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'''Human evolution''' is the [[evolution]]ary process within the [[Primate evolution|history]] of [[primate]]s that led to the emergence of ''[[Homo sapiens]]'' as a distinct species of the [[hominid]] family that includes all the [[great ape]]s.<ref>{{Cite web |date=June 8, 2024 |title=Human evolution: History, Stages, Timeline, Tree, Chart, & Facts |url= https://www.britannica.com/science/human-evolution |access-date= |website=Britannica.com}}</ref> This process involved the gradual development of traits such as [[Human skeletal changes due to bipedalism|human bipedalism]], [[dexterity]], and [[complex language]],<ref name="HallHallgrímsson2011c">{{cite book |last1=Hall |first1=Brian K. |author1-link=Brian K. Hall |last2=Hallgrímsson |first2=Benedikt |title=Strickberger's Evolution |url= https://books.google.com/books?id=CrJsNQ5wX8oC&pg=PA488 |date=2011 |publisher=[[Jones & Bartlett Publishers]] |isbn=978-1-4496-6390-2 |page=488}}</ref> as well as interbreeding with other [[hominins]] (a tribe of the [[Homininae|African hominid]] subfamily),<ref name=":3" /> indicating that human evolution was not linear but weblike.<ref name="Ackermann 2015" /><ref name="Denisovans & Neandertals">{{cite web |url= https://www.livinganthropologically.com/biological-anthropology/denisovans-neandertals-human-races/ |title=Denisovans and Neandertals: Rethinking Species Boundaries |website=Living Anthropologically |last=Antrosio |first=Jason |date=August 23, 2018 |access-date=August 25, 2018 |archive-date=August 1, 2020 |archive-url= https://web.archive.org/web/20200801180554/https://www.livinganthropologically.com/biological-anthropology/denisovans-neandertals-human-races/ |url-status=live}}</ref><ref name="Human Hybrids">{{cite magazine |last=Hammer |first=Michael F. |title=Human Hybrids |magazine=[[Scientific American]] |date=May 2013 |url= http://www.grochbiology.org/EarlyHominidInterbreeding.pdf |archive-url= https://web.archive.org/web/20180824034550/http://www.grochbiology.org/EarlyHominidInterbreeding.pdf |archive-date=August 24, 2018}}</ref><ref name="Mosaic humans, the hybrid species">{{cite magazine |last=Yong |first=Ed |title=Mosaic humans, the hybrid species |magazine=[[New Scientist]] |date=July 2011 |volume=211 |issue=2823 |pages=34–38 |doi=10.1016/S0262-4079(11)61839-3 |bibcode=2011NewSc.211...34Y |doi-access=free|issn = 0262-4079 }}</ref> The study of the origins of humans involves [[Interdisciplinary|several]] scientific disciplines, including [[Biological anthropology|physical]] and [[evolutionary anthropology]], [[paleontology]], and [[genetics]]; the field is also known by the terms '''anthropogeny''', '''anthropogenesis''', and '''anthropogony'''.<ref>{{cite journal |last=Heng |first=Henry H. Q. |date=May 2009 |title=The genome-centric concept: Resynthesis of evolutionary theory |journal=[[BioEssays]] |volume=31 |issue=5 |pages=512–525 |doi=10.1002/bies.200800182 |issn=0265-9247 |pmid=19334004 |s2cid=1336952}}</ref><ref name="Marlowe 54–67">{{Cite journal |last=Marlowe |first=Frank W. |date=April 13, 2005 |title=Hunter-gatherers and human evolution |journal=Evolutionary Anthropology: Issues, News, and Reviews |volume=14 |issue=2 |pages=54–67 |doi=10.1002/evan.20046 |s2cid=53489209}}</ref> (The latter two terms are sometimes used to refer to the related subject of [[hominization]].)
Primates diverged from other [[mammal]]s about {{Mya|85}} ([[Myr|mya]]), in the [[Late Cretaceous]] period, with their earliest [[fossil]]s appearing over 55 mya, during the [[Paleocene]].<ref>{{cite web |last=Tyson |first=Peter |date=July 1, 2008 |title=Meet Your Ancestors |work=[[Nova ScienceNow]] |publisher=[[PBS]] / [[WGBH Educational Foundation]] |url= https://www.pbs.org/wgbh/nova/evolution/meet-your-ancestors.html |access-date=April 18, 2015 |archive-date=March 8, 2021 |archive-url= https://web.archive.org/web/20210308195934/https://www.pbs.org/wgbh/nova/evolution/meet-your-ancestors.html |url-status=live}}</ref> Primates produced successive clades leading to the [[ape]] superfamily, which gave rise to the hominid and the [[gibbon]] families; these diverged some 15–20 mya. African and [[Ponginae|Asian hominids]] (including [[orangutan]]s) diverged about 14 mya. [[Hominini|Hominins]] (including the [[Australopithecine]] and [[Panina]] subtribes) parted from the [[Gorillini]] tribe ([[gorilla]]s) between 8 and 9 mya; Australopithecine (including the extinct biped ancestors of humans) separated from the ''Pan'' genus (containing [[chimpanzee]]s and [[bonobo]]s) 4–7 mya.<ref name=":4">{{cite web |url= https://www.science.org/content/article/bonobos-join-chimps-closest-human-relatives-rev2 |title=Bonobos Join Chimps as Closest Human Relatives |website=[[TimeTree]] |access-date=May 19, 2018 |date=June 13, 2012 |last=Gibbons |first=Ann |archive-date=September 13, 2021 |archive-url= https://web.archive.org/web/20210913223232/https://www.science.org/content/article/bonobos-join-chimps-closest-human-relatives-rev2 |url-status=live}}</ref> The ''[[Homo]]'' genus is evidenced by the appearance of ''[[H. habilis]]'' over 2 mya,{{efn|name=habilis}} while [[anatomically modern human]]s emerged in Africa approximately 300,000 years ago.
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The presence of other generalized non-cercopithecids of [[Middle Miocene]] from sites far distant, such as ''[[Otavipithecus]]'' from cave deposits in Namibia, and ''[[Pierolapithecus]]'' and ''[[Dryopithecus]]'' 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 [[Miocene]] hominoids, ''[[Oreopithecus]]'', 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 [[Ponginae]]){{efn|Not to be confused with [[Pongidae]], an obsolete family which grouped together [[orangutan]]s, [[gorilla]]s and [[chimpanzee]]s to separate them from humans}} 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 ''[[Sivapithecus]]'' from India and ''[[Griphopithecus]]'' from Turkey, dated to around 10 mya.{{sfn|Srivastava|2009|p=[https://books.google.com/books?id=kCerOsM8XMwC&pg=PA87 87]}}
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{{Human timeline}}
The earliest documented representative of the genus ''Homo'' is ''[[Homo habilis]]'', which evolved around {{Mya|2.8}},<ref name="autogenerated1">{{cite news |last=Ghosh |first=Pallab |author-link=Pallab Ghosh |date=March 4, 2015 |title='First human' discovered in Ethiopia |url= https://www.bbc.com/news/science-environment-31718336 |work=BBC News |location=London |access-date=April 19, 2015 |archive-date=April 18, 2015 |archive-url= https://web.archive.org/web/20150418032919/http://www.bbc.com/news/science-environment-31718336 |url-status=live}}</ref> 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 [[SRGAP2]] [[gene]] doubled, producing a more rapid wiring of the frontal cortex. During the next million years a process of rapid [[encephalization]] occurred, and with the arrival of ''[[Homo erectus]]'' and ''[[Homo ergaster]]'' in the [[fossil#Dating|fossil record]], cranial capacity had doubled to 850 cm<sup>3</sup>.{{sfn|Swisher|Curtis|Lewin|2001}} (Such an increase in human brain size is equivalent to each generation having 125,000 more [[neuron]]s 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 {{Mya|1.3|1.8}}.
According to the recent African origin theory, modern humans evolved in Africa possibly from ''[[H. heidelbergensis]]'', ''[[H. rhodesiensis]]'' or ''[[H. antecessor]]'' and migrated out of the continent some 50,000 to 100,000 years ago, gradually replacing local populations of ''H. erectus'', [[Denisova hominin]]s, ''[[H. floresiensis]]'', ''[[H. luzonensis]]'' and ''[[H. neanderthalensis]]'', whose ancestors had left Africa in earlier migrations.{{sfn|Stringer|1994|p=242}}{{sfn|McHenry|2009|p=265}}<ref>{{cite journal |title=Out of Africa Revisited |date=May 13, 2005 |journal=[[Science (journal)|Science]] |type=This Week in ''Science'' |volume=308 |issue=5724 |page=921 |doi=10.1126/science.2005.308.5724.twis |doi-access=free |s2cid=220100436 |issn=0036-8075}}</ref><ref>{{cite journal |last=Stringer |first=Chris |author-link=Chris Stringer |date=June 12, 2003 |title=Human evolution: Out of Ethiopia |journal=[[Nature (journal)|Nature]] |volume=423 |issue=6941 |pages=692–695 |doi=10.1038/423692a |issn=0028-0836 |pmid=12802315 |bibcode=2003Natur.423..692S |s2cid=26693109}}</ref><ref>{{cite web |url= http://www.actionbioscience.org/evolution/johanson.html |title=Origins of Modern Humans: Multiregional or Out of Africa? |last=Johanson |first=Donald |author-link=Donald Johanson |date=May 2001 |website=[[actionbioscience]] |publisher=[[American Institute of Biological Sciences]] |location=Washington, DC |access-date=November 23, 2009 |archive-date=November 14, 2010 |archive-url= https://web.archive.org/web/20101114081543/http://www.actionbioscience.org/evolution/johanson.html}}</ref> [[Archaic humans|Archaic ''Homo sapiens'']], the forerunner of [[anatomically modern humans]], evolved in the [[Middle Paleolithic]] between 400,000 and 250,000 years ago.<ref>{{cite press release |last1=Mixon |first1=Bobbie |last2=Ehardt |first2=Carolyn |last3=Hammer |first3=Michael |date=September 6, 2011 |title=Evolution's Past Is Modern Human's Present |url= https://www.nsf.gov/news/news_summ.jsp?org=NSF&cntn_id=121603&preview=false |publisher=[[National Science Foundation]] |id=Press Release 11-181 |access-date=April 20, 2015 |archive-date=December 17, 2014 |archive-url= https://web.archive.org/web/20141217084326/http://www.nsf.gov/news/news_summ.jsp?org=NSF&cntn_id=121603&preview=false |url-status=live}}</ref><ref>{{cite web |url= http://anthro.palomar.edu/homo2/mod_homo_4.htm |title=Early Modern ''Homo sapiens'' |last=O'Neil |first=Dennis |website=Evolution of Modern Humans: A Survey of the Biological and Cultural Evolution of Archaic and Modern Homo sapiens |publisher=[[Palomar College]] |location=San Marcos, California |type=Tutorial |access-date=April 20, 2015 |archive-date=April 30, 2015 |archive-url= https://web.archive.org/web/20150430142627/http://anthro.palomar.edu/homo2/mod_homo_4.htm}}</ref><ref>{{cite journal |title=Fossil Reanalysis Pushes Back Origin of ''Homo sapiens'' |url= http://www.scientificamerican.com/article/fossil-reanalysis-pushes/ |date=February 17, 2005 |journal=[[Scientific American]] |issn=0036-8733 |access-date=April 20, 2015 |archive-date=January 15, 2016 |archive-url= https://web.archive.org/web/20160115000603/http://www.scientificamerican.com/article/fossil-reanalysis-pushes/ |url-status=live}}</ref> Recent [[DNA]] evidence suggests that several [[haplotype]]s of [[Neanderthal]] origin are present among all non-African populations, and Neanderthals and other hominins, such as Denisovans, may have contributed up to 6% of their [[genome]] to present-day humans, suggestive of a [[Interbreeding between archaic and modern humans|limited interbreeding between these species]].<ref name="pmid21179161">{{cite journal |last1=Reich |first1=David |author1-link=David Reich (geneticist) |last2=Green |first2=Richard E. |last3=Kircher |first3=Martin |last4=Krause |first4=Johannes |last5=Patterson |first5=Nick |last6=Durand |first6=Eric Y. |last7=Viola |first7=Bence |last8=Briggs |first8=Adrian W. |last9=Stenzel |first9=Udo |last10=Johnson |first10=Philip L. F. |last11=Maricic |first11=Tomislav |last12=Good |first12=Jeffrey M. |last13=Marques-Bonet |first13=Tomas |last14=Alkan |first14=Can |last15=Fu |first15=Qiaomei |last16=Mallick |first16=Swapan |last17=Li |first17=Heng |last18=Meyer |first18=Matthias |last19=Eichler |first19=Evan E. |last20=Stoneking |first20=Mark |last21=Richards |first21=Michael |last22=Talamo |first22=Sahra |last23=Shunkov |first23=Michael V. |last24=Derevianko |first24=Anatoli P. |last25=Hublin |first25=Jean-Jacques |last26=Kelso |first26=Janet |last27=Slatkin |first27=Montgomery |last28=Pääbo |first28=Svante |author28-link=Svante Pääbo |display-authors=3 |date=December 23, 2010 |title=Genetic history of an archaic hominin group from Denisova Cave in Siberia |journal=[[Nature (journal)|Nature]] |volume=468 |issue=7327 |pages=1053–1060 |bibcode=2010Natur.468.1053R |doi=10.1038/nature09710 |issn=0028-0836 |pmid=21179161 |pmc=4306417 |hdl=10230/25596}}</ref><ref name="pmid20439435">{{cite journal |last=Noonan |first=James P. |date=May 2010 |title=Neanderthal genomics and the evolution of modern humans |journal=[[Genome Research]] |volume=20 |issue=5 |pages=547–553 |doi=10.1101/gr.076000.108 |issn=1088-9051 |pmc=2860157 |pmid=20439435}}</ref><ref name="10.1126/science.1209202">{{cite journal |last1=Abi-Rached |first1=Laurent |last2=Jobin |first2=Matthew J. |last3=Kulkarni |first3=Subhash |last4=McWhinnie |first4=A. |last5=Dalva |first5=K. |last6=Gragert |first6=L. |last7=Babrzadeh |first7=F. |last8=Gharizadeh |first8=B. |last9=Luo |first9=M. |last10=Plummer |first10=F. A. |last11=Kimani |first11=J. |last12=Carrington |first12=M. |last13=Middleton |first13=D. |last14=Rajalingam |first14=R. |last15=Beksac |first15=M. |last16=Marsh |first16=S. G. E. |last17=Maiers |first17=M. |last18=Guethlein |first18=L. A. |last19=Tavoularis |first19=S. |last20=Little |first20=A.-M. |last21=Green |first21=R. E. |last22=Norman |first22=P. J. |last23=Parham |first23=P. |display-authors=3 |date=October 7, 2011 |title=The Shaping of Modern Human Immune Systems by Multiregional Admixture with Archaic Humans |journal=[[Science (journal)|Science]] |volume=334 |issue=6052 |pages=89–94 |bibcode=2011Sci...334...89A |doi=10.1126/science.1209202 |issn=0036-8075 |pmc=3677943 |pmid=21868630}}</ref> According to some anthropologists, the transition to [[behavioral modernity]] with the development of symbolic culture, language, and specialized [[lithic technology]] happened around 50,000 years ago (beginning of the [[Upper Paleolithic]]),
{{cite journal |doi=10.1126/science.aao6266 |pmid=28971970 |volume=358 |title=Southern African ancient genomes estimate modern human divergence to 350,000 to 260,000 years ago |date=2017 |journal=[[Science (journal)|Science]] |pages=652–655 |last1=Schlebusch |first1=C. M. |last2=Malmström |first2=H. |last3=Günther |first3=T. |last4=Sjödin |first4=P. |last5=Coutinho |first5=A. |last6=Edlund |first6=H. |last7=Munters |first7=A. R. |last8=Vicente |first8=M. |last9=Steyn |first9=M. |last10=Soodyall |first10=H. |last11=Lombard |first11=M. |last12=Jakobsson |first12=M. |issue=6363 |bibcode=2017Sci...358..652S |s2cid=206663925 |doi-access=free}}, [https://d2ufo47lrtsv5s.cloudfront.net/content/sci/early/2017/09/27/science.aao6266/F3.large.jpg Fig. 3] {{Webarchive|url= https://web.archive.org/web/20180114130711/https://d2ufo47lrtsv5s.cloudfront.net/content/sci/early/2017/09/27/science.aao6266/F3.large.jpg |date=January 14, 2018}} (''H. sapiens'' divergence times) and
{{cite journal |last=Stringer |first=C. |title=What makes a modern human |journal=[[Nature (journal)|Nature]] |date=2012 |volume=485 |issue=7396 |pages=33–35 |doi=10.1038/485033a |pmid=22552077 |bibcode=2012Natur.485...33S |s2cid=4420496 |doi-access=free}} (archaic admixture).</ref>
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[[Bipedalism]], (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 ''[[Sahelanthropus]]''<ref name="Brunet2002">{{cite journal |last1=Brunet |first1=M. |last2=Guy |first2=F. |last3=Pilbeam |first3=D. |last4=Mackaye |first4=H. |last5=Likius |first5=A. |last6=Ahounta |first6=D. |last7=Beauvilain |first7=A. |last8=Blondel |first8=C. |last9=Bocherens |first9=H. |last10=Boisserie |first10=J. |last11=De Bonis |first11=L. |last12=Coppens |first12=Y. |last13=Dejax |first13=J. |last14=Denys |first14=C. |last15=Duringer |first15=P. |last16=Eisenmann |first16=V. |last17=Fanone |first17=G. |last18=Fronty |first18=P. |last19=Geraads |first19=D. |last20=Lehmann |first20=T. |last21=Lihoreau |first21=F. |last22=Louchart |first22=A. |last23=Mahamat |first23=A. |last24=Merceron |first24=G. |last25=Mouchelin |first25=G. |last26=Otero |first26=O. |last27=Pelaez Campomanes |first27=P. |last28=Ponce De Leon |first28=M. |last29=Rage |first29=J. |last30=Sapanet |first30=M. |last31=Schuster |first31=M. |last32=Sudre |first32=J. |last33=Tassy |first33=P. |last34=Valentin |first34=X. |last35=Vignaud |first35=P. |last36=Viriot |first36=L. |last37=Zazzo |first37=A. |last38=Zollikofer |first38=C. |display-authors=6 |author1-link=Michel Brunet (paleontologist) |author3-link=David Pilbeam |date=July 11, 2002 |title=A new hominid from the Upper Miocene of Chad, Central Africa |journal=[[Nature (journal)|Nature]] |volume=418 |issue=6894 |pages=145–151 |doi=10.1038/nature00879 |issn=0028-0836 |pmid=12110880 |bibcode=2002Natur.418..145B |s2cid=1316969 |url= http://doc.rero.ch/record/13388/files/PAL_E190.pdf |access-date=February 20, 2023 |archive-date=February 25, 2023 |archive-url= https://web.archive.org/web/20230225204437/https://doc.rero.ch/record/13388/files/PAL_E190.pdf |url-status=live}}}{{collapsible list |title=Full list of authors |bullets=true |Michel Brunet |Franck Guy |David Pilbeam |Hassane Taisso Mackaye |Andossa Likius |Djimdoumalbaye Ahounta |Alain Beauvilain |Cécile Blondel |Hervé Bocherens |Jean-Renaud Boisserie |Louis De Bonis |Yves Coppens |Jean Dejax |Christiane Denys |Philippe Duringer |Véra Eisenmann |Gongdibé Fanone |Pierre Fronty |Denis Geraads |Thomas Lehmann |Fabrice Lihoreau |Antoine Louchart |Adoum Mahamat |Gildas Merceron |Guy Mouchelin |Olga Otero |Pablo Pelaez Campomanes |Marcia Ponce De Leon |Jean-Claude Rage |Michel Sapanet |Mathieu Schuster |Jean Sudre |Pascal Tassy |Xavier Valentin |Patrick Vignaud |Laurent Viriot |Antoine Zazzo |Christoph Zollikofer}}</ref> or ''[[Orrorin]]'', both of which arose some 6 to 7 million years ago. The non-bipedal knuckle-walkers, the [[gorilla]]s 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. ''[[Ardipithecus]]'', a full biped, arose approximately 5.6 million years ago.<ref>{{cite journal |title=Ardipithecus ramidus and the Paleobiology of Early Hominids |journal=[[Science (journal)|Science]] |volume=326 |issue=5949 |pages=75–86 |date=2009 |last1=White |first1=T. D. |last2=Asfaw |first2=B. |last3=Beyene |first3=Y. |last4=Haile-Selassie |first4=Y. |last5=Lovejoy |first5=C. O. |last6=Suwa |first6=G. |last7=Woldegabriel |first7=G. |pmid=19810190 |bibcode=2009Sci...326...75W |s2cid=20189444 |doi=10.1126/science.1175802}}</ref>
The early bipeds eventually evolved into the australopithecines and still later into the genus ''[[Homo]]''. 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,<ref name="Kwang Hyun 2015 929–934">{{Cite journal |last=Kwang Hyun |first=Ko |date=2015 |title=Origins of Bipedalism |journal=Brazilian Archives of Biology and Technology |doi=10.1590/S1516-89132015060399 |volume=58 |issue=6 |pages=929–934 |arxiv=1508.02739 |bibcode=2015arXiv150802739K |s2cid=761213}}</ref> 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.<ref name="Kwang Hyun 2015 929–934" />{{sfn|DeSalle|Tattersall|2008|p=[https://books.google.com/books?id=Bf4Sitw7YaIC&pg=PA146 146]}}{{sfn|Curry|2008|pp=106–109}} A 2007 study provides support for the hypothesis that bipedalism evolved because it used less energy than quadrupedal knuckle-walking.<ref>{{cite news |date=July 17, 2007 |title=Study Identifies Energy Efficiency as Reason for Evolution of Upright Walking |work=[[ScienceDaily]] |url= https://www.sciencedaily.com/releases/2007/07/070716191140.htm |access-date=April 9, 2015 |archive-date=May 4, 2015 |archive-url= https://web.archive.org/web/20150504174649/http://www.sciencedaily.com/releases/2007/07/070716191140.htm |url-status=live}}<br />{{cite web |url= http://uanews.org/story/study-identifies-energy-efficiency-reason-evolution-upright-walking |title=Study identifies energy efficiency as reason for evolution of upright walking |date=July 16, 2007 |website=UANews |publisher=[[University of Arizona]] |location=Tucson |access-date=April 23, 2015 |archive-date=July 3, 2022 |archive-url= https://web.archive.org/web/20220703195556/http://uanews.org/story/study-identifies-energy-efficiency-reason-evolution-upright-walking |url-status=
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 [[Human vertebral column|vertebral column]], feet and ankles, and skull.{{sfn|Aiello|Dean|1990}} The [[femur]] 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 [[lumbar vertebrae]] 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 [[foramen magnum]] migrated under the skull and more anterior.{{sfn|Kondo|1985}}
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* [[Archaeogenetics]]
* [[Dual inheritance theory]]
* [[Evolution of human intelligence]]
* [[Evolution of morality]]
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