Editing Early modern human

{{short description|Old Stone Age ''Homo sapiens''}}
{{about|the first Homo sapiens, specifically during the [[Middle Paleolithic]]|a broader perspective on the human species|Human}}
{{pp-vandalism|small=yes}}
[[File:Homo sapiens sapiens (Fundort Jebel Irhoud Marokko).jpg|thumb|Reconstruction of early ''Homo sapiens'' from [[Jebel Irhoud]], Morocco {{c.|315 000 years BP}}]]
'''Early modern human''' ('''EMH'''), or '''anatomically modern human''' ('''AMH'''),<ref>{{cite book|last1=Nitecki|first1=Matthew H|url=https://books.google.com/books?id=tzb5BwAAQBAJ|title=Origins of Anatomically Modern Humans|last2=Nitecki|first2=Doris V|publisher=Springer|year=1994|isbn=1489915079}}</ref> are terms used to distinguish ''[[Homo sapiens]]'' ([[Homo sapiens sapiens|sometimes]] ''Homo sapiens sapiens'') that are [[Human anatomy|anatomically]] consistent with the [[Human variability|range of phenotypes]] seen in contemporary humans, from extinct [[Archaic humans|archaic human]] species (of which some are at times also identified with, but only one, prefix ''sapiens''). This distinction is useful especially for times and regions where anatomically modern and archaic humans co-existed, for example, in [[Paleolithic Europe]]. Among the oldest known remains of ''Homo sapiens'' are those found at the [[Omo remains|Omo-Kibish I]] archaeological site in south-western [[Ethiopia]], dating to about 233,000<ref name="Vidal22">{{Cite journal |last1=Vidal |first1=Celine M. |last2=Lane |first2=Christine S. |author-link2=Christine Lane |last3=Asfawrossen |first3=Asrat |display-authors=etal |date=Jan 2022 |title=Age of the oldest known Homo sapiens from eastern Africa |journal=Nature |volume=601 |issue=7894 |pages=579–583 |bibcode=2022Natur.601..579V |doi=10.1038/s41586-021-04275-8 |pmc=8791829 |pmid=35022610}}</ref> to 196,000 years ago,<ref name="Hammond 199–219">{{Cite journal|last1=Hammond|first1=Ashley S.|last2=Royer|first2=Danielle F.|last3=Fleagle|first3=John G.|date=Jul 2017|title=The Omo-Kibish I pelvis|journal=Journal of Human Evolution|volume=108|pages=199–219|doi=10.1016/j.jhevol.2017.04.004|issn=1095-8606|pmid=28552208|doi-access=free|bibcode=2017JHumE.108..199H }}</ref> the [[Florisbad Skull]] founded at the [[Florisbad archaeological and paleontological site]] in South Africa, dating to about 259,000 years ago{{cn|date=January 2025}}, and the [[Jebel Irhoud]] site in Morocco, dated about 315,000 years ago.{{cn|date=January 2025}}

[[Lists of extinct species|Extinct species]] of the genus ''[[Homo]]'' include ''[[Homo erectus]]'' (extant from roughly 2 to 0.1&nbsp;million years ago) and a number of other species (by some authors considered [[Human subspecies|subspecies]] of either ''H. sapiens'' or ''H. erectus''). The divergence of the lineage leading to ''H. sapiens'' out of ancestral ''H. erectus'' (or an intermediate species such as ''[[Homo antecessor]]'') is estimated to have occurred in Africa roughly 500,000 years ago. The earliest fossil evidence of early modern humans appears in [[Africa]] around 300,000 years ago, with the earliest genetic splits among modern people, according to some evidence, dating to around the same time.<ref name="NAT-20190910">{{cite journal |last1=Mounier |first1=Aurélien |last2=Lahr |first2=Marta |title=Deciphering African late middle Pleistocene hominin diversity and the origin of our species |journal=[[Nature Communications]] |volume=10 |issue=1 |page=3406 |doi=10.1038/s41467-019-11213-w |pmid=31506422 |pmc=6736881 |year=2019 |bibcode=2019NatCo..10.3406M }}</ref><ref name=":02">{{Cite journal|last1=Scerri|first1=Eleanor M. L.|last2=Thomas|first2=Mark G.|last3=Manica|first3=Andrea|last4=Gunz|first4=Philipp|last5=Stock|first5=Jay T.|last6=Stringer|first6=Chris|last7=Grove|first7=Matt|last8=Groucutt|first8=Huw S.|last9=Timmermann|first9=Axel|author-link9= Axel Timmermann|last10=Rightmire|first10=G. Philip|last11=d'Errico|first11=Francesco|date=2018-08-01|title=Did Our Species Evolve in Subdivided Populations across Africa, and Why Does It Matter?|url= |journal=Trends in Ecology & Evolution|language=en|volume=33|issue=8|pages=582–594|doi=10.1016/j.tree.2018.05.005|issn=0169-5347|pmid=30007846|pmc=6092560|bibcode=2018TEcoE..33..582S }}</ref>{{refn|group=note|name=350kiloyearsAgo|Based on Schlebusch et al., "Southern African ancient genomes estimate modern human divergence to 350,000 to 260,000 years ago",<ref name=Schlebusch350-260>{{cite journal |last=Schlebusch |display-authors=etal |title=Southern African ancient genomes estimate modern human divergence to 350,000 to 260,000 years ago |journal=Science |volume=358 |issue=6363 |date=3 November 2017 |pages=652–655 |doi=10.1126/science.aao6266 |pmid=28971970 |bibcode=2017Sci...358..652S |doi-access=free }}</ref> [https://d2ufo47lrtsv5s.cloudfront.net/content/sci/early/2017/09/27/science.aao6266/F3.large.jpg Fig. 3] (''H. sapiens'' divergence times) and Stringer (2012),<ref name=Stringer2012>{{cite journal|last=Stringer |first=C |title=What makes a modern human |journal=Nature |year=2012 |volume=485 |issue=7396 |pages=33–35 |doi=10.1038/485033a |pmid=22552077 |bibcode=2012Natur.485...33S |s2cid=4420496 |doi-access=free }}</ref> (archaic admixture).}}<ref name=":2">{{Cite journal|last1=Neubauer|first1=Simon|last2=Hublin|first2=Jean-Jacques|last3=Gunz|first3=Philipp|date=2018-01-01|title=The evolution of modern human brain shape|journal=Science Advances|language=en|volume=4|issue=1|pages=eaao5961|doi=10.1126/sciadv.aao5961|issn=2375-2548|pmid=29376123|pmc=5783678|bibcode=2018SciA....4.5961N}}</ref> Sustained [[archaic human admixture with modern humans]] is known to have taken place both in Africa and (following the [[Recent African origin of modern humans|recent Out-Of-Africa expansion]]) in Eurasia, between about 100,000 and 30,000 years ago.<ref name=Harrod2014>{{cite journal|url= https://www.academia.edu/34411084 |title=Harrod (2014) Suppl File Table 1 mtDNA language myth Database rev May 17 2019.doc |journal=Mother Tongue |last1=Harrod |first1=James }}</ref>

==Name and taxonomy==
{{further|Homo|Names for the human species}}
{{main|Human taxonomy}}
{{Human timeline}}
The [[binomial nomenclature|binomial name]] ''Homo sapiens'' was coined by [[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]].<ref>{{cite book|last=Linné |first=Carl von |title=Systema naturæ. Regnum animale |year=1758 |pages=18, 20 |url=https://www.biodiversitylibrary.org/item/80764#page/28/mode/1up |edition=10th|access-date=2019-05-06|publisher=Sumptibus Guilielmi Engelmann }}</ref> The [[Latin]] noun ''[[wikt:homo#Latin|homō]]'' (genitive ''hominis'') means "human being", while the participle ''[[wikt:sapiens#Latin|sapiēns]]'' means "discerning, wise, sensible".

The species was initially thought to have emerged from a predecessor within the genus ''Homo'' around 300,000 to 200,000 years ago.{{refn|group=note|This is a matter of convention (rather than a factual dispute), and there is no universal consensus on terminology. Some scholars include humans of up to 600,000 years ago under the same species. See Bryant (2003), p. 811.<ref>{{cite book|url= https://books.google.com/books?id=3z9EpgisKOgC |title=Handbook of Death and Dying |last=Bryant |first=Clifton D |year=2003 |publisher=SAGE |isbn=0761925147}}</ref> See also Tattersall (2012), Page 82 (''cf''. Unfortunately this consensus in principle hardly clarifies matters much in practice. For there is no agreement on what the 'qualities of a man' actually are," [...]).<ref>{{cite book|url= https://books.google.com/books?id=h5PGjJW8FLoC |title=Masters of the Planet: The Search for Our Human Origins |last=Tattersall |first=Ian |year=2012 |publisher=St Martin's Press |isbn=978-1137000385}}</ref>}} A problem with the morphological classification of "anatomically modern" was that it would not have included certain extant populations. For this reason, a lineage-based ([[cladistic]]) definition of ''H. sapiens'' has been suggested, in which ''H. sapiens'' would by definition refer to the modern human lineage following the split from the Neanderthal lineage. Such a cladistic definition would extend the age of ''H. sapiens'' to over 500,000 years.{{refn|group=note|Werdelin<ref>{{cite book|first1=Lars |last1=Werdelin |first2=William Joseph |last2=Sanders |title=Cenozoic Mammals of Africa |year=2010 |url= https://books.google.com/books?id=6c8lDQAAQBAJ&pg=PA517 |page=517 |publisher=Univ of California Press|isbn=978-0520257214 }}</ref> citing Lieberman et al.<ref>{{cite journal|first1=DE |last1=Lieberman |first2=BM |last2=McBratney |first3=G |last3=Krovitz |title=The evolution and development of cranial form in ''Homo sapiens'' |journal=PNAS |year=2002 |volume=99 |issue=3 |pages=1134–1139 |doi=10.1073/pnas.022440799|pmid=11805284 |pmc=122156 |bibcode=2002PNAS...99.1134L |doi-access=free }}</ref>}}

Estimates for the split between the Homo sapiens line and combined [[Neanderthal]]/[[Denisovan]] line range from between 503,000 and 565,000 years ago;<ref name="Hajdinjak2018">{{Cite journal|last1=Hajdinjak|first1=Mateja|last2=Fu|first2=Qiaomei|last3=Hübner|first3=Alexander|last4=Petr|first4=Martin|last5=Mafessoni|first5=Fabrizio|last6=Grote|first6=Steffi|last7=Skoglund|first7=Pontus|last8=Narasimham|first8=Vagheesh|last9=Rougier|first9=Hélène|last10=Crevecoeur|first10=Isabelle|last11=Semal|first11=Patrick|display-authors=4|date=2018-03-01|title=Reconstructing the genetic history of late Neanderthals|journal=[[Nature (journal)|Nature]]|volume=555|issue=7698|pages=652–656|bibcode=2018Natur.555..652H|doi=10.1038/nature26151|issn=1476-4687|pmc=6485383|pmid=29562232|first14=Jean-Jacques|last30=Pääbo|last27=Reich|first27=David|last28=Prüfer|first28=Kay|last29=Meyer|first29=Matthias|first31=Janet|first30=Svante|last31=Kelso|last26=Patterson|last14=Hublin|first13=Sahra|last13=Talamo|first12=Marie|last12=Soressi|first26=Nick|first25=Montgomery|last15=Gušić|last20=Posth|first15=Ivan|last16=Kućan|first16=Željko|last17=Rudan|last18=Golovanova|first18=Liubov V.|last19=Doronichev|first19=Vladimir B.|first20=Cosimo|last25=Slatkin|last21=Krause|first21=Johannes|last22=Korlević|first22=Petra|last23=Nagel|first23=Sarah|last24=Nickel|first24=Birgit|first17=Pavao}}</ref> between 550,000 and 765,000 years ago;<ref name="Meyer2016">{{Cite journal|last1=Meyer|first1=Matthias|last2=Arsuaga|first2=Juan-Luis|last3=de Filippo|first3=Cesare|last4=Nagel|first4=Sarah|last5=Aximu-Petri|first5=Ayinuer|last6=Nickel|first6=Birgit|last7=Martínez|first7=Ignacio|last8=Gracia|first8=Ana|last9=de Castro|first9=José María Bermúdez|last10=Carbonell|first10=Eudald|last11=Viola|first11=Bence|display-authors=4|date=2016-03-01|title=Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins|journal=[[Nature (journal)|Nature]]|volume=531|issue=7595|pages=504–507|bibcode=2016Natur.531..504M|doi=10.1038/nature17405|issn=1476-4687|pmid=26976447|last14=Pääbo|first13=Kay|last13=Prüfer|first12=Janet|last12=Kelso|first14=Svante|s2cid=4467094}}</ref> and (based on rates of dental evolution) possibly more than 800,000 years ago.<ref name="Gómez-Robles2019">{{Cite journal|last=Gómez-Robles|first=Aida|date=2019-05-01|title=Dental evolutionary rates and its implications for the Neanderthal–modern human divergence|journal=[[Science Advances]]|volume=5|issue=5|pages=1268|bibcode=2019SciA....5.1268G|doi=10.1126/sciadv.aaw1268|issn=2375-2548|pmc=6520022|pmid=31106274}}</ref>

Extant human populations have historically been divided into [[Human subspecies|subspecies]], but since around the 1980s all extant groups have tended to be subsumed into a single species, ''H. sapiens'', avoiding division into subspecies altogether.{{refn|group=note|The history of claimed or proposed subspecies of ''H. sapiens'' is complicated and fraught with controversy. The only widely recognized archaic subspecies{{citation needed|date=February 2019}} is ''[[Homo sapiens idaltu|H. sapiens idaltu]]'' (2003). The name ''H. s. sapiens'' is due to [[Carl Linnaeus|Linnaeus]] ([[10th edition of Systema Naturae|1758]]), and refers by definition the subspecies of which Linnaeus himself is the type specimen. However, Linnaeus postulated four other extant subspecies, viz. ''H. s. afer'', ''H. s. americanus'',  ''H. s. asiaticus'' and ''H. s. ferus'' for Africans, Americans, Asians and [[Malay race|Malay]]. This classification remained in common usage until the mid 20th century, sometimes alongside ''H. s. tasmanianus'' for Australians. See, for example, Bailey, 1946;<ref>{{cite book|url= https://books.google.com/books?id=y3AxAAAAMAAJ |first=John Wendell |last=Bailey |title=The Mammals of Virginia |year=1946 |page=356}}</ref> Hall, 1946.<ref name=Hall>{{cite journal|last=Hall |first=E |year=1946 |title=Zoological Subspecies of Man at the Peace Table |journal=Journal of Mammalogy |volume=27 |issue=4 |pages=358–364 |doi=10.2307/1375342|jstor=1375342 |pmid=20247535 }}</ref> The division of extant human populations into taxonomic subspecies was gradually given up in the 1970s (for example, [[Grzimek's Animal Life Encyclopedia]]<ref>{{cite encyclopedia|url= https://books.google.com/books?id=ZHNMAQAAIAAJ |title=Grzimek's Animal Life Encyclopedia |volume=11 |page=55 |year=1970|isbn=978-0442784782 |last1=Grzimek |first1=Bernhard |publisher=Van Nostrand Reinhold Company }}</ref>).}}

Some sources show Neanderthals (''H. neanderthalensis'') as a subspecies (''H. sapiens neanderthalensis'').<ref name="HublinOrigin">{{cite journal |doi=10.1073/pnas.0904119106 |title=The origin of Neandertals |year=2009 |last1=Hublin |first1=J. J. |journal=Proceedings of the National Academy of Sciences |volume=106 |issue=38 |pages=16022–16027 |pmid=19805257 |jstor=40485013 |bibcode=2009PNAS..10616022H |pmc=2752594|doi-access=free }}</ref><ref name="HFMcN">{{Cite journal |title=Neanderthal taxonomy reconsidered: implications of 3D primate models of intra- and interspecific differences |pmid=14745010 |last1=Harvati |first1=K. |last2=Frost |first2=S.R. |last3=McNulty |first3=K.P. |date=2004 |doi=10.1073/pnas.0308085100 |pmc=337021 |volume=101 |issue=5 |journal=Proc. Natl. Acad. Sci. U.S.A. |pages=1147–1152|bibcode=2004PNAS..101.1147H |doi-access=free }}</ref> Similarly, the discovered specimens of the ''[[Homo rhodesiensis|H. rhodesiensis]]'' species have been classified by some as a subspecies (''H. sapiens rhodesiensis''), although it remains more common to treat these last two as separate species within the genus ''Homo'' rather than as subspecies within ''H. sapiens''.<ref>{{cite encyclopedia |title= ''Homo neanderthalensis'' King, 1864|encyclopedia= Wiley-Blackwell Encyclopedia of Human Evolution|year= 2013 |publisher= Wiley-Blackwell |location= Chichester, West Sussex|pages=328–331}}</ref>

All humans are considered to be a part of the subspecies ''[[Homo sapiens sapiens|H. sapiens sapiens]]'',<ref name="britannica-H.-sapiens-sapiens">{{Cite web|last=Rafferty|first=John P.|title=Homo sapiens sapiens|url=https://www.britannica.com/topic/Homo-sapiens-sapiens|access-date=2020-08-11|website=Encyclopedia Britannica|language=en}}</ref> a designation which has been a matter of debate since a species is usually not given a subspecies category unless there is evidence of multiple distinct subspecies.<ref name="britannica-H.-sapiens-sapiens" />

== Age and speciation process ==
{{further|Human evolution|Homo|Multiregional origin of modern humans|Timeline of human evolution|Early human migrations}}

=== Derivation from ''H. erectus'' ===
{{further|Homo ergaster|Homo antecessor|Homo heidelbergensis|Homo rhodesiensis|Acheulean}}

The divergence of the lineage that would lead to ''H. sapiens'' out of [[archaic human]] varieties derived from ''H. erectus'', is estimated as having taken place over 500,000 years ago (marking the split of the ''H. sapiens'' lineage from ancestors shared with other known archaic hominins).<ref name=":2" /><ref name=":02"/> But the oldest split among modern human populations (such as the [[Khoisan]] split from other groups) has been recently dated to between 350,000 and 260,000 years ago,<ref>{{cite bioRxiv |title=Ancient genomes from southern Africa pushes modern human divergence beyond 260,000 years ago |first1=Carina M. |last1=Schlebusch |first2=Helena |last2=Malmström |first3=Torsten |last3=Günther |first4=Per |last4=Sjödin |first5=Alexandra |last5=Coutinho |first6=Hanna |last6=Edlund |first7=Arielle R. |last7=Munters |first8=Maryna |last8=Steyn |first9=Himla |last9=Soodyall |first10=Marlize |last10=Lombard |first11=Mattias |last11=Jakobsson |date=5 June 2017 |biorxiv=10.1101/145409}}</ref><ref>{{Cite journal |last1=Schlebusch |first1=Carina M. |last2=Malmström |first2=Helena |last3=Günther |first3=Torsten |last4=Sjödin |first4=Per |last5=Coutinho |first5=Alexandra |last6=Edlund |first6=Hanna |last7=Munters |first7=Arielle R. |last8=Vicente |first8=Mário |last9=Steyn |first9=Maryna |last10=Soodyall |first10=Himla |last11=Lombard |first11=Marlize |date=2017-11-03 |title=Southern African ancient genomes estimate modern human divergence to 350,000 to 260,000 years ago |journal=[[Science (journal)|Science]] |language=en |volume=358 |issue=6363 |pages=652–655 |doi=10.1126/science.aao6266 |issn=0036-8075 |pmid=28971970 |bibcode=2017Sci...358..652S |doi-access=free}}</ref> and the earliest known examples of ''H. sapiens'' fossils also date to about that period, including the [[Jebel Irhoud]] remains from Morocco (ca. 300,000 or 350–280,000 years ago),<ref name="NAT-20170607a"/> the [[Florisbad Skull]] from South Africa (ca. 259,000 years ago), and the [[Omo remains]] from Ethiopia (ca. 195,000, or, as more recently dated, ca. 233,000 years ago).<ref>{{harvp|Stringer|2016|p=20150237}}; {{harvp|Sample|2017}}; {{harvp|Hublin|Ben-Ncer|Bailey|Freidline|2017|pp=289–292}}; {{harvp|Scerri|2018|pp=582–594}}</ref><ref name="Vidal22"/>

An mtDNA study in 2019 proposed an origin of modern humans in Botswana (and a Khoisan split) of around 200,000 years.<ref name="NAT-201910283">{{cite journal|last=Chan|first=Eva, K. F.|date=28 October 2019|title=Human origins in a southern African palaeo-wetland and first migrations|journal=[[Nature (journal)|Nature]]|volume=857|issue=7781|pages=185–189|doi=10.1038/s41586-019-1714-1|pmid=31659339|bibcode=2019Natur.575..185C|s2cid=204946938}}</ref> However, this proposal has been widely criticized by scholars,<ref name="Sample">{{Cite news|url=https://www.theguardian.com/science/2019/oct/28/ancestral-home-of-modern-humans-is-in-botswana-study-finds|title=Ancestral home of modern humans is in Botswana, study finds|last=Sample|first=Ian|date=28 October 2019|work=The Guardian|access-date=29 October 2019|language=en-GB|issn=0261-3077}}</ref><ref name="SA-20191028">{{cite news|url=https://www.sciencealert.com/new-study-finds-the-ancestral-homeland-of-all-humans-alive-today|title=New Study Pinpoints The Ancestral Homeland of All Humans Alive Today|last=Woodward|first=Aylin|date=28 October 2019|work=ScienceAlert.com|access-date=29 October 2019}}</ref><ref name="The Atlantic-600826">{{cite news|url=https://www.theatlantic.com/science/archive/2019/10/controversial-study-pinpoints-humanitys-homeland/600826/|title=Has Humanity's Homeland Been Found?|last=Yong|first=Ed|date=28 October 2019|work=[[The Atlantic]]|access-date=28 October 2019|author-link=Ed Yong}}</ref> with the recent evidence overall (genetic, fossil, and archaeological) supporting an origin for ''H. sapiens'' approximately 100,000 years earlier and in a broader region of Africa than the study proposes.<ref name="The Atlantic-600826" />

In September 2019, scientists proposed that the earliest ''H. sapiens'' (and last common human ancestor to modern humans) arose between 350,000 and 260,000 years ago through a merging of populations in [[East Africa|East]] and [[South Africa]].<ref name="NYT-20190910">{{cite news |last=Zimmer |first=Carl |author-link=Carl Zimmer |title=Scientists Find the Skull of Humanity's Ancestor – on a Computer – By comparing fossils and CT scans, researchers say they have reconstructed the skull of the last common forebear of modern humans. |url=https://www.nytimes.com/2019/09/10/science/human-ancestor-skull-computer.html |date=10 September 2019 |work=[[The New York Times]] |access-date=10 September 2019 }}</ref><ref name="NAT-20190910"/>

An alternative suggestion defines ''H. sapiens'' [[Cladistics|cladistically]] as including the lineage of modern humans since the split from the lineage of [[Neanderthal]]s, roughly 500,000 to 800,000 years ago.

The time of divergence between archaic ''H. sapiens'' and ancestors of Neanderthals and Denisovans caused by a [[genetic bottleneck]] of the latter was dated at 744,000 years ago, combined with repeated early admixture events and [[Denisovan]]s diverging from Neanderthals 300 generations after their split from ''H. sapiens'', as calculated by Rogers et al. (2017).<ref>{{cite journal |last1=Rogers |first1=Alan R. |last2=Bohlender |first2=Ryan J. |last3=Huff |first3=Chad D. |title=Early history of Neanderthals and Denisovans |journal=Proceedings of the National Academy of Sciences |date=12 September 2017 |volume=114 |issue=37 |pages=9859–9863 |doi=10.1073/pnas.1706426114 |pmid=28784789 |pmc=5604018|bibcode=2017PNAS..114.9859R |doi-access=free }}</ref>

The derivation of a comparatively homogeneous single species of ''H. sapiens'' from more diverse varieties of [[archaic humans]] (all of which were descended from the [[Out of Africa I|early dispersal]] of ''H. erectus'' some 1.8&nbsp;million years ago) was debated in terms of two competing models during the 1980s: "[[recent African origin]]" postulated the emergence of ''H. sapiens'' from a single source population in Africa, which expanded and led to the extinction of all other human varieties, while the "[[multiregional evolution]]" model postulated the survival of regional forms of archaic humans, gradually converging into the [[human genetic variation|modern human varieties]] by the mechanism of [[cline (population genetics)|clinal variation]], via [[genetic drift]], [[gene flow]] and [[Natural selection|selection]] throughout the Pleistocene.<ref name=Wolpoff>{{cite journal |last1=Wolpoff |first1=M. H. |last2=Spuhler |first2=J. N. |last3=Smith |first3=F. H. |last4=Radovcic |first4=J. |last5=Pope |first5=G. |last6=Frayer |first6=D. W. |last7=Eckhardt |first7=R. |last8=Clark |first8=G. |year=1988 |title=Modern Human Origins |journal=Science |volume=241 |issue=4867|pages=772–774 |doi=10.1126/science.3136545 |pmid=3136545|bibcode=1988Sci...241..772W |s2cid=5223638 }}</ref>

Since the 2000s, the availability of data from [[archaeogenetics]] and [[population genetics]] has led to the emergence of a much more detailed picture, intermediate between the two competing scenarios outlined above: The [[recent African origin|recent Out-of-Africa]] expansion accounts for the predominant part of modern human ancestry, while there were also significant [[Archaic human admixture with modern humans|admixture events]] with regional archaic humans.<ref name=Draft>{{cite journal |last1=Green |first1=RE |last2=Krause |first2=J |last3=Briggs |first3=Adrian W. |last4=Maricic |first4=Tomislav |last5=Stenzel |first5=Udo |last6=Kircher |first6=Martin |last7=Patterson |first7=Nick |last8=Li |first8=Heng |last9=Zhai |first9=Weiwei |last10=Fritz |first10=Markus Hsi-Yang |last11=Hansen |first11=Nancy F. |last12=Durand |first12=Eric Y. |last13=Malaspinas |first13=Anna-Sapfo |last14=Jensen |first14=Jeffrey D. |last15=Marques-Bonet |first15=Tomas |last16=Alkan |first16=Can |last17=Prüfer |first17=Kay |last18=Meyer |first18=Matthias |last19=Burbano |first19=Hernán A. |last20=Good |first20=Jeffrey M. |last21=Schultz |first21=Rigo |last22=Aximu-Petri |first22=Ayinuer |last23=Butthof |first23=Anne |last24=Höber |first24=Barbara |last25=Höffner |first25=Barbara |last26=Siegemund |first26=Madlen |last27=Weihmann |first27=Antje |last28=Nusbaum |first28=Chad |last29=Lander |first29=Eric S. |last30=Russ |first30=Carsten |name-list-style=vanc |date=May 2010 |title=A draft sequence of the Neandertal genome |journal=Science |volume=328 |issue=5979|pages=710–722 |doi=10.1126/science.1188021 |pmid=20448178 |bibcode=2010Sci...328..710G |display-authors=29 |pmc=5100745}}</ref><ref>{{cite journal |last1=Reich |first1=D |last2=Patterson |first2=Nick |last3=Kircher |first3=Martin |last4=Delfin |first4=Frederick |last5=Nandineni |first5=Madhusudan R. |last6=Pugach |first6=Irina |last7=Ko |first7=Albert Min-Shan |last8=Ko |first8=Ying-Chin |last9=Jinam |first9=Timothy A. |last10=Phipps |first10=Maude E. |last11=Saitou |first11=Naruya |last12=Wollstein |first12=Andreas |last13=Kayser |first13=Manfred |last14=Pääbo |first14=Svante |last15=Stoneking |first15=Mark |name-list-style=vanc |year=2011 |title=Denisova admixture and the first modern human dispersals into southeast Asia and oceania |journal=Am J Hum Genet |volume=89 |issue=4 |pages=516–528 |doi=10.1016/j.ajhg.2011.09.005 |pmid=21944045 |pmc=3188841}}</ref>

Since the 1970s, the Omo remains, originally dated to some 195,000 years ago, have often been taken as the conventional cut-off point for the emergence of "anatomically modern humans". Since the 2000s, the discovery of older remains with comparable characteristics, and the discovery of ongoing hybridization between "modern" and "archaic" populations after the time of the Omo remains, have opened up a renewed debate on the age of ''H. sapiens'' in journalistic publications.<ref>{{cite web|url= https://www.nsf.gov/news/news_summ.jsp?cntn_id=102968 |title=New Clues Add 40,000 Years to Age of Human Species |publisher=NSF – National Science Foundation|website=www.nsf.gov}}</ref><ref>{{cite news|url= http://news.bbc.co.uk/2/hi/science/nature/4269299.stm |work=BBC News |title=Age of ancient humans reassessed |date=February 16, 2005 |access-date=April 10, 2010}}</ref><ref>{{cite web|url= https://www.sciencedaily.com/releases/2005/02/050223122209.htm |title=The Oldest Homo Sapiens: Fossils Push Human Emergence Back To 195,000 Years Ago |date=February 28, 2005 |access-date=2019-05-06 |website=ScienceDaily}}</ref><ref>{{cite journal |author=Alemseged, Z. |author2=Coppens, Y. |author3=Geraads, D. |title=Hominid cranium from Homo: Description and taxonomy of Homo-323-1976-896 |journal=Am J Phys Anthropol |volume=117 |issue=2 |pages=103–112 |year=2002 |pmid=11815945 |doi=10.1002/ajpa.10032|url=http://doc.rero.ch/record/13324/files/PAL_E59.pdf |archive-url=https://web.archive.org/web/20200718021120/http://doc.rero.ch/record/13324/files/PAL_E59.pdf |archive-date=2020-07-18 |url-status=live }}</ref><ref>{{cite journal |author1=Stoneking, Mark |author2=Soodyall, Himla |title=Human evolution and the mitochondrial genome |journal=Current Opinion in Genetics & Development |volume=6 |issue=6 |pages=731–736 |year=1996 |doi=10.1016/S0959-437X(96)80028-1|pmid=8994844 }}</ref> ''H. s. idaltu'', dated to 160,000 years ago, has been postulated as an extinct subspecies of ''H. sapiens'' in 2003.<ref>[http://www.anth.ucsb.edu/projects/human/ Human evolution: the fossil evidence in 3D], by Philip L. Walker and Edward H. Hagen, Dept. of Anthropology, University of California, Santa Barbara. Retrieved April 5, 2005.</ref><ref name="britannica-H.-sapiens-sapiens" /> ''H. neanderthalensis'', which became extinct about 40,000 years ago, was also at one point considered to be a subspecies, ''H. s. neanderthalensis''.<ref name="britannica-H.-sapiens-sapiens" />

''H. heidelbergensis'', dated 600,000 to 300,000 years ago, has long been thought to be a likely candidate for the last common ancestor of the Neanderthal and modern human lineages. However, genetic evidence from the [[Sima de los Huesos]] fossils published in 2016 seems to suggest that ''H. heidelbergensis'' in its entirety should be included in the Neanderthal lineage, as "pre-Neanderthal" or "early Neanderthal", while the divergence time between the Neanderthal and modern lineages has been pushed back to before the emergence of ''H. heidelbergensis'', to close to 800,000 years ago, the approximate time of disappearance of ''H. antecessor''.<ref>{{cite journal |last1=Meyer |first1=Matthias |last2=Arsuaga |first2=Juan-Luis |last3=de Filippo |first3=Cesare |last4=Nagel |first4=Sarah |last5=Aximu-Petri |first5=Ayinuer |last6=Nickel |first6=Birgit |last7=Martínez |first7=Ignacio |last8=Gracia |first8=Ana |last9=de Castro |first9=José María Bermúdez |last10=Carbonell |first10=Eudald |last11=Viola |first11=Bence |last12=Kelso |first12=Janet |last13=Prüfer |first13=Kay |last14=Pääbo |first14=Svante |title=Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins |journal=Nature |date=14 March 2016 |volume=531 |issue=7595 |pages=504–507 |doi=10.1038/nature17405 |pmid=26976447|bibcode=2016Natur.531..504M |s2cid=4467094 }}</ref><ref>{{cite journal |last1=Callaway |first1=Ewen |title=Oldest ancient-human DNA details dawn of Neanderthals |journal=Nature |date=14 March 2016 |volume=531 |issue=7594 |pages=296–286 |doi=10.1038/531286a|pmid=26983523 |bibcode=2016Natur.531..296C |s2cid=4459329 |doi-access=free }}</ref>

===Early ''Homo sapiens''===
{{further|Human subspecies|Middle Paleolithic|Mousterian|Interbreeding between archaic and modern humans|Homo sapiens idaltu|Skhul and Qafzeh hominins}}
[[File:Skhul.JPG|thumb|100 to 80 thousand year old [[Skhul V]] from [[Israel]]]]
The term [[Middle Paleolithic]] is intended to cover the time between the first emergence of ''H. sapiens'' (roughly 300,000 years ago) and the period held by some to mark the emergence of full [[behavioral modernity]] (roughly by 50,000 years ago, corresponding to the start of the [[Upper Paleolithic]]).

Many of the early modern human finds, like those of [[Jebel Irhoud]], [[Omo remains|Omo]], [[Homo sapiens idaltu|Herto]], [[Florisbad Skull|Florisbad]], [[Skhul remains|Skhul]], and [[Peștera cu Oase]] exhibit a mix of archaic and modern traits.<ref name="Oppenheimer">{{cite book |last=Oppenheimer |first=S. |title=Out of Eden: The Peopling of the World |year=2003 |publisher=Robinson |isbn=978-1841196978}}</ref><ref>{{Cite journal|doi=10.1073/pnas.2035108100 |last1=Trinkaus |first1=E. |last2=Moldovan |first2=O. |last3=Milota |first3=Ș. |last4=Bîlgăr |first4=A. |last5=Sarcina |first5=L. |last6=Athreya |first6=S. |last7=Bailey |first7=S. E. |last8=Rodrigo |first8=R. |last9=Gherase |first9=M. |last10=Higham |first10=T. |last11=Ramsey |first11=C. B. |last12=Van Der Plicht |first12=J. |year=2003 |title=An early modern human from Peștera cu Oase, Romania |journal=PNAS |volume=100 |issue=20|pages=11231–11236 |pmid=14504393 |pmc=208740 |bibcode=2003PNAS..10011231T |display-authors=8 |doi-access=free }}</ref><ref name="NAT-20170607a">{{cite journal |last=Callaway |first=Ewan |title=Oldest Homo sapiens fossil claim rewrites our species' history |url= http://www.nature.com/news/oldest-homo-sapiens-fossil-claim-rewrites-our-species-history-1.22114 |date=7 June 2017 |journal=[[Nature (journal)|Nature]] |doi=10.1038/nature.2017.22114 |access-date=11 June 2017 }}</ref> Skhul V, for example, has prominent brow ridges and a projecting face. However, the [[Neurocranium|brain case]] is quite rounded and distinct from that of the Neanderthals and is similar to the brain case of modern humans. It is uncertain whether the robust traits of some of the early modern humans like Skhul V reflects [[Archaic human admixture with modern humans|mixed ancestry]] or retention of older traits.<ref name="Reich et al.">{{Cite journal|first1=David |last1=Reich |first2=Richard E. |last2=Green |first3=Martin |last3=Kircher |first4=Johannes |last4=Krause |first5=Nick |last5=Patterson |first6=Eric Y. |last6=Durand |first7=Bence |last7=Viola |first8=Adrian W. |last8=Briggs |first9=Udo |last9=Stenzel |first10=Philip L. F. |last10=Johnson |first11=Tomislav |last11=Maricic |first12=Jeffrey M. |last12=Good |first13=Tomas |last13=marques-Bonet |first14=Can |last14=Alkan |first15=Qiaomei |last15=Fu |first16=Swapan |last16=Mallick |first17=Heng |last17=Li |first18=Matthias |last18=Meyer |first19=Evan E. |last19=Eichler |first20=Mark |last20=Stoneking |first21=Michael |last21=Richards |first22=Sahra |last22=Talamo |first23=Michael V. |last23=Shunkov |first24=Anatoli P. |last24=Derevianko |first25=Jean-Jacques |last25=Hublin |first26=Janet |last26=Kelso |first27=Montgomery |last27=Slatkin |name-list-style=amp|first28=Svante |last28=Pääbo |display-authors=8 |year=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 |doi=10.1038/nature09710 |pmid=21179161|bibcode= 2010Natur.468.1053R |pmc=4306417 |hdl=10230/25596}}</ref><ref>{{cite journal|last=Trinkaus |first=Erik |title=Early modern humans |journal=Annual Review of Anthropology |date=October 2005 |volume=34 |issue=1 |pages=207–230 |doi=10.1146/annurev.anthro.34.030905.154913|s2cid=9039428 |s2cid-access=free}}</ref>

The "gracile" or lightly built skeleton of anatomically modern humans has been connected to a change in behavior, including increased cooperation and "resource transport".<ref>{{cite book|author1=Meldrum, Jeff |author2=Hilton, Charles E. |title=From Biped to Strider: The Emergence of Modern Human Walking, Running, and Resource Transport |url=https://books.google.com/books?id=IfIWVrxg-hEC |date= 2004 |publisher=Springer Science & Business Media |isbn=978-0306480003}}</ref><ref>{{cite book |author1=Vonk, Jennifer |author2=Shackelford, Todd K. |title=The Oxford Handbook of Comparative Evolutionary Psychology |url=https://books.google.com/books?id=btS8XyqTY6MC&pg=PA429 |date= 2012 |publisher=Oxford University Press, US |isbn=978-0199738182 |pages=429–}}</ref>

There is evidence that the characteristic human brain development, especially the prefrontal cortex, was due to "an exceptional acceleration of [[metabolome]] evolution ... paralleled by a drastic reduction in muscle strength. The observed rapid metabolic changes in brain and muscle, together with the unique human cognitive skills and low muscle performance, might reflect parallel mechanisms in human evolution."<ref>{{cite journal |doi=10.1371/journal.pbio.1001871 |pmid=24866127 |title=Exceptional Evolutionary Divergence of Human Muscle and Brain Metabolomes Parallels Human Cognitive and Physical Uniqueness |journal=PLOS Biology |volume=12 |issue=5 |pages=e1001871 |year=2014 |last1=Bozek |first1=Katarzyna |last2=Wei |first2=Yuning |last3=Yan |first3=Zheng |last4=Liu |first4=Xiling |last5=Xiong |first5=Jieyi |last6=Sugimoto |first6=Masahiro |last7=Tomita |first7=Masaru |last8=Pääbo |first8=Svante |last9=Pieszek |first9=Raik |last10=Sherwood |first10=Chet C. |last11=Hof |first11=Patrick R. |last12=Ely |first12=John J. |last13=Steinhauser |first13=Dirk |last14=Willmitzer |first14=Lothar |last15=Bangsbo |first15=Jens |last16=Hansson |first16=Ola |last17=Call |first17=Josep |last18=Giavalisco |first18=Patrick |last19=Khaitovich |first19=Philipp |pmc=4035273 |doi-access=free }}</ref> The [[Schöningen spears]] and their correlation of finds are evidence that complex technological skills already existed 300,000 years ago, and are the first obvious proof of an active [[Big game hunting|(big game) hunt]]. ''H. heidelbergensis'' already had intellectual and cognitive skills like anticipatory planning, thinking and acting that so far have only been attributed to modern man.<ref>{{cite encyclopedia|last=Thieme |first=H |year=2007 |title=Der große Wurf von Schöningen: Das neue Bild zur Kultur des frühen Menschen |pages=224–328 |encyclopedia=Die Schöninger Speere – Mensch und Jagd vor 400 000 Jahren |publisher=Konrad Theiss Verlag |isbn=978-3896460400}}</ref><ref>{{cite encyclopedia|last=Haidle |first=M.N. |year=2006 |title=Menschenaffen? Affenmenschen? Mensch! Kognition und Sprache im Altpaläolithikum |pages=69–97 |editor-last=Conard |editor-first=N.J. |encyclopedia=Woher kommt der Mensch |publisher=Attempto Verlag |isbn=3893083812}}</ref>

The ongoing admixture events within anatomically modern human populations make it difficult to estimate the age of the matrilinear and patrilinear most recent common ancestors of modern populations ([[Mitochondrial Eve]] and [[Y-chromosomal Adam]]). Estimates of the age of Y-chromosomal Adam have been pushed back significantly with the discovery of an ancient Y-chromosomal lineage in 2013, to likely beyond 300,000 years ago.{{refn|group=note|(95% confidence interval 237–581 kya)<ref>{{Cite journal |last1=Mendez |first1=Fernando |last2=Krahn |first2=Thomas |last3=Schrack |first3=Bonnie |last4=Krahn |first4=Astrid-Maria |last5=Veeramah |first5=Krishna |last6=Woerner |first6=August |last7=Fomine |first7=Forka Leypey Mathew |last8=Bradman |first8=Neil |last9=Thomas |first9=Mark |title=An African American paternal lineage adds an extremely ancient root to the human Y chromosome phylogenetic tree |journal=[[American Journal of Human Genetics]] |date=7 March 2013 |doi=10.1016/j.ajhg.2013.02.002 |url=http://haplogroup-a.com/Ancient-Root-AJHG2013.pdf |volume=92 |issue=3 |pages=454–459 |pmid=23453668 |pmc=3591855 |access-date=21 April 2018 |archive-date=24 September 2019 |archive-url=https://web.archive.org/web/20190924160717/http://haplogroup-a.com/Ancient-Root-AJHG2013.pdf |url-status=dead }}</ref>}} There have, however, been no reports of the survival of Y-chromosomal or mitochondrial DNA clearly deriving from archaic humans (which would push back the age of the most recent patrilinear or matrilinear ancestor beyond 500,000 years).<ref>{{cite journal |vauthors=Krings M, Stone A, Schmitz RW, Krainitzki H, Stoneking M, Pääbo S |title=Neandertal DNA sequences and the origin of modern humans |journal=Cell |volume=90 |issue=1 |pages=19–30 |date=July 1997 |pmid=9230299 |doi=10.1016/S0092-8674(00)80310-4|hdl=11858/00-001M-0000-0025-0960-8 |s2cid=13581775 |hdl-access=free}}</ref><ref>{{cite web |url=https://www.science.org/content/article/no-neandertals-gene-pool |last=Hill |first=Deborah |date=16 March 2004 |title=No Neandertals in the Gene Pool |publisher=Science |access-date=6 May 2019}}</ref><ref>{{cite journal |last1=Serre |year=2004 |title=No evidence of Neandertal mtDNA contribution to early modern humans |journal=[[PLOS Biology]] |volume=2 |issue=3 |pages=313–317 |pmid=15024415 |doi=10.1371/journal.pbio.0020057 |last2=Langaney |first2=A. |last3=Chech |first3=M. |last4=Teschler-Nicola |first4=M. |last5=Paunovic |first5=M. |last6=Mennecier |first6=P. |last7=Hofreiter |first7=M. |last8=Possnert |first8=G. |last9=Pääbo |first9=S. |pmc=368159 |first1=D. |author4-link=Maria Teschler-Nicola |doi-access=free }}</ref>

Fossil teeth found at [[Qesem Cave]] (Israel) and dated to between 400,000 and 200,000 years ago have been compared to the dental material from the younger (120,000–80,000 years ago) [[Skhul and Qafzeh hominins]].{{refn|group=note|"Although none of the Qesem teeth shows a suite of Neanderthal characters, a few traits may suggest some affinities with members of the Neanderthal evolutionary lineage. However, the balance of the evidence suggests a closer similarity with the Skhul/Qafzeh dental material, although many of these resemblances likely represent plesiomorphous features."<ref>{{cite journal|last1=Hershkovitz |first1=I |last2=Smith |first2=P |last3=Sarig |first3=R |last4=Quam |first4=R |last5=Rodríguez |first5=L |last6=García |first6=R |last7=Arsuaga |first7=JL |last8=Barkai |first8=R |last9=Gopher |first9=A |title=Middle pleistocene dental remains from Qesem Cave (Israel) |journal=American Journal of Physical Anthropology |year=2011 |volume=144 |issue=4 |pages=575–592 |doi=10.1002/ajpa.21446 |pmid=21404234 |s2cid=3106938}}</ref>}}

== Dispersal and archaic admixture ==
{{further|Recent African origin of modern humans|Southern Dispersal|Early human migrations|List of first human settlements}}
{{further|Interbreeding between archaic and modern humans}}
Dispersal of early ''H. sapiens'' begins soon after its emergence, as evidenced by the North African [[Jebel Irhoud]] finds (dated to around 315,000 years ago).<ref name="NAT-20170607a"/>{{sfn|Hublin|Ben-Ncer|Bailey|Freidline|2017|pp=289–292}} There is indirect evidence for ''H. sapiens'' presence in West Asia around 270,000 years ago.<ref name="NC-20170704">{{cite journal |last=Posth |first=Cosimo |display-authors=etal |title=Deeply divergent archaic mitochondrial genome provides lower time boundary for African gene flow into Neanderthals |date=4 July 2017 |journal=[[Nature Communications]] |volume=8 |page=16046 |doi=10.1038/ncomms16046 |pmid=28675384 |pmc=5500885 |bibcode=2017NatCo...816046P}}</ref>

The [[Florisbad Skull]] from Florisbad, South Africa, dated to about 259,000 years ago, has also been classified as representing early ''H. sapiens''.{{sfn|Stringer|2016|p=20150237}}{{sfn|Sample|2017}}{{harvp|Scerri|2018|pp=582–594}}<ref name="NAT-20190910"/>

In September 2019, scientists proposed that the earliest ''H. sapiens'' (and last common human ancestor to modern humans) arose between 350,000 and 260,000 years ago through a merging of populations in [[East Africa|East]] and [[South Africa]].<ref name="NYT-20190910"/><ref name="NAT-20190910"/>

Among extant populations, the [[Khoi-San]] (or "[[Capoid]]") hunters-gatherers of Southern Africa may represent the human population with the earliest possible divergence within the group ''Homo sapiens sapiens''. Their separation time has been estimated in a 2017 study to be between 350 and 260,000 years ago, compatible with the estimated age of early ''H. sapiens''. The study states that the deep split-time estimation of 350 to 260 thousand years ago is consistent with the archaeological estimate for the onset of the Middle Stone Age across sub-Saharan Africa and coincides with archaic ''H. sapiens'' in southern Africa represented by, for example, the Florisbad skull dating to 259 (± 35) thousand years ago.<ref name=Schlebusch350-260 />

''H. s. idaltu'', found at [[Middle Awash]] in Ethiopia, lived about 160,000 years ago,<ref>{{cite journal |last1=White |first1=Tim D. |last2=Asfaw |first2=Berhane |last3=Degusta |first3=David |last4=Gilbert |first4=Henry |last5=Richards |first5=Gary D. |last6=Suwa |first6=Gen |last7=Howell |first7=Clark F. |date=June 2003 |title=Pleistocene ''Homo sapiens'' from Middle Awash, Ethiopia |journal=[[Nature (journal)|Nature]] |volume=423 |issue=6941 |pages=742–747 |pmid=12802332 |doi=10.1038/nature01669 |bibcode=2003Natur.423..742W |s2cid=4432091}}</ref> and ''H. sapiens'' lived at Omo Kibish in Ethiopia about 233,000-195,000 years ago.<ref name="USER:CALR">{{cite magazine |title=Fossil Reanalysis Pushes Back Origin of ''Homo sapiens'' |magazine=[[Scientific American]] |date=17 February 2005 |access-date=6 May 2019 |url=https://www.scientificamerican.com/article/fossil-reanalysis-pushes/}}</ref><ref name="Vidal22"/> Two fossils from Guomde, Kenya, dated to at least (and likely more than) 180,000 years ago{{sfn|Stringer|2016|p=20150237}} and (more precisely) to 300–270,000 years ago,<ref name="NAT-20190910"/> have been tentatively assigned to ''H. sapiens'' and similarities have been noted between them and the Omo Kibbish remains.{{sfn|Stringer|2016|p=20150237}} Fossil evidence for modern human presence in West Asia is ascertained for 177,000 years ago,<ref>{{cite web |url=https://www.ibtimes.co.in/177000-year-old-jawbone-fossil-discovered-israel-oldest-human-remains-found-outside-africa-758401 |title=A 177,000-year-old jawbone fossil discovered in Israel is oldest human remains found outside Africa |first=Ankita |last=Mehta |date=26 January 2018 |website=International Business Times |access-date=6 May 2019}}</ref> and disputed fossil evidence suggests expansion as far as East Asia by 120,000 years ago.<ref name="SCI-20171208">{{cite journal |last1=Bae |first1=Christopher J. |last2=Douka |first2=Katerina |last3=Petraglia |first3=Michael D. |title=On the origin of modern humans: Asian perspectives |journal=[[Science (journal)|Science]] |date=8 December 2017 |volume=358 |issue=6368 |page=eaai9067 |doi=10.1126/science.aai9067 |pmid=29217544 |doi-access=free }}</ref><ref name="QZ-20171210">{{cite web |last=Kuo |first=Lily |title=Early humans migrated out of Africa much earlier than we thought |url= https://qz.com/1151816/early-humans-migrated-out-of-africa-much-earlier-than-we-thought/ |date=10 December 2017 |website=[[Quartz (publication)|Quartz]] |access-date=6 May 2019}}</ref>

In July 2019, anthropologists reported the discovery of 210,000 year old remains of a ''H. sapiens'' and 170,000 year old remains of a ''H. neanderthalensis'' in [[Apidima Cave]], [[Peloponnese]], [[Greece]], more than 150,000 years older than previous ''H. sapiens'' finds in Europe.<ref name="NYT-20190710">{{cite news |last=Zimmer |first=Carl |author-link=Carl Zimmer |title=A Skull Bone Discovered in Greece May Alter the Story of Human Prehistory – The bone, found in a cave, is the oldest modern human fossil ever discovered in Europe. It hints that humans began leaving Africa far earlier than once thought. |url=https://www.nytimes.com/2019/07/10/science/skull-neanderthal-human-europe-greece.html |date=10 July 2019 |work=[[The New York Times]] |access-date=11 July 2019 }}</ref><ref name="PHYS-20190710">{{cite news |author=Staff |title='Oldest remains' outside Africa reset human migration clock |url=https://phys.org/news/2019-07-oldest-africa-reset-human-migration.html |date=10 July 2019 |work=[[Phys.org]] |access-date=10 July 2019 }}</ref><ref name="NAT-20190710">{{cite journal |last=Harvati |first=Katerina |display-authors=etal |title=Apidima Cave fossils provide earliest evidence of Homo sapiens in Eurasia |date=10 July 2019 |journal=[[Nature (journal)|Nature]] |volume=571 |issue=7766 |pages=500–504 |doi=10.1038/s41586-019-1376-z |pmid=31292546 |s2cid=195873640 |url=https://zenodo.org/record/6646855 }}</ref>

A significant dispersal event, within Africa and to West Asia, is associated with the African [[megadrought]]s during [[Marine Isotope Stage 5|MIS 5]], beginning 130,000 years ago.<ref>{{cite journal |last1=Rito |first1=T. |last2=Richards |first2=M. B. |last3=Fernandes |first3=V. |last4=Alshamali |first4=F. |last5=Cerny |first5=V. |last6=Pereira |first6=L. |last7=Soares |first7=P. |title=The first modern human dispersals across Africa |journal=[[PLOS ONE]] |year=2013 |volume=8 |issue=11 |page=e80031 |doi=10.1371/journal.pone.0080031 |pmid=24236171 |pmc=3827445 |bibcode=2013PLoSO...880031R|doi-access=free}}</ref> A 2011 study located the origin of basal population of contemporary human populations at 130,000 years ago, with the Khoi-San representing an "ancestral population cluster" located in southwestern Africa (near the coastal border of [[Namibia]] and [[Angola]]).<ref>{{cite journal |last1=Henn |first1=Brenna |last2=Gignoux |first2=Christopher R. |last3=Jobin |first3=Matthew |year=2011 |title=Hunter-gatherer genomic diversity suggests a southern African origin for modern humans |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=108 |issue=13 |pages=5154–5162 |doi=10.1073/pnas.1017511108 |bibcode=2011PNAS..108.5154H |pmid=21383195 |pmc=3069156|doi-access=free}}</ref>

[[File:Ksar Akil Fossils.jpg|thumb|Layer sequence at [[Ksar Akil]] in the [[Levantine corridor]], and discovery of two fossils of ''Homo sapiens'', dated to 40,800 to 39,200 years BP for "Egbert",<ref name=":1" /> and 42,400–41,700 BP for "Ethelruda".<ref name=":1">{{Cite journal |last1=Higham |first1=Thomas F. G. |last2=Wesselingh |first2=Frank P. |last3=Hedges |first3=Robert E. M. |last4=Bergman |first4=Christopher A. |last5=Douka |first5=Katerina |date=11 September 2013 |title=Chronology of Ksar Akil (Lebanon) and Implications for the Colonization of Europe by Anatomically Modern Humans |journal=[[PLOS ONE]] |language=en |volume=8 |issue=9 |pages=e72931 |doi=10.1371/journal.pone.0072931 |issn=1932-6203 |pmc=3770606 |pmid=24039825 |bibcode=2013PLoSO...872931D |doi-access=free}}</ref>]]
While early modern human expansion in [[Sub-Saharan Africa]] before 130 kya persisted, early expansion to North Africa and Asia appears to have mostly disappeared by the end of MIS5 (75,000 years ago), and is known only from fossil evidence and from [[archaic admixture]]. Eurasia was re-populated by early modern humans in the so-called [[Recent African origin of modern humans|"recent out-of-Africa migration"]] post-dating MIS5, beginning around 70,000–50,000 years ago.<ref>{{harvp|Posth, et al., 2016}}; {{harvp|Kamin, et al., 2015}}; {{harvp|Vai, et al., 2019}}; {{harvp|Haber, et al., 2019}}</ref> In this expansion, bearers of [[Haplogroup L3 (mtDNA)|mt-DNA haplogroup L3]] left East Africa, likely reaching Arabia via the [[Bab-el-Mandeb]], and in the [[Great Coastal Migration]] spread to South Asia, Maritime South Asia and Oceania between 65,000 and 50,000 years ago,<ref>{{cite journal |first1=Chris |last1=Clarkson |first2=Zenobia |last2=Jacobs |first3=Colin |last3=Pardoe |year=2017 |title=Human occupation of northern Australia by 65,000 years ago |journal=[[Nature (journal)|Nature]] |doi=10.1038/nature22968 |pmid=28726833 |volume=547 |issue=7663 |pages=306–310 |bibcode=2017Natur.547..306C |hdl=2440/107043 |s2cid=205257212 |url=https://digital.library.adelaide.edu.au/dspace/bitstream/2440/107043/2/hdl_107043.pdf |archive-url=https://web.archive.org/web/20190428141305/https://digital.library.adelaide.edu.au/dspace/bitstream/2440/107043/2/hdl_107043.pdf |archive-date=2019-04-28 |url-status=live |hdl-access=free}}</ref><ref>{{cite news |last1=St. Fleu |first1=Nicholas |title=Humans First Arrived in Australia 65,000 Years Ago, Study Suggests |url=https://www.nytimes.com/2017/07/19/science/humans-reached-australia-aboriginal-65000-years.html |newspaper=[[The New York Times]] |date=July 19, 2017}}</ref><ref name="Wood_2017">{{Cite journal |last=Wood |first=Rachel |name-list-style=vanc |date=2017-09-02 |title=Comments on the chronology of Madjedbebe |journal=Australian Archaeology |volume=83 |issue=3 |pages=172–174 |doi=10.1080/03122417.2017.1408545 |s2cid=148777016 |issn=0312-2417}}</ref><ref>{{cite journal |vauthors=O'Connell JF, Allen J, Williams MA, Williams AN, Turney CS, Spooner NA, Kamminga J, Brown G, Cooper A |display-authors=6 |title=Homo sapiens first reach Southeast Asia and Sahul? |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=115 |issue=34 |pages=8482–8490 |date=August 2018 |pmid=30082377 |pmc=6112744 |doi=10.1073/pnas.1808385115 |doi-access=free}}</ref> while [[European early modern humans|Europe]], [[East Asia|East]] and [[North Asia]] were reached by about 45,000 years ago. Some evidence suggests that an early wave of humans may have reached [[Pleistocene peopling of the Americas|the Americas]] by about 40,000–25,000 years ago.{{citation needed|date=July 2020}}

Evidence for the overwhelming contribution of this "recent" ([[Haplogroup L3 (mtDNA)|L3]]-derived) expansion to all non-African populations was established based on [[mitochondrial DNA]], combined with evidence based on [[physical anthropology]] of archaic [[Biological specimen|specimens]], during the 1990s and 2000s,{{refn|group=note|"Currently available genetic and archaeological evidence is generally interpreted as supportive of a recent single origin of modern humans in East Africa."<ref>{{cite journal |last1=Liu |first1=Hua |display-authors=etal |year=2006 |title=A Geographically Explicit Genetic Model of Worldwide Human-Settlement History |doi=10.1086/505436 |journal=The American Journal of Human Genetics |volume=79 |issue=2|pages=230–237 |pmid=16826514 |pmc=1559480}}</ref>}}<ref>{{cite journal |title=Out of Africa Revisited |doi=10.1126/science.308.5724.921g |date=2005-05-13 |volume=308 |issue=5724 |journal=Science |page=921g|s2cid=220100436 }}</ref> and has also been supported by [[Y DNA]] and [[autosome|autosomal DNA]].{{sfn|Haber, et al., 2019}} The assumption of complete replacement has been revised in the 2010s with the discovery of [[archaic human admixture with modern humans|admixture events]] ([[introgression]]) of populations of ''H. sapiens'' with populations of archaic humans over the period of between roughly 100,000 and 30,000 years ago, both in Eurasia and in Sub-Saharan Africa. [[Neanderthal admixture]], in the range of 1–4%, is found in all modern populations outside of Africa, including in Europeans, Asians, Papua New Guineans, Australian Aboriginals, Native Americans, and other non-Africans.<ref name="SankararamanMallick2016">{{cite journal|last1=Sankararaman |first1=Sriram |last2=Mallick |first2=Swapan |last3=Patterson |first3=Nick |last4=Reich |first4=David |author4-link=David Reich (geneticist) |title=The Combined Landscape of Denisovan and Neanderthal Ancestry in Present-Day Humans |journal=Current Biology |volume=26 |issue=9 |year=2016 |pages=1241–1247 |issn=0960-9822 |doi=10.1016/j.cub.2016.03.037 |pmid=27032491 |pmc=4864120|bibcode=2016CBio...26.1241S }}</ref><ref name=Draft /> This suggests that interbreeding between Neanderthals and anatomically modern humans took place after the [[Southern Dispersal|recent "out of Africa" migration]], likely between 60,000 and 40,000 years ago.<ref>{{cite journal |date=October 17, 2012|title=North African Populations Carry the Signature of Admixture with Neandertals |journal=[[PLOS ONE]] |doi=10.1371/journal.pone.0047765 |volume=7 |issue=10 |page=e47765 |pmid=23082212 |pmc=3474783 |last1=Sánchez-Quinto |first1=F |last2=Botigué |first2=LR |last3=Civit |first3=S |last4=Arenas |first4=C |last5=Avila-Arcos |first5=MC |last6=Bustamante |first6=CD |last7=Comas |first7=D |last8=Lalueza-Fox |first8=C |bibcode=2012PLoSO...747765S|doi-access=free }}</ref><ref>{{cite journal |date=October 23, 2014 |title= Genome sequence of a 45,000-year-old modern human from western Siberia |journal=[[Nature (journal)|Nature]] |volume=514 |issue= 7523 |pages= 445–449 |doi= 10.1038/nature13810 |pmid=25341783 |pmc=4753769 |last1=Fu |first1=Q |last2=Li |first2=H |last3=Moorjani |first3=P |last4=Jay |first4=F |last5=Slepchenko |first5=SM |last6=Bondarev |first6=AA |last7=Johnson |first7=PL |last8=Aximu-Petri |first8=A |last9=Prüfer |first9=K |last10=de Filippo |first10=C |last11=Meyer |first11=M |last12=Zwyns |first12=N |last13=Salazar-García |first13=DC |last14=Kuzmin |first14=YV |last15=Keates |first15=SG |last16=Kosintsev |first16=PA |last17=Razhev |first17=DI |last18=Richards |first18=MP |last19=Peristov |first19=NV |last20=Lachmann |first20=M |last21=Douka |first21=K |last22=Higham |first22=TF |last23=Slatkin |first23=M |last24=Hublin |first24=JJ |last25=Reich |first25=D |last26=Kelso |first26=J |last27=Viola |first27=TB |last28=Pääbo |first28=S |bibcode= 2014Natur.514..445F}}</ref><ref>{{cite magazine |last=Brahic |first=Catherine |url= https://www.newscientist.com/article/dn24988-humanitys-forgotten-return-to-africa-revealed-in-dna/ |title=Humanity's forgotten return to Africa revealed in DNA |magazine=[[The New Scientist]] |date=February 3, 2014 |access-date=2019-05-06}}</ref> Recent admixture analyses have added to the complexity, finding that Eastern Neanderthals derive up to 2% of their ancestry from anatomically modern humans who left Africa some 100 [[wikt:kya|kya]].<ref name="nature.com">{{cite journal |last1=Kuhlwilm |first1=Martin |title=Ancient gene flow from early modern humans into Eastern Neanderthals |journal=Nature |date=17 February 2016 |volume=530 |issue=7591 |pages=429–433 |doi=10.1038/nature16544 |pmid=26886800 |pmc=4933530 |bibcode=2016Natur.530..429K }}</ref> The extent of [[Neanderthal admixture]] (and [[introgression]] of genes acquired by admixture) varies significantly between contemporary racial groups, being absent in Africans, intermediate in Europeans and highest in East Asians. Certain genes related to UV-light adaptation introgressed from Neanderthals have been found to have been selected for in East Asians specifically from 45,000 years ago until around 5,000 years ago.<ref name=dinch3>{{cite journal |last=Ding |first=Q. |author2=Hu, Y. |author3= Xu, S. |author4= Wang, J. |author5= Jin, L. |title=Neanderthal Introgression at Chromosome 3p21.31 was Under Positive Natural Selection in East Asians |journal=[[Molecular Biology and Evolution]] |year=2014 |orig-date=Online 2013 |volume=31 |issue=3 |pages=683–695 |doi=10.1093/molbev/mst260 |pmid=24336922|doi-access=free}}</ref> The extent of archaic admixture is of the order of about 1% to 4% in Europeans and East Asians, and highest among [[Melanesians]] (the last also having [[Denisova hominin]] admixture at 4% to 6% in addition to neanderthal admixture).<ref name=Draft /><ref name="Reich et al." /> Cumulatively, about 20% of the Neanderthal genome is estimated to remain present spread in contemporary populations.<ref name=vern14res>{{cite journal |last=Vernot |first=B. |author2=Akey, J. M. |title=Resurrecting Surviving Neandertal Lineages from Modern Human Genomes |journal=[[Science (journal)|Science]] |date=2014 |volume=343 |issue=6174 |pages=1017–1021 |doi=10.1126/science.1245938 |pmid=24476670 |bibcode=2014Sci...343.1017V |s2cid=23003860|doi-access=free }}</ref>

In September 2019, scientists reported the computerized determination, based on 260 [[CT scan]]s, of a virtual [[Human skull|skull shape]] of the last common human ancestor to modern humans/''H. sapiens'', representative of the earliest modern humans, and suggested that modern humans arose between 350,000 and 260,000 years ago through a merging of populations in [[East Africa|East]] and [[South Africa]] while [[North Africa|North-African]] fossils may represent a population which introgressed into Neandertals during the LMP.<ref name="NYT-20190910" /><ref name="NAT-20190910" />

According to a study published in 2020, there are indications that 2% to 19% (or about ≃6.6 and ≃7.0%) of the DNA of four [[West Africa]]n populations may have come from an unknown archaic hominin which split from the ancestor of humans and Neanderthals between 360 kya to 1.02 mya.<ref>{{Cite journal|doi=10.1126/sciadv.aax5097|title=Recovering signals of ghost archaic introgression in African populations|year=2020|last1=Durvasula|first1=Arun|last2=Sankararaman|first2=Sriram|journal=Science Advances|volume=6|issue=7|pages=eaax5097|pmid=32095519|pmc=7015685|bibcode=2020SciA....6.5097D}}</ref>

== Anatomy ==
{{see also|Human anatomy|Human physical appearance|Anthropometry}}
[[File:Anatomically Modern Humans archaeological remains, Europe and Africa, directly dated, calibrated carbon dates as of 2013.jpg|thumb|Known archaeological remains of anatomically modern humans in Europe and Africa, directly dated, calibrated carbon dates as of 2013.<ref name=":1"/>]]
Generally, modern humans are more lightly built (or more "gracile") than the more "robust" [[archaic humans]]. Nevertheless, contemporary humans exhibit high [[Human physical appearance#Physiological differences|variability in many physiological traits]], and may exhibit remarkable "robustness". There are still a number of physiological details which can be taken as reliably differentiating the physiology of [[Neanderthals]] vs. anatomically modern humans.

=== Anatomical modernity ===
The term "anatomically modern humans" (AMH) is used with varying scope depending on context, to distinguish "anatomically modern" ''Homo sapiens'' from [[archaic humans]] such as Neanderthals and Middle and [[Lower Paleolithic]] hominins with transitional features intermediate between ''H. erectus'', Neanderthals and early AMH called ''archaic Homo sapiens''.<ref>{{Cite book |url=https://books.google.com/books?id=1OU-DgAAQBAJ&pg=PA358 |title=Processes in Human Evolution: The Journey from Early Hominins to Neanderthals and Modern Humans |isbn=978-0198739906 |last1=Ayala |first1=Francisco José |last2=Conde |first2=Camilo José Cela |year=2017 |publisher=Oxford University Press |via=[[Google Books]]}}</ref> In a convention popular in the 1990s, Neanderthals were classified as a [[human subspecies|subspecies]] of ''H. sapiens'', as ''H. s. neanderthalensis'', while AMH (or [[European early modern humans]], EEMH) was taken to refer to "[[Cro-Magnon]]" or ''H. s. sapiens''. Under this nomenclature (Neanderthals considered ''H. sapiens''), the term "anatomically modern ''Homo sapiens''" (AMHS) has also been used to refer to EEMH ("Cro-Magnons").<ref>{{cite book |last=Schopf |first=J. William |title=Major Events in the History of Life|url= https://books.google.com/books?id=py01HMuAIh4C&pg=PA168 |year=1992 |publisher=Jones & Bartlett Learning |isbn=978-0867202687 |pages=168– |via=[[Google Books]]}}</ref> It has since become more common to designate Neanderthals as a separate species, ''H. neanderthalensis'', so that AMH in the European context refers to ''H. sapiens'', but the question is by no means resolved.{{refn|group=note|This is a question of conventional terminology, not one of a factual disagreement. Pääbo (2014) frames this as a debate that is unresolvable in principle, "since there is no definition of species perfectly describing the case."<ref>{{cite book |last=Pääbo |first=Svante |author-link=Svante Pääbo |title=Neanderthal Man: In Search of Lost Genomes |publisher=[[Basic Books]] |location=New York |year=2014 |page=237}}</ref>}}

In this more narrow definition of ''H. sapiens'', the subspecies ''[[Homo sapiens idaltu]]'', discovered in 2003, also falls under the umbrella of "anatomically modern".<ref>{{cite web |url=https://www.berkeley.edu/news/media/releases/2003/06/11_idaltu.shtml |first=Robert |last=Sanders |publisher=UC Berkeley News |title=160,000-year-old fossilized skulls uncovered in Ethiopia are oldest anatomically modern humans |date=11 June 2003 |access-date=2019-05-07}}</ref> The recognition of ''H. sapiens idaltu'' as a [[human subspecies|valid subspecies]] of the anatomically modern human lineage would justify the description of contemporary humans with the subspecies name ''Homo sapiens sapiens''.<ref name="White03">{{Cite journal |last1=White |first1=Tim D. |author-link=Tim White (anthropologist) |last2=Asfaw |first2=B. |last3=DeGusta |first3=D. |last4=Gilbert |first4=H. |last5=Richards |first5=G. D. |last6=Suwa |first6=G. |last7=Howell |first7=F. C. |year=2003 |title=Pleistocene ''Homo sapiens'' from Middle Awash, Ethiopia |journal=[[Nature (journal)|Nature]] |volume=423 |issue=6491 |pages=742–747 |bibcode=2003Natur.423..742W |doi=10.1038/nature01669 |pmid=12802332 |s2cid=4432091}}</ref> However, biological anthropologist [[Chris Stringer]] does not consider ''idaltu'' distinct enough within ''H. sapiens'' to warrant its own subspecies designation.<ref>{{Cite journal |title= Human evolution: Out of Ethiopia |journal=[[Nature (journal)|Nature]] |volume=423 |issue=6941 |pages=693–695 |date=12 June 2003 |bibcode=2003Natur.423..692S |last1= Stringer |first1=Chris |doi=10.1038/423692a |pmid=12802315 |s2cid=26693109}}</ref>{{sfn|Stringer|2016|p=20150237}}

A further division of AMH into "early" or "robust" vs. "post-glacial" or "[[gracile]]" subtypes has since been used for convenience. The emergence of "gracile AMH" is taken to reflect a process towards a smaller and more fine-boned skeleton beginning around 50,000–30,000 years ago.<ref>{{cite journal |doi=10.1073/pnas.0707650104 |title=Recent acceleration of human adaptive evolution |journal=[[Proceedings of the National Academy of Sciences]] |volume=104 |issue=52 |pages=20753–20758 |year=2007 |last1=Hawks |first1=J. |last2=Wang |first2=E. T. |last3=Cochran |first3=G. M. |last4=Harpending |first4=H. C. |last5=Moyzis |first5=R. K. |bibcode=2007PNAS..10420753H |pmid=18087044 |pmc=2410101 |doi-access=free}}</ref>

=== Braincase anatomy ===
{{further|Brain size}}
[[File:Sapiens neanderthal comparison en.png|thumb|upright=1.35|Anatomical comparison of skulls of ''H. sapiens'' (left) and ''H. neanderthalensis'' (right)<br />(in [[Cleveland Museum of Natural History]])<br />Features compared are the [[neurocranium|braincase]] shape, [[forehead]], [[browridge]], [[nasal bone|nasal bone projection]], [[cheek bone|cheek bone angulation]], [[chin]] and [[occipital bone|occipital contour]]]]
The cranium lacks a pronounced [[occipital bun]] in the neck, a bulge that anchored considerable neck muscles in Neanderthals. Modern humans, even the earlier ones, generally have a larger fore-brain than the archaic people, so that the brain sits above rather than behind the eyes. This will usually (though not always) give a higher forehead, and reduced [[supraorbital ridge|brow ridge]]. Early modern people and some living people do however have quite pronounced brow ridges, but they differ from those of archaic forms by having both a [[supraorbital foramen]] or notch, forming a groove through the ridge above each eye.<ref>{{cite journal|last=Bhupendra |first=P. |title=Forehead Anatomy|url= https://emedicine.medscape.com/article/834862-overview |website=Medscape references |access-date=2019-05-06|date=April 2019 }}</ref> This splits the ridge into a central part and two distal parts. In current humans, often only the central section of the ridge is preserved (if it is preserved at all). This contrasts with archaic humans, where the brow ridge is pronounced and unbroken.<ref>{{cite web|title=How to ID a modern human?|url= http://www.nhm.ac.uk/about-us/news/2012/may/how-to-id-a-modern-human109960.html |website=News, 2012|publisher=[[Natural History Museum, London]]|access-date=11 December 2013}}</ref>

Modern humans commonly have a steep, even vertical [[forehead]] whereas their predecessors had foreheads that sloped strongly backwards.<ref>{{cite encyclopedia|title=Encarta, Human Evolution |url=http://encarta.msn.com/encyclopedia_761566394_9/human_evolution.html |encyclopedia= Encarta |archive-url= https://web.archive.org/web/20091029044339/http://encarta.msn.com/encyclopedia_761566394_9/Human_Evolution.html |archive-date=29 October 2009 |url-status=dead |df=dmy }}</ref> According to [[Desmond Morris]], the vertical forehead in humans plays an important role in human communication through [[eyebrow]] movements and forehead skin wrinkling.<ref>{{cite book|title=The Naked Woman: A Study of the Female Body |chapter=The Brow |chapter-url= https://books.google.com/books?id=Wa9zntiEKeAC&pg=PA22 |first=Desmond |last=Morris |author-link=Desmond Morris |year=2007 |publisher=Macmillan |isbn=978-0312338534}}</ref>

[[Brain size]] in both Neanderthals and AMH is significantly larger on average (but overlapping in range) than brain size in ''H. erectus''. Neanderthal and AMH brain sizes are in the same range, but there are differences in the relative sizes of individual brain areas, with significantly larger visual systems in Neanderthals than in AMH.<ref>{{Cite journal|last1=Pearce |first1=Eiluned |last2=Stringer |first2=Chris |last3=Dunbar |first3=R. I. M. |date=2013-05-07 |title=New insights into differences in brain organization between Neanderthals and anatomically modern humans|journal=Proceedings of the Royal Society of London B: Biological Sciences |volume=280 |issue=1758 |page=20130168 |doi=10.1098/rspb.2013.0168 |issn=0962-8452 |pmc=3619466 |pmid=23486442}}</ref>{{refn|group=note|Contemporary human endocranial volume averages at {{cvt|1350|cm3}}, with significant differences between populations, global group means range {{cvt|1085|-|1580|cm3}}.<ref>{{cite journal|last1=Smith |first1=C. L. |last2=Beals |first2=K. L. |title=Cultural correlates with cranial capacity |journal=American Anthropologist |date=1990 |volume=92 |pages=193–200 |doi=10.1525/aa.1990.92.1.02a00150|s2cid=162406199 |s2cid-access=free}}</ref> Neanderthal average is close to {{cvt|1450|cm3}} (male average {{cvt|1600|cm3}}, female average {{cvt|1300|cm3}}), with a range extending up to {{cvt|1736|cm3}} ([[Amud 1]]).<ref>{{cite encyclopedia |last=Stringer |first=C |editor-last=Foley |editor-first=R |encyclopedia=Hominid evolution and community ecology |title=Human evolution and biological adaptation in the Pleistocene |year=1984 |publisher=Academic Press |location=New York |isbn=978-0122619205}}</ref>}}

===Jaw anatomy===
Compared to archaic people, anatomically modern humans have smaller, differently shaped teeth.<ref name="Townsend G, Richards L, Hughes T 2003 350–5">{{Cite journal|vauthors=Townsend G, Richards L, Hughes T |title=Molar intercuspal dimensions: genetic input to phenotypic variation |journal=Journal of Dental Research |volume=82 |issue=5 |pages=350–355 |date=May 2003 |pmid=12709500 |doi=10.1177/154405910308200505|s2cid=26123427 }}</ref><ref>{{Cite journal|author=Keith A |title=Problems relating to the Teeth of the Earlier Forms of Prehistoric Man |journal=Proceedings of the Royal Society of Medicine |volume=6 |issue=Odontol Sect |pages=103–124 |year=1913 |pmid=19977113 |pmc=2005996|doi=10.1177/003591571300601018 }}</ref> This results in a smaller, more receded dentary, making the rest of the jaw-line stand out, giving an often quite prominent chin. The central part of the mandible forming the chin carries a triangularly shaped area forming the apex of the chin called the '''[[Chin|mental Trigon]]''', not found in archaic humans.<ref>{{cite book|last1=Tattersall |first1=Jeffrey H |last2=Schwartz |first2=Ian |title=The human fossil record Craniodental Morphology of Genus Homo (Africa and Asia) (vol 2) |date=2003 |publisher=Wiley-Liss |isbn=978-0471319283 |pages=327–328}}</ref> Particularly in living populations, the use of fire and tools requires fewer jaw muscles, giving slender, more gracile jaws. Compared to archaic people, modern humans have smaller, lower faces.

===Body skeleton structure===
The body skeletons of even the earliest and most robustly built modern humans were less robust than those of Neanderthals (and from what little we know from Denisovans), having essentially modern proportions. Particularly regarding the long bones of the limbs, the distal bones (the [[Radius (bone)|radius]]/[[ulna]] and [[tibia]]/[[fibula]]) are nearly the same size or slightly shorter than the proximal bones (the [[humerus]] and [[femur]]). In ancient people, particularly Neanderthals, the distal bones were shorter, usually thought to be an adaptation to cold climate.<ref>{{cite journal|last=Steegmann|first=A. Theodore|author2=Cerny, Frank J.|author3= Holliday, Trenton W.|title=Neandertal cold adaptation: Physiological and energetic factors|journal=American Journal of Human Biology|year=2002|volume=14|issue=5|pages=566–583|doi=10.1002/ajhb.10070|pmid=12203812|s2cid=2437566}}</ref> The same adaptation is found in some modern people living in the polar regions.<ref>{{cite journal|last=Stock|first=J.T.|title=Hunter-gatherer postcranial robusticity relative to patterns of mobility, climatic adaptation, and selection for tissue economy|journal=American Journal of Physical Anthropology|date=October 2006|volume=131|issue=2|pages=194–204|doi=10.1002/ajpa.20398|pmid=16596600}}</ref>

[[Human height|Height]] ranges overlap between Neanderthals and AMH, with Neanderthal averages cited as {{cvt|164|to|168|cm|0}} and {{cvt|152|to|156|cm|0}} for males and females, respectively, which is largely identical to pre-industrial average heights for AMH.{{refn|group=note|"Based on 45 long bones from maximally 14 males and 7 females, Neanderthals' height averages between 164 and 168 (males) resp. 152 to 156 cm (females). This height is indeed 12–14 cm lower than the height of post-WWII Europeans, but compared to Europeans some 20,000 or 100 years ago, it is practically identical or even slightly higher."<ref name="Helmuth1998">{{Cite journal|author=Helmuth H |title=Body height, body mass and surface area of the Neanderthals |journal=Zeitschrift für Morphologie und Anthropologie |volume=82 |issue=1 |pages=1–12 |year=1998 |doi=10.1127/zma/82/1998/1 |doi-broken-date=1 November 2024 |pmid=9850627 }}</ref>}} [[List of average human height worldwide|Contemporary national averages]] range between {{cvt|158|to|184|cm|0}} in males and {{cvt|147|to|172|cm|0}} in females. Neanderthal ranges approximate the contemporary height distribution measured among [[Malay people]], for one.{{refn|group=note|Malay, 20–24 (N{{=}} m:749 f:893, Median{{=}} m:{{Height|cm=166}} f:{{Height|cm=155}}, SD{{=}} m:{{Height|cm=6.46}} f:{{Height|cm=6.04}})<ref>{{cite journal|url= http://www.e-mjm.org/2000/v55n1/Body_Weight.pdf |archive-url=https://web.archive.org/web/20170224180141/http://www.e-mjm.org/2000/v55n1/Body_Weight.pdf |archive-date=2017-02-24 |url-status=live |title=Distribution of Body Weight, Height and Body Mass Index in a National Sample of Malaysian Adults |pmid=11072496 |volume=55 |issue=1 |date=March 2000 |journal=Med. J. Malaysia |pages=108–128 |vauthors=Lim TO, Ding LM, Zaki M, etal}}</ref>}}

==Recent evolution==
{{main|Recent human evolution}}
{{further|Human genetic variation|Race and genetics|Sexual selection in humans}}
[[File:Em - Homo sapiens man model - 3.jpg|thumb|210px|Reconstruction of a modern man from southwestern Europe {{c.|30 000 years BP}}, London Natural History Museum.]]
Following the [[peopling of Africa]] some 130,000 years ago, and the [[Coastal migration|recent Out-of-Africa]] expansion some 70,000 to 50,000 years ago, some sub-populations of ''H. sapiens'' had been essentially [[geographic isolation|isolated]] for tens of thousands of years prior to the early modern [[Age of Discovery]]. Combined with [[Archaic human admixture with modern humans|archaic admixture]] this has resulted in significant [[human genetic variation|genetic variation]], which in some instances has been shown to be the result of [[directional selection]] taking place over the past 15,000 years, i.e., significantly later than possible archaic admixture events.<ref>{{cite news|url= https://www.nytimes.com/2006/03/07/science/07evolve.html |title=Still Evolving, Human Genes Tell New Story |last=Wade |first=N |date=2006-03-07 |access-date=2008-07-10 |newspaper=The New York Times }}</ref>

Some climatic adaptations, such as [[high-altitude adaptation in humans]], are thought to have been acquired by archaic admixture. [[Introgression]] of genetic variants acquired by [[Neanderthal admixture]] have different distributions in [[Caucasoid|European]] and [[Mongoloid|East Asians]], reflecting differences in recent selective pressures. A 2014 study reported that Neanderthal-derived variants found in East Asian populations showed clustering in functional groups related to [[Complement system|immune]] and [[Haematopoiesis|haematopoietic pathways]], while European populations showed clustering in functional groups related to the [[Catabolism|lipid catabolic process]].{{refn|group=note|"Specifically, genes in the LCP [lipid catabolic process] term had the greatest excess of NLS in populations of European descent, with an average NLS frequency of 20.8±2.6% versus 5.9±0.08% genome wide (two-sided t-test, P<0.0001, n{{=}}379 Europeans and n{{=}}246 Africans). Further, among examined out-of-Africa human populations, the excess of NLS [Neanderthal-like genomic sites] in LCP genes was only observed in individuals of European descent: the average NLS frequency in Asians is 6.7±0.7% in LCP genes versus 6.2±0.06% genome wide."<ref>{{cite journal|first1=E |last1=Khrameeva |first2=K |last2=Bozek |first3=L |last3=He |first4=Z |last4=Yan |first5=X |last5=Jiang |first6=Y |last6=Wei |first7=K |last7=Tang |first8=MS |last8=Gelfand |first9=K |last9=Prüfer |first10=J |last10=Kelso |first11=S |last11=Pääbo |first12=P |last12=Giavalisco |first13=M |last13=Lachmann |first14=P |last14=Khaitovich |title=Neanderthal ancestry drives evolution of lipid catabolism in contemporary Europeans |journal=Nature Communications |volume=5 |page=3584 |number=3584 |year=2014 |doi=10.1038/ncomms4584|pmid=24690587 |pmc=3988804 |bibcode=2014NatCo...5.3584K }}</ref>}} A 2017 study found correlation of [[Neanderthal admixture]] in phenotypic traits in modern European populations.<ref>Michael Dannemann 1 and Janet Kelso, "The Contribution of Neanderthals to Phenotypic Variation in Modern Humans", ''The American Journal of Human Genetics 101'', 578–589, October 5, 2017.</ref>

Physiological or phenotypical changes have been traced to Upper Paleolithic mutations, such as the East Asian variant of the ''[[EDAR]]'' gene, dated to c. 35,000 years ago.{{refn|group=note|Traits affected by the mutation are sweat glands, teeth, hair thickness and breast tissue.<ref>{{cite journal|last1=Kamberov |first1=Yana G |url= |title=Modeling Recent Human Evolution in Mice by Expression of a Selected EDAR Variant |journal=Cell |volume=152 |issue=4 |pages=691–702 |date=14 February 2013 |doi=10.1016/j.cell.2013.01.016|pmid=23415220 |pmc=3575602 }}</ref><ref>{{cite news|url= https://www.nytimes.com/2013/02/15/science/studying-recent-human-evolution-at-the-genetic-level.html |title=East Asian Physical Traits Linked to 35,000-Year-Old Mutation |newspaper=The New York Times |date=14 February 2013 |last=Wade |first=Nicholas |access-date=2019-05-06}}</ref>}}

Recent divergence of Eurasian lineages was sped up significantly during the [[Last Glacial Maximum]] (LGM), the [[Mesolithic]] and the [[Neolithic]], due to increased selection pressures and due to [[founder effect]]s associated with [[prehistoric migrations|migration]].<ref name=Beleza2012>{{cite journal |title=The timing of pigmentation lightening in Europeans |journal=Molecular Biology and Evolution |year=2012 |doi=10.1093/molbev/mss207 |pmid=22923467 |pmc=3525146 |volume=30 |issue=1 |pages=24–35 |last1=Beleza |first1=Sandra |last2=Santos |first2=A. M. |last3=McEvoy |first3=B. |last4=Alves |first4=I. |last5=Martinho |first5=C. |last6=Cameron |first6=E. |last7=Shriver |first7=M. D. |last8=Parra |first8=E. J. |last9=Rocha |first9=J.}}</ref> Alleles predictive of [[light skin]] have been found in [[Neanderthal genome|Neanderthals]],<ref name=Lalu>{{cite journal|last1=Lalueza-Fox |title=A melanocortin-1 receptor allele suggests varying pigmentation among Neanderthals |journal=Science |year=2007 |volume=318 |issue=5855 |pages=1453–1455 |pmid=17962522 |last2=Römpler |first2=H |last3=Caramelli |first3=D |last4=Stäubert |first4=C |last5=Catalano |first5=G |last6=Hughes |first6=D |last7=Rohland |first7=N |last8=Pilli |first8=E |last9=Longo |first9=L |last10=Condemi |first10=S |last11=de la Rasilla |first11=M |last12=Fortea |first12=J |last13=Rosas |first13=A |last14=Stoneking |first14=M |last15=Schöneberg |first15=T |last16=Bertranpetit |first16=J |last17=Hofreiter |first17=M |doi=10.1126/science.1147417 |display-authors=etal |bibcode=2007Sci...318.1453L|s2cid=10087710 |s2cid-access=free}}</ref> but the alleles for light skin in Europeans and East Asians, associated with [[KITLG]] and [[Agouti signalling peptide|ASIP]], are ({{as of|2012|lc=y}}) thought to have not been acquired by archaic admixture but recent mutations since the LGM.<ref name=Beleza2012 /> Phenotypes associated with the "[[white people|white]]" or "[[Caucasian race|Caucasian]]" populations of Western Eurasian stock emerge during the LGM, from about 19,000 years ago. Average [[cranial capacity]] in modern human populations varies in the range of 1,200 to 1,450&nbsp;cm<sup>3</sup> for adult males. Larger cranial volume is associated with climatic region, the largest averages being found in populations of [[Siberia]] and the [[Arctic]].{{refn|group=note|"We offer an alternative hypothesis that suggests that hominid expansion into regions of cold climate produced change in head shape. Such change in shape contributed to the increased cranial volume. Bioclimatic effects directly upon body size (and indirectly upon brain size) in combination with cranial globularity appear to be a fairly powerful explanation of ethnic group differences." (figure in Beals, p304)<ref name=Beals1984>{{cite journal|first1=Kenneth L |last1=Beals |first2=Courtland L |last2=Smith |first3=Stephen M |last3=Dodd |title=Brain Size, Cranial Morphology, Climate, and Time Machines |journal=Current Anthropology |volume=25 |number=3 |year=1984 |pages=301–330 |doi=10.1086/203138|s2cid=86147507 }}</ref>}}<ref>{{Cite journal|url= http://connection.ebscohost.com/c/articles/69706893/morphological-adaptation-climate-modern-homo-sapiens-crania-importance-basicranial-breadth |archive-url= https://web.archive.org/web/20141011075329/http://connection.ebscohost.com/c/articles/69706893/morphological-adaptation-climate-modern-homo-sapiens-crania-importance-basicranial-breadth |url-status= dead |archive-date= 2014-10-11 |title=Morphological Adaptation to Climate in Modern Homo sapiens Crania: The Importance of Basicranial Breadth |first1=Wioletta |last1=Nowaczewska |first2=Pawel |last2=Dabrowski |last3=Kuźmiński |first3=Lukasz |journal=Collegium Antropologicum |year=2011 |volume=35 |issue=3 |pages=625–636 |pmid=22053534 }}</ref> Both [[Neanderthal]] and [[EEMH]] had somewhat larger cranial volumes on average than modern Europeans, suggesting the relaxation of selection pressures for larger brain volume after the end of the LGM.<ref name=Beals1984 />

Examples for still later adaptations related to [[Neolithic Revolution|agriculture]] and [[animal domestication]] including [[East Asian race|East Asian]] types of [[ADH1B]] associated with [[Oryza sativa#Continental East Asia|rice domestication]],<ref>{{cite journal | last1 = Peng | first1 = Y. | display-authors = etal | year = 2010 | title = The ADH1B Arg47His polymorphism in East Asian populations and expansion of rice domestication in history | journal = BMC Evolutionary Biology | volume = 10 | issue = 1 | page = 15 | doi = 10.1186/1471-2148-10-15 | pmid = 20089146 | pmc = 2823730 | doi-access = free | bibcode = 2010BMCEE..10...15P }}</ref> or [[lactase persistence]],<ref>{{Cite journal|last1=Ségurel |first1=Laure |last2=Bon |first2=Céline |date=2017 |title=On the Evolution of Lactase Persistence in Humans |journal=Annual Review of Genomics and Human Genetics |volume=18 |issue=1 |pages=297–319 |doi=10.1146/annurev-genom-091416-035340 |pmid=28426286}}</ref><ref>{{Cite journal|last1=Ingram |first1=Catherine J. E. |last2=Mulcare |first2=Charlotte A. |last3=Itan |first3=Yuval |last4=Thomas |first4=Mark G. |last5=Swallow |first5=Dallas M. |date=2008-11-26 |title=Lactose digestion and the evolutionary genetics of lactase persistence |journal=Human Genetics |language=en |volume=124 |issue=6 |pages=579–591 |doi=10.1007/s00439-008-0593-6 |pmid=19034520 |s2cid=3329285 |issn=0340-6717}}</ref> are due to recent selection pressures.

An even more recent adaptation has been proposed for the Austronesian [[Sama-Bajau]], developed under selection pressures associated with subsisting on [[freediving]] over the past thousand years or so.<ref>{{cite journal|last1= Ilardo |first1=M. A. |last2= Moltke |first2= I. |last3= Korneliussen |first3=T. S. |last4= Cheng |first4= J. |last5= Stern |first5=A. J. |last6= Racimo |first6= F. |last7=de Barros Damgaard |first7= P. |last8= Sikora |first8= M. |last9= Seguin-Orlando |first9= A. |last10= Rasmussen |first10= S. |last11=van den Munckhof |first11=I. C. L. |last12=ter Horst |first12= R. |last13= Joosten |first13=L. A. B. |last14=Netea |first14=M. G. |last15=Salingkat |first15=S. |last16=Nielsen |first16=R. |last17=Willerslev |first17=E. |title=Physiological and Genetic Adaptations to Diving in Sea Nomads |journal= Cell |volume= 173 |issue= 3 |date= 2018-04-18 |pages= 569–580.e15 |doi= 10.1016/j.cell.2018.03.054 |pmid=29677510|doi-access= free }}</ref><ref>{{cite journal|last1=Gislén |first1=A |last2=Dacke |first2=M |last3=Kröger |first3=RH |last4=Abrahamsson |first4=M |last5=Nilsson |first5=DE |last6=Warrant |first6=EJ |title=Superior Underwater Vision in a Human Population of Sea Gypsies |journal=Current Biology |year=2003 |volume=13 |issue=10 |pages=833–836 |doi=10.1016/S0960-9822(03)00290-2|pmid=12747831 |s2cid=18731746 |doi-access=free |bibcode=2003CBio...13..833G }}</ref>

==Behavioral modernity==
{{main|Behavioral modernity}}
[[File:Lithic Industries at Blombos Cave, Southern Cape, South Africa (c. 105 – 90 Ka).jpg|thumb|upright=1.5|Lithic Industries of early ''Homo sapiens'' at [[Blombos Cave]] (M3 phase, MIS 5), Southern Cape, South Africa (c. 105,000 – 90,000 years old)]]
[[Behavioral modernity]], involving the development of [[origin of language|language]], [[Paleolithic Art|figurative art]] and early forms of [[Paleolithic religion|religion]] (etc.) is taken to have arisen before 40,000 years ago, marking the beginning of the [[Upper Paleolithic]] (in African contexts also known as the [[Later Stone Age]]).<ref name="Klein 1995">{{cite journal |last=Klein |first=Richard |title=Anatomy, behavior, and modern human origins |journal=Journal of World Prehistory |date=1995 |volume=9 |issue=2 |pages=167–198 |doi=10.1007/bf02221838|s2cid=10402296 }}</ref>

There is considerable debate regarding whether the earliest anatomically modern humans behaved similarly to recent or existing humans. [[Behavioral modernity]] is taken to include fully developed [[Origin of language|language]] (requiring the capacity for [[abstract thought]]), [[Art of the Upper Paleolithic|artistic expression]], early forms of [[Paleolithic religion|religious behavior]],<ref>{{cite book|author=Feierman, Jay R. |page=220|title=The Biology of Religious Behavior: The Evolutionary Origins of Faith and Religion|url=https://books.google.com/books?id=mOLGXhzAXhsC |year=2009|publisher=ABC-CLIO|isbn=978-0313364303}}</ref> increased cooperation and the formation of early settlements, and the production of articulated tools from [[lithic core]]s, bone or antler. The term [[Upper Paleolithic]] is intended to cover the period since the [[Coastal migration|rapid expansion]] of modern humans throughout Eurasia, which coincides with the first appearance of [[Paleolithic art]] such as [[cave paintings]] and the development of technological innovation such as the [[spear-thrower]]. The Upper Paleolithic begins around 50,000 to 40,000 years ago, and also coincides with the disappearance of archaic humans such as the [[Neanderthal]]s.

[[File:Blombos point white.JPG|thumb|left|Bifacial silcrete point of early ''Homo sapiens'', from M1 phase (71,000 BCE) layer of [[Blombos Cave]], South Africa]]
The term "behavioral modernity" is somewhat disputed. It is most often used for the set of characteristics marking the Upper Paleolithic, but some scholars use "behavioral modernity" for the emergence of ''H. sapiens'' around 200,000 years ago,<ref>Soressi M. (2005) [http://www.eva.mpg.de/evolution/staff/soressi/pdf/Soressi2005_ToolsToSymbols.pdf Late Mousterian lithic technology. Its implications for the pace of the emergence of behavioural modernity and the relationship between behavioural modernity and biological modernity], pp. 389–417 in L. Backwell et F. d'Errico (eds.) ''From Tools to Symbols'', Johannesburg: University of Witswatersand Press. {{ISBN|1868144178}}.</ref> while others use the term for the rapid developments occurring around 50,000 years ago.<ref>''Companion encyclopedia of archaeology'' (1999). Routledge. {{ISBN|0415213304}}. Vol. 2. p. 763 (''cf''., ... "effectively limited to [[Organic matter|organic samples]]" [ed. [[organic compound]]s ] "or [[biogenic|biogenic carbonate]]s that date to less than 50 ka (50,000 years ago)."). See also: [[Later Stone Age]] and [[Upper Paleolithic]].</ref><ref name="mellars">{{cite journal |author-link=Paul Mellars |last=Mellars |first=Paul |title=Why did modern human populations disperse from Africa ca. 60,000 years ago? |year=2006 |doi=10.1073/pnas.0510792103 |journal=Proceedings of the National Academy of Sciences |volume=103 |pages=9381–9386 |pmid=16772383 |issue=25 |pmc=1480416 |bibcode=2006PNAS..103.9381M|doi-access=free }}</ref><ref>{{cite journal |last1=Shea |first1=John |title=Homo sapiens Is As Homo sapiens Was |journal=Current Anthropology |date=2011 |volume=52 |issue=1 |pages=1–35 |doi=10.1086/658067|s2cid=142517998 }}</ref> It has been proposed that the emergence of behavioral modernity was a gradual process.<ref>{{cite journal |last1=McBrearty |first1=Sally |last2=Brooks |first2=Allison |date=2000 |title=The revolution that wasn't: a new interpretation of the origin of modern human behavior |journal=Journal of Human Evolution |volume=39 |issue=5 |pages=453–563 |doi=10.1006/jhev.2000.0435 |pmid=11102266|bibcode=2000JHumE..39..453M |s2cid=42968840 }}</ref><ref>{{cite journal |last1=Henshilwood |first1=Christopher |last2=Marean |first2=Curtis |date=2003 |title=The Origin of Modern Human Behavior: Critique of the Models and Their Test Implications |journal=Current Anthropology |volume=44 |issue=5 |pages=627–651 |doi=10.1086/377665|pmid=14971366 |s2cid=11081605 }}</ref><ref>{{cite journal |last1=Marean |first1=Curtis |title=Early human use of marine resources and pigment in South Africa during the Middle Pleistocene |journal=Nature |date=2007 |volume=449 |issue=7164 |display-authors=etal |doi=10.1038/nature06204 |pages=905–908 |pmid=17943129 |bibcode=2007Natur.449..905M|s2cid=4387442 |url=http://doc.rero.ch/record/15550/files/PAL_E2962.pdf }}</ref><ref>{{cite journal |last1=Powell |first1=Adam |title=Late Pleistocene Demography and the Appearance of Modern Human Behavior |journal=Science |date=2009 |volume=324 |issue=5932 |pages=1298–1301 |display-authors=etal |doi=10.1126/science.1170165 |bibcode=2009Sci...324.1298P |pmid=19498164|s2cid=206518315 |url=http://doc.rero.ch/record/210393/files/PAL_E4401.pdf |archive-url=https://web.archive.org/web/20170829071205/http://doc.rero.ch/record/210393/files/PAL_E4401.pdf |archive-date=2017-08-29 |url-status=live }}</ref><ref>{{cite journal |last1=Premo |first1=Luke |last2=Kuhn |first2=Steve |title=Modeling Effects of Local Extinctions on Culture Change and Diversity in the Paleolithic |journal=PLOS ONE |date=2010 |volume=5 |issue=12 |doi=10.1371/journal.pone.0015582 |pmid=21179418 |pages=e15582 |bibcode=2010PLoSO...515582P |pmc=3003693|doi-access=free }}</ref>

===Examples of behavioral modernity===
[[File:Claimed Oldest Known Drawing by Human Hands Discovered in South African Cave.jpg|thumb|Claimed "[[History of art#Lower and Middle Paleolithic|oldest known drawing by human hands]]", discovered in [[Blombos Cave]] in [[South Africa]]. Estimated to be a 73,000-year-old work of a ''Homo sapiens''.<ref name="NYT-20180912">{{cite news |last=St. Fleur |first=Nicholas|title=Oldest Known Drawing by Human Hands Discovered in South African Cave |url=https://www.nytimes.com/2018/09/12/science/oldest-drawing-ever-found.html |date=12 September 2018 |work=[[The New York Times]] |access-date=15 September 2018 }}</ref>]]

The equivalent of the Eurasian Upper Paleolithic in African archaeology is known as the [[Later Stone Age]], also beginning roughly 40,000 years ago. While most clear evidence for behavioral modernity uncovered from the later 19th century was from Europe, such as the [[Venus figurine]]s and other artefacts from the [[Aurignacian]], more recent archaeological research has shown that all essential elements of the kind of material culture typical of contemporary [[San people|San]] hunter-gatherers in [[Southern Africa]] was also present by at least 40,000 years ago, including digging sticks of similar materials used today, [[ostrich egg]] shell beads, bone [[arrow]] heads with individual maker's marks etched and embedded with red [[ochre]], and poison applicators.<ref>{{Cite journal |doi=10.1073/pnas.1204213109 |pmid=22847420 |title=Early evidence of San material culture represented by organic artifacts from Border Cave, South Africa |journal=Proceedings of the National Academy of Sciences |volume=109 |issue=33 |pages=13214–13219 |year=2012 |last1=d'Errico |first1=F. |last2=Backwell |first2=L. |last3=Villa |first3=P. |last4=Degano |first4=I. |last5=Lucejko |first5=J. J. |last6=Bamford |first6=M. K. |last7=Higham |first7=T. F. G. |last8=Colombini |first8=M. P. |last9=Beaumont |first9=P. B. |bibcode=2012PNAS..10913214D |pmc=3421171|doi-access=free }}</ref> There is also a suggestion that "pressure flaking best explains the morphology of lithic artifacts recovered from the c. 75-ka Middle Stone Age levels at [[Blombos Cave]], South Africa. The technique was used during the final shaping of Still Bay bifacial points made on heat‐treated silcrete."<ref>{{Cite journal |doi=10.1126/science.1195550 |pmid=21030655 |title=Early Use of Pressure Flaking on Lithic Artifacts at Blombos Cave, South Africa |journal=Science |volume=330 |issue=6004 |pages=659–662 |year=2010 |last1=Mourre |first1=V. |last2=Villa |first2=P. |last3=Henshilwood |first3=C. S. |bibcode=2010Sci...330..659M|s2cid=34833884 }}</ref> Both pressure flaking and heat treatment of materials were previously thought to have occurred much later in prehistory, and both indicate a behaviourally modern sophistication in the use of natural materials. Further reports of research on cave sites along the southern African coast indicate that "the debate as to when cultural and cognitive characteristics typical of modern humans first appeared" may be coming to an end, as "advanced technologies with elaborate chains of production" which "often demand high-fidelity transmission and thus language" have been found at the South African [[Pinnacle Point]] Site 5–6. These have been dated to approximately 71,000 years ago. The researchers suggest that their research "shows that [[microlithic technology]] originated early in South Africa by 71 kya, evolved over a vast time span (c. 11,000 years), and was typically coupled to complex heat treatment that persisted for nearly 100,000 years. Advanced technologies in [[Africa]] were early and enduring; a small sample of excavated sites in Africa is the best explanation for any perceived 'flickering' pattern."<ref name="auto2">{{cite journal |doi=10.1038/nature11660 |pmid=23135405 |title=An early and enduring advanced technology originating 71,000 years ago in South Africa |journal=Nature |volume=491 |issue=7425 |pages=590–593 |year=2012 |last1=Brown |first1=Kyle S. |last2=Marean |first2=Curtis W. |last3=Jacobs |first3=Zenobia |last4=Schoville |first4=Benjamin J. |last5=Oestmo |first5=Simen |last6=Fisher |first6=Erich C. |last7=Bernatchez |first7=Jocelyn |last8=Karkanas |first8=Panagiotis |last9=Matthews |first9=Thalassa |bibcode=2012Natur.491..590B|s2cid=4323569 }}</ref> Increases in behavioral complexity have been speculated to have been linked to an earlier climatic change to much drier conditions between 135,000 and 75,000 years ago.<ref>{{cite journal |doi=10.1073/pnas.0703874104 |title=East African megadroughts between 135 and 75 thousand years ago and bearing on early-modern human origins |journal=Proceedings of the National Academy of Sciences |volume=104 |issue=42 |pages=16416–16421 |year=2007 |last1=Scholz |first1=C. A. |last2=Johnson |first2=T. C. |last3=Cohen |first3=A. S. |last4=King |first4=J. W. |last5=Peck |first5=J. A. |last6=Overpeck |first6=J. T. |last7=Talbot |first7=M. R. |last8=Brown |first8=E. T. |last9=Kalindekafe |first9=L. |last10=Amoako |first10=P. Y. O. |last11=Lyons |first11=R. P. |last12=Shanahan |first12=T. M. |last13=Castaneda |first13=I. S. |last14=Heil |first14=C. W. |last15=Forman |first15=S. L. |last16=McHargue |first16=L. R. |last17=Beuning |first17=K. R. |last18=Gomez |first18=J. |last19=Pierson |first19=J. |bibcode=2007PNAS..10416416S |pmid=17785420 |pmc=1964544|doi-access=free }}</ref> This might have led to human groups who were seeking refuge from the inland droughts, expanded along the coastal marshes rich in shellfish and other resources. Since sea levels were low due to so much water tied up in [[glacier]]s, such marshlands would have occurred all along the southern coasts of Eurasia. The use of [[raft]]s and boats may well have facilitated exploration of offshore islands and travel along the coast, and eventually permitted expansion to New Guinea and then to [[Australia]].<ref>{{cite book|title=The Journey of Man: A Genetic Odyssey|author=Wells, Spencer|url=https://archive.org/details/journeyofmangene00well|isbn=978-0691115320|year=2003|url-access=registration|publisher=Princeton, N.J. : Princeton University Press}}</ref>

In addition, a variety of other evidence of abstract imagery, widened subsistence strategies, and other "modern" behaviors has been discovered in Africa, especially South, North, and East Africa, predating 50,000 years ago (with some predating 100,000 years ago). The Blombos Cave site in South Africa, for example, is famous for rectangular slabs of ochre engraved with [[geometric]] designs. Using multiple dating techniques, the site was confirmed to be around 77,000 and 100,000–75,000 years old.<ref name="Henshilwood 2002">{{cite journal |last1=Henshilwood |first1=Christopher |title=Emergence of Modern Human Behavior: Middle Stone Age Engravings from South Africa |journal=Science |date=2002 |volume=295 |issue=5558 |pages=1278–1280 |display-authors=etal |doi=10.1126/science.1067575 |pmid=11786608 |bibcode=2002Sci...295.1278H |s2cid=31169551 }}</ref><ref name="Henshilwood et al. 2009">{{cite journal |doi=10.1016/j.jhevol.2009.01.005 |pmid=19487016 |title=Engraved ochres from the Middle Stone Age levels at Blombos Cave, South Africa |journal=Journal of Human Evolution |volume=57 |issue=1 |pages=27–47 |year=2009 |last1=Henshilwood |first1=Christopher S. |last2=d'Errico |first2=Francesco |last3=Watts |first3=Ian|bibcode=2009JHumE..57...27H }}</ref> Ostrich egg shell containers engraved with geometric designs dating to 60,000 years ago were found at [[Diepkloof Rock Shelter|Diepkloof]], South Africa.<ref name="Texier">{{cite journal | last1 = Texier | first1 = PJ | last2 = Porraz | first2 = G | last3 = Parkington | first3 = J | last4 = Rigaud | first4 = JP | last5 = Poggenpoel | first5 = C | last6 = Miller | first6 = C | last7 = Tribolo | first7 = C | last8 = Cartwright | first8 = C | last9 = Coudenneau | first9 = A | last10 = Klein | first10 = R | last11 = Steele | first11 = T | last12 = Verna | first12 = C | year = 2010 | title = A Howiesons Poort tradition of engraving ostrich eggshell containers dated to 60,000 years ago at Diepkloof Rock Shelter, South Africa | journal =   Proceedings of the National Academy of Sciences| volume = 107 | issue = 14| pages = 6180–6185 | doi = 10.1073/pnas.0913047107 | pmid = 20194764 | pmc = 2851956 | bibcode = 2010PNAS..107.6180T | doi-access = free }}</ref> Beads and other personal ornamentation have been found from Morocco which might be as much as 130,000 years old; as well, the Cave of Hearths in South Africa has yielded a number of beads dating from significantly prior to 50,000 years ago,<ref name="McBrearty Brooks 2000">{{cite journal |last1=McBrearty |first1=Sally |last2=Brooks |first2=Allison |date=2000 |title=The revolution that wasn't: a new interpretation of the origin of modern human behavior |journal=Journal of Human Evolution |volume=39 |issue=5 |pages=453–563 |doi=10.1006/jhev.2000.0435 |pmid=11102266 |bibcode=2000JHumE..39..453M |s2cid=42968840 }}</ref> and shell beads dating to about 75,000 years ago have been found at Blombos Cave, South Africa.<ref name="Henshilwood et al. 2004">{{cite journal | last1 = Henshilwood | first1 = Christopher S. | author-link = Christopher Henshilwood | display-authors = etal | year = 2004 | title = Middle Stone Age shell beads from South Africa | journal = Science | volume = 304 | issue = 5669| page = 404 | doi = 10.1126/science.1095905 | pmid = 15087540 | s2cid = 32356688 }}</ref><ref name="d'Errico et al. 2005">{{cite journal | last1 = d'Errico | first1 = Francesco | display-authors = etal | year = 2005 | title = Nassarius kraussianus shell beads from Blombos Cave: evidence for symbolic behaviour in the Middle Stone Age | journal = Journal of Human Evolution | volume = 48 | issue = 1| pages = 3–24 | doi = 10.1016/j.jhevol.2004.09.002 | pmid = 15656934 | bibcode = 2005JHumE..48....3D }}</ref><ref name="Vanhaeren et al. 2013">{{cite journal | last1 = Vanhaeren | first1 = Marian | display-authors = etal | year = 2013 | title = Thinking strings: Additional evidence for personal ornament use in the Middle Stone Age at Blombos Cave, South Africa | journal = Journal of Human Evolution | volume = 64 | issue = 6| pages = 500–517 | doi = 10.1016/j.jhevol.2013.02.001 | pmid = 23498114 | bibcode = 2013JHumE..64..500V }}</ref> Specialized projectile weapons as well have been found at various sites in Middle Stone Age Africa, including bone and stone arrowheads at South African sites such as [[Sibudu Cave]] (along with an early bone needle also found at Sibudu) dating approximately 72,000–60,000 years ago<ref name="Backwell">{{cite journal | last1 = Backwell | first1 = L | last2 = d'Errico | first2 = F | last3 = Wadley | first3 = L | year = 2008 | title = Middle Stone Age bone tools from the Howiesons Poort layers, Sibudu Cave, South Africa | journal = Journal of Archaeological Science | volume = 35 | issue = 6| pages = 1566–1580 | doi = 10.1016/j.jas.2007.11.006 | bibcode = 2008JArSc..35.1566B }}</ref><ref>{{cite journal |last1=Wadley |first1=Lyn |year=2008 |title=The Howieson's Poort industry of Sibudu Cave |journal= South African Archaeological Society Goodwin Series |volume=10}}</ref><ref name="Lombard">{{Cite journal |title=Indications of bow and stone-tipped arrow use 64,000 years ago in KwaZulu-Natal, South Africa |journal=Antiquity |volume=84 |issue=325 |pages=635–648 |year=2010 |doi=10.1017/S0003598X00100134 |vauthors=Lombard M, Phillips L|s2cid=162438490 }}</ref><ref name="Lombard M">{{Cite journal|title=Quartz-tipped arrows older than 60 ka: further use-trace evidence from Sibudu, Kwa-Zulu-Natal, South Africa|journal=Journal of Archaeological Science|year=2011|doi=10.1016/j.jas.2011.04.001 |vauthors=Lombard M|volume=38|issue=8|pages=1918–1930|bibcode=2011JArSc..38.1918L }}</ref><ref name="Backwell2018">{{cite journal | last1 = Backwell | first1 = L | last2 = Bradfield | first2 = J | last3 = Carlson | first3 = KJ | last4 = Jashashvili | first4 = T | last5 = Wadley | first5 = L | last6 = d'Errico | first6 = F | year = 2018 | title = The antiquity of bow-and-arrow technology: evidence from Middle Stone Age layers at Sibudu Cave | journal = Journal of Archaeological Science | volume = 92 | issue = 362| pages = 289–303 | doi = 10.15184/aqy.2018.11 | doi-access = free | hdl = 11336/81248 | hdl-access = free }}</ref> some of which may have been tipped with poisons,<ref name="Lombard2020">{{Cite journal|title=The tip cross-sectional areas of poisoned bone arrowheads from southern Africa |journal=Journal of Archaeological Science: Reports |volume=33 |year=2020 |doi= 10.1016/j.jasrep.2020.102477 |vauthors=Lombard M|page=102477 |bibcode=2020JArSR..33j2477L |s2cid=224889105 }}</ref> and bone harpoons at the Central African site of Katanda dating ca. 90,000 years ago.<ref>{{cite journal |title=A middle stone age worked bone industry from Katanda, Upper Semliki Valley, Zaire |date=28 April 1995 |last=Yellen |first=JE |author2=AS Brooks |author3=E Cornelissen |author4=MJ Mehlman |author5=K Stewart |journal=Science |volume=268 |pages=553–556 |issue=5210 |doi=10.1126/science.7725100 |pmid=7725100|bibcode=1995Sci...268..553Y }}</ref> Evidence also exists for the systematic heat treating of silcrete stone to increase its flake-ability for the purpose of toolmaking, beginning approximately 164,000 years ago at the South African site of Pinnacle Point and becoming common there for the creation of microlithic tools at about 72,000 years ago.<ref>{{citation|last1=Brown|first1=Kyle S. |last2= Marean| first2= Curtis W. |last3= Herries |first3= Andy I.R. |last4= Jacobs |first4= Zenobia |last5=Tribolo |first5= Chantal |last6= Braun |first6=David |last7=Roberts |first7= David L. |last8=Meyer |first8=Michael C. |author9=Bernatchez, J. |date=14 August 2009 |title= Fire as an Engineering Tool of Early Modern Humans| journal= Science |volume= 325 |issue=5942 |pages= 859–862 |doi=10.1126/science.1175028 |pmid=19679810|bibcode=2009Sci...325..859B |hdl=11422/11102 |s2cid=43916405 |hdl-access=free }}</ref><ref name="auto2"/>

In 2008, an ochre processing workshop likely for the production of paints was uncovered dating to ca. 100,000 years ago at Blombos Cave, South Africa. Analysis shows that a liquefied pigment-rich mixture was produced and stored in the two abalone shells, and that ochre, bone, charcoal, grindstones and hammer-stones also formed a composite part of the toolkits. Evidence for the complexity of the task includes procuring and combining raw materials from various sources (implying they had a mental template of the process they would follow), possibly using pyrotechnology to facilitate fat extraction from bone, using a probable recipe to produce the compound, and the use of shell containers for mixing and storage for later use.<ref name="bbc.com-15257259">{{cite news |last=Amos |first=Jonathan |author-link=Jonathan Amos |title=A Cultural Leap at the Dawn of Humanity – Ancient 'paint factory' unearthed |url= https://www.bbc.com/news/science-environment-15257259 |date=13 October 2011 |work=[[BBC News]] |access-date=13 October 2011 }}</ref><ref name="Washington Post-2011/10/12/gIQApyHrhL">{{cite news |last=Vastag |first=Brian |author-link=Brian Vastag |title=South African cave yields paint from dawn of humanity |url= https://www.washingtonpost.com/national/health-science/african-cave-yields-paint-from-dawn-of-humanity/2011/10/12/gIQApyHrhL_story.html |date=13 October 2011 |newspaper=[[The Washington Post]] |access-date=13 October 2011 }}</ref><ref name="Henshilwood et al. 2011">{{cite journal | last1 = Henshilwood | first1 = Christopher S. | display-authors = etal | year = 2011 | title = A 100,000-Year-Old Ochre-Processing Workshop at Blombos Cave, South Africa | journal = Science | volume = 334 | issue = 6053| pages = 219–222 | doi = 10.1126/science.1211535 | pmid = 21998386 | bibcode = 2011Sci...334..219H | s2cid = 40455940 }}</ref> Modern behaviors, such as the making of shell beads, bone tools and arrows, and the use of ochre pigment, are evident at a Kenyan site by 78,000-67,000 years ago.<ref>Shipton C, d'Errico F, Petraglia M, et al. (2018). 78,000-year-old record of Middle and Later Stone Age innovation in an East African tropical forest. Nature Communications</ref> Evidence of early stone-tipped projectile weapons (a characteristic tool of ''Homo sapiens''), the stone tips of javelins or throwing spears, were discovered in 2013 at the Ethiopian site of [[Gademotta]], and date to around 279,000 years ago.<ref name="SahlePLOS1">{{Cite journal |last1=Sahle |first1=Y. |last2=Hutchings |first2=W. K. |last3=Braun |first3=D. R. |last4=Sealy |first4=J. C. |last5=Morgan |first5=L. E. |last6=Negash |first6=A. |last7=Atnafu |first7=B. |editor1-last=Petraglia |editor1-first=Michael D |title=Earliest Stone-Tipped Projectiles from the Ethiopian Rift Date to >279,000 Years Ago |doi=10.1371/journal.pone.0078092 |journal=PLOS ONE |volume=8 |issue=11 |pages=e78092 |year=2013 |pmid=24236011 |pmc=3827237 |bibcode=2013PLoSO...878092S |doi-access=free }}</ref>

Expanding subsistence strategies beyond big-game hunting and the consequential diversity in tool types have been noted as signs of behavioral modernity. A number of South African sites have shown an early reliance on aquatic resources from fish to shellfish. [[Pinnacle Point]], in particular, shows exploitation of marine resources as early as 120,000 years ago, perhaps in response to more arid conditions inland.<ref name="Marean et al 2007">{{cite journal|last1=Marean|first1=Curtis|title=Early human use of marine resources and pigment in South Africa during the Middle Pleistocene|journal=Nature|date=2007|volume=449|issue=7164|display-authors=etal|doi=10.1038/nature06204|pages=905–908|pmid=17943129|bibcode=2007Natur.449..905M|s2cid=4387442|url=http://doc.rero.ch/record/15550/files/PAL_E2962.pdf }}</ref> Establishing a reliance on predictable shellfish deposits, for example, could reduce mobility and facilitate complex social systems and symbolic behavior. Blombos Cave and Site 440 in Sudan both show evidence of fishing as well. Taphonomic change in fish skeletons from Blombos Cave have been interpreted as capture of live fish, clearly an intentional human behavior.<ref name="McBrearty Brooks 2000" />

Humans in North Africa ([[Nazlet Sabaha]], [[Egypt]]) are known to have dabbled in [[chert]] [[mining]], as early as ≈100,000 years ago, for the construction of [[stone tool]]s.<ref>{{Cite web |url=https://www.promine.com/blog/5-oldest-mines-in-the-world-a-casual-survey |title=5 Oldest Mines in the World: A Casual Survey |access-date=2019-09-11 |archive-url=https://web.archive.org/web/20190105054434/https://www.promine.com/blog/5-oldest-mines-in-the-world-a-casual-survey |archive-date=2019-01-05 |url-status=dead }}</ref><ref name="Records2015">{{cite book|author=Guinness World Records|title=Guinness World Records 2016|url=https://books.google.com/books?id=f896CgAAQBAJ&pg=PA27|date=10 September 2015|publisher=Guinness World Records|isbn=978-1910561034|page=27}}</ref>

Evidence was found in 2018, dating to about 320,000 years ago at the site of [[Olorgesailie]] in Kenya, of the early emergence of modern behaviors including: the trade and long-distance transportation of resources (such as obsidian), the use of pigments, and the possible making of projectile points. The authors of three 2018 studies on the site observe that the evidence of these behaviors is roughly contemporary with the earliest known ''Homo sapiens'' fossil remains from Africa (such as at Jebel Irhoud and Florisbad), and they suggest that complex and modern behaviors began in Africa around the time of the emergence of ''Homo sapiens''.<ref name="NPR-593591796">{{cite news |last=Chatterjee |first=Rhitu |author-link=List of NPR personnel |title=Scientists Are Amazed By Stone Age Tools They Dug Up In Kenya |url=https://www.npr.org/sections/goatsandsoda/2018/03/15/593591796/scientists-are-amazed-by-stone-age-tools-they-dug-up-in-kenya |date=15 March 2018 |work=[[NPR]] |access-date=15 March 2018 }}</ref><ref name="The Atlantic-555674">{{cite news |last=Yong |first=Ed |author-link=Ed Yong |title=A Cultural Leap at the Dawn of Humanity – New finds from Kenya suggest that humans used long-distance trade networks, sophisticated tools, and symbolic pigments right from the dawn of our species. |url=https://www.theatlantic.com/science/archive/2018/03/a-deeper-origin-of-complex-human-cultures/555674/ |date=15 March 2018 |work=[[The Atlantic]] |access-date=15 March 2018 }}</ref><ref name="Brooks">{{Cite journal|title=Long-distance stone transport and pigment use in the earliest Middle Stone Age|journal=Science|volume=360|issue=6384|pages=90–94|year=2018|doi = 10.1126/science.aao2646|pmid=29545508|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|bibcode=2018Sci...360...90B|doi-access=free}}</ref>

In 2019, further evidence of Middle Stone Age complex projectile weapons in Africa was found at Aduma, Ethiopia, dated 100,000–80,000 years ago, in the form of points considered likely to belong to darts delivered by spear throwers.<ref name="Sahle">{{Cite journal|title=Assessment of complex projectiles in the early Late Pleistocene at Aduma, Ethiopia|journal=PLOS ONE|volume=14|issue=5|pages=e0216716|year=2018|doi = 10.1371/journal.pone.0216716|pmid=31071181|pmc=6508696|vauthors=Sahle Y, Brooks AS|bibcode=2019PLoSO..1416716S|doi-access=free}}</ref>

==Pace of progress during ''Homo sapiens'' history==
{{See also|Sapient paradox|The 10,000 Year Explosion}}
''Homo sapiens'' technological and cultural progress appears to have been very much faster in recent millennia than in [[Middle Paleolithic|''Homo sapiens'' early periods]]. The pace of development may indeed have accelerated, due to massively larger population (so more humans extant to think of innovations), more communication and sharing of ideas among human populations, and the accumulation of thinking tools. However it may also be that the pace of advancements always looks relatively faster to humans in the time they live, because previous advances are unrecognised.<ref>{{cite magazine |last1=Douglas |first1=Kate |title=Puzzles of Evolution: Why was technological development so slow? |url=https://www.newscientist.com/article/mg21328571-400-puzzles-of-evolution-why-was-technological-development-so-slow/ |magazine=[[New Scientist]] |access-date=May 4, 2022 |archive-url=https://web.archive.org/web/20160413061933/https://www.newscientist.com/article/mg21328571-400-puzzles-of-evolution-why-was-technological-development-so-slow/ |archive-date=April 13, 2016 |date=March 24, 2012 |url-status=unfit}}</ref>

== Notes ==
{{reflist|group=note}}

== References ==
{{reflist}}

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* {{cite journal |vauthors=Vai S, Sarno S, Lari M, Luiselli D, Manzi G, Gallinaro M, Mataich S, Hübner A, Modi A, Pilli E, Tafuri MA, Caramelli D, di Lernia S |date=March 2019 |title=Ancestral mitochondrial N lineage from the Neolithic 'green' Sahara |journal=Sci Rep |volume=9 |issue=1 |page=3530 |doi=10.1038/s41598-019-39802-1 |pmc=6401177 |pmid=30837540 |bibcode=2019NatSR...9.3530V |ref={{sfnref|Vai, et al., 2019}}}}
{{refend}}

== Further reading ==
* {{cite book |title=Who We Are And How We Got Here – Ancient DNA and the New Science of the Human Past |first=David |last=Reich |publisher=[[Pantheon Books]] |year=2018 |isbn=978-1101870327 }}<ref name="NYT-20180420">{{cite news |last=Diamond |first=Jared |title=A Brand-New Version of Our Origin Story |url=https://www.nytimes.com/2018/04/20/books/review/david-reich-who-we-are-how-we-got-here.html |date=April 20, 2018 |newspaper=[[The New York Times]] |access-date=April 23, 2018 }}</ref>

== External links ==
* [http://humanorigins.si.edu/evidence/human-evolution-timeline-interactive Human Timeline (Interactive)] – [[Smithsonian Institution|Smithsonian]], [[National Museum of Natural History]] (August 2016).
* {{wikispecies-inline}}

{{Human Evolution|state=collapsed}}
{{Big History}}
{{portal bar|Evolutionary biology}}

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[[Category:Humans]]
[[Category:Anatomically modern humans| ]]
[[Category:Mammals described in 1758]]
[[Category:Taxa named by Carl Linnaeus]]
[[Category:Tool-using mammals]]