Editing Early modern human (section)

== 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>