Editing Early modern human (section)

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

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

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

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

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

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

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

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

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

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

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

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

Many of the early modern human finds, like those of [[Jebel Irhoud]], [[Omo remains|Omo]], [[Homo sapiens idaltu|Herto]], [[Florisbad Skull|Florisbad]], [[Skhul remains|Skhul]], and [[Peștera cu Oase]] exhibit a mix of archaic and modern traits.<ref name="Oppenheimer">{{cite book |last=Oppenheimer |first=S. |title=Out of Eden: The Peopling of the World |year=2003 |publisher=Robinson |isbn=978-1841196978}}</ref><ref>{{Cite journal|doi=10.1073/pnas.2035108100 |last1=Trinkaus |first1=E. |last2=Moldovan |first2=O. |last3=Milota |first3=Ș. |last4=Bîlgăr |first4=A. |last5=Sarcina |first5=L. |last6=Athreya |first6=S. |last7=Bailey |first7=S. E. |last8=Rodrigo |first8=R. |last9=Gherase |first9=M. |last10=Higham |first10=T. |last11=Ramsey |first11=C. B. |last12=Van Der Plicht |first12=J. |year=2003 |title=An early modern human from Peștera cu Oase, Romania |journal=PNAS |volume=100 |issue=20|pages=11231–11236 |pmid=14504393 |pmc=208740 |bibcode=2003PNAS..10011231T |display-authors=8 |doi-access=free }}</ref><ref name="NAT-20170607a">{{cite journal |last=Callaway |first=Ewan |title=Oldest Homo sapiens fossil claim rewrites our species' history |url= http://www.nature.com/news/oldest-homo-sapiens-fossil-claim-rewrites-our-species-history-1.22114 |date=7 June 2017 |journal=[[Nature (journal)|Nature]] |doi=10.1038/nature.2017.22114 |access-date=11 June 2017 }}</ref> Skhul V, for example, has prominent brow ridges and a projecting face. However, the [[Neurocranium|brain case]] is quite rounded and distinct from that of the Neanderthals and is similar to the brain case of modern humans. It is uncertain whether the robust traits of some of the early modern humans like Skhul V reflects [[Archaic human admixture with modern humans|mixed ancestry]] or retention of older traits.<ref name="Reich et al.">{{Cite journal|first1=David |last1=Reich |first2=Richard E. |last2=Green |first3=Martin |last3=Kircher |first4=Johannes |last4=Krause |first5=Nick |last5=Patterson |first6=Eric Y. |last6=Durand |first7=Bence |last7=Viola |first8=Adrian W. |last8=Briggs |first9=Udo |last9=Stenzel |first10=Philip L. F. |last10=Johnson |first11=Tomislav |last11=Maricic |first12=Jeffrey M. |last12=Good |first13=Tomas |last13=marques-Bonet |first14=Can |last14=Alkan |first15=Qiaomei |last15=Fu |first16=Swapan |last16=Mallick |first17=Heng |last17=Li |first18=Matthias |last18=Meyer |first19=Evan E. |last19=Eichler |first20=Mark |last20=Stoneking |first21=Michael |last21=Richards |first22=Sahra |last22=Talamo |first23=Michael V. |last23=Shunkov |first24=Anatoli P. |last24=Derevianko |first25=Jean-Jacques |last25=Hublin |first26=Janet |last26=Kelso |first27=Montgomery |last27=Slatkin |name-list-style=amp|first28=Svante |last28=Pääbo |display-authors=8 |year=2010 |title=Genetic history of an archaic hominin group from Denisova Cave in Siberia |journal=[[Nature (journal)|Nature]] |volume=468 |issue=7327 |pages=1053–1060 |doi=10.1038/nature09710 |pmid=21179161|bibcode= 2010Natur.468.1053R |pmc=4306417 |hdl=10230/25596}}</ref><ref>{{cite journal|last=Trinkaus |first=Erik |title=Early modern humans |journal=Annual Review of Anthropology |date=October 2005 |volume=34 |issue=1 |pages=207–230 |doi=10.1146/annurev.anthro.34.030905.154913|s2cid=9039428 |s2cid-access=free}}</ref>

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

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

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

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