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{{Short description|"Out of Africa" theory of the early migration of humans}}
{{about|modern humans|migrations of early humans|Early hominin expansions out of Africa}}
{{About|modern humans|migrations of early humans|Early expansions of hominins out of Africa}}
{{See also|Early human migrations}}
{{redirects here|Single origin hypothesis|a single origin of humans in the general sense|Monogenism}}
{{use dmy dates|date=July 2016}}
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[[File:Map-of-human-migrations.jpg|thumb|400px|right|Map of the migration of modern humans out of Africa, based on [[mitochondrial DNA]]. Colored rings indicate thousand years before present.]]
{{Use dmy dates|date=September 2024}}
In [[paleoanthropology]], the '''recent African origin of modern humans''', also called the "'''Out of Africa'''" '''theory''' ('''OOA'''), '''recent single-origin hypothesis''' ('''RSOH'''), '''replacement hypothesis''', or '''recent African origin model''' ('''RAO'''), is the dominant{{r|pmid16826514Quo|pmid12802315}} model of the geographic origin and [[Early human migrations|early migration]] of [[anatomically modern human]]s (''[[Homo sapiens]]'').


[[File:Spreading homo sapiens la.svg|thumb|upright=1.5|Successive dispersals (labeled in [[Before Present|years before present]]) of <br />{{color box|yellow||}} ''[[Homo erectus]]'' greatest extent (yellow)<br />{{color box|#e4ca30}} ''[[Neanderthal|Homo neanderthalensis]]'' greatest extent (ochre) <br />{{color box|#e9252c}} ''[[Human|Homo sapiens]]'' (red)]]
The model proposes a "single origin" of ''[[Homo sapiens]]'' in the taxonomic sense, precluding
[[File:Expansion of early modern humans from Africa.jpg|thumb|upright=1.5|Expansion of early modern humans from Africa through the [[Near East]]]]
[[parallel evolution]] of traits considered [[anatomically modern humans|anatomically modern]] in other regions,{{r|pmid10766948}} but not precluding limited [[Archaic human admixture with modern humans|admixture]] between ''H. sapiens'' and archaic humans in Europe and Asia.{{refn|group=note|name="JurmainKilgore2008"|From 1984 to 2003, an alternative scientific hypothesis was the [[multiregional origin of modern humans]], which envisioned a wave of ''Homo sapiens'' migrating earlier from Africa and interbreeding with local ''[[Homo erectus]]'' populations in varied regions of the globe.{{cite book|author1=Robert Jurmain |author2=Lynn Kilgore |author3=Wenda Trevathan |title=Essentials of Physical Anthropology |url=https://books.google.com/books?id=TSaSPza9LMYC&pg=PA266 |accessdate=14 June 2011 |date=20 March 2008 |publisher=Cengage Learning |isbn=978-0-495-50939-4 |pages=266–}}}}
In [[paleoanthropology]], the '''recent African origin of modern humans''' or the "'''Out of Africa'''" '''theory''' ('''OOA'''){{Efn|Also called the '''recent single-origin hypothesis''' ('''RSOH'''), '''replacement hypothesis''', or '''recent African origin model''' ('''RAO''').}} is the most widely accepted{{r|pmid16826514Quo|pmid12802315}}<ref name="Stringer2012">{{cite book
''H. sapiens'' most likely developed in the [[Horn of Africa]] between 300,000 and 200,000 years ago.
|vauthors=Stringer C |title=Lone Survivors: How We Came to Be the Only Humans on Earth|url=https://books.google.com/books?id=zRxfKetC1gMC&pg=PA26 |year= 2012 |publisher=Henry Holt and Company |isbn=978-1429973441 |page=26 |author-link=Chris Stringer |via=[[Google Books]]}}</ref> model of the geographic origin and [[Early human migrations|early migration]] of [[early modern human|anatomically modern humans]] (''[[Human|Homo sapiens]]''). It follows the [[early expansions of hominins out of Africa]], accomplished by ''[[Homo erectus]]'' and then ''[[Neanderthal|Homo neanderthalensis]]''.
The "recent African origin" model proposes that all modern non-African populations are substantially descended from populations of ''H. sapiens'' that left Africa after that time.


The model proposes a "[[monogenism|single origin]]" of ''[[Human|Homo sapiens]]'' in the [[taxonomic]] sense, precluding [[parallel evolution]] in other regions of traits considered [[anatomically modern humans|anatomically modern]],{{r|pmid10766948}} but not precluding multiple [[Archaic human admixture with modern humans|admixture]] between ''H. sapiens'' and [[archaic humans]] in Europe and Asia.{{Efn|From 1984 to 2003, an alternative scientific hypothesis was the [[multiregional origin of modern humans]], which envisioned a wave of ''Homo sapiens'' migrating earlier from Africa and interbreeding with local ''[[Homo erectus]]'' populations in varied regions of the globe.{{cite book |vauthors = Jurmain R, Kilgore L, Trevathan W |title=Essentials of Physical Anthropology |url=https://books.google.com/books?id=TSaSPza9LMYC&pg=PA266 |access-date=14 June 2011 |year=2008 |publisher=[[Cengage Learning]] |isbn=978-0495509394 |pages=266–}}}}<ref>{{cite journal |vauthors=Mafessoni F |title=Encounters with archaic hominins |journal=[[Nature Ecology & Evolution]] |volume=3 |issue=1 |pages=14–15 |date=January 2019 |pmid=30478304 |doi=10.1038/s41559-018-0729-6 |s2cid=53783648}}</ref><ref>{{cite journal |vauthors=Villanea FA, Schraiber JG |title=Multiple episodes of interbreeding between Neanderthal and modern humans |journal=[[Nature Ecology & Evolution]] |volume=3 |issue=1 |pages=39–44 |date=January 2019 |pmid=30478305 |pmc=6309227 |doi=10.1038/s41559-018-0735-8}}</ref> ''H. sapiens'' most likely developed in the [[Horn of Africa]] between 300,000 and 200,000 years ago,<ref name="EA-20190320">{{cite news |author=University of Huddersfield |title=Researchers shed new light on the origins of modern humans – The work, published in Nature, confirms a dispersal of Homo sapiens from southern to eastern Africa immediately preceded the out-of-Africa migration |url=https://www.eurekalert.org/pub_releases/2019-03/uoh-nrs032019.php |date=20 March 2019 |work=[[EurekAlert!]] |access-date=23 March 2019 |author-link=University of Huddersfield}}</ref><ref name="SR-20190318">{{cite journal |vauthors=Rito T, Vieira D, Silva M, Conde-Sousa E, Pereira L, Mellars P, Richards MB, Soares P |display-authors=6 |title=A dispersal of Homo sapiens from southern to eastern Africa immediately preceded the out-of-Africa migration |journal=[[Scientific Reports]] |volume=9 |issue=1 |page=4728 |date=March 2019 |pmid=30894612 |pmc=6426877 |doi=10.1038/s41598-019-41176-3 |bibcode=2019NatSR...9.4728R}}</ref> although an alternative hypothesis argues that diverse morphological features of ''H. sapiens'' appeared locally in different parts of Africa and converged due to [[gene flow]] between different populations within the same period.<ref>{{cite journal |vauthors=Scerri EM, Chikhi L, Thomas MG |title=Beyond multiregional and simple out-of-Africa models of human evolution |journal=[[Nature Ecology & Evolution]] |volume=3 |issue=10 |pages=1370–1372 |date=October 2019 |pmid=31548642 |doi=10.1038/s41559-019-0992-1 |bibcode=2019NatEE...3.1370S |hdl=10400.7/954 |s2cid=202733639 |hdl-access=free}}</ref><ref>{{cite journal |vauthors=Scerri EM, Thomas MG, Manica A, Gunz P, Stock JT, Stringer C, Grove M, Groucutt HS, Timmermann A, Rightmire GP, d'Errico F, Tryon CA, Drake NA, Brooks AS, Dennell RW, Durbin R, Henn BM, Lee-Thorp J, deMenocal P, Petraglia MD, Thompson JC, Scally A, Chikhi L |display-authors=6 |title=Did Our Species Evolve in Subdivided Populations across Africa, and Why Does It Matter? |journal=[[Trends (journals)|Trends in Ecology & Evolution]] |volume=33 |issue=8 |pages=582–594 |date=August 2018 |pmid=30007846 |pmc=6092560 |doi=10.1016/j.tree.2018.05.005|bibcode=2018TEcoE..33..582S }}</ref> The "recent African origin" model proposes that all modern non-African populations are substantially descended from populations of ''H. sapiens'' that left Africa after that time.
There were at least several "out-of-Africa" dispersals of modern humans, possibly beginning as early as 270,000 years ago, and certainly during 130,000 to 115,000 ago via northern Africa.{{r|pmid21273486|pmid21212332|pmid21601174|pmid17372199}}<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 |url=http://science.sciencemag.org/content/358/6368/eaai9067 |journal=[[Science (journal)|Science]] |date=8 December 2017 |volume=358 |issue=6368 |page=eaai9067 |doi=10.1126/science.aai9067 |accessdate=10 December 2017 }}</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 |work=[[Quartz (publication)|Quartz]] |accessdate=10 December 2017 }}</ref>
These early waves appear to have mostly died out or retreated by 80,000 years ago.<ref name="Liu2015">{{harvp|Liu, Martinón-Torres et al.|2015}}.<br>See also [http://dienekes.blogspot.nl/2015/10/modern-humans-in-china-80000-years-ago.html ''Modern humans in China ~80,000 years ago (?)''], Dieneks' Anthropology Blog.</ref>


There were at least several "out-of-Africa" dispersals of modern humans, possibly beginning as early as 270,000 years ago, including 215,000 years ago to at least Greece,<ref name="NYT-20190710">{{cite news |vauthors=Zimmer C |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 |vauthors=Harvati K, Röding C, Bosman AM, Karakostis FA, Grün R, Stringer C, Karkanas P, Thompson NC, Koutoulidis V, Moulopoulos LA, Gorgoulis VG, Kouloukoussa M |display-authors=6 |title=Apidima Cave fossils provide earliest evidence of Homo sapiens in Eurasia |journal=[[Nature (journal)|Nature]] |volume=571 |issue=7766 |pages=500–504 |date=July 2019 |pmid=31292546 |doi=10.1038/s41586-019-1376-z |s2cid=195873640 |url=https://zenodo.org/record/6646855}}</ref> and certainly via northern Africa and the Arabian Peninsula about 130,000 to 115,000 years ago.{{refn|{{r|pmid21273486|pmid21212332|pmid21601174|pmid17372199}}<ref name="SCI-20171208">{{cite journal |vauthors=Bae CJ, Douka K, Petraglia MD |title=On the origin of modern humans: Asian perspectives |journal=[[Science (journal)|Science]] |volume=358 |issue=6368 |pages=eaai9067 |date=December 2017 |pmid=29217544 |doi=10.1126/science.aai9067 |doi-access=free}}</ref><ref name="QZ-20171210">{{cite web |vauthors=Kuo L |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 |work=[[Quartz (publication)|Quartz]] |access-date=10 December 2017}}</ref>}} There is evidence that modern humans had reached China around 80,000 years ago.<ref name="Liu2015">{{harvp|Liu, Martinón-Torres et al.|2015}}.<br />See also [http://dienekes.blogspot.nl/2015/10/modern-humans-in-china-80000-years-ago.html ''Modern humans in China ~80,000 years ago (?)''], Dieneks' Anthropology Blog.</ref> Practically all of these early waves seem to have gone extinct or retreated back, and present-day humans outside Africa descend mainly from a single expansion about 70,000–50,000 years ago,{{sfnp|Finlayson|2009|p=68}}{{sfnp|Liu, Prugnolle et al.|2006}}<ref name="Haber Jones etal 2019">{{cite journal |vauthors=Haber M, Jones AL, Connell BA, Arciero E, Yang H, Thomas MG, Xue Y, Tyler-Smith C |display-authors=6 |title=A Rare Deep-Rooting D0 African Y-Chromosomal Haplogroup and Its Implications for the Expansion of Modern Humans Out of Africa |journal=[[Genetics (journal)|Genetics]] |volume=212 |issue=4 |pages=1421–1428 |date=August 2019 |pmid=31196864 |pmc=6707464 |doi=10.1534/genetics.119.302368}}</ref><ref name="EA-20190320" /><ref name="SR-20190318" /><ref name="Posth">{{cite journal |title=Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-Africans and a Late Glacial Population Turnover in Europe |journal=[[Current Biology]] |volume=26 |issue=6 |pages=827–833 |year=2016 |doi=10.1016/j.cub.2016.01.037 |pmid=26853362 |vauthors=Posth C, Renaud G, Mittnik M, Drucker DG, Rougier H, Cupillard C, Valentin F, Thevenet C, Furtwängler A, Wißing C, Francken M, Malina M, Bolus M, Lari M, Gigli E, Capecchi G, Crevecoeur I, Beauval C, Flas D, Germonpré M, van der Plicht J, Cottiaux R, Gély B, Ronchitelli A, Wehrberger K, Grigorescu D, Svoboda J, Semal P, Caramelli D, Bocherens H, Harvati K, Conard NJ, Haak W, Powell A, Krause J |bibcode=2016CBio...26..827P |hdl=2440/114930 |s2cid=140098861 |display-authors=6 |hdl-access=free}}</ref><ref>{{cite journal |vauthors=Karmin M, Saag L, Vicente M, Wilson Sayres MA, Järve M, Talas UG, Rootsi S, Ilumäe AM, Mägi R, Mitt M, Pagani L, Puurand T, Faltyskova Z, Clemente F, Cardona A, Metspalu E, Sahakyan H, Yunusbayev B, Hudjashov G, DeGiorgio M, Loogväli EL, Eichstaedt C, Eelmets M, Chaubey G, Tambets K, Litvinov S, Mormina M, Xue Y, Ayub Q, Zoraqi G, Korneliussen TS, Akhatova F, Lachance J, Tishkoff S, Momynaliev K, Ricaut FX, Kusuma P, Razafindrazaka H, Pierron D, Cox MP, Sultana GN, Willerslev R, Muller C, Westaway M, Lambert D, Skaro V, Kovačevic L, Turdikulova S, Dalimova D, Khusainova R, Trofimova N, Akhmetova V, Khidiyatova I, Lichman DV, Isakova J, Pocheshkhova E, Sabitov Z, Barashkov NA, Nymadawa P, Mihailov E, Seng JW, Evseeva I, Migliano AB, Abdullah S, Andriadze G, Primorac D, Atramentova L, Utevska O, Yepiskoposyan L, Marjanovic D, Kushniarevich A, Behar DM, Gilissen C, Vissers L, Veltman JA, Balanovska E, Derenko M, Malyarchuk B, Metspalu A, Fedorova S, Eriksson A, Manica A, Mendez FL, Karafet TM, Veeramah KR, Bradman N, Hammer MF, Osipova LP, Balanovsky O, Khusnutdinova EK, Johnsen K, Remm M, Thomas MG, Tyler-Smith C, Underhill PA, Willerslev E, Nielsen R, Metspalu M, Villems R, Kivisild T |display-authors=6 |title=A recent bottleneck of Y chromosome diversity coincides with a global change in culture |journal=[[Genome Research]] |volume=25 |issue=4 |pages=459–466 |date=April 2015 |pmid=25770088 |pmc=4381518 |doi=10.1101/gr.186684.114}}</ref>{{excessive citations inline|date=January 2024}} via the so-called "[[Southern Dispersal|Southern Route]]". These humans spread rapidly along the coast of Asia and reached [[Prehistoric Australia|Australia]] by around 65,000–50,000 years ago,<ref name=Clarkson2017>{{cite journal |vauthors=Clarkson C, Jacobs Z, Marwick B, Fullagar R, Wallis L, Smith M, Roberts RG, Hayes E, Lowe K, Carah X, Florin SA, McNeil J, Cox D, Arnold LJ, Hua Q, Huntley J, Brand HE, Manne T, Fairbairn A, Shulmeister J, Lyle L, Salinas M, Page M, Connell K, Park G, Norman K, Murphy T, Pardoe C |display-authors=6 |title=Human occupation of northern Australia by 65,000 years ago |journal=[[Nature (journal)|Nature]] |volume=547 |issue=7663 |pages=306–310 |date=July 2017 |pmid=28726833 |doi=10.1038/nature22968 |bibcode=2017Natur.547..306C |hdl=2440/107043 |s2cid=205257212 |hdl-access=free}}</ref><ref name=StFleur2017>{{cite news | last=St. Fleur | first=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 |work=[[The New York Times]] |date=19 July 2017}}</ref>{{Efn|{{harvp|McChesney|2015}}: "...genetic evidence suggests that a small band with the marker M168 migrated out of Africa along the coasts of the Arabian Peninsula and India, through Indonesia, and reached Australia very early, between 60,000 and 50,000 years ago. This very early migration into Australia is also supported by Rasmussen et al. (2011)."}} (though some researchers question the earlier Australian dates and place the arrival of humans there at 50,000 years ago at earliest,<ref name="Wood_2017">{{Cite journal |vauthors=Wood R |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 name="Homo sapiens first reach Southeast">{{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 |bibcode=2018PNAS..115.8482O |doi-access=free}}</ref> while others have suggested that these first settlers of Australia may represent an older wave before the more significant out of Africa migration and thus not necessarily be ancestral to the region's later inhabitants<ref name="Haber Jones etal 2019" />) while Europe was populated by an early offshoot which settled the Near East and Europe less than 55,000 years ago.{{sfn|McChesney|2015}}<ref name=Macaulay2005>{{harvp|Macaulay et al.|2005}}.</ref><ref name=Posth2016>{{harvp|Posth et al.|2016}}.<br />See also {{cite web |url=http://dienekes.blogspot.nl/2016/02/mtdna-from-55-hunter-gatherers-across.html |title=mtDNA from 55 hunter-gatherers across 35,000 years in Europe |website=Dienekes' Anthroplogy Blog|date=8 February 2016 }}</ref>
The most significant "recent" wave took place about 70,000 years ago, via the so-called "[[Southern Dispersal|Southern Route]]", spreading rapidly along the coast of Asia and reaching [[Prehistoric Australia|Australia]] by around 65,000–50,000 years ago.<ref name=Clarkson2017>{{cite journal | last1 = Clarkson | first1 = Chris | display-authors = 1 | last2 = et al | year = 2017 | title = Human occupation of northern Australia by 65,000 years ago | url = https://www.nature.com/nature/journal/v547/n7663/full/nature22968.html | journal = Nature | volume = 547| issue = | pages = 306–310| doi = 10.1038/nature22968 }}</ref><ref name=StFleur2017>{{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|publisher=New York Times|date=July 19, 2017}}</ref>{{refn|group=note|name=McChesney2015_short|McChesney: "...genetic evidence suggests that a small band with the marker M168 migrated out of Africa along the coasts of the Arabian Peninsula and India, through Indonesia, and reached Australia very early, between 60,000 and 50,000 years ago. This very early migration into Australia is also supported by Rasmussen et al. (2011).}} while Europe was populated by an early offshoot which settled the Near East and Europe less than 55,000 years ago.{{sfn|Young McChesney|2015}}<ref name=Macaulay2005>{{harvp|Macaulay et al.|2005}}.</ref><ref name=Posth2016>{{harvp|Posth et al.|2016}}.<br>See also [http://dienekes.blogspot.nl/2016/02/mtdna-from-55-hunter-gatherers-across.html ''mtDNA from 55 hunter-gatherers across 35,000 years in Europe''], Dienekes' Anthroplogy Bog.</ref>


In the 2010s, studies in [[population genetics]] have uncovered evidence of [[Archaic human admixture with modern humans|interbreeding]] of ''H. sapiens'' with [[archaic humans]] both in Africa and in Eurasia,<ref name="pruf13comal">{{cite journal| last=Prüfer| first=K.|author2=Racimo, F. |author3=Patterson, N. |author4=Jay, F. |author5=Sankararaman, S. |author6=Sawyer, S. | title=The complete genome sequence of a Neanderthal from the Altai Mountains| journal=Nature| date=2014 | origyear=Online 2013 | volume=505| issue=7481| pages=43–49| doi=10.1038/nature12886|display-authors=etal | bibcode=2014Natur.505...43P | pmid=24352235 | pmc=4031459}}</ref> which means that all modern population groups, both African
In the 2010s, studies in [[population genetics]] uncovered evidence of [[Archaic human admixture with modern humans|interbreeding]] that occurred between ''H. sapiens'' and [[archaic humans]] in Eurasia, Oceania and Africa,<ref name="pruf13comal">{{cite journal |vauthors=Prüfer K, Racimo F, Patterson N, Jay F, Sankararaman S, Sawyer S, Heinze A, Renaud G, Sudmant PH, de Filippo C, Li H, Mallick S, Dannemann M, Fu Q, Kircher M, Kuhlwilm M, Lachmann M, Meyer M, Ongyerth M, Siebauer M, Theunert C, Tandon A, Moorjani P, Pickrell J, Mullikin JC, Vohr SH, Green RE, Hellmann I, Johnson PL, Blanche H, Cann H, Kitzman JO, Shendure J, Eichler EE, Lein ES, Bakken TE, Golovanova LV, Doronichev VB, Shunkov MV, Derevianko AP, Viola B, Slatkin M, Reich D, Kelso J, Pääbo S |display-authors=6 |title=The complete genome sequence of a Neanderthal from the Altai Mountains |journal=[[Nature (journal)|Nature]] |volume=505 |issue=7481 |pages=43–49 |date=January 2014 |pmid=24352235 |pmc=4031459 |doi=10.1038/nature12886 |orig-date=Online 2013 |bibcode=2014Natur.505...43P}}</ref><ref name="lac12adaafr">{{cite journal |vauthors=Lachance J, Vernot B, Elbers CC, Ferwerda B, Froment A, Bodo JM, Lema G, Fu W, Nyambo TB, Rebbeck TR, Zhang K, Akey JM, Tishkoff SA |display-authors=6 |title=Evolutionary history and adaptation from high-coverage whole-genome sequences of diverse African hunter-gatherers |journal=[[Cell (journal)|Cell]] |volume=150 |issue=3 |pages=457–469 |date=August 2012 |pmid=22840920 |pmc=3426505 |doi=10.1016/j.cell.2012.07.009}}</ref><ref name="hamgenev">{{cite journal |vauthors=Hammer MF, Woerner AE, Mendez FL, Watkins JC, Wall JD |title=Genetic evidence for archaic admixture in Africa |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=108 |issue=37 |pages=15123–15128 |date=September 2011 |pmid=21896735 |pmc=3174671 |doi=10.1073/pnas.1109300108 |bibcode=2011PNAS..10815123H |doi-access=free}}</ref> indicating that modern population groups, while mostly derived from early ''H. sapiens'', are to a lesser extent also descended from regional variants of archaic humans.
and non-African, while mostly derived from early ''H. sapiens'', to a lesser extent are also descended from regional variants of [[archaic humans]].


==Proposed waves==
== Proposed waves ==
:''See [[Early hominin expansions out of Africa]] for archaic humans (''H. erectus, H. heidelbergensis, Neanderthals, Denisovans).''
{{see|Early expansions of hominins out of Africa|Skhul and Qafzeh hominins}}
[[File:Ksar Akil Fossils.jpg|thumb|upright=1.3|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="Douka_2013" /> and 42,400–41,700 BP for "Ethelruda".<ref name="Douka_2013">{{cite journal |vauthors=Douka K, Bergman CA, Hedges RE, Wesselingh FP, Higham TF |title=Chronology of Ksar Akil (Lebanon) and implications for the colonization of Europe by anatomically modern humans |journal=[[PLOS ONE]] |volume=8 |issue=9 |pages=e72931 |date=2013-09-11 |pmid=24039825 |pmc=3770606 |doi=10.1371/journal.pone.0072931 |bibcode=2013PLoSO...872931D |doi-access=free}}</ref>]]
{{see|Skhul and Qafzeh hominins}}
"Recent African origin," or ''Out of Africa II'', refers to the migration of [[anatomically modern humans]] (''[[Homo sapiens]]'') out of Africa after their emergence at c. 300,000 to 200,000 years ago, in contrast to "[[Out of Africa I]]", the migration of archaic humans from Africa to Eurasia between roughly 1.8 to 0.5 million years ago.
"Recent African origin", or ''Out of Africa II'', refers to the migration of [[anatomically modern humans]] (''[[Homo sapiens]]'') out of Africa after their emergence at c. 300,000 to 200,000 years ago, in contrast to "[[Out of Africa I]]", which refers to the migration of archaic humans from Africa to Eurasia from before 1.8 and up to 0.5 million years ago. [[Omo remains|Omo-Kibish I (Omo I)]] from southern Ethiopia is the oldest anatomically modern Homo sapiens skeleton currently known
(around 233,000 years old).<ref name="Vidal22">{{cite journal |vauthors=Vidal CM, Lane CS, Asrat A, Barfod DN, Mark DF, Tomlinson EL, Tadesse AZ, Yirgu G, Deino A, Hutchison W, Mounier A, Oppenheimer C |display-authors=6 |title=Age of the oldest known Homo sapiens from eastern Africa |journal=[[Nature (journal)|Nature]] |volume=601 |issue=7894 |pages=579–583 |date=January 2022 |pmid=35022610 |pmc=8791829 |doi=10.1038/s41586-021-04275-8 |bibcode=2022Natur.601..579V}}</ref> There are even older ''Homo sapiens'' fossils from [[Jebel Irhoud]] in Morocco which exhibit a mixture of modern and archaic features at around 315,000 years old.<ref>{{cite journal |vauthors=Hublin JJ, Ben-Ncer A, Bailey SE, Freidline SE, Neubauer S, Skinner MM, Bergmann I, Le Cabec A, Benazzi S, Harvati K, Gunz P |display-authors=6 |title=New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens |journal=[[Nature (journal)|Nature]] |volume=546 |issue=7657 |pages=289–292 |date=June 2017 |pmid=28593953 |doi=10.1038/nature22336 |bibcode=2017Natur.546..289H |s2cid=256771372 |url=https://kar.kent.ac.uk/62267/1/Submission_288356_1_art_file_2637492_j96j1b.pdf}}</ref>


Since the early 21st century, the picture of "recent single-origin" migrations has become significantly more complex, not just due to the discovery of modern-archaic admixture but also due to the increasing evidence that the "recent out-of-Africa" migration took place in a number of waves spread over a long time period.
Since the beginning of the 21st century, the picture of "recent single-origin" migrations has become significantly more complex, due to the discovery of modern-archaic admixture and the increasing evidence that the "recent out-of-Africa" migration took place in waves over a long time. As of 2010, there were two main accepted dispersal routes for the out-of-Africa migration of early anatomically modern humans, the "Northern Route" (via Nile Valley and Sinai) and the "Southern Route" via the [[Bab-el-Mandeb]] strait.{{sfnp|Beyin|2011}}
As of 2010, there were two main accepted dispersal route for the out-of-Africa migration of early anatomically modern humans: via the "Northern Route" (via Nile Valley and Sinai) and the "Southern Route"
via the [[Bab al Mandab]] strait.{{sfnp|Beyin|2011}}


*Posth et al. (2017) suggest that early ''Homo sapiens'', or "another species in Africa closely related to us," might have first migrated out of Africa around 270,000 years ago.<ref name="NC-20170704">{{cite journal |author=Posth, Cosimo et al. |title=Deeply divergent archaic mitochondrial genome provides lower time boundary for African gene flow into Neanderthals |url=https://www.nature.com/articles/ncomms16046 |date=4 July 2017 |journal=[[Nature Communications]] |doi=10.1038/ncomms16046 |accessdate=4 July 2017 |volume=8 |page=16046}}; see also {{cite news |last=Zimmer |first=Carl |authorlink=Carl Zimmer |title=In Neanderthal DNA, Signs of a Mysterious Human Migration |url=https://www.nytimes.com/2017/07/04/science/neanderthals-dna-homo-sapiens-human-evolution.html |date=4 July 2017 |work=[[New York Times]] |accessdate=4 July 2017 }}.</ref>
* Posth et al. (2017) suggest that early ''Homo sapiens'', or "another species in Africa closely related to us", might have first migrated out of Africa around 270,000 years ago based on the closer affinity within Neanderthals' [[Mitochondrial DNA|mitochondrial genomes]] to ''Homo sapiens'' than [[Denisovans]].<ref name="NC-20170704">{{cite journal |vauthors=Posth C, Wißing C, Kitagawa K, Pagani L, van Holstein L, Racimo F, Wehrberger K, Conard NJ, Kind CJ, Bocherens H, Krause J |display-authors=6 |title=Deeply divergent archaic mitochondrial genome provides lower time boundary for African gene flow into Neanderthals |journal=[[Nature Communications]] |volume=8 |pages=16046 |date=July 2017 |pmid=28675384 |pmc=5500885 |doi=10.1038/ncomms16046 |bibcode=2017NatCo...816046P}}; see also {{cite news |vauthors=Zimmer C |author-link=Carl Zimmer |title=In Neanderthal DNA, Signs of a Mysterious Human Migration |url=https://www.nytimes.com/2017/07/04/science/neanderthals-dna-homo-sapiens-human-evolution.html |date=4 July 2017 |work=[[The New York Times]] |access-date=4 July 2017}}.</ref>
*Finds at [[Misliya cave]], which include a partial jawbone with eight teeth have been dated to around 185,000 years ago. Layers dating from between 250,000 and 140,000 years ago in the same cave contained tools of the [[Levallois technique|Levallois]] type which could put the date of the first migration even earlier if the tools can be associated with the modern human jawbone finds.<ref>{{Cite news|url=https://www.sciencedaily.com/releases/2018/01/180125140923.htm|title=Scientists discover oldest known modern human fossil outside of Africa: Analysis of fossil suggests Homo sapiens left Africa at least 50,000 years earlier than previously thought|work=ScienceDaily|access-date=2018-01-27|language=en}}</ref><ref>{{Cite news|url=http://www.bbc.co.uk/news/science-environment-42817323|title=Modern humans left Africa much earlier|last=Ghosh|first=Pallab|date=2018|work=BBC News|access-date=2018-01-27|language=en-GB}}</ref>
*An Eastward Dispersal from Northeast Africa to Arabia during 150–130 kya based on the finds at [[Jebel Faya]] dated to 127 kya (discovered in 2011).{{r|pmid21273486}}{{r|pmid21212332}} Possibly related to this wave are the finds from [[Zhirendong]] cave, Southern China, dated to more than 100 kya.{{sfnp|Beyin|2011}} Other evidence of modern human presence in China has been dated to 80,000 years ago.{{r|Liu2015}}
*The most significant dispersal took place around 70,000 years ago via the so-called [[Southern Dispersal|Southern Route]], either before{{sfnp|Appenzeller|2012}} or after{{r|Macaulay2005|Posth2016}} the [[Toba catastrophe theory|Toba event]], which happened between 69,000 and 77,000 years ago.{{sfnp|Appenzeller|2012}} This dispersal followed the southern coastline of Asia, and reached Australia around 65,000-50,000 years ago. Western Asia was "re-occupied" by a different derivation from this wave around 50,000 years ago, and Europe was populated from Western Asia beginning around 43,000 years ago.{{sfnp|Beyin|2011}}
*{{harvp|Wells|2003}} describes an additional wave of migration after the southern coastal route, namely a northern migration into Europe at circa 45,000 years ago.{{refn|group=note|name=McChesney2015|Kay Young McChesney: "Wells (2003) divided the descendants of men who left Africa into a genealogical tree with 11 lineages. Each genetic marker represents a single-point mutation (SNP) at a specific place in the genome. First, genetic evidence suggests that a small band with the marker M168 migrated out of Africa along the coasts of the Arabian Peninsula and India, through Indonesia, and reached Australia very early, between 60,000 and 50,000 years ago. This very early migration into Australia is also supported by Rasmussen et al. (2011). Second, a group bearing the marker M89 moved out of northeastern Africa into the Middle East 45,000 years ago. From there, the M89 group split into two groups. One group that developed the marker M9 went into Asia about 40,000 years ago. The Asian (M9) group split three ways: into Central Asia (M45), 35,000 years ago; into India (M20), 30,000 years ago; and into China (M122), 10,000 years ago. The Central Asian (M45) group split into two groups: toward Europe (M173), 30,000 years ago and toward Siberia (M242), 20,000 years ago. Finally, the Siberian group (M242) went on to populate North and South America (M3), about 10,000 years ago.{{sfn|Young McChesney|2015}}}} This possibility is ruled out by {{harvp|Macaulay et al.|2005}} and {{harvp|Posth et al.|2016}}, arguing for a single coastal dispersal, with an early offshoot into Europe.


* Fossil evidence also points to an early ''Homo sapiens'' migration with the oldest known fossil coming from [[Apidima Cave]] in Greece and dated at 210,000 years ago.<ref>{{cite journal |vauthors=Harvati K, Röding C, Bosman AM, Karakosti FA, Grün R, Stringer C, Karkanas P, Thompson NC, Koutoulidis V, Moulopoulous LA, Gorgoulis VG, Kouloukoussa M |display-authors=6 |title=Apidima Cave fossils provide earliest evidence of Homo sapiens in Eurasia |journal=[[Nature (journal)|Nature]] |volume=571 |pages=500–504 |date=July 2019 |issue=7766 |doi=10.1038/s41586-019-1376-z|pmid=31292546 |s2cid=195873640 |url=https://zenodo.org/record/6646855}}</ref> Finds at [[Misliya cave]], which include a partial jawbone with eight teeth, have been dated to around 185,000 years ago. Layers dating from between 250,000 and 140,000 years ago in the same cave contained tools of the [[Levallois technique|Levallois]] type which could put the date of the first migration even earlier if the tools can be associated with the modern human jawbone finds.<ref>{{Cite news|url=https://www.sciencedaily.com/releases/2018/01/180125140923.htm |title=Scientists discover oldest known modern human fossil outside of Africa: Analysis of fossil suggests Homo sapiens left Africa at least 50,000 years earlier than previously thought |work=ScienceDaily |access-date=2018-01-27 |language=en}}</ref><ref>{{Cite news |url=https://www.bbc.co.uk/news/science-environment-42817323 |title=Modern humans left Africa much earlier |vauthors=Ghosh P |date=2018 |work=[[BBC News]] |access-date=2018-01-27 |language=en-GB}}</ref><ref>{{cite journal |vauthors=Hershkovitz I, Weber GW, Quam R, Duval M, Grün R, Kinsley L, Ayalon A, Bar-Matthews M, Valladas H, Mercier N, Arsuaga JL, Martinón-Torres M, Bermúdez de Castro JM, Fornai C, Martín-Francés L, Sarig R, May H, Krenn VA, Slon V, Rodríguez L, García R, Lorenzo C, Carretero JM, Frumkin A, Shahack-Gross R, Bar-Yosef Mayer DE, Cui Y, Wu X, Peled N, Groman-Yaroslavski I, Weissbrod L, Yeshurun R, Tsatskin A, Zaidner Y, Weinstein-Evron M |display-authors=6 |title=The earliest modern humans outside Africa |journal=[[Science (journal)|Science]] |volume=359 |issue=6374 |pages=456–459 |date=January 2018 |pmid=29371468 |doi=10.1126/science.aap8369 |bibcode=2018Sci...359..456H |hdl=10072/372670 |s2cid=206664380 |hdl-access=free}}</ref>
==Northern Route dispersal{{anchor|Early northern Africa dispersal}}==
{{see|Skhul and Qafzeh hominins}}
Beginning 135,000 years ago, tropical Africa experienced [[megadrought]]s which drove the humans from the land and towards the sea shores, and forced them to cross over to other continents.<ref name="U of AZ"/>{{refn|group=note|The researchers used radiocarbon dating techniques on pollen grains trapped in lake-bottom mud to establish vegetation over the ages of the [[Malawi]] lake in Africa, taking samples at 300-year-intervals. Samples from the megadrought times had little pollen or charcoal, suggesting sparse vegetation with little to burn. The area around [[Lake Malawi]], today heavily forested, was a desert approximately 135,000 to 90,000 years ago.<ref name="U of AZ">{{cite web | url=https://uanews.arizona.edu/story/newfound-ancient-african-megadroughts-may-have-driven-evolution-of-humans-and-fish| title= Newfound Ancient African Megadroughts May Have Driven Evolution of Humans and Fish. The findings provide new insights into humans' migration out of Africa and the evolution of fishes in Africa's Great Lakes.| publisher=The University of Arizona| language=English|date=8 October 2007 | accessdate=25 September 2017 | author=Mari N. Jensen}}</ref>}}


* An eastward dispersal from Northeast Africa to Arabia 150,000–130,000 years ago is based on the stone tools finds at [[Jebel Faya]] dated to 127,000 years ago (discovered in 2011), although fossil evidence in the area is significantly later at 85,000 years ago.<ref name=pmid21273486 /> <ref>{{cite journal |vauthors=Groucutt HS, Grün R, Zalmout IS, Drake NA, Armitage SJ, Candy I, Clark-Wilson R, Louys J, Breeze PS, Duval M, Buck LT, Kivell TL, Pomeroy E, Stephens NB, Stock JT, Stewart M, Price GJ, Kinsley L, Sung WW, Alsharekh A, Al-Omari A, Zahir M, Memesh AM, Abdulshakoor AJ, Al-Masari AM, Bahameem AA, Al Murayyi KS, Zahrani B, Scerri EM, Petraglia MD |display-authors=6 |title=Homo sapiens in Arabia by 85,000 years ago |journal=[[Nature Ecology & Evolution]] |volume=2 |pages=800–809 |date=May 2018 |issue=5 |doi=10.1038/s41559-018-0518-2 |pmid=29632352 |pmc=5935238|bibcode=2018NatEE...2..800G }}</ref> Possibly related to this wave are the finds from [[Zhirendong]] cave, Southern China, dated to more than 100,000 years ago.<ref>{{cite journal |vauthors=Liu W, Jin CZ, Zhang YQ, Cai YJ, Xing S, Wu XJ, Cheng H, Edwards RL, Pan WS, Qin DG, An ZS, Trinkaus E, Wu XZ |display-authors=6 |title=Human remains from Zhirendong, South China, and modern human emergence in East Asia |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=107 |issue=45 |pages=19201–19206 |date=November 2010 |pmid=20974952 |pmc=2984215 |doi=10.1073/pnas.1014386107 |bibcode=2010PNAS..10719201L |doi-access=free}}</ref> Other evidence of modern human presence in China has been dated to 80,000 years ago.{{r|Liu2015}}
Modern humans crossed the Straits of Bab el Mandab in the southern [[Red Sea]], and moved along the green coastlines around Arabia, and thence to the rest of Eurasia. Fossils of early ''Homo sapiens'' were found in [[Qafzeh]] cave in Israel and have been dated 80,000 to 100,000 years ago. These humans seem to have either become extinct or retreated back to Africa 70,000 to 80,000 years ago, possibly replaced by southbound Neanderthals escaping the colder regions of ice-age Europe.{{sfnp|Finlayson|2009|p=68}} Hua Liu ''et al.'' analyzed autosomal microsatellite markers dating to about 56,000 years ago. They interpret the [[paleontological]] fossil as an isolated early offshoot that retracted back to Africa.{{sfnp|Liu, Prugnolle et al.|2006}}

* The most significant out of Africa dispersal took place around 50,000–70,000 years ago via the so-called [[Southern Dispersal|Southern Route]], either before{{sfnp|Appenzeller|2012}} or after{{r|Macaulay2005|Posth2016}} the [[Toba catastrophe theory|Toba event]], which happened between 69,000 and 77,000 years ago.{{sfnp|Appenzeller|2012}} This dispersal followed the southern coastline of Asia and reached Australia around 65,000–50,000 years ago or according to some research, by 50,000 years ago at earliest.<ref name="Wood_2017" /><ref name="Homo sapiens first reach Southeast" /> Western Asia was "re-occupied" by a different derivation from this wave around 50,000 years ago and Europe was populated from Western Asia beginning around 43,000 years ago.{{sfnp|Beyin|2011}}

* {{harvp|Wells|2003}} describes an additional wave of migration after the southern coastal route, a northern migration into Europe about 45,000 years ago.{{Efn|{{harvp|McChesney|2015}}: "Wells (2003) divided the descendants of men who left Africa into a genealogical tree with 11 lineages. Each genetic marker represents a single-point mutation (SNP) at a specific place in the genome. First, genetic evidence suggests that a small band with the marker M168 migrated out of Africa along the coasts of the Arabian Peninsula and India, through Indonesia, and reached Australia very early, between 60,000 and 50,000 years ago. This very early migration into Australia is also supported by Rasmussen et al. (2011). Second, a group bearing the marker M89 moved out of northeastern Africa into the Middle East 45,000 years ago. From there, the M89 group split into two groups. One group that developed the marker M9 went into Asia about 40,000 years ago. The Asian (M9) group split three ways: into Central Asia (M45), 35,000 years ago; into India (M20), 30,000 years ago; and into China (M122), 10,000 years ago. The Central Asian (M45) group split into two groups: toward Europe (M173), 30,000 years ago and toward Siberia (M242), 20,000 years ago. Finally, the Siberian group (M242) went on to populate North and South America (M3), about 10,000 years ago.{{sfn|McChesney|2015}}}} This possibility is ruled out by {{harvp|Macaulay et al.|2005}} and {{harvp|Posth et al.|2016}}, who argue for a single coastal dispersal, with an early offshoot into Europe.

== Northern Route dispersal{{anchor|Early northern Africa dispersal}} ==
{{see|Skhul and Qafzeh hominins}}
[[File:Anatomically Modern Humans archaeological remains, Europe and Africa, directly dated, calibrated carbon dates as of 2013.jpg|thumb|270px|Anatomically Modern Humans known archaeological remains in Europe and Africa, directly dated, calibrated carbon dates as of 2013.<ref name="Douka_2013" />]]
Beginning 135,000 years ago, tropical Africa experienced [[megadrought]]s which drove humans from the land and towards the sea shores, and forced them to cross over to other continents.<ref name="U of AZ" />{{Efn|The researchers used radiocarbon dating techniques on pollen grains trapped in lake-bottom mud to establish vegetation over the ages of the [[Malawi]] lake in Africa, taking samples at 300-year-intervals. Samples from the megadrought times had little pollen or charcoal, suggesting sparse vegetation with little to burn. The area around [[Lake Malawi]], today heavily forested, was a desert approximately 135,000 to 90,000 years ago.<ref name="U of AZ">{{cite web |url=https://uanews.arizona.edu/story/newfound-ancient-african-megadroughts-may-have-driven-evolution-of-humans-and-fish |title=Newfound Ancient African Megadroughts May Have Driven Evolution of Humans and Fish. The findings provide new insights into humans' migration out of Africa and the evolution of fishes in Africa's Great Lakes |publisher=The [[University of Arizona]] |language=en |date=8 October 2007 |access-date=25 September 2017 |vauthors=Jensen MN}}</ref>}}


Fossils of early ''Homo sapiens'' were found in [[Qafzeh]] and [[Es-Skhul Cave]]s in Israel and have been dated to 80,000 to 120,000 years ago.<ref>{{cite journal |vauthors=Stringer CB, Grün R, Schwarcz HP, Goldberg P |title=ESR dates for the hominid burial site of Es Skhul in Israel |journal=[[Nature (journal)|Nature]] |volume=338 |issue=6218 |pages=756–758 |date=April 1989 |pmid=2541339 |doi=10.1038/338756a0 |bibcode=1989Natur.338..756S |s2cid=4332370}}</ref><ref>{{cite journal |vauthors=Grün R, Stringer C, McDermott F, Nathan R, Porat N, Robertson S, Taylor L, Mortimer G, Eggins S, McCulloch M |display-authors=6 |title=U-series and ESR analyses of bones and teeth relating to the human burials from Skhul |journal=[[Journal of Human Evolution]] |volume=49 |issue=3 |pages=316–334 |date=September 2005 |pmid=15970310 |doi=10.1016/j.jhevol.2005.04.006|bibcode=2005JHumE..49..316G }}</ref> These humans seem to have either become extinct or retreated back to Africa 70,000 to 80,000 years ago, possibly replaced by southbound Neanderthals escaping the colder regions of ice-age Europe.{{sfnp|Finlayson|2009|p=68}} Hua Liu ''et al.'' analyzed autosomal microsatellite markers dating to about 56,000 years ago. They interpret the [[paleontological]] fossil as an isolated early offshoot that retracted back to Africa.{{sfnp|Liu, Prugnolle et al.|2006}}
The discovery of stone tools in the [[United Arab Emirates]] in 2011 indicated the presence of modern humans at least 100,000 and 125,000 years ago,{{r|pmid21273486}} leading to a resurgence of the "long-neglected" North African route.{{r|pmid21212332}}<ref>{{cite journal|title = Earliest evidence for the structure of ''Homo sapiens'' populations in Africa|url = http://www.sciencedirect.com/science/article/pii/S0277379114003023|journal = Quaternary Science Reviews|date = 1 October 2014|pages = 207–216|volume = 101|doi = 10.1016/j.quascirev.2014.07.019|first = Eleanor M. L.|last = Scerri|first2 = Nick A.|last2 = Drake|first3 = Richard|last3 = Jennings|first4 = Huw S.|last4 = Groucutt}}</ref>{{r|pmid21601174|pmid17372199}}


The discovery of stone tools in the [[United Arab Emirates]] in 2011 at the [[Jebel Faya|Faya-1 site]] in [[Mleiha Archaeological Centre|Mleiha]], [[Emirate of Sharjah|Sharjah]], indicated the presence of modern humans at least 125,000 years ago,{{r|pmid21273486}} leading to a resurgence of the "long-neglected" North African route.{{r|pmid21212332}}<ref>{{cite journal | last1=Scerri | first1=Eleanor M.L. | last2=Drake | first2=Nick A. | last3=Jennings | first3=Richard | last4=Groucutt | first4=Huw S. | title=Earliest evidence for the structure of Homo sapiens populations in Africa | journal=Quaternary Science Reviews | volume=101 | date=2014 | doi=10.1016/j.quascirev.2014.07.019 | pages=207–216|bibcode=2014QSRv..101..207S}}</ref>{{r|pmid21601174|pmid17372199}} This new understanding of the role of the Arabian dispersal began to change following results from archaeological and genetic studies stressing the importance of southern Arabia as a corridor for human expansions out of Africa.<ref>{{cite journal | last1=Bretzke | first1=Knut | last2=Armitage | first2=Simon J. | last3=Parker | first3=Adrian G. | last4=Walkington | first4=Helen | last5=Uerpmann | first5=Hans-Peter | title=The environmental context of Paleolithic settlement at Jebel Faya, Emirate Sharjah, UAE | journal=Quaternary International | volume=300 | date=2013 | doi=10.1016/j.quaint.2013.01.028 | pages=83–93|bibcode=2013QuInt.300...83B}}</ref>
In [[History of Oman|Oman]], a site was discovered by Bien Joven in 2011 containing more than 100 surface scatters of stone tools belonging to the late Nubian Complex, known previously only from [[Archaeological record|archaeological excavations]] in the [[Sudan]]. Two optically stimulated luminescence age estimates place the Arabian Nubian Complex at approximately 106,000 years old. This provides evidence for a distinct [[stone age]] technocomplex in southern Arabia, around the earlier part of the [[Marine Isotope Stage 5]].<ref>{{cite journal | pmc = 3227647 | pmid=22140561 | doi=10.1371/journal.pone.0028239 | volume=6 | title=The Nubian Complex of Dhofar, Oman: an African middle stone age industry in Southern Arabia | year=2011 | journal=PLoS ONE | pages=e28239 | last1 = Rose | first1 = JI | last2 = Usik | first2 = VI | last3 = Marks | first3 = AE | last4 = Hilbert | first4 = YH | last5 = Galletti | first5 = CS | last6 = Parton | first6 = A | last7 = Geiling | first7 = JM | last8 = Cerný | first8 = V | last9 = Morley | first9 = MW | last10 = Roberts | first10 = RG}}</ref>


In [[History of Oman|Oman]], a site was discovered by Bien Joven in 2011 containing more than 100 surface scatters of stone tools belonging to the late Nubian Complex, known previously only from [[Archaeological record|archaeological excavations]] in the [[Sudan]]. Two optically stimulated luminescence age estimates placed the Arabian Nubian Complex at approximately 106,000 years old. This provides evidence for a distinct [[Stone Age]] technocomplex in southern Arabia, around the earlier part of the [[Marine Isotope Stage 5]].<ref>{{cite journal |vauthors=Rose JI, Usik VI, Marks AE, Hilbert YH, Galletti CS, Parton A, Geiling JM, Cerný V, Morley MW, Roberts RG |display-authors=6 |title=The Nubian Complex of Dhofar, Oman: an African middle Stone Age industry in Southern Arabia |journal=[[PLOS ONE]] |volume=6 |issue=11 |pages=e28239 |year=2011 |pmid=22140561 |pmc=3227647 |doi=10.1371/journal.pone.0028239 |bibcode=2011PLoSO...628239R |doi-access=free}}</ref>
According to Kuhlwilm and his co-authors, [[Neanderthal]]s [[Archaic human admixture with modern humans|contributed to modern humans genetically]] around 100,000 years ago, from humans which split off from other modern humans around 200,000 years ago.<ref name=DAB-EastNeandertals>{{harvp|Kuhlwilm et al.|2016}}.<br>See also [http://dienekes.blogspot.nl/2016/02/ancestors-of-eastern-neandertals.html ''Ancestors of Eastern Neandertals admixed with modern humans 100 thousand years ago''], Dienekes'Anthropology Blog.</ref> They found that "the ancestors of Neanderthals from the Altai Mountains and early modern humans met and interbred, possibly in the Near East, many thousands of years earlier than previously thought".{{r|DAB-EastNeandertals}} According to co-author Ilan Gronau, "This actually complements archaeological evidence of the presence of early modern humans out of Africa around and before 100 ka by providing the first genetic evidence of such populations."{{r|DAB-EastNeandertals}} Similar [[genetic admixture]] events have been [[Archaic human admixture with modern humans|noted in other regions]] as well.<ref name="Denovisans">{{Cite journal|url =http://www.sciencemag.org/news/2017/03/ancient-skulls-may-belong-elusive-humans-called-denisovans |title = Ancient skulls may belong to elusive humans called Denisovans|last = Gibbons|first = Ann|date = 2 March 2017|journal = Science |accessdate = 25 September 2017}}</ref>


According to Kuhlwilm and his co-authors, [[Neanderthal]]s [[Archaic human admixture with modern humans|contributed genetically]] to modern humans then living outside of Africa around 100,000 years ago: humans which had already split off from other modern humans around 200,000 years ago, and this early wave of modern humans outside Africa also contributed genetically to the Altai Neanderthals.<ref name="DAB-EastNeandertals">{{harvp|Kuhlwilm et al.|2016}}.<br />See also [http://dienekes.blogspot.nl/2016/02/ancestors-of-eastern-neandertals.html ''Ancestors of Eastern Neandertals admixed with modern humans 100 thousand years ago''], Dienekes'Anthropology Blog.</ref> They found that "the ancestors of Neanderthals from the [[Altai Mountains]] and early modern humans met and interbred, possibly in the Near East, many thousands of years earlier than previously thought".{{r|"DAB-EastNeandertals"}} According to co-author Ilan Gronau, "This actually complements archaeological evidence of the presence of early modern humans out of Africa around and before 100,000 years ago by providing the first genetic evidence of such populations."{{r|"DAB-EastNeandertals"}} Similar [[genetic admixture]] events have been [[Archaic human admixture with modern humans|noted in other regions]] as well.<ref name="Denovisans">{{Cite journal |url=https://www.science.org/content/article/ancient-skulls-may-belong-elusive-humans-called-denisovans |title=Ancient skulls may belong to elusive humans called Denisovans |vauthors=Gibbons A |date=2 March 2017 |journal=[[Science (journal)|Science]] |doi=10.1126/science.aal0846 |access-date=25 September 2017}}</ref>
In China, the [[Liujiang man]] ({{zh|c=柳江人}}) is among the earliest modern humans found in [[East Asia]].<ref name="liujiang hominid site">{{Cite journal | last1 = Shen | first1 = G. | last2 = Wang | first2 = W. | last3 = Wang | first3 = Q. | last4 = Zhao | first4 = J. | last5 = Collerson | first5 = K. | last6 = Zhou | first6 = C. | last7 = Tobias | first7 = P. V. | title = U-Series dating of Liujiang hominid site in Guangxi, Southern China | doi = 10.1006/jhev.2002.0601 | journal = Journal of Human Evolution | volume = 43 | issue = 6 | pages = 817–829 | year = 2002 | pmid = 12473485| pmc = }}</ref>
The date most commonly attributed to the remains is 67,000 years ago.<ref name=":0">{{Cite journal|url = |title = A Late Pleistocene Human Skeleton from Liujiang, China Suggests Regional Population Variation in Sexual Dimorphism in the Human Pelvis|last = Rosenburg|first = Karen|date = 2002|journal = Variability and Evolution|doi = |pmid = |access-date = }}</ref> High rates of variability yielded by various dating techniques carried out by different researchers place the most widely accepted range of dates with 67,000 BP as a minimum, but does not rule out dates as old as 159,000 BP.<ref name=":0" /> {{harvp|Liu, Martinón-Torres et al.|2015}} claim that modern human teeth have been found in China dating to at least 80,000 years ago.<ref name=":1">{{Cite journal|url = |title = Fossil Record of Early Modern Humans in East Asia|last = Kaifu|first = Yousuke|date = 2012|journal = Palaeoenvironmental Changes and Human Dispersals in North and East Asia during MIS3 and MIS2|doi = |pmid = |access-date = }}</ref>


==Southern Route dispersal{{anchor|Southern Dispersal}}==
== Southern Route dispersal{{anchor|Southern Dispersal}} ==
{{main|Great Coastal Migration}}
{{main|Southern Dispersal}}
{{see|List of first human settlements}}
{{see|List of first human settlements}}

===Coastal route===
=== Coastal route ===
[[File:Red Sea2.png|thumb|[[Red Sea]] crossing]]
[[File:Red Sea2.png|thumb|[[Red Sea]] crossing]]
By some 70,000 years ago, a part of the bearers of mitochondrial haplogroup [[Haplogroup L3 (mtDNA)|L3]] migrated from [[East Africa]] into the [[Near East]]. It has been estimated that from a population of 2,000 to 5,000 individuals in Africa, only a small group, possibly as few as 150 to 1,000 people, crossed the Red Sea.<ref>{{cite journal|year=2003|last=Zhivotovsky|title=Features of Evolution and Expansion of Modern Humans, Inferred from Genomewide Microsatellite Markers | pmc=1180270 |volume=72 |issue=5 |pmid=12690579 |last2=Rosenberg |first2=NA| last3=Feldman |first3=MW |pages=1171–86 |doi=10.1086/375120|journal=American Journal of Human Genetics|display-authors=etal}}</ref><ref>{{cite web |year=2008 |first=Gary |last=Stix |url=http://www.scientificamerican.com/article.cfm?id=the-migration-history-of-humans|title=The Migration History of Humans: DNA Study Traces Human Origins Across the Continents|accessdate=14 June 2011}}</ref> The group that crossed the Red Sea travelled along the coastal route around [[Arabia]] and [[Persia]] to India, which appears to be the first major settling point.{{r|pmid15339343}} {{harvp|Wells|2003}} argued for the route along the southern coastline of Asia, across about {{convert|250|km|0|abbr=out}}{{dubious|date=February 2018}}, reaching Australia by around 50,000 years ago.
By some 50–70,000 years ago, a subset of the bearers of mitochondrial haplogroup [[Haplogroup L3 (mtDNA)|L3]] migrated from [[East Africa]] into the [[Near East]]. It has been estimated that from a population of 2,000 to 5,000 individuals in Africa, only a small group, possibly as few as 150 to 1,000 people, crossed the Red Sea.<ref>{{cite journal |vauthors=Zhivotovsky LA, Rosenberg NA, Feldman MW |title=Features of evolution and expansion of modern humans, inferred from genomewide microsatellite markers |journal=[[American Journal of Human Genetics]] |volume=72 |issue=5 |pages=1171–1186 |date=May 2003 |pmid=12690579 |pmc=1180270 |doi=10.1086/375120}}</ref><ref>{{cite web |year=2008 |vauthors=Stix G |url=http://www.scientificamerican.com/article.cfm?id=the-migration-history-of-humans |title=The Migration History of Humans: DNA Study Traces Human Origins Across the Continents |website=[[Scientific American]] |access-date=14 June 2011}}</ref> The group that crossed the Red Sea travelled along the coastal route around [[Arabia]] and the [[Persian Plateau]] to India, which appears to have been the first major settling point.{{r|pmid15339343}} {{harvp|Wells|2003}} argued for the route along the southern coastline of Asia, across about {{convert|250|km|0|abbr=out}}, reaching Australia by around 50,000 years ago.
[[File:Migration routes of modern humans (2023).png|thumb|Migration routes of modern humans, showing the northern route populating Western Eurasia, and the southern/coastal route populating Eastern Eurasia.]]
Today at the [[Bab-el-Mandeb straits]], the [[Red Sea]] is about {{convert|20|km|0|abbr=out}} wide, but 50,000 years ago sea levels were {{convert|70|m|0|abbr=on}} lower (owing to glaciation) and the water channel was much narrower. Though the straits were never completely closed, they were narrow enough to have enabled crossing using simple rafts, and there may have been islands in between.{{sfnp|Beyin|2011}}<ref>{{cite journal |vauthors=Fernandes CA, Rohling EJ, Siddall M |title=Absence of post-Miocene Red Sea land bridges: biogeographic implications |date=June 2006|doi=10.1111/j.1365-2699.2006.01478.x |journal=[[Journal of Biogeography]] |issue=6 |volume=33 |pages=961–966 |doi-access=free|bibcode=2006JBiog..33..961F }}</ref> Shell [[middens]] 125,000 years old have been found in [[Eritrea]],<ref name="pmid10811218">{{cite journal |vauthors=Walter RC, Buffler RT, Bruggemann JH, Guillaume MM, Berhe SM, Negassi B, Libsekal Y, Cheng H, Edwards RL, von Cosel R, Néraudeau D, Gagnon M |display-authors=6 |title=Early human occupation of the Red Sea coast of Eritrea during the last interglacial |journal=[[Nature (journal)|Nature]] |volume=405 |issue=6782 |pages=65–69 |date=May 2000 |pmid=10811218 |doi=10.1038/35011048 |bibcode=2000Natur.405...65W |s2cid=4417823}}</ref> indicating that the diet of early humans included seafood obtained by [[beachcombing]].


==== Toba eruption ====
Today at the [[Bab-el-Mandeb straits]], the [[Red Sea]] is about {{convert|20|km|0|abbr=out}} wide but 50,000 years ago sea levels were {{convert|70|m|0|abbr=on}} lower (owing to glaciation) and the water was much narrower. Though the straits were never completely closed, they were narrow enough and there may have been islands in between to have enabled crossing using simple rafts.<ref>{{cite journal|title = Absence of post-Miocene Red Sea land bridges: biogeographic implications|date = June 2006|doi = 10.1111/j.1365-2699.2006.01478.x|url = http://onlinelibrary.wiley.com/doi/10.1111/j.1365-2699.2006.01478.x/full|last = Fernandes et. al|journal = Journal of Biogeography|issue = 6|volume = 33|pages = 961–966}}</ref>{{sfnp|Beyin|2011}} Shell [[middens]] 125,000 years old have been found in [[Eritrea]],<ref name="pmid10811218">{{cite journal |vauthors=Walter RC, Buffler RT, Bruggemann JH, Guillaume MM, Berhe SM, Negassi B, Libsekal Y, Cheng H, Edwards RL, von Cosel R, Néraudeau D, Gagnon M | title = Early human occupation of the Red Sea coast of Eritrea during the last interglacial | journal = Nature | volume = 405 | issue = 6782 | pages = 65–9 |date=May 2000 | pmid = 10811218 | doi = 10.1038/35011048 | ref = harv }}</ref> indicating the diet of early humans included seafood obtained by [[beachcombing]].
{{Main|Toba catastrophe theory}}
The dating of the Southern Dispersal is a matter of dispute.{{sfnp|Appenzeller|2012}} It may have happened either pre- or post-Toba, a catastrophic volcanic eruption that took place between 69,000 and 77,000 years ago at the site of present-day [[Lake Toba]] in Sumatra, Indonesia. Stone tools discovered below the layers of ash deposited in India may point to a pre-Toba dispersal but the source of the tools is disputed.{{sfnp|Appenzeller|2012}} An indication for post-Toba is haplo-group L3, that originated before the dispersal of humans out of Africa and can be dated to 60,000–70,000 years ago, "suggesting that humanity left Africa a few thousand years after Toba".{{sfnp|Appenzeller|2012}} Some research showing slower than expected genetic mutations in human DNA was published in 2012, indicating a revised dating for the migration to between 90,000 and 130,000 years ago.<ref name="ourtruedawn">{{cite journal |vauthors=Catherine B |title=Our True Dawn |journal=[[New Scientist]] |issue=2892 |pages=34–37 |date=24 November 2012 |volume=216 |doi=10.1016/S0262-4079(12)63018-8 |bibcode=2012NewSc.216...34B |issn=0262-4079}}</ref> Some more recent research suggests a migration out-of-Africa of around 50,000-65,000 years ago of the ancestors of modern non-African populations, similar to most previous estimates.<ref name="Haber Jones etal 2019" /><ref>{{cite journal |vauthors=Karmin M, Saag L, Vicente M, Wilson Sayres MA, Järve M, etal |title=A recent bottleneck of Y chromosome diversity coincides with a global change in culture |journal=[[Genome Research]] |volume=25 |issue=4 |pages=459–466 |date=April 2015 |pmid=25770088 |pmc=4381518 |doi=10.1101/gr.186684.114}}</ref><ref name="Vai">{{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 |display-authors=6 |title=Ancestral mitochondrial N lineage from the Neolithic 'green' Sahara |journal=[[Scientific Reports]] |volume=9 |issue=1 |pages=3530 |date=March 2019 |pmid=30837540 |pmc=6401177 |doi=10.1038/s41598-019-39802-1 |bibcode=2019NatSR...9.3530V}}</ref>


=== Western Asia ===
{{See also|Toba catastrophe theory}}
Following the fossils dating 80,000 to 120,000 years ago from [[Qafzeh Cave|Qafzeh]] and [[Skhul Cave|Es-Skhul Caves]] in Israel there are no ''H. sapiens'' fossils in the [[Levant]] until the [[Manot 1]] fossil from [[Manot Cave]] in Israel, dated to 54,700 years ago,<ref>{{harvp|Hershkovitz et al.|2015}}<br />See also {{cite web |date=29 January 2015 |title=55,000-Year-Old Skull Fossil Sheds New Light on Human Migration out of Africa |url=http://www.sci-news.com/othersciences/anthropology/science-55000-year-old-skull-fossil-manot-cave-israel-02443.html |website=Science News}}</ref> though the dating was questioned by {{harvp|Groucutt et al.|2015}}. The lack of fossils and stone tool industries that can be safely associated with modern humans in the Levant has been taken to suggest that modern humans were outcompeted by Neanderthals until around 55,000 years ago, who would have placed a barrier on modern human dispersal out of Africa through the Northern Route.<ref>{{cite journal | last=Shea | first=John J. | title=Neandertals, competition, and the origin of modern human behavior in the Levant | journal=Evolutionary Anthropology: Issues, News, and Reviews | volume=12 | issue=4 | date=2003 | issn=1060-1538 | doi=10.1002/evan.10101 | pages=173–187}}</ref>{{fv|date=June 2023}} Climate reconstructions also support a Southern Route dispersal of modern humans as the [[Bab-el-Mandeb]] strait experienced a climate more conductive to human migration than the northern landbridge to the Levant during the major human dispersal out of Africa.<ref>{{cite journal |vauthors=Beyer RM, Krapp M, Eriksson A, Manica A |title=Climatic windows for human migration out of Africa in the past 300,000 years |journal=[[Nature Communications]] |volume=12 |issue=1 |pages=4889 |date=August 2021 |pmid=34429408 |pmc=8384873 |doi=10.1038/s41467-021-24779-1 |bibcode=2021NatCo..12.4889B}}</ref>
The dating of the Southern Dispersal is a matter of dispute.{{sfnp|Appenzeller|2012}} It may have happened either pre- or post-Toba, a catastrophic volcanic eruption that took place between 69,000 and 77,000 years ago at the site of present-day [[Lake Toba]]. Stone tools discovered below the layers of ash disposed in India may point to a pre-Toba dispersal but the source of the tools is disputed.{{sfnp|Appenzeller|2012}} An indication for post-Toba is haplo-group L3, that originated before the dispersal of humans out of Africa and can be dated to 60,000–70,000 years ago, "suggesting that humanity left Africa a few thousand years after Toba".{{sfnp|Appenzeller|2012}} New research showing slower than expected genetic mutations in human DNA was published in 2012, indicating a revised dating for the migration to between 90,000 and 130,000 years ago.<ref name="ourtruedawn">{{cite journal | author = Catherine Brahic | title = Our True Dawn | journal = New Scientist | issue = 2892 | pages = 34–7 | date = 24 November 2012 | ISSN= 0262-4079 | publisher = Reed Business Information }}</ref>


A 2023 study proposed that Eurasians and Africans genetically diverged ~100,000 years ago. Main Eurasians then lived in the Saudi Peninsula, genetically isolated from at least 85 kya, before expanding north 54 kya. For reference, Homo sapiens and Neanderthals diverged ~500 kya.<ref>{{Cite journal |last1=Tobler |first1=Raymond |last2=Souilmi |first2=Yassine |last3=Huber |first3=Christian D. |last4=Bean |first4=Nigel |last5=Turney |first5=Chris S. M. |last6=Grey |first6=Shane T. |last7=Cooper |first7=Alan |date=2023-05-30 |title=The role of genetic selection and climatic factors in the dispersal of anatomically modern humans out of Africa |journal=Proceedings of the National Academy of Sciences |language=en |volume=120 |issue=22 |pages=e2213061120 |doi=10.1073/pnas.2213061120 |issn=0027-8424 |pmc=10235988 |pmid=37220274|bibcode=2023PNAS..12013061T }}</ref>
===Western Asia===


=== Oceania ===
A fossil of a modern human dated to 54,700 years ago was found in [[Manot Cave]] in Israel, named [[Manot 1]],<ref>{{harvp|Hershkovitz et al.|2015}}<br>See also [http://www.sci-news.com/othersciences/anthropology/science-55000-year-old-skull-fossil-manot-cave-israel-02443.html 55,000-Year-Old Skull Fossil Sheds New Light on Human Migration out of Africa], Science News.</ref> though the dating was questioned by {{harvp|Groucutt et al.|2015}}.
It is thought that Australia was inhabited around 65,000–50,000 years ago. As of 2017, the earliest evidence of humans in Australia is at least 65,000 years old,<ref name=Clarkson2017/><ref name=StFleur2017 /> while McChesney stated that
{{blockquote|...genetic evidence suggests that a small band with the marker M168 migrated out of Africa along the coasts of the Arabian Peninsula and India, through Indonesia, and reached Australia very early, between 60,000 and 50,000 years ago. This very early migration into Australia is also supported by Rasmussen et al. (2011).{{sfn|McChesney|2015}}}}


Fossils from [[Mungo Lake remains|Lake Mungo]], Australia, have been dated to about 42,000 years ago.<ref>{{cite journal |vauthors=Bowler JM, Johnston H, Olley JM, Prescott JR, Roberts RG, Shawcross W, Spooner NA |title=New ages for human occupation and climatic change at Lake Mungo, Australia |journal=[[Nature (journal)|Nature]] |volume=421 |issue=6925 |pages=837–640 |date=February 2003 |pmid=12594511 |doi=10.1038/nature01383 |bibcode=2003Natur.421..837B |s2cid=4365526}}</ref><ref name="doisj.quascirev.2005.07.022">{{cite journal|vauthors=Olleya JM, Roberts RG, Yoshida H, Bowler JM |title=Single-grain optical dating of grave-infill associated with human burials at Lake Mungo, Australia |journal=[[Quaternary Science Reviews]] |volume=25 |issue=19–20 |year=2006 |pages=2469–2474 |doi=10.1016/j.quascirev.2005.07.022 |bibcode=2006QSRv...25.2469O}}</ref> Other fossils from a site called [[Madjedbebe]] have been dated to at least 65,000 years ago,<ref>{{cite journal |display-authors=2 |last1=Clarkson |first1=Chris |last2=Jacobs |first2=Zenobia |last3=Marwick |first3=Ben |last4=Fullagar |first4=Richard |last5=Wallis |first5=Lynley |last6=Smith |first6=Mike |last7=Roberts |first7=Richard G. |last8=Hayes |first8=Elspeth |last9=Lowe |first9=Kelsey |last10=Carah |first10=Xavier |last11=Florin |first11=S. Anna |last12=McNeil |first12=Jessica |last13=Cox |first13=Delyth |last14=Arnold |first14=Lee J. |last15=Hua |first15=Quan |last16=Huntley |first16=Jillian |last17=Brand |first17=Helen E. A. |last18=Manne |first18=Tiina |last19=Fairbairn |first19=Andrew |last20=Shulmeister |first20=James |last21=Lyle |first21=Lindsey |last22=Salinas |first22=Makiah |last23=Page |first23=Mara |last24=Connell |first24=Kate |last25=Park |first25=Gayoung |last26=Norman |first26=Kasih |last27=Murphy |first27=Tessa |last28=Pardoe |first28=Colin |title=Human occupation of northern Australia by 65,000 years ago |journal=[[Nature (journal)|Nature]] |date=19 July 2017 |volume=547 |issue=7663 |pages=306–310 |doi=10.1038/nature22968 |pmid=28726833 |hdl=2440/107043 |bibcode=2017Natur.547..306C |s2cid=205257212 |url=https://digital.library.adelaide.edu.au/dspace/bitstream/2440/107043/2/hdl_107043.pdf |hdl-access=free}}</ref><ref>{{cite journal |last1=Gibbons |first1=Ann |title=The first Australians arrived early |journal=[[Science (journal)|Science]] |date=20 July 2017 |volume=357 |issue=6348 |pages=238–239 |doi=10.1126/science.357.6348.238 |pmid=28729491 |bibcode=2017Sci...357..238G}}</ref> though some researchers doubt this early estimate and date the Madjedbebe fossils at about 50,000 years ago at the oldest.<ref name="Wood_2017" /><ref name="Homo sapiens first reach Southeast" />
===South-Asia and Australia===


Phylogenetic data suggests that an early Eastern Eurasian (Eastern non-African) meta-population trifurcated somewhere in [[South Asia|eastern South Asia]], and gave rise to the Australo-Papuans, the Ancient Ancestral South Indians (AASI), as well as East/Southeast Asians, although Papuans may have also received some gene flow from an earlier group (xOoA), around 2%,<ref>{{Cite web |title=Almost all living people outside of Africa trace back to a single migration more than 50,000 years ago |url=https://www.science.org/content/article/almost-all-living-people-outside-africa-trace-back-single-migration-more-50000-years |access-date=2022-08-19 |website=www.science.org |language=en}}</ref> next to additional archaic admixture in the [[Sahul]] region.<ref name="Yang">{{cite journal |vauthors=Yang M |title=A genetic history of migration, diversification, and admixture in Asia |journal=Human Population Genetics and Genomics |date=2022-01-06 |pages=1–32 |doi=10.47248/hpgg2202010001 |doi-access=free}}</ref><ref name="auto6">Genetics and material culture support repeated expansions into Paleolithic Eurasia from a population hub out of Africa, Vallini et al. 2022 (April 4, 2022) Quote: "''Taken together with a lower bound of the final settlement of Sahul at 37 kya it is reasonable to describe Papuans as either an almost even mixture between East-Eurasians and a lineage basal to West and East-Eurasians which occurred sometimes between 45 and 38kya, or as a sister lineage of East-Eurasians with or without a minor basal OoA or xOoA contribution. We here chose to parsimoniously describe Papuans as a simple sister group of Tianyuan, cautioning that this may be just one out of six equifinal possibilities.''"</ref>
It is thought that Australia was inhabited around 65,000-50,000 years ago. As of 2017, the earliest evidence of humans in Australia is at least 65,000 years old,<ref name=Clarkson2017/><ref name=StFleur2017 /> while McChesney stated that
{{quote|...genetic evidence suggests that a small band with the marker M168 migrated out of Africa along the coasts of the Arabian Peninsula and India, through Indonesia, and reached Australia very early, between 60,000 and 50,000 years ago. This very early migration into Australia is also supported by Rasmussen et al. (2011).{{sfn|Young McChesney|2015}}}}


According to one study, Papuans could have either formed from a mixture between an East Eurasian lineage and lineage basal to West and East Asians, or as a sister lineage of East Asians with or without a minor basal OoA or xOoA contribution.<ref>Genetics and material culture support repeated expansions into Paleolithic Eurasia from a population hub out of Afri, Vallini et al. 2021 (October 15, 2021)
Fossils from [[Mungo Lake remains|Lake Mungo, Australia]], have been dated to about 42,000 years ago.<ref>{{cite journal |last1=Bowler |first1=James M. |last2=Johnston |first2=Harvey |last3=Olley |first3=John M. |last4=Prescott |first4=John R. |last5=Roberts |first5=Richard G. |last6=Shawcross |first6=Wilfred |last7=Spooner |first7=Nigel A. |year=2003 |title=New ages for human occupation and climatic change at Lake Mungo, Australia |journal=Nature |volume=421 |issue=6925 |pp=837–40 |pmid=12594511 |doi=10.1038/nature01383 |bibcode=2003Natur.421..837B }}</ref><ref name = "doisj.quascirev.2005.07.022">{{cite journal|vauthors=Olleya JM, Roberts RG, Yoshida H, Bowler JM |title =Single-grain optical dating of grave-infill associated with human burials at Lake Mungo, Australia | journal =Quaternary Science Reviews |volume = 25 |issue =19–20 | year = 2006 | pages = 2469–2474 | pmid = | doi = 10.1016/j.quascirev.2005.07.022 | url=| ref = harv |bibcode = 2006QSRv...25.2469O }}</ref> Other fossils from a site called [[Madjedbebe]] have been dated to at least 65,000 years ago.<ref name=StFleur2017/>
Quote: "''Taken together with a lower bound of the final settlement of Sahul at 37 kya (the date of the deepest population splits estimated by 1) it is reasonable to describe Papuans as either an almost even mixture between East''
Asians and a lineage basal to West and East Asians occurred sometimes between 45 and
38kya, or as a sister lineage of East Asians with or without a minor basal OoA or xOoA
contribution. ''"''</ref>


A Holocene hunter-gatherer sample (Leang_Panninge) from [[South Sulawesi]] was found to be genetically in between East-Eurasians and Australo-Papuans. The sample could be modeled as ~50% Papuan-related and ~50% Basal-East Asian-related (Andamanese Onge or Tianyuan). The authors concluded that Basal-East Asian ancestry was far more widespread and the peopling of Insular Southeast Asia and Oceania was more complex than previously anticipated.<ref>Genomic insights into the human population history of Australia and New Guinea, University of Cambridge, Bergström et al. 2018</ref><ref>{{cite journal |vauthors=Carlhoff S, Duli A, Nägele K, Nur M, Skov L, Sumantri I, Oktaviana AA, Hakim B, Burhan B, Syahdar FA, McGahan DP, Bulbeck D, Perston YL, Newman K, Saiful AM, Ririmasse M, Chia S, Pulubuhu DA, Jeong C, Peter BM, Prüfer K, Powell A, Krause J, Posth C, Brumm A |display-authors=6 |title=Genome of a middle Holocene hunter-gatherer from Wallacea |journal=[[Nature (journal)|Nature]] |volume=596 |issue=7873 |pages=543–547 |date=August 2021 |pmid=34433944 |pmc=8387238 |doi=10.1038/s41586-021-03823-6 |bibcode=2021Natur.596..543C |quote=The [[qpGraph]] analysis confirmed this branching pattern, with the Leang Panninge individual branching off from the Near Oceanian clade after the Denisovan gene flow, although with the most supported topology indicating around 50% of a basal East Asian component contributing to the Leang Panninge genome (Fig. 3c, Supplementary Figs. 7–11).}}</ref>[[File:PCA calculated on present-day individuals from eastern Eurasia and Near Oceania.png|thumb|PCA calculated on present-day and ancient individuals from eastern Eurasia and Oceania. PC1 (23,8%) distinguishes East-Eurasians and Australo-Melanesians, while PC2 (6,3%) differentiates East-Eurasians along a North to South cline.]]
===East Asia===
[[File:Principal component analysis of ancient and present-day individuals from worldwide populations.png|thumb|Principal component analysis (PCA) of ancient and modern day individuals from worldwide populations. Oceanians (Aboriginal Australians and Papuans) are most differentiated from both East-Eurasians and West-Eurasians.]]


=== East and Southeast Asia ===
[[Tianyuan man]] from [[China]] has a probable date range between 38,000 and 42,000 years ago, while [[Liujiang man]] from the same region has a probable date range between 67,000 and 159,000 years ago. According to 2013 DNA tests, Tianyuan man is related "to many present-day [[Asian people|Asians]] and [[Indigenous peoples of the Americas|Native Americans]]".<ref>{{cite web|url=http://www.mpg.de/6842535/dna-Tianyuan-cave|title=A relative from the Tianyuan Cave|publisher=[[Max Planck Society]]|date=2013-01-21}}</ref><ref name=daily>{{cite web|url=https://www.sciencedaily.com/releases/2013/01/130121161802.htm|title=A relative from the Tianyuan Cave: Humans living 40,000 years ago likely related to many present-day Asians and Native Americans|work=[[Science Daily]]|date=2013-01-21}}</ref><ref>{{cite web|url=http://www.sci-news.com/othersciences/anthropology/article00842.html|title=DNA Analysis Reveals Common Origin of Tianyuan Humans and Native Americans, Asians|work=Sci-News|date=2013-01-24}}</ref><ref>{{cite web|url=https://www.sciencenews.org/article/ancient-human-dna-suggests-minimal-interbreeding|title=Ancient human DNA suggests minimal interbreeding|publisher=''[[Science News]]''|date=2013-01-21}}</ref><ref>{{cite web|url=http://cavingnews.com/20130131-ancient-bone-dna-shows-ancestry-of-modern-asians-native-americans|title=Ancient Bone DNA Shows Ancestry of Modern Asians & Native Americans|publisher=Caving News|date=2013-01-31}}</ref> Tianyuan is similar in [[morphology (biology)|morphology]] to [[Minatogawa Man]], modern humans dated between 17,000 and 19,000 years ago and found on [[Okinawa]] Island, [[Japan]].<ref>{{cite journal |last1=Hu |first1=Yaowu |last2=Shang |first2=Hong |last3=Tong |first3=Haowen |last4=Nehlich |first4=Olaf |last5=Liu |first5=Wu |last6=Zhao |first6=Chaohong |last7=Yu |first7=Jincheng |last8=Wang |first8=Changsui |last9=Trinkaus |first9=Erik |last10=Richards |first10=Michael P. |date=July 2009 |title=Stable isotope dietary analysis of the Tianyuan 1 early modern human |journal=Proc. Natl. Acad. Sci. U.S.A. |volume=106 |issue=27 |pp=10971–4 |pmid=19581579 |pmc=2706269 |doi=10.1073/pnas.0904826106 |bibcode=2009PNAS..10610971H }}</ref><ref>{{cite journal |last=Brown |first=Peter |date=August 1992 |title=Recent human evolution in East Asia and Australasia |journal=Philos. Trans. R. Soc. Lond. B Biol. Sci. |volume=337 |issue=1280 |pp=235–42 |pmid=1357698 |doi=10.1098/rstb.1992.0101 }}</ref>
In China, the [[Liujiang man]] ({{zh|c=柳江人}}) is among the earliest modern humans found in [[East Asia]].<ref name="liujiang hominid site3">{{cite journal |vauthors=Shen G, Wang W, Wang Q, Zhao J, Collerson K, Zhou C, Tobias PV |title=U-Series dating of Liujiang hominid site in Guangxi, Southern China |journal=[[Journal of Human Evolution]] |volume=43 |issue=6 |pages=817–829 |date=December 2002 |pmid=12473485 |doi=10.1006/jhev.2002.0601|bibcode=2002JHumE..43..817S }}</ref> The date most commonly attributed to the remains is 67,000 years ago.<ref name="Rosenburg 2002">{{Cite journal |vauthors=Rosenburg K |date=2002 |title=A Late Pleistocene Human Skeleton from Liujiang, China Suggests Regional Population Variation in Sexual Dimorphism in the Human Pelvis |journal=[[Variability and Evolution]]}}</ref> High rates of variability yielded by various dating techniques carried out by different researchers place the most widely accepted range of dates with 67,000 BP as a minimum, but do not rule out dates as old as 159,000 BP.<ref name="Rosenburg 2002" /> {{harvp|Liu, Martinón-Torres et al.|2015}} claim that modern human teeth have been found in China dating to at least 80,000 years ago.<ref name="Kaifu_20123">{{cite journal | last1=Kaifu | first1=Yousuke | last2=Fujita | first2=Masaki | title=Fossil record of early modern humans in East Asia | journal=Quaternary International | volume=248 | date=2012 | doi=10.1016/j.quaint.2011.02.017 | pages=2–11|bibcode=2012QuInt.248....2K}}</ref>


[[Tianyuan man]] from China has a probable date range between 38,000 and 42,000 years ago, while [[Liujiang man]] from the same region has a probable date range between 67,000 and 159,000 years ago. According to 2013 DNA tests, Tianyuan man is related "to many present-day [[Asian people|Asians]] and [[Indigenous peoples of the Americas|Native Americans]]".<ref>{{cite web |url=http://www.mpg.de/6842535/dna-Tianyuan-cave |title=A relative from the Tianyuan Cave |publisher=[[Max Planck Society]] |date=2013-01-21}}</ref><ref name="daily">{{cite web |url=https://www.sciencedaily.com/releases/2013/01/130121161802.htm |title=A relative from the Tianyuan Cave: Humans living 40,000 years ago likely related to many present-day Asians and Native Americans |work=[[Science Daily]] |date=2013-01-21}}</ref><ref>{{cite web|url=http://www.sci-news.com/othersciences/anthropology/article00842.html |title=DNA Analysis Reveals Common Origin of Tianyuan Humans and Native Americans, Asians |work=Sci-News |date=2013-01-24}}</ref><ref>{{cite magazine |url=https://www.sciencenews.org/article/ancient-human-dna-suggests-minimal-interbreeding |title=Ancient human DNA suggests minimal interbreeding |magazine=[[Science News]] |date=2013-01-21}}</ref><ref>{{cite web |url=http://cavingnews.com/20130131-ancient-bone-dna-shows-ancestry-of-modern-asians-native-americans |title=Ancient Bone DNA Shows Ancestry of Modern Asians & Native Americans |publisher=Caving News |date=2013-01-31}}</ref> Tianyuan is similar in [[morphology (biology)|morphology]] to Liujiang man, and some [[Jōmon period]] modern humans found in Japan, as well as modern East and Southeast Asians.<ref>{{cite journal |vauthors=Hu Y, Shang H, Tong H, Nehlich O, Liu W, Zhao C, Yu J, Wang C, Trinkaus E, Richards MP |display-authors=6 |title=Stable isotope dietary analysis of the Tianyuan 1 early modern human |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=106 |issue=27 |pages=10971–10974 |date=July 2009 |pmid=19581579 |pmc=2706269 |doi=10.1073/pnas.0904826106 |doi-access=free |bibcode=2009PNAS..10610971H}}</ref><ref>{{cite journal |vauthors=Brown P |title=Recent human evolution in East Asia and Australasia |journal=Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences |volume=337 |issue=1280 |pages=235–242 |date=August 1992 |pmid=1357698 |doi=10.1098/rstb.1992.0101 |bibcode=1992RSPTB.337..235B}}</ref><ref>{{cite journal |vauthors=Curnoe D, Xueping J, Herries AI, Kanning B, Taçon PS, Zhende B, Fink D, Yunsheng Z, Hellstrom J, Yun L, Cassis G, Bing S, Wroe S, Shi H, Parr WC, Shengmin H, Rogers N |display-authors=6 |title=Human remains from the Pleistocene-Holocene transition of southwest China suggest a complex evolutionary history for East Asians |journal=[[PLOS ONE]] |volume=7 |issue=3 |pages=e31918 |date=2012-03-14 |pmid=22431968 |pmc=3303470 |doi=10.1371/journal.pone.0031918 |doi-access=free |bibcode=2012PLoSO...731918C}}</ref>
===Europe===


A 2021 study about the population history of Eastern Eurasia, concluded that distinctive [[East Asian people|Basal-East Asian]] (East-Eurasian) ancestry originated in [[Mainland Southeast Asia]] at ~50,000BC from a distinct southern Himalayan route, and expanded through multiple migration waves southwards and northwards respectively.<ref name="Larena">{{cite journal |vauthors=Larena M, Sanchez-Quinto F, Sjödin P, McKenna J, Ebeo C, Reyes R, Casel O, Huang JY, Hagada KP, Guilay D, Reyes J, Allian FP, Mori V, Azarcon LS, Manera A, Terando C, Jamero L, Sireg G, Manginsay-Tremedal R, Labos MS, Vilar RD, Latiph A, Saway RL, Marte E, Magbanua P, Morales A, Java I, Reveche R, Barrios B, Burton E, Salon JC, Kels MJ, Albano A, Cruz-Angeles RB, Molanida E, Granehäll L, Vicente M, Edlund H, Loo JH, Trejaut J, Ho SY, Reid L, Malmström H, Schlebusch C, Lambeck K, Endicott P, Jakobsson M |display-authors=6 |title=Multiple migrations to the Philippines during the last 50,000 years |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=118 |issue=13 |pages=e2026132118 |date=March 2021 |pmid=33753512 |pmc=8020671 |doi=10.1073/pnas.2026132118 |bibcode=2021PNAS..11826132L |doi-access=free}}</ref>
According to {{harvp|Macaulay et al.|2005}}, an early offshoot from the southern dispersal with haplogroup N followed the Nile from East Africa, heading northwards and crossing into [[Asia]] through the [[Sinai]]. This group then branched, some moving into Europe and others heading east into Asia.{{r|Macaulay2005}} This hypothesis is supported by the relatively late date of the arrival of modern humans in Europe as well as by archaeological and DNA evidence.{{r|Macaulay2005}} Based on an analysis of 55 human mitochondrial genomes (mtDNAs) of hunter-gatherers, {{harvp|Posth et al.|2016}} argue for a "rapid single dispersal of all non-Africans less than 55,000 years ago."


Genetic studies concluded that [[Indigenous peoples of the Americas|Native Americans]] descended from a single founding population that initially split from a Basal-East Asian source population in Mainland Southeast Asia around 36,000 years ago, at the same time at which the proper [[Jōmon people]] split from Basal-East Asians, either together with Ancestral Native Americans or during a separate expansion wave. They also show that the basal northern and southern Native American branches, to which all other Indigenous peoples belong, diverged around 16,000 years ago.<ref>{{cite journal |vauthors=Moreno-Mayar JV, Potter BA, Vinner L, Steinrücken M, Rasmussen S, Terhorst J, Kamm JA, Albrechtsen A, Malaspinas AS, Sikora M, Reuther JD, Irish JD, Malhi RS, Orlando L, Song YS, Nielsen R, Meltzer DJ, Willerslev E |display-authors=6 |title=Terminal Pleistocene Alaskan genome reveals first founding population of Native Americans |journal=[[Nature (journal)|Nature]] |volume=553 |issue=7687 |pages=203–207 |date=January 2018 |pmid=29323294 |doi=10.1038/nature25173 |s2cid=4454580 |bibcode=2018Natur.553..203M |url=https://researchonline.ljmu.ac.uk/id/eprint/7887/1/UpwardSun_Nature%20paper%20MS%20DEC17.pdf}}</ref><ref>{{cite journal |vauthors=Gakuhari T, Nakagome S, Rasmussen S, Allentoft ME, Sato T, Korneliussen T, Chuinneagáin BN, Matsumae H, Koganebuchi K, Schmidt R, Mizushima S, Kondo O, Shigehara N, Yoneda M, Kimura R, Ishida H, Masuyama T, Yamada Y, Tajima A, Shibata H, Toyoda A, Tsurumoto T, Wakebe T, Shitara H, Hanihara T, Willerslev E, Sikora M, Oota H |display-authors=6 |title=Ancient Jomon genome sequence analysis sheds light on migration patterns of early East Asian populations |journal=[[Communications Biology]] |volume=3 |issue=1 |pages=437 |date=August 2020 |pmid=32843717 |pmc=7447786 |doi=10.1038/s42003-020-01162-2}}</ref> An indigenous American sample from 16,000BC in [[Idaho]], which is craniometrically similar to modern Native Americans as well as [[Paleosiberian peoples|Paleosiberias]], was found to have largely East-Eurasian ancestry and showed high affinity with contemporary East Asians, as well as Jōmon period samples of Japan, confirming that Ancestral Native Americans split from an East-Eurasian source population in Eastern Siberia.<ref>{{cite journal |vauthors=Davis LG, Madsen DB, Becerra-Valdivia L, Higham T, Sisson DA, Skinner SM, Stueber D, Nyers AJ, Keen-Zebert A, Neudorf C, Cheyney M, Izuho M, Iizuka F, Burns SR, Epps CW, Willis SC, Buvit I |display-authors=6 |title=Late Upper Paleolithic occupation at Cooper's Ferry, Idaho ~16,000 years ago |language=EN |journal=[[Science (journal)|Science]] |volume=365 |issue=6456 |pages=891–897 |date=August 2019 |pmid=31467216 |doi=10.1126/science.aax9830 |quote=We interpret this temporal and technological affinity to signal a cultural connection with Upper Paleolithic northeastern Asia, which complements current evidence of shared genetic heritage between late Pleistocene peoples of northern Japan and North America. |s2cid=201672463 |doi-access=free |bibcode=2019Sci...365..891D}}</ref>
==Genetic reconstruction==


=== Mitochondrial haplogroups ===
=== Europe ===
According to {{harvp|Macaulay et al.|2005}}, an early offshoot from the southern dispersal with haplogroup N followed the Nile from East Africa, heading northwards and crossing into Asia through the [[Sinai Peninsula|Sinai]]. This group then branched, some moving into Europe and others heading east into Asia.{{r|Macaulay2005}} This hypothesis is supported by the relatively late date of the arrival of modern humans in Europe as well as by archaeological and DNA evidence.{{r|Macaulay2005}} Based on an analysis of 55 human mitochondrial genomes (mtDNAs) of hunter-gatherers, {{harvp|Posth et al.|2016}} argue for a "rapid single dispersal of all non-Africans less than 55,000 years ago". By 45,000 years ago, modern humans are known to have reached northwestern Europe.<ref>{{Cite journal |last1=Mylopotamitaki |first1=Dorothea |last2=Weiss |first2=Marcel |last3=Fewlass |first3=Helen |last4=Zavala |first4=Elena Irene |last5=Rougier |first5=Hélène |last6=Sümer |first6=Arev Pelin |last7=Hajdinjak |first7=Mateja |last8=Smith |first8=Geoff M. |last9=Ruebens |first9=Karen |last10=Sinet-Mathiot |first10=Virginie |last11=Pederzani |first11=Sarah |last12=Essel |first12=Elena |last13=Harking |first13=Florian S. |last14=Xia |first14=Huan |last15=Hansen |first15=Jakob |date=31 January 2024 |title=Homo sapiens reached the higher latitudes of Europe by 45,000 years ago |journal=[[Nature (journal)|Nature]] |language=en |volume=626 |issue=7998 |pages=341–346 |doi=10.1038/s41586-023-06923-7 |issn=1476-4687 |last16=Kirchner |first16=André |last17=Lauer |first17=Tobias  |last18=Stahlschmidt |first18=Mareike |last19=Hein |first19=Michael |last20=Talamo |first20=Sahra |last21=Wacker |first21=Lukas |last22=Meller |first22=Harald |last23=Dietl |first23=Holger |last24=Orschiedt |first24=Jörg |last25=Olsen |first25=Jesper V. |last26=Zeberg |first26=Hugo |last27=Prüfer |first27=Kay |last28=Krause |first28=Johannes |last29=Meyer |first29=Matthias |last30=Welker |first30=Frido |last31=McPherron |first31=Shannon P. |last32=Schüler |first32=Tim |last33=HublinHublin |first33=Jean-Jacques|pmid=38297117 |pmc=10849966 |bibcode=2024Natur.626..341M }}</ref>


== Genetic reconstruction ==
====Within Africa====
=== Mitochondrial haplogroups ===
==== Within Africa ====
{{Further|Most recent common ancestor|Archaeogenetics|Human mitochondrial DNA haplogroup}}
{{Further|Most recent common ancestor|Archaeogenetics|Human mitochondrial DNA haplogroup}}


[[File:African Mitochondrial descent.PNG|thumb|Map of early diversification of modern humans according to [[Mitochondrial DNA|mitochondrial]] [[population genetics]] ''(see: [[Macro-haplogroup L (mtDNA)|Haplogroup L]])''.]]
[[File:African Mitochondrial descent.PNG|thumb|Map of early diversification of modern humans according to [[Mitochondrial DNA|mitochondrial]] [[population genetics]] ''(see: [[Macro-haplogroup L (mtDNA)|Haplogroup L]])''.]]
The first lineage to branch off from [[Mitochondrial Eve]] is [[Haplogroup L0 (mtDNA)|L0]]. This haplogroup is found in high proportions among the [[san people|San]] of Southern Africa and the [[Sandawe people|Sandawe]] of East Africa. It is also found among the [[Mbuti]] people.{{r|pmid17194802|pmid10739760}} These groups branched off early in human history and have remained relatively genetically isolated since then. [[Haplogroup L1 (mtDNA)|Haplogroups L1]], [[Haplogroup L2 (mtDNA)|L2]] and [[Haplogroup L3 (mtDNA)|L3]] are descendants of L1-6 and are largely confined to Africa. The macro haplogroups [[Haplogroup M (mtDNA)|M]] and [[Haplogroup N (mtDNA)|N]], which are the lineages of the rest of the world outside Africa, descend from L3. L3 is about 84,000 years old and haplogroup M and N are about 63,000 years old.{{r|Macaulay2005}} The relationship between such gene trees and demographic history is still debated when applied to dispersals.{{sfnp|Groucutt et al.|2015}}
The first lineage to branch off from [[Mitochondrial Eve]] was [[Haplogroup L0|L0]]. This haplogroup is found in high proportions among the [[san people|San]] of Southern Africa and the [[Sandawe people|Sandawe]] of East Africa. It is also found among the [[Mbuti]] people.{{r|pmid17194802|pmid10739760}} These groups branched off early in human history and have remained relatively genetically isolated since then. [[Haplogroup L1 (mtDNA)|Haplogroups L1]], [[Haplogroup L2 (mtDNA)|L2]], and [[Haplogroup L3 (mtDNA)|L3]] are descendants of L1–L6, and are largely confined to Africa. The macro haplogroups [[Haplogroup M (mtDNA)|M]] and [[Haplogroup N (mtDNA)|N]], which are the lineages of the rest of the world outside Africa, descend from L3. L3 is about 70,000 years old, while haplogroups M and N are about 65–55,000 years old.<ref name="mbe.oxfordjournals.org">{{cite journal |vauthors=Soares P, Alshamali F, Pereira JB, Fernandes V, Silva NM, Afonso C, Costa MD, Musilová E, Macaulay V, Richards MB, Cerny V, Pereira L |display-authors=6 |title=The Expansion of mtDNA Haplogroup L3 within and out of Africa |journal=[[Molecular Biology and Evolution]] |volume=29 |issue=3 |pages=915–927 |date=March 2012 |pmid=22096215 |doi=10.1093/molbev/msr245 |doi-access=free}}</ref><ref name="Vai" /> The relationship between such gene trees and demographic history is still debated when applied to dispersals.{{sfnp|Groucutt et al.|2015}}


Of all the lineages present in Africa, only the female descendants of one lineage, [[haplogroup L3 (mtDNA)|mtDNA haplogroup L3]], are found outside Africa. If there had been several migrations, one would expect descendants of more than one lineage to be found. L3's female descendants, the [[Haplogroup M (mtDNA)|M]] and [[Haplogroup N (mtDNA)|N]] haplogroup lineages, are found in very low frequencies in Africa (although [[Haplogroup M (mtDNA)#Haplogroup M1|haplogroup M1]] populations are very ancient and diversified in [[North Africa|North]] and [[Horn of Africa|North-east Africa]]) and appear to be more recent arrivals. A possible explanation is that these mutations occurred in East Africa shortly before the exodus and became the dominant haplogroups after the departure through the [[founder effect]]. Alternatively, the mutations may have arisen shortly afterwards.
Of all the lineages present in Africa, the female descendants of only one lineage, [[haplogroup L3 (mtDNA)|mtDNA haplogroup L3]], are found outside Africa. If there had been several migrations, one would expect descendants of more than one lineage to be found. L3's female descendants, the [[Haplogroup M (mtDNA)|M]] and [[Haplogroup N (mtDNA)|N]] haplogroup lineages, are found in very low frequencies in Africa (although [[Haplogroup M (mtDNA)#Haplogroup M1|haplogroup M1]] populations are very ancient and diversified in [[North Africa|North]] and [[Horn of Africa|North-east Africa]]) and appear to be more recent arrivals.{{citation needed|date=September 2020}} A possible explanation is that these mutations occurred in East Africa shortly before the exodus and became the dominant haplogroups thereafter by means of the [[founder effect]]. Alternatively, the mutations may have arisen shortly afterwards.


====Southern Route and haplogroups M and N====
==== Southern Route and haplogroups M and N ====
Results from mtDNA collected from aboriginal Malaysians called [[Orang Asli]] indicate that the haplogroups M and N share characteristics with original African groups from approximately 85,000 years ago, and share characteristics with sub-haplogroups found in coastal south-east Asian regions, such as Australasia, the Indian subcontinent and throughout continental Asia, which had dispersed and separated from their African progenitor approximately 65,000 years ago. This southern coastal dispersal would have occurred before the dispersal through the Levant approximately 45,000 years ago.{{r|Macaulay2005}} This hypothesis attempts to explain why haplogroup N is predominant in Europe and why haplogroup M is absent in Europe. Evidence of the coastal migration is thought to have been destroyed by the rise in sea levels during the [[Holocene]] epoch.<ref>{{cite journal |vauthors=Maca-Meyer N, González AM, Larruga JM, Flores C, Cabrera VM |title=Major genomic mitochondrial lineages delineate early human expansions |journal=[[BMC Genetics]] |volume=2 |pages=13 |date=13 August 2001 |pmid=11553319 |pmc=55343 |doi=10.1186/1471-2156-2-13 |doi-access=free }}</ref> Alternatively, a small European founder population that had expressed haplogroup M and N at first, could have lost haplogroup M through random [[genetic drift]] resulting from a [[Population bottleneck|bottleneck]] (i.e. a [[founder effect]]).


The group that crossed the Red Sea travelled along the coastal route around [[Arabia]] and [[Persian Plateau|Persia]] until reaching India.{{r|pmid15339343}} [[Haplogroup M (mtDNA)|Haplogroup M]] is found in high frequencies along the southern coastal regions of Pakistan and India and it has the greatest diversity in India, indicating that it is here where the mutation may have occurred.{{r|pmid15339343}} Sixty percent of the Indian population belong to [[Haplogroup M (mtDNA)|Haplogroup M]]. The indigenous people of the [[Andaman Islands]] also belong to the M lineage. The Andamanese are thought to be offshoots of some of the earliest inhabitants in Asia because of their long isolation from the mainland. They are evidence of the coastal route of early settlers that extends from India to [[Thailand]] and Indonesia all the way to eastern [[New Guinea]]. Since M is found in high frequencies in highlanders from New Guinea and the Andamanese and New Guineans have dark skin and [[Hair|Afro-textured hair]], some scientists think they are all part of the same wave of migrants who departed across the Red Sea ~60,000 years ago in the [[Great Coastal Migration]]. The proportion of haplogroup M increases eastwards from [[Arabia]] to India; in eastern India, M outnumbers N by a ratio of 3:1. Crossing into Southeast Asia, haplogroup N (mostly in the form of derivatives of its R subclade) reappears as the predominant lineage.{{citation needed|date=October 2017}} M is predominant in East Asia, but amongst [[Indigenous Australians]], N is the more common lineage.{{citation needed|date=October 2017}} This haphazard distribution of Haplogroup N from Europe to Australia can be explained by [[founder effect]]s and [[population bottleneck]]s.<ref name="pmid12840039">{{cite journal |vauthors=Ingman M, Gyllensten U |title=Mitochondrial genome variation and evolutionary history of Australian and New Guinean aborigines |journal=[[Genome Research]] |volume=13 |issue=7 |pages=1600–1606 |date=July 2003 |pmid=12840039 |pmc=403733 |doi=10.1101/gr.686603}}</ref>
Results from mtDNA collected from aboriginal Malaysians called [[Orang Asli]] and the creation of a phylogentic tree indicate that the hapologroups M and N share characteristics with original African groups from approximately 85,000 years ago and share characteristics with sub-haplogroups among coastal south-east Asian regions, such as Australasia, the Indian subcontinent and throughout continental Asia, which had dispersed and separated from its African origins approximately 65,000 years ago. This southern coastal dispersion would have occurred before the dispersion through the Levant approximately 45,000 years ago.{{r|Macaulay2005}} This hypothesis attempts to explain why haplogroup N is predominant in Europe and why haplogroup M is absent in Europe. Evidence of the coastal migration is thought to have been destroyed by the rise in sea levels during the [[Holocene]] epoch.<ref>{{cite journal |last1=Maca-Meyer |first1=Nicole |last2=González |first2=Ana M. |last3=Larruga |first3=José M. |last4=Flores |first4=Carlos |last5=Cabrera |first5=Vicente M. |date=13 August 2001 |title=Major genomic mitochondrial lineages delineate early human expansions |journal=BMC Genet. |volume=2 |p=13 |url=http://bmcgenet.biomedcentral.com/articles/10.1186/1471-2156-2-13 |doi=10.1186/1471-2156-2-13 |pmid=11553319 |pmc=55343}}</ref> Alternatively, a small European founder population that had expressed haplogroup M and N at first, could have lost haplogroup M through random [[genetic drift]] resulting from a [[Population bottleneck|bottleneck]] (i.e. a [[founder effect]]).
[[File:Simplifed_Y_tree_is_shown_as_reference_for_colours.png|thumb|300x300px|The earliest-branching non-African paternal lineages (C, D, F) after the Out-of-Africa event (a), and their deepest divergence among modern day East or Southeast Asia (b), suggesting rapid coastal expansions. Simplified Y-chromosome tree is shown as reference for colours.<ref>{{Cite journal |vauthors=Hallast P, Agdzhoyan A, Balanovsky O, Xue Y, Tyler-Smith C |date=February 2021 |title=A Southeast Asian origin for present-day non-African human Y chromosomes |journal=Human Genetics |volume=140 |issue=2 |pages=299–307 |doi=10.1007/s00439-020-02204-9 |pmc=7864842 |pmid=32666166}}</ref>]]


=== Autosomal DNA ===
The group that crossed the Red Sea travelled along the coastal route around [[Arabia]] and [[Persia]] until reaching India.{{r|pmid15339343}} [[Haplogroup M (mtDNA)|Haplogroup M]] is found in high frequencies along the southern coastal regions of [[Pakistan]] and India and it has the greatest diversity in India, indicating that it is here where the mutation may have occurred.{{r|pmid15339343}} Sixty percent of the Indian population belong to [[Haplogroup M (mtDNA)|Haplogroup M]]. The indigenous people of the [[Andaman Islands]] also belong to the M lineage. The Andamanese are thought to be offshoots of some of the earliest inhabitants in Asia because of their long isolation from the mainland. They are evidence of the coastal route of early settlers that extends from India to [[Thailand]] and Indonesia all the way to [[Papua New Guinea]]. Since M is found in high frequencies in highlanders from New Guinea and the Andamanese and New Guineans have dark skin and [[Hair|Afro-textured hair]], some scientists think they are all part of the same wave of migrants who departed across the Red Sea ~60,000 years ago in the [[Great Coastal Migration]]. The proportion of haplogroup M increases eastwards from [[Arabia]] to India; in eastern India, M outnumbers N by a ratio of 3:1. Crossing into Southeast Asia, haplogroup N (mostly in the form of derivatives of its R subclade) reappears as the predominant lineage.{{citation needed|date=October 2017}} M is predominant in East Asia, but amongst [[Indigenous Australians]], N is the more common lineage.{{citation needed|date=October 2017}} This haphazard distribution of Haplogroup N from Europe to Australia can be explained by [[founder effect]]s and [[population bottleneck]]s.<ref name="pmid12840039">{{cite journal |vauthors=Ingman M, Gyllensten U | title = Mitochondrial genome variation and evolutionary history of Australian and New Guinean aborigines | journal = Genome Res. | volume = 13 | issue = 7 | pages = 1600–6 |date=July 2003 | pmid = 12840039 | pmc = 403733 | doi = 10.1101/gr.686603 | ref = harv }}</ref>
{{see|Human genetic variation}}


A 2002 study of African, European, and Asian populations, found greater genetic diversity among Africans than among Eurasians, and that genetic diversity among Eurasians is largely a subset of that among Africans, supporting the out of Africa model.<ref>{{cite journal |vauthors=Yu N, Chen FC, Ota S, Jorde LB, Pamilo P, Patthy L, Ramsay M, Jenkins T, Shyue SK, Li WH |display-authors=6 |title=Larger genetic differences within africans than between Africans and Eurasians |journal=[[Genetics (journal)|Genetics]] |volume=161 |issue=1 |pages=269–274 |date=May 2002 |doi=10.1093/genetics/161.1.269 |pmid=12019240 |pmc=1462113 |url=http://www.genetics.org/content/161/1/269.full |access-date=7 April 2013}}</ref> A large study by Coop ''et al''. (2009) found evidence for [[natural selection]] in [[autosome|autosomal]] DNA outside of Africa. The study distinguishes non-African sweeps (notably [[Stem cell factor|KITLG]] variants associated with [[skin color]]), West-Eurasian sweeps ([[SLC24A5]]) and East-Asian sweeps ([[Melanocortin 1 receptor|MC1R]], relevant to skin color). Based on this evidence, the study concluded that human populations encountered novel selective pressures as they expanded out of Africa.<ref name="coop2009">{{cite journal |vauthors=Coop G, Pickrell JK, Novembre J, Kudaravalli S, Li J, Absher D, Myers RM, Cavalli-Sforza LL, Feldman MW, Pritchard JK |display-authors=6 |title=The role of geography in human adaptation |journal=[[PLOS Genetics]] |volume=5 |issue=6 |pages=e1000500 |date=June 2009 |pmid=19503611 |pmc=2685456 |doi=10.1371/journal.pgen.1000500 |veditors=Schierup MH |doi-access=free }};
===Autosomal DNA===
summary in {{cite book |vauthors=Racimo F, Schraiber JG, Casey F, Huerta-Sanchez E |chapter=Directional Selection and Adaptation |veditors=Kliman RM |title=Encyclopedia of Evolutionary Biology |date=2016 |chapter-url=https://books.google.com/books?id=_r4OCAAAQBAJ&pg=PA451 |page=451 |doi=10.1016/B978-0-12-800049-6.00028-7 |isbn=978-0128004265 |publisher=[[Academic Press]] |via=[[Google Books]]}}</ref> [[MC1R]] and its relation to skin color had already been discussed by {{harvp|Harding et al.|2000|p=1355}}. According to this study, Papua New Guineans continued to be exposed to selection for dark skin color so that, although these groups are distinct from Africans in other places, the allele for dark skin color shared by contemporary Africans, Andamanese and New Guineans is an archaism. {{harvp|Endicott et al.|2003}} suggest [[convergent evolution]]. A 2014 study by Gurdasani et al. indicates that the higher genetic diversity in Africa was further increased in some regions by relatively recent Eurasian migrations affecting parts of Africa.<ref>{{cite journal |vauthors=Gurdasani D, Carstensen T, Tekola-Ayele F, Pagani L, Tachmazidou I, Hatzikotoulas K, Karthikeyan S, Iles L, Pollard MO, Choudhury A, Ritchie GR, Xue Y, Asimit J, Nsubuga RN, Young EH, Pomilla C, Kivinen K, Rockett K, Kamali A, Doumatey AP, Asiki G, Seeley J, Sisay-Joof F, Jallow M, Tollman S, Mekonnen E, Ekong R, Oljira T, Bradman N, Bojang K, Ramsay M, Adeyemo A, Bekele E, Motala A, Norris SA, Pirie F, Kaleebu P, Kwiatkowski D, Tyler-Smith C, Rotimi C, Zeggini E, Sandhu MS |display-authors=6 |title=The African Genome Variation Project shapes medical genetics in Africa |journal=[[Nature (journal)|Nature]] |volume=517 |issue=7,534 |pages=327–332 |date=January 2015 |pmid=25470054 |pmc=4297536 |doi=10.1038/nature13997 |bibcode=2015Natur.517..327G}}</ref>


=== Pathogen DNA ===
{{see|Human genetic variation|Human genetic clustering}}
Another promising route towards reconstructing human genetic genealogy is via the [[JC virus]] (JCV), a type of human [[polyomavirus]] which is carried by 70–90 percent of humans and which is usually transmitted vertically, from parents to offspring, suggesting codivergence with human populations. For this reason, JCV has been used as a genetic marker for human evolution and migration.<ref>{{cite book |vauthors=Matisoo-Smith E, Horsburgh KA |title=DNA for Archaeologists |publisher=[[Routledge]] |date=2016}}</ref> This method does not appear to be reliable for the migration out of Africa; in contrast to human genetics, JCV strains associated with African populations are not basal. From this {{harvp|Shackelton et al.|2006}} conclude that either a basal African strain of JCV has become extinct or that the original infection with JCV post-dates the migration from Africa.


=== Admixture of archaic and modern humans ===
A 2002 study of African, European and Asian populations, found greater genetic diversity among Africans than among Eurasians, and that genetic diversity among Eurasians is largely a subset of that among Africans, supporting the out of Africa model.<ref>{{cite journal|url=http://www.genetics.org/content/161/1/269.full|title=Larger Genetic Differences Within Africans Than Between Africans and Eurasians|first=Ning|last=Yu|journal=Genetics|publisher=Genetics Society of America|date=May 2002|accessdate=7 April 2013|displayauthors=etal }}</ref> A large study by Coop ''et al''. (2009) found evidence for [[natural selection]] in [[autosome|autosomal]] DNA outside of Africa. The study distinguishes non-African sweeps (notably [[Stem cell factor|KITLG]] variants associated with [[skin color]]), West-Eurasian sweeps ([[SLC24A5]]) and East-Asian sweeps ([[Melanocortin 1 receptor|MC1R]], relevant to skin color). Based on this evidence, the study concluded that human populations encountered novel selective pressures as they expanded out of Africa.<ref name="coop2009">{{cite journal | title = The role of geography in human adaptation | journal = PLoS Genet. | volume = 5 | issue = 6 | pages = e1000500 |date=June 2009 | pmid = 19503611 | pmc = 2685456 | doi = 10.1371/journal.pgen.1000500 | ref = harv | editor1-last = Schierup | editor1-first = Mikkel H. | last2 = Pickrell | last3 = Novembre | last4 = Kudaravalli | last5 = Li | last6 = Absher | last7 = Myers | last8 = Cavalli-Sforza | last9 = Feldman | last10 = Pritchard |name-list-format=vanc|author1 = Coop G}};
{{main|Interbreeding between archaic and modern humans}}
summary in Kliman (ed.), ''Encyclopedia of Evolutionary Biology'' (2016), [https://books.google.com/books?id=_r4OCAAAQBAJ&pg=PA451#v=onepage&q&f=false p. 451]</ref> [[MC1R]] and its relation to skin color had already been discussed by {{harvp|Liu, Harding et al.|2000|p=135}}. According to this study, Papua New Guineans continued to be exposed to selection for dark skin color so that, although these groups are distinct from Africans in other places, the allele for dark skin color shared by contemporary Africans, Andamanese and New Guineans is an archaism. {{harvp|Endicott et al.|2003}} suggest [[convergent evolution]]. A 2014 study by Gurdasani et al. indicate that higher genetic diversity in Africa was caused by relatively recent Eurasian migrations ''into'' Africa.<ref>{{cite journal|url=http://www.nature.com/nature/journal/v517/n7534/full/nature13997.html|title=The African Genome Variation Project shapes medical genetics in Africa|first=Deepti|last=Gurdasani|journal=Nature|date=July 2015|accessdate=14 June 2017|displayauthors=etal|volume=517|doi=10.1038/nature13997|pages=327–332|pmc=4297536}}</ref>
Evidence for [[archaic human]] species (descended from ''[[Homo heidelbergensis]]'') having [[Archaic human admixture with modern humans|interbred]] with modern humans outside of Africa, was discovered in the 2010s. This concerns primarily [[Neanderthals|Neanderthal]] admixture in all modern populations except for [[Sub-Saharan Africans]] but evidence has also been presented for [[Denisova hominin]] admixture in [[Australasia]] (i.e. in [[Melanesians]], [[Aboriginal Australians]] and some [[Negritos]]).<ref>{{cite journal |vauthors=Rasmussen M, Guo X, Wang Y, Lohmueller KE, Rasmussen S, Albrechtsen A, Skotte L, Lindgreen S, Metspalu M, Jombart T, Kivisild T, Zhai W, Eriksson A, Manica A, Orlando L, De La Vega FM, Tridico S, Metspalu E, Nielsen K, Ávila-Arcos MC, Moreno-Mayar JV, Muller C, Dortch J, Gilbert MT, Lund O, Wesolowska A, Karmin M, Weinert LA, Wang B, Li J, Tai S, Xiao F, Hanihara T, van Driem G, Jha AR, Ricaut FX, de Knijff P, Migliano AB, Gallego Romero I, Kristiansen K, Lambert DM, Brunak S, Forster P, Brinkmann B, Nehlich O, Bunce M, Richards M, Gupta R, Bustamante CD, Krogh A, Foley RA, Lahr MM, Balloux F, Sicheritz-Pontén T, Villems R, Nielsen R, Wang J, Willerslev E |display-authors=6 |title=An Aboriginal Australian genome reveals separate human dispersals into Asia |journal=[[Science (journal)|Science]] |volume=334 |issue=6052 |pages=94–98 |date=October 2011 |pmid=21940856 |pmc=3991479 |doi=10.1126/science.1211177 |bibcode=2011Sci...334...94R}}</ref> The rate of Neanderthal admixture to European and Asian populations as of 2017 has been estimated at between about 2–3%.<ref>East Asians 2.3–2.6%, Western Eurasians 1.8–2.4% ({{cite journal |vauthors=Prüfer K, de Filippo C, Grote S, Mafessoni F, Korlević P, Hajdinjak M, Vernot B, Skov L, Hsieh P, Peyrégne S, Reher D, Hopfe C, Nagel S, Maricic T, Fu Q, Theunert C, Rogers R, Skoglund P, Chintalapati M, Dannemann M, Nelson BJ, Key FM, Rudan P, Kućan Ž, Gušić I, Golovanova LV, Doronichev VB, Patterson N, Reich D, Eichler EE, Slatkin M, Schierup MH, Andrés AM, Kelso J, Meyer M, Pääbo S |display-authors=6 |title=A high-coverage Neandertal genome from Vindija Cave in Croatia |journal=[[Science (journal)|Science]] |volume=358 |issue=6363 |pages=655–658 |date=November 2017 |pmid=28982794 |pmc=6185897 |doi=10.1126/science.aao1887 |bibcode=2017Sci...358..655P}})</ref>


Archaic admixture in some Sub-Saharan African populations hunter-gatherer groups ([[Aka people|Biaka]] Pygmies and [[San people|San]]), derived from archaic hominins that broke away from the modern human lineage around 700,000 years ago, was discovered in 2011. The rate of admixture was estimated at 2%.<ref name="hamgenev" /> Admixture from archaic hominins of still earlier divergence times, estimated at 1.2 to 1.3 million years ago, was found in [[Pygmy peoples|Pygmies]], [[Hadza people|Hadza]] and five [[Sandawe people|Sandawe]] in 2012.<ref>{{cite journal |vauthors=Callaway E |title=Hunter-gatherer genomes a trove of genetic diversity |journal=[[Nature (journal)|Nature]] |url=http://www.nature.com/news/hunter-gatherer-genomes-a-trove-of-genetic-diversity-1.11076 |doi=10.1038/nature.2012.11076 |date=2012 |s2cid=87081207}}</ref><ref name="lac12adaafr" />
===Pathogen DNA===
Another promising route towards reconstructing human genetic genealogy is via the [[JC virus]] (JCV), a type of human [[polyomavirus]] which is carried by 70–90 percent of humans and which is usually transmitted vertically, from parents to offspring, suggesting codivergence with human populations. For this reason, JCV has been used as a genetic marker for human evolution and migration.<ref>Elizabeth Matisoo-Smith, K. Ann Horsburgh, ''DNA for Archaeologists'', Routledge (2016).</ref> This method does not appear to be reliable for the migration out of Africa, in contrast to human genetics, JCV strains associated with African populations are not basal. From this {{harvp|Shackelton et al.|2006}} conclude that either a basal African strain of JCV has become extinct or that the original infection with JCV post-dates the migration from Africa.


From an analysis of [[Mucin 7]], a highly divergent haplotype that has an estimated coalescence time with other variants around 4.5 million years BP and is specific to African populations, it is inferred to have been derived from interbreeding between African modern and archaic humans.<ref>{{cite journal |vauthors=Xu D, Pavlidis P, Taskent RO, Alachiotis N, Flanagan C, DeGiorgio M, Blekhman R, Ruhl S, Gokcumen O |display-authors=6 |title=Archaic Hominin Introgression in Africa Contributes to Functional Salivary MUC7 Genetic Variation |journal=[[Molecular Biology and Evolution]] |volume=34 |issue=10 |pages=2704–2715 |date=October 2017 |pmid=28957509 |pmc=5850612 |doi=10.1093/molbev/msx206}}</ref>
===Admixture of archaic and modern humans===
{{main|Archaic human admixture with modern humans}}
Evidence for [[archaic human]] species (descended from ''[[Homo heidelbergensis]]'') having [[Archaic human admixture with modern humans|interbred]] with modern humans outside of Africa, was discovered in the 2010s. This concerns primarily [[Neanderthals|Neanderthal]] admixture in all modern populations except for [[Sub-Saharan Africans]] but evidence has also been presented for [[Denisova hominin]] admixture in [[Australasia]] (i.e. in [[Melanesians]], [[Aboriginal Australians]] and some [[Negritos]]).<ref>{{cite journal | last=Rasmussen | first=M. |author2=Guo, X. |author3=Wang, Y. |author4=Lohmueller, K.E. |author5=Rasmussen, S. |author6=Albrechtsen, A. | title=An Aboriginal Australian Genome Reveals Separate Human Dispersals into Asia | journal=Science | year=2011 | volume=334 | issue=6052 | pages=94–98 | doi=10.1126/science.1211177|display-authors=etal | pmid=21940856 | pmc=3991479}}</ref>


A study published in 2020 found that the [[Yoruba people|Yoruba]] and [[Mende people|Mende]] populations of West Africa derive between 2% and 19% of their genome from an as-yet unidentified archaic hominin population that likely diverged before the split of modern humans and the ancestors of Neanderthals and Denisovans.<ref>{{cite journal | vauthors = Durvasula A, Sankararaman S | title = Recovering signals of ghost archaic introgression in African populations | journal = Science Advances | volume = 6 | issue = 7 | date = February 2020 | pages = eaax5097 | doi = 10.1126/sciadv.aax5097| pmid = 32095519 | pmc = 7015685 | bibcode = 2020SciA....6.5097D | doi-access = free }}</ref>
The rate of admixture of Neanderthal admixture to European and Asian populations as of 2017 has been estimated at between about 2%&ndash;3%.<ref>East Asians 2.3-2.6%, Western Eurasians 1.8-2.4% ({{cite journal| last1=Prüfer|first1=K.| last2=de Filippo|first2=C.| last3=Grote|first3=S.| last4=Mafessoni|first4=F.| last5=Korlević|first5=P.| last6=Hajdinjak|first6=M.|title=A high-coverage Neandertal genome from Vindija Cave in Croatia| journal=Science| date=2017| doi=10.1126/science.aao1887 |display-authors=etal| volume=358| pages=655–658}})</ref>


=== Stone tools ===
Archaic admixture in some Sub-Saharan African populations hunter-gatherer groups ([[Aka people|Biaka]] Pygmies and [[San people|San]]), derived from archaic hominins that broke away from the modern human lineage around 700,000 years, was discovered in 2011. The rate of admixture was estimated at around 2%.<ref name=hamgenev>{{cite journal | last=Hammer | first=M.F.|author2=Woerner, A.E.|author3= Mendez, F.L.|author4= Watkins, J.C.|author5= Wall, J.D. | title=Genetic evidence for archaic admixture in Africa | journal=Proceedings of the National Academy of Sciences | year=2011 | volume=108 | issue=37 | pages=15123–15128 | doi=10.1073/pnas.1109300108 | pmid=21896735 | pmc=3174671}}</ref>
{{see|Aterian|Baradostian|Microlith}}
Admixture from archaic hominins of still earlier divergence times, estimated at 1.2 to 1.3 million years ago, was found in [[Pygmy peoples|Pygmies]], [[Hadza people|Hadza]] and five [[Sandawe people|Sandawe]] in 2012.<ref>{{cite web | last=Callaway | first=E. | title=Hunter-gatherer genomes a trove of genetic diversity | work=Nature | url=http://www.nature.com/news/hunter-gatherer-genomes-a-trove-of-genetic-diversity-1.11076 | doi=10.1038/nature.2012.11076 | date=2012}}</ref><ref name=lac12adaafr>{{cite journal| last=Lachance| first=J.|author2=Vernot, B. |author3=Elbers, C.C. |author4=Ferwerda, B. |author5=Froment, A. |author6=Bodo, J.M. | title=Evolutionary History and Adaptation from High-Coverage Whole-Genome Sequences of Diverse African Hunter-Gatherers| journal=Cell| date= 2012| volume=150| issue=3| pages=457–469| doi=10.1016/j.cell.2012.07.009|display-authors=etal | pmid=22840920 | pmc=3426505}}</ref>
Archaic admixture in West African agricultural populations ([[Mende people|Mende]] and [[Yoruba people|Yoruba]]) was found in 2017.<ref>{{cite journal|author1=Xu, D.| author2 = Pavlidis, P. | author3= Taskent, O.R. | author4= Alachiotis, N. | author5= Flanagan, C. | author6= DeGiorgio, M. | title=Archaic Hominin Introgression in Africa Contributes to Functional Salivary MUC7 Genetic Variation| journal=Molecular Biology and Evolution |date=2017 |volume=34|issue=10| pages=2704–2715|url=https://academic.oup.com/mbe/article/34/10/2704/3988100/Archaic-Hominin-Introgression-in-Africa | display-authors=etal | doi=10.1093/molbev/msx206}}</ref><ref>{{cite journal|last1=Skoglund et al.|title=Reconstructing Prehistoric African Population Structure|journal=Cell|date=21 September 2017|volume=171|pages=59–71|doi=10.1016/j.cell.2017.08.049|url=http://www.cell.com/cell/fulltext/S0092-8674(17)31008-5|accessdate=6 November 2017}}</ref>


In addition to genetic analysis, Petraglia ''et al.'' also examines the small stone tools ([[microlith]]ic materials) from the Indian subcontinent and explains the expansion of population based on the reconstruction of paleoenvironment. He proposed that the stone tools could be dated to 35 ka in South Asia, and the new technology might be influenced by environmental change and population pressure.<ref>{{cite journal |vauthors=Petraglia M, Clarkson C, Boivin N, Haslam M, Korisettar R, Chaubey G, Ditchfield P, Fuller D, James H, Jones S, Kivisild T, Koshy J, Lahr MM, Metspalu M, Roberts R, Arnold L |display-authors=6 |title=Population increase and environmental deterioration correspond with microlithic innovations in South Asia ca. 35,000 years ago |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=106 |issue=30 |pages=12261–12266 |date=July 2009 |pmid=19620737 |pmc=2718386 |doi=10.1073/pnas.0810842106 |bibcode=2009PNAS..10612261P |doi-access=free}}</ref>
===Stone tools===


== History of the theory ==
{{see|Aterian|Baradostian|Microlith}}
=== Classical paleoanthropology ===
{{Further|Timeline of human evolution|Paleoanthropology#History of paleoanthropology}}
[[File:Huxley - Mans Place in Nature.jpg|thumb|upright=1.25|The frontispiece to Huxley's ''[[Evidence as to Man's Place in Nature]]'' (1863): the image compares the skeleton of a human to other apes.]]
[[File:Asian origin thesis for L3.jpg|thumb|The possibility of an origin of [[Haplogroup L3 (mtDNA)|L3]] in Asia was proposed by Cabrera et al. (2018).<ref>{{cite journal|vauthors=Cabrera VM, Marrero P, Abu-Amero KK, Larruga JM |date=June 2018 |title=Carriers of mitochondrial DNA macrohaplogroup L3 basal lineages migrated back to Africa from Asia around 70,000 years ago |journal=[[BMC Evolutionary Biology]] |volume=18 |issue=1 |pages=98 |doi=10.1186/s12862-018-1211-4 |pmc=6009813 |pmid=29921229 |biorxiv=10.1101/233502 |doi-access=free |bibcode=2018BMCEE..18...98C }}</ref><br />'''a:''' Exit of the [[Haplogroup L3 (mtDNA)|L3]] precursor to Eurasia. '''b:''' Return to Africa and expansion to Asia of basal L3 lineages with subsequent differentiation in both continents.]]


The cladistic relationship of humans with the African [[apes]] was suggested by [[Charles Darwin]] after studying the behaviour of African [[apes]], one of which was displayed at the [[London Zoo]].<ref name="Lafreniere2010">{{cite book |vauthors=Lafreniere P |title=Adaptive Origins: Evolution and Human Development |url=https://books.google.com/books?id=3F-Ms0mWKVYC&pg=PA90 |access-date=14 June 2011 |date=2010 |publisher=[[Taylor & Francis]] |isbn=978-0805860122 |pages=90 |via=[[Google Books]]}}</ref> The anatomist [[Thomas Huxley]] had also supported the hypothesis and suggested that African apes have a close evolutionary relationship with humans.<ref name="isbn0-470-01315-X">{{cite book |vauthors=Robinson D, Ash PM |title=The Emergence of Humans: An Exploration of the Evolutionary Timeline |publisher=[[Wiley (publishers)|Wiley]] |location=New York |year=2010 |isbn=978-0470013151}}</ref> These views were opposed by the German biologist [[Ernst Haeckel]], who was a proponent of the [[Out of Asia theory]]. Haeckel argued that humans were more closely related to the primates of South-east Asia and rejected Darwin's African hypothesis.<ref name="isbn0-520-24827-9">{{cite book |vauthors=Palmer D |title=Prehistoric Past Revealed: The Four Billion Year History of Life on Earth |publisher=[[University of California Press]] |location=Berkeley |year=2006 |isbn=978-0520248274 |page=43}}</ref><ref name="isbn1-85109-418-0">{{cite book |vauthors=Regal B |title=Human evolution: a guide to the debates |publisher=[[ABC-CLIO]] |location=Santa Barbara, Calif |year=2004 |isbn=978-1851094189 |pages=73–75}}</ref>
In addition to genetic analysis, Petraglia ''et al.'' also examines the small stone tools ([[microlith]]ic materials) from Indian subcontinent and explains the expansion of population based on the reconstruction of paleoenvironment. He proposed that the stone tools could be dated to 35 ka in South Asia, and the new technology might be influenced by environmental change and population pressure.<ref>{{cite journal | last1 = Petraglia | first1 = M. | last2 = Clarkson | first2 = C. | last3 = Boivin | first3 = N. | last4 = Haslam | first4 = M. | last5 = Korisettar | first5 = R. | last6 = Chaubey | first6 = G. | last7 = Arnold | first7 = L. | year = 2009 | title = Population increase and environmental deterioration correspond with microlithic innovations in South Asia ca. 35,000 years ago | journal = Proceedings of the National Academy of Sciences | volume = 106 | issue = 30| pages = 12261–12266 | doi=10.1073/pnas.0810842106 | pmid=19620737 | pmc=2718386 }}</ref>


In ''[[The Descent of Man]]'', Darwin speculated that humans had descended from apes, which still had small brains but walked upright, freeing their hands for uses which favoured intelligence; he thought such apes were African:
==History of the theory==


{{blockquote|In each great region of the world the living [[mammals]] are closely related to the extinct species of the same region. It is, therefore, probable that Africa was formerly inhabited by extinct apes closely allied to the [[gorilla]] and [[chimpanzee]]; and as these two species are now man's nearest allies, it is somewhat more probable that our early progenitors lived on the African continent than elsewhere. But it is useless to speculate on this subject, for an ape nearly as large as a man, namely the [[Dryopithecus]] of Lartet, which was closely allied to the anthropomorphous [[Hylobates]], existed in Europe during the [[Upper Miocene]] period; and since so remote a period the earth has certainly undergone many great revolutions, and there has been ample time for migration on the largest scale.|Charles Darwin|Descent of Man<ref>{{cite web |url=http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F937.1&pageseq=212 |title=''The Descent of Man'' Chapter 6 – "On the Affinities and Genealogy of Man" |website=Darwin-online.org.uk |access-date=11 January 2011}}</ref>}}
===Classical paleoanthropology===


In 1871, there were hardly any human fossils of ancient hominins available. Almost fifty years later, Darwin's speculation was supported when anthropologists began finding fossils of ancient small-brained hominins in several areas of Africa ([[list of hominina fossils]]). The hypothesis of ''recent'' (as opposed to [[Out of Africa I|archaic]]) African origin developed in the 20th century. The "Recent African origin" of modern humans means "single origin" (monogenism) and has been used in various contexts as an antonym to polygenism. The debate in anthropology had swung in favour of monogenism by the mid-20th century. Isolated proponents of polygenism held forth in the mid-20th century, such as [[Carleton S. Coon|Carleton Coon]], who thought as late as 1962 that ''H. sapiens'' arose five times from ''H. erectus'' in five places.<ref name=Jackson_2001>{{cite journal |vauthors=Jackson JP Jr |title='In Ways Unacademical': The Reception of Carleton S. Coon's The Origin of Races |journal=[[Journal of the History of Biology]] |year=2001 |volume=34 |issue=2 |pages=247–285 |doi=10.1023/A:1010366015968 |s2cid=86739986 |url=http://comm.colorado.edu/~jacksonj/research/coon.pdf |archive-url=https://web.archive.org/web/20130514075459/http://comm.colorado.edu/~jacksonj/research/coon.pdf |archive-date=14 May 2013 |df=dmy-all}}</ref>
{{further|Timeline of human evolution|Paleoanthropology#History_of_paleoanthropology}}


=== Multiregional origin hypothesis ===
[[File:Huxley - Mans Place in Nature.jpg|thumb|The frontispiece to Huxley's ''[[Evidence as to Man's Place in Nature]]'' (1863): the image compares the skeleton of a human to other apes.]]
{{Main|Multiregional origin of modern humans}}
The cladistic relationship of humans with the African [[apes]] was suggested by [[Charles Darwin]] after studying the behaviour of African [[apes]], one of which was displayed at the [[London Zoo]].<ref name="Lafreniere2010">{{cite book|author=Peter Lafreniere|title=Adaptive Origins: Evolution and Human Development|url=https://books.google.com/books?id=3F-Ms0mWKVYC&pg=PA90|accessdate=14 June 2011|date=22 September 2010|publisher=Taylor & Francis|isbn=978-0-8058-6012-2|page=90}}</ref> The anatomist [[Thomas Huxley]] had also supported the hypothesis and suggested that African apes have a close evolutionary relationship with humans.<ref name="isbn0-470-01315-X">{{cite book |vauthors=Robinson D, Ash PM | title = The Emergence of Humans: An Exploration of the Evolutionary Timeline | publisher = Wiley | location = New York | year = 2010 | pages = | isbn = 0-470-01315-X }}</ref> These views were opposed by the German biologist [[Ernst Haeckel]], who was a proponent of the [[Out of Asia theory]]. Haeckel argued that humans were more closely related to the primates of South-east Asia and rejected Darwin's African hypothesis.<ref name="isbn0-520-24827-9">{{cite book | author = Palmer D | title = Prehistoric Past Revealed: The Four Billion Year History of Life on Earth | publisher = University of California Press | location = Berkeley | year = 2006 | pages = | isbn = 0-520-24827-9 | page = 43 }}</ref><ref name="isbn1-85109-418-0">{{cite book | author = Regal B | title = Human evolution: a guide to the debates | publisher = ABC-CLIO | location = Santa Barbara, Calif | year = 2004 | isbn = 1-85109-418-0 | pages = 73–75 }}</ref>


The historical alternative to the recent origin model is the [[multiregional origin of modern humans]], initially proposed by [[Milford Wolpoff]] in the 1980s. This view proposes that the derivation of anatomically modern human populations from ''H. erectus'' at the beginning of the [[Pleistocene]] 1.8 million years BP, has taken place within a continuous world population. The hypothesis necessarily rejects the assumption of an [[species barrier|infertility barrier]] between ancient Eurasian and African populations of ''Homo''. The hypothesis was controversially debated during the late 1980s and the 1990s.<ref>{{cite journal |vauthors=Stringer CB, Andrews P |title=Genetic and fossil evidence for the origin of modern humans |journal=[[Science (journal)|Science]] |volume=239 |issue=4845 |pages=1263–1268 |date=March 1988 |pmid=3125610 |doi=10.1126/science.3125610 |bibcode=1988Sci...239.1263S}}
In the ''[[Descent of Man]]'', Darwin speculated that humans had descended from apes, which still had small brains but walked upright, freeing their hands for uses which favoured intelligence; he thought such apes were African:
{{cite journal |vauthors=Stringer C, Bräuer G |year=1994 |title=Methods, misreading, and bias |journal=[[American Anthropologist]] |volume=96 |issue=2 |pages=416–424 |doi=10.1525/aa.1994.96.2.02a00080}}<br />
{{cite book |vauthors=Stringer CB |date=1992 |chapter=Replacement, continuity and the origin of Homo sapiens |title=Continuity or replacement? Controversies in Homo sapiens evolution |veditors=Smith FH |location=Rotterdam |publisher=Balkema |pages=9–24}}<br />
{{cite book |vauthors=Bräuer G, Stringer C |date=1997 |chapter=Models, polarization, and perspectives on modern human origins |title=Conceptual issues in modern human origins research |location=New York |publisher=Aldine de Gruyter |pages=191–201}}</ref> The now-current terminology of "recent-origin" and "Out of Africa" became current in the context of this debate in the 1990s.<ref>{{cite journal |vauthors=Wu L |title=The dental continuity of humans in China from Pleistocene to Holocene, and the origin of mongoloids |journal=Quaternary Geology |date=1997 |volume=21 |pages=24–32 |url=https://books.google.com/books?id=QZym919tNigC&pg=PA24 |isbn=978-9067642439}}</ref> Originally seen as an antithetical alternative to the recent origin model, the multiregional hypothesis in its original "strong" form is obsolete, while its various modified weaker variants have become variants of a view of "recent origin" combined with [[Archaic human admixture with modern humans|archaic admixture]].<ref>{{cite journal |vauthors=Stringer C |year=2001 |title=Modern human origins – distinguishing the models |journal=[[African Archaeological Review]] |volume=18 |issue=2 |pages=67–75 |doi=10.1023/A:1011079908461 |s2cid=161991922}}</ref> Stringer (2014) distinguishes the original or "classic" Multiregional model as having existed from 1984 (its formulation) until 2003, to a "weak" post-2003 variant that has "shifted close to that of the Assimilation Model".<ref>{{cite journal |vauthors=Stringer C |title=Modern human origins: progress and prospects |journal=[[Philosophical Transactions of the Royal Society B|Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences]] |volume=357 |issue=1420 |pages=563–579 |date=April 2002 |pmid=12028792 |pmc=1692961 |doi=10.1098/rstb.2001.1057}}</ref><ref>{{cite journal |vauthors=Stringer C |title=Why we are not all multiregionalists now |journal=[[Trends in Ecology & Evolution]] |volume=29 |issue=5 |pages=248–251 |date=May 2014 |pmid=24702983 |doi=10.1016/j.tree.2014.03.001 |doi-access=free|bibcode=2014TEcoE..29..248S }}</ref>


=== Mitochondrial analyses ===
{{quote|In each great region of the world the living [[mammals]] are closely related to the extinct species of the same region. It is, therefore, probable that Africa was formerly inhabited by extinct apes closely allied to the [[gorilla]] and [[chimpanzee]]; and as these two species are now man's nearest allies, it is somewhat more probable that our early progenitors lived on the African continent than elsewhere. But it is useless to speculate on this subject, for an ape nearly as large as a man, namely the [[Dryopithecus]] of Lartet, which was closely allied to the anthropomorphous [[Hylobates]], existed in Europe during the [[Upper Miocene]] period; and since so remote a period the earth has certainly undergone many great revolutions, and there has been ample time for migration on the largest scale.|Charles Darwin|Descent of Man<ref>{{cite web | url = http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F937.1&pageseq=212 | title = The descent of man Chapter 6 – On the Affinities and Genealogy of Man | publisher = Darwin-online.org.uk | accessdate = 11 January 2011 }}</ref>}}
{{further|Mitochondrial Eve}}
In the 1980s, [[Allan Wilson (biologist)|Allan Wilson]] together with [[Rebecca L. Cann]] and [[Mark Stoneking]] worked on genetic dating of the matrilineal most recent common ancestor of modern human populations (dubbed "[[Mitochondrial Eve]]"). To identify informative [[genetic marker]]s for tracking human evolutionary history, Wilson concentrated on [[mitochondrial DNA]] (mtDNA), which is maternally inherited. This DNA material mutates quickly, making it easy to plot changes over relatively short times. With his discovery that human mtDNA is genetically much less diverse than chimpanzee mtDNA, Wilson concluded that modern human populations had diverged recently from a single population while older human species such as [[Neanderthal]]s and ''[[Homo erectus]]'' had become extinct.<ref name="url_Allan_Wilson">{{cite web |url=http://www.nzedge.com/heroes/wilson.html |title=Allan Wilson: Revolutionary Evolutionist |work=New Zealanders Heroes}}</ref> With the advent of [[archaeogenetics]] in the 1990s, the dating of mitochondrial and [[Human Y-chromosome DNA haplogroup|Y-chromosomal haplogroups]] became possible with some confidence. By 1999, estimates ranged around 150,000 years for the [[mt-MRCA]] and 60,000 to 70,000 years for the migration out of Africa.<ref>{{cite journal |vauthors=Wallace DC, Brown MD, Lott MT |title=Mitochondrial DNA variation in human evolution and disease |journal=[[Gene (journal)|Gene]] |volume=238 |issue=1 |pages=211–230 |date=September 1999 |pmid=10570998 |doi=10.1016/S0378-1119(99)00295-4}} "evidence that our species arose in Africa about 150,000 years before present (YBP), migrated out of Africa into Asia about 60,000 to 70,000 YBP and into Europe about 40,000 to 50,000 YBP, and migrated from Asia and possibly Europe to the Americas about 20,000 to 30,000 YBP."</ref>


From 2000 to 2003, there was controversy about the mitochondrial DNA of "[[Mungo Man|Mungo Man 3]]" (LM3) and its possible bearing on the [[Multiregional origin of modern humans|multiregional hypothesis]]. LM3 was found to have more than the expected number of sequence differences when compared to modern human DNA ([[Cambridge Reference Sequence|CRS]]).<ref name="Adcock">{{cite journal |vauthors=Adcock GJ, Dennis ES, Easteal S, Huttley GA, Jermiin LS, Peacock WJ, Thorne A |title=Mitochondrial DNA sequences in ancient Australians: Implications for modern human origins |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=98 |issue=2 |pages=537–542 |date=January 2001 |pmid=11209053 |pmc=14622 |doi=10.1073/pnas.98.2.537 |bibcode=2001PNAS...98..537A |doi-access=free}}</ref> Comparison of the mitochondrial DNA with that of ancient and modern [[Aboriginal Australians|aborigines]], led to the conclusion that Mungo Man fell outside the range of genetic variation seen in Aboriginal Australians and was used to support the multiregional origin hypothesis. A reanalysis of LM3 and other ancient specimens from the area published in 2016, showed it to be akin to modern Aboriginal Australian sequences, inconsistent with the results of the earlier study.<ref>{{cite journal |vauthors=Heupink TH, Subramanian S, Wright JL, Endicott P, Westaway MC, Huynen L, Parson W, Millar CD, Willerslev E, Lambert DM |display-authors=6 |title=Ancient mtDNA sequences from the First Australians revisited |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=113 |issue=25 |pages=6892–6897 |date=June 2016 |pmid=27274055 |pmc=4922152 |doi=10.1073/pnas.1521066113 |bibcode=2016PNAS..113.6892H |doi-access=free}}</ref>
In 1871 there were hardly any human fossils of ancient hominins available. Almost fifty years later, Darwin's speculation was supported when anthropologists began finding fossils of ancient small-brained hominins in several areas of Africa ([[list of hominina fossils]]). The hypothesis of ''recent'' (as opposed to [[Out of Africa I|archaic]]) African origin developed in the 20th century. The "Recent African origin" of modern humans means "single origin" (monogenism) and has been used in various contexts as an antonym to polygenism. The debate in anthropology had swung in favour of monogenism by the mid-20th century. Isolated proponents of polygenism held forth in the mid-20th century, such as [[Carleton S. Coon|Carleton Coon]], who thought as late as 1962 that ''H. sapiens'' arose five times from ''H. erectus'' in five places.<ref name=Jackson_2001>{{cite journal | author = Jackson JP Jr | title = 'In Ways Unacademical': The Reception of Carleton S. Coon's The Origin of Races | journal = Journal of the History of Biology | year = 2001 | volume = 34 | issue = 2 | pages = 247–285 | doi = 10.1023/A:1010366015968 | url = http://comm.colorado.edu/~jacksonj/research/coon.pdf | format = pdf | deadurl = yes | archiveurl = https://web.archive.org/web/20130514075459/http://comm.colorado.edu/~jacksonj/research/coon.pdf | archivedate = 14 May 2013 | df = dmy-all }}</ref>


=== Y-chromosome analyses ===
===Multiregional origin hypothesis===
{{further|Y-chromosomal Adam}}
[[File:World Map of Y-DNA Haplogroups.png|thumb|upright=1.3|Map of Y-chromosome haplogroups – dominant haplogroups in pre-colonial populations with proposed migrations routes]]


The [[Y chromosome]], which is paternally inherited, does not go through much recombination and thus stays largely the same after inheritance. Similar to Mitochondrial Eve, this could be studied to track the male most recent common ancestor ("[[Y-chromosomal Adam]]" or Y-MRCA).<ref>{{Cite book |last1=Parsons |first1=Paul |title=50 Ideas You Really Need to Know: Science |last2=Dixon |first2=Gail |publisher=[[Quercus]] |year=2016 |isbn=978-1-78429-614-8 |location=London |pages=127 |language=en}}</ref>
{{Main article|Multiregional origin of modern humans}}


The most basal lineages have been detected in [[West Africa|West]], [[Northwest Africa|Northwest]] and [[Central Africa]], suggesting plausibility for the Y-MRCA living in the general region of "Central-Northwest Africa".<ref name="pmid21601174" />
The historical alternative to the recent origin model is the [[multiregional origin of modern humans]], initially proposed by [[Milford Wolpoff]] in the 1980s. This view proposes that the derivation of anatomically modern human populations from ''H. erectus'' at the beginning of the [[Pleistocene]] 1.8 million years BP, has taken place within a continuous world population. The hypothesis necessarily rejects the assumption of an [[species barrier|infertility barrier]] between ancient Eurasian and African populations of ''Homo''. The hypothesis was controversially debated during the late 1980s and the 1990s.<ref>{{cite journal | last1 = Stringer | first1 = C. B. | last2 = Andrews | first2 = P. | year = 1988 | title = Genetic and fossil evidence for the origin of modern humans | url = http://www.sciencemag.org/content/239/4845/1263.abstract | journal = Science | volume = 239 | issue = 4845| pages = 1263–1268 | doi=10.1126/science.3125610 | pmid=3125610}}
{{cite journal | last1 = Stringer | first1 = C. | last2 = Bräuer | first2 = G. | year = 1994 | title = Methods, misreading, and bias | url = | journal = American Anthropologist | volume = 96 | issue = 2| pages = 416–424 | doi=10.1525/aa.1994.96.2.02a00080}}<br>
Stringer, C. B. (1992). "Replacement, continuity and the origin of Homo sapiens". In: ''Continuity or replacement? Controversies in Homo sapiens evolution''. F. H. Smith (ed). Rotterdam: Balkema. pp. 9–24.<br>
Bräuer, G.; Stringer, C. (1997). "Models, polarization, and perspectives on modern human origins". In: ''Conceptual issues in modern human origins research''. New York: Aldine de Gruyter. pp. 191–201.</ref> The now-current terminology of "recent-origin" and "Out of Africa" became current in the context of this debate in the 1990s.<ref>Liu Wu
in Zhisheng, Weijian Zhou (eds.), ''Quaternary Geology'' VSP (1997), [https://books.google.com/books?id=QZym919tNigC&pg=PA24 p. 24].</ref> Originally seen as an antithetical alternative to the recent origin model, the multiregional hypothesis in its original "strong" form is obsolete, while its various modified weaker variants have become variants of a view of "recent origin" combined with [[Archaic human admixture with modern humans|archaic admixture]].<ref>{{cite journal | last1 = Stringer | first1 = C. | year = 2001 | title = Modern human origins—distinguishing the models | url = | journal = Afr. Archaeol. Rev | volume = 18 | issue = | pages = 67–75 }}</ref> Stringer (2014) distinguishes the original or "classic" Multiregional model as having existed from 1984 (its formulation) until 2003, to a "weak" post-2003 variant that has "shifted close to that of the Assimilation Model".<ref>{{cite journal | last1 = Stringer | first1 = C | year = 2002 | title = Modern human origins: progress and prospects | url = | journal = Philosophical Transactions of the Royal Society of London | volume = 357 | issue = 1420| pages = 563–579 | doi=10.1098/rstb.2001.1057}}</ref><ref>{{cite journal | last1 = Stringer | first1 = C. | year = 2014 | title = Why we are not all multiregionalists now | url = http://www.sciencedirect.com/science/article/pii/S0169534714000470 | journal = Trends in Ecology & Evolution | volume = 29 | issue = 5| pages = 248–251 | doi=10.1016/j.tree.2014.03.001}}</ref>


A [[Stanford University School of Medicine]] study was done by comparing Y-chromosome sequences and mtDNA in 69 men from different geographic regions and constructing a family tree. It was found that the Y-MRCA lived between 120,000 and 156,000, and the Mitochondrial Eve lived between 99,000 and 148,000 years ago, which not only predates some proposed waves of migration, but also meant that both lived in the [[Africa|African continent]] around the same time period.<ref>{{Cite journal |last1=Poznik |first1=G. David |last2=Henn |first2=Brenna M. |last3=Yee |first3=Muh-Ching |last4=Sliwerska |first4=Elzbieta |last5=Euskirchen |first5=Ghia M. |last6=Lin |first6=Alice A. |last7=Snyder |first7=Michael |last8=Quintana-Murci |first8=Lluis |last9=Kidd |first9=Jeffrey M. |last10=Underhill |first10=Peter A. |last11=Bustamante |first11=Carlos D. |date=2013-08-02 |title=Sequencing Y Chromosomes Resolves Discrepancy in Time to Common Ancestor of Males Versus Females |journal=Science |language=en |volume=341 |issue=6145 |pages=562–565 |doi=10.1126/science.1237619 |issn=0036-8075 |pmc=4032117 |pmid=23908239|bibcode=2013Sci...341..562P }}</ref>
===Genetics===
In the 1980s, [[Allan Wilson]] together with [[Rebecca L. Cann]] and [[Mark Stoneking]] worked on genetic dating of the matrilineal most recent common ancestor of modern human populations (dubbed "[[Mitochondrial Eve]]"). To identify informative [[genetic marker]]s for tracking human evolutionary history, Wilson concentrated on [[mitochondrial DNA]] (mtDNA), passed from mother to child. This DNA material mutates quickly, making it easy to plot changes over relatively short times. With his discovery that human mtDNA is genetically much less diverse than chimpanzee mtDNA, Wilson concluded that modern human populations had diverged recently from a single population while older human species such as [[Neanderthal]]s and ''[[Homo erectus]]'' had become extinct.<ref name="url_Allan_Wilson">{{cite web | url = http://www.nzedge.com/heroes/wilson.html | title = Allan Wilson: Revolutionary Evolutionist | work = New Zealanders Heroes | accessdate = }}</ref> With the advent of [[archaeogenetics]] in the 1990s, the dating of mitochondrial and [[Human Y-chromosome DNA haplogroup|Y-chromosomal haplogroups]] became possible with some confidence. By 1999, estimates ranged around 150,000 years for the [[mt-MRCA]] and 60,000 to 70,000 years for the migration out of Africa.<ref>{{cite journal |last1=Wallace |first1=D |last2=Brown |first2=MD |last3=Lott |first3=MT |title=Mitochondrial DNA variation in human evolution and disease |journal=Gene |volume=238 |issue=1 |pages=211–30 |year=1999 |pmid=10570998 |doi=10.1016/S0378-1119(99)00295-4}} "evidence that our species arose in Africa about 150 000 years before present (YBP), migrated out of Africa into Asia about 60 000 to 70 000 YBP and into Europe about 40 000 to 50 000 YBP, and migrated from Asia and possibly Europe to the Americas about 20 000 to 30 000 YBP."</ref>


Another study finds a plausible placement in "the north-western quadrant of the African continent" for the emergence of the A1b haplogroup.<ref>{{cite journal |vauthors=Scozzari R, Massaia A, D'Atanasio E, Myres NM, Perego UA, Trombetta B, Cruciani F |title=Molecular dissection of the basal clades in the human Y chromosome phylogenetic tree |journal=[[PLOS ONE]] |volume=7 |issue=11 |pages=e49170 |year=2012 |pmid=23145109 |pmc=3492319 |doi=10.1371/journal.pone.0049170 |veditors=Caramelli D |bibcode=2012PLoSO...749170S |doi-access=free}}</ref> The 2013 report of [[haplogroup A00]] found among the [[Mbo people (Cameroon)|Mbo people]] of western present-day [[Cameroon]] is also compatible with this picture.<ref name="Mendez13">{{cite journal |vauthors=Mendez FL, Krahn T, Schrack B, Krahn AM, Veeramah KR, Woerner AE, Fomine FL, Bradman N, Thomas MG, Karafet TM, Hammer MF |display-authors=6 |title=An African American paternal lineage adds an extremely ancient root to the human Y chromosome phylogenetic tree |journal=[[American Journal of Human Genetics]] |volume=92 |issue=3 |pages=454–459 |date=March 2013 |pmid=23453668 |pmc=3591855 |doi=10.1016/j.ajhg.2013.02.002 |url=http://haplogroup-a.com/Ancient-Root-AJHG2013.pdf}}</ref>
From 2000–2003, there was controversy about the mitochondrial DNA of "[[Mungo Man|Mungo Man 3]]" (LM3) and its possible bearing on the multiregional hypothesis. LM3 was found to have more than the expected number of sequence differences when compared to modern human DNA ([[Cambridge Reference Sequence|CRS]]).<ref name="Adcock">{{cite journal|vauthors=Adcock GJ, Dennis ES, Easteal S, Huttley GA, Jarmiin LS, Peacock WJ, Thorne A |title = Mitochondrial DNA sequences in ancient Australians: Implications for modern human origins | journal = PNAS |volume= 98|issue = 2|pages=537–542 |year =2001|pmid = 11209053|doi = 10.1073/pnas.98.2.537|pmc = 14622 |url=http://www.pnas.org/content/98/2/537.full?sid=8f289c70-86e5-43cf-9d98-2c7b22612d47}}</ref> Comparison of the mitochondrial DNA with that of ancient and modern aborigines, led to the conclusion that Mungo Man fell outside the range of genetic variation seen in Aboriginal Australians and was used to support the multiregional origin hypothesis. A reanalysis on LM3 and other ancient specimens from the area published in 2016, showed it to be akin to modern Aboriginal Australian sequences, inconsistent with the results of the earlier study.<ref>{{cite journal| vauthors=Heupink TH, Subramanian S, Wright JL, Endicott P, Carrington Westaway M, Huynen L, Parson W, Millar C, Willerslev E, Lambert DM | title= Ancient mtDNA sequences from the First Australians revisited| journal=PNAS|volume= 113|issue = |pages=6892–7 |year =2016|doi = 10.1073/pnas.1521066113 |url=http://www.pnas.org/content/early/2016/06/01/1521066113.full#T1 | pmid=27274055 | pmc=4922152}}</ref>


The revision of Y-chromosomal phylogeny since 2011 has affected estimates for the likely geographical origin of Y-MRCA as well as estimates on time depth. By the same reasoning, future discovery of presently-unknown archaic haplogroups in living people would again lead to such revisions. In particular, the possible presence of between 1% and 4% [[Archaic human admixture with modern humans|Neanderthal-derived DNA]] in Eurasian genomes implies that the (unlikely) event of a discovery of a single living Eurasian male exhibiting a Neanderthal patrilineal line would immediately push back T-MRCA ("time to MRCA") to at least twice its current estimate. However, the discovery of a Neanderthal Y-chromosome by Mendez ''et al''. was tempered by a 2016 study that suggests the extinction of Neanderthal patrilineages, as the lineage inferred from the Neanderthal sequence is outside of the range of [[anatomically modern humans|contemporary human]] genetic variation.<ref>{{cite journal |vauthors=Mendez FL, Poznik GD, Castellano S, Bustamante CD |title=The Divergence of Neandertal and Modern Human Y Chromosomes |journal=[[American Journal of Human Genetics]] |volume=98 |issue=4 |pages=728–734 |date=April 2016 |pmid=27058445 |pmc=4833433 |doi=10.1016/j.ajhg.2016.02.023}}</ref> Questions of geographical origin would become part of the debate on Neanderthal evolution from ''[[Homo erectus]]''.
==See also==

== See also ==
{{Div col}}
{{Div col}}
* {{annotated link|Behavioral modernity}}
* [[Archaeogenetics of the Near East]]
* {{annotated link|Southern Dispersal}}
* [[Behavioral modernity]]
* {{annotated link|Dawn of Humanity|''Dawn of Humanity'' (2015 PBS film)}}
* [[Coastal migration]]
* {{annotated link|Early human migrations}}
* [[Dawn of Humanity (film)|''Dawn of Humanity'' (2015 PBS film)]]
* {{annotated link|Genetics and archaeogenetics of South Asia}}
* [[Early human migrations]]
* {{annotated link|Genetic history of Europe}}
* [[Genetics and archaeogenetics of South Asia]]
* [[Genetic history of Europe]]
* {{annotated link|Genetic history of Indigenous peoples of the Americas}}
* [[Genetic history of indigenous peoples of the Americas]]
* {{annotated link|Genetic history of Italy}}
* [[Genetic history of Italy]]
* {{annotated link|Genetic history of North Africa}}
* [[Genetic history of North Africa]]
* {{annotated link|Genetic history of the British Isles}}
* [[Genetic history of the British Isles]]
* {{annotated link|Genetic history of the Iberian Peninsula}}
* [[Genetic history of the Iberian Peninsula]]
* {{annotated link|Genetic history of the Middle East}}
* [[Hofmeyr Skull]]
* {{annotated link|Hofmeyr Skull}}
* {{annotated link|Human evolution}}
* {{annotated link|Human origins (disambiguation)|Human origins}}
* [[Template:Human timeline|Human timeline]]
* [[Template:Human timeline|Human timeline]]
* [[Identical ancestors point]]
* {{annotated link|Identical ancestors point}}
* [[Indo-Aryan migration theory]]
* {{annotated link|Indo-Aryan migrations}}
* [[Sahara pump theory]]
* {{annotated link|Sahara pump theory}}
* ''[[The Incredible Human Journey]]''
* {{annotated link|The Incredible Human Journey|''The Incredible Human Journey''}}
* [[Timeline of human evolution]]
* {{annotated link|Timeline of human evolution}}
* [[Human origins (disambiguation)|Human origins]]
* [[Human evolution]]
{{div col end}}
{{div col end}}


==Notes==
== Notes ==
{{Notelist}}
{{reflist|group=note|35em}}


==References==
== References ==
{{reflist|colwidth=25em|refs=
{{reflist|colwidth=32em|refs=


<ref name=pmid16826514Quo>{{harvp|Liu, Prugnolle et al.|2006}}. "Currently available genetic and archaeological evidence is supportive of a recent single origin of modern humans in East Africa. However, this is where the consensus on human settlement history ends, and considerable uncertainty clouds any more detailed aspect of human colonization history."</ref>
<ref name=pmid16826514Quo>{{harvp|Liu, Prugnolle et al.|2006}}. "Currently available genetic and archaeological evidence is supportive of a recent single origin of modern humans in East Africa. However, this is where the consensus on human settlement history ends, and considerable uncertainty clouds any more detailed aspect of human colonization history."</ref>


<ref name=pmid12802315>{{cite journal |author=Stringer, Chris |date=June 2003 |title=Human evolution: Out of Ethiopia |journal=[[Nature (journal)|Nature]] |volume=423 |issue= 6941 |pp =692–3, 695 |pmid=12802315 |doi=10.1038/423692a |bibcode=2003Natur.423..692S }}</ref>
<ref name=pmid12802315>{{cite journal |vauthors=Stringer C |title=Human evolution: Out of Ethiopia |journal=[[Nature (journal)|Nature]] |volume=423 |issue=6941 |pages=692–693, 695 |date=June 2003 |pmid=12802315 |doi=10.1038/423692a |bibcode=2003Natur.423..692S |s2cid=26693109}}</ref>


<ref name=pmid10766948>{{cite journal |last1=Wolpoff |first1=Milford H. |last2=Hawks |first2=John |last3=Caspari |first3=Rachel |date=May 2000 |title=Multiregional, not multiple origins |volume=112 |issue=1 |journal=[[American Journal of Physical Anthropology|Am. J. Phys. Anthropol.]] |pp=129–36 |pmid=10766948 |doi=10.1002/(SICI)1096-8644(200005)112:1<129::AID-AJPA11>3.0.CO;2-K }}</ref>
<ref name=pmid10766948>{{cite journal |vauthors=Wolpoff MH, Hawks J, Caspari R |title=Multiregional, not multiple origins |journal=American Journal of Physical Anthropology |volume=112 |issue=1 |pages=129–136 |date=May 2000 |pmid=10766948 |doi=10.1002/(SICI)1096-8644(200005)112:1<129::AID-AJPA11>3.0.CO;2-K |url=https://deepblue.lib.umich.edu/bitstream/2027.42/34270/1/11_ftp.pdf |hdl=2027.42/34270 |hdl-access=free}}</ref>


<ref name=pmid21273486>{{cite journal |last1=Armitage |first1=Simon J. |last2=Jasim |first2=Sabah A. |last3=Marks |first3=Anthony E. |last4=Parker |first4=Adrian G. |last5=Usik |first5=Vitaly I. |last6=Uerpmann |first6=Hans-Peter |date=January 2011 |title=The southern route "out of Africa": evidence for an early expansion of modern humans into Arabia |journal=[[Science (journal)|Science]] |volume=331 |issue=6016 |pp=453–6 |pmid=21273486 |doi=10.1126/science.1199113 |bibcode=2011Sci...331..453A }}</ref>
<ref name=pmid21273486>{{cite journal |vauthors=Armitage SJ, Jasim SA, Marks AE, Parker AG, Usik VI, Uerpmann HP |title=The southern route "out of Africa": evidence for an early expansion of modern humans into Arabia |journal=[[Science (journal)|Science]] |volume=331 |issue=6016 |pages=453–456 |date=January 2011 |pmid=21273486 |doi=10.1126/science.1199113 |bibcode=2011Sci...331..453A |s2cid=20296624}}</ref>


<ref name=pmid21212332>{{cite journal |author=Balter Michael |date=January 2011 |title=Was North Africa the launch pad for modern human migrations? |journal=Science |volume=331 |issue=6013 |pp= 20–3 |pmid=21212332| doi=10.1126/science.331.6013.20 |bibcode=2011Sci...331...20B |url=https://www.springer.com/cda/content/document/cda_downloaddocument/North+Africa+(+Aterian)+possible+source+of+Eurasian+modern+humans--Balter+Science+news.pdf?SGWID=0-0-45-1058837-p173624756 }}</ref>
<ref name=pmid21212332>{{cite journal |vauthors=Balter M |title=Was North Africa the launch pad for modern human migrations? |journal=[[Science (journal)|Science]] |volume=331 |issue=6013 |pages=20–23 |date=January 2011 |pmid=21212332 |doi=10.1126/science.331.6013.20 |url=https://www.springer.com/cda/content/document/cda_downloaddocument/North+Africa+(+Aterian)+possible+source+of+Eurasian+modern+humans--Balter+Science+news.pdf?SGWID=0-0-45-1058837-p173624756 |bibcode=2011Sci...331...20B}}</ref>


<ref name=pmid21601174>{{cite journal |last1=Cruciani |first1=Fulvio |last2=Trombetta |first2=Beniamino |last3=Massaia |first3=Andrea |last4=Destro-Bisol |first4=Giovanni |last5=Sellitto |first5=Daniele |last6=Scozzari |first6=Rosaria |date=June 2011 |title=A revised root for the human Y chromosomal phylogenetic tree: the origin of patrilineal diversity in Africa |journal=[[American Journal of Human Genetics|AJHG]] |volume=88 |issue=6 |pp= 814–8 |pmid=21601174 |pmc=3113241 |doi=10.1016/j.ajhg.2011.05.002 }}</ref>
<ref name=pmid21601174>{{cite journal |vauthors=Cruciani F, Trombetta B, Massaia A, Destro-Bisol G, Sellitto D, Scozzari R |title=A revised root for the human Y chromosomal phylogenetic tree: the origin of patrilineal diversity in Africa |journal=[[American Journal of Human Genetics]] |volume=88 |issue=6 |pages=814–818 |date=June 2011 |pmid=21601174 |pmc=3113241 |doi=10.1016/j.ajhg.2011.05.002}}</ref>


<ref name=pmid17372199>{{cite journal |last1=Smith |first1=Tanya M. |last2=Tafforeau |first2=Paul |last3=Reid |first3=Donald J. |last4=Grün |first4=Rainer |last5=Eggins |first5=Stephen |last6=Boutakiout |first6=Mohamed |last7=Hublin |first7=Jean-Jacques |date=April 2007 |title=Earliest evidence of modern human life history in North African early ''Homo sapiens'' |journal=[[Proceedings of the National Academy of Sciences of the United States of America|PNAS]] |volume=104 |issue=15 |pp=6128–33 |pmid=17372199 |pmc=1828706 |doi=10.1073/pnas.0700747104 |bibcode= 2007PNAS..104.6128S }}</ref>
<ref name=pmid17372199>{{cite journal |vauthors=Smith TM, Tafforeau P, Reid DJ, Grün R, Eggins S, Boutakiout M, Hublin JJ |title=Earliest evidence of modern human life history in North African early Homo sapiens |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=104 |issue=15 |pages=6128–6133 |date=April 2007 |pmid=17372199 |pmc=1828706 |doi=10.1073/pnas.0700747104 |bibcode=2007PNAS..104.6128S |doi-access=free}}</ref>


<ref name=pmid15339343>{{cite journal |vauthors=Metspalu M, Kivisild T, Metspalu E, Parik J, Hudjashov G, Kaldma K, Serk P, Karmin M, Behar DM, Gilbert MT, Endicott P, Mastana S, Papiha SS, Skorecki K, Torroni A, Villems R |date=August 2004 |title=Most of the extant mtDNA boundaries in south and southwest Asia were likely shaped during the initial settlement of Eurasia by anatomically modern humans |journal=BMC Genet. |volume=5 |issue = |p=26 |pmid=15339343 |pmc=516768 |doi=10.1186/1471-2156-5-26 }}</ref>
<ref name=pmid15339343>{{cite journal |vauthors=Metspalu M, Kivisild T, Metspalu E, Parik J, Hudjashov G, Kaldma K, Serk P, Karmin M, Behar DM, Gilbert MT, Endicott P, Mastana S, Papiha SS, Skorecki K, Torroni A, Villems R |display-authors=6 |title=Most of the extant mtDNA boundaries in south and southwest Asia were likely shaped during the initial settlement of Eurasia by anatomically modern humans |journal=[[BMC Genetics]] |volume=5 |pages=26 |date=August 2004 |pmid=15339343 |pmc=516768 |doi=10.1186/1471-2156-5-26 |doi-access=free }}</ref>


<ref name=pmid17194802>{{cite journal |last1=Gonder |first1=Mary K. |last2=Mortensen |first2=Holly M. |last3=Reed |first3=Floyd A. |last4=De Sousa |first4=Alexandra |last5=Tishkoff |first5=Sarah A. |date=March 2007 |title=Whole-mtDNA genome sequence analysis of ancient African lineages |journal=[[Molecular Biology and Evolution|Mol. Biol. Evol.]] |volume=24 |issue=3 |pp=757–68 |pmid=17194802 |doi=10.1093/molbev/msl209 }}</ref>
<ref name=pmid17194802>{{cite journal |vauthors=Gonder MK, Mortensen HM, Reed FA, de Sousa A, Tishkoff SA |title=Whole-mtDNA genome sequence analysis of ancient African lineages |journal=[[Molecular Biology and Evolution]] |volume=24 |issue=3 |pages=757–768 |date=March 2007 |pmid=17194802 |doi=10.1093/molbev/msl209 |doi-access=free}}</ref>


<ref name=pmid10739760>{{cite journal |last1=Chen |first1=Yu-Sheng |last2=Olckers |first2=Antonel |last3=Schurr |first3=Theodore G. |last4=Kogelnik |first4=Andreas M. |last5=Huoponen |first5=Kirsi |last6=Wallace |first6=Douglas C. |date=April 2000 |title=mtDNA variation in the South African Kung and Khwe-and their genetic relationships to other African populations |journal=Am. J. Hum. Genet. |volume=66 |issue=4 |pp=1362–83 |pmid=10739760 |pmc=1288201 |doi=10.1086/302848 }}</ref>
<ref name=pmid10739760>{{cite journal |vauthors=Chen YS, Olckers A, Schurr TG, Kogelnik AM, Huoponen K, Wallace DC |title=mtDNA variation in the South African Kung and Khwe-and their genetic relationships to other African populations |journal=[[American Journal of Human Genetics]] |volume=66 |issue=4 |pages=1362–1383 |date=April 2000 |pmid=10739760 |pmc=1288201 |doi=10.1086/302848}}</ref>


}}
}}


==Sources==
== Sources ==
{{refbegin}}
{{refbegin|32em}}
* {{cite journal |last=Appenzeller |first=Tim |year=2012 |title=Human migrations: Eastern odyssey. Humans had spread across Asia by 50,000 years ago. Everything else about our original exodus from Africa is up for debate. |journal=Nature |volume=485 |issue=7396 |url=http://www.nature.com/news/human-migrations-eastern-odyssey-1.10560 |ref=harv }}
* {{cite journal |vauthors=Appenzeller T |year=2012 |title=Human migrations: Eastern odyssey. Humans had spread across Asia by 50,000 years ago. Everything else about our original exodus from Africa is up for debate. |journal=[[Nature (journal)|Nature]] |volume=485 |issue=7396 |url=http://www.nature.com/news/human-migrations-eastern-odyssey-1.10560}}
* {{cite journal |last=Beyin |first=Amanuel |year=2011 |title=Upper Pleistocene Human Dispersals out of Africa: A Review of the Current State of the Debate |journal=International Journal of Evolutionary Biology |volume=2011 |issue=615094 |pp=1–17 |doi=10.4061/2011/615094 |ref=harv }}
* {{cite journal |vauthors=Beyin A |title=Upper Pleistocene Human Dispersals out of Africa: A Review of the Current State of the Debate |journal=[[International Journal of Evolutionary Biology]] |volume=2011 |issue=615094 |pages=615094 |year=2011 |pmid=21716744 |pmc=3119552 |doi=10.4061/2011/615094 |doi-access=free }}
* {{cite journal |last1=Endicott |first1=Phillip|last2=Gilbert |first2=M. Thomas P. |last3=Stringer |first3=Chris |last4=Lalueza-Fox |first4=Carles |last5=Willerslev |first5=Eske |last6=Hansen |first6=Anders J. |last7=Cooper |first7=Alan |date=January 2003 |title=The genetic origins of the Andaman Islanders |journal=[[American Journal of Human Genetics|AJHG]] |volume=72 |issue=1 |pp=178–84 |pmid=12478481 |pmc=378623 |doi=10.1086/345487 |ref=CITEREFEndicott et al.2003 }}
* {{cite journal |vauthors=Endicott P, Gilbert MT, Stringer C, Lalueza-Fox C, Willerslev E, Hansen AJ, Cooper A |title=The genetic origins of the Andaman Islanders |journal=[[American Journal of Human Genetics]] |volume=72 |issue=1 |pages=178–184 |date=January 2003 |pmid=12478481 |pmc=378623 |doi=10.1086/345487 |ref=CITEREFEndicott et al.2003}}
* {{cite book |last=Finlayson |first=Clive |year=2009 |title=The humans who went extinct: why Neanderthals died out and we survived |url=https://books.google.com/books?id=EzBV3OPb5mAC&pg=PA68 |publisher=Oxford University Press US |isbn=978-0-19-923918-4 |ref=harv }}
* {{cite book |vauthors=Finlayson C |year=2009 |title=The humans who went extinct: why Neanderthals died out and we survived |url=https://books.google.com/books?id=EzBV3OPb5mAC&pg=PA68 |publisher=[[Oxford University Press]] US |isbn=978-0199239184 |via=[[Google Books]]}}
* {{cite journal |last=Groucutt |first=Huw S. |year=2015 |title=Rethinking the dispersal of ''Homo sapiens'' out of Africa |journal=Evolutionary Anthropology |volume=24 |issue=4 - July/August |pp=149–164 |display-authors=etal |doi=10.1002/evan.21455 |ref=CITEREFGroucutt et al.2015 }}
* {{cite journal |vauthors=Groucutt HS, Petraglia MD, Bailey G, Scerri EM, Parton A, Clark-Balzan L, Jennings RP, Lewis L, Blinkhorn J, Drake NA, Breeze PS, Inglis RH, Devès MH, Meredith-Williams M, Boivin N, Thomas MG, Scally A |display-authors=6 |title=Rethinking the dispersal of Homo sapiens out of Africa |journal=[[Evolutionary Anthropology]] |volume=24 |issue=4 |pages=149–164 |year=2015 |pmid=26267436 |pmc=6715448 |doi=10.1002/evan.21455 |ref=CITEREFGroucutt et al.2015}}
* {{cite journal |last1=Harding |first1=Rosalind M. |last2=Healy |first2=Eugene |last3=Ray |first3=Amanda J. |last4=Ellis |first4=Nichola S. |last5=Flanagan |first5=Niamh |last6=Todd |first6=Carol |last7=Dixon |first7=Craig |last8=Sajantila |first8=Antti |last9=Jackson |first9=Ian J. |last10=Birch-Machin |first10=Mark A. |last11=Rees |first11=Jonathan L. |date=April 2000 |title=Evidence for variable selective pressures at MC1R |journal=[[American Journal of Human Genetics|AJHG]] |volume=66 |issue=4 |pp= 1351–61 |pmid=10733465 |pmc=1288200 |doi=10.1086/302863 |ref=CITEREFHarding et al.2000 }}
* {{cite journal |vauthors=Harding RM, Healy E, Ray AJ, Ellis NS, Flanagan N, Todd C, Dixon C, Sajantila A, Jackson IJ, Birch-Machin MA, Rees JL |display-authors=6 |title=Evidence for variable selective pressures at MC1R |journal=[[American Journal of Human Genetics]] |volume=66 |issue=4 |pages=1351–1361 |date=April 2000 |pmid=10733465 |pmc=1288200 |doi=10.1086/302863 |ref=CITEREFHarding et al.2000}}
* {{cite journal |last1=Hershkovitz |first1=Israel |year=2015 |last2=Marder |first2=Ofer |last3=Ayalon |first3=Avner |last4=Bar-Matthews |first4=Miryam |last5=Yasur |first5=Gal |last6=Boaretto |first6=Elisabetta |last7=Caracuta |first7=Valentina |last8=Alex |first8=Bridget |last9=Frumkin |first9=Amos |last10=Goder-Goldberger |first10=Mae |last11=Gunz |first11=Philipp |last12=Holloway |first12=Ralph L. |last13=Latimer |first13=Bruce |last14=Lavi |first14=Ron |last15=Matthews |first15=Alan |last16=Slon |first16=Viviane |last17=Mayer |first17=Daniella Bar-Yosef |last18=Berna |first18=Francesco |last19=Bar-Oz |first19=Guy |last20=Yeshurun |first20=Reuven |last21=May |first21=Hila |last22=Hans |first22=Mark G. |last23=Weber |first23=Gerhard W. |last24=Barzilai |first24=Omry |title=Levantine cranium from Manot Cave (Israel) foreshadows the first European modern humans |journal=Nature |volume=520 |doi=10.1038/nature14134 |pmid=25629628 |displayauthors=8 |pp=216–9 |ref=CITEREFHershkovitz et al.2015 }}
* {{cite journal |vauthors=Hershkovitz I, Marder O, Ayalon A, Bar-Matthews M, Yasur G, Boaretto E, Caracuta V, Alex B, Frumkin A, Goder-Goldberger M, Gunz P, Holloway RL, Latimer B, Lavi R, Matthews A, Slon V, Mayer DB, Berna F, Bar-Oz G, Yeshurun R, May H, Hans MG, Weber GW, Barzilai O |display-authors=6 |title=Levantine cranium from Manot Cave (Israel) foreshadows the first European modern humans |journal=[[Nature (journal)|Nature]] |volume=520 |issue=7546 |pages=216–219 |date=April 2015 |pmid=25629628 |doi=10.1038/nature14134 |ref=CITEREFHershkovitz et al.2015 |bibcode=2015Natur.520..216H |s2cid=4386123}}
* {{Cite journal |last1=Kuhlwilm |first1=Martin |year=2016 |title=Ancient gene flow from early modern humans into Eastern Neanderthals |journal=Nature |volume=530 |pp=429–33 |doi=10.1038/nature16544 |display-authors=etal |ref=CITEREFKuhlwilm et al.2016 |pmid=26886800 |pmc=4933530}}
* {{cite journal |vauthors=Kuhlwilm M, Gronau I, Hubisz MJ, de Filippo C, Prado-Martinez J, Kircher M, Fu Q, Burbano HA, Lalueza-Fox C, de la Rasilla M, Rosas A, Rudan P, Brajkovic D, Kucan Ž, Gušic I, Marques-Bonet T, Andrés AM, Viola B, Pääbo S, Meyer M, Siepel A, Castellano S |display-authors=6 |title=Ancient gene flow from early modern humans into Eastern Neanderthals |journal=[[Nature (journal)|Nature]] |volume=530 |issue=7591 |pages=429–433 |date=February 2016 |pmid=26886800 |pmc=4933530 |doi=10.1038/nature16544 |ref=CITEREFKuhlwilm et al.2016 |bibcode=2016Natur.530..429K}}
* {{cite journal |last1=Liu |first1=Hua |last2=Prugnolle |first2=Franck |last3=Manica |first3=Andrea |last4=Balloux |first4=François |date=August 2006 |title=A geographically explicit genetic model of worldwide human-settlement history |journal=[[The American Journal of Human Genetics|AJHG]] |volume=79 |issue=2 |pp=230–7 |pmid=16826514 |pmc=1559480 |doi=10.1086/505436 |ref=CITEREFLiu, Prugnolle et al.2006}}
* {{cite journal |vauthors=Liu H, Prugnolle F, Manica A, Balloux F |title=A geographically explicit genetic model of worldwide human-settlement history |journal=[[American Journal of Human Genetics]] |volume=79 |issue=2 |pages=230–237 |date=August 2006 |pmid=16826514 |pmc=1559480 |doi=10.1086/505436 |ref=CITEREFLiu, Prugnolle et al.2006}}
* {{cite journal |last1=Liu |first1=Wu |last2=Martinón-Torres |first2=María |last3=Cai |first3=Yan-jun |last4=Xing |first4=Song |last5=Tong |first5=Hao-wen |last6=Pei |first6=Shu-wen |last7=Sier |first7=Mark Jan |last8=Wu |first8=Xiao-hong |last9=Edwards |first9=R. Lawrence |year=2015 |title=The earliest unequivocally modern humans in southern China |journal=Nature |doi=10.1038/nature15696 |volume=526 |pp=696–9 |url=http://www.nature.com/doifinder/10.1038/nature15696 |ref=CITEREFLiu, Martinón-Torres et al.2015 |pmid=26466566}}
* {{cite journal |vauthors=Liu W, Martinón-Torres M, Cai YJ, Xing S, Tong HW, Pei SW, Sier MJ, Wu XH, Edwards RL, Cheng H, Li YY, Yang XX, de Castro JM, Wu XJ |display-authors=6 |title=The earliest unequivocally modern humans in southern China |journal=[[Nature (journal)|Nature]] |volume=526 |issue=7575 |pages=696–699 |date=October 2015 |pmid=26466566 |doi=10.1038/nature15696 |url=http://discovery.ucl.ac.uk/1472397/1/Nature%20merged%20file.pdf |ref=CITEREFLiu, Martinón-Torres et al.2015 |bibcode=2015Natur.526..696L |s2cid=205246146}}
* {{cite journal |last1=Macaulay |first1=Vincent |last2=Hill |first2=Catherine |last3=Achilli |first3=Alessandro |last4=Rengo |first4=Chiara |last5=Clarke |first5=Douglas |last6=Meehan |first6=William |last7=Blackburn |first7=James |last8=Semino |first8=Ornella |last9=Scozzari |first9=Rosaria |last10=Cruciani |first10=Fulvio |last11=Taha |first11=Adi |last12=Shaari |first12=Norazila Kassim |last13=Raja |first13=Joseph Maripa |last14=Ismail |first14=Patimah |last15=Zainuddin |first15=Zafarina |last16=Goodwin |first16=William |last17=Bulbeck |first17=David |last18=Bandelt |first18=Hans-Jürgen |last19=Oppenheimer |first19=Stephen |last20=Torroni |first20=Antonio |last21=Richards |first21=Martin |date=13 May 2005 |title=Single, Rapid Coastal Settlement of Asia Revealed by Analysis of Complete Mitochondrial Genomes |journal=Science |volume=308 |issue=5724 |pp=1034–6 |doi=10.1126/science.1109792 |pmid=15890885 |ref=CITEREFMacaulay et al.2005 }}
* {{cite journal |vauthors=Macaulay V, Hill C, Achilli A, Rengo C, Clarke D, Meehan W, Blackburn J, Semino O, Scozzari R, Cruciani F, Taha A, Shaari NK, Raja JM, Ismail P, Zainuddin Z, Goodwin W, Bulbeck D, Bandelt HJ, Oppenheimer S, Torroni A, Richards M |display-authors=6 |title=Single, rapid coastal settlement of Asia revealed by analysis of complete mitochondrial genomes |journal=[[Science (journal)|Science]] |volume=308 |issue=5724 |pages=1034–1036 |date=May 2005 |pmid=15890885 |doi=10.1126/science.1109792 |ref=CITEREFMacaulay et al.2005 |bibcode=2005Sci...308.1034M |s2cid=31243109 |url=http://psasir.upm.edu.my/id/eprint/40255/1/Single%2C%20rapid%20coastal%20settlement%20of%20Asia%20revealed%20by%20analysis%20of%20complete%20mitochondrial%20genomes.pdf}}
* {{cite journal |vauthors=McChesney KY |year=2015 |title=Teaching Diversity. The Science You Need to Know to Explain Why Race Is Not Biological |journal=SAGE Open |doi=10.1177/2158244015611712 |doi-access=free |volume=5 |issue=4}}
* {{cite book |last=Meredith |first=Martin |authorlink=Martin Meredith |year=2011 |title=Born in Africa: The Quest for the Origins of Human Life |publisher=PublicAffairs |location=New York City |isbn=1-58648-663-2 |url=https://books.google.com/books?id=WrR9OShae2wC&pg=PT148 |ref=harv }}
* {{cite book |vauthors=Meredith M |author-link=Martin Meredith |year=2011 |title=Born in Africa: The Quest for the Origins of Human Life |publisher=PublicAffairs |location=New York |isbn=978-1586486631 |url=https://books.google.com/books?id=WrR9OShae2wC&pg=PT148 |via=[[Google Books]]}}
* {{cite journal |last1=Posth |first1=Cosimo |last2=Renaud |first2=Gabriel |last3=Mittnik |first3=Alissa |last4=Drucker |first4=Dorothée G. |year=2016 |title=Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-Africans and a Late Glacial Population Turnover in Europe |journal=Current Biology |display-authors=etal |ref=CITEREFPosth et al.2016 |url=http://www.cell.com/current-biology/abstract/S0960-9822%2816%2900087-7?_returnURL=http%3A%2F%2Flinkinghub.elsevier.com%2Fretrieve%2Fpii%2FS0960982216000877%3Fshowall%3Dtrue |doi=10.1016/j.cub.2016.01.037 |volume=26 |pages=827–833 |pmid=26853362}}
* {{cite journal |vauthors=Posth C, Renaud G, Mittnik A, Drucker DG, Rougier H, Cupillard C, Valentin F, Thevenet C, Furtwängler A, Wißing C, Francken M, Malina M, Bolus M, Lari M, Gigli E, Capecchi G, Crevecoeur I, Beauval C, Flas D, Germonpré M, van der Plicht J, Cottiaux R, Gély B, Ronchitelli A, Wehrberger K, Grigorescu D, Svoboda J, Semal P, Caramelli D, Bocherens H, Harvati K, Conard NJ, Haak W, Powell A, Krause J |display-authors=6 |title=Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-Africans and a Late Glacial Population Turnover in Europe |journal=[[Current Biology]] |volume=26 |issue=6 |pages=827–833 |date=March 2016 |pmid=26853362 |doi=10.1016/j.cub.2016.01.037 |bibcode=2016CBio...26..827P |hdl=2440/114930 |s2cid=140098861 |ref=CITEREFPosth et al.2016 |hdl-access=free}}
* {{cite journal |last1=Shackelton |first1=Laura A. |last2=Rambaut |first2=Andrew |last3=Pybus |first3=Oliver G. |last4=Holmes |first4=Edward C. |year=2006 |title=JC Virus Evolution and Its Association with Human Populations |journal=Journal of Virology |volume=80 |pp=9928–33 |doi=10.1128/JVI.00441-06 |pmid=17005670 |pmc=1617318 |ref=CITEREFShackelton et al.2006 }}
* {{cite journal |last1=Shen |first1=Guanjun |last2=Wang |first2=Wei |last3=Wang |first3=Qian |last4=Zhao |first4=Jianxin |last5=Collerson |first5=Kenneth |last6=Zhou |first6=Chunlin |last7=Tobias |first7=Phillip V. |year=2002 |title=U-Series dating of Liujiang hominid site in Guangxi, Southern China |journal=[[Journal of Human Evolution|J. Hum. Evol.]] |volume=43 |issue=6 |pages=817–29 |pmid=12473485 |doi=10.1006/jhev.2002.0601 |ref=CITEREFShen, Wang et al.2002 }}
* {{cite journal |vauthors=Shackelton LA, Rambaut A, Pybus OG, Holmes EC |title=JC virus evolution and its association with human populations |journal=[[Journal of Virology]] |volume=80 |issue=20 |pages=9928–9933 |date=October 2006 |pmid=17005670 |pmc=1617318 |doi=10.1128/JVI.00441-06 |ref=CITEREFShackelton et al.2006}}
* {{cite journal |vauthors=Shen G, Wang W, Wang Q, Zhao J, Collerson K, Zhou C, Tobias PV |title=U-Series dating of Liujiang hominid site in Guangxi, Southern China |journal=[[Journal of Human Evolution]] |volume=43 |issue=6 |pages=817–829 |date=December 2002 |pmid=12473485 |doi=10.1006/jhev.2002.0601 |bibcode=2002JHumE..43..817S |ref=CITEREFShen, Wang et al.2002}}
* {{Citation | last =Young McChesney | first =Kai | year =2015 | title =Teaching Diversity. The Science You Need to Know to Explain Why Race Is Not Biological | journal =SAGE open | doi =10.1177/2158244015611712 | url =http://sgo.sagepub.com/content/5/4/2158244015611712 | volume=5 | pages=2158244015611712}}
* {{cite book |last1=Wells |first1=Spencer |authorlink=Spencer Wells |year=2003 |orig-year=2002 |title=[[The Journey of Man|The Journey of Man: A Genetic Odyssey]] |publisher=Random House Trade Paperbacks|location=New York|isbn=0-8129-7146-9 |ref=harv }}
* {{cite book |vauthors=Wells S |author-link=Spencer Wells |year=2003 |orig-date=2002 |title=The Journey of Man: A Genetic Odyssey |publisher=Random House Trade Paperbacks |location=New York |isbn=978-0812971460 |title-link=The Journey of Man}}
{{refend}}
{{refend}}


'''Further reading'''
== Further reading ==
{{refbegin}}
{{refbegin| 30em}}
* {{cite book | author = Stringer, Chris | title = The Origin of Our Species | edition = | publisher = Allen Lane | location = London | year = 2011 | pages = | isbn = 978-1-84614-140-9 | doi = | url = | accessdate = }}
* {{cite book |vauthors=Stringer C |title=The Origin of Our Species |publisher=Allen Lane |location=London |year=2011 |isbn=978-1846141409}}
* {{cite book | author = Wells, Spencer | title = Deep ancestry: inside the Genographic Project | edition = | publisher = National Geographic | location = Washington, D.C | year = 2006 | pages = | isbn = 0-7922-6215-8 | doi = | url = | accessdate = }}
* {{cite book |vauthors=Wells S |title=Deep ancestry: inside the Genographic Project |publisher=[[National Geographic]] |location=Washington, D.C. |year=2006 |isbn=978-0792262152}}
* {{cite book | author = Wade, N. | year = 2006 | title = Before the Dawn : Recovering the Lost History of Our Ancestors | edition = | publisher = Penguin Press HC, The | isbn = 1-59420-079-3}}
* {{cite book |vauthors=Wade N |year=2006 |title=Before the Dawn: Recovering the Lost History of Our Ancestors |publisher=Penguin Press|isbn=978-1594200793}}
* {{cite book | author = Sykes, Bryan | year = 2004| title = [[The Seven Daughters of Eve]]: The Science That Reveals Our Genetic Ancestry | edition = | publisher = Corgi Adult | pages = | isbn = 0-552-15218-8 | doi = | url = | accessdate = }}
* {{cite book |vauthors=Sykes B |year=2004 |title=The Seven Daughters of Eve: The Science That Reveals Our Genetic Ancestry |publisher=Corgi Adult |isbn=978-0552152181 |title-link=The Seven Daughters of Eve}}
{{refend}}
{{refend}}


==External links==
== External links ==
* Encyclopædia Britannica, ''[http://www.britannica.com/EBchecked/topic/275670/human-evolution Human Evolution]''
* Encyclopædia Britannica, ''[http://www.britannica.com/EBchecked/topic/275670/human-evolution Human Evolution]''
* [http://humanorigins.si.edu/evidence/human-evolution-timeline-interactive Human Timeline (Interactive)] – [[Smithsonian Institution|Smithsonian]], [[National Museum of Natural History]] (August 2016).
* [http://humanorigins.si.edu/evidence/human-evolution-timeline-interactive Human Timeline (Interactive)] – [[Smithsonian Institution|Smithsonian]], [[National Museum of Natural History]] (August 2016).
* [https://www.nature.com/articles/s41598-021-98618-0 Age constraints for the Trachilos footprints from Crete] [[Nature (journal)|Nature]] (October 2021).


{{Early human migrations}}
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{{Human Evolution}}
{{Human Evolution}}
{{Human genetics}}
{{Human genetics}}
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{{DEFAULTSORT:Recent African Origin Of Modern Humans}}
[[Category:Recent African origin of modern humans| ]]
[[Category:Recent African origin of modern humans| ]]
[[Category:Prehistoric migrations]]
[[Category:Prehistoric migrations]]

Latest revision as of 05:22, 20 December 2024

Successive dispersals (labeled in years before present) of
  Homo erectus greatest extent (yellow)
  Homo neanderthalensis greatest extent (ochre)
  Homo sapiens (red)
Expansion of early modern humans from Africa through the Near East

In paleoanthropology, the recent African origin of modern humans or the "Out of Africa" theory (OOA)[a] is the most widely accepted[1][2][3] model of the geographic origin and early migration of anatomically modern humans (Homo sapiens). It follows the early expansions of hominins out of Africa, accomplished by Homo erectus and then Homo neanderthalensis.

The model proposes a "single origin" of Homo sapiens in the taxonomic sense, precluding parallel evolution in other regions of traits considered anatomically modern,[4] but not precluding multiple admixture between H. sapiens and archaic humans in Europe and Asia.[b][5][6] H. sapiens most likely developed in the Horn of Africa between 300,000 and 200,000 years ago,[7][8] although an alternative hypothesis argues that diverse morphological features of H. sapiens appeared locally in different parts of Africa and converged due to gene flow between different populations within the same period.[9][10] The "recent African origin" model proposes that all modern non-African populations are substantially descended from populations of H. sapiens that left Africa after that time.

There were at least several "out-of-Africa" dispersals of modern humans, possibly beginning as early as 270,000 years ago, including 215,000 years ago to at least Greece,[11][12][13] and certainly via northern Africa and the Arabian Peninsula about 130,000 to 115,000 years ago.[20] There is evidence that modern humans had reached China around 80,000 years ago.[21] Practically all of these early waves seem to have gone extinct or retreated back, and present-day humans outside Africa descend mainly from a single expansion about 70,000–50,000 years ago,[22][23][24][7][8][25][26][excessive citations] via the so-called "Southern Route". These humans spread rapidly along the coast of Asia and reached Australia by around 65,000–50,000 years ago,[27][28][c] (though some researchers question the earlier Australian dates and place the arrival of humans there at 50,000 years ago at earliest,[29][30] while others have suggested that these first settlers of Australia may represent an older wave before the more significant out of Africa migration and thus not necessarily be ancestral to the region's later inhabitants[24]) while Europe was populated by an early offshoot which settled the Near East and Europe less than 55,000 years ago.[31][32][33]

In the 2010s, studies in population genetics uncovered evidence of interbreeding that occurred between H. sapiens and archaic humans in Eurasia, Oceania and Africa,[34][35][36] indicating that modern population groups, while mostly derived from early H. sapiens, are to a lesser extent also descended from regional variants of archaic humans.

Proposed waves

[edit]
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",[37] and 42,400–41,700 BP for "Ethelruda".[37]

"Recent African origin", or Out of Africa II, refers to the migration of anatomically modern humans (Homo sapiens) out of Africa after their emergence at c. 300,000 to 200,000 years ago, in contrast to "Out of Africa I", which refers to the migration of archaic humans from Africa to Eurasia from before 1.8 and up to 0.5 million years ago. Omo-Kibish I (Omo I) from southern Ethiopia is the oldest anatomically modern Homo sapiens skeleton currently known (around 233,000 years old).[38] There are even older Homo sapiens fossils from Jebel Irhoud in Morocco which exhibit a mixture of modern and archaic features at around 315,000 years old.[39]

Since the beginning of the 21st century, the picture of "recent single-origin" migrations has become significantly more complex, due to the discovery of modern-archaic admixture and the increasing evidence that the "recent out-of-Africa" migration took place in waves over a long time. As of 2010, there were two main accepted dispersal routes for the out-of-Africa migration of early anatomically modern humans, the "Northern Route" (via Nile Valley and Sinai) and the "Southern Route" via the Bab-el-Mandeb strait.[40]

  • Posth et al. (2017) suggest that early Homo sapiens, or "another species in Africa closely related to us", might have first migrated out of Africa around 270,000 years ago based on the closer affinity within Neanderthals' mitochondrial genomes to Homo sapiens than Denisovans.[41]
  • Fossil evidence also points to an early Homo sapiens migration with the oldest known fossil coming from Apidima Cave in Greece and dated at 210,000 years ago.[42] Finds at Misliya cave, which include a partial jawbone with eight teeth, have been dated to around 185,000 years ago. Layers dating from between 250,000 and 140,000 years ago in the same cave contained tools of the Levallois type which could put the date of the first migration even earlier if the tools can be associated with the modern human jawbone finds.[43][44][45]
  • An eastward dispersal from Northeast Africa to Arabia 150,000–130,000 years ago is based on the stone tools finds at Jebel Faya dated to 127,000 years ago (discovered in 2011), although fossil evidence in the area is significantly later at 85,000 years ago.[14] [46] Possibly related to this wave are the finds from Zhirendong cave, Southern China, dated to more than 100,000 years ago.[47] Other evidence of modern human presence in China has been dated to 80,000 years ago.[21]
  • The most significant out of Africa dispersal took place around 50,000–70,000 years ago via the so-called Southern Route, either before[48] or after[32][33] the Toba event, which happened between 69,000 and 77,000 years ago.[48] This dispersal followed the southern coastline of Asia and reached Australia around 65,000–50,000 years ago or according to some research, by 50,000 years ago at earliest.[29][30] Western Asia was "re-occupied" by a different derivation from this wave around 50,000 years ago and Europe was populated from Western Asia beginning around 43,000 years ago.[40]
  • Wells (2003) describes an additional wave of migration after the southern coastal route, a northern migration into Europe about 45,000 years ago.[d] This possibility is ruled out by Macaulay et al. (2005) and Posth et al. (2016), who argue for a single coastal dispersal, with an early offshoot into Europe.

Northern Route dispersal

[edit]
Anatomically Modern Humans known archaeological remains in Europe and Africa, directly dated, calibrated carbon dates as of 2013.[37]

Beginning 135,000 years ago, tropical Africa experienced megadroughts which drove humans from the land and towards the sea shores, and forced them to cross over to other continents.[49][e]

Fossils of early Homo sapiens were found in Qafzeh and Es-Skhul Caves in Israel and have been dated to 80,000 to 120,000 years ago.[50][51] These humans seem to have either become extinct or retreated back to Africa 70,000 to 80,000 years ago, possibly replaced by southbound Neanderthals escaping the colder regions of ice-age Europe.[22] Hua Liu et al. analyzed autosomal microsatellite markers dating to about 56,000 years ago. They interpret the paleontological fossil as an isolated early offshoot that retracted back to Africa.[23]

The discovery of stone tools in the United Arab Emirates in 2011 at the Faya-1 site in Mleiha, Sharjah, indicated the presence of modern humans at least 125,000 years ago,[14] leading to a resurgence of the "long-neglected" North African route.[15][52][16][17] This new understanding of the role of the Arabian dispersal began to change following results from archaeological and genetic studies stressing the importance of southern Arabia as a corridor for human expansions out of Africa.[53]

In Oman, a site was discovered by Bien Joven in 2011 containing more than 100 surface scatters of stone tools belonging to the late Nubian Complex, known previously only from archaeological excavations in the Sudan. Two optically stimulated luminescence age estimates placed the Arabian Nubian Complex at approximately 106,000 years old. This provides evidence for a distinct Stone Age technocomplex in southern Arabia, around the earlier part of the Marine Isotope Stage 5.[54]

According to Kuhlwilm and his co-authors, Neanderthals contributed genetically to modern humans then living outside of Africa around 100,000 years ago: humans which had already split off from other modern humans around 200,000 years ago, and this early wave of modern humans outside Africa also contributed genetically to the Altai Neanderthals.[55] They found that "the ancestors of Neanderthals from the Altai Mountains and early modern humans met and interbred, possibly in the Near East, many thousands of years earlier than previously thought".[55] According to co-author Ilan Gronau, "This actually complements archaeological evidence of the presence of early modern humans out of Africa around and before 100,000 years ago by providing the first genetic evidence of such populations."[55] Similar genetic admixture events have been noted in other regions as well.[56]

Southern Route dispersal

[edit]

Coastal route

[edit]
Red Sea crossing

By some 50–70,000 years ago, a subset of the bearers of mitochondrial haplogroup L3 migrated from East Africa into the Near East. It has been estimated that from a population of 2,000 to 5,000 individuals in Africa, only a small group, possibly as few as 150 to 1,000 people, crossed the Red Sea.[57][58] The group that crossed the Red Sea travelled along the coastal route around Arabia and the Persian Plateau to India, which appears to have been the first major settling point.[59] Wells (2003) argued for the route along the southern coastline of Asia, across about 250 kilometres (155 mi), reaching Australia by around 50,000 years ago.

Migration routes of modern humans, showing the northern route populating Western Eurasia, and the southern/coastal route populating Eastern Eurasia.

Today at the Bab-el-Mandeb straits, the Red Sea is about 20 kilometres (12 mi) wide, but 50,000 years ago sea levels were 70 m (230 ft) lower (owing to glaciation) and the water channel was much narrower. Though the straits were never completely closed, they were narrow enough to have enabled crossing using simple rafts, and there may have been islands in between.[40][60] Shell middens 125,000 years old have been found in Eritrea,[61] indicating that the diet of early humans included seafood obtained by beachcombing.

Toba eruption

[edit]

The dating of the Southern Dispersal is a matter of dispute.[48] It may have happened either pre- or post-Toba, a catastrophic volcanic eruption that took place between 69,000 and 77,000 years ago at the site of present-day Lake Toba in Sumatra, Indonesia. Stone tools discovered below the layers of ash deposited in India may point to a pre-Toba dispersal but the source of the tools is disputed.[48] An indication for post-Toba is haplo-group L3, that originated before the dispersal of humans out of Africa and can be dated to 60,000–70,000 years ago, "suggesting that humanity left Africa a few thousand years after Toba".[48] Some research showing slower than expected genetic mutations in human DNA was published in 2012, indicating a revised dating for the migration to between 90,000 and 130,000 years ago.[62] Some more recent research suggests a migration out-of-Africa of around 50,000-65,000 years ago of the ancestors of modern non-African populations, similar to most previous estimates.[24][63][64]

Western Asia

[edit]

Following the fossils dating 80,000 to 120,000 years ago from Qafzeh and Es-Skhul Caves in Israel there are no H. sapiens fossils in the Levant until the Manot 1 fossil from Manot Cave in Israel, dated to 54,700 years ago,[65] though the dating was questioned by Groucutt et al. (2015). The lack of fossils and stone tool industries that can be safely associated with modern humans in the Levant has been taken to suggest that modern humans were outcompeted by Neanderthals until around 55,000 years ago, who would have placed a barrier on modern human dispersal out of Africa through the Northern Route.[66][failed verification] Climate reconstructions also support a Southern Route dispersal of modern humans as the Bab-el-Mandeb strait experienced a climate more conductive to human migration than the northern landbridge to the Levant during the major human dispersal out of Africa.[67]

A 2023 study proposed that Eurasians and Africans genetically diverged ~100,000 years ago. Main Eurasians then lived in the Saudi Peninsula, genetically isolated from at least 85 kya, before expanding north 54 kya. For reference, Homo sapiens and Neanderthals diverged ~500 kya.[68]

Oceania

[edit]

It is thought that Australia was inhabited around 65,000–50,000 years ago. As of 2017, the earliest evidence of humans in Australia is at least 65,000 years old,[27][28] while McChesney stated that

...genetic evidence suggests that a small band with the marker M168 migrated out of Africa along the coasts of the Arabian Peninsula and India, through Indonesia, and reached Australia very early, between 60,000 and 50,000 years ago. This very early migration into Australia is also supported by Rasmussen et al. (2011).[31]

Fossils from Lake Mungo, Australia, have been dated to about 42,000 years ago.[69][70] Other fossils from a site called Madjedbebe have been dated to at least 65,000 years ago,[71][72] though some researchers doubt this early estimate and date the Madjedbebe fossils at about 50,000 years ago at the oldest.[29][30]

Phylogenetic data suggests that an early Eastern Eurasian (Eastern non-African) meta-population trifurcated somewhere in eastern South Asia, and gave rise to the Australo-Papuans, the Ancient Ancestral South Indians (AASI), as well as East/Southeast Asians, although Papuans may have also received some gene flow from an earlier group (xOoA), around 2%,[73] next to additional archaic admixture in the Sahul region.[74][75]

According to one study, Papuans could have either formed from a mixture between an East Eurasian lineage and lineage basal to West and East Asians, or as a sister lineage of East Asians with or without a minor basal OoA or xOoA contribution.[76]

A Holocene hunter-gatherer sample (Leang_Panninge) from South Sulawesi was found to be genetically in between East-Eurasians and Australo-Papuans. The sample could be modeled as ~50% Papuan-related and ~50% Basal-East Asian-related (Andamanese Onge or Tianyuan). The authors concluded that Basal-East Asian ancestry was far more widespread and the peopling of Insular Southeast Asia and Oceania was more complex than previously anticipated.[77][78]

PCA calculated on present-day and ancient individuals from eastern Eurasia and Oceania. PC1 (23,8%) distinguishes East-Eurasians and Australo-Melanesians, while PC2 (6,3%) differentiates East-Eurasians along a North to South cline.
Principal component analysis (PCA) of ancient and modern day individuals from worldwide populations. Oceanians (Aboriginal Australians and Papuans) are most differentiated from both East-Eurasians and West-Eurasians.

East and Southeast Asia

[edit]

In China, the Liujiang man (Chinese: 柳江人) is among the earliest modern humans found in East Asia.[79] The date most commonly attributed to the remains is 67,000 years ago.[80] High rates of variability yielded by various dating techniques carried out by different researchers place the most widely accepted range of dates with 67,000 BP as a minimum, but do not rule out dates as old as 159,000 BP.[80] Liu, Martinón-Torres et al. (2015) claim that modern human teeth have been found in China dating to at least 80,000 years ago.[81]

Tianyuan man from China has a probable date range between 38,000 and 42,000 years ago, while Liujiang man from the same region has a probable date range between 67,000 and 159,000 years ago. According to 2013 DNA tests, Tianyuan man is related "to many present-day Asians and Native Americans".[82][83][84][85][86] Tianyuan is similar in morphology to Liujiang man, and some Jōmon period modern humans found in Japan, as well as modern East and Southeast Asians.[87][88][89]

A 2021 study about the population history of Eastern Eurasia, concluded that distinctive Basal-East Asian (East-Eurasian) ancestry originated in Mainland Southeast Asia at ~50,000BC from a distinct southern Himalayan route, and expanded through multiple migration waves southwards and northwards respectively.[90]

Genetic studies concluded that Native Americans descended from a single founding population that initially split from a Basal-East Asian source population in Mainland Southeast Asia around 36,000 years ago, at the same time at which the proper Jōmon people split from Basal-East Asians, either together with Ancestral Native Americans or during a separate expansion wave. They also show that the basal northern and southern Native American branches, to which all other Indigenous peoples belong, diverged around 16,000 years ago.[91][92] An indigenous American sample from 16,000BC in Idaho, which is craniometrically similar to modern Native Americans as well as Paleosiberias, was found to have largely East-Eurasian ancestry and showed high affinity with contemporary East Asians, as well as Jōmon period samples of Japan, confirming that Ancestral Native Americans split from an East-Eurasian source population in Eastern Siberia.[93]

Europe

[edit]

According to Macaulay et al. (2005), an early offshoot from the southern dispersal with haplogroup N followed the Nile from East Africa, heading northwards and crossing into Asia through the Sinai. This group then branched, some moving into Europe and others heading east into Asia.[32] This hypothesis is supported by the relatively late date of the arrival of modern humans in Europe as well as by archaeological and DNA evidence.[32] Based on an analysis of 55 human mitochondrial genomes (mtDNAs) of hunter-gatherers, Posth et al. (2016) argue for a "rapid single dispersal of all non-Africans less than 55,000 years ago". By 45,000 years ago, modern humans are known to have reached northwestern Europe.[94]

Genetic reconstruction

[edit]

Mitochondrial haplogroups

[edit]

Within Africa

[edit]
Map of early diversification of modern humans according to mitochondrial population genetics (see: Haplogroup L).

The first lineage to branch off from Mitochondrial Eve was L0. This haplogroup is found in high proportions among the San of Southern Africa and the Sandawe of East Africa. It is also found among the Mbuti people.[95][96] These groups branched off early in human history and have remained relatively genetically isolated since then. Haplogroups L1, L2, and L3 are descendants of L1–L6, and are largely confined to Africa. The macro haplogroups M and N, which are the lineages of the rest of the world outside Africa, descend from L3. L3 is about 70,000 years old, while haplogroups M and N are about 65–55,000 years old.[97][64] The relationship between such gene trees and demographic history is still debated when applied to dispersals.[98]

Of all the lineages present in Africa, the female descendants of only one lineage, mtDNA haplogroup L3, are found outside Africa. If there had been several migrations, one would expect descendants of more than one lineage to be found. L3's female descendants, the M and N haplogroup lineages, are found in very low frequencies in Africa (although haplogroup M1 populations are very ancient and diversified in North and North-east Africa) and appear to be more recent arrivals.[citation needed] A possible explanation is that these mutations occurred in East Africa shortly before the exodus and became the dominant haplogroups thereafter by means of the founder effect. Alternatively, the mutations may have arisen shortly afterwards.

Southern Route and haplogroups M and N

[edit]

Results from mtDNA collected from aboriginal Malaysians called Orang Asli indicate that the haplogroups M and N share characteristics with original African groups from approximately 85,000 years ago, and share characteristics with sub-haplogroups found in coastal south-east Asian regions, such as Australasia, the Indian subcontinent and throughout continental Asia, which had dispersed and separated from their African progenitor approximately 65,000 years ago. This southern coastal dispersal would have occurred before the dispersal through the Levant approximately 45,000 years ago.[32] This hypothesis attempts to explain why haplogroup N is predominant in Europe and why haplogroup M is absent in Europe. Evidence of the coastal migration is thought to have been destroyed by the rise in sea levels during the Holocene epoch.[99] Alternatively, a small European founder population that had expressed haplogroup M and N at first, could have lost haplogroup M through random genetic drift resulting from a bottleneck (i.e. a founder effect).

The group that crossed the Red Sea travelled along the coastal route around Arabia and Persia until reaching India.[59] Haplogroup M is found in high frequencies along the southern coastal regions of Pakistan and India and it has the greatest diversity in India, indicating that it is here where the mutation may have occurred.[59] Sixty percent of the Indian population belong to Haplogroup M. The indigenous people of the Andaman Islands also belong to the M lineage. The Andamanese are thought to be offshoots of some of the earliest inhabitants in Asia because of their long isolation from the mainland. They are evidence of the coastal route of early settlers that extends from India to Thailand and Indonesia all the way to eastern New Guinea. Since M is found in high frequencies in highlanders from New Guinea and the Andamanese and New Guineans have dark skin and Afro-textured hair, some scientists think they are all part of the same wave of migrants who departed across the Red Sea ~60,000 years ago in the Great Coastal Migration. The proportion of haplogroup M increases eastwards from Arabia to India; in eastern India, M outnumbers N by a ratio of 3:1. Crossing into Southeast Asia, haplogroup N (mostly in the form of derivatives of its R subclade) reappears as the predominant lineage.[citation needed] M is predominant in East Asia, but amongst Indigenous Australians, N is the more common lineage.[citation needed] This haphazard distribution of Haplogroup N from Europe to Australia can be explained by founder effects and population bottlenecks.[100]

The earliest-branching non-African paternal lineages (C, D, F) after the Out-of-Africa event (a), and their deepest divergence among modern day East or Southeast Asia (b), suggesting rapid coastal expansions. Simplified Y-chromosome tree is shown as reference for colours.[101]

Autosomal DNA

[edit]

A 2002 study of African, European, and Asian populations, found greater genetic diversity among Africans than among Eurasians, and that genetic diversity among Eurasians is largely a subset of that among Africans, supporting the out of Africa model.[102] A large study by Coop et al. (2009) found evidence for natural selection in autosomal DNA outside of Africa. The study distinguishes non-African sweeps (notably KITLG variants associated with skin color), West-Eurasian sweeps (SLC24A5) and East-Asian sweeps (MC1R, relevant to skin color). Based on this evidence, the study concluded that human populations encountered novel selective pressures as they expanded out of Africa.[103] MC1R and its relation to skin color had already been discussed by Harding et al. (2000), p. 1355. According to this study, Papua New Guineans continued to be exposed to selection for dark skin color so that, although these groups are distinct from Africans in other places, the allele for dark skin color shared by contemporary Africans, Andamanese and New Guineans is an archaism. Endicott et al. (2003) suggest convergent evolution. A 2014 study by Gurdasani et al. indicates that the higher genetic diversity in Africa was further increased in some regions by relatively recent Eurasian migrations affecting parts of Africa.[104]

Pathogen DNA

[edit]

Another promising route towards reconstructing human genetic genealogy is via the JC virus (JCV), a type of human polyomavirus which is carried by 70–90 percent of humans and which is usually transmitted vertically, from parents to offspring, suggesting codivergence with human populations. For this reason, JCV has been used as a genetic marker for human evolution and migration.[105] This method does not appear to be reliable for the migration out of Africa; in contrast to human genetics, JCV strains associated with African populations are not basal. From this Shackelton et al. (2006) conclude that either a basal African strain of JCV has become extinct or that the original infection with JCV post-dates the migration from Africa.

Admixture of archaic and modern humans

[edit]

Evidence for archaic human species (descended from Homo heidelbergensis) having interbred with modern humans outside of Africa, was discovered in the 2010s. This concerns primarily Neanderthal admixture in all modern populations except for Sub-Saharan Africans but evidence has also been presented for Denisova hominin admixture in Australasia (i.e. in Melanesians, Aboriginal Australians and some Negritos).[106] The rate of Neanderthal admixture to European and Asian populations as of 2017 has been estimated at between about 2–3%.[107]

Archaic admixture in some Sub-Saharan African populations hunter-gatherer groups (Biaka Pygmies and San), derived from archaic hominins that broke away from the modern human lineage around 700,000 years ago, was discovered in 2011. The rate of admixture was estimated at 2%.[36] Admixture from archaic hominins of still earlier divergence times, estimated at 1.2 to 1.3 million years ago, was found in Pygmies, Hadza and five Sandawe in 2012.[108][35]

From an analysis of Mucin 7, a highly divergent haplotype that has an estimated coalescence time with other variants around 4.5 million years BP and is specific to African populations, it is inferred to have been derived from interbreeding between African modern and archaic humans.[109]

A study published in 2020 found that the Yoruba and Mende populations of West Africa derive between 2% and 19% of their genome from an as-yet unidentified archaic hominin population that likely diverged before the split of modern humans and the ancestors of Neanderthals and Denisovans.[110]

Stone tools

[edit]

In addition to genetic analysis, Petraglia et al. also examines the small stone tools (microlithic materials) from the Indian subcontinent and explains the expansion of population based on the reconstruction of paleoenvironment. He proposed that the stone tools could be dated to 35 ka in South Asia, and the new technology might be influenced by environmental change and population pressure.[111]

History of the theory

[edit]

Classical paleoanthropology

[edit]
The frontispiece to Huxley's Evidence as to Man's Place in Nature (1863): the image compares the skeleton of a human to other apes.
The possibility of an origin of L3 in Asia was proposed by Cabrera et al. (2018).[112]
a: Exit of the L3 precursor to Eurasia. b: Return to Africa and expansion to Asia of basal L3 lineages with subsequent differentiation in both continents.

The cladistic relationship of humans with the African apes was suggested by Charles Darwin after studying the behaviour of African apes, one of which was displayed at the London Zoo.[113] The anatomist Thomas Huxley had also supported the hypothesis and suggested that African apes have a close evolutionary relationship with humans.[114] These views were opposed by the German biologist Ernst Haeckel, who was a proponent of the Out of Asia theory. Haeckel argued that humans were more closely related to the primates of South-east Asia and rejected Darwin's African hypothesis.[115][116]

In The Descent of Man, Darwin speculated that humans had descended from apes, which still had small brains but walked upright, freeing their hands for uses which favoured intelligence; he thought such apes were African:

In each great region of the world the living mammals are closely related to the extinct species of the same region. It is, therefore, probable that Africa was formerly inhabited by extinct apes closely allied to the gorilla and chimpanzee; and as these two species are now man's nearest allies, it is somewhat more probable that our early progenitors lived on the African continent than elsewhere. But it is useless to speculate on this subject, for an ape nearly as large as a man, namely the Dryopithecus of Lartet, which was closely allied to the anthropomorphous Hylobates, existed in Europe during the Upper Miocene period; and since so remote a period the earth has certainly undergone many great revolutions, and there has been ample time for migration on the largest scale.

— Charles Darwin, Descent of Man[117]

In 1871, there were hardly any human fossils of ancient hominins available. Almost fifty years later, Darwin's speculation was supported when anthropologists began finding fossils of ancient small-brained hominins in several areas of Africa (list of hominina fossils). The hypothesis of recent (as opposed to archaic) African origin developed in the 20th century. The "Recent African origin" of modern humans means "single origin" (monogenism) and has been used in various contexts as an antonym to polygenism. The debate in anthropology had swung in favour of monogenism by the mid-20th century. Isolated proponents of polygenism held forth in the mid-20th century, such as Carleton Coon, who thought as late as 1962 that H. sapiens arose five times from H. erectus in five places.[118]

Multiregional origin hypothesis

[edit]

The historical alternative to the recent origin model is the multiregional origin of modern humans, initially proposed by Milford Wolpoff in the 1980s. This view proposes that the derivation of anatomically modern human populations from H. erectus at the beginning of the Pleistocene 1.8 million years BP, has taken place within a continuous world population. The hypothesis necessarily rejects the assumption of an infertility barrier between ancient Eurasian and African populations of Homo. The hypothesis was controversially debated during the late 1980s and the 1990s.[119] The now-current terminology of "recent-origin" and "Out of Africa" became current in the context of this debate in the 1990s.[120] Originally seen as an antithetical alternative to the recent origin model, the multiregional hypothesis in its original "strong" form is obsolete, while its various modified weaker variants have become variants of a view of "recent origin" combined with archaic admixture.[121] Stringer (2014) distinguishes the original or "classic" Multiregional model as having existed from 1984 (its formulation) until 2003, to a "weak" post-2003 variant that has "shifted close to that of the Assimilation Model".[122][123]

Mitochondrial analyses

[edit]

In the 1980s, Allan Wilson together with Rebecca L. Cann and Mark Stoneking worked on genetic dating of the matrilineal most recent common ancestor of modern human populations (dubbed "Mitochondrial Eve"). To identify informative genetic markers for tracking human evolutionary history, Wilson concentrated on mitochondrial DNA (mtDNA), which is maternally inherited. This DNA material mutates quickly, making it easy to plot changes over relatively short times. With his discovery that human mtDNA is genetically much less diverse than chimpanzee mtDNA, Wilson concluded that modern human populations had diverged recently from a single population while older human species such as Neanderthals and Homo erectus had become extinct.[124] With the advent of archaeogenetics in the 1990s, the dating of mitochondrial and Y-chromosomal haplogroups became possible with some confidence. By 1999, estimates ranged around 150,000 years for the mt-MRCA and 60,000 to 70,000 years for the migration out of Africa.[125]

From 2000 to 2003, there was controversy about the mitochondrial DNA of "Mungo Man 3" (LM3) and its possible bearing on the multiregional hypothesis. LM3 was found to have more than the expected number of sequence differences when compared to modern human DNA (CRS).[126] Comparison of the mitochondrial DNA with that of ancient and modern aborigines, led to the conclusion that Mungo Man fell outside the range of genetic variation seen in Aboriginal Australians and was used to support the multiregional origin hypothesis. A reanalysis of LM3 and other ancient specimens from the area published in 2016, showed it to be akin to modern Aboriginal Australian sequences, inconsistent with the results of the earlier study.[127]

Y-chromosome analyses

[edit]
Map of Y-chromosome haplogroups – dominant haplogroups in pre-colonial populations with proposed migrations routes

The Y chromosome, which is paternally inherited, does not go through much recombination and thus stays largely the same after inheritance. Similar to Mitochondrial Eve, this could be studied to track the male most recent common ancestor ("Y-chromosomal Adam" or Y-MRCA).[128]

The most basal lineages have been detected in West, Northwest and Central Africa, suggesting plausibility for the Y-MRCA living in the general region of "Central-Northwest Africa".[16]

A Stanford University School of Medicine study was done by comparing Y-chromosome sequences and mtDNA in 69 men from different geographic regions and constructing a family tree. It was found that the Y-MRCA lived between 120,000 and 156,000, and the Mitochondrial Eve lived between 99,000 and 148,000 years ago, which not only predates some proposed waves of migration, but also meant that both lived in the African continent around the same time period.[129]

Another study finds a plausible placement in "the north-western quadrant of the African continent" for the emergence of the A1b haplogroup.[130] The 2013 report of haplogroup A00 found among the Mbo people of western present-day Cameroon is also compatible with this picture.[131]

The revision of Y-chromosomal phylogeny since 2011 has affected estimates for the likely geographical origin of Y-MRCA as well as estimates on time depth. By the same reasoning, future discovery of presently-unknown archaic haplogroups in living people would again lead to such revisions. In particular, the possible presence of between 1% and 4% Neanderthal-derived DNA in Eurasian genomes implies that the (unlikely) event of a discovery of a single living Eurasian male exhibiting a Neanderthal patrilineal line would immediately push back T-MRCA ("time to MRCA") to at least twice its current estimate. However, the discovery of a Neanderthal Y-chromosome by Mendez et al. was tempered by a 2016 study that suggests the extinction of Neanderthal patrilineages, as the lineage inferred from the Neanderthal sequence is outside of the range of contemporary human genetic variation.[132] Questions of geographical origin would become part of the debate on Neanderthal evolution from Homo erectus.

See also

[edit]

Notes

[edit]
  1. ^ Also called the recent single-origin hypothesis (RSOH), replacement hypothesis, or recent African origin model (RAO).
  2. ^ From 1984 to 2003, an alternative scientific hypothesis was the multiregional origin of modern humans, which envisioned a wave of Homo sapiens migrating earlier from Africa and interbreeding with local Homo erectus populations in varied regions of the globe.Jurmain R, Kilgore L, Trevathan W (2008). Essentials of Physical Anthropology. Cengage Learning. pp. 266–. ISBN 978-0495509394. Retrieved 14 June 2011.
  3. ^ McChesney (2015): "...genetic evidence suggests that a small band with the marker M168 migrated out of Africa along the coasts of the Arabian Peninsula and India, through Indonesia, and reached Australia very early, between 60,000 and 50,000 years ago. This very early migration into Australia is also supported by Rasmussen et al. (2011)."
  4. ^ McChesney (2015): "Wells (2003) divided the descendants of men who left Africa into a genealogical tree with 11 lineages. Each genetic marker represents a single-point mutation (SNP) at a specific place in the genome. First, genetic evidence suggests that a small band with the marker M168 migrated out of Africa along the coasts of the Arabian Peninsula and India, through Indonesia, and reached Australia very early, between 60,000 and 50,000 years ago. This very early migration into Australia is also supported by Rasmussen et al. (2011). Second, a group bearing the marker M89 moved out of northeastern Africa into the Middle East 45,000 years ago. From there, the M89 group split into two groups. One group that developed the marker M9 went into Asia about 40,000 years ago. The Asian (M9) group split three ways: into Central Asia (M45), 35,000 years ago; into India (M20), 30,000 years ago; and into China (M122), 10,000 years ago. The Central Asian (M45) group split into two groups: toward Europe (M173), 30,000 years ago and toward Siberia (M242), 20,000 years ago. Finally, the Siberian group (M242) went on to populate North and South America (M3), about 10,000 years ago.[31]
  5. ^ The researchers used radiocarbon dating techniques on pollen grains trapped in lake-bottom mud to establish vegetation over the ages of the Malawi lake in Africa, taking samples at 300-year-intervals. Samples from the megadrought times had little pollen or charcoal, suggesting sparse vegetation with little to burn. The area around Lake Malawi, today heavily forested, was a desert approximately 135,000 to 90,000 years ago.[49]

References

[edit]
  1. ^ Liu, Prugnolle et al. (2006). "Currently available genetic and archaeological evidence is supportive of a recent single origin of modern humans in East Africa. However, this is where the consensus on human settlement history ends, and considerable uncertainty clouds any more detailed aspect of human colonization history."
  2. ^ Stringer C (June 2003). "Human evolution: Out of Ethiopia". Nature. 423 (6941): 692–693, 695. Bibcode:2003Natur.423..692S. doi:10.1038/423692a. PMID 12802315. S2CID 26693109.
  3. ^ Stringer C (2012). Lone Survivors: How We Came to Be the Only Humans on Earth. Henry Holt and Company. p. 26. ISBN 978-1429973441 – via Google Books.
  4. ^ Wolpoff MH, Hawks J, Caspari R (May 2000). "Multiregional, not multiple origins" (PDF). American Journal of Physical Anthropology. 112 (1): 129–136. doi:10.1002/(SICI)1096-8644(200005)112:1<129::AID-AJPA11>3.0.CO;2-K. hdl:2027.42/34270. PMID 10766948.
  5. ^ Mafessoni F (January 2019). "Encounters with archaic hominins". Nature Ecology & Evolution. 3 (1): 14–15. doi:10.1038/s41559-018-0729-6. PMID 30478304. S2CID 53783648.
  6. ^ Villanea FA, Schraiber JG (January 2019). "Multiple episodes of interbreeding between Neanderthal and modern humans". Nature Ecology & Evolution. 3 (1): 39–44. doi:10.1038/s41559-018-0735-8. PMC 6309227. PMID 30478305.
  7. ^ a b University of Huddersfield (20 March 2019). "Researchers shed new light on the origins of modern humans – The work, published in Nature, confirms a dispersal of Homo sapiens from southern to eastern Africa immediately preceded the out-of-Africa migration". EurekAlert!. Retrieved 23 March 2019.
  8. ^ a b Rito T, Vieira D, Silva M, Conde-Sousa E, Pereira L, Mellars P, et al. (March 2019). "A dispersal of Homo sapiens from southern to eastern Africa immediately preceded the out-of-Africa migration". Scientific Reports. 9 (1): 4728. Bibcode:2019NatSR...9.4728R. doi:10.1038/s41598-019-41176-3. PMC 6426877. PMID 30894612.
  9. ^ Scerri EM, Chikhi L, Thomas MG (October 2019). "Beyond multiregional and simple out-of-Africa models of human evolution". Nature Ecology & Evolution. 3 (10): 1370–1372. Bibcode:2019NatEE...3.1370S. doi:10.1038/s41559-019-0992-1. hdl:10400.7/954. PMID 31548642. S2CID 202733639.
  10. ^ Scerri EM, Thomas MG, Manica A, Gunz P, Stock JT, Stringer C, et al. (August 2018). "Did Our Species Evolve in Subdivided Populations across Africa, and Why Does It Matter?". Trends in Ecology & Evolution. 33 (8): 582–594. Bibcode:2018TEcoE..33..582S. doi:10.1016/j.tree.2018.05.005. PMC 6092560. PMID 30007846.
  11. ^ Zimmer C (10 July 2019). "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". The New York Times. Retrieved 11 July 2019.
  12. ^ Staff (10 July 2019). "'Oldest remains' outside Africa reset human migration clock". Phys.org. Retrieved 10 July 2019.
  13. ^ Harvati K, Röding C, Bosman AM, Karakostis FA, Grün R, Stringer C, et al. (July 2019). "Apidima Cave fossils provide earliest evidence of Homo sapiens in Eurasia". Nature. 571 (7766): 500–504. doi:10.1038/s41586-019-1376-z. PMID 31292546. S2CID 195873640.
  14. ^ a b c Armitage SJ, Jasim SA, Marks AE, Parker AG, Usik VI, Uerpmann HP (January 2011). "The southern route "out of Africa": evidence for an early expansion of modern humans into Arabia". Science. 331 (6016): 453–456. Bibcode:2011Sci...331..453A. doi:10.1126/science.1199113. PMID 21273486. S2CID 20296624.
  15. ^ a b Balter M (January 2011). "Was North Africa the launch pad for modern human migrations?" (PDF). Science. 331 (6013): 20–23. Bibcode:2011Sci...331...20B. doi:10.1126/science.331.6013.20. PMID 21212332.
  16. ^ a b c Cruciani F, Trombetta B, Massaia A, Destro-Bisol G, Sellitto D, Scozzari R (June 2011). "A revised root for the human Y chromosomal phylogenetic tree: the origin of patrilineal diversity in Africa". American Journal of Human Genetics. 88 (6): 814–818. doi:10.1016/j.ajhg.2011.05.002. PMC 3113241. PMID 21601174.
  17. ^ a b Smith TM, Tafforeau P, Reid DJ, Grün R, Eggins S, Boutakiout M, Hublin JJ (April 2007). "Earliest evidence of modern human life history in North African early Homo sapiens". Proceedings of the National Academy of Sciences of the United States of America. 104 (15): 6128–6133. Bibcode:2007PNAS..104.6128S. doi:10.1073/pnas.0700747104. PMC 1828706. PMID 17372199.
  18. ^ Bae CJ, Douka K, Petraglia MD (December 2017). "On the origin of modern humans: Asian perspectives". Science. 358 (6368): eaai9067. doi:10.1126/science.aai9067. PMID 29217544.
  19. ^ Kuo L (10 December 2017). "Early humans migrated out of Africa much earlier than we thought". Quartz. Retrieved 10 December 2017.
  20. ^ [14][15][16][17][18][19]
  21. ^ a b Liu, Martinón-Torres et al. (2015).
    See also Modern humans in China ~80,000 years ago (?), Dieneks' Anthropology Blog.
  22. ^ a b Finlayson (2009), p. 68.
  23. ^ a b Liu, Prugnolle et al. (2006).
  24. ^ a b c Haber M, Jones AL, Connell BA, Arciero E, Yang H, Thomas MG, et al. (August 2019). "A Rare Deep-Rooting D0 African Y-Chromosomal Haplogroup and Its Implications for the Expansion of Modern Humans Out of Africa". Genetics. 212 (4): 1421–1428. doi:10.1534/genetics.119.302368. PMC 6707464. PMID 31196864.
  25. ^ Posth C, Renaud G, Mittnik M, Drucker DG, Rougier H, Cupillard C, et al. (2016). "Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-Africans and a Late Glacial Population Turnover in Europe". Current Biology. 26 (6): 827–833. Bibcode:2016CBio...26..827P. doi:10.1016/j.cub.2016.01.037. hdl:2440/114930. PMID 26853362. S2CID 140098861.
  26. ^ Karmin M, Saag L, Vicente M, Wilson Sayres MA, Järve M, Talas UG, et al. (April 2015). "A recent bottleneck of Y chromosome diversity coincides with a global change in culture". Genome Research. 25 (4): 459–466. doi:10.1101/gr.186684.114. PMC 4381518. PMID 25770088.
  27. ^ a b Clarkson C, Jacobs Z, Marwick B, Fullagar R, Wallis L, Smith M, et al. (July 2017). "Human occupation of northern Australia by 65,000 years ago". Nature. 547 (7663): 306–310. Bibcode:2017Natur.547..306C. doi:10.1038/nature22968. hdl:2440/107043. PMID 28726833. S2CID 205257212.
  28. ^ a b St. Fleur, Nicholas (19 July 2017). "Humans First Arrived in Australia 65,000 Years Ago, Study Suggests". The New York Times.
  29. ^ a b c Wood R (2 September 2017). "Comments on the chronology of Madjedbebe". Australian Archaeology. 83 (3): 172–174. doi:10.1080/03122417.2017.1408545. ISSN 0312-2417. S2CID 148777016.
  30. ^ a b c O'Connell JF, Allen J, Williams MA, Williams AN, Turney CS, Spooner NA, et al. (August 2018). "Homo sapiens first reach Southeast Asia and Sahul?". Proceedings of the National Academy of Sciences of the United States of America. 115 (34): 8482–8490. Bibcode:2018PNAS..115.8482O. doi:10.1073/pnas.1808385115. PMC 6112744. PMID 30082377.
  31. ^ a b c McChesney 2015.
  32. ^ a b c d e Macaulay et al. (2005).
  33. ^ a b Posth et al. (2016).
    See also "mtDNA from 55 hunter-gatherers across 35,000 years in Europe". Dienekes' Anthroplogy Blog. 8 February 2016.
  34. ^ Prüfer K, Racimo F, Patterson N, Jay F, Sankararaman S, Sawyer S, et al. (January 2014) [Online 2013]. "The complete genome sequence of a Neanderthal from the Altai Mountains". Nature. 505 (7481): 43–49. Bibcode:2014Natur.505...43P. doi:10.1038/nature12886. PMC 4031459. PMID 24352235.
  35. ^ a b Lachance J, Vernot B, Elbers CC, Ferwerda B, Froment A, Bodo JM, et al. (August 2012). "Evolutionary history and adaptation from high-coverage whole-genome sequences of diverse African hunter-gatherers". Cell. 150 (3): 457–469. doi:10.1016/j.cell.2012.07.009. PMC 3426505. PMID 22840920.
  36. ^ a b Hammer MF, Woerner AE, Mendez FL, Watkins JC, Wall JD (September 2011). "Genetic evidence for archaic admixture in Africa". Proceedings of the National Academy of Sciences of the United States of America. 108 (37): 15123–15128. Bibcode:2011PNAS..10815123H. doi:10.1073/pnas.1109300108. PMC 3174671. PMID 21896735.
  37. ^ a b c Douka K, Bergman CA, Hedges RE, Wesselingh FP, Higham TF (11 September 2013). "Chronology of Ksar Akil (Lebanon) and implications for the colonization of Europe by anatomically modern humans". PLOS ONE. 8 (9): e72931. Bibcode:2013PLoSO...872931D. doi:10.1371/journal.pone.0072931. PMC 3770606. PMID 24039825.
  38. ^ Vidal CM, Lane CS, Asrat A, Barfod DN, Mark DF, Tomlinson EL, et al. (January 2022). "Age of the oldest known Homo sapiens from eastern Africa". Nature. 601 (7894): 579–583. Bibcode:2022Natur.601..579V. doi:10.1038/s41586-021-04275-8. PMC 8791829. PMID 35022610.
  39. ^ Hublin JJ, Ben-Ncer A, Bailey SE, Freidline SE, Neubauer S, Skinner MM, et al. (June 2017). "New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens" (PDF). Nature. 546 (7657): 289–292. Bibcode:2017Natur.546..289H. doi:10.1038/nature22336. PMID 28593953. S2CID 256771372.
  40. ^ a b c Beyin (2011).
  41. ^ Posth C, Wißing C, Kitagawa K, Pagani L, van Holstein L, Racimo F, et al. (July 2017). "Deeply divergent archaic mitochondrial genome provides lower time boundary for African gene flow into Neanderthals". Nature Communications. 8: 16046. Bibcode:2017NatCo...816046P. doi:10.1038/ncomms16046. PMC 5500885. PMID 28675384.; see also Zimmer C (4 July 2017). "In Neanderthal DNA, Signs of a Mysterious Human Migration". The New York Times. Retrieved 4 July 2017..
  42. ^ Harvati K, Röding C, Bosman AM, Karakosti FA, Grün R, Stringer C, et al. (July 2019). "Apidima Cave fossils provide earliest evidence of Homo sapiens in Eurasia". Nature. 571 (7766): 500–504. doi:10.1038/s41586-019-1376-z. PMID 31292546. S2CID 195873640.
  43. ^ "Scientists discover oldest known modern human fossil outside of Africa: Analysis of fossil suggests Homo sapiens left Africa at least 50,000 years earlier than previously thought". ScienceDaily. Retrieved 27 January 2018.
  44. ^ Ghosh P (2018). "Modern humans left Africa much earlier". BBC News. Retrieved 27 January 2018.
  45. ^ Hershkovitz I, Weber GW, Quam R, Duval M, Grün R, Kinsley L, et al. (January 2018). "The earliest modern humans outside Africa". Science. 359 (6374): 456–459. Bibcode:2018Sci...359..456H. doi:10.1126/science.aap8369. hdl:10072/372670. PMID 29371468. S2CID 206664380.
  46. ^ Groucutt HS, Grün R, Zalmout IS, Drake NA, Armitage SJ, Candy I, et al. (May 2018). "Homo sapiens in Arabia by 85,000 years ago". Nature Ecology & Evolution. 2 (5): 800–809. Bibcode:2018NatEE...2..800G. doi:10.1038/s41559-018-0518-2. PMC 5935238. PMID 29632352.
  47. ^ Liu W, Jin CZ, Zhang YQ, Cai YJ, Xing S, Wu XJ, et al. (November 2010). "Human remains from Zhirendong, South China, and modern human emergence in East Asia". Proceedings of the National Academy of Sciences of the United States of America. 107 (45): 19201–19206. Bibcode:2010PNAS..10719201L. doi:10.1073/pnas.1014386107. PMC 2984215. PMID 20974952.
  48. ^ a b c d e Appenzeller (2012).
  49. ^ a b Jensen MN (8 October 2007). "Newfound Ancient African Megadroughts May Have Driven Evolution of Humans and Fish. The findings provide new insights into humans' migration out of Africa and the evolution of fishes in Africa's Great Lakes". The University of Arizona. Retrieved 25 September 2017.
  50. ^ Stringer CB, Grün R, Schwarcz HP, Goldberg P (April 1989). "ESR dates for the hominid burial site of Es Skhul in Israel". Nature. 338 (6218): 756–758. Bibcode:1989Natur.338..756S. doi:10.1038/338756a0. PMID 2541339. S2CID 4332370.
  51. ^ Grün R, Stringer C, McDermott F, Nathan R, Porat N, Robertson S, et al. (September 2005). "U-series and ESR analyses of bones and teeth relating to the human burials from Skhul". Journal of Human Evolution. 49 (3): 316–334. Bibcode:2005JHumE..49..316G. doi:10.1016/j.jhevol.2005.04.006. PMID 15970310.
  52. ^ Scerri, Eleanor M.L.; Drake, Nick A.; Jennings, Richard; Groucutt, Huw S. (2014). "Earliest evidence for the structure of Homo sapiens populations in Africa". Quaternary Science Reviews. 101: 207–216. Bibcode:2014QSRv..101..207S. doi:10.1016/j.quascirev.2014.07.019.
  53. ^ Bretzke, Knut; Armitage, Simon J.; Parker, Adrian G.; Walkington, Helen; Uerpmann, Hans-Peter (2013). "The environmental context of Paleolithic settlement at Jebel Faya, Emirate Sharjah, UAE". Quaternary International. 300: 83–93. Bibcode:2013QuInt.300...83B. doi:10.1016/j.quaint.2013.01.028.
  54. ^ Rose JI, Usik VI, Marks AE, Hilbert YH, Galletti CS, Parton A, et al. (2011). "The Nubian Complex of Dhofar, Oman: an African middle Stone Age industry in Southern Arabia". PLOS ONE. 6 (11): e28239. Bibcode:2011PLoSO...628239R. doi:10.1371/journal.pone.0028239. PMC 3227647. PMID 22140561.
  55. ^ a b c Kuhlwilm et al. (2016).
    See also Ancestors of Eastern Neandertals admixed with modern humans 100 thousand years ago, Dienekes'Anthropology Blog.
  56. ^ Gibbons A (2 March 2017). "Ancient skulls may belong to elusive humans called Denisovans". Science. doi:10.1126/science.aal0846. Retrieved 25 September 2017.
  57. ^ Zhivotovsky LA, Rosenberg NA, Feldman MW (May 2003). "Features of evolution and expansion of modern humans, inferred from genomewide microsatellite markers". American Journal of Human Genetics. 72 (5): 1171–1186. doi:10.1086/375120. PMC 1180270. PMID 12690579.
  58. ^ Stix G (2008). "The Migration History of Humans: DNA Study Traces Human Origins Across the Continents". Scientific American. Retrieved 14 June 2011.
  59. ^ a b c Metspalu M, Kivisild T, Metspalu E, Parik J, Hudjashov G, Kaldma K, et al. (August 2004). "Most of the extant mtDNA boundaries in south and southwest Asia were likely shaped during the initial settlement of Eurasia by anatomically modern humans". BMC Genetics. 5: 26. doi:10.1186/1471-2156-5-26. PMC 516768. PMID 15339343.
  60. ^ Fernandes CA, Rohling EJ, Siddall M (June 2006). "Absence of post-Miocene Red Sea land bridges: biogeographic implications". Journal of Biogeography. 33 (6): 961–966. Bibcode:2006JBiog..33..961F. doi:10.1111/j.1365-2699.2006.01478.x.
  61. ^ Walter RC, Buffler RT, Bruggemann JH, Guillaume MM, Berhe SM, Negassi B, et al. (May 2000). "Early human occupation of the Red Sea coast of Eritrea during the last interglacial". Nature. 405 (6782): 65–69. Bibcode:2000Natur.405...65W. doi:10.1038/35011048. PMID 10811218. S2CID 4417823.
  62. ^ Catherine B (24 November 2012). "Our True Dawn". New Scientist. 216 (2892): 34–37. Bibcode:2012NewSc.216...34B. doi:10.1016/S0262-4079(12)63018-8. ISSN 0262-4079.
  63. ^ Karmin M, Saag L, Vicente M, Wilson Sayres MA, Järve M, et al. (April 2015). "A recent bottleneck of Y chromosome diversity coincides with a global change in culture". Genome Research. 25 (4): 459–466. doi:10.1101/gr.186684.114. PMC 4381518. PMID 25770088.
  64. ^ a b Vai S, Sarno S, Lari M, Luiselli D, Manzi G, Gallinaro M, et al. (March 2019). "Ancestral mitochondrial N lineage from the Neolithic 'green' Sahara". Scientific Reports. 9 (1): 3530. Bibcode:2019NatSR...9.3530V. doi:10.1038/s41598-019-39802-1. PMC 6401177. PMID 30837540.
  65. ^ Hershkovitz et al. (2015)
    See also "55,000-Year-Old Skull Fossil Sheds New Light on Human Migration out of Africa". Science News. 29 January 2015.
  66. ^ Shea, John J. (2003). "Neandertals, competition, and the origin of modern human behavior in the Levant". Evolutionary Anthropology: Issues, News, and Reviews. 12 (4): 173–187. doi:10.1002/evan.10101. ISSN 1060-1538.
  67. ^ Beyer RM, Krapp M, Eriksson A, Manica A (August 2021). "Climatic windows for human migration out of Africa in the past 300,000 years". Nature Communications. 12 (1): 4889. Bibcode:2021NatCo..12.4889B. doi:10.1038/s41467-021-24779-1. PMC 8384873. PMID 34429408.
  68. ^ Tobler, Raymond; Souilmi, Yassine; Huber, Christian D.; Bean, Nigel; Turney, Chris S. M.; Grey, Shane T.; Cooper, Alan (30 May 2023). "The role of genetic selection and climatic factors in the dispersal of anatomically modern humans out of Africa". Proceedings of the National Academy of Sciences. 120 (22): e2213061120. Bibcode:2023PNAS..12013061T. doi:10.1073/pnas.2213061120. ISSN 0027-8424. PMC 10235988. PMID 37220274.
  69. ^ Bowler JM, Johnston H, Olley JM, Prescott JR, Roberts RG, Shawcross W, Spooner NA (February 2003). "New ages for human occupation and climatic change at Lake Mungo, Australia". Nature. 421 (6925): 837–640. Bibcode:2003Natur.421..837B. doi:10.1038/nature01383. PMID 12594511. S2CID 4365526.
  70. ^ Olleya JM, Roberts RG, Yoshida H, Bowler JM (2006). "Single-grain optical dating of grave-infill associated with human burials at Lake Mungo, Australia". Quaternary Science Reviews. 25 (19–20): 2469–2474. Bibcode:2006QSRv...25.2469O. doi:10.1016/j.quascirev.2005.07.022.
  71. ^ Clarkson, Chris; Jacobs, Zenobia; et al. (19 July 2017). "Human occupation of northern Australia by 65,000 years ago" (PDF). Nature. 547 (7663): 306–310. Bibcode:2017Natur.547..306C. doi:10.1038/nature22968. hdl:2440/107043. PMID 28726833. S2CID 205257212.
  72. ^ Gibbons, Ann (20 July 2017). "The first Australians arrived early". Science. 357 (6348): 238–239. Bibcode:2017Sci...357..238G. doi:10.1126/science.357.6348.238. PMID 28729491.
  73. ^ "Almost all living people outside of Africa trace back to a single migration more than 50,000 years ago". www.science.org. Retrieved 19 August 2022.
  74. ^ Yang M (6 January 2022). "A genetic history of migration, diversification, and admixture in Asia". Human Population Genetics and Genomics: 1–32. doi:10.47248/hpgg2202010001.
  75. ^ Genetics and material culture support repeated expansions into Paleolithic Eurasia from a population hub out of Africa, Vallini et al. 2022 (April 4, 2022) Quote: "Taken together with a lower bound of the final settlement of Sahul at 37 kya it is reasonable to describe Papuans as either an almost even mixture between East-Eurasians and a lineage basal to West and East-Eurasians which occurred sometimes between 45 and 38kya, or as a sister lineage of East-Eurasians with or without a minor basal OoA or xOoA contribution. We here chose to parsimoniously describe Papuans as a simple sister group of Tianyuan, cautioning that this may be just one out of six equifinal possibilities."
  76. ^ Genetics and material culture support repeated expansions into Paleolithic Eurasia from a population hub out of Afri, Vallini et al. 2021 (October 15, 2021) Quote: "Taken together with a lower bound of the final settlement of Sahul at 37 kya (the date of the deepest population splits estimated by 1) it is reasonable to describe Papuans as either an almost even mixture between East Asians and a lineage basal to West and East Asians occurred sometimes between 45 and 38kya, or as a sister lineage of East Asians with or without a minor basal OoA or xOoA contribution. "
  77. ^ Genomic insights into the human population history of Australia and New Guinea, University of Cambridge, Bergström et al. 2018
  78. ^ Carlhoff S, Duli A, Nägele K, Nur M, Skov L, Sumantri I, et al. (August 2021). "Genome of a middle Holocene hunter-gatherer from Wallacea". Nature. 596 (7873): 543–547. Bibcode:2021Natur.596..543C. doi:10.1038/s41586-021-03823-6. PMC 8387238. PMID 34433944. The qpGraph analysis confirmed this branching pattern, with the Leang Panninge individual branching off from the Near Oceanian clade after the Denisovan gene flow, although with the most supported topology indicating around 50% of a basal East Asian component contributing to the Leang Panninge genome (Fig. 3c, Supplementary Figs. 7–11).
  79. ^ Shen G, Wang W, Wang Q, Zhao J, Collerson K, Zhou C, Tobias PV (December 2002). "U-Series dating of Liujiang hominid site in Guangxi, Southern China". Journal of Human Evolution. 43 (6): 817–829. Bibcode:2002JHumE..43..817S. doi:10.1006/jhev.2002.0601. PMID 12473485.
  80. ^ a b Rosenburg K (2002). "A Late Pleistocene Human Skeleton from Liujiang, China Suggests Regional Population Variation in Sexual Dimorphism in the Human Pelvis". Variability and Evolution.
  81. ^ Kaifu, Yousuke; Fujita, Masaki (2012). "Fossil record of early modern humans in East Asia". Quaternary International. 248: 2–11. Bibcode:2012QuInt.248....2K. doi:10.1016/j.quaint.2011.02.017.
  82. ^ "A relative from the Tianyuan Cave". Max Planck Society. 21 January 2013.
  83. ^ "A relative from the Tianyuan Cave: Humans living 40,000 years ago likely related to many present-day Asians and Native Americans". Science Daily. 21 January 2013.
  84. ^ "DNA Analysis Reveals Common Origin of Tianyuan Humans and Native Americans, Asians". Sci-News. 24 January 2013.
  85. ^ "Ancient human DNA suggests minimal interbreeding". Science News. 21 January 2013.
  86. ^ "Ancient Bone DNA Shows Ancestry of Modern Asians & Native Americans". Caving News. 31 January 2013.
  87. ^ Hu Y, Shang H, Tong H, Nehlich O, Liu W, Zhao C, et al. (July 2009). "Stable isotope dietary analysis of the Tianyuan 1 early modern human". Proceedings of the National Academy of Sciences of the United States of America. 106 (27): 10971–10974. Bibcode:2009PNAS..10610971H. doi:10.1073/pnas.0904826106. PMC 2706269. PMID 19581579.
  88. ^ Brown P (August 1992). "Recent human evolution in East Asia and Australasia". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 337 (1280): 235–242. Bibcode:1992RSPTB.337..235B. doi:10.1098/rstb.1992.0101. PMID 1357698.
  89. ^ Curnoe D, Xueping J, Herries AI, Kanning B, Taçon PS, Zhende B, et al. (14 March 2012). "Human remains from the Pleistocene-Holocene transition of southwest China suggest a complex evolutionary history for East Asians". PLOS ONE. 7 (3): e31918. Bibcode:2012PLoSO...731918C. doi:10.1371/journal.pone.0031918. PMC 3303470. PMID 22431968.
  90. ^ Larena M, Sanchez-Quinto F, Sjödin P, McKenna J, Ebeo C, Reyes R, et al. (March 2021). "Multiple migrations to the Philippines during the last 50,000 years". Proceedings of the National Academy of Sciences of the United States of America. 118 (13): e2026132118. Bibcode:2021PNAS..11826132L. doi:10.1073/pnas.2026132118. PMC 8020671. PMID 33753512.
  91. ^ Moreno-Mayar JV, Potter BA, Vinner L, Steinrücken M, Rasmussen S, Terhorst J, et al. (January 2018). "Terminal Pleistocene Alaskan genome reveals first founding population of Native Americans" (PDF). Nature. 553 (7687): 203–207. Bibcode:2018Natur.553..203M. doi:10.1038/nature25173. PMID 29323294. S2CID 4454580.
  92. ^ Gakuhari T, Nakagome S, Rasmussen S, Allentoft ME, Sato T, Korneliussen T, et al. (August 2020). "Ancient Jomon genome sequence analysis sheds light on migration patterns of early East Asian populations". Communications Biology. 3 (1): 437. doi:10.1038/s42003-020-01162-2. PMC 7447786. PMID 32843717.
  93. ^ Davis LG, Madsen DB, Becerra-Valdivia L, Higham T, Sisson DA, Skinner SM, et al. (August 2019). "Late Upper Paleolithic occupation at Cooper's Ferry, Idaho ~16,000 years ago". Science. 365 (6456): 891–897. Bibcode:2019Sci...365..891D. doi:10.1126/science.aax9830. PMID 31467216. S2CID 201672463. We interpret this temporal and technological affinity to signal a cultural connection with Upper Paleolithic northeastern Asia, which complements current evidence of shared genetic heritage between late Pleistocene peoples of northern Japan and North America.
  94. ^ Mylopotamitaki, Dorothea; Weiss, Marcel; Fewlass, Helen; Zavala, Elena Irene; Rougier, Hélène; Sümer, Arev Pelin; Hajdinjak, Mateja; Smith, Geoff M.; Ruebens, Karen; Sinet-Mathiot, Virginie; Pederzani, Sarah; Essel, Elena; Harking, Florian S.; Xia, Huan; Hansen, Jakob; Kirchner, André; Lauer, Tobias; Stahlschmidt, Mareike; Hein, Michael; Talamo, Sahra; Wacker, Lukas; Meller, Harald; Dietl, Holger; Orschiedt, Jörg; Olsen, Jesper V.; Zeberg, Hugo; Prüfer, Kay; Krause, Johannes; Meyer, Matthias; Welker, Frido; McPherron, Shannon P.; Schüler, Tim; HublinHublin, Jean-Jacques (31 January 2024). "Homo sapiens reached the higher latitudes of Europe by 45,000 years ago". Nature. 626 (7998): 341–346. Bibcode:2024Natur.626..341M. doi:10.1038/s41586-023-06923-7. ISSN 1476-4687. PMC 10849966. PMID 38297117.
  95. ^ Gonder MK, Mortensen HM, Reed FA, de Sousa A, Tishkoff SA (March 2007). "Whole-mtDNA genome sequence analysis of ancient African lineages". Molecular Biology and Evolution. 24 (3): 757–768. doi:10.1093/molbev/msl209. PMID 17194802.
  96. ^ Chen YS, Olckers A, Schurr TG, Kogelnik AM, Huoponen K, Wallace DC (April 2000). "mtDNA variation in the South African Kung and Khwe-and their genetic relationships to other African populations". American Journal of Human Genetics. 66 (4): 1362–1383. doi:10.1086/302848. PMC 1288201. PMID 10739760.
  97. ^ Soares P, Alshamali F, Pereira JB, Fernandes V, Silva NM, Afonso C, et al. (March 2012). "The Expansion of mtDNA Haplogroup L3 within and out of Africa". Molecular Biology and Evolution. 29 (3): 915–927. doi:10.1093/molbev/msr245. PMID 22096215.
  98. ^ Groucutt et al. (2015).
  99. ^ Maca-Meyer N, González AM, Larruga JM, Flores C, Cabrera VM (13 August 2001). "Major genomic mitochondrial lineages delineate early human expansions". BMC Genetics. 2: 13. doi:10.1186/1471-2156-2-13. PMC 55343. PMID 11553319.
  100. ^ Ingman M, Gyllensten U (July 2003). "Mitochondrial genome variation and evolutionary history of Australian and New Guinean aborigines". Genome Research. 13 (7): 1600–1606. doi:10.1101/gr.686603. PMC 403733. PMID 12840039.
  101. ^ Hallast P, Agdzhoyan A, Balanovsky O, Xue Y, Tyler-Smith C (February 2021). "A Southeast Asian origin for present-day non-African human Y chromosomes". Human Genetics. 140 (2): 299–307. doi:10.1007/s00439-020-02204-9. PMC 7864842. PMID 32666166.
  102. ^ Yu N, Chen FC, Ota S, Jorde LB, Pamilo P, Patthy L, et al. (May 2002). "Larger genetic differences within africans than between Africans and Eurasians". Genetics. 161 (1): 269–274. doi:10.1093/genetics/161.1.269. PMC 1462113. PMID 12019240. Retrieved 7 April 2013.
  103. ^ Coop G, Pickrell JK, Novembre J, Kudaravalli S, Li J, Absher D, et al. (June 2009). Schierup MH (ed.). "The role of geography in human adaptation". PLOS Genetics. 5 (6): e1000500. doi:10.1371/journal.pgen.1000500. PMC 2685456. PMID 19503611.; summary in Racimo F, Schraiber JG, Casey F, Huerta-Sanchez E (2016). "Directional Selection and Adaptation". In Kliman RM (ed.). Encyclopedia of Evolutionary Biology. Academic Press. p. 451. doi:10.1016/B978-0-12-800049-6.00028-7. ISBN 978-0128004265 – via Google Books.
  104. ^ Gurdasani D, Carstensen T, Tekola-Ayele F, Pagani L, Tachmazidou I, Hatzikotoulas K, et al. (January 2015). "The African Genome Variation Project shapes medical genetics in Africa". Nature. 517 (7, 534): 327–332. Bibcode:2015Natur.517..327G. doi:10.1038/nature13997. PMC 4297536. PMID 25470054.
  105. ^ Matisoo-Smith E, Horsburgh KA (2016). DNA for Archaeologists. Routledge.
  106. ^ Rasmussen M, Guo X, Wang Y, Lohmueller KE, Rasmussen S, Albrechtsen A, et al. (October 2011). "An Aboriginal Australian genome reveals separate human dispersals into Asia". Science. 334 (6052): 94–98. Bibcode:2011Sci...334...94R. doi:10.1126/science.1211177. PMC 3991479. PMID 21940856.
  107. ^ East Asians 2.3–2.6%, Western Eurasians 1.8–2.4% (Prüfer K, de Filippo C, Grote S, Mafessoni F, Korlević P, Hajdinjak M, et al. (November 2017). "A high-coverage Neandertal genome from Vindija Cave in Croatia". Science. 358 (6363): 655–658. Bibcode:2017Sci...358..655P. doi:10.1126/science.aao1887. PMC 6185897. PMID 28982794.)
  108. ^ Callaway E (2012). "Hunter-gatherer genomes a trove of genetic diversity". Nature. doi:10.1038/nature.2012.11076. S2CID 87081207.
  109. ^ Xu D, Pavlidis P, Taskent RO, Alachiotis N, Flanagan C, DeGiorgio M, et al. (October 2017). "Archaic Hominin Introgression in Africa Contributes to Functional Salivary MUC7 Genetic Variation". Molecular Biology and Evolution. 34 (10): 2704–2715. doi:10.1093/molbev/msx206. PMC 5850612. PMID 28957509.
  110. ^ Durvasula A, Sankararaman S (February 2020). "Recovering signals of ghost archaic introgression in African populations". Science Advances. 6 (7): eaax5097. Bibcode:2020SciA....6.5097D. doi:10.1126/sciadv.aax5097. PMC 7015685. PMID 32095519.
  111. ^ Petraglia M, Clarkson C, Boivin N, Haslam M, Korisettar R, Chaubey G, et al. (July 2009). "Population increase and environmental deterioration correspond with microlithic innovations in South Asia ca. 35,000 years ago". Proceedings of the National Academy of Sciences of the United States of America. 106 (30): 12261–12266. Bibcode:2009PNAS..10612261P. doi:10.1073/pnas.0810842106. PMC 2718386. PMID 19620737.
  112. ^ Cabrera VM, Marrero P, Abu-Amero KK, Larruga JM (June 2018). "Carriers of mitochondrial DNA macrohaplogroup L3 basal lineages migrated back to Africa from Asia around 70,000 years ago". BMC Evolutionary Biology. 18 (1): 98. Bibcode:2018BMCEE..18...98C. bioRxiv 10.1101/233502. doi:10.1186/s12862-018-1211-4. PMC 6009813. PMID 29921229.
  113. ^ Lafreniere P (2010). Adaptive Origins: Evolution and Human Development. Taylor & Francis. p. 90. ISBN 978-0805860122. Retrieved 14 June 2011 – via Google Books.
  114. ^ Robinson D, Ash PM (2010). The Emergence of Humans: An Exploration of the Evolutionary Timeline. New York: Wiley. ISBN 978-0470013151.
  115. ^ Palmer D (2006). Prehistoric Past Revealed: The Four Billion Year History of Life on Earth. Berkeley: University of California Press. p. 43. ISBN 978-0520248274.
  116. ^ Regal B (2004). Human evolution: a guide to the debates. Santa Barbara, Calif: ABC-CLIO. pp. 73–75. ISBN 978-1851094189.
  117. ^ "The Descent of Man Chapter 6 – "On the Affinities and Genealogy of Man"". Darwin-online.org.uk. Retrieved 11 January 2011.
  118. ^ Jackson JP Jr (2001). "'In Ways Unacademical': The Reception of Carleton S. Coon's The Origin of Races" (PDF). Journal of the History of Biology. 34 (2): 247–285. doi:10.1023/A:1010366015968. S2CID 86739986. Archived from the original (PDF) on 14 May 2013.
  119. ^ Stringer CB, Andrews P (March 1988). "Genetic and fossil evidence for the origin of modern humans". Science. 239 (4845): 1263–1268. Bibcode:1988Sci...239.1263S. doi:10.1126/science.3125610. PMID 3125610. Stringer C, Bräuer G (1994). "Methods, misreading, and bias". American Anthropologist. 96 (2): 416–424. doi:10.1525/aa.1994.96.2.02a00080.
    Stringer CB (1992). "Replacement, continuity and the origin of Homo sapiens". In Smith FH (ed.). Continuity or replacement? Controversies in Homo sapiens evolution. Rotterdam: Balkema. pp. 9–24.
    Bräuer G, Stringer C (1997). "Models, polarization, and perspectives on modern human origins". Conceptual issues in modern human origins research. New York: Aldine de Gruyter. pp. 191–201.
  120. ^ Wu L (1997). "The dental continuity of humans in China from Pleistocene to Holocene, and the origin of mongoloids". Quaternary Geology. 21: 24–32. ISBN 978-9067642439.
  121. ^ Stringer C (2001). "Modern human origins – distinguishing the models". African Archaeological Review. 18 (2): 67–75. doi:10.1023/A:1011079908461. S2CID 161991922.
  122. ^ Stringer C (April 2002). "Modern human origins: progress and prospects". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 357 (1420): 563–579. doi:10.1098/rstb.2001.1057. PMC 1692961. PMID 12028792.
  123. ^ Stringer C (May 2014). "Why we are not all multiregionalists now". Trends in Ecology & Evolution. 29 (5): 248–251. Bibcode:2014TEcoE..29..248S. doi:10.1016/j.tree.2014.03.001. PMID 24702983.
  124. ^ "Allan Wilson: Revolutionary Evolutionist". New Zealanders Heroes.
  125. ^ Wallace DC, Brown MD, Lott MT (September 1999). "Mitochondrial DNA variation in human evolution and disease". Gene. 238 (1): 211–230. doi:10.1016/S0378-1119(99)00295-4. PMID 10570998. "evidence that our species arose in Africa about 150,000 years before present (YBP), migrated out of Africa into Asia about 60,000 to 70,000 YBP and into Europe about 40,000 to 50,000 YBP, and migrated from Asia and possibly Europe to the Americas about 20,000 to 30,000 YBP."
  126. ^ Adcock GJ, Dennis ES, Easteal S, Huttley GA, Jermiin LS, Peacock WJ, Thorne A (January 2001). "Mitochondrial DNA sequences in ancient Australians: Implications for modern human origins". Proceedings of the National Academy of Sciences of the United States of America. 98 (2): 537–542. Bibcode:2001PNAS...98..537A. doi:10.1073/pnas.98.2.537. PMC 14622. PMID 11209053.
  127. ^ Heupink TH, Subramanian S, Wright JL, Endicott P, Westaway MC, Huynen L, et al. (June 2016). "Ancient mtDNA sequences from the First Australians revisited". Proceedings of the National Academy of Sciences of the United States of America. 113 (25): 6892–6897. Bibcode:2016PNAS..113.6892H. doi:10.1073/pnas.1521066113. PMC 4922152. PMID 27274055.
  128. ^ Parsons, Paul; Dixon, Gail (2016). 50 Ideas You Really Need to Know: Science. London: Quercus. p. 127. ISBN 978-1-78429-614-8.
  129. ^ Poznik, G. David; Henn, Brenna M.; Yee, Muh-Ching; Sliwerska, Elzbieta; Euskirchen, Ghia M.; Lin, Alice A.; Snyder, Michael; Quintana-Murci, Lluis; Kidd, Jeffrey M.; Underhill, Peter A.; Bustamante, Carlos D. (2 August 2013). "Sequencing Y Chromosomes Resolves Discrepancy in Time to Common Ancestor of Males Versus Females". Science. 341 (6145): 562–565. Bibcode:2013Sci...341..562P. doi:10.1126/science.1237619. ISSN 0036-8075. PMC 4032117. PMID 23908239.
  130. ^ Scozzari R, Massaia A, D'Atanasio E, Myres NM, Perego UA, Trombetta B, Cruciani F (2012). Caramelli D (ed.). "Molecular dissection of the basal clades in the human Y chromosome phylogenetic tree". PLOS ONE. 7 (11): e49170. Bibcode:2012PLoSO...749170S. doi:10.1371/journal.pone.0049170. PMC 3492319. PMID 23145109.
  131. ^ Mendez FL, Krahn T, Schrack B, Krahn AM, Veeramah KR, Woerner AE, et al. (March 2013). "An African American paternal lineage adds an extremely ancient root to the human Y chromosome phylogenetic tree" (PDF). American Journal of Human Genetics. 92 (3): 454–459. doi:10.1016/j.ajhg.2013.02.002. PMC 3591855. PMID 23453668.
  132. ^ Mendez FL, Poznik GD, Castellano S, Bustamante CD (April 2016). "The Divergence of Neandertal and Modern Human Y Chromosomes". American Journal of Human Genetics. 98 (4): 728–734. doi:10.1016/j.ajhg.2016.02.023. PMC 4833433. PMID 27058445.

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