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{{Redirect|Neandertal|the location in Germany|Neandertal (valley)}}{{Redirect|Neanderthal Man|the 1970 single by English pop rock band Hotlegs|Neanderthal Man (song)}}{{short description|Extinct species of the genus Homo}}{{Use mdy dates|date=October 2018}}{{Speciesbox| taxon = Homo neanderthalensis| extinct = yes| name = Neanderthal< 35 PeÈ™tera cu Oase 37-42 ka, Mladeč 31 ka, PeÈ™tera Muierii 30 ka (n/s),NEWS, Jacqui, Hayes, November 2, 2006, Humans and Neanderthals interbred,weblink Cosmos, May 17, 2009, dead,weblink" title="">weblink February 22, 2009, Lagar Velho 1 24.5 ka.

In popular culture

Neanderthals have been portrayed in popular culture including appearances in literature, visual media and comedy. Early 20th century artistic interpretations often presented Neanderthals as beastly creatures, emphasising hairiness and a rough, dark complexion.Neanderthal image by Kupka, based on Boule, 1909, in Humanity's Journeys Dr. Kathryn Denning, 2005. Retrieved March 17, 2012.

See also

{{col div}}
  • {{annotated link|Homo floresiensis}}
  • {{annotated link|Homo luzonensis}}
  • {{annotated link|Denisovan}}
  • {{annotated link|Early human migrations}}
  • {{annotated link|List of Neanderthal sites}}
  • {{annotated link|Neanderthals in Gibraltar}}




  • JOURNAL, 10.1006/jpho.2002.0170, The potential Neandertal vowel space was as large as that of modern humans, 2002, Boë, Louis-Jean, Heim, Jean-Louis, Honda, Kiyoshi, Maeda, Shinji, Journal of Phonetics, 30, 3, 465–84,
  • JOURNAL, 10.1016/j.wocn.2005.07.002, Current views on Neanderthal speech capabilities: A reply to Boe et al. (2002), October 2007, Philip, Lieberman, Journal of Phonetics, 35, 4, 552–63,
  • JOURNAL, 10.1371/journal.pbio.0020057, No Evidence of Neandertal mtDNA Contribution to Early Modern Humans, 2004, Serre, David, Langaney, André, Chech, Mario, Teschler-Nicola, Maria, Paunovic, Maja, Mennecier, Philippe, Hofreiter, Michael, Possnert, Göran, Pääbo, Svante, PLoS Biology, 2, 3, e57, 15024415, 368159,
  • JOURNAL, 10.1038/nature03585, Direct dating of Early Upper Palaeolithic human remains from Mladeč, 2005, Wild, Eva M., Teschler-Nicola, Maria, Kutschera, Walter, Steier, Peter, Trinkaus, Erik, Wanek, Wolfgang, Nature, 435, 7040, 332–35, 15902255, 2005Natur.435..332W,
  • JOURNAL, 10.1371/journal.pone.0008880,weblink ScienceDaily, January 27, 2010, Pego do Diabo (Loures, Portugal): Dating the Emergence of Anatomical Modernity in Westernmost Eurasia, 2010, Hawks, John, Zilhão, João, Davis, Simon J. M., Duarte, Cidália, Soares, António M. M., Steier, Peter, Wild, Eva, PLOS ONE, 5, e8880, 20111705, 1, 2811729, 2010PLoSO...5.8880Z,
  • BOOK, Anatoliy P., Derev'anko, William Roger, Powers, Demitri Boris, Shimkin, The Paleolithic of Siberia: new discoveries and interpretations, Institute of Anthropology and Ethnography, Novosibirsk, 1998, 978-0-252-02052-0, 36461622,
  • BOOK, Jonathan I., Lunine, Earth: Evolution of a Habitable World, Cambridge University Press, 2013, 978-0-521-85001-8, 327,weblink harv,

Further reading

  • BOOK, Who We Are And How We Got Here – Ancient DNA and the New Science of the Human Past, Who We Are and How We Got Here, David, Reich, David Reich (geneticist), Pantheon Books, 2018, 978-1101870327,
  • NEWS, Diamond, Jared, A Brand-New Version of Our Origin Story,weblink April 20, 2018, The New York Times, April 23, 2018,

External links

{{Commons category|Homo neanderthalensis}}{{Wikispecies|Homo neanderthalensis}}{{NIE Poster|Neanderthal Man}}
  • WEB

, C. David, Kreger, June 30, 2000
,weblink Homo neanderthalensis,, May 23, 2009,
  • WEB

, Dennis, O'Neil, May 12, 2009
,weblink Evolution of Modern Humans: Neandertals, May 23, 2009,

, Homo neanderthalensis
, The Smithsonian Institution
,weblink 2010-02-14
  • WEB

, Neanderthal DNA
, International Society of Genetic Genealogy
, dead
,weblink" title="">weblink
, June 17, 2006
, : Includes Neanderthal mtDNA sequences {{Prehistoric technology}}{{Human Evolution}}{{Homo neanderthalensis|state=expanded}}{{Taxonbar|from=Q40171}}{{Authority control}}

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Eastern Philosophy
History of Philosophy
M.R.M. Parrott
Middle Pleistocene>Middle–Late Pleistocene {{Fossil range0.04}} | image= Homo sapiens neanderthalensis.jpg| image_caption = Late Neanderthal skull(La Chapelle-aux-Saints 1)| image2 = Neanderthalensis.jpg| image2_upright = 0.4| image2_caption = An approximate reconstruction of a Neanderthal skeleton. The central rib cage, including the sternum, and parts of the pelvis are from modern humans.William King (geologist)>King, 1864Homo stupidus (Ernst Haeckel>Haeckel 1866)Homo mousteriensis (Hermann Klaatsch 1909)ROMEOTITLE=ECCE HOMO!:A LEXICON OF MANLOCATION=AMSTERDAMPAGE=92ISBN=978-9027220066, Palaeoanthropus neanderthalensisCAMP>FIRST1=C. LFIRST2=H. J.FIRST3=R. H.PUBLISHER=THE GEOLOGICAL SOCIETY OF AMERICA, INC.YEAR=1964URL=HTTPS://BOOKS.GOOGLE.COM/BOOKS?ID=MR_CXCXO8H8C&PG=PA556, 9780813710921, | range_map = Range_of_NeanderthalsAColoured.pngEurope (blue), Neanderthals in Southwest Asia>Southwest Asia (orange), Uzbekistan (green), and the Altai Mountains (violet).}}Neanderthals ({{IPAc-en||n|i|ˈ|æ|n|d|É™r|t|ɑː|l|,_|n|eɪ|-|,_|-|θ|ɔː|l}};BOOK, Wells, John, John C. Wells, Longman Pronunciation Dictionary, Pearson Longman, 3rd, 3 April 2008, 978-1-4058-8118-0, {{IPA-de|neˈ(Ê”)andɐtaːl|lang}};{{Citation|last1=Krech|first1=Eva Maria|last2=Stock|first2=Eberhard|last3=Hirschfeld|first3=Ursula|last4=Anders|first4=Lutz-Christian|year=2009|title=Deutsches Aussprachewörterbuch|location=Berlin, New York|publisher=Walter de Gruyter|isbn=978-3-11-018202-6}}BOOK,weblink 561, Duden - Das Aussprachewörterbuch: Betonung und Aussprache von über 132.000 Wörtern und Namen, de, 2015, Bibliographisches Institut GmbH, 9783411911516, Homo neanderthalensis or Homo sapiens neanderthalensis)BOOK, Pääbo, Svante, Neanderthal Man: In Search of Lost Genomes, Basic Books, New York, 2014, 237, are an extinct species or subspecies of archaic humans in the genus Homo, who lived within Eurasia from circa 400,000 until 40,000 years ago.JOURNAL, T. Higham, K. Douka, R. Wood, C.B. Ramsey, F. Brock, L. Basell, M. Camps, A. Arrizabalaga, J. Baena, C. Barroso-Ruíz, C. Bergman, C. Boitard, P. Boscato, M. Caparrós, N.J. Conard, C. Draily, A. Froment, B. Galván, P. Gambassini, A. Garcia-Moreno, S. Grimaldi, P. Haesaerts, B. Holt, M.-J. Iriarte-Chiapusso, A. Jelinek, J.F. Jordá Pardo, J.-M. Maíllo-Fernández, A. Marom, J. Maroto, M. Menéndez, L. Metz, E. Morin, A. Moroni, F. Negrino, E. Panagopoulou, M. Peresani, S. Pirson, M. de la Rasilla, J. Riel-Salvatore, A. Ronchitelli, D. Santamaria, P. Semal, L. Slimak, J. Soler, N. Soler, A. Villaluenga, R. Pinhasi, R. Jacobi, 2014, The timing and spatiotemporal patterning of Neanderthal disappearance, Nature, 512, 7514, 306–09,weblink 10.1038/nature13621, 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., 25143113, 2014Natur.512..306H, JOURNAL, T. Higham, 2011, European Middle and Upper Palaeolithic radiocarbon dates are often older than they look: problems with previous dates and some remedies, Antiquity, 85, 327, 235–49,weblink 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., 10.1017/s0003598x00067570, JOURNAL, R. Pinhasi, T.F.G. Higham, L.V. Golovanova, V.B. Doronichev, 2011, Revised age of late Neanderthal occupation and the end of the Middle Paleolithic in the northern Caucasus, Proceedings of the National Academy of Sciences USA, 108, 21, 8611–16, The direct date of the fossil (39,700 ± 1,100 14C 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 ka 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 ka cal BP in any other region of Western Eurasia, including the Caucasus., 10.1073/pnas.1018938108, 21555570, 2011PNAS..108.8611P, 3102382, JOURNAL, B. Galván, C.M. Hernández, C. Mallol, N. Mercier, A. Sistiaga, V. Soler, 2014, New evidence of early Neanderthal disappearance in the Iberian Peninsula, Journal of Human Evolution, 75, 16–27,weblink 10.1016/j.jhevol.2014.06.002, 25016565, NEWS, McKie, Robin, June 2, 2013, Why did the Neanderthals die out?,weblink The Guardian, April 6, 2017, "It was once thought we appeared in Europe around 35,000 years ago and that we coexisted with Neanderthals for thousands of years after that. They may have hung on in pockets â€“ including caves in Gibraltar â€“ until 28,000 years ago [said Chris Stringer]" Previous research on Neanderthal sites which suggested that they were more recent than 40,000 years old appears to be wrong," said Stringer. "That is a key finding that will be discussed at the conference."[...] However, scientists have set out to get round these problems. At Oxford University, scientists led by Tom Higham have developed new methods to remove contamination and have been able to make much more precise radiocarbon dating for this period., Currently the earliest fossils of Neanderthals in Europe are dated between 450,000 and 430,000 years ago, and thereafter Neanderthals expanded into Southwest and Central Asia. They are known from numerous fossils, as well as stone tool assemblages. Almost all assemblages younger than 160,000 years are of the so-called Mousterian techno-complex, which is characterised by tools made out of stone flakes.BOOK, Shaw, Ian, Jameson, Robert, A Dictionary of Archaeology, 1999, Blackwell, 978-0-631-17423-3, 408,weblink August 1, 2016, The type specimen is Neanderthal 1, found in Neander Valley in the German Rhineland, in 1856.Compared to modern humans, Neanderthals were stockier, with shorter legs and bigger bodies. In conformance with Bergmann's rule, as well as Allen's rule, this was likely an adaptation to preserve heat in cold climates. Male and female Neanderthals had cranial capacities averaging {{convert|1600|cm3|abbr=on}} and {{convert|1300|cm3|abbr=on}}, respectively,ENCYCLOPEDIA, Stringer, C., Foley, R., Hominid evolution and community ecology, Human evolution and biological adaptation in the Pleistocene, 1984, Academic Press, New York, 978-0122619205, ENCYCLOPEDIA, Holloway, R.L., Delson, E., Ancestors: The hard evidence, The poor brain of Homo sapiens neanderthalensis: see what you please, 1985, Alan R. Liss, New York, 978-0471843764, ;extending to {{convert|1736|cm3|abbr=on}} in the male Amud 1.JOURNAL, Amano, H., Kikuchi, T., Morita, Y., Kondo, O., Suzuki, Hiromasa, Ponce de Leon, M. S., Zollikofer, C.P.E., Bastir, M., Stringer, C., Ogihara, N., 5, August 2015, Virtual Reconstruction of the Neanderthal Amud 1 Cranium, American Journal of Physical Anthropology, 158, 2, 185–97, 10.1002/ajpa.22777, 26249757, 10261/123419, within the range of the values for anatomically modern humans.JOURNAL, Beals, Kenneth, Smith, Courtland, Dodd, Stephen, Brain Size, Cranial Morphology, Climate, and Time Machines, Current Anthropology, 12, 3, 301–30, 1984, 10.1086/203138,weblink Average males stood around {{Convert|164|to|168|cm|0|abbr=on}} and females {{Convert|152|to|156|cm|0|abbr=on}} tall.JOURNAL, Helmuth H, Body height, body mass and surface area of the Neanderthals, Zeitschrift für Morphologie und Anthropologie, 82, 1, 1–12, 1998, 9850627, There has been growing evidence for admixture between Neanderthals and anatomically modern humans, reflected in the genomes of all modern non-African populations but not in the genomes of most sub-Saharan Africans.JOURNAL, Draft full sequence of Neanderthal Genome, 7 May 2010, 10.1126/science.1188021, 20448178, 328, 5979, 5100745, Green RE, Krause J, Briggs AW, Maricic T, Stenzel U, Kircher M, Patterson N, Li H, Zhai W, Fritz MH, Hansen NF, Durand EY, Malaspinas AS, Jensen JD, Marques-Bonet T, Alkan C, Prüfer K, Meyer M, Burbano HA, Good JM, Schultz R, Aximu-Petri A, Butthof A, Höber B, Höffner B, Siegemund M, Weihmann A, Nusbaum C, Lander ES, Russ C, Novod N, Affourtit J, Egholm M, Verna C, Rudan P, Brajkovic D, Kucan Z, Gusic I, Doronichev VB, Golovanova LV, Lalueza-Fox C, de la Rasilla M, Fortea J, Rosas A, Schmitz RW, Johnson PL, Eichler EE, Falush D, Birney E, Mullikin JC, Slatkin M, Nielsen R, Kelso J, Lachmann M, Reich D, Pääbo S, Science, 710–22, 2010Sci...328..710G, JOURNAL, Sankararaman, Sriram, Mallick, Swapan, Patterson, Nick, Reich, David, David Reich (geneticist), The Combined Landscape of Denisovan and Neanderthal Ancestry in Present-Day Humans, Current Biology, 26, 9, 2016, 1241–47, 0960-9822, 10.1016/j.cub.2016.03.037, 27032491, 4864120, JOURNAL, S. Sankararaman, S. Mallick, M. Dannemann, K. Prüfer, J. Kelso, N. Patterson, D. Reich, The landscape of Neandertal ancestry in present-day humans, 4072735, 10.1038/nature12961, 24476815, Nature, 507, 7492, 354–57, 2014, The proportion of Neanderthal-derived ancestry is estimated to be around 1–4% of the modern Eurasian genome. This suggests that some interbreeding between Neanderthals and anatomically modern humans took place after the recent "out of Africa" migration of around 50-70,000 years ago.JOURNAL, October 17, 2012, North African Populations Carry the Signature of Admixture with Neandertals, PLOS ONE, 10.1371/journal.pone.0047765, 7, 10, e47765, 23082212, 3474783, Sánchez-Quinto, F, Botigué, LR, Civit, S, Arenas, C, Avila-Arcos, MC, Bustamante, CD, Comas, D, Lalueza-Fox, C, 2012PLoSO...747765S, JOURNAL, October 23, 2014, Genome sequence of a 45,000-year-old modern human from western Siberia, Nature, 514, 7523, 445–49, 10.1038/nature13810, 25341783, 4753769, Fu, Q, Li, H, Moorjani, P, Jay, F, Slepchenko, SM, Bondarev, AA, Johnson, PL, Aximu-Petri, A, Prüfer, K, de Filippo, C, Meyer, M, Zwyns, N, Salazar-García, DC, Kuzmin, YV, Keates, SG, Kosintsev, PA, Razhev, DI, Richards, MP, Peristov, NV, Lachmann, M, Douka, K, Higham, TF, Slatkin, M, Hublin, JJ, Reich, D, Kelso, J, Viola, TB, Pääbo, S, 2014Natur.514..445F, Brahic, Catherine. "Humanity's forgotten return to Africa revealed in DNA", The New Scientist (February 3, 2014). Recent admixture analyses have added to the complexity, finding that Eastern Neanderthals derived up to 2% of their ancestry from an earlier wave of anatomically modern humans who left Africa some 100,000 years ago.JOURNAL, Kuhlwilm, Martin, Ancient gene flow from early modern humans into Eastern Neanderthals, Nature, 17 February 2016, 530, 7591, 429–433, 10.1038/nature16544, 26886800, 4933530, 2016Natur.530..429K,

Name and classification

{{see|Human taxonomy}}Neanderthals are named after one of the first sites where their fossils were discovered in the mid-19th century in the Neander Valley, just east of Düsseldorf, at the time in the Rhine Province of the Kingdom of Prussia (now in North Rhine-Westphalia, Germany). The valley itself was named for Joachim Neander, Neander being the Hellenized form of the surname Neumann ("new man").JOURNAL, Krings, Matthias, Stone, Anne, Schmitz, Ralf W, Krainitzki, Heike, Stoneking, Mark, Pääbo, Svante, Neandertal DNA Sequences and the Origin of Modern Humans, Cell, 90, 1, 1997, 19–30, 0092-8674, 9230299, 10.1016/S0092-8674(00)80310-4, JOURNAL, M. Krings, 1999, DNA sequence of the mitochondrial hypervariable region II from the Neandertal type specimen, Proceedings of the National Academy of Sciences USA, 96, 10, 5581–85, 10.1073/pnas.96.10.5581, 10318927, 1999PNAS...96.5581K, 21903, JOURNAL, P. Beerli, S.V. Edwards, 2002, When did Neanderthals and modern humans diverge?,weblink Evolutionary Anthropology, 11, S1, 60–63, 10.1002/evan.10058, The German spelling of Thal (valley) was current in the 19th century; it is now spelled Tal.{{efn|The German noun is cognate with English (:wikt:dale|dale).The German /t/ phoneme was frequently spelled th throughout the 15th to 19th centuries; Tal became standardized with the German spelling reform of 1901, thus the German name Neandertal for both the valley and species.}}JOURNAL, Howell, F. Clark, 2816956, The evolutionary significance of variation and varieties of 'Neanderthal' man, The Quarterly Review of Biology, 32, 4, 1957, 330–47, 13506025, 10.1086/401978, Foley, Tim. TalkOrigins Archive. "Neanderthal or Neandertal?". 2005.Neanderthal 1 was known as the "Neanderthal cranium" or "Neanderthal skull" in anthropological literature, and the individual reconstructed on the basis of the skull was occasionally called "the Neanderthal man".The binomial name Homo neanderthalensis—extending the name "Neanderthal man" from the individual type specimen to the entire group—was first proposed by the Anglo-Irish geologist William King in a paper read to the British Association in 1863,JOURNAL, King, W., 1864, On the Neanderthal Skull, or reasons for believing it to belong to the Clydian Period and to a species different from that represented by man, Report of the British Association for the Advancement of Science, Notices and Abstracts, Newcastle-upon-Tyne, 1863, 81–82,weblink 2018-10-21, {{Citation |last1=Murray |first1=John |last2=Nasheuer |first2=Heinz Peter |last3=Seoighe |first3=Cathal |last4=McCormack |first4=Grace P. |last5=Williams |first5=D. Michael |last6=Harper |first6=David A.T. |date=2015 |title=The Contribution of William King to the Early Development of Palaeoanthropology |journal=Irish Journal of Earth Sciences |volume=33 |pages=1–16 |doi=10.3318/ijes.2015.33.1 |lastauthoramp=yes }} although in the following year he stated that the specimen was not human and rejected the name.JOURNAL, King, William, The Reputed Fossil Man of the Neanderthal, The Quarterly Journal of Science, Jan 1864, 1, 96,weblink King's name had priority over the proposal put forward in 1866 by Ernst Haeckel, Homo stupidus.Popular English usage of "Neanderthal" as shorthand for "Neanderthal man", as in "the Neanderthals" or "a Neanderthal", emerged in the popular literature of the 1920s.Inter alia, Boys' Life, p. 18. January 1924.Since the historical spelling -th- in German represents the phoneme /t/, not the fricative /θ/, standard British pronunciation of "Neanderthal" is with /t/ (IPA: {{IPA|/niːˈændÉ™rtɑːl/}}).BOOK, The Oxford Illustrated Dictionary, 564, Oxford University Press, 1976, 1975, Great Britain, (tahl), WEB,weblink Neanderthal adjective â€“ definition in British English Dictionary & Thesaurus,, January 8, 2013, January 22, 2013, WEB,weblink Oxford Learner's Dictionaries â€“ Find pronunciation, clear meanings and definitions of words,, {{dead link|date=July 2016}}BOOK, Kurtén, Björn, Dance of the Tiger: A Novel of the Ice Age, University of California Press, October 10, 1995, xxi, 978-0-520-20277-1,weblink May 9, 2012, JOURNAL, ...And Etymology, Carl J., Pollet, Science News, 140, 12, September 21, 1991, 191, 3975867, 10.2307/3975867, Because of the usual sound represented by the digraph ⟨th⟩ in English, "Neanderthal" is also pronounced with the voiceless fricative /θ/ (as {{IPA|/niːˈændÉ™rθɔːl/}}).WEB,weblink Neanderthal | Define Neanderthal at,, January 22, 2013, {{multiple image| align = right| direction =vertical| header=| total_width=300| image1 = Neanderthal distribution.jpg| caption1 = Distribution of the Neanderthal, and main sites.| image2 = Le Moustier.jpg| caption2 = Reconstitution of Le Moustier Homo neanderthalensis by Charles R. Knight| footer=}}The spelling Neandertal is occasionally seen in English, even in scientific publications.Since "Neanderthal", or "Neandertal", is a common name, there is no authoritative prescription on its spelling, unlike the spelling of the binominal name H. neanderthalensis, which is predicated by King 1864.ENCYCLOPEDIA, Neanderthal, Wiley-Blackwell Encyclopedia of Human Evolution, 2013, Wiley-Blackwell, Chichester, West Sussex, The common name in German is invariably Neandertaler (lit. "of the valley of Neander"), not Neandertal,but the spelling of the name of the Neander Valley itself (Neandertal vs. Neanderthal) has been affected by thespecies name, the names of the Neanderthal Museum and of Neanderthal station persisting with pre-1900 orthography.WEB,weblink Neandertal oder Neanderthal? â€“ Was ist denn nun richtig?,, Neanderthal museum, February 1, 2017, Heute sollten Ortsbezeichnungen das 'Neandertal' ohne 'h' bezeichnen. Alle Namen, die sich auf den prähistorischen Menschen beziehen, führen das 'h'. [Today one should write for place names 'Neandertal' without an 'h'. All names related to the prehistoric humans keep the 'h'.], Ever since the discovery of the Neanderthal fossils, expert opinion has been divided as to whether Neanderthals should be considered a separate species (Homo neanderthalensis) or a subspecies (Homo sapiens neanderthalensis) relative to modern humans.JOURNAL, 10.1073/pnas.96.13.7117, Hominids and hybrids: The place of Neanderthals in human evolution, 1999, Tattersall, Ian, Schwartz, Jeffrey H., Proceedings of the National Academy of Sciences, 96, 13, 7117–19, 48019, 1999PNAS...96.7117T, 10377375, 33580, JOURNAL, Duarte, C., Mauricio, J., Pettitt, P. B., The early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern human emergence in Iberia, Proceedings of the National Academy of Sciences, 96, 13, 1999, 7604–09, 0027-8424, 10.1073/pnas.96.13.7604, 10377462, 1999PNAS...96.7604D, 22133, JOURNAL, 10.1073/pnas.0904119106, The origin of Neandertals, 2009, Hublin, J. J., Proceedings of the National Academy of Sciences, 106, 38, 16022–27, 19805257, 40485013, 2009PNAS..10616022H, 2752594, JOURNAL, Neanderthal taxonomy reconsidered: implications of 3D primate models of intra- and interspecific differences, 14745010, Harvati, K., Frost, S.R., McNulty, K.P., 2004, 10.1073/pnas.0308085100, 337021, 101, 5, Proc. Natl. Acad. Sci. USA, 1147–52, 2004PNAS..101.1147H, Svante Pääbo (2014) described such "taxonomic wars" as unresolveable in principle, "since there is no definition of species perfectly describing the case."The question depends on the definition of Homo sapiens as a chronospecies, which has also been in flux throughout the 20th century. Authorities preferring classification of Neanderthals as subspecies have introduced the subspecies name Homo sapiens sapiens for the anatomically modern Cro-Magnon population which lived in Europe at the same time as Neanderthals, while authorities preferring classification as separate species use Homo sapiens as equivalent to "anatomically modern humans".During the early 20th century, a prevailing view of Neanderthals as "simian", influenced by Arthur Keith and Marcellin Boule,WEB,weblink L'homme fossile de La Chapelle-aux-Saints – full text: Volume VI (pp. 11–172), Volume VII (pp. 21–56), Volume VIII (pp. 1–70), 1911–13, Royal College of Surgeons of England, July 26, 2016, tended to exaggerate the anatomical differences between Neanderthals and Cro-Magnons.Beginning in the 1930s, revised reconstructions of Neanderthals increasingly emphasized the similarity rather than differences from modern humans. From the 1940s throughout the 1970s, it was increasingly common to use the subspecies classification ofHomo sapiens neanderthalensis vs. Homo sapiens sapiens.JOURNAL, Our Neandertal Brethren: Why They Were Not a Separate Species, Scientific American, 303, 2, 34, August 1, 2010, 10.1038/scientificamerican0810-34, 20684369, Shermer, Michael, The hypothesis of "multiregional origin" of modern man was formulated in the 1980s on such grounds, arguing for the presence of an unbroken succession of fossil sites in both Europe and Asia.JOURNAL, Wolpoff, MH, Hawks, J, John D. Hawks, Caspari, R, 2000, Multiregional, not multiple origins, American Journal of Physical Anthropology, 112, 1, 129–36,weblink 10.1002/(SICI)1096-8644(200005)112:13.0.CO;2-K, 10766948, 2027.42/34270, Hybridization between Neanderthals and Cro Magnon had been suggested on skeletal and craniological grounds since the early 20th century, and found increasing support in the later 20th century,BOOK, Cairney, Christopher Thomas, Clans and Families of Ireland and Scotland, an Ethnography of the Gael, 1989, McFarland, London, 978-0899503622, 14,weblink until Neanderthal admixture was found to be present in modern populations' genomes in the 2010s.


{{See|Human evolution#Neanderthal and Denisovan|Accretion model of Neanderthal origins|Neanderthal genetics}}{{see|Homo antecessor|Homo heidelbergensis}}{{multiple image|perrow=2|total_width=280
caption1=Stage 1: Early pre-Neanderthal, possibly Homo erectus, (Tautavel Man, 450 ka)caption2=Stage 2: Archaic Neanderthal (Miguelón, 430 ka)caption3=Stage 3: Intermediate Neanderthal (Saccopastore I, 130 ka) width5 = 150 | caption4 = Stage 4: Classic European Neanderthal (La Chapelle-aux-Saints 1, 50 ka)Accretion model of Neanderthal origins>accretion model.}}Both Neanderthals and anatomically modern humans were initially thought to have evolved from Homo erectus between 300,000 and 200,000 years ago.H. erectus had emerged around 1.8 million years ago, and had long been present, in various subspecies throughout Eurasia.The divergence time between the Neanderthal and archaic Homo sapiens lineages is estimated to be between 800,000 and 400,000 years ago.The more recent time depth has been suggested by Endicott et al. (2010)410–440 kaJOURNAL, P. Endicott, S.Y.W. Ho, Chris Stringer, C. Stringer, 2010, Using genetic evidence to evaluate four palaeoanthropological hypotheses for the timing of Neanderthal and modern human origins, Journal of Human Evolution, 59, 1, 87–95,weblink 10.1016/j.jhevol.2010.04.005, 20510437, and Rieux et al. (2014).295–498 ka.JOURNAL, A. Rieux, 2014, Improved calibration of the human mitochondrial clock using ancient genomes, Molecular Biology and Evolution, 31, 10, 2780–92, 10.1093/molbev/msu222, 25100861, 4166928, The time of divergence between archaic Homo sapiens and ancestors of Neanderthals and Denisovans caused by a population bottleneck of the latter was dated at 744,000 years ago, combined with repeated early admixture events and Denisovans diverging from Neanderthals 300 generations after their split from Homo sapiens, was calculated by Rogers et al. (2017).WEB, Fenton, Bruce R.,weblink Neanderthal Genome Study Reveals Homo sapiens Lineage Diverged 744,000 Years Ago, Huge Gap in Fossil Record Indicated,, August 8, 2017, October 2, 2018, Alan R. Rogers, Ryan J. Bohlender, Chad D. Huff, "Early history of Neanderthals and Denisovans", PNAS 114 (37), September 12, 2017, 9859–63, {{doi|10.1073/pnas.1706426114}};see also:Jordana Cepelewicz, "Genetics Spills Secrets From Neanderthals' Lost History," Quanta Magazine, September 18, 2017."The dating of that schism between the Neanderthals and the Denisovans is surprising because previous research had pegged it as much more recent: a 2016 study, for instance, set it at only 450,000 years ago. An earlier separation means we should expect to find many more fossils of both eventually. It also changes the interpretation of some fossils that have been found. Take the large-brained hominid bones belonging to a species called Homo heidelbergensis, which lived in Europe and Asia around 600,000 years ago. Paleoanthropologists have disagreed about how they relate to other human groups, some positing they were ancestors of both modern humans and Neanderthals, others that they were a nonancestral species replaced by the Neanderthals, who spread across Europe."Homo 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 about 600,000 to 800,000 years ago, the approximate age of Homo antecessor.Matthias Meyer, Juan-Luis Arsuaga, Cesare de Filippo, Sarah Nagel, Ayinuer Aximu-Petri, Birgit Nickel, Ignacio Martínez, Ana Gracia, José María Bermúdez de Castro, Eudald Carbonell, Bence Viola, Janet Kelso, Kay Prüfer & Svante Pääbo, "Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins", Nature 531, pp. 504–07 (March 24, 2016), {{doi|10.1038/nature17405}}.Ewen Callaway, "Oldest ancient-human DNA details dawn of Neanderthals" Sequence of 430,000-year-old DNA pushes back divergence of humans and Neanderthals", Nature News, March 14, 2016.The taxonomic distinction between H. heidelbergensis and Neanderthals is mostly due to a fossil gap in Europe between300,000 and 243,000 years ago (MIS 8). "Neanderthals", by conventions, are fossils which date to after this gap.ARTICLE, D. Dean, On the phylogenetic position of the pre-Neandertal specimen from Reilingen, Germany, J.-J. Hublin, R. Holloway, R. Ziegler, 1998, Journal of Human Evolution, 34, 5, 485–508, 10.1006/jhev.1998.0214, The quality of the fossil record greatly increases from 130,000 years ago onwards. Specimens younger than this date make up the bulk of known Neanderthal skeletons and were the first whose anatomy was comprehensively studied.ARTICLE, B. Vandermeersch, M.D. Garralda, Continuity or Discontinuity in the Peopling of Europe: One Hundred and Fifty Years of Neanderthal Study, Springer Netherlands, S. Condemi, G.-C. Weniger, Vertebrate Paleobiology and Paleoanthropology, 113–25, 2011, 10.1007/978-94-007-0492-3_10, ENCYCLOPEDIA, Neanderthal Lifeways, Subsistence and Technology: One Hundred and Fifty Years of Neanderthal Study, Springer, N.J. Conard, J. Richter, 2, 7–14, 2011, 10.1007/978-94-007-0415-2_2, Neanderthal Lifeways, Subsistence and Technology, Vertebrate Paleobiology and Paleoanthropology, 19, 978-9400704145, In morphological studies, the term "classic Neanderthal" may be used in a narrower sense for Neanderthals younger than 71,000 years old (MIS 4 and 3).{{clear}}


Neanderthals lived alongside modern humans until their extinction between 40,000–30,000 years ago,JOURNAL, Agusti, Jordi, Rubio-Campillo, Xavier, February 2017, Were Neanderthals responsible for their own extinction?, Quaternary International, 431, 232–237, 10.1016/j.quaint.2016.02.017, 2017QuInt.431..232A, and share a common ancestor which could tell us more about how our microbiome evolved.JOURNAL, Weyrich, Laura, etal, 2017, Neanderthal behavior, diet, and disease inferred from ancient DNA in Dental Calculus., Nature, 544, 7650, 357–361, 2017Natur.544..357W, 10.1038/nature21674, 28273061, 10261/152016, Using dental calculus, calcified bone that traps microorganisms, researchers can understand how ancient human microbiomes may have evolved.JOURNAL, Schnorr, Stephanie, etal, 2016, Insights into human evolution from ancient and contemporary microbiome studies, Current Opinions in Genetics and Development, 41, 14–26, 10.1016/j.gde.2016.07.003, 27507098, 5534492, Based on a 16s shotgun sequence of dental calculus found in neanderthal specimens, researchers have found a large portion of neanderthal oral microbiome contained Actinobacteria, Firmicutes, Bacteroidetes, Proteobacteria, much like modern humans, but also Euryarchaeota, fungi, and some oral pathogens that modern humans lack.The diet of neanderthals depended on the environment they lived in. Neanderthal remains recovered from Spy Cave, Belgium and examined using dental calculus, indicate neanderthals in this area had a meat-based diet, including woolly rhinoceros and wild sheep. In comparison, remains found in El Sidrón Cave, Spain, indicate the presence of a large amount of plant material such as nuts and moss, as well as mushrooms in the diet. Researchers determined that the difference in diets contributed to the neanderthal microbiota, and meat-based diet caused the most variation. According to fecal biomarkers, neanderthals were able to convert cholesterol to coprostanol at a high rate, much like modern humans, because of the bacteria present in their gut.JOURNAL, Sistiaga, Ainara, etal, June 25, 2014, The Neanderthal Meal: A New Perspective Using Faecal Biomarkers, PLOS ONE, 9, 6, e101045, 10.1371/journal.pone.0101045, 24963925, 4071062, 2014PLoSO...9j1045S,

Habitat and range

{{Further|Southwest Asian Neanderthals|List of Neanderthal sites}}File:Homo Neanderthalensis Tabun 1 Mount Carmel Israel About 1200,000-50,000 BP.jpg|thumb|left|upright|The southernmost Neanderthal: Homo neandertalensis fossil from Tabun Cave, Israel. 120.000–50.000 BC. Israel MuseumIsrael Museum(File:Neanderthal Range.png|thumb|Approximate Neanderthal range; pre-Neanderthal and early Neanderthal range shown in purple, late Neanderthal range in blue.)Early Neanderthals, living before the Eemian interglacial (130 ka), are poorly known and come mostly from European sites. From 130 ka onwards, the quality of the fossil record increases dramatically. From then on, Neanderthal remains are found in Western, Central, Eastern, and Mediterranean Europe,PRESS RELEASE, Ancient tooth provides evidence of Neanderthal movement, Durham University, February 11, 2008,weblink May 18, 2009, as well as Southwest, Central, and Northern Asia up to the Altai Mountains in Siberia. No Neanderthal has ever been found outside Central to Western Eurasia, namely neither to the south of 30°N (Shuqba, Levant), nor east of 85°E (Denisova, Siberia).File:Carte Neandertaliens.jpg|thumb|Sites where "classic Neanderthal" fossils (70–40 kya) have been found. Ice sheets during the Last Glacial MaximumLast Glacial MaximumThe limit of their northern range appears to have been south of53°N (Bontnewydd, Wales),NEWS, Nicholas, Wade, Nicholas Wade, October 2, 2007, Fossil DNA Expands Neanderthal Range,weblink The New York Times, May 18, 2009, NEWS, Kate, Ravilious, October 1, 2007, Neandertals Ranged Much Farther East Than Thought,weblink National Geographic Society, May 18, 2009, although it is difficult to assess because glacial advances destroy most human remains, the Bontnewydd tooth being exceptional. Middle Palaeolithic artefacts have been found up to 60°N on the Russian plains.JOURNAL, Pavlov P, Roebroeks W, Svendsen JI, The Pleistocene colonization of northeastern Europe: a report on recent research, Journal of Human Evolution, 47, 1–2, 3–17, 2004, 15288521, 10.1016/j.jhevol.2004.05.002, Slimak, L., et al. (2012). "Response to 'Comment on Late Mousterian Persistence near the Arctic Circle{{'"}} Science, 335 (6065), 167 {{doi|10.1126/science.1210211}}Zwyns, N., et al. (2012). "Comment on 'Late Mousterian Persistence near the Arctic Circle{{'"}} Science, 335 (6065), 167 {{doi|10.1126/science.1209908}}Total Neanderthal effective population size has been estimated at close to 15,000 individuals (corresponding to a total population of roughly 150,000 individuals), living in small, isolated, inbred groups.Fabrizio Mafessoni, Kay Prüfer, "Better support for a small effective population size of Neandertals and a long shared history of Neandertals and Denisovans", PNAS 114 (48), November 28, 2017, E10256–E10257, {{doi|10.1073/pnas.1716918114}}.Alan R. Rogers, Ryan J. Bohlender, Chad D. Huff, "Early history of Neanderthals and Denisovans", PNAS 114 (37), September 12, 2017, 9859–63, {{doi|10.1073/pnas.1706426114}}.Analysis of the genomic DNA from three locations suggests that about 120,000 years ago there were genetically distinct Neanderthal populations in Western Europe and Siberia. Later specimens show that the western population not only still populated Europe through 40,000 years ago, but also had spread east, being present at the Siberian locale by 90,000 years ago.JOURNAL, Peyrégne, Stéphane, et al., Nuclear DNA from two early Neandertals reveals 80,000 years of genetic continuity in Europe, Science Advances, 5, 6, eaaw5873, 2019, 10.1126/sciadv.aaw5873, {{clear}}


File:Sapiens neanderthal comparison.jpg|thumb|upright=1.5|Anatomical comparison of skulls of Homo sapiens (Oase 1, left) and Homo neanderthalensis (right)(Cleveland Museum of Natural History).Features compared are the braincase shape, forehead, browridge, nasal bone projection, cheek bone angulation, chin and occipital contour.]](File:Neanderthal cranial anatomy.jpg|frameless|right|upright=1.75)File:Comparison of faces of Homo sapiens and Neanderthal.jpg|thumb|upright=1.5|Comparison of faces of early European Homo sapiens (left) and Homo neanderthalensis (right) based on forensic facial reconstructions exhibited at the Neanderthal MuseumNeanderthal MuseumNeanderthal anatomy differed from modern humans in that they had a more robust build and distinctive morphological features, especially on the cranium, which gradually accumulated more derived aspects as it was described by Marcellin Boule,JOURNAL, Pierre-François, Puech, Bernard, Puech, L'Homme de Neanderthal par Paul Dardé : L'Homme Primitif,weblink French, Neanderthal Man by Paul Dardé: Primitive Man,, July 5, 2017, particularly in certain isolated geographic regions. These include shorter limb proportions, a wider, barrel-shaped rib cage, a reduced chin, sloping forehead, and a large nose, being at the modern human higher end in both width and length,{{efn|There are modern humans with noses as wide as those of Neanderthals and modern humans with similar nose lengths, but none with both Neanderthal nose width and nose length.}} and started somewhat higher on the face than in modern humans. The Neanderthal skull is typically more elongated and less globular than that of anatomically modern humans, and features a notable occipital bun. Inherited Neanderthal DNA variants may subtly influence the skull shape of living people.JOURNAL, Fisher, Simon E., Grabe, Hans J., Macciardi, Fabio, Pääbo, Svante, Franke, Barbara, Hublin, Jean-Jacques, Pourcain, Beate St, Kelso, Janet, Manzi, Giorgio, 2019-01-07, Neandertal Introgression Sheds Light on Modern Human Endocranial Globularity,weblink Current Biology, English, 29, 1, 120–127.e5, 10.1016/j.cub.2018.10.065, 0960-9822, 30554901, 6380688, Neanderthals were much stronger than modern humans, with particularly strong arms and hands, while they were comparable in height; based on 45 long bones from 14 males and 7 females, three different methods of estimating height produced averages for Neanderthal males from {{Convert|164|to|168|cm|0|abbr=on}} and {{Convert|152|to|156|cm|0|abbr=on}} for females. Samples of 26 specimens found an average weight of {{Convert|77.6|kg|lbs|abbr=on}} for males and {{Convert|66.4|kg|lbs|abbr=on}} for females.JOURNAL, Froehle, Andrew W, Churchill, Steven E, 2009, Energetic Competition Between Neandertals and Anatomically Modern Humans, PaleoAnthropology, 96–116,weblink October 31, 2011, Neanderthals are known for their large cranial capacity, which at {{Convert|1600|cm3|cuin|abbr=on}} is larger on average than that of modern humans. One study has found that drainage of the dural venous sinuses (low pressure blood vessels that run between the meninges and skull leading down through the skull) in the occipital lobe region of Neanderthal brains appears more asymmetric than other hominid brains.WEB,weblink El cerebro neandertal era más asimétrico que el del 'Homo sapiens', SINC Servicio de Información y Noticias Científicas, 2011-11-07, Three-dimensional computer-assisted reconstructions of Neanderthal infants based on fossils from Russia and Syria indicated that Neanderthal and modern human brains were the same size at birth, but that by adulthood, the Neanderthal brain was larger than the modern human brain.WEB,weblink Neanderthal Brain Size at Birth Sheds Light on Human Evolution, David Maxwell, Braun, National Geographic, September 9, 2008, September 19, 2009, They had almost the same degree of encephalisation (i.e. brain-to-body-size ratio) as modern humans.Silberman, Neil. The Oxford Companion to Archaeology, p. 455 (Oxford University Press 2012): "[I]t is with the Neanderthals that we see the full achievement, for the first time, of the degree of encephalization (brain to body size ratio) that characterizes modern humans."Abramiuk, Marc. The Foundations of Cognitive Archaeology, p. 199 (MIT Press 2012): "the encephalization quotient was slightly smaller".Three-dimensional reconstructions of nasal cavities and computational fluid dynamics techniques have found that Neanderthals and modern humans both adapted their noses (independently and in a convergent way) to help breathe in cold and dry conditions.JOURNAL, de Azevedo, S., González, M. F., Cintas, C., Ramallo, V., Quinto-Sánchez, M., Márquez, F., Hünemeier, T., Paschetta, C., Ruderman, A., Navarro, P., Pazos, B., Silva de Cerqueira, C., Velan, O., Ramírez-Rozzi, F., Calvo, N., Castro, H., Paz, R.R., González-José, R., October 30, 2017, Nasal airflow simulations suggest convergent adaptation in Neanderthals and modern humans, Proceedings of the National Academy of Sciences, 10.1073/pnas.1703790114, 0027-8424, 29087302, 114, 47, 12442–12447, 5703271, The large nose seen in Neanderthals, as well as Homo heidelbergensis, affected the shape of the skull and the muscle attachments, and gave them a weaker bite force than in modern humans.NEWS,weblink Your face is probably more primitive than a Neanderthal's, Richard, Gray, February 15, 2017, BBC News, Larger eye sockets and larger areas of the brain devoted to vision suggest that their eyesight may have been better than that of modern humans.WEB,weblink Neanderthal brains focused on vision and movement leaving less room for social networking, Science Daily, March 19, 2013, Dental remains from two Italian sites indicate that Neanderthal dental features had evolved by around 450,000 years ago during the Middle Pleistocene epoch.JOURNAL, Zanolli, Clément, Martinón-Torres, María, Bernardini, Federico, Boschian, Giovanni, Coppa, Alfredo, Dreossi, Diego, Mancini, Lucia, Martínez de Pinillos, Marina, Martín-Francés, Laura, Bermúdez de Castro, José María, Tozzi, Carlo, Tuniz, Claudio, Macchiarelli, Roberto, The Middle Pleistocene (MIS 12) human dental remains from Fontana Ranuccio (Latium) and Visogliano (Friuli-Venezia Giulia), Italy. A comparative high resolution endostructural assessment, PLOS ONE, Plos, 3 October 2018, e0189773, 13, 10, 10.1371/journal.pone.0189773, 30281595, 6169847, 2018PLoSO..1389773Z, Two Neanderthal specimens from Italy and Spain were found to have an allele of the melanocortin 1 receptor (MC1R) with reduced activity. This receptor plays a role in mammalian pigmentation, and the activity of the novel allele in Neanderthals was found to be reduced sufficiently to allow for visibly lighter pigment expression.JOURNAL, Lalueza-Fox, C., Rompler, H., Caramelli, D., Staubert, C., Catalano, G., Hughes, D., Rohland, N., Pilli, E., Longo, L., Condemi, S., de la Rasilla, M., Fortea, J., Rosas, A., Stoneking, M., Schoneberg, T., Bertranpetit, J., Hofreiter, M., A Melanocortin 1 Receptor Allele Suggests Varying Pigmentation Among Neanderthals, Science, 318, 17962522, 5855, 2007, 1453–55,weblink 2007Sci...318.1453L, 0036-8075, 10.1126/science.1147417, September 10, 2017, Although not found in the small European sample studied by Lalueza et al., a larger study found that the derived variant was present at 70% frequency in Taiwanese Aborigines, 50% frequency in Cheyenne Native Americans, 30% frequency in Han Chinese, and 5% frequency in Europeans.JOURNAL, Q., Ding, Neanderthal origin of the haplotypes carrying the functional variant Val92Met in the MC1R in modern humans., Molecular Biology and Evolution, 31, 8, 1994–2003, 10 June 2014, 10.1093/molbev/msu180, 24916031, It is therefore unclear whether this loss-of-function variant is responsible for any other traits other than lightening the skin (such as red or blonde hair). This allele was not found in the Croatian or Altai Neanderthal specimens subjected to whole-genome sequencing, nor have the MC1R variants known to cause red hair in modern humans, though the Altai specimen was polymorphic for another variant MC1R allele of unknown effect.JOURNAL, Dannerman, Micheal, Kelso, Janet, The Contribution of Neanderthals to Phenotypic Variation in Modern Humans, American Journal of Human Genetics, 101, 4, P578-589, 2017, 10.1016/j.ajhg.2017.09.010, 28985494, 5630192, Genomic analysis of three Croatian specimens for the alleles of numerous genes that affect pigment in modern humans showed the Neanderthals to have more of the alleles that produce dark pigment in modern humans than those producing reduced pigmentation. Based on this they concluded these Neanderthals had darker hair, skin and eye coloration than modern Europeans. Skin pigmentation prediction for archaic humans is controversial, as there are no living samples in which to evaluate the effect of SNP variants, and, with the tested samples coming from a single Neanderthal population, they may not be representative of the diversity across Neanderthals' full geographic range.WEB, Watson, Traci, Were Some Neandertals Brown-Eyed Girls?,, 19 March 2012, 8 March 2019,weblink The overall shorter limbs and in general more stout body proportions of Neanderthals may have been an adaptation to colder climates. In comparison to modern humans, Neanderthals were more suited for sprinting and pouncing activities rather than endurance running, which would have been adaptive in the forests and woodlands that seem to have been their preferred environment. Genomic evidence possibly points to a higher proportion of fast-twitch muscle fiber in the Neanderthal.WEB,weblink Palaeoecological and genetic evidence for Neanderthal power locomotion as an adaptation to a woodland environment, ResearchGate, en, 2019-01-30, Evidence suggests that Neanderthals walked upright much like modern humans.JOURNAL, Haeusler, Martin, et al., Morphology, pathology, and the vertebral posture of the La Chapelle-aux-Saints Neandertal, 25 February 2019, Proceedings of the National Academy of Sciences of the United States of America, 116, 11, 4923–4927, 10.1073/pnas.1820745116, 30804177, 6421410, NEWS, Cassella, Carly, We Have Been Wrong About a Key Feature of Neanderthals' Appearance,weblink 1 March 2019,, 1 March 2019, {{clear}}


{{See also|Pleistocene human diet}}File:Pointe levallois Beuzeville MHNT PRE.2009.0.203.2.fond.jpg|thumb|right|Levallois point – Beuzeville, FranceFranceNeanderthals made stone tools, used fire,JOURNAL, Heyes, Peter, Anastasakis, Konstantinos, de Jong, Wiebren, Selection and Use of Manganese Dioxide by Neanderthals, Scientific Reports, 6, 2016, 22159, 2045-2322, 10.1038/srep22159, 26922901, 4770591, 2016NatSR...622159H, and were hunters. This is the extent of the consensus on their behaviour. It had long been debated whether Neanderthals were hunters or scavengers, but the discovery of the pre-Neanderthal Schöningen wooden spears in Germany helped settle the debate in favour of hunting. A Levallois point embedded in the vertebrae of a wild ass indicated that a javelin had been thrown with a parabolic trajectory to disable the animal.A Levallois point embedded in the vertebra of a wild ass (Equus africanus): hafting, projectiles and Mousterian hunting weaponsEric Boëda, Antiquity, Volume 73, Issue 280 June 1999 , pp. 394–402 Most available evidence suggests they were apex predators,JOURNAL, 10.1016/j.jhevol.2005.03.003, Isotopic evidence for diet and subsistence pattern of the Saint-Césaire I Neanderthal: Review and use of a multi-source mixing model, 2005, Bocherens, Hervé, Drucker, Dorothée G., Billiou, Daniel, Patou-Mathis, Marylène, Vandermeersch, Bernard, Journal of Human Evolution, 49, 71–87, 15869783, 1, and fed on red deer, reindeer, ibex, wild boar, aurochs and on occasion mammoth, straight-tusked elephant and rhinoceros.NEWS,weblink French dig up Neanderthal 'butcher's shop', Lichfield, John, The New Zealand Herald, September 30, 2006, JOURNAL, 10.1371/journal.pone.0014769, Molar Macrowear Reveals Neanderthal Eco-Geographic Dietary Variation, 2011, Rosenberg, Karen, Fiorenza, Luca, Benazzi, Stefano, Tausch, Jeremy, Kullmer, Ottmar, Bromage, Timothy G., Schrenk, Friedemann, PLOS ONE, 6, 3, e14769, 21445243, 3060801, 2011PLoSO...614769F, They appear to have occasionally used vegetables as fall-back food,JOURNAL, 2, 108, 486–91, 2010, 3021051, 21187393, Proceedings of the National Academy of Sciences, 10.1073/pnas.1016868108, Brooks, A. G., A. S., Piperno, Microfossils in calculus demonstrate consumption of plants and cooked foods in Neanderthal diets (Shanidar III, Iraq; Spy I and II, Belgium), D. R., Henry, 2011PNAS..108..486H, revealed by isotope analysis of their teeth and study of their coprolites (fossilised faeces).JOURNAL, 10.1073/pnas.120178997, Neanderthal diet at Vindija and Neanderthal predation: The evidence from stable isotopes, 2000, Richards, Michael P., Pettitt, Paul B., Trinkaus, Erik, Smith, Fred H., Paunović, Maja, Karavanić, Ivor, Proceedings of the National Academy of Sciences, 97, 13, 7663–66, 10852955, 122870, 2000pnas...97.7663r, 16602, Dental analysis of specimens from Spy, Belgium and El Sidrón, SpainJOURNAL, Neanderthal behaviour, diet, and disease inferred from ancient DNA in dental calculus, 544, 7650, 357–361, Nature, 10.1038/nature21674, 28273061, 2017, Weyrich, Laura S., Duchene, Sebastian, Soubrier, Julien, Arriola, Luis, Llamas, Bastien, Breen, James, Morris, Alan G., Alt, Kurt W., Caramelli, David, Dresely, Veit, Farrell, Milly, Farrer, Andrew G., Francken, Michael, Gully, Neville, Haak, Wolfgang, Hardy, Karen, Harvati, Katerina, Held, Petra, Holmes, Edward C., Kaidonis, John, Lalueza-Fox, Carles, de la Rasilla, Marco, Rosas, Antonio, Semal, Patrick, Soltysiak, Arkadiusz, Townsend, Grant, Usai, Donatella, Wahl, Joachim, Huson, Daniel H., Dobney, Keith, 29, 2017Natur.544..357W, 10261/152016, suggested that these Neanderthals had a wide-ranging diet, with no evidence at all that the El Sidrón Neanderthals were carnivorous, instead living on "a mixture of forest moss, pine nuts and a mushroom known as split gill".WEB,weblink Neanderthal dental tartar reveals plant-based diet â€“ and drugs, The Guardian, March 8, 2017, Nicola Davis, March 9, 2017, Nonetheless, isotope studies of Neanderthals from two French sites showed similar profiles to other carnivores, suggesting that these populations may have eaten meat.JOURNAL, Jaouen, Klervia, etal, Exceptionally high δ15N values in collagen single amino acids confirm Neandertals as high-trophic level carnivores, 19 February 2019, Proceedings of the National Academy of Sciences of the United States of America, 116, 11, 4928–4933, 10.1073/pnas.1814087116, 30782806, 6421459, NEWS, Yika, Bob, Isotopes found in bones suggest Neanderthals were fresh meat eaters,weblink 19 February 2019,, 19 February 2019, NEWS, Max Planck Institute for Evolutionary Anthropology, Neanderthals' main food source was definitely meat – Isotope analyses performed on single amino acids in Neanderthals' collagen samples shed new light on their debated diet,weblink 19 February 2019, Science Daily, 21 February 2019, The Neanderthal skeleton suggests they consumed {{convert|100|to|350|kcal|kJ|abbr=on}} more per day than modern male humans of {{convert|68.5|kg|abbr=on}} and females of {{convert|59.2|kg|abbr=on}}.(File:Reconstruction of Neanderthal woman.jpg|thumb|Reconstruction of a Neanderthal woman.JOURNAL, Cro-Magnons Conquered Europe, but Left Neanderthals Alone, PLOS Biology, 30 November 2004, 2, 12, e449, 10.1371/journal.pbio.0020449, en, 1545-7885, 532398, )The size and distribution of Neanderthal sites, along with genetic evidence, suggests Neanderthals lived in much smaller and more sparsely distributed groups than anatomically-modern Homo sapiens.JOURNAL,weblink Shaw, Kate, Sheer Numbers Gave Early Humans Edge Over Neanderthals, Wired,, July 29, 2011, WEB,weblink Vergano, Dan, Neanderthals Lived in Small, Isolated Populations, Gene Analysis Shows, National Geographic, April 22, 2014, The bones of twelve Neanderthals were discovered at El Sidrón cave in northwestern Spain. They are thought to have been a group killed and butchered about 50,000 years ago. Analysis of the mtDNA showed that the three adult males belonged to the same maternal lineage, while the three adult females belonged to different ones. This suggests a social structure where males remained in the same social group and females "married out".BOOK, Ian, Tattersall, Ian Tattersall, The Strange Case of the Rickety Cossack and other Cautionary Tales from Human Evolution, 202, Palgrave Macmillan, 2015, 978-1-137-27889-0, The bones of the El Sidrón group show signs of defleshing, suggesting that they were victims of cannibalism. The St. Césaire 1 skeleton from La Roche à Pierrot, France, showed a healed fracture on top of the skull apparently caused by a deep blade wound, suggesting interpersonal violence.JOURNAL, Zollikofer, C. P. E., Ponce de Leon, M. S., Vandermeersch, B., Lévêque, F., 2002, Evidence for Interpersonal Violence in the St. Césaire Neanderthal, PNAS, 99, 9, 6444–48, 10.1073/pnas.082111899, 11972028, 122968, 2002PNAS...99.6444Z, Shanidar 3, an adult male dated to the late middle Paleolithic, was found to have a rib lesion characteristic of projectile weapon injuries, which some anthropologists consider evidence for interspecies conflict.JOURNAL, Churchill, Steven E., Franciscus, Robert G., McKean-Peraza, Hilary A., Daniel, Julie A., Warren, Brittany R., August 2009, Shanidar 3 Neandertal rib puncture wound and paleolithic weaponry, Journal of Human Evolution, 57, 2, 163–178, 10.1016/j.jhevol.2009.05.010, 1095-8606, 19615713, Neanderthals suffered a high rate of traumatic injury, with by some estimates 79% of specimens showing evidence of healed major trauma.JOURNAL, Nakahashi, W., March 2017, The effect of trauma on Neanderthal culture: A mathematical analysis, Homo: Internationale Zeitschrift Fur die Vergleichende Forschung Am Menschen, 68, 2, 83–100, 10.1016/j.jchb.2017.02.001, 1618-1301, 28238406, It was thus theorized that Neanderthals employed a riskier and possibly less sophisticated hunting strategy. However, rates of cranial trauma are not significantly different between Neanderthal and middle paleolithic Anatomically Modern Human samples.JOURNAL, Harvati, Katerina, Wahl, Joachim, Anthes, Nils, Beier, Judith, November 2018, Similar cranial trauma prevalence among Neanderthals and Upper Palaeolithic modern humans, Nature, 563, 7733, 686–690, 10.1038/s41586-018-0696-8, 30429606, 1476-4687, 2018Natur.563..686B, Both populations evidently cared for the injured and had some degree of medical knowledge.It was long believed that an adhesive (birch bark tar) made by Neanderthals required to follow a complex recipe, and that it thus showed complex cognitive skills and cultural transmission. A study from 2019 showed that birch tar production can instead be a very simple process - merely involving the burning of birch bark near smooth vertical surfaces in open air conditions.JOURNAL, Schmidt, P., Blessing, M., Rageot, M., Iovita, R., Pfleging, J., Nickel, K. G.; Righetti, L. & Tennie, C., Birch tar extraction does not prove Neanderthal behavioral complexity, PNAS, 10.1073/pnas.1911137116, Claims that Neanderthals deliberately buried their dead, and if they did, whether such burials had any symbolic meaning,BOOK, Stringer, C., Gamble, C., 1993, In Search of the Neanderthals,weblink London, Thames and Hudson, 978-0500050705, registration, {{rp|158–60}} are heavily contested.JOURNAL, Rendu W, Beauval C, Crevecoeur I, Bayle P, Balzeau A, Bismuth T, Bourguignon L, Delfour G, Faivre JP, Lacrampe-Cuyaubère F, Muth X, Pasty S, Semal P, Tavormina C, Todisco D, Turq A, Maureille B, 2016, Let the dead speak...comments on Dibble et al.'s reply to 'Evidence supporting an intentional burial at La Chapelle-aux-Saints', Journal of Archaeological Science, 69, 12–20, 10.1016/j.jas.2016.02.006, JOURNAL, Gargett, R.H., 1989, Grave Shortcomings: The Evidence for Neandertal Burial, Current Anthropology, 30, 2, 157–90, 10.1086/203725, JOURNAL, Gargett, R.H., 1999, Middle Palaeolithic burial is not a dead issue: the view from Qafzeh, Saint-Césaire, Kebara, Amud, and Dederiyeh, Journal of Human Evolution, 37, 1, 27–90, 10.1006/jhev.1999.0301, 10375476, The debate on deliberate Neanderthal burials has been active since the 1908 discovery of the well-preserved Chapelle-aux-Saints 1 skeleton in a small hole in a cave in southwestern France. In this controversy's most recent installment, a team of French researchers reinvestigated the Chapelle-aux-Saints cave and in January 2014 reasserted the century-old claim that the 1908 Neanderthal specimen had been deliberately buried,JOURNAL, Rendu W, Beauval C, Crevecoeur I, Bayle P, Balzeau A, Bismuth T, Bourguignon L, Delfour G, Faivre JP, Lacrampe-Cuyaubère F, Tavormina C, Todisco D, Turq A, Maureille B, January 2014, Evidence supporting an intentional Neandertal burial at La Chapelle-aux-Saints, Proceedings of the National Academy of Sciences, 111, 1, 81–86, 10.1073/pnas.1316780110, 24344286, 2014PNAS..111...81R, 3890882, and this has in turn been heavily criticised.JOURNAL, H., Dibble, V., Aldeias, P., Goldberg, D., Sandgathe, T. E., Steele, 2015, A critical look at evidence from La Chapelle-aux-Saints supporting an intentional burial, Journal of Archaeological Science, 53, 649–57, 10.1016/j.jas.2014.04.019, According to archaeologist John F. Hoffecker:JOURNAL, Hoffecker, JF, 2009, The spread of modern humans in Europe, Proc. Natl. Acad. Sci. USA, 106, 38, 16040–16045, 10.1073/pnas.0903446106, 19571003, 2009PNAS..10616040H, 2752585, – References 399 Hoffecker JF, Cleghorn ...{{clear}}Evidence of likely Neanderthal presence in Kefallinia and Zakynthos and the discovery of early Palaeolithic artefacts in Crete may suggest that Neanderthals used boats and mastered seafaring.Ferentinos et al., 2012, Early seafaring activity in the southern Ionian Islands, Mediterranean Sea. Journal of Archaeological Science 39, pp. 2167-2176.weblink 2012, Evidence suggests Neanderthals took to boats before modern humans,weblink et al., 2011, Dating Palaeolithic sites in southwestern Crete, Greece. Journal of Quaternary Science, 26, pp. 553-560.weblink


{{see also|Bear worship|History of art}}(File:NHM - Neandertaler Modell 1.jpg|thumb|left|Artist's reconstruction of a Neanderthal man with child)File:Possible Neandertal Jewelry White-Tailed Eagle Claws with striations at the Neanderthal site of Krapina, Croatia, 130,000 BP.jpg|thumb|Proposed Neanderthal jewelry: white-tailed eagle claw with striations at the Neanderthal site of Krapina, (Croatia]], circa 130,000 BP.JOURNAL, Frayer, David W., Radovčić, Jakov, Sršen, Ankica Oros, Radovčić, Davorka, Evidence for Neandertal Jewelry: Modified White-Tailed Eagle Claws at Krapina, PLOS ONE, 11 March 2015, 10, 3, e0119802, 10.1371/journal.pone.0119802, en, 1932-6203, 4356571, )Whether Neanderthals created art and used adornments, which would indicate a capability for complex symbolic thought, remains unresolved. A 2010 paper on radiocarbon dates cast doubt on the association of Châtelperronian beads with Neanderthals,JOURNAL, 10.1073/pnas.1007963107, 20956292, 2996711, Chronology of the Grotte du Renne (France) and implications for the context of ornaments and human remains within the Chatelperronian, Proceedings of the National Academy of Sciences, 107, 47, 20234–20239, 2010, Higham, T., Jacobi, R., Julien, M., David, F., Basell, L., Wood, R., Davies, W., Ramsey, C. B., and Paul Mellars considered the evidence for symbolic behaviour to have been refuted.JOURNAL, 10.1073/pnas.1014588107, 21078972, 2996706, Neanderthal symbolism and ornament manufacture: The bursting of a bubble?, Proceedings of the National Academy of Sciences, 107, 47, 20147–20148, 2010, Mellars, P., 2010PNAS..10720147M, This conclusion, however, is controversial, and others such as Jean-Jacques Hublin and colleagues have re-dated material associated with the Châtelperronian artifactsJOURNAL, J.-J. Hublin, S. Talamo, M. Julien, F. David, N. Connet, P. Bodu, B. Vandermeersch, M.P. Richards, Radiocarbon dates from the Grotte du Renne and Saint-Césaire support a Neandertal origin for the Châtelperronian, Proceedings of the National Academy of Sciences USA, 109, 46, 18743–18748, 10.1073/pnas.1212924109, 23112183, 3503158, 2012PNAS..10918743H, 2012, and used proteomic evidence to restate the challenged association with Neanderthals.JOURNAL, F. Welkera, M. Hajdinjak, S. Talamo, K. Jaouen, M. Dannemann, F. David, M. Julien, M. Meyer, J. Kelso, I. Barnes, S. Brace, P. Kamminga, R. Fischer, B.M. Kessler, J.R. Stewart, S. Pääbo, M.J. Collins, J.-J. Hublin, Palaeoproteomic evidence identifies archaic hominins associated with the Châtelperronian at the Grotte du Renne, Proceedings of the National Academy of Sciences USA, 113, 40, 11162–67, 10.1073/pnas.1605834113, 27638212, 5056053, 2016, A large number of other claims of Neanderthal art, adornment, and structures have been made. These are often taken by the media as showing Neanderthals were capable of symbolic thought,NEWS, ScienceDaily, Neanderthals were not inferior to modern humans, study finds, April 30, 2014,weblink NEWS, Sci News, E., de Lazaro, January 18, 2017, Neanderthals Capable of Incorporating Symbolic Objects into Their Culture, Discovery Suggests,weblink or were "mental equals" to anatomically modern humans.MAGAZINE, Scientific American, January 8, 2010, C.Q. Choi, Heavy Brows, High Art?: Newly Unearthed Painted Shells Show Neandertals Were Homo sapiens' Mental Equals,weblink NEWS, The Guardian, Neanderthals were not less intelligent than modern humans, scientists find, I. Sample,weblink April 30, 2014, As evidence of symbolism, none of them are widely accepted,MAGAZINE, N. Branan, 2010,weblink Neandertal Symbolism: Evidence Suggests a Biological Basis for Symbolic Thought, Scientific American, although the same is true for Middle Palaeolithic anatomically modern humans. Among many others:File:Decorated raven bone from the Zaskalnaya VI (Kolosovskaya) Neanderthal site, Crimea, Micoquian industry dated to between 38 and 43 cal kyr BP.jpg|thumb|Proposed symbolic endeavour: incision-decorated raven bone from the Zaskalnaya VI (Kolosovskaya) Neanderthal site, Crimea, MicoquianMicoquian
  • Flower pollen on the body of pre-Neanderthal Shanidar 4, Iraq, had in 1975 been argued to be a flower burial.JOURNAL, R. S. Solecki, Shanidar IV, a Neanderthal Flower Burial in Northern Iraq, 190, 4217, 880–81, 1975, 10.1126/science.190.4217.880, Science, 1975Sci...190..880S, Once popular, this theory is no longer accepted.JOURNAL, D.J. Sommer, The Shanidar IV 'Flower Burial': a Re-evaluation of Neanderthal Burial Ritual, Cambridge Archaeological Journal, 9, 1, 127–29, 1999, 10.1017/s0959774300015249, JOURNAL, The Neanderthal Dead, exploring mortuary variability in Middle Paleolithic Eurasia, Paul B. Pettitt, 2002, Before Farming, 1, 4,
  • Bird bones were argued to show evidence for feather plucking in a 2012 study examining 1,699 ancient sites across Eurasia, which the authors controversiallyMAGAZINE, Did Neanderthals use feathers for fashion?,weblink New Scientist, June 16, 2017, took to mean Neanderthals wore bird feathers as personal adornments.JOURNAL, Birds of a Feather: Neanderthal Exploitation of Raptors and Corvids, 10.1371/journal.pone.0045927, 23029321, 3444460, PLOS ONE, September 17, 2012, 7, 9, e45927, Finlayson, Clive, Brown, Kimberly, Blasco, Ruth, Rosell, Jordi, Negro, Juan José, Bortolotti, Gary R, Finlayson, Geraldine, Sánchez Marco, Antonio, Giles Pacheco, Francisco, Rodríguez Vidal, Joaquín, Carrión, José S, Fa, Darren A, Rodríguez Llanes, José M, 2012PLoSO...745927F,
  • Deep scratches were found in 2012 on a cave floor underlying Neanderthal layer in Gorham's Cave, Gibraltar, which some have controversiallyJOURNAL, E. Callaway, Neanderthals made some of Europe's oldest art, 2014, Nature News, 10.1038/nature.2014.15805,weblink interpreted as art.NEWS,weblink Neanderthal 'artwork' found in Gibraltar cave, September 1, 2014, BBC News, Paul, Rincon,
  • Two 176,000-year-old stalagmite ring structures, several metres wide, were reported in 2016 more than 300 metres from the entrance within Bruniquel Cave, France. The authors claim artificial lighting would have been required as this part of the cave is beyond the reach of daylight and that the structures had been made by early Neanderthals, the only humans in Europe at this time.JOURNAL, 10.1038/nature18291, Early Neanderthal Constructions deep in Bruniquel Cave in Southwestern France, Jaubert, Jacques, Verheyden, Sophie, Genty, Dominique, Soulier, Michel, Cheng, Hai, Blamart, Dominique, Burlet, Christian, Camus, Hubert, Delaby, Serge, Deldicque, Damien, Edwards, R. Lawrence, Ferrier, Catherine, Lacrampe-Cuyaubère, François, Lévêque, François, Maksud, Frédéric, Mora, Pascal, Muth, Xavier, Régnier, Édouard, Rouzaud, Jean-Noël, Santos, Frédéric, Nature, 534, 7605, June 2, 2016, online May 25, 2016, 0028-0836, 111–14, 2016Natur.534..111J, 27251286,
  • In 2015, a study argued that a number of 130,000-year-old eagle talons found in a cache near Krapina, Croatia along with Neanderthal bones, had been modified to be used as jewelry.JOURNAL, Radovčić, D., SrÅ¡en, A. O., Radovčić, J., Frayer, D. W., 2015, Evidence for Neandertal jewelry: Modified white-tailed eagle claws at Krapina, PLOS ONE, 10, 3, e0119802, 10.1371/journal.pone.0119802, 25760648, 4356571, 2015PLoSO..1019802R, JOURNAL,weblink 10.1038/nature.2015.17095, Neanderthals wore eagle talons as jewellery, Nature, 2015, Callaway, Ewen,
All of these appeared only in single locations. Yet in 2018, using uranium-thorium dating methods,JOURNAL, Pike, A. W., Hoffmann, D. L., Pettitt, P. B., García-Diez, M., Zilhão, J., 2017, Dating Palaeolithic cave art: Why U–Th is the way to go, Quaternary International, 432, 41–49, 10.1016/j.quaint.2015.12.013, 2017QuInt.432...41P, red painted symbols comprising a scalariform (ladder shape), a negative hand stencil, and red lines and dots on the cave walls of three Spanish caves {{convert|700|km|abbr=on}} apart were dated to at least 64,000 years old.JOURNAL, D. L. Hoffmann, C. D. Standish, M. García-Diez, P. B. Pettitt, J. A. Milton, J. Zilhão, J. J. Alcolea-González, P. Cantalejo-Duarte, H. Collado, R. de Balbín, M. Lorblanchet, J. Ramos-Muñoz, G.-Ch. Weniger, A. W. G. Pike, 2018, U-Th dating of carbonate crusts reveals Neandertal origin of Iberian cave art, Science, 359, 6378, 912–15, 10.1126/science.aap7778, 29472483, 2018Sci...359..912H, If the dating is correct, they were painted before the time anatomically modern humans are thought to have arrived in Europe. Paleoanthropologist John D. Hawks argues these findings demonstrate Neanderthals were capable of symbolic behaviour previously thought to be unique to modern humans.NEWS,weblink Ancient cave paintings turn out to be by Neanderthals, not modern humans, Researchers also found perforated seashell beads and pigments that are at least 115,000 years old in Cueva de los Aviones, a cave in southeastern Spain.WEB,weblink World's Oldest Cave Art Found—And Neanderthals Made It, 2018-02-22, National Geographic News, 2019-06-28,

Interbreeding with archaic and modern humans{{anchor|Interbreeding}}

File:Homo-Stammbaum, Version Stringer-en.svg|upright=1.25|thumb|right|Chris Stringer's hypothesis of the family tree of genus Homo, published 2012 in Nature â€“ the horizontal axis represents geographic location, and the vertical axis represents time in millions of years ago.{{efn|Homo floresiensis originated in an unknown location from unknown ancestors and reached remote parts of Indonesia. Homo erectus spread from Africa to western Asia, then east Asia and Indonesia; its presence in Europe is uncertain, but it gave rise to Homo antecessor, found in Spain. Homo heidelbergensis originated from Homo erectus in an unknown location and dispersed across Africa, southern Asia and southern Europe (other scientists interpret fossils, here named heidelbergensis, as late erectus). Homo sapiens sapiens spread from Africa to western Asia and then to Europe and southern Asia, eventually reaching Australia and the Americas. In addition to Neanderthals and Denisovans, a third (gene flow]] of archaic Africa origin is indicated at the right.JOURNAL, 10.1038/485033a, 2012Natur.485...33S, 22552077, Evolution: What makes a modern human, 2012, Nature, Stringer, Chris, 485, 7396, 33–35, }})An alternative proposal regarding the fate of Neanderthals is that rather than being replaced by modern humans and going extinct, Neanderthals were absorbed into the Cro-Magnon population by interbreeding. This would be counter to strict versions of theory of recent African origin of modern humans, since it would imply that at least part of the genome of Europeans would descend from Neanderthals. Evidence of interbreeding has been detected in genomes of ancient and modern humans, though this could have resulted from occasional interbreeding and introgression, rather than population absorption.

Pre-2010 interbreeding hypotheses

Until the early 1950s, most scholars thought Neanderthals were not in the ancestry of living humans.JOURNAL, Boule, Marcellin, 1911–1913, L'homme fossile de La Chapelle-aux-Saints, Annales de Paléontologie, 6–8, French, {{rp|232–34}}JOURNAL, Neanderthal Man Not Our Ancestor, G.E. Smith, Scientific American, 10.1038/scientificamerican0828-112, 139, 2, 1928, 112–15, 1928SciAm.139..112S, {{paywall}} Nevertheless, in 1904 Thomas H. Huxley saw among Frisians the presence of what he suspected to be Neanderthaloid skeletal and cranial characteristics as an evolutionary development from Neanderthal rather than as a result of interbreeding, saying that "the blond long-heads may exhibit one of the lines of evolution of the men of the Neanderthaloid type," yet he raised the possibility that the Frisians alternatively "may be the result of the admixture of the blond long-heads with Neanderthal men," thus separating "blond" from "Neanderthaloid."WEB, Huxley, Thomas H., Man's Place in Nature and Other Anthropological Essays,weblink Google Books, J. A. Hill, 1906, Hans Peder Steensby proposed interbreeding in 1907 in the article Race studies in Denmark. He strongly emphasised that all living humans are of mixed origins.WEB, H. P., Steensby, Racestudier i Danmark, Race Studies in Denmark, Danish,weblink Geographical Journal, Royal Library, Denmark, 1907, July 6, 2017, He held that this would best fit observations, and challenged the widespread idea that Neanderthals were ape-like or inferior. Basing his argument primarily on cranial data, he noted that the Danes, like the Frisians and the Dutch, exhibit some Neanderthaloid characteristics, and felt it was reasonable to "assume something was inherited" and that Neanderthals "are among our ancestors."Carleton Stevens Coon in 1962 found it likely, based upon evidence from cranial data and material culture, that Neanderthal and Upper Paleolithic peoples either interbred or that the newcomers reworked Neanderthal implements "into their own kind of tools."BOOK, Coon, Carleton Stevens, The Origin of races, 1962, Knopf, New York, 548–49, Christopher Thomas Cairney in 1989 went further, laying out a rationale for hybridisation and adding a broader discussion of physical characteristics as well as commentary on interbreeding and its importance to adaptive European phenotypes. Cairney specifically discussed the "intermixture of racial elements" and "hybridisation."By the early 2000s, the majority of scholars supported the Out of Africa hypothesis,{{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."JOURNAL, Stringer, Chris, June 2003, Human evolution: Out of Ethiopia, Nature (journal), Nature, 423, 6941, 692–93, 695, 12802315, 10.1038/423692a, 2003Natur.423..692S, according to which anatomically modern humans left Africa about 50,000 years ago and replaced Neanderthals with little or no interbreeding.Yet some scholars still argued for hybridisation with Neanderthals. The most vocal proponent of the hybridisation hypothesis was Erik Trinkaus of Washington University.Dan Jones: The Neanderthal within., New Scientist 193.2007, H. 2593 (March 3), 28–32. Modern Humans, Neanderthals May Have Interbred{{dead link|date=February 2018|bot=medic}}{{cbignore|bot=medic}} ; Humans and Neanderthals interbred {{webarchive |url= |date=February 22, 2009}} Trinkaus claimed various fossils as products of hybridised populations, including the skeleton of a child found at Lagar Velho in PortugalWEB, Jim, Foley, The Lagar Velho 1 Skeleton,weblink Fossil Hominids FAQ, TalkOrigins Archive, July 31, 2000, July 6, 2017, WEB, Ian, Sample, Life on the edge: was a Gibraltar cave last outpost of the lost neanderthal?,weblink The Guardian, September 13, 2006, July 6, 2017, WEB,weblink Not a lasting last for the Neandertals, John D. Hawks, john hawks weblog, September 13, 2006, July 6, 2017, and the Peștera Muierii skeletons from Romania.JOURNAL, 10.1073/pnas.0608443103, Early modern humans from the Pestera Muierii, Baia de Fier, Romania, 2006, Soficaru, Andrei, Dobos, Adrian, Trinkaus, Erik, Proceedings of the National Academy of Sciences, 103, 46, 17196–201, 30052409, 2006PNAS..10317196S, 17085588, 1859909,

Genetic evidence

{{see|Neanderthal genetics}}File:Le Moustier skull in Berlin reconstitution.jpg|thumb|Le Moustier Neanderthal facial reconstitution, Neues MuseumNeues MuseumFile:Figure of Neanderthal, by Borivoje Žuža.jpg|thumb|Neanderthal figure representation, Museum of Republika SrpskaRepublika SrpskaIn 2010, geneticists announced that interbreeding had likely taken place,NEWS, Nicholas, Wade,weblink Genetic Data and Fossil Evidence Tell Differing Tales of Human Origins, The New York Times, July 26, 2012, May 5, 2015, a result confirmed in 2012.BOOK, Svante Pääbo, 2014, Neanderthal Man: In Search of Lost Genomes, New York, Basic Books, 978-0465054954, Neanderthal Man: In Search of Lost Genomes, Svante Pääbo, {{page needed|date=November 2017}} The genomes of all non-Africans (such as Europeans, East Asians, and Oceanians including Papua New GuineansJOURNAL, Draft full sequence of Neanderthal Genome, 7 May 2010, 10.1126/science.1188021, 20448178, 328, 5979, 5100745, Green RE, Krause J, Briggs AW, Maricic T, Stenzel U, Kircher M, Patterson N, Li H, Zhai W, Fritz MH, Hansen NF, Durand EY, Malaspinas AS, Jensen JD, Marques-Bonet T, Alkan C, Prüfer K, Meyer M, Burbano HA, Good JM, Schultz R, Aximu-Petri A, Butthof A, Höber B, Höffner B, Siegemund M, Weihmann A, Nusbaum C, Lander ES, Russ C, Novod N, Affourtit J, Egholm M, Verna C, Rudan P, Brajkovic D, Kucan Z, Gusic I, Doronichev VB, Golovanova LV, Lalueza-Fox C, de la Rasilla M, Fortea J, Rosas A, Schmitz RW, Johnson PL, Eichler EE, Falush D, Birney E, Mullikin JC, Slatkin M, Nielsen R, Kelso J, Lachmann M, Reich D, Pääbo S, Science, 710–22, 2010Sci...328..710G, ) include portions that are of Neanderthal origin,JOURNAL, 10.1093/molbev/msr024, An X-Linked Haplotype of Neandertal Origin is Present Among All Non-African Populations, 2011, Yotova, V., Lefebvre, J.-F., Moreau, C., Gbeha, E., Hovhannesyan, K., Bourgeois, S., Bédarida, S., Azevedo, L., Amorim, A., Sarkisian, T., Avogbe, P. H., Chabi, N., Dicko, M. H., Kou' Santa Amouzou, E. S., Sanni, A., Roberts-Thomson, J., Boettcher, B., Scott, R. J., Labuda, D., Molecular Biology and Evolution, 28, 7, 1957–62, 21266489, WEB, Jennifer, Viegas,weblink All Non-Africans Part Neanderthal, Genetics Confirm, DNews, July 18, 2011, a share estimated in 2014 to 1.5–2.1%.JOURNAL, K. Prüfer, etal, 2014, The complete genome sequence of a Neanderthal from the Altai Mountains, Nature, 505, 7481, 43–49, 10.1038/nature12886, 24352235, 4031459, 2014Natur.505...43P, This DNA is absent in Sub-Saharan Africans (Yoruba people and San subjects). Ötzi the iceman, Europe's oldest preserved mummy, was found to possess an even higher percentage of Neanderthal ancestry.WEB,weblink Neandertal ancestry 'Iced', john hawks weblog, August 15, 2012, July 6, 2017, The two percent of Neanderthal DNA in Europeans and Asians is not the same in all Europeans and Asians: in all, approximately 20% of the Neanderthal genome appears to survive in the modern human gene pool.JOURNAL, Resurrecting Surviving Neandertal Lineages from Modern Human Genomes, Science, 343, 6174, 1017–21, January 29, 2014, 2014Sci...343.1017V, Vernot, Benjamin, Akey, Joshua M., 10.1126/science.1245938, 24476670, Genomic studies suggest that modern humans mated with at least two groups of archaic humans: Neanderthals and Denisovans.NEWS, Mitchell, Alanna, DNA Turning Human Story into a Tell-All,weblink January 30, 2012, NY Times, January 31, 2012, Some researchers suggest admixture of 3.4–7.9% in modern humans of non-African ancestry, rejecting the hypothesis of ancestral population structure.JOURNAL, 1307.8263, Maximum likelihood evidence for Neandertal admixture in Eurasian populations from three genomes, Konrad, Lohse, Laurent A. F., Frantz, Populations and Evolution, 2013arXiv1307.8263L, 1307, 2013, 8263, , and others later suggesting Neanderthal admixture of 1.5-2.1%.JOURNAL, K. Prüfer, etal, 2014, The complete genome sequence of a Neanderthal from the Altai Mountains, Nature, 505, 7481, 43–49, 10.1038/nature12886, 24352235, 4031459, 2014Natur.505...43P, Detractors have argued and continue to argue that the signal of Neanderthal interbreeding may be due to ancient African substructure, meaning that the similarity is only a remnant of a common ancestor of both Neanderthals and modern humans and not the result of interbreeding.JOURNAL, Lowery, Robert K., Uribe, Gabriel, Jimenez, Eric B., Weiss, Mark A., Herrera, Kristian J., Regueiro, Maria, Herrera, Rene J., Neanderthal and Denisova genetic affinities with contemporary humans: Introgression versus common ancestral polymorphisms, Gene, 530, 1, 2013, 83–94, 0378-1119, 10.1016/j.gene.2013.06.005, 23872234, WEB,weblink Study casts doubt on human-Neanderthal interbreeding theory, Alok, Jha, The Guardian, August 14, 2012, February 19, 2015, John D. Hawks has argued that the genetic similarity to Neanderthals may indeed be the result of both structure and interbreeding, as opposed to just one or the other.JOURNAL, 10.1146/annurev-anthro-092412-155548, Significance of Neandertal and Denisovan Genomes in Human Evolution, 2013, Hawks, John, Annual Review of Anthropology, 42, 433–49, An approximately 40,000 year old anatomically-modern human skeleton from Peștera cu Oase, Romania, was found in 2015 to have a much larger proportion of DNA matching the Neanderthal genome than seen in humans of today, and this was estimated to have resulted from an interbreeding event as few as four generations earlier. However, this hybrid Romanian population does not appear to have made a substantial contribution to the genomes of later Europeans.NEWS, Ancient Romanian Could Have Had Neanderthal Great-Great-Grandfather, Reuters, NBC News, July 22, 2015, August 25, 2018,weblink JOURNAL, Qiaomei Fu, Mateja Hajdinjak, Oana Teodora Moldovan, Silviu Constantin, Swapan Mallick, Pontus Skoglund, Nick Patterson, Nadin Rohland, Iosif Lazaridis, Birgit Nickel, Bence Viola, Kay Prüfer, Matthias Meyer, Janet Kelso, David Reich, Svante Pääbo, An early modern human from Romania with a recent Neanderthal ancestor, Nature, August 13, 2015, 524, 7564, 216–219, 10.1038/nature14558, 26098372, 4537386, 2015Natur.524..216F, While some modern human nuclear DNA has been linked to the extinct Neanderthals, no mitochondrial DNA of Neanderthal origin has been detected, which in primates is almost always maternally transmitted.JOURNAL, Luo, Shiyu, Valencia, C. Alexander, Zhang, Jinglan, Lee, Ni-Chung, Slone, Jesse, Gui, Baoheng, Wang, Xinjian, Li, Zhuo, Dell, Sarah, 2018-11-21, Biparental Inheritance of Mitochondrial DNA in Humans, Proceedings of the National Academy of Sciences, 115, 51, en, 13039–13044, 10.1073/pnas.1810946115, 0027-8424, 30478036, 6304937, This observation has prompted the hypothesis that whereas female humans interbreeding with male Neanderthals were able to generate fertile offspring, the progeny of female Neanderthals who mated with male humans were either rare, absent or sterile.JOURNAL, Paul H., Mason, Roger V., Short, Neanderthal-human Hybrids, Hypothesis, 2011, 9, e1, 10.5779/hypothesis.v9i1.215, Eastern Neanderthals from the Altai show evidence of an introgression from modern humans not seen in western Neanderthals. This contribution to their genome derived from a modern human population that diverged from most other modern humans about 120 kya and expanded from Africa into Eurasia, but which was later largely replaced by a second expansion of modern humans out of Africa after 75,000 years ago that gave rise to modern Eurasians, although 2% of the genome of New Guineans derives from this earlier dispersal.JOURNAL, Pagani, Luca, Genomic analyses inform on migration events during the peopling of Eurasia, Nature, 13 October 2016, 538, 7624, 238–242, 2016Natur.538..238P, 10.1038/nature19792, 27654910, 5164938, Kuhlwilm et al. argue that the admixture between this early modern human group, modern Eurasians, and Neanderthals took place in Southern Arabia or the Levant and that the introgressed Siberian Neanderthals had spread there from the Middle East.

Interbreeding with Denisovans

Sequencing of the genome of a Denisovan, a distinct but related archaic hominin, from the Denisova cave in the Siberian Altai region has shown that 17% of its genome represents Neanderthal DNA.JOURNAL, 10.1126/science.340.6134.799, 23687020, More Genomes from Denisova Cave Show Mixing of Early Human Groups, Science, 340, 6134, 799, 2013, Pennisi, E, This Neanderthal DNA present in the Denisovan genome more closely resembled that found in the genome of a 120,000-year-old Neanderthal bone found in the same cave than that of Neanderthals from the Vindija cave in Croatia or the Mezmaiskaya cave in the Caucasus, suggesting that the gene flow came from a local interbreeding. However, the complete genome sequencing of DNA from a 90,000-year-old bone fragment, Denisova 11, showed it to have belonged to a Denisovan-Neanderthal hybrid whose father was a typical Denisovan with the Altai Neanderthal component dating to an interbreeding more than 300 generations earlier, but the specimen's mother was a Neanderthal belonging to a population more closely related to the Vindija Neanderthal than to the sequenced Altai Neanderthal genome. This suggests mobility or turnover among the distinct Neanderthal populations.WEB, Briggs, Helen, Cave girl was half Neanderthal, half Denisovan, BBC news, August 22, 2018, August 22, 2018,weblink WEB, Marshall, Michael, Prehistoric girl had parents belonging to different human species,weblink August 22, 2018, new Scientist, August 22, 2018, JOURNAL, Warren, Matthew, Mum's a Neanderthal, Dad's a Denisovan: First discovery of an ancient-human hybrid, Nature, 560, 7719, 417–418, 2018, 10.1038/d41586-018-06004-0, 30135540, 2018Natur.560..417W, NEWS, Netburn, Deborah, Found: An ancient hominin hybrid who had a Neanderthal for a mother and a Denisovan for a father, Los Angeles Times, August 22, 2018, August 25, 2018,weblink

mtDNA phylogeny

The mtDNA phylogeny of the Neanderthals, Denisovans, and modern humans produces a puzzling picture. Based on mtDNA sequences of Neanderthals from the Vindija and Altai sites, the most recent common ancestor of modern humans and Neanderthals lived approximately 440,000 years ago, while that between Neanderthals and Denisovans is far back as 830,000 years. This contrasts with nuclear genome analysis, in which the Neanderthals and Denisovans are sister groups.JOURNAL, Paabo, Svante, A mitochondrial genome sequence of a hominin from Sima de los Huesos, Nature, 505, 16 January 2014, 7483, 403–406, 2014Natur.505..403M, 10.1038/nature12788,weblink The mtDNA of the 400,000-year-old Sima de los Huesos sample, thought to be ancestral to Western European Neanderthals, is much closer to Denisovan mtDNA. Paabo et al. suggest several alternative explanations for this, including genetic substructure within the populations, introgression of mtDNA from one group to another, or incorrect taxonomic placement of the Sima de los Huesos hominins. Peyrégne et al. again suggested genetic substructure or introgression as possible explanations for an approximately 124,000-year-old German Neanderthal specimen with mtDNA that diverged from that of other Neanderthal specimens (except for Sima de Huesos) about 270,000 years ago, while its genomic DNA was consistent with divergence less than 150,000 years ago.


According to a 2014 study by Thomas Higham and colleagues of organic samples from European sites, Neanderthals died out in Europe between 41,000 and 39,000 years ago.{{efn|Higham et al did not study samples from sites outside Europe and they stated that further work was required to rule out later survival at Gorhams Cave, Gibraltar.}} New dating in Iberia, where Neanderthal dates as late as 24,000 years had been reported before, now suggests evidence of Neanderthal survival in the peninsula after 42,000 years ago is almost non-existent.(File:CurratExcoffierNeandethalmtDNA.jpg|thumb|upright=1.5|mtDNA-based simulation of modern human expansion in Europe starting 1,600 generations ago. Neanderthal range in light greyJOURNAL, 10.1371/journal.pbio.0020421, Modern Humans Did Not Admix with Neanderthals during Their Range Expansion into Europe, 2004, Currat, Mathias, Excoffier, Laurent, PLoS Biology, 2, 12, e421, 15562317, 532389, )Anatomically modern humans arrived in Mediterranean Europe between 45,000 and 43,000 years ago, so the two different human populations shared Europe for several thousand years.JOURNAL, Stein, Richard A, October 1, 2015, Copy Number Analysis Starts to Add Up,weblink Gen. Eng. Biotechnol. News, Genetic Engineering & Biotechnology News, Paper, 35, 17, 20, Neanderthals, which are thought to have come into conteact with modern humans approximately 80,000 years ago, appear to have survived until about 35,000 years ago in some regions of Europe., {{paywall}} The exact nature of biological and cultural interaction between Neanderthals and other human groups is contested.JOURNAL, Finlayson, C., Carrión, J. S., Rapid ecological turnover and its impact on Neanderthal and other human populations, Trends in Ecology & Evolution, 22, 4, 213–22, April 2007, 17300854, 10.1016/j.tree.2007.02.001, Possible scenarios for the extinction of the Neanderthals are:
  1. Neanderthals were a separate species from modern humans, and became extinct (because of climate change or interaction with modern humans) and were replaced by modern humans moving into their habitat between 45,000 and 40,000 years ago.NEWS, October 24, 2007, First genocide of human beings occurred 30,000 years ago,weblink Pravda, May 18, 2009, Jared Diamond has suggested a scenario of violent conflict and displacement.BOOK, Jared M., Diamond, Jared Diamond, The third chimpanzee: the evolution and future of the human animal, HarperCollins, New York City, 1992, 52, 978-0-06-098403-8, 60088352,
  2. Neanderthals were a contemporary subspecies that bred with modern humans and disappeared through absorption (interbreeding theory).
  3. Volcanic catastrophe: see Campanian Ignimbrite Eruption

Climate change

File:Skeleton and restoration model of Neanderthal La Ferrassie 1.jpg|thumb|left|Skeleton and restoration model of the La Ferrassie 1 Neanderthal man (National Museum of Nature and ScienceNational Museum of Nature and ScienceAbout 55,000 years ago, the climate began to fluctuate wildly from extreme cold conditions to mild cold and back in a matter of decades. Neanderthal bodies were well-suited for survival in a cold climate—their stocky chests and limbs stored body heat better than the Cro-Magnons. Neanderthals died out in Europe between 41,000 and 39,000 years ago, coinciding with the start of a very cold period.NEWS,weblink New dates rewrite Neanderthal story, BBC News, Pallab, Ghosh, August 20, 2014, NEWS, Kate, Wong,weblink The Mysterious Downfall of the Neandertals, Scientific American, August 1, 2009, Raw material sourcing and the examination of faunal remains found in the southern Caucasus suggest that modern humans may have had a survival advantage, being able to use social networks to acquire resources from a greater area. In both the Late Middle Palaeolithic and Early Upper Palaeolithic more than 95% of stone artifacts were drawn from local material, suggesting Neanderthals restricted themselves to more local sources.JOURNAL, Adler, Daniel S., Bar-Oz, Guy, Belfer-Cohen, Anna, Bar-Yosef, Ofer, Ahead of the Game: Middle and Upper Palaeolithic Hunting Behaviors in the Southern Caucasus, Current Anthropology, 2006, 47, 1, 89–118, 10.1086/432455, 10.1086/432455,weblink

Coexistence with modern humans

The exact nature of biological and cultural interaction between Neanderthals and other human groups has been contested. The dating of the period of overlap has been repeatedly revised. In 2011, this period of coexistence was extended when what were previously thought to be Neanderthal baby teeth, unearthed in 1964 from the Grotta del Cavallo in Italy, were identified as the oldest modern human remains discovered anywhere in Europe, dating from between 43,000 and 45,000 years ago.NEWS, Wilford, John Noble, John Noble Wilford, Fossil Teeth Put Humans in Europe Earlier Than Thought,weblink November 2, 2011, New York Times, August 27, 2014, At that time, Neanderthal samples from Iberia were thought to have dated as recently as 24,000 years ago, but a 2014 study of Neanderthal bones and tools led to a redating of these samples, and indicated that Neanderthals died out in Europe between 41,000 and 39,000 years ago, with the two different human populations sharing Europe only for about 5,000 years. However, a 2019 reanalysis of skull fragments from southern Greece assumed to have belonged to Neanderthals concluded that they belonged to a modern human, and dated them to 210,000 years ago, vastly extending the period of potential overlap in Europe between the two groups.NEWS, Zimmer, Carl, Carl Zimmer, 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.,weblink 10 July 2019, The New York Times, 11 July 2019, NEWS, Staff, 'Oldest remains' outside Africa reset human migration clock,weblink 10 July 2019,, 10 July 2019, JOURNAL, Harvati, Katerina, et al., Apidima Cave fossils provide earliest evidence of Homo sapiens in Eurasia,weblink 10 July 2019, Nature (journal), Nature, 10.1038/s41586-019-1376-z, 10 July 2019, File:Expansion of early modern humans from Africa.jpg|thumb|upright=1.5|Replacement of Neanderthals by early modern humansearly modern humansNeanderthals inhabited that continent long before the arrival of modern humans. These modern humans may have introduced a disease that contributed to the extinction of Neanderthals,{{Citation needed|date=July 2019}} and that may be added to other recent explanations for their extinction. When Neanderthal ancestors left Africa potentially as early as over 800,000 years ago they adapted to the pathogens in their European environment, unlike modern humans, who adapted to African pathogens. This transcontinental movement is known as the Out of Africa model. If contact between humans and Neanderthals occurred in Europe and Asia the first contact may have been devastating to the Neanderthal population, because they would have had little, if any, immunity to the African pathogens.{{Citation needed|date=July 2019}} More recent historical events in Eurasia and the Americas show a similar pattern, whereby the unintentional introduction of viral or bacterial pathogens to unprepared populations has led to mass mortality and local population extinction.Wolff, H. "Result Filters." National Center for Biotechnology Information. U.S. National Library of Medicine, July 2010. Web. October 22, 2014. The most well-known example of this is the arrival of Christopher Columbus to the New World, which brought and introduced foreign diseases when he and his crew arrived to a native population who had no immunity.Anthropologist Pat Shipman suggested that domestication of the dog could have played a role in Neanderthals' extinction.NEWS, McKie, Robin, March 1, 2015, How hunting with wolves helped humans outsmart the Neanderthals,weblink The Guardian, May 5, 2016,

History of research{{anchor|Discovery}}

{{Multiple image | align = right | direction = vertical | width = 100
| image1 = Schmerling Planche I.jpg| width1 = | caption1 = Engis 2, child (1829)
| image2 =Gibraltar_Skull_(1)a.jpg| width2 = | caption2 = Gibraltar 1, female (1848)
| image3 = Neander1.jpg| width3 = | caption3 = Neanderthal 1, male (upper skull 1856, left-cheek 2000)
| image4 = Spy Skull.jpg | width4 = | caption4 = Spy 2 skull, sex unclearARTICLE, S., Genovés, 1954, The problem of the sex of certain fossil hominids, with special reference to the Neandertal skeletons from Spy, The Journal of the Royal Anthropological Institute of Great Britain and Ireland, 84, 1/2, 131–44, 10.2307/2844006, 2844006, (1886)
width5 = | caption5 = Krapina 3, female (1899)}}Neanderthal fossils were first discovered in 1829 in the Engis caves (the partial skull dubbed Engis 2), in present-day Belgium by Philippe-Charles Schmerling and the Gibraltar 1 skull in 1848 in the Forbes' Quarry, Gibraltar. These finds were not, at the time,recognized as representing an archaic form of humans.The first discovery which was recognized as representing an archaic form of humans was made in August 1856, three years before Charles Darwin's On the Origin of Species was published.This was the discovery of the type specimen, Neanderthal 1, in a limestone quarry (Feldhofer Cave), located in Neandertal Valley in the German Rhineland, about {{convert|12|km|mi|0|abbr=on}} east of Düsseldorf).The find consisted of a skull cap, two femora, three bones of the right arm, two of the left arm, parts of the left ilium, fragments of a scapula, and ribs. The workers who recovered the objects originally thought them to be the remains of a cave bear. However, they eventually gave the material to amateur naturalist Johann Carl Fuhlrott, who turned the fossils over to anatomist Hermann Schaaffhausen.WEB,weblink Ausflug und Urlaub im Kreis Mettmann, September 21, 2017,, de, December 5, 2017, WEB,weblink Stadt Erkrath,, December 5, 2017, WEB, Homo neanderthalensis,weblink Smithsonian Institution, May 18, 2009, dead,weblink" title="">weblink May 21, 2009, To date, the bones of over 400 Neanderthals have been found.WEB, New Evidence on the Role of Climate in Neanderthal Extinction,weblink Science Daily,
  • 1829: A damaged skull of a Neanderthal child, Engis 2, is discovered in Engis, Netherlands (now Belgium).
  • 1848: A female Neanderthal skull, Gibraltar 1, is found in Forbes' Quarry, Gibraltar, but its importance is not recognised.
  • 1856: Limestone miners discover the Neanderthal-type specimen, Neanderthal 1, in Neandertal, western Prussia (now Germany).
  • 1864: William King is the first to recognise Neanderthal 1 as belonging to a separate species, for which he gives the scientific name Homo neanderthalensis. He then changed his mind on placing it in the genus Homo, arguing that the upper skull was different enough to warrant a separate genus since, to him, it had likely been "incapable of moral and theistic conceptions."
  • 1880: The mandible of a Neanderthal child is discovered in a secure context in Å ipka cave, in the Austro-Hungarian Empire (now the Czech Republic), associated with cultural debris, including hearths, Mousterian tools, and bones of extinct animals.
  • 1886: Two well-preserved Neanderthal skeletons are found at Spy, Belgium, making the hypothesis that Neanderthal 1 was only a diseased modern human difficult to sustain.ENCYCLOPEDIA, Encyclopedia Britannica, 1982, Neanderthal Man, 15th,
  • 1899: Sand excavation workers find hundreds of fragmentary Neanderthal remains representing at least 12 and likely as much as 70 individuals on a hill in Krapina, in the Austro-Hungarian Empire (now Croatia).
  • 1908: A very well preserved Neanderthal, La Chapelle-aux-Saints 1, is found in its eponymous site in France,WEB,weblink Neanderthal – Homo neanderthalensis – Details – Key Fossils – La Chapelle-aux-Saints, Encyclopedia of Life

, July 18, 2016, said by the excavators to be a burial, a claim still heatedly contested. For historical reasons it remains the most famous Neanderthal skeleton.{{rp|15}}
  • 1912: Marcellin Boule publishes his now discredited influential study of Neanderthal skeletal morphology based on La Chapelle-aux-Saints 1.
  • 1953–1957: Ten Neanderthal skeletons are excavated in Shanidar Cave, Iraqi Kurdistan, by Ralph Solecki and colleagues.
  • 1975: Erik Trinkaus's study of Neanderthal feet strongly argues that Neanderthals walked like modern humans.
  • 1981: The site of Bontnewydd, Wales yielded an early Neanderthal tooth, the most north-western Neanderthal remain ever.
  • 1987: Israeli Neanderthal Kebara 2 is dated (by TL and ESR) to 60,000 BP, thus later than the Israeli anatomically modern humans dated to 90,000 and 80,000 BP at Qafzeh and Skhul.
  • 1997: Matthias Krings et al. are the first to amplify Neanderthal mitochondrial DNA (mtDNA) using a specimen from Feldhofer grotto in the Neander valley.
  • 2005: The Max Planck Institute for Evolutionary Anthropology and associated institutions launch the Neanderthal genome project to sequence the Neanderthal nuclear genome.JOURNAL, Green, Richard E., Krause, Johannes, Ptak, Susan E., Briggs, Adrian W., Ronan, Michael T., Simons, Jan F., Du, Lei, Egholm, Michael, Rothberg, Jonathan M., Paunovic, Maja, Pääbo, Svante, Analysis of one million base pairs of Neanderthal DNA, Nature, 444, 7117, 2006, 330–36, 0028-0836, 2006Natur.444..330G, 17108958, 10.1038/nature05336,
  • 2010: Discovery of Neanderthal admixture in the genome of modern populations.JOURNAL, Sankararaman, S., Patterson, N., Li, H., Pääbo, S., Reich, D, Akey, J.M., The Date of Interbreeding between Neandertals and Modern Humans, PLoS Genetics, 2012, 8, 10, e1002947, 10.1371/journal.pgen.1002947, 23055938, 3464203, JOURNAL, Yang, M.A., Malaspinas, A.S., Durand, E.Y., Slatkin, M., Ancient Structure in Africa Unlikely to Explain Neanderthal and Non-African Genetic Similarity, Molecular Biology and Evolution, 2012, 29, 10, 2987–95, 10.1093/molbev/mss117, 22513287, 3457770,
  • 2014: A comprehensive dating of Neanderthal bones and tools from hundreds of sites in Europe dates the disappearance of Neanderthals to 41,000 and 39,000 years ago.BOOK,weblink The Human Lineage by Matt Cartmill, Fred H. Smith, 2009-03-30, September 27, 2015, 9780471214915, Cartmill, Matt, Smith, Fred H.,
  • 2018: Report on the complete genomic sequence of Denisova 11, a first generation of Neanderthal-Denisovan hybrid.
  • 2019: 257 hominin footprints were revealed by archaeologists at Le Rozel (Normandy, France). This finding, the vast majority of which consists of young footprints, has proved that Neanderthals lived in this area 80,000 years ago.WEB,weblink French Neanderthals Had Lots of Kids, Fossil Footprints Show, HAARETZ, live, JOURNAL, Cliquet, Dominique, Laisné, Gilles, Verna, Christine, Berillon, Gilles, Duveau, Jérémy, 2019-09-09, The composition of a Neandertal social group revealed by the hominin footprints at Le Rozel (Normandy, France),weblink Proceedings of the National Academy of Sciences, en, 201901789, 10.1073/pnas.1901789116, 0027-8424, 31501334,


Notable European Neanderthals

{{Multiple image | align = right | direction = vertical | width=100
| image1 = Ferrassie skull.jpg| width1 = | caption1 = La Ferrassie 1, skull cast
| image2 = Le_Moustier_1_before_being_bombed.jpg| width2 = | caption2 = Le Moustier 1 in 1909
| image3 = Museum_für_Vor-_und_Frühgeschichte_Berlin_069A.jpg| width3 = | caption3 = Le Moustier 1 in 2011
| image4 = Shanidar_skull.jpg | width4 = | caption4 = Shanidar 1, skull cast
| image5 = Neanderthal-burial.gif | width5 = | caption5 = Kebara 2
| image6 = Skull of Teshik-Tash Boy.jpg| width6 = | caption6 = Teshik-Tash 1
}}Remains of more than 300 European Neanderthals have been found. For the most important, see List of human evolution fossils.
  • Neanderthal 1: The first human bones recognised as showing a non-modern anatomy. Discovered in 1856 in a limestone quarry at the Feldhofer grotto in Neanderthal, Western Prussia (now Germany), they consist of a skull cap, the two femora, the three right arm bones, two left arm bones, the ilium, and fragments of a scapula and ribs.
  • La Chapelle-aux-Saints 1: Called the Old Man, a fossilised skeleton discovered in La Chapelle-aux-Saints, France, by A. and J. Bouyssonie, and L. Bardon in 1908. Characteristics include a low vaulted cranium and large browridge typical of Neanderthals. Estimated to be about 60,000 years old, the specimen was severely arthritic and had lost all his teeth long before death, leading some to suggest he was cared for by others.
  • La Ferrassie 1: A fossilised skull discovered in La Ferrassie, France, by R. Capitan in 1909. It is estimated to be 70,000 years old. Its characteristics include a large occipital bun, low-vaulted cranium and heavily worn teeth.
  • Le Moustier 1: One of the rare nearly complete Neanderthal skeletons to be discovered, it was excavated by a German team in 1908, at Peyzac-le-Moustier, France. Sold to a Berlin museum, the post cranial skeleton was bombed and mostly destroyed in 1945, and parts of the mid face were lost sometime after then. The skull, estimated to be less than 45,000 years old, includes a large nasal cavity and a less developed brow ridge and occipital bun than seen in other Neanderthals. The Mousterian tool techno-complex is named after its discovery site.

Notable Southwest Asian Neanderthals

Remains of more than 70 Southwest Asian Neanderthals have been found. For a complete list see List of Southwest Asian Neanderthals.
  • Shanidar 1 to 10: Eight Neanderthals and two pre-Neanderthals (Shanidar 2 and 4) were discovered in the Zagros Mountains in Iraqi Kurdistan. One of the skeletons, Shanidar 4, was once thought to have been buried with flowers, a theory no longer accepted. To Paul B. Pettitt the "deliberate placement of flowers has now been convincingly eliminated", since "[a] recent examination of the microfauna from the strata into which the grave was cut suggests that the pollen was deposited by the burrowing rodent Meriones tersicus, which is common in the Shanidar microfauna and whose burrowing activity can be observed today".The Neanderthal Dead, exploring mortuary variability in middle paleolithic eurasia. Paul B. Pettitt (2002)
  • Amud 1: A male adult Neanderthal, dated to roughly 55,000 BP, and one of several found in a cave at Nahal Amud, Israel. At {{Convert|178|cm|0|abbr=on}}, it is the tallest known Neanderthal. It also has the largest cranial capacity of all extinct hominins: 1,736 cm3.BOOK, Dmitra, Papagianni, Michael, Morse, The Neanderthals Rediscovered, Thames & Hudson, 2013, 978-0-500-05177-1, WEB,weblink Homo neanderthalensis – The Neanderthals, Australian Museum, June 26, 2014,
  • Kebara 2: A male adult post-cranial skeleton, dated to roughly 60,000 BP, that was discovered in 1983 in Kebara Cave, Israel. It has been studied extensively, for its hyoid, ribcage, and pelvis are much better preserved than in all other Neanderthal specimens.

Notable Central Asian Neanderthal

  • Teshik-Tash 1: An 8–11-year-old skeleton discovered in Uzbekistan by Okladnikov in 1938. This is the only fairly complete skeleton discovered to the east of Iraq. Okladnikov claimed it was a deliberate burial, but this is debated.


This section describes bones with Neanderthal traits in chronological order.

Mixed with H. heidelbergensis traits

  • > 350 ka: Sima de los Huesos c. 500:350 ka agoJOURNAL, 10.1006/jasc.2002.0834, The Sima de los Huesos Hominids Date to Beyond U/Th Equilibrium (>350kyr) and Perhaps to 400–500kyr: New Radiometric Dates, 2003, Bischoff, James L., Shamp, etal, Journal of Archaeological Science, 30, 3, 275–80,weblink
JOURNAL, Arsuaga JL, Martínez I, Gracia A, Lorenzo C, The Sima de los Huesos crania (Sierra de Atapuerca, Spain). A comparative study, Journal of Human Evolution, 33, 2–3, 219–81, 1997, 9300343, 10.1006/jhev.1997.0133,
  • 350–200 ka: Pontnewydd 225 ka ago.
  • 200–135 ka: Atapuerca,WEB, C. David, Kreger, Homo neanderthalensis,weblink, May 16, 2009, VértesszÅ‘lÅ‘s, Ehringsdorf, Casal de' Pazzi, Biache, La Chaise, Montmaurin, Prince, Lazaret, Fontéchevade

H. neanderthalensis fossils

  • 130–50 ka: Krapina, Saccopastore skulls, Malarnaud, Altamura, Gánovce, Denisova, Okladnikov, Pech de l'Azé, Tabun 120–{{val|100|5}} ka,JOURNAL, 10.1038/363252a0, May 1993, Mcdermott, F, Grün, R, Stringer, Cb, Hawkesworth, Cj, Mass-spectrometric U-series dates for Israeli Neanderthal/early modern hominid sites, 363, 6426, 252–55, 0028-0836, 8387643, Nature, 1993Natur.363..252M, Shanidar 1 to 9 80–60 ka, La Ferrassie 1 70 ka, Kebara 60 ka, Régourdou, Mt. Circeo, Combe Grenal, Erd 50 ka, La Chapelle-aux Saints 1 60 ka, Amud I {{val|53|8}} ka,JOURNAL, 10.1002/gea.1017, August 2002, W. Jack, Rink, H.P., Schwarcz, H.K., Lee, J., Rees-Jones, R., Rabinovich, E., Hovers, Electron spin resonance (ESR) and thermal ionization mass spectrometric (TIMS) 230Th/234U dating of teeth in Middle Paleolithic layers at Amud Cave, Israel, 16, 6, 701–17, Geoarchaeology, JOURNAL, 10.1006/jasc.1998.0334, March 1999, Valladas, Hélène, N., Merciera, L., Frogeta, E., Hoversb, J.L., Joronc, W.H., Kimbeld, Y., Rak, TL Dates for the Neanderthal Site of the Amud Cave, Israel, 26, 3, 259–68, Journal of Archaeological Science, Teshik-Tash.
  • In radiocarbon range, > 50 ka: Le Moustier, Feldhofer, La Quina, l'Hortus, Kulna, Å ipka, Saint Césaire, Bacho Kiro, El Castillo, Bañolas, El Sidrón (48±3 cal ka),JOURNAL, 10.1111/j.1475-4754.2012.00671.x, March 20, 2012, R. E., Wood, T. F. G., Higham, T., de Torres, N., Tisnérate-Laborde, H., Valladas, J. E., Ortiz, C., Lalueza-Fox, S., Sánchez-Moral, J. C., Cañaveras, A., Rosas, D., Santamaría, M., de la Rasilla, A new date for the Neanderthals from El Sidrón Cave (Asturias, Northern Spain), 55, 1, 148–58, Archaeometry, 10261/79592, Arcy-sur-Cure, Châtelperron, Figueira Brava, Mezmaiskaya (41±1 cal ka), Zafarraya, Vindija, Velika Pećina.

H. s. sapiens with traits reminiscent of Neanderthals