Vol. 33. <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>978-0-521-66315-1</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2001037847">2001037847</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/47254191">47254191</a>.</cite></li>

<li><cite id="CITEREFHeizmannBegun2001" class="citation journal cs1">Heizmann, Elmar P. J.; Begun, David R. (November 2001). <a rel="nofollow" class="external text" href="http://doc.rero.ch/record/13528/files/PAL_E351.pdf">"The oldest Eurasian hominoid"</a> (PDF). Journal of Human Evolution. 41 (5): 463–481. <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/2001JHumE..41..463H">2001JHumE..41..463H</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1006%2Fjhev.2001.0495">10.1006/jhev.2001.0495</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/0047-2484">0047-2484</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/11681862">11681862</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:21230992">21230992</a>.</cite></li>

<li><cite id="CITEREFHillWard1988" class="citation journal cs1">Hill, Andrew; Ward, Steven (1988). <a rel="nofollow" class="external text" href="https://doi.org/10.1002%2Fajpa.1330310505">"Origin of the hominidae: The record of African large hominoid evolution between 14 my and 4 my"</a>. American Journal of Physical Anthropology. 31 (59): 49–83. <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/1988AJPA...31S..49H">1988AJPA...31S..49H</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1002%2Fajpa.1330310505">10.1002/ajpa.1330310505</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/0002-9483">0002-9483</a>.</cite></li>

<li><cite id="CITEREFHoagland1964" class="citation journal cs1">Hoagland, Hudson (1964). "Science and the New Humanism". <a href="Science_(journal)" title="Science (journal)">Science</a>. 143 (3602): 111–114. <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/1964Sci...143..111H">1964Sci...143..111H</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.143.3602.111">10.1126/science.143.3602.111</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/17781189">17781189</a>.</cite></li>

<li><cite id="CITEREFIjdoBaldiniWardReeders1991" class="citation journal cs1">Ijdo, Jacob W.; Baldini, Antonio; Ward, David C.; et al. (October 15, 1991). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC52649">"Origin of human chromosome 2: An ancestral telomere-telomere fusion"</a>. <a href="Proceedings_of_the_National_Academy_of_Sciences_of_the_United_States_of_America" title="Proceedings of the National Academy of Sciences of the United States of America">Proceedings of the National Academy of Sciences of the United States of America</a>. 88 (20): 9051–9055. <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/1991PNAS...88.9051I">1991PNAS...88.9051I</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.88.20.9051">10.1073/pnas.88.20.9051</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/0027-8424">0027-8424</a>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC52649">52649</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/1924367">1924367</a>.</cite> – two ancestral ape chromosomes fused to give rise to human chromosome 2</li>

<li><cite id="CITEREFJohansonWong2010" class="citation book cs1"><a href="Donald_Johanson" title="Donald Johanson">Johanson, Donald</a>; Wong, Kate (2010). Lucy's Legacy: The Quest for Human Origins. New York: <a href="Three_Rivers_Press" title="Three Rivers Press">Three Rivers Press</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-307-39640-2</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2010483830">2010483830</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/419801728">419801728</a>.</cite></li>

<li><cite id="CITEREFJonesMartinPilbeam1994" class="citation book cs1">Jones, Steve; Martin, Robert D.; Pilbeam, David, eds. (1994) [1992]. <a rel="nofollow" class="external text" href="https://archive.org/details/cambridgeencyclo00step">The Cambridge Encyclopedia of Human Evolution</a>. Foreword by Richard Dawkins. <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>978-0-521-32370-3</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/92018037">92018037</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/444512451">444512451</a>.</cite> (This book contains very useful, information-dense chapters on primate evolution in general, and human evolution in particular, including fossil history.)</li>

<li><cite id="CITEREFKawabataKaifu2020" class="citation book cs1">Kawabata, Hiroshi; Kaifu, Yousuke (2020). <a rel="nofollow" class="external text" href="https://www.jpicinternational.com/books/science/b78321546939314b4f3be37eeeba766ac05b6301.html">Lost in Evolution: Exploring Humanity's Path in Asia</a>. Tokyo: Japan Publishing Industry Foundation for Culture. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-4-86658-133-0</bdi>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20221020054826/https://www.jpicinternational.com/books/science/b78321546939314b4f3be37eeeba766ac05b6301.html">Archived</a> from the original on October 20, 2022. Retrieved October 20, 2022.</cite></li>

<li><cite id="CITEREFLeakeyLewin1992" class="citation book cs1">Leakey, Richard E.; Lewin, Roger (1992). Origins Reconsidered: In Search of What Makes us Human. New York: <a href="Doubleday_(publisher)" title="Doubleday (publisher)">Doubleday</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-385-41264-3</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/92006661">92006661</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/25373161">25373161</a>.</cite></li>

<li><cite id="CITEREFLewin1997" class="citation book cs1">Lewin, Roger (1997). Bones of Contention: Controversies in the Search for Human Origins (2nd ed.). <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>978-0-226-47651-3</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/97000972">97000972</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/36181117">36181117</a>.</cite></li>

<li><cite id="CITEREFMorwoodvan_Oosterzee2007" class="citation book cs1">Morwood, Mike; van Oosterzee, Penny (2007). <a rel="nofollow" class="external text" href="https://archive.org/details/newhumanstrange00morw">A New Human: The Startling Discovery and Strange Story of the 'Hobbits' of Flores, Indonesia</a>. New York: <a href="Smithsonian_Institution" title="Smithsonian Institution">Smithsonian</a> Books / <a href="HarperCollins" title="HarperCollins">HarperCollins</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-06-089908-0</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2006052267">2006052267</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/76481584">76481584</a>.</cite></li>

<li><cite id="CITEREFOppenheimer2003" class="citation book cs1"><a href="Stephen_Oppenheimer" title="Stephen Oppenheimer">Oppenheimer, Stephen</a> (2003). Out of Eden: The Peopling of the World. London: <a href="Constable_%26_Robinson" title="Constable & Robinson">Constable & Robinson</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-1-84119-697-8</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2005482222">2005482222</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/52195607">52195607</a>.</cite></li>

<li><cite id="CITEREFOvchinnikovGötherströmRomanovaKharitonov2000" class="citation journal cs1">Ovchinnikov, Igor V.; Götherström, Anders; Romanova, Galina P.; et al. (March 30, 2000). "Molecular analysis of Neanderthal DNA from the Northern Caucasus". <a href="Nature_(journal)" title="Nature (journal)">Nature</a>. 404 (6777): 490–493. <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/2000Natur.404..490O">2000Natur.404..490O</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%2F35006625">10.1038/35006625</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/0028-0836">0028-0836</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/10761915">10761915</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:3101375">3101375</a>.</cite></li>

<li><cite id="CITEREFRoberts2009" class="citation book cs1"><a href="Alice_Roberts" title="Alice Roberts">Roberts, Alice M.</a> (2009). The Incredible Human Journey: The Story of How We Colonised the Planet. London: <a href="Bloomsbury_Publishing" title="Bloomsbury Publishing">Bloomsbury Publishing</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-7475-9839-8</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/310156315">310156315</a>.</cite></li>

<li><cite id="CITEREFShreeve1995" class="citation book cs1">Shreeve, James (1995). <a rel="nofollow" class="external text" href="https://archive.org/details/neandertalenigma00shre_0">The Neandertal Enigma: Solving the Mystery of Modern Human Origins</a>. New York: <a href="William_Morrow_%26_Co." class="mw-redirect" title="William Morrow & Co.">William Morrow & Co.</a> <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-688-09407-2</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/95006337">95006337</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/32088673">32088673</a>.</cite></li>

<li><cite id="CITEREFStringer2011" class="citation book cs1">Stringer, Chris B. (2011). The Origin of Our Species. London: <a href="Penguin_Books" title="Penguin Books">Allen Lane</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-1-84614-140-9</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2011489742">2011489742</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/689522193">689522193</a>.</cite></li>

<li><cite id="CITEREFStringerAndrews2005" class="citation book cs1">Stringer, Chris B.; Andrews, Peter (2005). <a rel="nofollow" class="external text" href="https://archive.org/details/completeworldofh0000stri">The Complete World of Human Evolution</a>. London / New York: <a href="Thames_%26_Hudson" title="Thames & Hudson">Thames & Hudson</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-500-05132-0</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2004110563">2004110563</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/224377190">224377190</a>.</cite></li>

<li><cite id="CITEREFStringerMcKie1997" class="citation book cs1">Stringer, Christopher; McKie, Robin (1997). <a rel="nofollow" class="external text" href="https://archive.org/details/africanexodusori00stri">African Exodus: The Origins of Modern Humanity</a>. New York: <a href="Henry_Holt_%26_Co." class="mw-redirect" title="Henry Holt & Co.">Henry Holt & Co.</a> <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-8050-2759-4</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/96037718">96037718</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/36001167">36001167</a>.</cite></li>

<li><cite id="CITEREFSykes2020" class="citation book cs1">Sykes, Rebecca Wragg (2020). <a href="Kindred%3A_Neanderthal_Life%2C_Love%2C_Death_and_Art" title="Kindred: Neanderthal Life, Love, Death and Art">Kindred: Neanderthal Life, Love, Death and Art</a>. London: <a href="Bloomsbury_Publishing" title="Bloomsbury Publishing">Bloomsbury Sigma</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-1-4729-3749-0</bdi>.</cite></li>

<li><cite id="CITEREFTattersall2008" class="citation book cs1">Tattersall, Ian (2008). The Fossil Trail: How We Know What We Think We Know About Human Evolution (2nd ed.). <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>978-0-19-536766-9</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2008013654">2008013654</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/218188644">218188644</a>.</cite></li>

<li><cite id="CITEREFvan_Oosterzee1999" class="citation book cs1">van Oosterzee, Penny (1999). Dragon Bones: The Story of Peking Man. St Leonards, New South Wales: <a href="Allen_%26_Unwin" title="Allen & Unwin">Allen & Unwin</a> Australia. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-1-86508-123-6</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/00300421">00300421</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/45853997">45853997</a>.</cite></li>

<li><cite id="CITEREFWade2006" class="citation book cs1"><a href="Nicholas_Wade" title="Nicholas Wade">Wade, Nicholas</a> (2006). <a href="Before_the_Dawn_(Wade_book)" title="Before the Dawn (Wade book)">Before the Dawn: Recovering the Lost History of Our Ancestors</a>. New York: <a href="Penguin_Books" title="Penguin Books">Penguin</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-1-59420-079-3</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2005055293">2005055293</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/62282400">62282400</a>.</cite></li>

<li><cite id="CITEREFWalkerShipman1996" class="citation book cs1">Walker, Alan; Shipman, Pat (1996). The Wisdom of the Bones: In Search of Human Origins. 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>978-0-297-81670-6</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/35202130">35202130</a>.</cite></li>

<li><cite id="CITEREFWeissMann1985" class="citation book cs1">Weiss, Mark L.; Mann, Alan E. (1985). Human Biology and Behavior: An Anthropological Perspective (4th ed.). Boston: <a href="Little%2C_Brown_%26_Co." class="mw-redirect" title="Little, Brown & Co.">Little, Brown & Co.</a> <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-316-92894-6</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/85000158">85000158</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/11726796">11726796</a>.</cite> (This book contains very accessible descriptions of human and non-human primates, their evolution, and fossil history.)</li>

<li><cite id="CITEREFWells2003" class="citation book cs1"><a href="Spencer_Wells" title="Spencer Wells">Wells, Spencer</a> (2003) [Originally published 2002; Princeton, NJ: Princeton University Press]. The Journey of Man: A Genetic Odyssey (Random House trade paperback 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>978-0-8129-7146-0</bdi>. <a href="LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/2003066679">2003066679</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/53287806">53287806</a>.</cite></li></ul>

</div>

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

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<div class="side-box-text plainlist">Wikimedia Commons has media related to <a href="https://commons.wikimedia.org/wiki/Category:Human_evolution" class="extiw external" title="commons:Category:Human evolution">Human evolution</a>.</div></div>

</div>

<ul><li>"<a rel="nofollow" class="external text" href="https://www.scientificamerican.com/article/race-evolution-and-the-science-of-human-origins/">Race, Evolution and the Science of Human Origins</a>" by Allison Hopper, <a href="Scientific_American" title="Scientific American">Scientific American</a> (July 5, 2021).</li>

<li><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.bbc.co.uk/sn/prehistoric_life/human/human_evolution/index.shtml">"The evolution of man"</a>. BBC Science & Nature. Retrieved May 6, 2015.</cite></li>

<li><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://www.becominghuman.org/">"Becoming Human"</a>. <a href="Arizona_State_University" title="Arizona State University">Arizona State University</a>'s <a href="Institute_of_Human_Origins" title="Institute of Human Origins">Institute of Human Origins</a>. Retrieved May 6, 2015.</cite></li>

<li><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20150424155530/http://www.hhmi.org/biointeractive/bones-stones-and-genes-origin-modern-humans-0">"Bones, Stones and Genes: The Origin of Modern Humans"</a> (Video lecture series). <a href="Howard_Hughes_Medical_Institute" title="Howard Hughes Medical Institute">Howard Hughes Medical Institute</a>. Archived from <a rel="nofollow" class="external text" href="http://www.hhmi.org/biointeractive/bones-stones-and-genes-origin-modern-humans-0">the original</a> on April 24, 2015. Retrieved May 6, 2015.</cite></li>

<li><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://www.evolution-textbook.org/content/free/figures/ch25.html">"Evolution Figures: Chapter 25"</a>. Cold Spring Harbor Laboratory Press. Retrieved May 6, 2015.</cite> – Illustrations from the book Evolution (2007)</li>

<li><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://humanorigins.si.edu/">"Human Evolution"</a>. <a href="Smithsonian_Institution" title="Smithsonian Institution">Smithsonian Institution</a>'s Human Origins Program. Retrieved June 24, 2013.</cite></li>

<li><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20130618121818/http://archaeologyinfo.com/human-evolution-timeline/">"Human Evolution Timeline"</a>. ArchaeologyInfo.com. Archived from <a rel="nofollow" class="external text" href="http://archaeologyinfo.com/human-evolution-timeline/">the original</a> on June 18, 2013. Retrieved June 24, 2013.</cite></li>

<li><a rel="nofollow" class="external text" href="https://www.canal-u.tv/video/universite_du_havre/conference_mondiale_de_l_unitwin_unesco_sur_les_systemes_complexes_le_paradigme_de_l_homme_trace.20031">"Human Trace" video</a> (2015) <a href="Normandy_University" class="mw-redirect" title="Normandy University">Normandy University</a> UNIHAVRE, <a href="Centre_national_de_la_recherche_scientifique" class="mw-redirect" title="Centre national de la recherche scientifique">CNRS</a>, IDEES, E.Laboratory on Human Trace Unitwin Complex System Digital Campus UNESCO.</li>

<li><cite class="citation audio-visual cs1">Lambert, Tim (Producer) (June 24, 2015). <a rel="nofollow" class="external text" href="https://www.pbs.org/show/first-peoples/">First Peoples</a>. London: <a href="Wall_to_Wall_(production_company)" class="mw-redirect" title="Wall to Wall (production company)">Wall to Wall Television</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/910115743">910115743</a>. Retrieved July 18, 2015.</cite></li>

<li><a rel="nofollow" class="external text" href="http://yalebooks.com/yupbooks/book.asp?isbn=9780300182026">Shaping Humanity Video</a> 2013 <a href="Yale_University" title="Yale University">Yale University</a></li>

<li><a rel="nofollow" class="external text" href="http://humanorigins.si.edu/evidence/human-evolution-timeline-interactive">Human Timeline (Interactive)</a> – <a href="Smithsonian_Institution" title="Smithsonian Institution">Smithsonian</a>, <a href="National_Museum_of_Natural_History" title="National Museum of Natural History">National Museum of Natural History</a> (August 2016).</li>

<li><a rel="nofollow" class="external text" href="https://www.bbc.co.uk/programmes/p003hyfl">Human Evolution</a>, BBC Radio 4 discussion with Steve Jones, Fred Spoor & Margaret Clegg (In Our Time, February 16, 2006)</li>

<li><a rel="nofollow" class="external text" href="https://www.smithsonianmag.com/science-nature/essential-timeline-understanding-evolution-homo-sapiens-180976807/">Evolutionary Timeline of Home Sapiens</a> − <a href="Smithsonian_(magazine)" title="Smithsonian (magazine)">Smithsonian</a> (February 2021)</li>

<li><a rel="nofollow" class="external text" href="https://www.salon.com/2021/08/24/more-americans-believe-in-evolution/">History of Human Evolution in the United States</a> – <a href="Salon.com" title="Salon.com">Salon</a> (August 24, 2021)</li></ul>

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

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

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

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

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

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

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

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

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

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

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

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

<li><a href="Biological_engineering" title="Biological engineering">Biological engineering</a></li>

<li><a href="Mathematical_and_theoretical_biology" title="Mathematical and theoretical biology">Biomathematics</a></li>

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

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

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

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

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

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

<li><a href="Cell_biology" title="Cell biology">Cell biology</a></li>

<li><a href="Cellular_microbiology" title="Cellular microbiology">Cellular microbiology</a></li>

<li><a href="Chemical_biology" title="Chemical biology">Chemical biology</a></li>

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

<li><a href="Cognitive_biology" title="Cognitive biology">Cognitive biology</a></li>

<li><a href="Computational_biology" title="Computational biology">Computational biology</a></li>

<li><a href="Conservation_biology" title="Conservation biology">Conservation biology</a></li>

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

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

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

<li><a href="Developmental_biology" title="Developmental biology">Developmental biology</a></li>

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

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

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

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

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

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

<li><a href="Freshwater_biology" title="Freshwater biology">Freshwater biology</a></li>

<li><a href="Generative_biology" class="mw-redirect" title="Generative biology">Generative biology</a></li>

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

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

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

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

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

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

<li><a href="Human_biology" title="Human biology">Human biology</a></li>

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

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

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

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

<li><a href="Marine_biology" title="Marine biology">Marine biology</a></li>

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

<li><a href="Molecular_biology" title="Molecular biology">Molecular biology</a></li>

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

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

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

<li><a href="Nutritional_science" title="Nutritional science">Nutrition</a></li>

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

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

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

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

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

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

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

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

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

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

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

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

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

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

<li><a href="Quantum_biology" title="Quantum biology">Quantum biology</a></li>

<li><a href="Relational_biology" class="mw-redirect" title="Relational biology">Relational biology</a></li>

<li><a href="Reproductive_biology" title="Reproductive biology">Reproductive biology</a></li>

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

<li><a href="Structural_biology" title="Structural biology">Structural biology</a></li>

<li><a href="Synthetic_biology" title="Synthetic biology">Synthetic biology</a></li>

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

<li><a href="Systems_biology" title="Systems biology">Systems biology</a></li>

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

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

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

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

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

<li><a href="Welfare_biology" title="Welfare biology">Welfare biology</a></li>

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

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

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

<ul><li><a href="History_of_biology" title="History of biology">History of biology</a></li>

<li><a href="Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Nobel Prize in Physiology or Medicine</a></li>

<li><a href="Timeline_of_biology_and_organic_chemistry" title="Timeline of biology and organic chemistry">Timeline of biology and organic chemistry</a></li></ul>

</div></td></tr></tbody></table></div><div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1066933788">

/* start https://en.wikipedia.org/ */

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</style></div><div role="navigation" class="navbox" aria-labelledby="Biology858" 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="3"><div id="Biology858" style="font-size:114%;margin:0 4em"><a href="Biology" title="Biology">Biology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="3"><div>

<ul><li>Introduction (<a href="Introduction_to_genetics" title="Introduction to genetics">Genetics</a>, <a href="Introduction_to_evolution" title="Introduction to evolution">Evolution</a>)</li>

<li><a href="Outline_of_biology" title="Outline of biology">Outline</a></li>

<li><a href="History_of_biology" title="History of biology">History</a>

<ul><li><a href="Timeline_of_biology_and_organic_chemistry" title="Timeline of biology and organic chemistry">Timeline</a></li></ul></li>

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

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Biology" title="Biology">Biology</a></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%">Overview</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="Science" title="Science">Science</a></li>

<li>Properties (<a href="Adaptation" title="Adaptation">Adaptation</a>, <a href="Metabolism" title="Metabolism"> Energy processing</a>, <a href="Developmental_biology" title="Developmental biology">Growth</a>, <a href="Structure#Biological" title="Structure">Order</a>, <a href="Homeostasis" title="Homeostasis">Regulation</a>, <a href="Reproduction" title="Reproduction">Reproduction</a> (<a href="Self-replication" title="Self-replication">Self-replication</a>), <a href="Stimulus_(physiology)" title="Stimulus (physiology)">Response to environment</a>)</li>

<li><a href="Biological_organisation" title="Biological organisation">Hierarchy of life</a> (<a href="Atom" title="Atom">Atom</a> > <a href="Molecule" title="Molecule">Molecule</a> > <a href="Organelle" title="Organelle">Organelle</a> > <a href="Cell_(biology)" title="Cell (biology)">Cell</a> > <a href="Tissue_(biology)" title="Tissue (biology)">Tissue</a> > <a href="Organ_(biology)" title="Organ (biology)">Organ</a> > <a href="Biological_system" title="Biological system">Organ system</a> > <a href="Organism" title="Organism">Organism</a> > <a href="Population" title="Population">Population</a> > <a href="Biocoenosis" title="Biocoenosis">Community</a> > <a href="Ecosystem" title="Ecosystem">Ecosystem</a> > <a href="Biosphere" title="Biosphere">Biosphere</a>)</li>

<li><a href="Reductionism#In_science" title="Reductionism">Reductionistic</a></li>

<li><a href="Emergence" title="Emergence"> Emergent property</a></li>

<li><a href="Mechanical_philosophy" class="mw-redirect" title="Mechanical philosophy">Mechanistic</a></li>

<li><a href="Scientific_method" title="Scientific method">Scientific method</a></li>

<li><a href="Taxonomic_rank" title="Taxonomic rank">Taxonomic rank</a></li>

<li><a href="Scientific_theory" title="Scientific theory"> Theory</a></li>

<li><a href="Peer_review" title="Peer review">Peer review</a></li>

<li><a href="List_of_biology_journals" title="List of biology journals">Biology journals</a></li>

<li><a href="Common_name" title="Common name">Common name</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Chemical basis</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="Atom" title="Atom">Atoms</a></li>

<li><a href="Amino_acid" title="Amino acid">Amino acids</a></li>

<li><a href="Carbohydrate" title="Carbohydrate">Carbohydrates</a></li>

<li><a href="Chemical_bond" title="Chemical bond">Chemical bond</a></li>

<li><a href="Chemical_element" title="Chemical element">Chemical element</a></li>

<li><a href="Lipid" title="Lipid">Lipids</a></li>

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

<ul><li><a href="Quantum_biology" title="Quantum biology">Quantum</a></li></ul></li>

<li><a href="Molecule" title="Molecule">Molecules</a></li>

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

<li><a href="Nucleic_acid" title="Nucleic acid">Nucleic acids</a></li>

<li><a href="Organic_compound" title="Organic compound">Organic compounds</a></li>

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

<li><a href="Protein" title="Protein">Proteins</a></li>

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

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Cell_(biology)" title="Cell (biology)">Cells</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="Cell_cycle" title="Cell cycle">Cell cycle</a></li>

<li><a href="Cell_theory" title="Cell theory">Cell theory</a></li>

<li><a href="Cell_signaling" title="Cell signaling">Cell signaling</a></li>

<li><a href="Cellular_respiration" title="Cellular respiration">Cellular respiration</a></li>

<li><a href="Energy" title="Energy">Energy transformation</a></li>

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

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

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

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

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

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

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

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

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

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

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

<li><a href="Gene_expression" title="Gene expression">Gene expression</a></li>

<li><a href="Regulation_of_gene_expression" title="Regulation of gene expression">Gene regulation</a></li>

<li><a href="Genome" title="Genome">Genomes</a></li>

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

<li><a href="Post-transcriptional_modification" title="Post-transcriptional modification">Post-transcriptional modification</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="Adaptation" title="Adaptation">Adaptation</a></li>

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

<li><a href="Function_(biology)" title="Function (biology)">Function</a></li>

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

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

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

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

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

<li><a href="Natural_selection" title="Natural selection">Natural selection</a></li>

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

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

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

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Biodiversity" title="Biodiversity">Diversity</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="Archaea" title="Archaea">Archaea</a></li>

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

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

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

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

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

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

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

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Plant form and function</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="Epidermis_(botany)" title="Epidermis (botany)">Epidermis (botany)</a></li>

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

<li><a href="Ground_tissue" title="Ground tissue">Ground tissue</a></li>

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

<li><a href="Plant_stem" title="Plant stem">Plant stem</a></li>

<li><a href="Shoot_(botany)" title="Shoot (botany)">Shoot</a></li>

<li><a href="Vascular_plant" title="Vascular plant">Vascular plant</a></li>

<li><a href="Vascular_tissue" title="Vascular tissue">Vascular tissue</a></li>

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

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Animal form and function</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="Breathing" title="Breathing">Breathing</a></li>

<li><a href="Circulatory_system" title="Circulatory system">Circulatory system</a></li>

<li><a href="Endocrine_system" title="Endocrine system">Endocrine system</a></li>

<li><a href="Human_digestive_system" title="Human digestive system">Digestive system</a></li>

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

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

<li><a href="Internal_environment" title="Internal environment">Internal environment</a></li>

<li><a href="Muscular_system" title="Muscular system">Muscular system</a></li>

<li><a href="Nervous_system" title="Nervous system">Nervous system</a></li>

<li><a href="Reproductive_system" title="Reproductive system">Reproductive system</a></li>

<li><a href="Respiratory_system" title="Respiratory system">Respiratory system</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Ecology" title="Ecology">Ecology</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="Biogeochemical_cycle" title="Biogeochemical cycle">Biogeochemical cycle</a></li>

<li><a href="Biological_interaction" title="Biological interaction">Biological interaction</a></li>

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

<li><a href="Biome" title="Biome">Biomes</a></li>

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

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

<li><a href="Climate_change" title="Climate change">Climate change</a></li>

<li><a href="Community_(ecology)" title="Community (ecology)">Community</a></li>

<li><a href="Conservation_biology" title="Conservation biology">Conservation</a></li>

<li><a href="Ecosystem_ecology" title="Ecosystem ecology">Ecosystem</a></li>

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

<ul><li><a href="Ecological_niche" title="Ecological niche">niche</a></li></ul></li>

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

<li><a href="Population_ecology" title="Population ecology">Population dynamics</a></li>

<li><a href="Resource_(biology)" title="Resource (biology)">Resources</a></li></ul>

</div></td></tr></tbody></table><div></div></td><td class="noviewer navbox-image" rowspan="4" style="width:1px;padding:0 0 0 2px"><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="List_of_research_methods_in_biology" title="List of research methods in biology">Research methods</a></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%">Laboratory techniques</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="Genetic_engineering" title="Genetic engineering">Genetic engineering</a></li>

<li><a href="Transformation_(genetics)" class="mw-redirect" title="Transformation (genetics)">Transformation</a></li>

<li><a href="Gel_electrophoresis" title="Gel electrophoresis">Gel electrophoresis</a></li>

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

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

<li><a href="Cell_culture" title="Cell culture">Cell culture</a></li>

<li><a href="DNA_sequencing" title="DNA sequencing">DNA sequencing</a></li>

<li><a href="DNA_microarray" title="DNA microarray">DNA microarray</a></li>

<li><a href="Green_fluorescent_protein" title="Green fluorescent protein">Green fluorescent protein</a></li>

<li><a href="Vector_(molecular_biology)" title="Vector (molecular biology)">vector</a></li>

<li><a href="Enzyme_assay" title="Enzyme assay">Enzyme assay</a></li>

<li><a href="Protein_purification" title="Protein purification">Protein purification</a></li>

<li><a href="Western_blot" title="Western blot">Western blot</a></li>

<li><a href="Northern_blot" title="Northern blot">Northern blot</a></li>

<li><a href="Southern_blot" title="Southern blot">Southern blot</a></li>

<li><a href="Restriction_enzyme" title="Restriction enzyme">Restriction enzyme</a></li>

<li><a href="Polymerase_chain_reaction" title="Polymerase chain reaction">Polymerase chain reaction</a></li>

<li><a href="Two-hybrid_screening" title="Two-hybrid screening">Two-hybrid screening</a></li>

<li><a href="In_vitro" title="In vitro">in vitro</a></li>

<li><a href="In_silico" title="In silico">in silico</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Field techniques</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="Belt_transect" title="Belt transect">Belt transect</a></li>

<li><a href="Mark_and_recapture" title="Mark and recapture">mark and recapture</a></li>

<li><a href="Species_discovery_curve" title="Species discovery curve">species discovery curve</a></li></ul>

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

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

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

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

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

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

<li>Anthropogeny</li>