Editing Neanderthal extinction (section)

==Upper Palaeolithic Transition==
[[File:SpanienEbro.jpg|thumb|Map emphasising the [[Ebro River]] in northern Spain|alt=Black and white satellite image of the Iberian Peninsula, but the Ebro River valley at the Spain/France border uses red to blue colours to indicate topography and elevation]]
The extinction of Neanderthals was part of the broader [[Late Pleistocene extinctions|Late Pleistocene megafaunal extinction event]].<ref>{{Cite journal |last1=Hortolà |first1=Policarp |last2=Martínez-Navarro |first2=Bienvenido |date=May 8, 2013 |title=The Quaternary megafaunal extinction and the fate of Neanderthals: An integrative working hypothesis |url=https://www.sciencedirect.com/science/article/pii/S1040618212001188 |journal=[[Quaternary International]] |series=East meets West: First settlements and human evolution in Eurasia |volume=295 |pages=69–72 |doi=10.1016/j.quaint.2012.02.037 |bibcode=2013QuInt.295...69H |issn=1040-6182 |access-date=February 5, 2024 |via=Elsevier Science Direct|url-access=subscription }}</ref> Whatever the cause of their extinction, Neanderthals were replaced by modern humans, indicated by near full replacement of Middle Palaeolithic Mousterian stone technology with modern human Upper Palaeolithic Aurignacian stone technology across Europe (the Middle-to-Upper Palaeolithic Transition) from 41,000 to 39,000 years ago.<ref name="higham2014">{{cite journal |last1=Higham |first1=T. |last2=Douka |first2=K. |last3=Wood |first3=R. |last4=Ramsey |first4=C. B. |last5=Brock |first5=F. |last6=Basell |first6=L. |last7=Camps |first7=M. |last8=Arrizabalaga |first8=A. |last9=Baena |first9=J. |last10=Barroso-Ruíz |first10=C. |author11=C. Bergman |author12=C. Boitard |author13=P. Boscato |author14=M. Caparrós |author15=N.J. Conard |display-authors=etal |year=2014 |title=The timing and spatiotemporal patterning of Neanderthal disappearance |journal=[[Nature (journal)|Nature]] |volume=512 |issue=7514 |pages=306–309 |bibcode=2014Natur.512..306H |doi=10.1038/nature13621 |pmid=25143113 |s2cid=205239973 |hdl-access=free |author16=C. Draily |author17=A. Froment |author18=B. Galván |author19=P. Gambassini |author20=A. Garcia-Moreno |author21=S. Grimaldi |author22=P. Haesaerts |author23=B. Holt |author24=M.-J. Iriarte-Chiapusso |author25=A. Jelinek |author26=J.F. Jordá Pardo |author27=J.-M. Maíllo-Fernández |author28=A. Marom |author29=J. Maroto |author30=M. Menéndez |author31=L. Metz |author32=E. Morin |author33=A. Moroni |author34=F. Negrino |author35=E. Panagopoulou |author36=M. Peresani |author37=S. Pirson |author38=M. de la Rasilla |author39=J. Riel-Salvatore |author40=A. Ronchitelli |author41=D. Santamaria |author42=P. Semal |author43=L. Slimak |author44=J. Soler |author45=N. Soler |author46=A. Villaluenga |author47=R. Pinhasi |author48=R. Jacobi |quotation=We show that the Mousterian [the Neanderthal tool-making tradition] ended by 41,030–39,260 calibrated years BP (at 95.4% probability) across Europe. We also demonstrate that succeeding 'transitional' archaeological industries, one of which has been linked with Neanderthals (Châtelperronian), end at a similar time. |hdl=1885/75138}}</ref><ref name=higham2011>{{cite journal |first=T. |last=Higham |year=2011 |title=European Middle and Upper Palaeolithic radiocarbon dates are often older than they look: problems with previous dates and some remedies |journal=Antiquity |volume=85 |issue=327 |pages=235–249 |doi=10.1017/s0003598x00067570 |s2cid=163207571 |quotation=Few events of European prehistory are more important than the transition from ancient to modern humans about 40,000 years ago, a period that unfortunately lies near the limit of radiocarbon dating. This paper shows that as many as 70 per cent of the oldest radiocarbon dates in the literature may be too young, due to contamination by modern carbon.}}</ref><ref name="galvan2014">{{cite journal |last1=Galván |first1=B. |last2=Hernández |first2=C. M. |last3=Mallol |first3=C. |last4=Mercier |first4=N. |last5=Sistiaga |first5=A. |last6=Soler |first6=V. |year=2014 |title=New evidence of early Neanderthal disappearance in the Iberian Peninsula |journal=[[Journal of Human Evolution]] |volume=75 |pages=16–27 |doi=10.1016/j.jhevol.2014.06.002 |pmid=25016565|bibcode=2014JHumE..75...16G }}</ref><ref>{{Cite journal |last1=Agusti |first1=J. |last2=Rubio-Campillo |first2=X. |year=2017 |title=Were Neanderthals responsible for their own extinction? |journal=[[Quaternary International]] |volume=431 |pages=232–237 |bibcode=2017QuInt.431..232A |doi=10.1016/j.quaint.2016.02.017}}</ref> By between 44,200 and 40,600 BP, Neanderthals vanished from northwestern Europe.<ref>{{Cite journal |last1=Devièse |first1=Thibaut |last2=Abrams |first2=Grégory |last3=Hajdinjak |first3=Mateja |last4=Pirson |first4=Stéphane |last5=De Groote |first5=Isabelle |last6=Di Modica |first6=Kévin |last7=Toussaint |first7=Michel |last8=Fischer |first8=Valentin |last9=Comeskey |first9=Dan |last10=Spindler |first10=Luke |last11=Meyer |first11=Matthias |last12=Semal |first12=Patrick |last13=Higham |first13=Tom |date=March 23, 2021 |title=Reevaluating the timing of Neanderthal disappearance in Northwest Europe |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |language=en |volume=118 |issue=12 |doi=10.1073/pnas.2022466118 |doi-access=free |issn=0027-8424 |pmc=7999949 |pmid=33798098 |bibcode=2021PNAS..11822466D }}</ref> However, it is postulated that Iberian Neanderthals persisted until about 35,000 years ago, as indicated by the date range of transitional lithic assemblages—Châtelperronian, Uluzzian, [[Protoaurignacian]] and Early Aurignacian. The latter two are attributed to modern humans, but the former two have unconfirmed authorship, potentially products of Neanderthal/modern human cohabitation and cultural transmission. Further, the appearance of the Aurignacian south of the [[Ebro River]] has been dated to roughly 37,500 years ago, which has prompted the "Ebro Frontier" hypothesis which states that the river presented a geographic barrier preventing modern human immigration, and thus prolonging Neanderthal persistence.<ref name=Zilhao2017>{{cite journal |first1=J. |last1=Zilhão |first2=D. |last2=Anesin |first3=T. |last3=Aubry |display-authors=et al. |year=2017 |title=Precise dating of the Middle-to-Upper Paleolithic transition in Murcia (Spain) supports late Neandertal persistence in Iberia |journal=Heliyon |volume=3 |issue=1 |pages=E00435 |doi=10.1016/j.heliyon.2017.e00435 |doi-access=free |pmid=29188235 |pmc=5696381|bibcode=2017Heliy...300435Z }}</ref><ref name=Morales2019>{{cite journal |first1=J. I |last1=Morales |first2=A. |last2=Cebrià |first3=A. |last3=Burguet-Coca |display-authors=et al. |year=2019 |title=The Middle-to-Upper Paleolithic transition occupations from Cova Foradada (Calafell, NE Iberia) |journal=PLOS ONE |volume=14 |issue=5 |pages=e0215832 |doi=10.1371/journal.pone.0215832 |pmc=6522054 |pmid=31095578 |bibcode=2019PLoSO..1415832M|doi-access=free }}</ref> However, the dating of the Iberian Transition is debated, with a contested timing of 43,000–40,800 years ago at Cueva Bajondillo, Spain.<ref>{{cite journal |last1=Cortés-Sánches |first1=M. |last2=Jiménez-Espejo |first2=F. J. |last3=Simón-Vallejo |first3=M. D. |last4=Stringer |first4=C. |author4-link=Chris Stringer |display-authors=et al. |year=2019 |title=An early Aurignacian arrival in southwestern Europe |journal=[[Nature Ecology & Evolution]] |volume=3 |issue=2 |pages=207–212 |doi=10.1038/s41559-018-0753-6 |pmid=30664696 |bibcode=2019NatEE...3..207C |s2cid=58571988}}</ref><ref>{{cite journal |first1=L. |last1=Anderson |first2=N. |last2=Reynolds |first3=N. |last3=Teyssandier |s2cid=115153460 |year=2019 |title=No reliable evidence for a very early Aurignacian in Southern Iberia |journal=Nature Ecology and Evolution |volume=3 |issue=5 |page=713 |doi=10.1038/s41559-019-0885-3 |pmid=30988496 |doi-access=free|bibcode=2019NatEE...3..713A }}</ref><ref>{{cite journal |last=de la Peña |first=P. |year=2019 |title=Dating on its own cannot resolve hominin occupation patterns |journal=[[Nature Ecology and Evolution]] |volume=3 |issue=5 |page=712 |doi=10.1038/s41559-019-0886-2 |pmid=30988497 |s2cid=116861216 |doi-access=free|bibcode=2019NatEE...3..712D }}</ref><ref>{{cite journal |first1=M. |last1=Cortés-Sánches |first2=F. J. |last2=Jiménez-Espejo |first3=M. D. |last3=Simón-Vallejo |first4=C. |last4=Stringer |s2cid=115153407 |author4-link=Chris Stringer |display-authors=et al. |year=2019 |title=Reply to 'Dating on its own cannot resolve hominin occupation patterns' and 'No reliable evidence for a very early Aurignacian in Southern Iberia' |journal=Nature Ecology and Evolution |volume=3 |issue=5 |pages=714–715 |doi=10.1038/s41559-019-0887-1 |pmid=30988498|bibcode=2019NatEE...3..714C }}</ref> The Châtelperronian appears in northeastern Iberia about 42,500–41,600 years ago.<ref name=Zilhao2017/>

Some [[Neanderthals in Gibraltar]] were dated to much later than this—such as Zafarraya (30,000 years ago)<ref>{{cite journal |first1=M. |last1=Fontugne |first2=J. L. |last2=Reyss |first3=C. B. |last3=Ruis |first4=P. M. |last4=Lara |year=1995 |title=The Mousterian site of Zafarraya (Granada, Spain): dating and implications on the palaeolithic peopling processes of Western Europe |journal=Comptes Rendus de l'Académie des Sciences |volume=321 |issue=10 |pages=931–937}}</ref> and Gorham's Cave (28,000 years ago)<ref>{{cite journal |first1=C. |last1=Finlayson |author1-link=Clive Finlayson |first2=F. G. |last2=Pacheco |s2cid=4411186 |year=2006 |title=Late survival of Neanderthals at the southernmost extreme of Europe |journal=Nature |volume=443 |issue=7,113 |pages=850–853 |doi=10.1038/nature05195 |pmid=16971951 |bibcode=2006Natur.443..850F}}</ref>—which may be inaccurate as they were based on ambiguous artefacts instead of direct dating.<ref name=galvan2014/> A claim of Neanderthals surviving in a polar refuge in the [[Ural Mountains]]<ref name=slimak2011>{{cite journal |first1=L. |last1=Slimak |first2=J. I. |last2=Svendsen |first3=J. |last3=Mangerud |first4=H. |last4=Plisson |year=2011 |title=Late Mousterian persistence near the Arctic Circle |journal=Science |volume=332 |issue=6031 |pages=841–845 |doi=10.1126/science.1203866 |pmid=21566192 |jstor=29784275 |bibcode=2011Sci...332..841S |s2cid=24688365}}</ref> is loosely supported by Mousterian stone tools dating to 34,000 years ago from the northern Siberian Byzovaya site at a time when modern humans may not yet have colonised the northern reaches of Europe;<ref name=zwyns2012>{{cite journal |last=Zwyns |first=N. |year=2012 |title=Comment on Late Mousterian persistence near the Arctic Circle |journal=Science |volume=335 |issue=6065 |page=167 |doi=10.1126/science.1209908 |pmid=22246757 |bibcode=2012Sci...335..167Z |doi-access=free}}</ref> however, modern human remains are known from the nearby [[Mamontovaya Kurya]] site dating to 40,000 years ago.<ref>{{cite journal |doi=10.1038/35092552 |pmid=11544525 |title=Human presence in the European Arctic nearly 40,000 years ago |journal=Nature |volume=413 |issue=6,851 |pages=64–67 |year=2001 |last1=Pavlov |first1=P. |last2=Svendsen |first2=J. I. |last3=Indrelid |first3=S. |s2cid=1986562 |bibcode=2001Natur.413...64P}}</ref> Indirect dating of Neanderthals remains from [[Mezmaiskaya Cave]] reported a date of about 30,000 years ago, but direct dating instead yielded 39,700&thinsp;±1,100 years ago, more in line with trends exhibited in the rest of Europe.<ref name="pinhasi2011">{{cite journal |first1=R. |last1=Pinhasi |first2=T. F. G. |last2=Higham |first3=L. V. |last3=Golovanova |first4=V. B. |last4=Doronichev |year=2011 |title=Revised age of late Neanderthal occupation and the end of the Middle Palaeolithic in the northern Caucasus |journal=Proceedings of the National Academy of Sciences |volume=108 |issue=21 |pages=8611–8616 |quotation=The direct date of the fossil (39,700±1,100 <sup>14</sup>C BP) is in good agreement with the probability distribution function, indicating at a high level of probability that Neanderthals did not survive at Mezmaiskaya Cave after 39 kya cal BP. [...] This challenges previous claims for late Neanderthal survival in the northern Caucasus. [...] Our results confirm the lack of reliably dated Neanderthal fossils younger than ≈40 kya cal BP in any other region of Western Eurasia, including the Caucasus. |doi=10.1073/pnas.1018938108 |pmid=21555570 |bibcode=2011PNAS..108.8611P |pmc=3102382|doi-access=free }}</ref>

[[File:Bohunician scrapers.jpg|thumb|upright=1.35|[[Bohunician]] [[scraper (archaeology)|scrapers]] in the [[Moravian Museum]], Czech Republic]]
The earliest indication of Upper Palaeolithic modern human immigration into Europe is a series of modern human teeth with Neronian [[Industry (archaeology)|industry]] stone tools found at [[Mandrin Cave]], [[Malataverne]] in France, dated in 2022 to between 56,800 and 51,700 years ago.<ref>{{cite journal|first1=L.|last1=Slimak|first2=C.|last2=Zanolli|first3=T.|last3=Higham|display-authors=et al.|year=2022|title=Modern human incursion into Neanderthal territories 54,000 years ago at Mandrin, France|journal=Science Advances|volume=8|issue=6|pages=eabj9496 |doi=10.1126/sciadv.abj9496|pmid=35138885 |pmc=8827661 |bibcode=2022SciA....8J9496S }}</ref> The earliest bones in Europe date to roughly 45–43,000 years ago in Bulgaria,<ref>{{cite journal |last1=Hublin |first1=J.-J. |last2=Sirakov |first2=N. |display-authors=et al. |year=2020 |title=Initial Upper Palaeolithic ''Homo sapiens'' from Bacho Kiro Cave, Bulgaria |url=https://kar.kent.ac.uk/81524/1/Hublin%20et%20al%20_authors%20accepted.pdf |journal=[[Nature (journal)|Nature]] |volume=581 |issue=7808 |pages=299–302 |bibcode=2020Natur.581..299H |doi=10.1038/s41586-020-2259-z |pmid=32433609 |s2cid=218592678 |hdl=11585/770553}}</ref> Italy,<ref>{{Cite journal |last1=Benazzi |first1=S. |last2=Douka |first2=K. |last3=Fornai |first3=C. |last4=Bauer |first4=C.C. |last5=Kullmer |first5=O. |last6=Svoboda |first6=J.Í. |last7=Pap |first7=I. |last8=Mallegni |first8=F. |last9=Bayle |first9=P. |last10=Coquerelle |first10=M. |last11=Condemi |first11=S. |last12=Ronchitelli |first12=A. |last13=Harvati |first13=K. |last14=Weber |first14=G.W. |year=2011 |title=Early dispersal of modern humans in Europe and implications for Neanderthal behaviour |journal=[[Nature (journal)|Nature]] |volume=479 |issue=7374 |pages=525–528 |bibcode=2011Natur.479..525B |doi=10.1038/nature10617 |pmid=22048311 |s2cid=205226924}}</ref> and Britain.<ref>{{Cite journal |last1=Higham |first1=T. |last2=Compton |first2=T. |last3=Stringer |first3=C. |last4=Jacobi |first4=R. |last5=Shapiro |first5=B. |last6=Trinkaus |first6=E. |last7=Chandler |first7=B. |last8=Gröning |first8=F. |last9=Collins |first9=C. |last10=Hillson |first10=S. |last11=o’Higgins |first11=P. |last12=Fitzgerald |first12=C. |last13=Fagan |first13=M. |year=2011 |title=The earliest evidence for anatomically modern humans in northwestern Europe |journal=[[Nature (journal)|Nature]] |volume=479 |issue=7374 |pages=521–524 |bibcode=2011Natur.479..521H |doi=10.1038/nature10484 |pmid=22048314 |s2cid=4374023}}</ref> This wave of modern humans replaced Neanderthals.<ref name=higham2014/> However, Neanderthals and ''H. sapiens'' have a much longer contact history. DNA evidence indicates ''H. sapiens'' contact with Neanderthals and admixture as early as 120–100,000 years ago. A 2019 reanalysis of 210,000-year-old skull fragments from the Greek [[Apidima Cave]] assumed to have belonged to a Neanderthal concluded that they belonged to a modern human, and a Neanderthal skull dating to 170,000 years ago from the cave indicates ''H. sapiens'' were replaced by Neanderthals until returning about 40,000 years ago.<ref name="NAT-20190710">{{cite journal |last=Harvati |first=K. |display-authors=et al. |title=Apidima Cave fossils provide earliest evidence of ''Homo sapiens'' in Eurasia |year=2019 |journal=Nature |volume=571 |issue=7766 |pages=500–504 |doi=10.1038/s41586-019-1376-z |pmid=31292546 |s2cid=195873640|url=https://zenodo.org/record/6646855 }}</ref> This identification was refuted by a 2020 study.<ref>{{cite journal |first1=M.-A. |last1=de Lumley |first2=G. |last2=Guipert |first3=H. |last3=de Lumley |author3-link=Henry de Lumley |first4=N. |last4=Protopapa |first5=T. |last5=Pitsios |year=2020 |title=Apidima 1 and Apidima 2: two anteneandertal skulls in the Peloponnese, Greece |journal=L'Anthropologie |volume=124 |issue=1 |page=102743 |doi=10.1016/j.anthro.2019.102743|s2cid=216449347 }}</ref> Archaeological evidence suggests that Neanderthals displaced modern humans in the Near East around 100,000 years ago until about 60–50,000 years ago.{{sfn|Reich|2018}}