Editing Human evolution (section)

==== Dispersal of modern ''Homo sapiens'' ====
Up until the genetic evidence became available, there were two dominant models for the dispersal of modern humans. The [[Multiregional evolution|multiregional hypothesis]] proposed that the genus ''Homo'' contained only a single interconnected population as it does today (not separate species), and that its evolution took place worldwide continuously over the last couple of million years. This model was proposed in 1988 by [[Milford H. Wolpoff]].<ref>{{cite journal |last1=Wolpoff |first1=Milford H. |author1-link=Milford H. Wolpoff |last2=Hawks |first2=John |author2-link=John D. Hawks |last3=Caspari |first3=Rachel |date=May 2000 |title=Multiregional, Not Multiple Origins |journal=American Journal of Physical Anthropology |volume=112 |issue=1 |pages=129–136 |doi=10.1002/(SICI)1096-8644(200005)112:1<129::AID-AJPA11>3.0.CO;2-K |issn=0002-9483 |pmid=10766948 |url= http://www-personal.umich.edu/~wolpoff/Papers/Multiregional.PDF |hdl=2027.42/34270 |hdl-access=free |access-date=May 12, 2013 |archive-date=January 25, 2021 |archive-url= https://web.archive.org/web/20210125052356/http://www-personal.umich.edu/~wolpoff/Papers/Multiregional.PDF |url-status=live}}</ref><ref>{{cite journal |last1=Wolpoff |first1=M. H. |author1-link=Milford H. Wolpoff |last2=Spuhler |first2=J. N. |last3=Smith |first3=F. H. |last4=Radovcic |first4=J. |last5=Pope |first5=G. |last6=Frayer |first6=D. W. |last7=Eckhardt |first7=R. |last8=Clark |first8=G. |date=August 12, 1988 |title=Modern Human Origins |journal=[[Science (journal)|Science]] |volume=241 |issue=4867 |pages=772–774 |bibcode=1988Sci...241..772W |doi=10.1126/science.3136545 |issn=0036-8075 |pmid=3136545 |s2cid=5223638}}</ref> In contrast, the "out of Africa" model proposed that modern ''H.&nbsp;sapiens'' speciated in Africa recently (that is, approximately 200,000 years ago) and the subsequent migration through [[Eurasia]] resulted in the nearly complete replacement of other ''Homo'' species. This model has been developed by [[Chris Stringer]] and Peter Andrews.<ref>{{cite news |last=Owen |first=James |date=July 18, 2007 |url= http://news.nationalgeographic.com/news/2007/07/070718-african-origin.html |title=Modern Humans Came Out of Africa, 'Definitive' Study Says |work=National Geographic News |location=Washington, DC |publisher=[[National Geographic Society]] |access-date=May 14, 2011 |archive-date=June 27, 2018 |archive-url= https://web.archive.org/web/20180627005155/https://news.nationalgeographic.com/news/2007/07/070718-african-origin.html |url-status=dead}}</ref><ref>{{cite journal |last1=Stringer |first1=Chris B. |last2=Andrews |first2=Peter |date=March 11, 1988 |title=Genetic and fossil evidence for the origin of modern humans |journal=[[Science (journal)|Science]] |volume=239 |issue=4845 |pages=1263–1268 |issn=0036-8075 |bibcode=1988Sci...239.1263S |doi=10.1126/science.3125610 |pmid=3125610 |s2cid=39200956}}</ref>

Sequencing mtDNA and Y-DNA sampled from a wide range of indigenous populations revealed ancestral information relating to both male and female genetic heritage, and strengthened the "out of Africa" theory and weakened the views of multiregional evolutionism.{{sfn|Webster|2010|p=[https://books.google.com/books?id=cdiy1EhJVW0C&pg=PA53 53]}} Aligned in genetic tree differences were interpreted as supportive of a recent single origin.{{sfn|Ramachandran|Tang|Gutenkunst|Bustamante|2010|p=[https://books.google.com/books?id=FlfPSpBvKLgC&pg=PA606 606]}}

"Out of Africa" has thus gained much support from research using female mitochondrial DNA and the male [[Y chromosome]]. After analysing genealogy trees constructed using 133 types of mtDNA, researchers concluded that all were descended from a female African progenitor, dubbed [[Mitochondrial Eve]]. "Out of Africa" is also supported by the fact that mitochondrial genetic diversity is highest among African populations.<ref name="Cann">{{cite journal |last1=Cann |first1=Rebecca L. |author1-link=Rebecca L. Cann |last2=Stoneking |first2=Mark |author2-link=Mark Stoneking |last3=Wilson |first3=Allan C. |author3-link=Allan Wilson (biologist) |date=January 1, 1987 |title=Mitochondrial DNA and human evolution |url= http://artsci.wustl.edu/~landc/html/cann/ |journal=[[Nature (journal)|Nature]] |volume=325 |issue=6099 |pages=31–36 |bibcode=1987Natur.325...31C |doi=10.1038/325031a0 |issn=0028-0836 |pmid=3025745 |s2cid=4285418 |archive-url= https://web.archive.org/web/20100813121953/http://artsci.wustl.edu/~landc/html/cann/ |archive-date=August 13, 2010 |access-date=April 21, 2015}}</ref>

A broad study of African genetic diversity, headed by [[Sarah Tishkoff]], found the [[San people]] had the greatest genetic diversity among the 113 distinct populations sampled, making them one of 14 "ancestral population clusters". The research also located a possible origin of modern human migration in southwestern Africa, near the coastal border of [[Namibia]] and [[Angola]].<ref>{{cite news |last=Gill |first=Victoria |date=May 1, 2009 |title=Africa's genetic secrets unlocked |work=[[BBC News]] |location=London |publisher=BBC |url= http://news.bbc.co.uk/2/hi/science/nature/8027269.stm |access-date=June 8, 2011 |archive-date=May 14, 2020 |archive-url= https://web.archive.org/web/20200514095952/http://news.bbc.co.uk/2/hi/science/nature/8027269.stm |url-status=live}}</ref> The fossil evidence was insufficient for archaeologist [[Richard Leakey]] to resolve the debate about exactly where in Africa modern humans first appeared.{{sfn|Leakey|1994|pp=87–89}} Studies of [[haplogroup]]s in [[Human Y-chromosome DNA haplogroup|Y-chromosomal DNA]] and [[Human mitochondrial DNA haplogroup|mitochondrial DNA]] have largely supported a recent African origin.<ref>{{cite journal |last1=Jorde |first1=Lynn B. |last2=Bamshad |first2=Michael |last3=Rogers |first3=Alan R. |author3-link=Alan R. Rogers |date=February 1998 |title=Using mitochondrial and nuclear DNA markers to reconstruct human evolution |journal=BioEssays |volume=20 |issue=2 |pages=126–136 |doi=10.1002/(SICI)1521-1878(199802)20:2<126::AID-BIES5>3.0.CO;2-R |issn=0265-9247 |pmid=9631658 |s2cid=17203268}}</ref> All the evidence from autosomal DNA also predominantly supports a Recent African origin. However, evidence for [[Archaic human admixture with modern humans|archaic admixture in modern humans]], both in Africa and later, throughout Eurasia has recently been suggested by a number of studies.<ref>{{cite journal |last1=Wall |first1=Jeffrey D. |last2=Lohmueller |first2=Kirk E. |last3=Plagnol |first3=Vincent |date=August 2009 |title=Detecting Ancient Admixture and Estimating Demographic Parameters in Multiple Human Populations |pmid=19420049 |journal=Molecular Biology and Evolution |volume=26 |issue=8 |pages=1823–1827 |issn=0737-4038 |doi=10.1093/molbev/msp096 |pmc=2734152}}</ref>

Recent sequencing of Neanderthal<ref name="greenetal">{{cite journal |last1=Green |first1=R. E. |last2=Krause |first2=J. |last3=Briggs |first3=A. W. |last4=Maricic |first4=T. |last5=Stenzel |first5=U. |last6=Kircher |first6=M. |last7=Patterson |first7=N. |last8=Li |first8=H. |last9=Zhai |first9=W. |last10=Fritz |first10=M. H. Y. |last11=Hansen |first11=N. F. |last12=Durand |first12=E. Y. |last13=Malaspinas |first13=A. S. |last14=Jensen |first14=J. D. |last15=Marques-Bonet |first15=T. |last16=Alkan |first16=C. |last17=Prufer |first17=K. |last18=Meyer |first18=M. |last19=Burbano |first19=H. A. |last20=Good |first20=J. M. |last21=Schultz |first21=R. |last22=Aximu-Petri |first22=A. |last23=Butthof |first23=A. |last24=Höber |first24=B. |last25=Höffner |first25=B. |last26=Siegemund |first26=M. |last27=Weihmann |first27=A. |last28=Nusbaum |first28=C. |last29=Lander |first29=E. S. |last30=Russ |first30=C. |last31=Novod |first31=N. |last32=Affourtit |first32=J. |last33=Egholm |first33=M. |last34=Verna |first34=C. |last35=Rudan |first35=P. |last36=Brajkovic |first36=D. |last37=Kucan |first37=Z. |last38=Gusic |first38=I. |last39=Doronichev |first39=V. B. |last40=Golovanova |first40=L. V. |last41=Lalueza-Fox |first41=C. |last42=de la Rasilla |first42=M. |last43=Fortea |first43=J. |last44=Rosas |first44=A. |last45=Schmitz |first45=R. W. |last46=Johnson |first46=P. L. F. |last47=Eichler |first47=E. E. |last48=Falush |first48=D. |last49=Birney |first49=E. |last50=Mullikin |first50=J. C. |last51=Slatkin |first51=M. |last52=Nielsen |first52=R. |last53=Kelso |first53=J. |last54=Lachmann |first54=M. |last55=Reich |first55=D. |last56=Pääbo |first56=S. |display-authors=4 |date=May 7, 2010 |title=A Draft Sequence of the Neandertal Genome |journal=[[Science (journal)|Science]] |volume=328 |issue=5979 |pages=710–722 |bibcode=2010Sci...328..710G |doi=10.1126/science.1188021 |issn=0036-8075 |pmid=20448178 |pmc=5100745}}{{collapsible list |title=Full list of authors |bullets=true |Richard E. Green |Johannes Krause |Adrian W. Briggs |Tomislav Maricic |Udo Stenzel |Martin Kircher |Nick Patterson |Heng Li |Weiwei Zhai |Markus Hsi-Yang Fritz |Nancy F. Hansen |Eric Y. Durand |Anna-Sapfo Malaspinas |Jeffrey D. Jensen |Tomas Marques-Bonet |Can Alkan |Kay Prüfer |Matthias Meyer |Hernán A. Burbano |Jeffrey M. Good |Rigo Schultz |Ayinuer Aximu-Petri |Anne Butthof |Barbara Höber |Barbara Höffner |Madlen Siegemund |Antje Weihmann |Chad Nusbaum |Eric S. Lander |Carsten Russ |Nathaniel Novod |Jason Affourtit |Michael Egholm |Christine Verna |Pavao Rudan |Dejana Brajkovic |Željko Kucan |Ivan Gušic |Vladimir B. Doronichev |Liubov V. Golovanova |Carles Lalueza-Fox |Marco de la Rasilla |Javier Fortea |Antonio Rosas |Ralf W. Schmitz |Philip L. F. Johnson |Evan E. Eichler |Daniel Falush |Ewan Birney |James C. Mullikin |Montgomery Slatkin |Rasmus Nielsen |Janet Kelso |Michael Lachmann |David Reich |[[Svante Pääbo]]}}</ref> and Denisovan<ref name="pmid21179161" /> genomes shows that some admixture with these populations has occurred. All modern human groups outside Africa have 1–4% or (according to more recent research) about 1.5–2.6% Neanderthal [[allele]]s in their genome,<ref name="A high-coverage Neandertal genome f">{{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 (journal)|Science]] |date=2017 |doi=10.1126/science.aao1887 |pmid=28982794 |pmc=6185897 |display-authors=etal |volume=358 |issue=6363 |pages=655–658 |bibcode=2017Sci...358..655P}}</ref> and some [[Melanesians]] have an additional 4–6% of Denisovan alleles. These new results do not contradict the "out of Africa" model, except in its strictest interpretation, although they make the situation more complex. After recovery from a [[genetic bottleneck]] that some researchers speculate might be linked to the [[Toba catastrophe theory|Toba supervolcano catastrophe]], a fairly small group left Africa and interbred with Neanderthals, probably in the Middle East, on the Eurasian steppe or even in North Africa before their departure. Their still predominantly African descendants spread to populate the world. A fraction in turn interbred with Denisovans, probably in southeastern Asia, before populating Melanesia.<ref name="Reich_2011">{{cite journal |last1=Reich |first1=David |last2=Patterson |first2=Nick |last3=Kircher |first3=Martin |last4=Delfin |first4=Frederick |last5=Nandineni |first5=Madhusudan R. |last6=Pugach |first6=Irina |last7=Ko |first7=Albert Min-Shan |last8=Ko |first8=Ying-Chin |last9=Jinam |first9=Timothy A. |last10=Phipps |first10=Maude E. |last11=Saitou |first11=Naruya |last12=Wollstein |first12=Andreas |last13=Kayser |first13=Manfred |last14=Pääbo |first14=Svante |last15=Stoneking |first15=Mark |display-authors=3 |date=October 7, 2011 |title=Denisova Admixture and the First Modern Human Dispersals into Southeast Asia and Oceania |journal=American Journal of Human Genetics |volume=89 |issue=4 |pages=516–528 |doi=10.1016/j.ajhg.2011.09.005 |issn=0002-9297 |pmc=3188841 |pmid=21944045}}</ref> [[Human leukocyte antigen|HLA]] haplotypes of Neanderthal and Denisova origin have been identified in modern Eurasian and Oceanian populations.<ref name="10.1126/science.1209202" /> The Denisovan [[EPAS1]] gene has also been found in Tibetan populations.<ref>Huertha Sanchez, Emilia et al. (2014), "Altitude adaptation in Tibetans caused by introgression of Denisovan-like DNA" (Nature Vol 512, August 14, 2014)</ref> Studies of the human genome using machine learning have identified additional genetic contributions in Eurasians from an "unknown" ancestral population potentially related to the Neanderthal-Denisovan lineage.<ref>{{Cite web |url= https://www.sciencedaily.com/releases/2019/01/190116122650.htm |title=Artificial intelligence applied to the genome identifies an unknown human ancestor |website=[[ScienceDaily]] |access-date=January 17, 2019 |archive-date=January 16, 2019 |archive-url= https://web.archive.org/web/20190116213803/https://www.sciencedaily.com/releases/2019/01/190116122650.htm |url-status=live}}</ref>

[[File:Spreading homo sapiens la.svg|thumb|upright=1.5| A map of [[early human migrations]]]]

There are still differing theories on whether there was a single exodus from Africa or several. A multiple dispersal model involves the Southern Dispersal theory,<ref name="lahr">{{cite web |url= http://www.human-evol.cam.ac.uk/Projects/sdispersal/sdispersal.htm |title=Searching for traces of the Southern Dispersal |last1=Lahr |first1=Marta Mirazón |author1-link=Marta Mirazón Lahr |last2=Petraglia |first2=Mike |last3=Stokes |first3=Stephen |last4=Duller |first4=Geoff |display-authors=3 |website=Leverhulme Centre for Human Evolutionary Studies |publisher=[[University of Cambridge]] |archive-url= https://web.archive.org/web/20120510133600/http://www.human-evol.cam.ac.uk/Projects/sdispersal/sdispersal.htm |archive-date=May 10, 2012 |access-date=April 21, 2015}}</ref><ref>Simpson, Colin. [https://www.thenational.ae/uae/science/dna-evidence-suggests-the-whole-world-is-a-little-bit-arab-1.408619 DNA evidence suggests the whole world is a little bit Arab]. [[The National (Abu Dhabi)|The National]]. 16, June 2020. Archived at [https://web.archive.org/web/20200129100200/https://www.thenational.ae/uae/science/dna-evidence-suggests-the-whole-world-is-a-little-bit-arab-1.408619 The Wayback Machine]</ref><ref>{{cite journal |last1=Fernandes |first1=V. |last2=Alshamali |first2=F. |last3=Alves |first3=M. |display-authors=etal |title=The Arabian cradle: Mitochondrial relicts of the first steps along the southern route out of Africa |journal=[[American Journal of Human Genetics]] |date=2012 |volume=90 |issue=2 |pages=347–355 |doi=10.1016/j.ajhg.2011.12.010 |pmid=22284828 |pmc=3276663 }}</ref> which has gained support in recent years from genetic, linguistic and archaeological evidence. In this theory, there was a coastal dispersal of modern humans from the [[Horn of Africa]] crossing the Bab el Mandib to Yemen at a lower sea level around 70,000 years ago. This group helped to populate Southeast Asia and Oceania, explaining the discovery of early human sites in these areas much earlier than those in the [[Levant]].<ref name="lahr" /> This group seems to have been dependent upon marine resources for their survival.

[[Stephen Oppenheimer]] has proposed a second wave of humans may have later dispersed through the Persian Gulf oases, and the Zagros mountains into the Middle East. Alternatively it may have come across the [[Sinai Peninsula]] into Asia, from shortly after 50,000 yrs BP, resulting in the bulk of the human populations of Eurasia. It has been suggested that this second group possibly possessed a more sophisticated "big game hunting" tool technology and was less dependent on coastal food sources than the original group. Much of the evidence for the first group's expansion would have been destroyed by the rising sea levels at the end of each [[glacial maximum]].<ref name="lahr" /> The multiple dispersal model is contradicted by studies indicating that the populations of Eurasia and the populations of Southeast Asia and Oceania are all descended from the same mitochondrial DNA L3 [[Lineage (genetic)|lineages]], which support a single migration out of Africa that gave rise to all non-African populations.<ref>{{cite journal |last1=Macaulay |first1=Vincent |last2=Hill |first2=Catherine |last3=Achilli |first3=Alessandro |last4=Rengo |first4=Chiara |last5=Clarke |first5=D. |last6=Meehan |first6=W. |last7=Blackburn |first7=J. |last8=Semino |first8=O. |last9=Scozzari |first9=R. |last10=Cruciani |first10=F. |last11=Taha |first11=A. |last12=Shaari |first12=N. K. |last13=Raja |first13=J. M. |last14=Ismail |first14=P. |last15=Zainuddin |first15=Z. |last16=Goodwin |first16=W. |last17=Bulbeck |first17=D. |last18=Bandelt |first18=H. J. |last19=Oppenheimer |first19=S. |last20=Torroni |first20=A. |last21=Richards |first21=M. |display-authors=3 |date=May 13, 2005 |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 |bibcode=2005Sci...308.1034M |doi=10.1126/science.1109792 |issn=0036-8075 |pmid=15890885 |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 |url-status=live |access-date=August 20, 2019 |archive-url= https://web.archive.org/web/20220828110206/http://psasir.upm.edu.my/id/eprint/40255/1/Single%2C%20rapid%20coastal%20settlement%20of%20Asia%20revealed%20by%20analysis%20of%20complete%20mitochondrial%20genomes.pdf |archive-date=August 28, 2022}}</ref>

On the basis of the early date of Badoshan Iranian Aurignacian, Oppenheimer suggests that this second dispersal may have occurred with a pluvial period about 50,000 years before the present, with modern human big-game hunting cultures spreading up the Zagros Mountains, carrying modern human genomes from Oman, throughout the Persian Gulf, northward into Armenia and Anatolia, with a variant travelling south into Israel and to Cyrenicia.<ref name="oppenheimer">{{cite book |last=Oppenheimer |first=Stephen |date=2012 |title=Out of Eden: The Peopling of the World |publisher=Robinson |edition=New}}</ref>

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