Human

== Evolution ==
{{Main|Human evolution}}
Humans are apes ([[Hominoidea|superfamily Hominoidea]]).<ref>{{Cite book|vauthors=Tuttle RH |title=International Encyclopedia of Biological Anthropology |date=4 October 2018 |publisher=[[John Wiley & Sons, Inc.]] |isbn=978-1-118-58442-2 |veditors=Trevathan W, Cartmill M, Dufour D, Larsen C |place=[[Hoboken]], [[New Jersey]], [[United States]]|pages=1–2|language=en|chapter=Hominoidea: conceptual history|doi=10.1002/9781118584538.ieba0246|s2cid=240125199|author-link=Russell Tuttle|access-date=26 May 2021|chapter-url=https://doi.wiley.com/10.1002/9781118584538.ieba0246}}</ref> The [[Lineage (evolution)|lineage]] of apes that eventually gave rise to humans first split from [[gibbon]]s (family Hylobatidae) and [[orangutan]]s (genus ''Pongo''), then [[gorilla]]s (genus ''Gorilla''), and finally, [[chimpanzee]]s and [[bonobo]]s (genus ''[[Pan (genus)|Pan]]''). The last split, between the human and chimpanzee–bonobo lineages, took place around 8–4 million years ago, in the late [[Miocene]] epoch.<ref>{{cite journal |vauthors=Goodman M, Tagle DA, Fitch DH, Bailey W, Czelusniak J, Koop BF, Benson P, Slightom JL |display-authors=6 |title=Primate evolution at the DNA level and a classification of hominoids |journal=[[Journal of Molecular Evolution]] |volume=30 |issue=3 |pages=260–266 |date=March 1990 |pmid=2109087 |doi=10.1007/BF02099995 |s2cid=2112935 |bibcode=1990JMolE..30..260G}}</ref><ref name="Ruvolo1997">{{cite journal|vauthors=Ruvolo M |date=March 1997 |title=Molecular phylogeny of the hominoids: inferences from multiple independent DNA sequence data sets |journal=[[Molecular Biology and Evolution]] |volume=14 |issue=3 |pages=248–265|doi=10.1093/oxfordjournals.molbev.a025761|pmid=9066793|doi-access=free}}</ref> During this split, [[chromosome 2]] was formed from the joining of two other chromosomes, leaving humans with only 23 pairs of chromosomes, compared to 24 for the other apes.<ref name="fusion">{{cite web|title=Human Chromosome 2 is a fusion of two ancestral chromosomes |url=https://www.evolutionpages.com/chromosome_2.htm |archive-url=https://web.archive.org/web/20110809040210/https://www.evolutionpages.com/chromosome_2.htm |archive-date=9 August 2011 |access-date=18 May 2006 |work=Evolution pages |vauthors=MacAndrew A}}</ref> Following their split with chimpanzees and bonobos, the [[Hominini|hominins]] diversified into many species and at least two distinct genera. All but one of these lineages{{snd}}representing the genus ''[[Homo]]'' and its sole extant species ''Homo sapiens''{{snd}}are now extinct.<ref>{{Cite web |last=McNulty |first=Kieran P. |year=2016 |title=Hominin Taxonomy and Phylogeny: What's In A Name? |url=https://www.nature.com/scitable/knowledge/library/hominin-taxonomy-and-phylogeny-what-s-in-142102877/ |access-date=11 June 2022 |website=Nature Education Knowledge |language=en |archive-date=10 January 2016 |archive-url=https://web.archive.org/web/20160110013134/https://www.nature.com/scitable/knowledge/library/hominin-taxonomy-and-phylogeny-what-s-in-142102877/ |url-status=live}}</ref>

[[File:Lucy Skeleton.jpg|thumb|left|upright|Reconstruction of [[Lucy (Australopithecus)|Lucy]]'','' the first ''[[Australopithecus afarensis]]'' skeleton found]]
The genus ''Homo'' evolved from ''[[Australopithecus]]''.<ref>{{cite journal |vauthors=Strait DS |title=The Evolutionary History of the Australopiths |journal=Evolution: Education and Outreach |date=September 2010 |volume=3 |issue=3 |pages=341–352 |doi=10.1007/s12052-010-0249-6 |s2cid=31979188 |language=en |issn=1936-6434 |doi-access=free }}</ref><ref>{{cite journal |vauthors=Dunsworth HM |title=Origin of the Genus Homo |journal=Evolution: Education and Outreach |date=September 2010 |volume=3 |issue=3 |pages=353–366 |doi=10.1007/s12052-010-0247-8 |s2cid=43116946 |language=en |issn=1936-6434 |doi-access=free }}</ref> Though [[Human fossils|fossils]] from the transition are scarce, the earliest members of ''Homo'' share several key traits with ''Australopithecus''.<ref>{{cite journal |vauthors=Kimbel WH, Villmoare B |title=From Australopithecus to Homo: the transition that wasn't |journal=Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences |volume =371 |issue=1698 |page=20150248 |date=July 2016 |pmid=27298460 |pmc=4920303 |doi=10.1098/rstb.2015.0248 |s2cid=20267830}}</ref><ref name=Villmoare2015>{{cite journal |vauthors=Villmoare B, Kimbel WH, Seyoum C, Campisano CJ, DiMaggio EN, Rowan J, Braun DR, Arrowsmith JR, Reed KE |display-authors=6 |title=Paleoanthropology. Early Homo at 2.8 Ma from Ledi-Geraru, Afar, Ethiopia |journal=[[Science (journal)|Science]] |volume=347 |issue=6228 |pages=1352–1355 |date=March 2015 |pmid=25739410 |doi=10.1126/science.aaa1343 |doi-access=free |bibcode=2015Sci...347.1352V}}</ref> The earliest record of ''Homo'' is the 2.8 million-year-old specimen [[LD 350-1]] from [[Ethiopia]], and the earliest named species are ''[[Homo habilis]]'' and ''[[Homo rudolfensis]]'' which evolved by 2.3 million years ago.<ref name=Villmoare2015 /> ''[[Homo erectus|H. erectus]]'' (the African variant is sometimes called ''[[Homo ergaster|H. ergaster]]'') evolved 2 million years ago and was the first [[archaic human]] species to leave Africa and disperse across Eurasia.<ref>{{cite journal |vauthors=Zhu Z, Dennell R, Huang W, Wu Y, Qiu S, Yang S, Rao Z, Hou Y, Xie J, Han J, Ouyang T |display-authors=6 |title=Hominin occupation of the Chinese Loess Plateau since about 2.1 million years ago |journal=[[Nature (journal)|Nature]] |volume=559 |issue=7715 |pages=608–612 |date=July 2018 |pmid=29995848 | doi=10.1038/s41586-018-0299-4 |bibcode=2018Natur.559..608Z |s2cid =49670311}}</ref> ''H. erectus'' also was the first to evolve a characteristically human [[body plan]]. ''Homo sapiens'' emerged in Africa around 300,000 years ago from a species commonly designated as either ''[[Homo heidelbergensis|H. heidelbergensis]]'' or ''[[Homo rhodesiensis|H. rhodesiensis]]'', the descendants of ''H. erectus'' that remained in Africa.<ref>{{cite journal | vauthors=Hublin JJ, Ben-Ncer A, Bailey SE, Freidline SE, Neubauer S, Skinner MM, Bergmann I, Le Cabec A, Benazzi S, Harvati K, Gunz P |display-authors=6 |title=New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens |journal=[[Nature (journal)|Nature]] |volume=546 |issue=7657 |pages=289–292 |date=June 2017 |pmid=28593953 |doi=10.1038/nature22336 |bibcode=2017Natur.546..289H |s2cid=256771372 |url=https://kar.kent.ac.uk/62267/1/Submission_288356_1_art_file_2637492_j96j1b.pdf |access-date=30 July 2022 |archive-date=8 January 2020 |archive-url=https://web.archive.org/web/20200108234003/https://kar.kent.ac.uk/62267/1/Submission_288356_1_art_file_2637492_j96j1b.pdf |url-status=live}}</ref> ''H. sapiens'' migrated out of the continent, gradually replacing or interbreeding with local populations of archaic humans.<ref>{{cite journal |author=<!--Staff writer(s); no by-line.--> |date=13 May 2005 |title=Out of Africa Revisited |journal=[[Science (journal)|Science]] |type=This Week in ''Science'' |volume=308 |issue=5724 |page=921 |doi=10.1126/science.308.5724.921g |issn=0036-8075 |s2cid=220100436}}</ref><ref>{{cite journal |vauthors=Stringer C |title=Human evolution: Out of Ethiopia |journal=[[Nature (journal)|Nature]] |volume=423 |issue=6941 |pages=692–693, 695 |date=June 2003 |pmid=12802315 |doi=10.1038/423692a |s2cid=26693109 |author-link=Chris Stringer |bibcode=2003Natur.423..692S}}</ref><ref>{{cite web |vauthors=Johanson D |author-link=Donald Johanson |date=May 2001 |title=Origins of Modern Humans: Multiregional or Out of Africa? |url=https://www.actionbioscience.org/evolution/johanson.html |access-date=23 November 2009 |website=[[actionbioscience]] |publisher=[[American Institute of Biological Sciences]] |location=Washington, DC |archive-date=17 June 2021 |archive-url=https://web.archive.org/web/20210617010349/http://www.actionbioscience.org/evolution/johanson.html |url-status=dead}}</ref> Humans began exhibiting [[behavioral modernity]] about 160,000–70,000 years ago,<ref name="Marean et al 2007">{{cite journal |last1=Marean |first1=Curtis |display-authors=etal |date=2007 |title=Early human use of marine resources and pigment in South Africa during the Middle Pleistocene |journal=[[Nature (journal)|Nature]] |volume=449 |issue=7164 |pages=905–908 |bibcode=2007Natur.449..905M |doi=10.1038/nature06204 |pmid=17943129 |s2cid=4387442|url=http://doc.rero.ch/record/15550/files/PAL_E2962.pdf }}</ref> and possibly earlier.<ref name="Brooks">{{Cite journal |vauthors=Brooks AS, Yellen JE, Potts R, Behrensmeyer AK, Deino AL, Leslie DE, Ambrose SH, Ferguson JR, d'Errico F, Zipkin AM, Whittaker S, Post J, Veatch EG, Foecke K, Clark JB |year=2018 |title=Long-distance stone transport and pigment use in the earliest Middle Stone Age |journal=[[Science (journal)|Science]] |volume=360 |issue=6384 |pages=90–94 |bibcode=2018Sci...360...90B |doi=10.1126/science.aao2646 |pmid=29545508 |doi-access=free}}</ref>

The [[Recent African origin of modern humans|"out of Africa" migration]] took place in at least two waves, the first around 130,000 to 100,000 years ago, the second ([[Southern Dispersal]]) around 70,000 to 50,000 years ago.<ref name="Posth">{{cite journal |vauthors=Posth C, Renaud G, Mittnik A, Drucker DG, Rougier H, Cupillard C, Valentin F, Thevenet C, Furtwängler A, Wißing C, Francken M, Malina M, Bolus M, Lari M, Gigli E, Capecchi G, Crevecoeur I, Beauval C, Flas D, Germonpré M, van der Plicht J, Cottiaux R, Gély B, Ronchitelli A, Wehrberger K, Grigorescu D, Svoboda J, Semal P, Caramelli D, Bocherens H, Harvati K, Conard NJ, Haak W, Powell A, Krause J |display-authors=6 |title=Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-Africans and a Late Glacial Population Turnover in Europe |journal=[[Current Biology]] |volume=26 |issue=6 |pages=827–833 |date=March 2016 |pmid=26853362 |doi=10.1016/j.cub.2016.01.037 |bibcode=2016CBio...26..827P |hdl-access=free |s2cid=140098861 |hdl=2440/114930}}</ref><ref>{{cite journal |vauthors=Karmin M, Saag L, Vicente M, Wilson Sayres MA, Järve M, Talas UG, Rootsi S, Ilumäe AM, Mägi R, Mitt M, Pagani L, Puurand T, Faltyskova Z, Clemente F, Cardona A, Metspalu E, Sahakyan H, Yunusbayev B, Hudjashov G, DeGiorgio M, Loogväli EL, Eichstaedt C, Eelmets M, Chaubey G, Tambets K, Litvinov S, Mormina M, Xue Y, Ayub Q, Zoraqi G, Korneliussen TS, Akhatova F, Lachance J, Tishkoff S, Momynaliev K, Ricaut FX, Kusuma P, Razafindrazaka H, Pierron D, Cox MP, Sultana GN, Willerslev R, Muller C, Westaway M, Lambert D, Skaro V, Kovačevic L, Turdikulova S, Dalimova D, Khusainova R, Trofimova N, Akhmetova V, Khidiyatova I, Lichman DV, Isakova J, Pocheshkhova E, Sabitov Z, Barashkov NA, Nymadawa P, Mihailov E, Seng JW, Evseeva I, Migliano AB, Abdullah S, Andriadze G, Primorac D, Atramentova L, Utevska O, Yepiskoposyan L, Marjanovic D, Kushniarevich A, Behar DM, Gilissen C, Vissers L, Veltman JA, Balanovska E, Derenko M, Malyarchuk B, Metspalu A, Fedorova S, Eriksson A, Manica A, Mendez FL, Karafet TM, Veeramah KR, Bradman N, Hammer MF, Osipova LP, Balanovsky O, Khusnutdinova EK, Johnsen K, Remm M, Thomas MG, Tyler-Smith C, Underhill PA, Willerslev E, Nielsen R, Metspalu M, Villems R, Kivisild T |display-authors=6 |title=A recent bottleneck of Y chromosome diversity coincides with a global change in culture |journal=[[Genome Research]] |volume=25 |issue=4 |pages=459–466 |date=April 2015 |pmid=25770088 |pmc=4381518 |doi=10.1101/gr.186684.114}}</ref> ''H. sapiens'' proceeded to colonize all the continents and larger islands, arriving in [[Eurasia]] 125,000 years ago,<ref>{{cite journal |vauthors=Armitage SJ, Jasim SA, Marks AE, Parker AG, Usik VI, Uerpmann HP |title=The southern route "out of Africa": evidence for an early expansion of modern humans into Arabia |journal=[[Science (journal)|Science]] |volume=331 |issue=6016 |pages=453–456 |date=January 2011 |pmid=21273486 |doi=10.1126/science.1199113 |url=https://www.sciencenews.org/view/generic/id/69197/title/Hints_of_earlier_human_exit_from_Africa |url-status=live |access-date=1 May 2011 |bibcode=2011Sci...331..453A |s2cid=20296624 |archive-url=https://web.archive.org/web/20110427201317/https://www.sciencenews.org/view/generic/id/69197/title/Hints_of_earlier_human_exit_from_Africa |archive-date=27 April 2011}}</ref><ref>{{cite web |vauthors=Rincon P |url=https://www.bbc.co.uk/news/science-environment-12300228 |title=Humans 'left Africa much earlier' |archive-url=https://web.archive.org/web/20120809051349/https://www.bbc.co.uk/news/science-environment-12300228 |archive-date=9 August 2012 |work=[[BBC News]] |date=27 January 2011}}</ref> Australia around 65,000 years ago,<ref>{{cite journal |vauthors=Clarkson C, Jacobs Z, Marwick B, Fullagar R, Wallis L, Smith M, Roberts RG, Hayes E, Lowe K, Carah X, Florin SA, McNeil J, Cox D, Arnold LJ, Hua Q, Huntley J, Brand HE, Manne T, Fairbairn A, Shulmeister J, Lyle L, Salinas M, Page M, Connell K, Park G, Norman K, Murphy T, Pardoe C |display-authors=6 | title=Human occupation of northern Australia by 65,000 years ago |journal=[[Nature (journal)|Nature]] |volume=547 |issue=7663 |pages=306–310 |date=July 2017 |pmid=28726833 |doi=10.1038/nature22968 |bibcode=2017Natur.547..306C |s2cid=205257212 |hdl=2440/107043|hdl-access=free }}</ref> the Americas around 15,000 years ago, and remote islands such as [[Hawaiian Islands|Hawaii]], [[Easter Island]], [[Madagascar]], and [[New Zealand]] in the years 300 to 1280 CE.<ref name="Lowe">{{cite web|vauthors=Lowe DJ |year=2008 |title=Polynesian settlement of New Zealand and the impacts of volcanism on early Maori society: an update |url=https://researchcommons.waikato.ac.nz/bitstream/10289/2690/1/Lowe%202008%20Polynesian%20settlement%20guidebook.pdf |url-status=live |archive-url=https://web.archive.org/web/20100522032853/https://researchcommons.waikato.ac.nz/bitstream/10289/2690/1/Lowe%202008%20Polynesian%20settlement%20guidebook.pdf |archive-date=22 May 2010 |access-date=29 April 2010 |publisher=[[University of Waikato]]}}</ref><ref>{{cite journal |vauthors=Appenzeller T |title=Human migrations: Eastern odyssey |journal=[[Nature (journal)|Nature]] |volume=485 |issue=7396 |pages=24–26 |date=May 2012 |pmid=22552074 |doi=10.1038/485024a |bibcode=2012Natur.485...24A |doi-access=free}}</ref>

Human evolution was not a simple linear or branched progression but involved [[Interbreeding between archaic and modern humans|interbreeding between related species]].<ref name="pmid21179161" /><ref>{{cite journal |url=https://www.grochbiology.org/EarlyHominidInterbreeding.pdf |title=Human Hybrids |archive-url=https://web.archive.org/web/20180824034550/https://www.grochbiology.org/EarlyHominidInterbreeding.pdf |archive-date=24 August 2018 |vauthors=Hammer MF |journal=[[Scientific American]] |date=May 2013 |volume=308 |issue=5 |pages=66–71 |doi=10.1038/scientificamerican0513-66 |pmid=23627222 |bibcode=2013SciAm.308e..66H}}</ref><ref>{{cite journal |vauthors=Yong E |date=July 2011 |title=Mosaic humans, the hybrid species |journal=[[New Scientist]] |volume=211 |issue=2823 |pages=34–38 |bibcode=2011NewSc.211...34Y |doi=10.1016/S0262-4079(11)61839-3|doi-access=free }}</ref> Genomic research has shown that hybridization between substantially diverged lineages was common in human evolution.<ref name="Ackermann 2015">{{cite journal |vauthors=Ackermann RR, Mackay A, Arnold ML |date=October 2015 |title=The Hybrid Origin of "Modern" Humans |journal=[[Evolutionary Biology (journal)|Evolutionary Biology]] |volume=43 |issue=1 |pages=1–11 |doi=10.1007/s11692-015-9348-1 |s2cid=14329491}}</ref> [[DNA]] evidence suggests that several genes of [[Neanderthal]] origin are present among all non sub-Saharan-African populations, and Neanderthals and other hominins, such as [[Denisovan]]s, may have contributed up to 6% of their [[genome]] to present-day non sub-Saharan-African humans.<ref name="pmid21179161">{{cite journal |vauthors=Reich D, Green RE, Kircher M, Krause J, Patterson N, Durand EY, Viola B, Briggs AW, Stenzel U, Johnson PL, Maricic T, Good JM, Marques-Bonet T, Alkan C, Fu Q, Mallick S, Li H, Meyer M, Eichler EE, Stoneking M, Richards M, Talamo S, Shunkov MV, Derevianko AP, Hublin JJ, Kelso J, Slatkin M, Pääbo S |display-authors=6 |title=Genetic history of an archaic hominin group from Denisova Cave in Siberia |journal=[[Nature (journal)|Nature]] |volume=468 |issue=7327 |pages=1053–1060 |date=December 2010 |pmid=21179161 |pmc=4306417 |doi=10.1038/nature09710 |bibcode=2010Natur.468.1053R |hdl=10230/25596 |author-link1=David Reich (geneticist) }}</ref><ref name="pmid20439435">{{cite journal |vauthors=Noonan JP |title=Neanderthal genomics and the evolution of modern humans |journal=[[Genome Research]] |volume=20 |issue=5 |pages=547–553 |date=May 2010 |pmid=20439435 |pmc=2860157 |doi=10.1101/gr.076000.108}}</ref><ref name="10.1126/science.1209202">{{cite journal |vauthors=Abi-Rached L, Jobin MJ, Kulkarni S, McWhinnie A, Dalva K, Gragert L, Babrzadeh F, Gharizadeh B, Luo M, Plummer FA, Kimani J, Carrington M, Middleton D, Rajalingam R, Beksac M, Marsh SG, Maiers M, Guethlein LA, Tavoularis S, Little AM, Green RE, Norman PJ, Parham P |display-authors=6 |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 |date=October 2011 |pmid=21868630 |pmc=3677943 |doi=10.1126/science.1209202 |bibcode=2011Sci...334...89A}}</ref>

Human evolution is characterized by a number of [[morphology (biology)|morphological]], [[human development (biology)|developmental]], [[human physiology|physiological]], and [[Human behavior|behavioral]] changes that have taken place since the split between the [[chimpanzee–human last common ancestor|last common ancestor of humans and chimpanzees]]. The most significant of these adaptations are [[Prehistory_of_nakedness_and_clothing#Evolution_of_hairlessness|hairlessness]],<ref>{{cite journal |last1=Sandel |first1=Aaron A. |title=Brief communication: Hair density and body mass in mammals and the evolution of human hairlessness |journal=American Journal of Physical Anthropology |date=30 July 2013 |volume=152 |issue=1 |pages=145–150 |doi=10.1002/ajpa.22333 |pmid=23900811 |hdl=2027.42/99654 |url=https://onlinelibrary.wiley.com/doi/10.1002/ajpa.22333 |access-date=22 July 2023|hdl-access=free }}</ref> obligate bipedalism, increased brain size and decreased [[sexual dimorphism]] ([[neoteny]]). The relationship between all these changes is the subject of ongoing debate.<ref name="Boyd2003">{{cite book |vauthors=Boyd R, Silk JB |author1-link=Robert Boyd (anthropologist) |author2-link=Joan Silk |url=https://archive.org/details/howhumansevolved03edboyd |title=How Humans Evolved |publisher=[[W. W. Norton & Company|Norton]] |year=2003 |isbn=978-0-393-97854-4 |location=New York |url-access=registration}}</ref>
{{clade|{{clade
       |1=Hylobatidae ([[gibbon]]s)
       |label2=Hominidae (hominids, [[great ape]]s)
       |2={{clade|label1=[[Ponginae]] |1={{clade
         |label1=Pongo ([[orangutan]]s)
           |1={{clade
           |1=''[[Pongo abelii]]''
           |label2=
           |2={{clade
             |1=''[[Pongo tapanuliensis]]''
             |2=''[[Pongo pygmaeus]]''
             }}
           }} }}
         |label2=[[Homininae]] (hominines)
          |2={{clade|label1=[[Gorillini]]  |1={{clade
             |label1=Gorilla ([[gorilla]]s)
             |1={{clade
               |1=''[[Gorilla gorilla]]''
               |2=''[[Gorilla beringei]]''
             }} }}
             |label2=[[Hominini]] (hominins)
             |2={{clade
                |label1=[[Panina]]
                |1={{clade|label1=Pan ([[chimpanzee]]s)|1={{clade
                   |1=''[[Pan troglodytes]]''
                   |2=''[[Pan paniscus]]''
                }} }}
                |2={{clade|label1=[[Hominina]] (homininans)|1='''''Homo sapiens''''' (humans)}}
             }}
         }}
      }}
   }}|style1=font-size:80%; line-height:80%|label1=[[Hominoidea]] (hominoids, [[ape]]s)}}