Neanderthal

{{Short description|Extinct human species}} {{other uses|Neanderthal (disambiguation)}} {{For|Neanderthals and other related species in popular culture|Caveman}} {{good article}} {{Use British English|date=October 2024}} {{Use mdy dates|date=November 2025}}

{{Speciesbox | taxon = Homo neanderthalensis | extinct = yes | name = Neanderthal | fossil_range = [[Middle Pleistocene|Middle]]–[[Late Pleistocene]]<ref name=Richards2021/> {{Fossil range|0.24|0.04}}{{Period fossil range|Quaternary|0.24|0.04}} | image = Neanderthal at AMNH.jpg | image_caption = An approximate reconstruction of a Neanderthal skeleton. The central rib-cage (including the sternum) and parts of the pelvis are from modern humans. | image_alt = Slightly angled head-on view of a Neanderthal skeleton, stepping forward with the left leg | authority = [[William King (geologist)|King]], 1864 | synonyms = {{#invoke:collapsible list|main| title=''Homo''| *''H. stupidus'' [[Ernst Haeckel|Haeckel]], 1895<ref name=haeckel>{{cite book |url=https://archive.org/details/systematischephy03haec/page/601 |first=E. |last=Haeckel |author-link=Ernst Haeckel |year=1895 |title=Systematische Phylogenie: Wirbelthiere |publisher=G. Reimer |language=de |page=601}}</ref> *''H. europaeus primigenius'' Wilser, 1898 *''H. primigenius'' [[Gustav Albert Schwalbe|Schwalbe]], 1906<ref name=schwalbe>{{cite book |url=https://www.biodiversitylibrary.org/item/126001#page/13 |first=G. |last=Schwalbe |author-link=Gustav Albert Schwalbe |year=1906 |title=Studien zur Vorgeschichte des Menschen |language=de |trans-title=Studies on the pre-history of man |publisher=Stuttgart, E. Nägele |doi=10.5962/bhl.title.61918|hdl= 2027/uc1.b4298459}}</ref> *''H. antiquus'' Adloff, 1908 *''H. transprimigenius mousteriensis'' Farrer, 1908 *''H. mousteriensis hauseri'' [[Hermann Klaatsch|Klaatsch]] 1909<ref name=klaatsch>{{cite journal |url= https://gallica.bnf.fr/ark:/12148/bpt6k5864483g/f25.image. |first=H. |last=Klaatsch |author-link=Hermann Klaatsch |year=1909 |title=Preuves que l{{'}}''Homo Mousteriensis Hauseri'' appartient au type de Neandertal |trans-title=Evidence that ''Homo Mousteriensis Hauseri'' belongs to the Neanderthal type |language=fr |journal=L'Homme Préhistorique |volume=7 |pages=10–16}}</ref><ref name="romeo1979">{{cite book |last=Romeo |first=L. |title=Ecce Homo!: a lexicon of man |publisher=John Benjamins Publishing Company |year=1979 |page=92 |url=https://books.google.com/books?id=jj1iftPqCssC&pg=PA92 |isbn=90-272-2006-9}}</ref> *''H. priscus'' [[Wilhelm Krause (anatomist)|Krause]], 1909 *''H. chapellensis'' [[Hugo Berthold von Buttel-Reepen|von Buttel-Reepen]], 1911 *''H. calpicus'' [[Arthur Keith|Keith]], 1911 *''H. acheulensis moustieri'' Wiegers, 1915 *''H. lemousteriensis'' Wiegers, 1915 *''H. naulettensis'' Baudouin, 1916 *''H. sapiens neanderthalensis'' Kleinshmidt, 1922 *''H. heringsdorfensis'' Werthe, 1928 *''H. galilensis'' Joleaud, 1931 *''H. primigenius galilaeensis'' Sklerj, 1937 *''H. kiikobiensis'' Bontsch-Osmolovskii, 1940 *''H. sapiens krapinensis'' Campbell, 1962 *''H. erectus mapaensis'' Kurth, 1965 }} {{#invoke:collapsible list|main|title=''Palaeoanthropus''| *''P. neanderthalensis'' [[Theodore McCown|McCown]] and [[Sir Arthur Keith|Keith]], 1939<ref name=mccown1939>{{cite book |first1=T. |last1=McCown |first2=A. |last2=Keith |author-link2=Arthur Keith |year=1939 |title=The stone age of Mount Carmel. The fossil human remains from the Levalloisso-Mousterian |volume=2 |publisher=Clarenden Press}}</ref> *''P. heidelbergensis'' [[Theodore McCown|McCown]] and [[Sir Arthur Keith|Keith]], 1939<ref name=mccown1939/> *''P. ehringsdorfensis'' Paterson, 1940<ref name=mccown1939/> *''P. krapinensis'' Sergi, 1911<ref name=mccown1939/> *''P. palestinensis'' [[Theodore McCown|McCown]] and [[Sir Arthur Keith|Keith]], 1939<ref name=mccown1939/> *''P. europaeus'' Sergi, 1910 }} {{#invoke:collapsible list|main|title=''Protanthropus''| *''P. atavus'' [[Ernst Haeckel|Haeckel]], 1895 *''P. tabunensis'' Bonarelli, 1944 }} {{#invoke:collapsible list|main|title=''Acanthropus''| *''A. neanderthalensis'' Arldt, 1915 *''A. primigenius'' [[Othenio Abel|Abel]], 1920 *''A. neanderthalensis'' [[William Boyd Dawkins|Dawkins]], 1926 }} | synonyms_ref = <ref>{{cite book |url= https://books.google.com/books?id=jE7gBAAAQBAJ&pg=PA508 |first1=F. S. |last1=Szalay |first2=E. |last2=Delson |year=2013 |title=Evolutionary history of the Primates |publisher=Academic Press |page=508 |isbn=978-1-4832-8925-0}}</ref> }}

'''Neanderthals''' ({{IPAc-en||n|i|ˈ|æ|n|d|ə(r)|ˌ|t|ɑː|l|,_|n|eɪ|-|,_|-|ˌ|θ|ɑː|l}} {{respell|nee|AN|də(r)|TAHL|,_|nay-|,_|-|THAHL}};<ref>{{Cite LPD|3|page=534}}</ref> '''''Homo neanderthalensis''''' or sometimes '''''Homo sapiens neanderthalensis''''') are an [[extinction|extinct]] group of archaic [[Homo|humans]] who inhabited Europe and Western and Central Asia during the [[Middle Pleistocene|Middle]] to [[Late Pleistocene]]. [[Neanderthal extinction]] occurred roughly 40,000 years ago with the immigration of modern humans ([[Cro-Magnon]]s), but [[Neanderthals in Gibraltar]] may have persisted for thousands of years longer.

The first recognised Neanderthal [[fossil]], [[Neanderthal 1]], was discovered in 1856 in the [[Neander Valley]], Germany. At first, Neanderthal 1 was considered to be one of the [[racial hierarchy|lower races]] in accord with [[historical race concepts]]. As more fossils were discovered through the early 20th century, Neanderthals were characterised as a unique species of underdeveloped human, in particular by [[Marcellin Boule]]. By the mid-twentieth century, it was believed that [[human evolution]] progressed from an ape-like ancestor through a "Neanderthal phase" to modern humans. This gave way to the [[recent African origin of modern humans|"Out of Africa" theory]] in the 1970s. Sequencing of the [[Neanderthal genetics|Neanderthal genome]] in 2010 revealed that Neanderthals [[interbreeding between archaic and modern humans|interbred with modern humans]].

[[Neanderthal anatomy]] is characterised by a long and low skull, a heavy and rounded brow ridge (supraorbital torus), an [[occipital bun]] (bony projection) at the back of the skull, strong teeth and jaws, a wide chest, and short limbs. These traits gradually became more frequent through the [[Middle Pleistocene]] of Europe, possibly due to [[natural selection]] in a cold climate, as well as [[genetic drift]] when populations collapsed during [[glacial period]]s. Neanderthal specimens vary in height from {{cvt|147.5|to|177|cm|ftin|0}}, with average male dimensions estimated at {{cvt|165|cm|ftin}} and {{cvt|75|kg}}. While Neanderthal brain volume and ratio to body size averaged higher than any living human population — {{cvt|1640|cc}} for males and {{cvt|1460|cc}} for females — their brain organisation differed from modern humans in areas related to cognition and language, which could explain the comparative simplicity of [[Neanderthal behaviour]] to Cro-Magnons in the archaeological record.

Neanderthals maintained a low population and suffered [[inbreeding depression]], which may have impeded their ability to progress technologically. They produced [[Mousterian]] [[stone tool]]s (a [[Middle Palaeolithic]] [[industry (archaeology)|industry]]) and possibly wore blankets and [[poncho]]s. They maintained and might have [[control of fire by early humans|created fire]]. They predominantly ate whatever was abundant close to home, usually big game as well as plants and mushrooms. Neanderthals were frequently victims of major physical traumas and [[animal attack]]s. Examples of [[Art of the Middle Paleolithic|Palaeolithic art]] have been inconclusively attributed to Neanderthals, namely possible ornaments made from bird claws and feathers; collections of unusual objects including crystals and fossils; and engravings. Neanderthals buried their dead, but there is no clear indication that they believed in [[afterlife|life after death]].

==Etymology== Neanderthals are named after the [[Neandertal (valley)|Neander Valley]] in which the first identified specimen was found. The valley was spelled ''Neanderthal'' and the species was spelled ''Neanderthaler'' in German until the [[German Orthographic Conference of 1901|spelling reform of 1901]].{{efn|The German [[Standard German phonology#Consonants|/t/ phoneme]] was frequently spelled ''th'' from the 15th to 19th centuries until the [[German Orthographic Conference of 1901]]. The German spelling ''[[Dale (landform)#Etymology|Thal]]'' ("valley", a [[cognate]] of English ''[[:wikt:dale|dale]]'') changed to ''Tal'', and the ''h'' was also dropped from ''Neandertal'' for the valley and ''Neandertaler'' for the species.<ref name=Kreisstadt2017/>}} The spelling ''Neandertal'' for the species is occasionally seen in English, even in scientific publications, but the scientific name, ''H. neanderthalensis'', is always spelled with ''th'' according to the [[principle of priority]]. The vernacular name of the species in German is always ''Neandertaler'' ("inhabitant of the Neander Valley"), whereas ''Neandertal'' always refers to the valley.{{efn|In [[Mettmann]], "Neander Valley", there is a local idiosyncrasy in use of the outdated spellings with ''th'', such as with the [[Neanderthal Museum]] (but the name is in English [German would require ''Neandertalermuseum'']), the [[Neanderthal station]] (Bahnhof Neanderthal), and some other rare occasions meant for tourists. Beyond these, city convention is to use ''th'' when referring to the species.<ref name=Kreisstadt2017>{{cite web |url=https://www.mettmann.de/neandertal/schreibweise.php |title=Neandertal oder Neanderthal? Was ist denn nun richtig? |publisher=Kreisstadt Mettmann |archive-url=https://web.archive.org/web/20200809042411/https://www.mettmann.de/neandertal/schreibweise.php |trans-title=Neandertal or Neanderthal? So which is actually right? |access-date=February 1, 2017 |archive-date=August 9, 2020 |quote=Heute sollten Ortsbezeichnungen das 'Neandertal' ohne 'h' bezeichnen. Alle Namen, die sich auf den prähistorischen Menschen beziehen, führen das 'h'. (Nowadays, place names should refer to the Neander Valley ['Neandertal'] without an 'h'. All names referring to the prehistoric humans have the 'h'.)}}</ref>}}<ref name=Kreisstadt2017/> The valley itself was named after the late 17th century German theologian and hymn writer [[Joachim Neander]], who often visited the area.<ref name=Schmitz2002/> His grandfather, a musician, had changed the family name from the original German ''[[Neumann]]'' "new man" (cf. "[[Newman]]") to the Graeco-Roman form ''Neander'' (deriving with Greek {{wikt-lang|grc|ἀνήρ}} ''ănḗr'' "man"), following the fashion of the time.<ref name=EB1911>{{cite EB1911 |wstitle=Neander, Joachim |volume=19 |pages=320–321}}</ref>

The ''th'' in ''Neanderthal'' can be pronounced as {{IPA|/t/}} (hence {{IPAc-en|n|i|ˈ|æ|n|d|ər|t|ɑː|l}})<ref>{{cite web |url=https://www.collinsdictionary.com/dictionary/english/neanderthal |title=Neanderthal |website=Collins English Dictionary |access-date=February 18, 2020}}</ref> following the German convention<ref>{{cite magazine|first=B.|last=Alex|year=2016|title=Is It Neander-TAL or Neander-THAL?|magazine=[[Discover (magazine)|Discover]]|access-date=March 3, 2025|url=https://www.discovermagazine.com/planet-earth/is-it-neander-tal-or-neander-thal}}</ref> or Anglicized as fricative /[[voiceless dental fricative|θ]]/ (hence {{IPAc-en|n|i|ˈ|æ|n|d|ər|θ|ɑː|l}}), [[Spelling pronunciation|as per]] the standard [[Pronunciation of English ⟨th⟩|English pronunciation of ''th'']].<ref>{{cite web |url=https://www.merriam-webster.com/dictionary/Neanderthal |title=Neanderthal |website=Merriam-Webster Dictionary |access-date=February 18, 2020}}</ref><ref>{{cite web |url=https://www.ahdictionary.com/word/search.html?q=Neanderthal |title=Neanderthal |website=American Heritage Dictionary |access-date=February 18, 2020}}</ref>

[[Neanderthal 1]], the [[type specimen]], 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".<ref>{{cite book |url=https://archive.org/stream/lecturesonmanhi00huntgoog#page/n330/mode/2up |title=Lectures on man: his place in creation, and in the history of the earth |last=Vogt |first=K. C. |author-link=Carl Vogt |publisher=Longman, Green, Longman and Roberts |year=1864 |place=London |pages=302, 473}}</ref> In 1863, Irish geologist [[William King (geologist)|William King]] extended the name "Neanderthal man" from the individual specimen to the entire species.<ref name="King64">{{Cite journal |last1=King |first1=W. |author-link=William King (geologist) |year=1864 |title=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 |url=https://biodiversitylibrary.org/page/29371003 |journal=Report of the British Association for the Advancement of Science, Notices and Abstracts, Newcastle-upon-Tyne, 1863 |volume=33 |pages=81–82 |via=Biodiversity Heritage Library}}</ref><ref name="MurrNashSeoiMcCo15">{{cite journal |last1=Murray |first1=J. |title=The contribution of William King to the early development of palaeoanthropology |year=2015 |journal=Irish Journal of Earth Sciences |volume=33 |pages=1–16 |doi=10.3318/ijes.2015.33.1 |jstor=10.3318/ijes.2015.33.1 |last2=Nasheuer |first2=H. P. |last3=Seoighe |first3=C. |last4=McCormack |first4=G. P. |last5=Williams |first5=D. M. |last6=Harper |first6=D. A. T. |url=https://durham-repository.worktribe.com/output/1406049 }}</ref><ref>{{cite journal |first=A. K. |last=Winner |year=1964 |title=Terminology |journal=Current Anthropology |volume=5 |issue=2 |pages=119–122 |doi=10.1086/200469 |jstor=2739959 }}</ref>

==Taxonomy== {{See also|Human taxonomy}} ===Discovery=== [[File:Calotte crânienne, type de l'espèce Homo neanderthalensis, vallée de Néander.jpg|thumb|left|[[Calvaria (skull)|Skullcap]] of [[Neanderthal 1]], the [[type specimen]], at the [[Musée de l'Homme]], Paris|alt=A skullcap with a broad brow ridge and a large chip behind the right brow]] [[File:Neanderthal_excavation_site.JPG|alt=A grass field with 16 white-red-white-red poles spaced in diagonal lines, several plus-shaped stone blocks behind them, and a road is visible behind trees in the background|right|thumb|The site of [[Kleine Feldhofer Grotte]] where [[Neanderthal 1]] was discovered{{efn|After being mined for [[limestone]], the cave caved in and was lost by 1900. It was rediscovered in 1997 by archaeologists [[Ralf Schmitz (archaeologist)|Ralf Schmitz]] and [[Jürgen Thissen]].<ref name=Schmitz2002/>}}]] A number of Neanderthal fossils had been discovered before their antiquity was fully understood. The first Neanderthal remains—[[Engis 2]] (a skull)—were discovered in 1829 by Dutch/Belgian prehistorian [[Philippe-Charles Schmerling]] in the [[Schmerling Caves|Grottes d'Engis]], Belgium. He concluded that these "poorly developed" human remains must have been buried at the same time and by the same causes as the co-existing remains of extinct animal species.<ref>{{cite book |url=https://orbi.uliege.be/handle/2268/207986 |first=P. |last=Schmerling |author-link=Philippe-Charles Schmerling |year=1834 |title=Recherches sur les ossemens fossiles découverts dans les cavernes de la province de Liége |trans-title=Research on the fossil specimens discovered in the caves of Liège |publisher=P. J. Collardin |pages=30–32 |hdl=2268/207986}}</ref> In 1848, [[Gibraltar 1]] from [[Forbes' Quarry]] was presented to the Gibraltar Scientific Society by their Secretary Lieutenant Edmund Henry Réné Flint, but was thought to be a modern human skull.<ref>{{cite journal |first=A. |last=Menez |year=2018 |title=Custodian of the Gibraltar skull: the history of the Gibraltar Scientific Society |journal=Earth Sciences History |volume=37 |issue=1 |pages=34–62 |doi=10.17704/1944-6178-37.1.34|bibcode=2018ESHis..37...34M }}</ref>

In 1856, local schoolteacher [[Johann Carl Fuhlrott]] recognised bones from [[Kleine Feldhofer Grotte]] in Neander Valley—Neanderthal 1—as distinct from modern humans,{{efn|The bones were discovered by workers of [[Wilhelm Beckershoff]] and [[Friedrich Wilhelm Pieper]]. Initially, the workers threw the bones out as debris, but Beckershoff then told them to store the bones. Pieper asked Fuhlrott to come up to the cave and investigate the bones, which Beckershoff and Pieper believed belonged to a [[cave bear]].<ref name=Schmitz2002>{{cite journal |first1=R. W. |last1=Schmitz |first2=D. |last2=Serre |first3=G. |last3=Bonani |display-authors=et al. |year=2002 |title=The Neandertal type site revisited: interdisciplinary investigations of skeletal remains from the Neander Valley, Germany |journal=Proceedings of the National Academy of Sciences |volume=99 |issue=20 |pages=13342–13347 |doi=10.1073/pnas.192464099 |pmc=130635 |pmid=12232049 |bibcode=2002PNAS...9913342S|doi-access=free }}</ref>}} and gave them to German anthropologist [[Hermann Schaaffhausen]] to study in 1857. It comprised the cranium, thigh bones, right arm, left [[humerus]] and [[ulna]], left [[ilium (bone)|ilium]] (hip bone), part of the right [[shoulder blade]], and pieces of the [[rib]]s.<ref name="king1864">{{cite journal |first=W. |last=King |author-link=William King (geologist) |year=1864 |title=The reputed fossil man of the Neanderthal |journal=Quarterly Journal of Science |volume=1 |page=96 |url=https://biostor.org/reference/195152}}</ref><ref>{{cite journal |first=H. |last=Schaaffhausen |author-link=Hermann Schaaffhausen |year=1858 |title=Zur Kenntnis der ältesten Rassenschädel |journal=Archiv für Anatomie, Physiologie und Wissenschaftliche Medicin |pages=453–478 |language=de |trans-title=Acknowledging the oldest racial skull}}</ref>

===Research history=== [[File:Guide leaflet (1901) (14767192135).jpg|thumb|upright=1.8|[[William K. Gregory]]'s ''The Family Tree of Man'' exhibit at the [[American Museum of Natural History]], 1924 1) ''[[Notharctus]]'' 2) ''[[Propliopithecus]]'' 3) ''[[Dryopithecus]]'' 4) [[Java Man]] 5) [[Piltdown Man]] 6) [[Mauer 1|Heidelberg Man]] 7) '''Neanderthal Man''' 8) [[Cro-Magnon 1|Cro-Magnon Man]] 9) [[Blackfella|Australian Black-fellow]] (pejorative term for [[Aboriginal Australian]]s) 10) [[Hottentot (racial term)|Hottentot]] (pejorative term for the Southern African [[Khoisan]]) 11) [[Chinese people|Chinese]] 12) [[White Americans|American Caucasian]] ]] Following [[Charles Darwin]]'s 1859 ''[[On the Origin of Species]]'', Fuhlrott and Schaaffhausen argued that Neanderthal 1 represents a primitive [[racial hierarchy|lower]] human form, aligning more closely with non-human apes as well as [[Negroid]]s, [[Eskimo]]s, and [[Aboriginal Australian]]s (which were variably classified as separate species or [[human taxonomy#Homo sapiens subspecies|subspecies of human]] at the time).<ref name=drell2000/><ref name=king1864/><ref name=schlager>{{cite book |first1=S. |last1=Schlager |first2=U. |last2=Wittwer-Backofen |title=Handbook of Paleoanthropology |chapter=Images in Paleoanthropology: Facing Our Ancestors |year=2015 |publisher=Springer-Verlag Berlin Heidelberg |pages=1019–1027 |editor1-first=W. |editor1-last=Henke |editor2-first=I. |editor2-last=Tattersall |doi=10.1007/978-3-642-39979-4_70 |isbn=978-3-642-39978-7}}</ref><ref>{{cite journal |first=J. C. |last=Fuhlrott |author-link=Johann Carl Fuhlrott |url=https://www.naturhistorischerverein.de/neandertaler_ebook.pdf |title=Menschliche Überreste aus einer Felsengrotte des Düsselthales |trans-title=Human remains from a rock grotto in Düsseltal |journal=Verh Naturhist Ver Preuss Rheinl |year=1859 |volume=16 |pages=131–153 |language=de}}</ref>

The binomial name ''Homo neanderthalensis'' was first proposed by William King in a paper read to the 33rd [[British Science Association]] in 1863, formally recognising it as distinct from modern humans.<ref name="King642">{{Cite journal |last1=King |first1=W. |author-link=William King (geologist) |year=1864 |title=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 |url=https://biodiversitylibrary.org/page/29371003 |journal=Report of the British Association for the Advancement of Science, Notices and Abstracts, Newcastle-upon-Tyne, 1863 |volume=33 |pages=81–82 |via=Biodiversity Heritage Library}}</ref><ref name="MurrNashSeoiMcCo152">{{cite journal |last1=Murray |first1=J. |last2=Nasheuer |first2=H. P. |last3=Seoighe |first3=C. |last4=McCormack |first4=G. P. |last5=Williams |first5=D. M. |last6=Harper |first6=D. A. T. |year=2015 |title=The contribution of William King to the early development of palaeoanthropology |url=https://durham-repository.worktribe.com/output/1406049 |journal=Irish Journal of Earth Sciences |volume=33 |pages=1–16 |doi=10.3318/ijes.2015.33.1 |jstor=10.3318/ijes.2015.33.1}}</ref><ref>{{cite journal |last=Winner |first=A. K. |year=1964 |title=Terminology |journal=Current Anthropology |volume=5 |issue=2 |pages=119–122 |doi=10.1086/200469 |jstor=2739959}}</ref> However, in 1864 he recommended that Neanderthals and modern humans be classified in different genera as he compared the Neanderthal braincase to that of a chimpanzee and argued that they were "incapable of moral and [<nowiki/>[[Theism|theistic]]{{efn|King made a typo and wrote "theositic".}}] conceptions".<ref name="king18642">{{cite journal |last=King |first=W. |author-link=William King (geologist) |year=1864 |title=The reputed fossil man of the Neanderthal |url=https://biostor.org/reference/195152 |journal=Quarterly Journal of Science |volume=1 |page=96}}</ref>

The uniqueness of Neanderthal Man met opposition namely from the pathologist [[Rudolf Virchow]], who argued against defining new species based on only a single find. In 1872, Virchow erroneously interpreted Neanderthal characteristics as evidence of [[Dementia|senility]], disease, and malformation instead of archaicness,<ref>{{cite journal |last=Virchow |first=R. |author-link=Rudolf Virchow |trans-title=Examinations on the Neandertal skull |language=de |title=Untersuchung des Neanderthal-Schädels |journal=Verh Berl Anthrop Ges |year=1872 |volume=4 |pages=157–165}}</ref> which stalled Neanderthal research until the end of the century.<ref name="drell2000">{{cite journal |first=J. R. R. |last=Drell |year=2000 |title=Neanderthals: a history of interpretation |journal=Oxford Journal of Archaeology |volume=19 |issue=1 |pages=1–24 |doi=10.1111/1468-0092.00096 }}</ref><ref name="schlager" />

By the early 20th century, numerous other Neanderthal discoveries were made, establishing ''H. neanderthalensis'' as a legitimate species. At first, many palaeontologists considered Neanderthals to be an intermediary phase between modern humans and more apelike ancestors, as suggested by German anatomist [[Gustav Albert Schwalbe]]. This hypothesis was opposed by French palaeontologist [[Marcellin Boule]], who authored several publications starting in 1908 describing the French Neanderthal specimen [[La Chapelle-aux-Saints 1]] ("The Old Man") as a slouching, ape-like creature distantly related to modern man. Boule's ideas would define discussions of Neanderthals for some time.<ref name=drell2000/><ref name=boule>{{cite book |url=https://archive.org/details/b22463355 |first=M. |last=Boule |author-link=Marcellin Boule |year=1911 |title=L'homme fossile de La Chapelle-aux-Saints |trans-title=Fossil man from La Chapelle-aux-Saints |language=fr |publisher=Masson |pages=[https://archive.org/details/b22463355/page/n16 1]–62}}</ref><ref name=Reybrouck2002>{{cite journal |first=D. |last=Van Reybrouck |author-link=David Van Reybrouck |year=2002 |title=Boule's error: on the social context of scientific knowledge |journal=Antiquity |volume=76 |issue=291 |pages=158–164 |doi=10.1017/S0003598X00089936 }}</ref><ref name=Langdon2016>{{cite book |first=J. H. |last=Langdon |year=2016 |chapter=Case study 18. Neanderthals in the mirror: imagining our relatives |title=The science of human evolution: getting it right |publisher=Springer |isbn=978-3-319-41584-0}}</ref><ref name=sommer2006>{{cite journal |first=M. |last=Sommer |year=2006 |title=Mirror, mirror on the wall: Neanderthal as image and 'distortion' in early 20th-century French science and press |journal=Social Studies of Science |volume=36 |issue=2 |pages=207–240 |doi=10.1177/0306312706054527 }}</ref>

Boule suggested two different lineages existed in Ice Age Europe: a more evolved one descending from the British [[Piltdown Man]] (a hoax) to the French [[Grimaldi Man]] (a [[Cro-Magnon]]) which would culminate with modern Europeans; and a less evolved dead-end lineage leading from the German [[Mauer 1|Heidelberg Man]] to Neanderthal Man. As the focus of human origins shifted from Europe to East Asia ("Out of Asia" hypothesis) by the 1930s and 40s with discoveries such as [[Java Man]] and [[Peking Man]] (as well as the marginalisation of Piltdown Man), the question of a "Neanderthal phase" in human evolution once again became a topic of discussion. The definition of "Neanderthal" expanded to include several anatomically variable specimens around the [[Old World]]. Some specimens were described as "progressive" Neanderthals which would evolve into some local subspecies of ''H. sapiens'' ([[multiregional origin of modern humans|polycentricism]]), while the "classic" Neanderthals of the Western European [[Würm glaciation]] would not.<ref>{{cite journal|first1=F.|last1=Spencer|first2=F. H.|last2=Smith|year=1981|title=The significance of Aleš Hrdlička's 'Neanderthal phase of man': A historical and current assessment|journal=American Journal of Physical Anthropology|volume=56|issue=4|pages=435–459|doi=10.1002/ajpa.1330560417}}</ref>

In the 1970s, with the formulation of [[cladistics]] and the consequent refinement of the anatomical definitions of species, this "global morphological pattern" fell apart. The "Neanderthaloids" of Africa and East Asia were reclassified as distant relatives to ''H. neanderthalensis''.<ref name=Hublin2002/> At around the same time, the "Out of Asia" hypothesis was overturned by the [[recent African origin of modern humans|"Out of Africa" hypothesis]], which posited that all modern humans share a fully modern common ancestor ([[monogenism]]). There were two main schools of thought: modern humans competitively replaced all other archaic humans ("Replacement"), or extensively interbred with them while dispersing throughout the world ("Regional Continuity").<ref name=Holliday2014>{{cite journal|last1=Holliday|first1=T. W.|last2=Gautney|first2=J. R.|last3=Friedl|first3=L.|year=2014|title=Right for the Wrong Reasons|journal=Current Anthropology|volume=55|issue=6|pages=696–724|doi=10.1086/679068}}</ref> In 2010, the first mapping of the Neanderthal [[genome]] demonstrated that there was at least some [[interbreeding between archaic and modern humans]].<ref name=green>{{cite journal |last1=Green |first1=R. E. |last2=Krause |first2=J. |last3=Briggs |first3=A. W. |display-authors=3 |last4=Maricic |first4=T. |last5=Stenzel |first5=U. |last6=Kircher |first6=M. |last7=Patterson |first7=N. |last8=Li |first8=H. |last9=Zhai |first9=W. |last10=Fritz |first10=M. H. Y. |last11=Hansen |first11=N. F. |last12=Durand |first12=E. Y. |last13=Malaspinas |first13=A. S. |last14=Jensen |first14=J. D. |last15=Marques-Bonet |first15=T. |last16=Alkan |first16=C. |last17=Prüfer |first17=K. |last18=Meyer |first18=M. |last19=Burbano |first19=H. A. |last20=Good |first20=J. M. |last21=Schultz |first21=R. |last22=Aximu-Petri |first22=A. |last23=Butthof |first23=A. |last24=Hober |first24=B. |last25=Hoffner |first25=B. |last26=Siegemund |first26=M. |last27=Weihmann |first27=A. |last28=Nusbaum |first28=C. |last29=Lander |first29=E. S. |last30=Russ |first30=C. |last31=Novod |first31=N. |last32=Affourtit |first32=J. |last33=Egholm |first33=M. |last34=Verna |first34=C. |last35=Rudan |first35=P. |last36=Brajkovic |first36=D. |last37=Kucan |first37=Z. |last38=Gusic |first38=I. |last39=Doronichev |first39=V. B. |last40=Golovanova |first40=L. V. |last41=Lalueza-Fox |first41=C. |last42=de la Rasilla |first42=M. |last43=Fortea |first43=J. |last44=Rosas |first44=A. |last45=Schmitz |first45=R. W. |last46=Johnson |first46=P. L. F. |last47=Eichler |first47=E. E. |last48=Falush |first48=D. |last49=Birney |first49=E. |last50=Mullikin |first50=J. C. |last51=Slatkin |first51=M. |last52=Nielsen |first52=R. |last53=Kelso |first53=J. |last54=Lachmann |first54=M. |last55=Reich |first55=D. |last56=Pääbo |first56=S. |author-link56=Svante Pääbo |title=A draft sequence of the Neandertal genome |journal=Science |year= 2010 |volume=328 |issue=5979 |pages=710–722 |doi=10.1126/science.1188021 |pmid=20448178 |pmc=5100745 |bibcode=2010Sci...328..710G}}</ref> Subsequent genetic studies continue to raise questions on how Neanderthals should be classified relative to modern humans.<ref name=Meneganzin2024/>

===Classification=== Neanderthals can be classified as a unique species as ''H. neanderthalensis'', though some authors argue expanding the definition of ''H. sapiens'' to include other ancient humans, with combinations such as ''H. sapiens neanderthalensis'' ([[lumpers and splitters|splitters and lumpers]]). The latter opinion has generally been justified using [[Neanderthal genetics]], as well as inferences on the complexity of [[Neanderthal behaviour]] based on the archaeological record. While there seems to have been some genetic contact between these two groups, there are potential indicators of [[hybrid incompatibility]],{{efn|The [[X-chromosome]] carries far less archaic DNA than any [[autosome]], which has either been explained as [[hybrid incompatibility]] (the [[large-X effect]] — [[background selection]]) or male sex bias (hybrids were normally the children of a male Neanderthal and female modern human).<ref name="Chevy Huerta-Sánchez Ramachandran 2023 p. e1010399"/>}} which if true could justify species distinction. The crux of the issue lies in the vagueness of the term "species" (the [[species problem]]).<ref name=Meneganzin2024>{{cite journal|first1=A.|last1=Meneganzin|first2=C.|last2=Stringer|author2-link=Chris Stringer|year=2024|title=''Homo sapiens'', Neanderthals and speciation complexity in palaeoanthropology|journal=Evolutionary Journal of the Linnean Society|volume=3|issue=1|article-number=kzae033 |doi=10.1093/evolinnean/kzae033|doi-access=free}}</ref><ref name="Pääbo2014">{{cite book |title=Neanderthal man: in search of lost genomes |author-last=Pääbo |author-first=S. |publisher=Basic Books |year=2014 |location=New York |page=237 |author-link=Svante Pääbo |title-link=Neanderthal Man: In Search of Lost Genomes}}</ref><ref>{{cite journal |author-first=M. |author-last=Hofreiter |year=2011 |title=Drafting human ancestry: What does the Neanderthal genome tell us about hominid evolution? Commentary on Green et al. (2010) |journal=Human Biology |volume=83 |issue=1 |pages=1–11 |doi=10.3378/027.083.0101 |pmid=21453001 }}</ref>

Among identified archaic humans, Neanderthals are most closely related to [[Denisovan]]s based on [[nuclear DNA]] (nDNA) analyses. Denisovans are an enigmatic group of [[Late Pleistocene]] humans only recognisable by a genetic signature rather than anatomical landmarks.<ref name="Petr2020">{{Cite journal |last1=Petr |first1=Martin |last2=Hajdinjak |first2=Mateja |last3=Fu |first3=Qiaomei |last4=Essel |first4=Elena |last5=Rougier |first5=Hélène |last6=Crevecoeur |first6=Isabelle |last7=Semal |first7=Patrick |last8=Golovanova |first8=Liubov V. |last9=Doronichev |first9=Vladimir B. |last10=Lalueza-Fox |first10=Carles |last11=de la Rasilla |first11=Marco |last12=Rosas |first12=Antonio |last13=Shunkov |first13=Michael V. |last14=Kozlikin |first14=Maxim B. |last15=Derevianko |first15=Anatoli P. |year= 2020 |title=The evolutionary history of Neanderthal and Denisovan Y chromosomes |journal=Science |volume=369 |issue=6511 |pages=1653–1656 |doi=10.1126/science.abb6460 |pmid=32973032 |bibcode=2020Sci...369.1653P |hdl=21.11116/0000-0007-11C2-A |hdl-access=free }}</ref> Likely due to more recent interbreeding episodes, the [[mitochondrial DNA]] (mtDNA, passed down maternally)<ref name="Posth2017">{{cite journal |first1=C. |last1=Posth |first2=C. |last2=Wißing |first3=K. |last3=Kitagawa |display-authors=et al. |title=Deeply divergent archaic mitochondrial genome provides lower time boundary for African gene flow into Neanderthals |journal=Nature Communications |volume=8 |article-number=16046 |year=2017 |doi=10.1038/ncomms16046| pmid=28675384 | pmc=5500885 | bibcode=2017NatCo...816046P}}</ref> and [[Y-chromosome]] DNA (passed down paternally)<ref name="Petr2020"/> are more similar between Neanderthals and modern humans than between Neanderthals and Denisovans. Similarly, 430,000 year old fossils from the [[Sima de los Huesos]] are more closely related to Neanderthals in their nDNA, but their mtDNA aligns more closely with Denisovans.<ref name=meyer2016>{{cite journal |first1=M. |last1=Meyer |first2=J. |last2=Arsuaga |first3=C.|last3=de Filippo |first4=S. |last4=Nagel |title=Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins |journal=Nature |volume=531 |issue=7595 |pages=504–507 |year=2016 |doi=10.1038/nature17405 |pmid=26976447 |bibcode=2016Natur.531..504M |hdl=11858/00-001M-0000-002A-0B82-0 }}</ref>

A 2021 phylogeny of some [[Middle Pleistocene]] and Neanderthal fossils using [[tip dating]]:<ref name="Ni2021">{{Cite journal |last1=Ni |first1=Xijun |last2=Ji |first2=Qiang |last3=Wu |first3=Wensheng |last4=Shao |first4=Qingfeng |last5=Ji |first5=Yannan |last6=Zhang |first6=Chi |last7=Liang |first7=Lei |last8=Ge |first8=Junyi |last9=Guo |first9=Zhen |last10=Li |first10=Jinhua |last11=Li |first11=Qiang |year=2021 |title=Massive cranium from Harbin in northeastern China establishes a new Middle Pleistocene human lineage |journal=The Innovation |language=en |volume=2 |issue=3 |doi=10.1016/j.xinn.2021.100130 |pmc=8454562 |pmid=34557770 |last13=Stringer |first13=Chris |last12=Grün |first12=Rainer |article-number=100130 |bibcode=2021Innov...200130N}}</ref>

{{Clade|label1=''[[Homo]]''|{{Clade |1=''[[Homo habilis|H. habilis]]'' |2={{clade |1=''[[Homo erectus|H. erectus]]'' |2={{clade |1=Tighennif |2={{clade |1={{clade |1=[[Saldanha man|Saldanha]] |2={{clade |1={{clade |1=[[Petralona skull|Petralona]] |2=[[Kabwe 1]] }} |2={{clade |1=[[Ceprano Man|Ceprano]] |2=[[Bodo cranium|Bodo]] }} }} }} |2={{clade |1={{clade |1=[[Mauer 1]] |2=[[Tautavel Man|Arago]] }} |2={{clade |1={{clade |1=[[Narmada Human|Narmada]] |2=[[Maba Man|Maba]] }} |2={{clade |1=[[Ndutu cranium|Ndutu]] |2={{clade |label1='''Neanderthal clade''' |1={{clade |1=[[Sima de los Huesos]] |2={{clade |1=[[Saccopastore skulls|Saccopastore]] |2={{clade |1={{clade |1={{clade |1=[[Tabun Cave|Tabun]] |2=[[Gibraltar 1]] }} |2={{clade |1=[[Spy Cave|Spy]] |2=[[Neanderthal 1]] }} }} |2={{clade |1={{clade |1=[[Amud 1]] |2=Saint-Césaire }} |2={{clade |1=[[Shanidar Cave|Shanidar]] |2={{clade |1=[[La Chapelle-aux-Saints 1]] |2=[[La Ferrassie 1]] }} }} }} }} }} }} |2=''[[Human|H. sapiens]]'' }} }} }} }} }} }} }} }} }}

==Evolution== [[File:Skull5 complete right Sima de los Huesos.jpg|thumb|left|A "pre-Neanderthal" skull ("[[Miguelón]]") from [[Sima de los Huesos hominins|Sima de los Huesos]], Spain]] Typical Neanderthal skull traits appear in the European fossil record near the beginning of the [[Chibanian|Middle Pleistocene]], in specimens usually classified as ''[[Homo heidelbergensis|H. heidelbergensis]]''. These "pre-Neanderthals" seem to have gradually accreted these traits ("Neanderthalization") as populations adapted to the cold environment, evolving a "hyper-arctic" physique. [[Circumpolar peoples]] (namely [[Inuit]] groups) are often used as modern Neanderthal analogues to study "hyper-arctic" adaptations. Additionally, [[glacial period]]s may have forced populations into small [[refugium (population biology)|refugia]], reducing [[genetic diversity]], leading to the development of other typical Neanderthal traits through [[genetic drift]] or [[pleiotropy]].<ref name=Hublin2002/> The 120,000 to 140,000-year-old Israeli [[Nesher Ramla Homo|Nesher Ramla]] remains may represent one such source population which would recolonise Europe following the [[Penultimate Glacial Period]].<ref>{{cite journal |first1=I. |last1=Hershkovitz |first2=H. |last2=May |first3=R. |last3=Sarig |display-authors=et al. |year=2021 |title=A Middle Pleistocene ''Homo'' from Nesher Ramla, Israel |journal=Science |volume=372 |issue=6549 |pages=1424–1428 |doi=10.1126/science.abh3169 |bibcode=2021Sci...372.1424H |s2cid=235628111}}</ref>

The occurrence of typical Neanderthal traits in the Middle Pleistocene was highly variable even among individuals of the same population.<ref name=Hublin2002/> The speed of Neanderthalization may have also been impeded by gene flow between Western Europe and Africa; this is exemplified by anomalous specimens which lack typical Neanderthal traits, such as [[Ceprano Man]].<ref name=meyer2016/> The first recognisable "early Neanderthals" show up in the fossil record by the end of [[Marine Isotope Stage]] 7 (beginning roughly 243,000 years ago) and give way to "classic" or "late Neanderthals" by the end of [[Marine Isotope Stage 5e]]. This spans the [[Penultimate Glacial Period]] to the [[Last Interglacial]]. Some early Neanderthal teeth from [[Payré]], France, potentially date to MIS 8, but the dating is uncertain.<ref name=Richards2021>{{cite journal|first1=Gary D.|last1=Richards|first2=Gaspard|last2=Guipert|first3=Rebecca S.|last3=Jabbour|first4=Alban R.|last4=Defleur|title=Neanderthal cranial remains from Baume Moula-Guercy (Soyons, Ardèche, France)|journal=American Journal of Physical Anthropology|year=2021 |issn=1096-8644|pages=201–226|volume=175|issue=1|doi=10.1002/ajpa.24256 |pmid=33644865 |bibcode=2021AJPA..175..201R }}</ref><ref name=Hublin2002/>

Genetic data usually estimates that Neanderthals diverged from modern humans sometime during the early Middle Pleistocene. Neanderthals and Denisovans are more closely related to each other than they are to modern humans, meaning the Neanderthal/Denisovan split occurred sometime later.<ref name=meyer2016/><ref name="Prufer2014">{{cite journal |first=K. |last=Prüfer |display-authors=etal |year=2014 |title=The complete genome sequence of a Neanderthal from the Altai Mountains |journal=Nature |volume=505 |issue=7481 |pages=43–49|doi=10.1038/nature12886 |pmid=24352235 |pmc=4031459 |bibcode=2014Natur.505...43P}}</ref><ref>{{cite journal |first1=S. |last1=Sawyer |first2=G. |last2=Renaud |first3=B. |last3=Viola |first4=J. J. |last4=Hublin |year=2015 |title=Nuclear and mitochondrial DNA sequences from two Denisovan individuals |journal=Proceedings of the National Academy of Sciences |volume=112 |issue=51 |pages=15696–15700 |doi=10.1073/pnas.1519905112 |pmc=4697428 |pmid=26630009 |bibcode=2015PNAS..11215696S|doi-access=free }}</ref> Before splitting, Neanderthal/Denisovans (or "Neandersovans") migrating out of Africa into Europe apparently interbred with an unidentified "superarchaic" human species who were already present there; these superarchaics were the descendants of a very early migration out of Africa around 1.9 million years ago.<ref name=Rogers2020>{{cite journal |first1=A. R. |last1=Rogers |first2=N. S. |last2=Harris |first3=A. A. |last3=Achenbach |year=2020 |title=Neanderthal-Denisovan ancestors interbred with a distantly related hominin |journal=Science Advances |volume=6 |issue=8 |article-number=eaay5483 |doi=10.1126/sciadv.aay5483 |pmid=32128408 |pmc=7032934 |bibcode=2020SciA....6.5483R}}</ref>

Genetic data indicates that Neanderthals, at least after 100,000 years ago, maintained a small population with low genetic diversity, weakening [[natural selection]] and proliferating [[harmful mutation]]s. It is unclear how long European populations suffered this population stress, or to what extent it influenced Neanderthalization.<ref name=SanchezQuinto2015/>

==Demographics== {{Further|Neanderthals in Southwest Asia|Neanderthals in Gibraltar|List of Neanderthal sites}}

===Range=== [[File:Tabun 1 NMNH.jpg|alt=A skull missing most of the left side of the face from the mid-orbit to the teeth|left|thumb|Neanderthal skull from [[Tabun Cave]], Israel, at the [[Israel Museum]]]] The Neanderthals were the first human species to permanently occupy Europe.{{sfn|French|2021|p=133}} While pre-Neanderthals are mostly identified around Western Europe, classic Neanderthals are recorded across Europe as well as [[Neanderthals in Southwest Asia|Southwest]]<ref name=Hublin2002/> and Central Asia, up to the [[Altai Mountains]] in southern Siberia. Pre- and early Neanderthals seem to have continuously occupied only France, Spain, and Italy, although some appear to have moved out of this "core-area" to form temporary settlements eastward (without leaving Europe). Nonetheless, southwestern France has the highest density of sites for pre- and classic Neanderthals.<ref name=Serangeli>{{cite journal |first1=J. |last1=Serangeli |first2=M. |last2=Bolus |url=https://quartaer.obermaier-gesellschaft.de/pdfs/2008/2008_serangeli.pdf |year=2008 |title=Out of Europe - The dispersal of a successful European hominin form |journal=Quartär |volume=55 |pages=83–98 }}</ref>

The southernmost find was recorded at [[Shuqba Cave]], Levant;<ref>{{cite journal |first=J. |last=Callander |year=2004 |title=Dorothy Garrod's excavations in the Late Mousterian of Shukbah Cave in Palestine reconsidered |journal=Proceedings of the Prehistoric Society |volume=70 |pages=207–231 |doi=10.1017/S0079497X00001171 }}</ref> reports of Neanderthals from the North African [[Jebel Irhoud]]<ref>{{cite journal |last1=Smith |first1=T. M. |last2=Tafforeau |first2=P. |last3=Reid |first3=D. J. |display-authors=et al. |title=Earliest evidence of modern human life history in North African early ''Homo sapiens'' |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=104 |issue=15 |pages=6128–6133 |year=2007 |pmid=17372199 |pmc=1828706 |doi=10.1073/pnas.0700747104 |bibcode=2007PNAS..104.6128S|doi-access=free }}</ref> and [[Haua Fteah]]<ref>{{Cite journal |title=The chronostratigraphy of the Haua Fteah cave (Cyrenaica, northeast Libya) |first1=K. |last1=Douka |first2=Zenobia |last2=J. |first3=C. |last3=Lane |display-authors=et al. |year=2014 |journal=Journal of Human Evolution |doi=10.1016/j.jhevol.2013.10.001 |pmid = 24331954 |volume=66 |pages=39–63 |doi-access=free|bibcode=2014JHumE..66...39D }}</ref> have been reidentified as ''H. sapiens''. Their easternmost presence is recorded at [[Denisova Cave]], Siberia [[85th meridian east|85°E]]; the southeast Chinese [[Maba Man]], a skull, shares several physical attributes with Neanderthals, although these may be the result of [[convergent evolution]] rather than Neanderthals extending their range to the Pacific Ocean.<ref>{{cite journal |first1=X.-J. |last1=Wu |first2=E. |last2=Bruner |year=2016 |title=The endocranial anatomy of Maba 1 |journal=American Journal of Physical Anthropology |volume=160 |issue=4 |pages=633–643 |doi=10.1002/ajpa.22974 |pmid=26972814 |bibcode=2016AJPA..160..633W }}</ref> The northernmost bound is generally accepted to have been [[55th parallel north|55°N]], with unambiguous sites known between [[50th parallel north|50]]–[[53rd parallel north|53°N]], but this is difficult to assess because glacial advances destroy most human remains.<ref>{{cite journal |first1=T. K. |last1=Nielsen |first2=B. M. |last2=Benito |display-authors=et al. |year=2017 |title=Investigating Neanderthal dispersal above 55°N in Europe during the Last Interglacial Complex |journal=Quaternary International |volume=431 |pages=88–103 |doi=10.1016/j.quaint.2015.10.039 |bibcode=2017QuInt.431...88N}}</ref><ref>{{cite journal |first1=T. K. |last1=Nielsen |first2=F. |last2=Riede |year=2018 |title=On research history and Neanderthal occupation at its northern margins |journal=European Journal of Archaeology |volume=21 |issue=4 |pages=506–527 |doi=10.1017/eaa.2018.12 }}</ref> Middle Palaeolithic artefacts have been found up to 60°N on the Russian plains,<ref>{{Cite journal |last1=Pavlov |first1=P. |last2=Roebroeks |first2=W. |last3=Svendsen |first3=J. I. |title=The Pleistocene colonization of northeastern Europe: a report on recent research |journal=Journal of Human Evolution |volume=47 |issue=1–2 |pages=3–17 |year=2004 |pmid=15288521 |doi=10.1016/j.jhevol.2004.05.002 |bibcode=2004JHumE..47....3P }}</ref><ref name=slimak2011>{{cite journal |first1=L. |last1=Slimak |first2=J. I. |last2=Svendsen |first3=J. |last3=Mangerud |first4=H. |last4=Plisson |year=2011 |title=Late Mousterian persistence near the Arctic Circle |journal=Science |volume=332 |issue=6031 |pages=841–845 |doi=10.1126/science.1203866 |pmid=21566192 |jstor=29784275 |bibcode=2011Sci...332..841S }}</ref><ref name=slimak2012>{{cite journal |last=Slimak |first=L. |year=2012 |title=Response to "Comment on Late Mousterian persistence near the Arctic Circle" |journal=Science |volume=335 |issue=6065 |page=167 |doi=10.1126/science.1210211 |bibcode=2012Sci...335..167S |doi-access=free |pmid=22246757 }}</ref> but these are more likely attributed to modern humans.<ref name=zwyns2012>{{cite journal |last=Zwyns |first=N. |year=2012 |title=Comment on Late Mousterian persistence near the Arctic Circle |journal=Science |volume=335 |issue=6065 |page=167 |doi=10.1126/science.1209908 |pmid=22246757 |bibcode=2012Sci...335..167Z |doi-access=free}}</ref>

[[File:Weichsel-Würm-Glaciation.png|thumb|upright=1.35|Map of Europe 20,000 to 70,000 years ago during the [[Würm glaciation]]]] It is possible Neanderthal range expanded and contracted as the ice retreated and grew, respectively, to avoid [[permafrost]] areas, residing in certain refuge zones during glacial maxima.<ref name=bocquet2013/> Stable environments with mild mean annual temperatures may have been the most suitable Neanderthal habitats.<ref>{{Cite journal |last1=Pederzani |first1=Sarah |last2=Britton |first2=Kate |last3=Jones |first3=Jennifer Rose |last4=Agudo Pérez |first4=Lucía |last5=Geiling |first5=Jeanne Marie |last6=Marín-Arroyo |first6=Ana B. |date=July 17, 2023 |title=Late Pleistocene Neanderthal exploitation of stable and mosaic ecosystems in northern Iberia shown by multi-isotope evidence |journal=[[Quaternary Research]] |language=en |volume=116 |pages=108–132 |doi=10.1017/qua.2023.32 |bibcode=2023QuRes.116..108P |hdl=2164/21236 |hdl-access=free }}</ref>

===Population=== Like modern humans, Neanderthals probably descended from a very small population with an [[effective population]]—the number of individuals who can bear or father children—of 3,000 to 12,000 approximately. Neanderthals maintained this low population, proliferating weakly harmful genes due to the reduced effectivity of natural selection.<ref name=juric>{{cite journal |first1=I. |last1=Juric |first2=S. |last2=Aeschbacher |first3=G. |last3=Coop |year=2016 |title=The strength of selection against Neanderthal introgression |journal=PLOS Genetics |volume=12 |issue=11 |article-number=e1006340 |doi=10.1371/journal.pgen.1006340 |pmid=27824859 |pmc=5100956 |doi-access=free }}</ref><ref name=mafessoni2017>{{cite journal |first1=F. |last1=Mafessoni |first2=K. |last2=Prüfer |title=Better support for a small effective population size of Neandertals and a long shared history of Neandertals and Denisovans |journal=Proceedings of the National Academy of Sciences |volume=114 |issue=48 |year=2017 |pages=10256–10257 |doi=10.1073/pnas.1716918114 |pmid=29138326 |pmc=5715791|bibcode=2017PNAS..11410256M |doi-access=free }}</ref> Archaeological evidence suggests that the initial Cro-Magnon population was 10 times higher than Neanderthals.<ref>{{cite journal |first1=P. |last1=Mellars |first2=J. C. |last2=French |year=2011 |title=Tenfold population increase in Western Europe at the Neandertal-to-modern human transition |journal=Science |volume=333 |issue=6042 |pages=623–627 |doi=10.1126/science.1206930 |pmid=21798948 |bibcode=2011Sci...333..623M }}</ref>

Compared to Cro-Magnons, Neanderthals may have been at a demographic disadvantage due to a lower fertility rate, a higher infant mortality rate, or a combination of the two.<ref name=Trinkaus2011>{{cite journal |first=E. |last=Trinkaus |author-link=Erik Trinkaus |year=2011 |title=Late Pleistocene adult mortality patterns and modern human establishment |journal=Proceedings of the National Academy of Sciences |volume=108 |issue=4 |pages=1267–1271 |doi=10.1073/pnas.1018700108 |pmc=3029716 |pmid=21220336 |bibcode=2011PNAS..108.1267T |doi-access=free }}</ref><ref name=bocquet2013>{{cite journal |first1=J. |last1=Bocquet-Appel |first2=A. |last2=Degioanni |year=2013 |title=Neanderthal demographic estimates |journal=Current Anthropology |volume=54 |issue=S8 |pages=202–214 |doi=10.1086/673725 |bibcode=2013CurrA..54S.202B }}</ref> In a sample of 206 Neanderthals, based on the abundance of young and mature adults in comparison to other age demographics, about 80% of them above the age of 20 died before reaching 40. This high mortality rate was probably due to their high-stress environment.<ref name=trinkaus1995>{{cite journal |first=E. |last=Trinkaus |author-link=Erik Trinkaus |year=1995 |title=Neanderthal mortality patterns |journal=Journal of Archaeological Science |volume=22 |issue=1 |pages=121–142 |doi=10.1016/S0305-4403(95)80170-7|bibcode=1995JArSc..22..121T }}</ref> Infant mortality was estimated to have been very high for Neanderthals, about 43% in northern Eurasia.<ref name=pettitt2000>{{cite journal |first=R. B. |last=Pettitt |year=2000 |title=Neanderthal lifecycles: developmental and social phases in the lives of the last archaics |journal=World Archaeology |volume=31 |issue=3 |pages=351–366 |doi=10.1080/00438240009696926 |pmid=16475295 |jstor=125106 }}</ref>

==Anatomy== {{Main|Neanderthal anatomy}}

===Skull=== [[File:Neanderthal cranial anatomy.jpg|thumb|upright=1.5|Neanderthal skull features]] The Neanderthal skull has a flat and broad skullcap, rounded supraorbital torus (the buldge that forms the brow ridges), larger, wide [[orbit (anatomy)|orbits]] (eye sockets), a broad nose, mid-facial [[prognathism]] (the face projects far from the [[base of the skull]]), an "en bombe" (bomb-like) skull shape when viewed from the back, a [[fossa (anatomy)|fossa]] (depression) on the back of the skull below the level of the [[inion]] (suprainiac fossa), and an [[occipital bun]] (bony projection) at the back of the skull. Like those of other archaic humans, their jaws lack a true [[chin]].<ref name=Hublin2002>{{cite book|first=J.-J.|last=Hublin|editor1-last=Akazawa|editor1-first=T.|editor2-last=Aoki|editor2-first=K.|editor3-last=Bar-Yosef|editor3-first=O.|year=2002|title=Neandertals and Modern Humans in Western Asia|chapter=Climatic Changes, Paleogeography, and the Evolution of the Neandertals|doi=10.1007/b109961|isbn=0-306-45924-8}}</ref>

The Neanderthal braincase averages {{cvt|1,640|cm3}} for males and {{cvt|1,460|cm3}} for females,<ref name="holloway1985">{{cite encyclopedia |author-last=Holloway |author-first=R. L. |editor-last=Delson |editor-first=E. |encyclopedia=Ancestors: The hard evidence |title=The poor brain of ''Homo sapiens neanderthalensis'': see what you please |year=1985 |publisher=Alan R. Liss |isbn=0-471-84376-8}}</ref> which is significantly larger than the averages for all living populations.<ref name="SmithDodd1984">{{Cite journal |last1=Beals |first1=K. |last2=Smith |first2=C. |last3=Dodd |first3=S. |year=1984 |title=Brain size, cranial morphology, climate, and time machines |journal=Current Anthropology |volume=12 |issue=3 |pages=301–30 |doi=10.1086/203138 |jstor=2742800 }}</ref> The largest Neanderthal brain, [[Amud 1]], was calculated to be {{cvt|1,736|cm3}}, one of the largest ever recorded in humans.<ref name="amano2015">{{cite journal |last1=Amano |first1=H. |last2=Kikuchi |first2=T. |last3=Morita |first3=Y. |last4=Kondo |first4=O. |last5=Suzuki |first5=H. |last6=Ponce de Leon |first6=M. S. |last7=Zollikofer |first7=C.P.E. |last8=Bastir |first8=M. |last9=Stringer |first9=C. |last10=Ogihara |first10=N. |display-authors=5 |year=2015 |title=Virtual reconstruction of the Neanderthal Amud 1 cranium |journal=American Journal of Physical Anthropology |volume=158 |issue=2 |pages=185–197 |pmid=26249757 |hdl=10261/123419 |doi=10.1002/ajpa.22777 |bibcode=2015AJPA..158..185A }}</ref> Neanderthal brain organisation differs in areas related to cognition and language, which may be implicated in the comparative simplicity of Neanderthal behaviour to Cro-Magnons in the archaeological record.<ref name=Hublinetal2015>{{cite journal |last1=Hublin |first1=Jean-Jacques |last2=Neubauer |first2=Simon |last3=Gunz |first3=Philipp |date=2015 |title=Brain Ontogeny and Life History in Pleistocene Hominins |journal=Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences |volume=370 |issue=1663 |pages=1–11 |doi=10.1098/rstb.2014.0062 |pmid=25602066 |pmc=4305163 }}</ref><ref>{{cite journal |last1=Bastir |first1=Markus |last2=Rosas |first2=Antonio |last3=Lieberman |first3=Daniel E |last4=O'Higgins |first4=Paul |date=2008 |title=Middle Cranial Fossa Anatomy and the Origin of Modern Humans |journal=The Anatomical Record |volume=291 |issue=2 |pages=130–140 |doi=10.1002/ar.20636 |pmid=18213701 |doi-access=free }}</ref><ref>{{cite journal |last1=Gunz |first1=Philipp |last2=Maureille |first2=Bruno |last3=Hublin |first3=Jean-Jacques |date=2010 |title=Brain Development after Birth Differs between Neanderthals and Modern Humans |journal=Current Biology |volume=20 |issue=21 |pages=R921–R922 |doi=10.1016/j.cub.2010.10.018 |pmid=21056830 |s2cid=29295311 |doi-access=free |bibcode=2010CBio...20.R921G }}</ref>

Neanderthals had large and wide noses, probably an adaptation to warm greater quantities of cold air to fuel their assumed heightened metabolism and activity levels.<ref name=Azevedo2017>{{cite journal |last1=de Azevedo |first1=S. |last2=González |first2=M. F. |last3=Cintas |first3=C. |display-authors=3 |last4=Ramallo |first4=V. |last5=Quinto-Sánchez |first5=M. |last6=Márquez |first6=F. |last7=Hünemeier |first7=T. |last8=Paschetta |first8=C. |last9=Ruderman |first9=A. |last10=Navarro |first10=P. |last11=Pazos |first11=B. A. |last12=Silva de Cerqueira |first12=C. C. |last13=Velan |first13=O. |last14=Ramírez-Rozzi |first14=F. |last15=Calvo |first15=N. |last16=Castro |first16=H. G. |last17=Paz |first17=R. R. |last18=González-José |first18=R. |title=Nasal airflow simulations suggest convergent adaptation in Neanderthals and modern humans |journal=Proceedings of the National Academy of Sciences |year=2017 |volume=114 |issue=47 |pages=12442–12447 |doi=10.1073/pnas.1703790114 |pmid=29087302 |pmc=5703271|bibcode=2017PNAS..11412442D |doi-access=free }}</ref> A large nose does not necessarily equate to a better sense of smell, and neurologically, because the [[olfactory bulb]]s are smaller, Neanderthals may have had a poorer sense of smell and [[olfactory memory]] than modern humans.<ref>{{cite journal |last1=Bastir |first1=Markus |last2=Rosas |first2=Antonio |last3=Gunz |first3=Philipp |last4=Peña-Melian |first4=Angel |last5=Manzi |first5=Giorgio |last6=Harvati |first6=Katerina |last7=Kruszynski |first7=Robert |last8=Stringer |first8=Chris |last9=Hublin |first9=Jean-Jacques |date=2011 |title=Evolution of the Base of the Brain in Highly Encephalized Human Species |journal=Nature Communications |volume=2 |issue=2 |page=588 |doi=10.1038/ncomms1593 |pmid=22158443 |bibcode=2011NatCo...2..588B |doi-access=free |hdl=10261/123641 |hdl-access=free }}</ref>

The [[zygomatic bone|cheek bones]] are strong, the [[incisor]]s are large and [[shovel-shaped incisors|shovel-shaped]], the [[molar (tooth)|molars]] have a swollen [[tooth pulp]] ([[taurodontism]]), and there is a gap behind the molars ([[retromolar space]]). These dental traits are usually interpreted as a response to habitual heavy loading of the front teeth, either to process mechanically challenging or [[dental attrition|attritive]] foods, or because Neanderthals regularly used the mouth as a third hand.<ref name=clement2012>{{cite journal |first1=A. F. |last1=Clement |first2=S. W. |last2=Hillson |first3=L. C. |last3=Aiello |year=2012 |title=Tooth wear, Neanderthal facial morphology and the anterior dental loading hypothesis |journal=Journal of Human Evolution |volume=62 |issue=3 |pages=367–376 |doi=10.1016/j.jhevol.2011.11.014 |pmid=22341317 |doi-access=free|bibcode=2012JHumE..62..367C }}</ref>

===Build===

Though Neanderthals are often imagined as short and stocky, they were on average the same height as pre-industrial Europeans, or even slightly taller.<ref name="height">{{cite journal |last1=Helmuth |first1=H. |title=Body height, body mass and surface area of the Neanderthals |journal=Zeitschrift für Morphologie und Anthropologie |date=1998 |volume=82 |issue=1 |pages=1–12 |doi=10.1127/zma/82/1998/1 |pmid=9850627 |issn=0044-314X |quote="...it is surprising that many textbooks portray a wrong picture of Neanderthal height as being "very short" or "just over 5 feet". Based on 45 long bones from maximally 14 males and 7 females, Neanderthals' height averages between 164 and 168 (males) resp. 152 to 156 cm (females). This height is indeed 12-14 cm lower than the height of post-WWII Europeans, but compared to Europeans some 20,000 or 100 years ago, it is practically identical or even slightly higher."}}</ref> The [[body mass index]] of the average European Neanderthal was comparable to 20th century [[Canadians]] and [[Americans]].<ref name="height" /> Neanderthal body size varied by location; [[West Asian]] Neanderthals were taller and more slightly built than European Neanderthals.<ref>{{cite book |last1=Silberman |first1=Neil Asher |title=The Oxford Companion to Archaeology |date=November 2012 |publisher=OUP USA |isbn=978-0-19-973578-5 |page=32-33 |url=https://books.google.com/books?id=xeJMAgAAQBAJ&pg=RA1-PA32 |language=en}}</ref>

In a sample of 45 Neanderthal [[long bone]]s from 14 men and 7 women, the average height was {{cvt|164 to 168|cm|ftin|0}} for males and {{cvt|152 to 156|cm|ftin}} for females.<ref name="height"/> The fossil record shows that adult Neanderthals varied from about {{cvt|147.5|to|177|cm|ftin|0}} in height.<ref name=duveau2019>{{cite journal |first1=J. |last1=Duveau |first2=G. |last2=Berillon |first3=C. |last3=Verna |first4=G. |last4=Laisné |first5=D. |last5=Cliquet |year=2019 |title=The composition of a Neandertal social group revealed by the hominin footprints at Le Rozel (Normandy, France) |journal=Proceedings of the National Academy of Sciences |volume=116 |issue=39 |pages=19409–19414 |doi=10.1073/pnas.1901789116 |pmid=31501334 |pmc=6765299|bibcode=2019PNAS..11619409D |doi-access=free }}</ref> The average male [[body mass index]] was 26.9–28.3.<ref name="height" /><ref name=Froehle2009>{{cite journal |last1=Froehle |first1=Andrew W. |last2=Churchill |first2=Steven |title=Energetic Competition Between Neandertals and Anatomically Modern Humans |journal=PaleoAnthropology |date=2009 |pages=96–116 |url=https://corescholar.libraries.wright.edu/kinesiology_health/73/ }}</ref>

Neanderthal [[human pelvis|pelvic bones]] were extremely wide; with Neanderthal male hips being approximately 31% wider than those of modern humans males.<ref>{{cite book |last1=Kimbel |first1=William H. |last2=Martin |first2=Lawrence B. |title=Species, Species Concepts and Primate Evolution |date=18 December 2013 |page=525 |publisher=Springer Science & Business Media |isbn=978-1-4899-3745-2 |url=https://books.google.com/books?id=Knb1BwAAQBAJ&pg=PA525 |language=en}}</ref> As in all archaic species, there is no evidence of [[sexual dimorphism]] in the Neanderthal pelvis; both males and females had large pelvic bones relative to their body size, with larger males having the largest pelvises.<ref>{{cite book |title=Mosaic |date=1988 |publisher=The Foundation |page=45 |url=https://books.google.com/books?id=Aa_nQZuMEpcC&pg=PA45 |language=en}}</ref>

The Neanderthal chest was deep and wide, with a proportionally expansive [[thoracic cavity]], and possibly stronger [[lung]] performance. Neanderthals also had relatively more [[skeletal muscle#Force generation|fast-twitch muscle fibres]],<ref name=Bastir2022>{{cite journal|first1=M.|last1=Bastir|first2=J. M. G.|last2=Ruíz|first3=J.|last3=Rueda|first4=G. G.|last4=López|first5=M.|last5=Gómez-Recio|first6=B.|last6=Beyer|first7=A. F.|last7=San Juan|first8=E.|last8=Navarro|year=2022|title=Variation in human 3D trunk shape and its functional implications in hominin evolution|journal=Scientific Reports|volume=12|issue=1 |article-number=11762|doi=10.1038/s41598-022-15344-x|doi-access=free|pmc=9273616|pmid=35817835|bibcode=2022NatSR..1211762B }}</ref> and much higher [[calorie|caloric]] demands.<ref name=Froehle2009/> The limbs are proportionally short. The body plan has traditionally been explained as a "hyper-arctic" adaptation ([[Allen's rule]]).<ref>{{cite journal |first=T. W. |last=Holliday |year=1997 |title=Postcranial evidence of cold adaptation in European Neandertals |journal=American Journal of Physical Anthropology |volume=104 |issue=2 |pages=245–258 |doi=10.1002/(SICI)1096-8644(199710)104:2<245::AID-AJPA10>3.0.CO;2-# |pmid=9386830}}</ref><ref name=etrinkaus1981>{{cite book |first=E. |last=Trinkaus |author-link=Erik Trinkaus |year=1981 |title=Aspects of human evolution |chapter=Neanderthal limb proportions and cold adaptation |publisher=Taylor and Francis Ltd. |editor-first=C. B. |editor-last=Stringer}}</ref><ref name=weaver2009>{{cite journal |first=T. D. |last=Weaver |year=2009 |title=The meaning of Neandertal skeletal morphology |journal=Proceedings of the National Academy of Sciences |volume=106 |issue=38 |pages=16,028–16,033 |doi=10.1073/pnas.0903864106 |pmid=19805258 |pmc=2752516|doi-access=free |bibcode=2009PNAS..10616028W }}</ref> Neanderthals would also have been effective sprinters: stronger lungs, more fast-twitch muscle, and shorter limbs would have boosted efficiency.<ref name=Bastir2022/><ref name=stewart2019>{{cite journal |last1=Stewart |first1=J.R. |last2=García-Rodríguez |first2=O. |last3=Knul |first3=M.V. |last4=Sewell |first4=L. |last5=Montgomery |first5=H. |last6=Thomas |first6=M.G. |last7=Diekmann |first7=Y. |title=Palaeoecological and genetic evidence for Neanderthal power locomotion as an adaptation to a woodland environment |journal=Quaternary Science Reviews |year= 2019 |volume=217 |pages=310–315 |doi=10.1016/j.quascirev.2018.12.023 |bibcode=2019QSRv..217..310S |url=http://eprints.bournemouth.ac.uk/31956/21/Neanderthal%20locomotion%20%20-%20Final%20-%20Copy.pdf }}</ref>

Neanderthal pigmentation genes in modern non-[[African people|African]] people are associated with both lighter and darker skin tones, as well as lighter and darker hair colors, suggesting Neanderthals themselves may have had variable pigmentation.<ref>{{cite journal |last1=Dannemann |first1=Michael |last2=Kelso |first2=Janet |title=The Contribution of Neanderthals to Phenotypic Variation in Modern Humans |journal=The American Journal of Human Genetics |date=5 October 2017 |volume=101 |issue=4 |pages=578–589 |doi=10.1016/j.ajhg.2017.09.010 |pmid=28985494 |pmc=5630192 |language=English |issn=0002-9297}}</ref><ref name="Lalueza-FoxRompler2007">{{cite journal |last1=Lalueza-Fox |first1=C. |last2=Rompler |first2=H. |last3=Caramelli |first3=D. |display-authors=3 |last4=Staubert |first4=C. |last5=Catalano |first5=G. |last6=Hughes |first6=D. |last7=Rohland |first7=N. |last8=Pilli |first8=E. |last9=Longo |first9=L. |last10=Condemi |first10=S. |last11=de la Rasilla |first11=M. |last12=Fortea |first12=J. |last13=Rosas |first13=A. |last14=Stoneking |first14=M. |last15=Schoneberg |first15=T. |last16=Bertranpetit |first16=J. |last17=Hofreiter |first17=M. |title=A melanocortin 1 receptor allele suggests varying pigmentation among Neanderthals |journal=Science |year=2007 |volume=318 |issue=5855 |pages=1453–1455 |doi=10.1126/science.1147417 |pmid=17962522 |bibcode=2007Sci...318.1453L |s2cid=10087710 }}</ref><ref name=cerqueira2012>{{cite journal |first1=C. C. |last1=Cerqueira |first2=V. R. |last2=Piaxão-Côrtes |first3=F. M. B. |last3=Zambra |first4=T. |last4=Hünemeier |first5=M. |last5=Bortolini |year=2012 |title=Predicting ''Homo'' pigmentation phenotype through genomic data: From neanderthal to James Watson |journal=American Journal of Human Biology |volume=24 |issue=5 |pages=705–709 |doi=10.1002/ajhb.22263 |pmid=22411106 |s2cid=25853632}}</ref> Variants associated with red hair have been found in some Neanderthals, but they do not appear to have been common.<ref name=dannemann>{{cite journal |last1=Dannemann |first1=M. |last2=Kelso |first2=J. |title=The contribution of Neanderthals to phenotypic variation in modern humans |journal=The American Journal of Human Genetics |year=2017 |volume=101 |issue=4 |pages=584–585 |doi=10.1016/j.ajhg.2017.09.010 |pmid=28985494 |pmc=5630192}}</ref> Neanderthal skin and hair pigmentation variants are under positive selection in non-Africans, unlike most other Neanderthal genes, suggesting that the adaptive value of lighter features was beneficial for modern humans in [[Eurasia]], where there is significantly less sunlight as compared to [[Africa]].<ref>{{cite journal |last1=McArthur |first1=Evonne |last2=Rinker |first2=David C. |last3=Capra |first3=John A. |title=Quantifying the contribution of Neanderthal introgression to the heritability of complex traits |journal=Nature Communications |date=22 July 2021 |volume=12 |issue=1 |page=4481 |doi=10.1038/s41467-021-24582-y |pmid=34294692 |pmc=8298587 |bibcode=2021NatCo..12.4481M |language=en |issn=2041-1723}}</ref>

===Pathology=== Neanderthals suffered a high rate of traumatic injury, with an estimated 79–94% of specimens showing evidence of healed major trauma, of which 37–52% were severely injured, and 13–19% injured before reaching adulthood.<ref name=nakahashi>{{cite journal |last1=Nakahashi |first1=W. |title=The effect of trauma on Neanderthal culture: A mathematical analysis |journal=Homo |year= 2017 |volume=68 |issue=2 |pages=83–100 |doi=10.1016/j.jchb.2017.02.001 |pmid=28238406}}</ref> One extreme example is [[Shanidar 1]], who shows signs of an [[amputation]] of the right arm likely due to a [[nonunion]] after breaking a bone in adolescence, [[osteomyelitis]] (a bone infection) on the left [[clavicle]], an abnormal [[gait]], vision problems in the left eye, and possible hearing loss<ref name=Trinkaus2017>{{cite journal |first1=E. |last1=Trinkaus |author-link=Erik Trinkaus |first2=S |last2=Villotte |year=2017 |title=External auditory exostoses and hearing loss in the Shanidar 1 Neandertal |journal=PLOS ONE |volume=12 |issue=10 |article-number=e0186684 |doi=10.1371/journal.pone.0186684 |pmc=5650169 |pmid=29053746 |bibcode=2017PLoSO..1286684T|doi-access=free }}</ref> (perhaps [[swimmer's ear]]).<ref name=Trinkaus2019>{{cite journal |first1=E. |last1=Trinkaus |author-link=Erik Trinkaus |first2=M. |last2=Samsel |first3=S. |last3=Villotte |year=2019 |title=External auditory exostoses among western Eurasian late Middle and Late Pleistocene humans |journal=PLOS ONE |volume=14 |issue=8 |article-number=e0220464 |doi=10.1371/journal.pone.0220464 |pmid=31412053 |pmc=6693685 |bibcode=2019PLoSO..1420464T|doi-access=free }}</ref> The high trauma rate may be ascribed to a dangerous hunting strategy,<ref name=trinkaus1995>{{cite journal |first=E. |last=Trinkaus |author-link=Erik Trinkaus |year=1995 |title=Neanderthal mortality patterns |journal=Journal of Archaeological Science |volume=22 |issue=1 |pages=121–142 |doi=10.1016/S0305-4403(95)80170-7|bibcode=1995JArSc..22..121T }}</ref> or frequent animal attacks.<ref name=Camaros2016>{{cite journal |last1=Camarós |first1=Edgard |last2=Cueto |first2=Marián |last3=Lorenzo |first3=Carlos |last4=Villaverde |first4=Valentín |last5=Rivals |first5=Florent |title=Large carnivore attacks on hominins during the Pleistocene: a forensic approach with a Neanderthal example |journal=Archaeological and Anthropological Sciences |date=September 2016 |volume=8 |issue=3 |pages=635–646 |doi=10.1007/s12520-015-0248-1 |bibcode=2016ArAnS...8..635C |hdl=10550/54275 |hdl-access=free}}</ref>

Low population caused a low genetic diversity and probably inbreeding, which reduced the population's ability to filter out harmful mutations ([[inbreeding depression]]). It is unknown how this affected a single Neanderthal's genetic burden and, thus, if this caused a higher rate of [[birth defect]]s than in modern humans.<ref name=SanchezQuinto2015/>

==Culture== {{Main|Neanderthal behavior}}

===Social structure=== [[File:Neanderthal genetic subgroups.png|thumb|upright=1.35|Genetically, Neanderthals may be grouped into three distinct regions (above), dots indicate sampled specimens.<ref name=Fabre2009/>]] It is difficult to infer Neanderthal group size, but indirect data generally suggests small bands of 10 to 30 individuals.<ref name=hayden2012>{{cite journal |first=B. |last=Hayden |year=2012 |title=Neandertal social structure? |journal=Oxford Journal of Archaeology |volume=31 |issue=1 |pages=1–26 |doi=10.1111/j.1468-0092.2011.00376.x}}</ref> Bands likely moved between certain caves depending on the season, indicated by remains of seasonal materials, such as certain foods. They returned to the same locations generation after generation and some sites may have been used for more than a century.<ref name=Farizy1994>{{cite journal |first=C. |last=Farizy |year=1994 |title=Spatial patterning of Middle Paleolithic sites |journal=Journal of Anthropological Archaeology |volume=13 |issue=2 |pages=153–160 |doi=10.1006/jaar.1994.1010}}</ref> Neanderthals may have been outcompeting [[cave bear]]s for cave space.<ref>{{cite journal |first1=M. |last1=Stiller |first2=G. |last2=Baryshnikov |first3=H. |last3=Bocherens |year=2010 |title=Withering away—25,000 years of genetic decline preceded cave bear extinction |journal=Molecular Biology and Evolution |volume=27 |issue=5 |pages=975–978 |doi=10.1093/molbev/msq083 |pmid=20335279 |doi-access=free}}</ref> Intergroup movement may have been predominantly [[patrilocal residence|patrilocal]] (male relationships as the basis of groups with females from other groups entering for breeding).<ref>{{cite journal |first1=C. |last1=Lalueza-Fox |first2=A. |last2=Rosas |first3=A. |last3=Estalrrich |year=2011 |title=Genetic evidence for patrilocal mating behavior among Neandertal groups |journal=Proceedings of the National Academy of Sciences |volume=108 |issue=1 |pages=250–253 |doi=10.1073/pnas.1011553108 |pmid=21173265 |pmc=3017130|doi-access=free }}</ref>

Neanderthals maintained a low population across their range, which may have hindered their ability to maintain long-distance trade routes<ref name=pearce2013>{{cite journal |first1=E. |last1=Pearce |first2=C. |last2=Stringer |first3=R. I. M. |last3=Dunbar |year=2013 |title=New insights into differences in brain organization between Neanderthals and anatomically modern humans |journal=Proceedings of the Royal Society B |volume=280 |issue=1758 |article-number=20130168 |doi=10.1098/rspb.2013.0168 |pmc=3619466 |pmid=23486442}}</ref> and to avoid [[inbreeding]].<ref name=SanchezQuinto2015>{{cite journal |first1=F. |last1=Sánchez-Quinto |first2=C. |last2=Lalueza-Fox |year=2015 |title=Almost 20 years of Neanderthal palaeogenetics: adaptation, admixture, diversity, demography and extinction |journal=Philosophical Transactions of the Royal Society B |volume=370 |issue=1660 |article-number=20130374 |doi=10.1098/rstb.2013.0374 |pmc=4275882 |pmid=25487326}}</ref> They may have regularly interacted with closely neighbouring communities within a region, but not so often beyond.<ref name=Ruebens2013>{{cite journal |first=K. |last=Ruebens |year=2013 |title=Regional behaviour among late Neanderthal groups in Western Europe: A comparative assessment of late Middle Palaeolithic bifacial tool variability |journal=Journal of Human Evolution |volume=65 |issue=4 |pages=341–362 |doi=10.1016/j.jhevol.2013.06.009 |pmid=23928352 |bibcode=2013JHumE..65..341R }}</ref> Genetic analysis indicates there were at least three distinct geographical groups: Western Europe, the Mediterranean coast, and east of the Caucasus, with some migration among these regions.<ref name=Fabre2009>{{cite journal |first1=V. |last1=Fabre |first2=S. |last2=Condemi |first3=A. |last3=Degioanni |year=2009 |title=Genetic evidence of geographical groups among Neanderthals |journal=PLOS ONE |volume=4 |issue=4 |article-number=e5151 |doi=10.1371/journal.pone.0005151 |pmc=2664900 |pmid=19367332 |bibcode=2009PLoSO...4.5151F|doi-access=free }}</ref> [[File:Brno, Neanderthal Mother (detail of diorama).jpg|thumb|Neanderthal mother with child depicted in the [[Anthropos Pavilion]] of the [[Moravian Museum]]]] While the Cro-Magnons are usually assumed to have generally practised [[sexual division of labour]] with men hunting and women gathering such as in the preponderance of recent hunter-gatherer societies, it is unclear to what extent this may be applied to Neanderthals. Both Neanderthal men and women have similar traumatic injury patterns, which might imply that both sexes were involved in hunting. Dental wearing patterns among Neanderthals, on the other hand, could indicate men and women typically carried different items with their mouths, though these may not have been related to subsistence tasks. The women at El Sidrón Cave, Spain, may have been eating more seeds and nuts than the men. The lack of distinctive task specialization in Neanderthals has usually been linked to their small population and group size, falling short of the demographic threshold where task specialization becomes feasible — which may also explain the comparative simplicity of Neanderthal material culture.<ref>{{cite book|first1=Jennifer C.|last1=French|chapter=Sex, gender, and the division of labour in the European Middle and Upper Palaeolithic|title=The Routledge Handbook of Gender Archaeology|publisher=Routledge|year= 2024|location=London|isbn=978-1-003-25753-0|pages=161–174|doi=10.4324/9781003257530-14}}</ref>

===Food=== {{See also|Pleistocene human diet}} [[File:Homo sapiens neanderthalensis-Jäger.jpg|thumb|left|Reconstruction of a Neanderthal man butchering a goat at the [[Neanderthal Museum]]]] Though once thought of as [[scavenger]]s, Neanderthals are now considered [[apex predator]]s.<ref name="PNAS-20190210">{{cite journal |last1=Jaouen |first1=K. |display-authors=etal |title=Exceptionally high δ15N values in collagen single amino acids confirm Neandertals as high-trophic level carnivores |year=2019 |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=116 |issue=11 |pages=4928–4933 |doi=10.1073/pnas.1814087116 |pmid=30782806 |pmc=6421459|bibcode=2019PNAS..116.4928J |doi-access=free }}</ref> They appear to have eaten predominantly what was abundant within their immediate surroundings,{{sfn|Tattersall|2015}} consequently consuming across their range a wide array of meats and plants,<ref>{{Cite journal |last1=Hernaiz-García |first1=María |last2=Zanolli |first2=Clément |last3=Martín-Francés |first3=Laura |last4=Mazurier |first4=Arnaud |last5=Benazzi |first5=Stefano |last6=Sarig |first6=Rachel |last7=Fu |first7=Jing |last8=Kullmer |first8=Ottmar |last9=Fiorenza |first9=Luca |date=29 February 2024 |title=Masticatory habits of the adult Neanderthal individual BD 1 from La Chaise-de-Vouthon (France) |url=https://onlinelibrary.wiley.com/doi/full/10.1002/ajpa.24926 |journal=[[American Journal of Biological Anthropology]] |language=en |volume=184 |issue=1 |article-number=e24926 |doi=10.1002/ajpa.24926 |pmid=38420653 |issn=2692-7691 |access-date=16 March 2026 |via=Wiley Online Library}}</ref> the relative proportion of which varied substantially geographically.<ref>{{Cite journal |last1=Fiorenza|first1=Luca|last2=Benazzi|first2=Stefano|last3=Tausch|first3=Jeremy|last4=Kullmer|first4=Ottmar|last5=Bromage|first5=Timothy G.|last6=Schrenk|first6=Friedemann|date=18 March 2011|editor-last=Rosenberg|editor-first=Karen|title=Molar Macrowear Reveals Neanderthal Eco-Geographic Dietary Variation|journal=[[PLOS One]]|language=en|volume=6|issue=3|article-number=e14769|doi=10.1371/journal.pone.0014769|doi-access=free|issn=1932-6203|pmc=3060801|pmid=21445243 |bibcode=2011PLoSO...614769F }}</ref> Cro-Magnons, in contrast, seem to have maintained a more diverse diet even in settings where certain foods would have been harder to procure; for example, Neanderthals living in forests ate about the same proportion of foodplants as Cro-Magnons, but Neanderthals on open steppe (where foodplants are harder to find) ate far less foodplants.<ref name=dusseldorp2013/><ref name=Zaatari2016>{{cite journal |first1=S. |last1=El Zaatari |first2=Frederick E. |last2=Grine |first3=Peter Stuart |last3=Ungar |first4=J.-J. |last4=Hublin |year=2016 |title=Neandertal versus modern human dietary responses to climatic fluctuations |journal=[[PLOS One]] |volume=11 |issue=4 |article-number=e0153277 |doi=10.1371/journal.pone.0153277 |pmc=4847867 |pmid=27119336 |bibcode=2016PLoSO..1153277E|doi-access=free }}</ref>

In many European sites, prey items include [[red deer]], [[reindeer]], [[wild horse|horse]], [[aurochs]], [[ibex]], and [[steppe bison]]. [[Neanderthals in Southwest Asia]] more commonly hunted [[mountain gazelle]], [[Persian fallow deer]], [[wild goat]], and [[camel]]s.<ref name=dusseldorp2013/> They may have less frequently taken down larger [[Pleistocene megafauna]] whenever locally abundant, such as [[woolly mammoth]] and [[woolly rhinoceros]].<ref>{{cite journal|first1=G. M.|last1=Smith|title=Neanderthal megafaunal exploitation in Western Europe and its dietary implications: A contextual reassessment of La Cotte de St Brelade (Jersey)|journal=Journal of Human Evolution|year= 2015|issn=0047-2484|pages=181–201|volume=78|doi=10.1016/j.jhevol.2014.10.007|pmid=25454779 |bibcode=2015JHumE..78..181S }}</ref> At the 125,000 year old Neumark-Nord site, Germany, there is evidence of regular hunting of [[straight-tusked elephant]]s maybe every 5 to 6 years.<ref>{{Cite journal |last1=Gaudzinski-Windheuser |first1=Sabine |last2=Kindler |first2=Lutz |last3=MacDonald |first3=Katharine |last4=Roebroeks |first4=Wil |date=February 3, 2023 |title=Hunting and processing of straight-tusked elephants 125.000 years ago: Implications for Neanderthal behavior |journal=[[Science Advances]] |language=en |volume=9 |issue=5 |article-number=eadd8186 |doi=10.1126/sciadv.add8186 |issn=2375-2548 |pmc=9891704 |pmid=36724231 |bibcode=2023SciA....9D8186G }}</ref> Some waterside communities ate fish and shellfish—and at [[Vanguard Cave]], Gibraltar—dolphin and [[Mediterranean monk seal]].{{sfn|Brown|2011}} Neanderthals also hunted small game, and some caves show evidence of regular rabbit and tortoise consumption. At Gibraltar sites, there are butchered remains of 143 different bird species, many ground-dwelling such as the [[common quail]], [[corn crake]], [[woodlark]], and [[crested lark]].{{sfn|Brown|2011}} Neanderthals also consumed a variety of plants and mushrooms across their range — at [[Kebara Cave]], Israel, over 50 species of seeds, nuts, fruits, and cereals.<ref>{{Cite journal |date=2018 |title=Dental calculus indicates widespread plant use within the stable Neanderthal dietary niche |journal=Journal of Human Evolution |volume=119 |pages=27–41 |doi=10.1016/j.jhevol.2018.02.009 |issn=0047-2484 |hdl=10550/65536 |hdl-access=free |last1=Power |first1=R. C. |last2=Salazar-García |first2=D. C. |last3=Rubini |first3=M. |last4=Darlas |first4=A. |last5=Harvati |first5=K. |last6=Walker |first6=M. |last7=Hublin |first7=J. |last8=Henry |first8=A. G. |pmid=29685752|bibcode=2018JHumE.119...27P |s2cid=13831823 }}</ref><ref name=shipley2016>{{cite journal |first1=G. P. |last1=Shipley |last2=Kindscher |first2=K. |year=2016 |title=Evidence for the paleoethnobotany of the Neanderthal: a review of the literature |journal=[[Scientifica]] |volume=2016 |pages=1–12 |doi=10.1155/2016/8927654 |pmc=5098096 |pmid=27843675 |doi-access=free}}</ref>

Neanderthals possibly employed a wide range of food preparation techniques. At Cueva del Sidrón, Spain, Neanderthals may have been [[roasting]] and [[smoking (cooking)|smoking]] meat, and used certain plants—such as [[yarrow]] and [[camomile]]—for flavouring,<ref name=krief2015>{{cite journal|first1=S. |last1=Krief |first2=C. |last2=Daujeard |first3=M. |last3=Moncel |first4=N. |last4=Lamon |first5=V. |last5=Reynolds |year=2015 |title=Flavouring food: the contribution of chimpanzee behaviour to the understanding of Neanderthal calculus composition and plant use in Neanderthal diets |journal=[[Antiquity (journal)|Antiquity]] |volume=89 |issue=344 |pages=464–471 |doi=10.15184/aqy.2014.7 }}</ref> although these plants may have instead been used for their medicinal properties.<ref name=buckley>{{cite journal |first1=S. |last1=Buckley |first2=K. |last2=Hardy |first3=M. |last3=Huffman |year=2013 |title=Neanderthal self-medication in context |journal=[[Antiquity (journal)|Antiquity]] |volume=87 |issue=337 |pages=873–878 |doi=10.1017/S0003598X00049528 |doi-access=free }}</ref><ref name=hardy2012>{{cite journal |first1=K. |last1=Hardy |first2=S. |last2=Buckley |first3=M. J. |last3=Collins |first4=A. |last4=Estalrrich |year=2012 |title=Neanderthal medics? Evidence for food, cooking, and medicinal plants entrapped in dental calculus |journal=[[The Science of Nature]] |volume=99 |issue=8 |pages=617–626 |doi=10.1007/s00114-012-0942-0 |pmid=22806252 |bibcode=2012NW.....99..617H |hdl=10651/7043 }}</ref> At [[Gorham's Cave]], Gibraltar, Neanderthals may have been roasting [[pinecone]]s to access [[pine nut]]s,{{sfn|Brown|2011}} and at Gruta da Figueira Brava, [[Cancer pagurus|brown crabs]] to soften the shell before cracking them open.<ref>{{cite journal|first1=M.|last1=Nabais|first2=R.|last2=Portero|first3=J.|last3=Zilhão|year=2024|title=Neanderthal brown crab recipes: A combined approach using experimental, archaeological and ethnographic evidence|journal=[[Historical Biology]]|volume=36|issue=8|pages=1487–1495|doi=10.1080/08912963.2023.2220005|bibcode=2024HBio...36.1487N |hdl=10366/156668|hdl-access=free}}</ref> At [[Grotte du Lazaret]], France, a total of twenty-three red deer, six ibexes, three aurochs, and one [[roe deer]] appear to have been hunted in a single autumn hunting season, when strong male and female deer herds would group together for [[rut (mammalian reproduction)|rut]]. It is possible these Neanderthals were [[curing (food preservation)|curing]] and storing all this meat before winter set in.<ref name=Valensi2013>{{cite journal |first1=P. |last1=Valensi |first2=V. |last2=Michel |display-authors=et al. |year=2013 |title=New data on human behavior from a 160,000 year old Acheulean occupation level at Lazaret cave, south-east France: An archaeozoological approach |journal=Quaternary International |volume=316 |pages=123–139 |doi=10.1016/j.quaint.2013.10.034 |bibcode=2013QuInt.316..123V}}</ref> Neanderthals at Neumark-Nord may have been [[rendering (animal products)|rendering]] fat from animal bones to offset [[protein toxicity]].<ref>{{cite journal | doi=10.1126/sciadv.adv1257 | title=Large-scale processing of within-bone nutrients by Neanderthals, 125,000 years ago | date=2025 | last1=Kindler | first1=Lutz | last2=Gaudzinski-Windheuser | first2=Sabine | last3=Scherjon | first3=Fulco | last4=Garcia-Moreno | first4=Alejandro | last5=Smith | first5=Geoff M. | last6=Pop | first6=Eduard | last7=Speth | first7=John D. | last8=Roebroeks | first8=Wil | journal=[[Science Advances]] | volume=11 | issue=27 | article-number=eadv1257 | pmid=40601744 | pmc=12219469 | bibcode=2025SciA...11.1257K }}</ref> Likewise, calcium measurements show that Neanderthals at Grotte du Bison frequently consumed bone.<ref>{{Cite journal |last1=Dodat |first1=Pierre-Jean |last2=Albalat |first2=Emmanuelle |last3=Balter |first3=Vincent |last4=Couture-Veschambre |first4=Christine |last5=Hardy |first5=Maurice |last6=Henrion |first6=Juliette |last7=Holliday |first7=Trenton |last8=Maureille |first8=Bruno |date=August 2024 |title=Diverse bone-calcium isotope compositions in Neandertals suggest different dietary strategies |journal=[[Journal of Human Evolution]] |volume=193 |article-number=103566 |doi=10.1016/j.jhevol.2024.103566 |pmid=39029412 |bibcode=2024JHumE.19303566D |url=https://hal.science/hal-04659304 }}</ref>

Neanderthals competed with several large carnivores, but also seem to have hunted them down, namely [[Panthera spelaea|cave lions]] and [[wolf|wolves]],{{sfn|Shipman|2015|loc=pp. 120–143}} as well as cave and [[brown bear]] both in and out of [[hibernation]].<ref>{{cite journal |last1=Romandini |first1=M. |last2=Terlato |first2=G. |last3=Nannini |first3=N. |year=2018 |title=Bears and humans, a Neanderthal tale. Reconstructing uncommon behaviors from zooarchaeological evidence in southern Europe |journal=[[Journal of Archaeological Science]] |volume=90 |pages=71–91 |bibcode=2018JArSc..90...71R |doi=10.1016/j.jas.2017.12.004 |s2cid=53410125 |hdl-access=free |hdl=11392/2381729}}</ref> Neanderthals and other predators may have sometimes avoided competition by pursuing different prey, namely with [[cave hyena]]s<ref name=dusseldorp2013>{{cite book |last1=Dusseldorp |first1=Gerrit L. |title=Zooarchaeology and Modern Human Origins |chapter=Neanderthals and Cave Hyenas: Co-existence, Competition or Conflict? |series=Vertebrate Paleobiology and Paleoanthropology |date=2013 |pages=191–208 |doi=10.1007/978-94-007-6766-9_12 |isbn=978-94-007-6765-2 |doi-access=free }}</ref> and wolves ([[niche differentiation]]).<ref>{{Cite journal |last1=Ecker |first1=Michaela |last2=Bocherens |first2=Hervé |last3=Julien |first3=Marie-Anne |last4=Rivals |first4=Florent |last5=Raynal |first5=Jean-Paul |last6=Moncel |first6=Marie-Hélène |year=2013 |title=Middle Pleistocene ecology and Neanderthal subsistence: Insights from stable isotope analyses in Payre (Ardèche, southeastern France) |journal=[[Journal of Human Evolution]] |volume=65 |issue=4 |pages=363–373 |doi=10.1016/j.jhevol.2013.06.013 |pmid=23920410 |bibcode=2013JHumE..65..363E }}</ref> Neanderthals, nonetheless, were frequently victims of [[animal attack]]s.<ref name=Camaros2016/>

There are multiple instances of Neanderthals practicing [[Human cannibalism|cannibalism]], though it may have only been done in times of extreme food shortages, as in some [[List of incidents of cannibalism|cases in recorded human history]].<ref name=yravedra2015>{{cite journal |last1=Yustos |first1=Marta |last2=Yravedra Sainz de los Terreros |first2=José |date=2015 |title=Cannibalism in the Neanderthal world: an exhaustive revision |journal=Journal of Taphonomy |volume=13 |issue=1 |pages=33–52 }}{{predatory}}</ref>

===The arts=== {{See also|Prehistoric art}} [[File:Neandertal Jewelry (from PLoS).jpg|thumb|upright|Speculative reconstruction of [[white-tailed eagle]] talon jewellery from [[Krapina Neanderthal site|Krapina, Croatia]] (arrows indicate cut marks)]] Neanderthals collected non-functional, uniquely-shaped objects, namely shells, fossils, and gems. It is unclear if these objects were simply picked up for their aesthetic qualities, or if some symbolic significance was applied to them.<ref name=Moncel2012>{{cite journal |first1=M.-H. |last1=Moncel |first2=L. |last2=Chiotti |first3=C. |last3=Gaillard |first4=G. |last4=Onoratini |first5=D. |last5=Pleurdeau |year=2012 |title=Non utilitarian objects in the Palaeolithic: emergence of the sense of precious? |journal=Archaeology, Ethnology & Anthropology of Eurasia |volume=401 |pages=25–27|doi=10.1016/j.aeae.2012.05.004 }}</ref> Some shells may have been painted.<ref name=Hoffman>{{cite journal |author-first1=D. L. |author-last1=Hoffman |author-last2=Angelucci |author-first2=D. E. |author-last3=Villaverde |author-first3=V. |author-last4=Zapata |author-first4=Z. |author-last5=Zilhão |author-first5=J. |year=2018 |title=Symbolic use of marine shells and mineral pigments by Iberian Neandertals 115,000 years ago |journal=Science Advances |volume=4 |issue=2 |article-number=eaar5255 |doi=10.1126/sciadv.aar5255 |pmid=29507889 |pmc=5833998 |bibcode=2018SciA....4.5255H}}</ref> Gibraltarian palaeoanthropologists [[Clive Finlayson|Clive]] and [[Geraldine Finlayson]] suggested that Neanderthals used various bird parts as artistic media, especially black feathers.{{sfn|Finlayson|2019|loc=pp. 129–132}}<ref name=SFinlayson2019/> A 2020 study found evidence of a [[yarn weight|3-ply]] cord fragment made from conifer inner-bark fibres at Abri du Maras, France, which can be used to knit light items, such as strings for hanging beads. 115,000-year-old perforated shell beads from Cueva Antón were possibly strung together to make a necklace.<ref>{{Cite journal |last1=Hardy |first1=B. L. |last2=Moncel |first2=M.-H. |last3=Kerfant |first3=C. |last4=Lebon |first4=M. |last5=Bellot-Gurlet |first5=L. |last6=Mélard |first6=N. |date=April 9, 2020 |title=Direct evidence of Neanderthal fibre technology and its cognitive and behavioral implications |journal=Scientific Reports |language=en |volume=10 |issue=1 |article-number=4889 |doi=10.1038/s41598-020-61839-w |issn=2045-2322 |pmc=7145842 |pmid=32273518|bibcode=2020NatSR..10.4889H }}</ref>

There are several instances of nondescript engravings and scratches on flints, bones, pebbles, and stone slabs — {{as of|2014|lc=y}}, 63 purported engravings have been reported from 27 different European and Middle Eastern Lower-to-Middle Palaeolithic sites. It is debated if these were made with symbolic intent.<ref name=Majkic2018>{{cite journal |first1=A. |last1=Majkić |first2=F. |last2=d'Errico |first3=V. |last3=Stepanchuk |year=2018 |title=Assessing the significance of Palaeolithic engraved cortexes. A case study from the Mousterian site of Kiik-Koba, Crimea |journal=PLOS ONE |volume=13 |issue=5 |article-number=e0195049 |doi=10.1371/journal.pone.0195049 |pmid=29718916 |pmc=5931501 |bibcode=2018PLoSO..1395049M|doi-access=free }}</ref> Neanderthals may have produced [[finger fluting]]s on the walls of La Roche-Cotard over 57,000 years ago.<ref>{{cite journal|first1=J.-C.|last1=Marquet|first2=T. H.|last2=Freiesleben|first3=K. J.|last3=Thomsen|year=2023|title=The earliest unambiguous Neanderthal engravings on cave walls: La Roche-Cotard, Loire Valley, France|journal=PLOS ONE|volume=18|issue=6|article-number=e0286568|doi=10.1371/journal.pone.0286568|doi-access=free|pmid=37343032 |pmc=10284424 |bibcode=2023PLoSO..1886568M }}</ref>

Neanderthals used ochre, a [[clay earth pigment]]. It is unclear if this constitutes evidence of artmaking because, while modern humans have used red ochre for decorative or symbolic colouration, they have also used ochre as medicine, hide tanning agent, food preservative, and insect repellent.<ref name="Roebrooks2012">{{cite journal |display-authors=et al. |year=2012 |title=Use of red ochre by early Neandertals |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=109 |issue=6 |pages=1889–1894 |bibcode=2012PNAS..109.1889R |doi=10.1073/pnas.1112261109 |pmc=3277516 |pmid=22308348 |doi-access=free |author-last1=Roebroeks |author-first1=W. |author-last2=Sier |author-first2=M. J. |author-last3=Nielsen |author-first3=T. K.}}</ref>

The 43,000-year-old [[Divje Babe flute]] (a cave bear [[femur]]) from Slovenia has been attributed by some researchers to Neanderthals, though its status as a [[Paleolithic flute|Palaeolithic flute]] is heavily disputed. Many researchers consider it to be most likely the product of a carnivorous animal chewing the bone.<ref name="Morley06">{{cite journal |last1=Morley |first1=Iain |title=Mousterian Musicianship? The Case of the Divje Babe I Bone |journal=Oxford Journal of Archaeology |date=November 2006 |volume=25 |issue=4 |pages=317–333 |doi=10.1111/j.1468-0092.2006.00264.x }}</ref>

===Technology=== [[File:Mousterian point.png|left|thumb|[[Mousterian]] point|alt=A thin, black triangular rock]] Neanderthals manufactured [[Middle Palaeolithic]] [[stone tool]]s, and are associated with the [[Mousterian]] [[industry (archaeology)|industry]], specifically the [[Levallois technique]]. After developing this technology from the [[Acheulean]] industry,<ref name=Lycett2013>{{cite journal |last1=Lycett |first1=S. J. |last2=von Cramon-Taubadel |first2=N. |title=A 3D morphometric analysis of surface geometry in Levallois cores: patterns of stability and variability across regions and their implications |journal=Journal of Archaeological Science |year=2013 |volume=40 |issue=3 |pages=1508–1517 |doi=10.1016/j.jas.2012.11.005|bibcode=2013JArSc..40.1508L }}</ref> there is a 150,000 year stagnation in Neanderthal stone tool innovation. Stalled technological growth may have followed from their low population, impeding complex ideas from being spread across their range or passed down generationally.<ref name=bocquet2013/><ref name=nakahashi>{{cite journal |last1=Nakahashi |first1=W. |title=The effect of trauma on Neanderthal culture: A mathematical analysis |journal=Homo |year= 2017 |volume=68 |issue=2 |pages=83–100 |doi=10.1016/j.jchb.2017.02.001 |pmid=28238406}}</ref> Neanderthals normally collected raw materials from a nearby source, no more than {{cvt|5|km}}.<ref name=hayden2012/> Some communities were also making tools from shells<ref name=Villa2020>{{cite journal |first1=P. |last1=Villa |first2=S. |last2=Soriano |first3=L. |last3=Pollarolo |year=2020 |title=Neandertals on the beach: use of marine resources at Grotta dei Moscerini (Latium, Italy) |journal=PLOS ONE |volume=15 |issue=1 |article-number=e0226690 |doi=10.1371/journal.pone.0226690 |pmid=31940356 |pmc=6961883 |bibcode=2020PLoSO..1526690V|doi-access=free }}</ref> and bone.<ref>{{cite journal |first1=N. L. |last1=Martisius |first2=F. |last2=Welker |first3=T. |last3=Dogandžić |display-authors=et al. |year=2020 |title=Non-destructive ZooMS identification reveals strategic bone tool raw material selection by Neandertals |journal=Scientific Reports |volume=10 |issue=1 |page=7746 |doi=10.1038/s41598-020-64358-w |pmid=32385291 |pmc=7210944 |bibcode=2020NatSR..10.7746M}}</ref> They may have [[hafting|hafted]] tips onto spears using [[birch bark tar]].<ref>{{cite journal |first1=I. |last1=Degano |first2=S. |last2=Soriano |first3=P. |last3=Villa |first4=L. |last4=Pollarolo |first5=J. J. |last5=Lukejko |first6=Z. |last6=Jacobs |first7=K. |last7=Douka |first8=S. |last8=Vitagliano |first9=C. |last9=Tozzi |year=2019 |title=Hafting of Middle Paleolithic tools in Latium (central Italy): new data from Fossellone and Sant'Agostino caves |journal=PLOS ONE |volume=14 |issue=6 |article-number=e0213473 |doi=10.1371/journal.pone.0213473 |pmc=6586293 |pmid=31220106 |bibcode=2019PLoSO..1413473D|doi-access=free }}</ref> European populations had also been manufacturing wood spears, namely the 400,000 year old British [[Clacton Spear]]; 300,000 year old German [[Schöningen spears]]; and 120,000 year old German [[Lehringen spear|Lehringen Spear]],<ref>{{cite journal|last=Allington-Jones|first=L|year=2015|title=The Clacton Spear: The Last One Hundred Years|journal=Archaeological Journal|volume=172|issue=2|pages=277–278|doi=10.1080/00665983.2015.1008839|hdl=10141/622351|hdl-access=free}}</ref> including both likely thrown (Schöningen)<ref>{{Cite journal |last1=Schoch |first1=Werner H. |last2=Bigga |first2=Gerlinde |last3=Böhner |first3=Utz |last4=Richter |first4=Pascale |last5=Terberger |first5=Thomas |date=December 2015 |title=New insights on the wooden weapons from the Paleolithic site of Schöningen |journal=Journal of Human Evolution |language=en |volume=89 |pages=214–225 |doi=10.1016/j.jhevol.2015.08.004 |pmid=26442632 |bibcode=2015JHumE..89..214S }}</ref> and thrusting (Lehringen) types.<ref name="Yew">Milks, A. (2020) ''[https://sticks-and-stones.blog/wp-content/uploads/2022/11/milks-2021.pdf Yew wood, would you? An exploration of the selection of wood for Pleistocene spears].'' In: Berihuete-Azorin, M., Martin Seijo, M., Lopez-Bulto, O. and Pique, R. (eds.) The Missing Woodland Resources: Archaeobotanical studies of the use of plant raw materials. Advances in Archaeobotany, 6 (6). Barkhuis Publishing, Groningen, pp. 5-22. {{ISBN|978-94-93194-35-9}}</ref> It has been suggested that Neanderthals likely specifically selected particular wood types (such as [[European yew]] in the case of the Clacton and Lehringen spears) for manufacturing spears for their beneficial material properties.<ref name="Yew" />

Neanderthals invented the earliest [[dry distillation]] process to produce [[birch tar]], a substance with adhesive and medicinal properties.<ref name="birch">{{cite journal |last1=Schmidt |first1=Patrick |last2=Koch |first2=Tabea J. |last3=Blessing |first3=Matthias A. |last4=Karakostis |first4=F. Alexandros |last5=Harvati |first5=Katerina |last6=Dresely |first6=Veit |last7=Charrié-Duhaut |first7=Armelle |title=Production method of the Königsaue birch tar documents cumulative culture in Neanderthals |journal=Archaeological and Anthropological Sciences |date=2023 |volume=15 |issue=6 |page=84 |doi=10.1007/s12520-023-01789-2 |pmid=37228449 |pmc=10202989 |bibcode=2023ArAnS..15...84S |issn=1866-9557}}</ref> A complex underground apparatus was used to distill birch tar from masses of bark, which suggests that a high degree of cultural innovation and evolution took place in the European [[Middle Paleolithic]] period.<ref name="birch" /> While Neanderthals may have been using this process 200,000 years ago,<ref name="birch" /> the oldest evidence for dry distillation among modern humans only dates to 20,000 years ago.<ref>{{cite web |last1=Puiu |first1=Tibi |title=Neanderthals were distilling tar 200,000 years ago well before humans |url=https://www.zmescience.com/science/news-science/neanderthals-distilled-tar/ |website=ZME Science |date=4 September 2017}}</ref>

Many Neanderthal sites have evidence of fire, some for extended periods of time.<ref name="Sorensen_etal2018">{{Cite journal |last1=Sorensen |first1=A. C. |last2=Claud |first2=E. |last3=Soressi |first3=M. |year=2018 |title=Neandertal fire-making technology inferred from microwear analysis |journal=Scientific Reports |volume=8 |issue=1 |page=10065 |doi=10.1038/s41598-018-28342-9 |pmid=30026576 |pmc=6053370 |issn=2045-2322 |bibcode=2018NatSR...810065S}}</ref><ref name=Brittingham2019>{{cite journal |first1=A. |last1=Brittingham |first2=M. T. |last2=Hren |first3=G. |last3=Hartman |first4=K. N. |last4=Wilkinson |first5=C. |last5=Mallol |first6=B. |last6=Gasparyan |first7=D. S. |last7=Adler |year=2019 |title=Geochemical evidence for the control of fire by Middle Palaeolithic hominins |journal=Scientific Reports |volume=9 |issue=1 |page=15368 |doi=10.1038/s41598-019-51433-0 |pmid=31653870 |pmc=6814844 |bibcode=2019NatSR...915368B}}</ref><ref>{{cite journal |last1=Heyes |first1=P. J. |last2=Anastasakis |first2=K. |last3=de Jong |first3=W. |last4=van Hoesel |first4=A.|last5=Roebroeks |first5=W. |last6=Soressi |first6=M. |title=Selection and Use of Manganese Dioxide by Neanderthals |journal=Scientific Reports |year= 2016 |volume=6 |issue=1 |page=22,159 |doi=10.1038/srep22159 |pmid=26922901 |pmc=4770591 |bibcode=2016NatSR...622159H}}</ref> They may have been using fire for cooking, keeping warm, and deterring predators.<ref>{{cite journal|journal=PLOS ONE|title=Formation processes, fire use, and patterns of human occupation across the Middle Palaeolithic (MIS 5a-5b) of Gruta da Oliveira (Almonda karst system, Torres Novas, Portugal)| date=2023 | doi=10.1371/journal.pone.0292075 | doi-access=free | last1=Angelucci | first1=Diego E. | last2=Nabais | first2=Mariana | last3=Zilhão | first3=João | volume=18 | issue=10 | article-number=e0292075 |pmid=37819902 | pmc=10566745 |bibcode=2023PLoSO..1892075A }}</ref> They were also capable of zoning areas for specific activities, such as for knapping, butchering, [[hearth]]s, and wood storage.<ref name=hayden2012/> At [[Capellades|Abric Romaní rock shelter]], Spain, Neanderthals may have maintained eight evenly spaced hearths lined up against the rock wall, likely used to stay warm while sleeping, with one person sleeping on either side of the fire.<ref name=kedar2019>{{cite journal |last1=Kedar |first1=Y. |last2=Barkai |first2=R. |title=The significance of air circulation and hearth location at Paleolithic cave sites |journal=Open Quaternary |year=2019 |volume=5 |issue=1 |page=4 |doi=10.5334/oq.52 |doi-access=free}}</ref>

The only known Neanderthal tools that could have been used to fashion clothes are hide [[scraper (archaeology)|scrapers]] as no bone sewing-needles and [[stitching awl]]s have been found as in Cro-Magnon sites. Hide scrapers could have been used to make items similar to blankets or [[poncho]]s. There is no direct evidence that Neanderthals could make fitted clothes from animal hide.<ref name=collard2016>{{cite journal |last1=Collard |first1=M. |last2=Tarle |first2=L. |last3=Sandgathe |first3=D. |last4=Allan |first4=A. |title=Faunal evidence for a difference in clothing use between Neanderthals and early modern humans in Europe |journal=Journal of Anthropological Archaeology |year=2016 |volume=44 |pages=235–246 |doi=10.1016/j.jaa.2016.07.010 |bibcode=2016JAnAr..44..235C |hdl=2164/9989 |hdl-access=free}}</ref><ref name=wales2012>{{cite journal |last=Wales |first=N. |title=Modeling Neanderthal clothing using ethnographic analogues |journal=Journal of Human Evolution |year=2012 |volume=63 |issue=6 |pages=781–795 |doi=10.1016/j.jhevol.2012.08.006 |pmid=23084621|bibcode=2012JHumE..63..781W }}</ref> Unfitted clothes would have limited range of mobility while dressed, and decreased the time Neanderthals could spend unprotected from the elements away from shelters.<ref name=Mukhopadhyay2025>{{cite journal |last1=Mukhopadhyay |first1=A. |last2=Panovska |first2=S. |last3=Garvey |first3=R. |last4=Liemohn |first4=M. W. |last5=Ganjushkina |first5=N. |last6=Brenner |first6=A. |last7=Usoskin |first7=I. |last8=Balikhin |first8=M. |last9=Welling |first9=D. T. |year=2025 |title=Wandering of the auroral oval 41,000 Years Ago |journal=[[Science Advances]] |volume=11 |issue=16 |article-number=eadq7275 |doi=10.1126/sciadv.adq7275 |pmid=40238891 |doi-access=free|pmc=12002135 |bibcode=2025SciA...11.7275M }}</ref> Anterior [[dental microwear]] of Neanderthals living in open environments is similar to that of the modern Ipiutak and Nunavut people, who are known to use their anterior teeth for clamping while preparing hides, suggesting that Neanderthals may have engaged in similar behaviour.<ref>{{cite journal |last1=Krueger |first1=Kristin L. |last2=Ungar |first2=Peter S. |last3=Guatelli-Steinberg |first3=Debbie |last4=Hublin |first4=Jean-Jacques |last5=Pérez-Pérez |first5=Alejandro |last6=Trinkaus |first6=Erik |last7=Willman |first7=John C. |title=Anterior dental microwear textures show habitat-driven variability in Neandertal behavior |journal=Journal of Human Evolution |date=April 2017 |volume=105 |pages=13–23 |doi=10.1016/j.jhevol.2017.01.004 |pmid=28366197 |bibcode=2017JHumE.105...13K }}</ref>

Neanderthals appear to have lived lives of frequent traumatic injury and recovery, indicating the setting of [[splint (medicine)|splints]] and dressing of major wounds. By and large, they appear to have avoided severe infections, indicating long-term treatment. Their knowledge of [[medicinal plants#Prehistoric times|medicinal plants]] was comparable to that of Cro-Magnons.<ref name=Spikins2019>{{cite journal |first1=P. |last1=Spikins |first2=A. |last2=Needham |first3=B. |last3=Wright |year=2019 |title=Living to fight another day: The ecological and evolutionary significance of Neanderthal healthcare |journal=Quaternary Science Reviews |volume=217 |pages=98–118 |doi=10.1016/j.quascirev.2018.08.011 |bibcode=2019QSRv..217...98S |doi-access=free}}</ref>

In 2026, a study published in ''PLOS One'' described a Neanderthal molar recovered from Chagyrskaya Cave in southwestern Siberia, dating to approximately 59,000 years ago, which bears evidence of an intentional dental procedure. Analysis of the tooth, designated Chagyrskaya 64, revealed a deep hole drilled into the chewing surface using a fine-pointed stone tool, extending into the pulp chamber in a manner consistent with cavity intervention to relieve pain — representing the earliest known instance of dental cavity treatment in human evolutionary history. Subsequent wear on the tooth indicates the individual survived the procedure.<ref>{{cite web |url=https://edition.cnn.com/2026/05/13/science/neanderthal-dentistry-stone-drill |title=59,000-year-old Neanderthal tooth shows earliest evidence of dentistry |publisher=CNN |date=2026-05-13 |access-date=2026-05-14}}</ref>

Stone tools on various Greek islands could indicate early seafaring through the Mediterranean, employing simple reed boats for one-day crossings,<ref name=ferentinos2012>{{cite journal |last1=Ferentinos |first1=G. |last2=Gkioni |first2=M. |last3=Geraga |first3=M. |last4=Papatheodorou |first4=G. |year=2012 |title=Early seafaring activity in the southern Ionian Islands, Mediterranean Sea |journal=Journal of Archaeological Science |volume=39 |issue=7 |pages=2167–2176 |doi=10.1016/j.jas.2012.01.032 |bibcode=2011JQS....26..553S}}</ref> but the evidence for such a big claim is limited.{{sfn|Broodbank|2013|pp=107-108}}

===Language=== {{See also|Origin of language|Origin of speech}} It is unclear if Neanderthals had the capacity for complex language, but some researchers have argued that Neanderthals required complex communications to discuss locations, hunting and gathering, and tool-making techniques in order to survive in their harsh environment.<ref name=dediu2018>{{cite journal |first1=D. |last1=Dediu |first2=S. C. |last2=Levinson |year=2018 |title=Neanderthal language revisited: not only us |journal=Current Opinion in Behavioral Sciences |volume=21 |pages=49–55 |doi=10.1016/j.cobeha.2018.01.001 |hdl=21.11116/0000-0000-1667-4 |hdl-access=free}}</ref><ref>{{cite journal |last=Johansson |first=S. |year=2015 |title=Language abilities in Neanderthals |journal=Annual Review of Linguistics |volume=1 |pages=311–322 |doi=10.1146/annurev-linguist-030514-124945|doi-access=free }}</ref><ref name="Whiting2018">{{cite journal |last1=Whiting |first1=K. |last2=Konstantakos |first2=L. |last3=Sadler |first3=G. |last4=Gill |first4=C. |year=2018 |title=Were Neanderthals rational? A stoic approach |journal=Humanities |volume=7 |issue=2 |page=39 |doi=10.3390/h7020039 |doi-access=free}}</ref> In experiments with modern humans, the [[Levallois technique]] can be taught with purely [[observational learning]] without spoken instruction.<ref>{{cite journal|author1=K. Ohnuma|author2=K. Aoki|author3=T. Akazawa|title=Transmission of tool-making through verbal and non-verbal communication: Preliminary experiments in Levallois flake production|journal=[[Journal of Anthropological Sciences]]|volume=105|issue=3|pages=159–68|year=1997|doi=10.1537/ase.105.159|doi-access=free}}</ref>

While the [[hyoid bone]] (a bone that supports the tongue) is almost identical to that of modern humans, this does not provide insight into the entire vocal tract.<ref>{{cite journal|author1=J.T. Laitman|author2=.S. Reidenberg|author3=D.R. Friedland|author4=P.J. Gannon|year=1991| title=What sayeth thou Neanderthal? A look at the evolution of their vocal tract and speech| journal=American Journal of Physical Anthropology| volume=34|issue=S12|page=109|doi=10.1002/ajpa.1330340505|doi-access=free}}</ref> Neanderthals had the [[FOXP2]] gene, which is associated with speech and language development, but not the modern human variant.<ref>{{cite journal |last1=Mozzi |first1=A. |last2=Forni |first2=D. |last3=Clerici |first3=M. |last4=Pozzoli |first4=U. |last5=Mascheretti |first5=S. |year=2016 |title=The evolutionary history of genes involved in spoken and written language: beyond FOXP2 |journal=Scientific Reports |volume=6 |article-number=22157 |bibcode=2016NatSR...622157M |doi=10.1038/srep22157 |pmc=4766443 |pmid=26912479}}</ref>

===Burials and religion=== {{See also|Paleolithic religion|History of religion}} [[File:Neandertal scratches.jpg|thumb|Engraved flint from a Neanderthal grave at {{ill|Kiik-Koba|de}}, Crimea]] Neanderthals, probably uncommonly, buried their dead. This may explain the abundance of fossil remains.{{sfn|Tattersall|2015}} The behaviour is not indicative of a religious belief of [[afterlife|life after death]] because it could also have had [[ritual behavior in animals|non-symbolic motivations]].<ref name=wunn2001/><ref name="PettittOHA2017"/> The dead were buried in simple, shallow graves and pits,<ref name="PettittOHA2017">{{cite book |last1=Pettitt |first1=Paul |title=The Oxford Handbook of Archaeology |date=2017 |publisher=Oxford University Press |location=New York |isbn=978-0-19-885520-0 |pages=354–355 |url=https://global.oup.com/academic/product/the-oxford-handbook-of-archaeology-9780198855200?cc=us&lang=en& |access-date=July 30, 2022}}</ref> but special care seems to have been given to child graves. The graves of children and infants, especially, are associated with grave goods such as artefacts and bones.<ref name=Spikins2014>{{cite journal |first1=P. |last1=Spikins |first2=G. |last2=Hitchens |first3=A. |last3=Needham |display-authors=et al. |year=2014 |title=The Cradle of Thought: Growth, Learning, Play and Attachment in Neanderthal Children |journal=Oxford Journal of Archaeology |volume=33 |issue=2 |pages=111–134 |doi=10.1111/ojoa.12030 |url=http://eprints.whiterose.ac.uk/83027/1/NeanderthalChildhood_OA_Images_sml.pdf}}</ref> Some sites with multiple well-preserved Neanderthal skeletons may represent [[cemeteries]].<ref name="PettittOHA2017"/>

One grave in Shanidar Cave, Iraq, was associated with the pollen of several flowers that may have been in bloom at the time of deposition—yarrow, [[Centaurea|centaury]], [[ragwort]], [[grape hyacinth]], [[joint pine]] and [[hollyhock]].<ref>{{cite journal |last=Leroi-Gourhan |first=A. |title=The flowers found with Shanidar IV, a Neanderthal burial in Iraq |journal=Science |year=1975 |volume=190 |issue=4214 |pages=562–564 |doi=10.1126/science.190.4214.562 |bibcode=1975Sci...190..562L |s2cid=140686473}}</ref> The medicinal properties of the plants led American archaeologist [[Ralph Solecki]] to claim that the man buried was a leader, healer, or [[shaman]], and that "the association of flowers with Neanderthals adds a whole new dimension to our knowledge of his humanness, indicating that he had 'soul{{'"}}.<ref>{{cite journal |last=Solecki |first=R. S. |title=Shanidar IV: a Neanderthal flower burial in northern Iraq |journal=Science |year=1975 |volume=190 |issue=4217 |pages=880–881 |doi=10.1126/science.190.4217.880 |bibcode=1975Sci...190..880S |s2cid=71625677}}</ref> It is also possible the pollen was deposited by a small burrowing rodent after the man's death.<ref>{{cite journal |last=Sommer |first=J. D. |title=The Shanidar IV 'flower burial': a re-evaluation of Neanderthal burial ritual |journal=Cambridge Archaeological Journal |year=1999 |volume=9 |issue=1 |pages=127–129 |doi=10.1017/s0959774300015249|s2cid=162496872 }}</ref>

Neanderthals were once thought to have ritually killed and eaten cave bears or other Neanderthals, but the evidence is circumstantial.<ref name=wunn2001>{{cite journal |first=I. |last=Wunn |year=2001 |title=Cave bear worship in the Paleolithic |journal=Cadernos do Laboratorio Xeolóxico de Laxe |volume=26 |pages=457–463 |url=https://www.udc.es/files/iux/almacen/articulos/cd26_art32.pdf}}</ref> In 2019, the Finlaysons reported that Neanderthals disproportionately butchered the [[golden eagle]] over any [[bird of prey]] or [[corvid]] species, and speculated that Neanderthals viewed the golden eagle as a symbol of power like some recent modern human societies did.<ref name=SFinlayson2019>{{cite journal |first1=S. |last1=Finlayson |first2=G. |last2=Finlayson |author2-link=Geraldine Finlayson |first3=F. G. |last3=Guzman |first4=C. |last4=Finlayson |author4-link=Clive Finlayson |year=2019 |title=Neanderthals and the cult of the sun bird |journal=Quaternary Science Reviews |volume=217 |pages=217–224 |doi=10.1016/j.quascirev.2019.04.010 |bibcode=2019QSRv..217..217F|s2cid=149949579 }}</ref>

==Interbreeding== {{Main|Interbreeding between archaic and modern humans|Neanderthal genetics}}

[[File:Homo sapiens sapiens, Oase, Rumänien (Daniela Hitzemann).jpg|thumb|upright|left|Reconstruction of [[Oase 2]] with around 7.3% Neanderthal DNA (from an ancestor 4–6 generations back)<ref name="fu">{{cite journal |first1=Q. |last1=Fu |first2=M. |last2=Hajdinjak |first3=O. T. |last3= Moldovan |display-authors=3 |first4=S. |last4=Constantin |first5=S. |last5= Mallick |first6=Pontus |last6=Skoglund |first7=N. |last7=Patterson |first8=N. |last8=Rohland |first9=I. |last9=Lazaridis |first10=B. |last10=Nickel |first11=B. |last11=Viola |first12=Kay |last12=Prüfer |first13=M. |last13=Meyer |first14=J. |last14=Kelso |first15=D |last15=Reich |first16=S. |last16=Pääbo |author-link16=Svante Pääbo |title=An early modern human from Romania with a recent Neanderthal ancestor |journal=Nature |year=2015 |volume=524 |issue=7564 |pages=216–219 |doi=10.1038/nature14558 |pmid=26098372 |pmc=4537386 |bibcode=2015Natur.524..216F}}</ref>|alt=A dark-skinned man with black, shiny hair going down to his shoulders, a slight moustache, a goatee, brown eyes, weak eyebrows, wearing a tailored shirt and holding a long spear to support himself]]

The first Neanderthal genome sequence was published in 2010, and strongly indicated interbreeding between Neanderthals and early modern humans.<ref name=green/> Neanderthal-derived genes descend from at least 2 interbreeding episodes outside of Africa: one about 250,000 years ago and another 40,000 to 54,000 years ago. Interbreeding also occurred in other populations which are not ancestral to any living person.<ref name="2023Harris">{{cite journal |author1=Daniel Harris |author2=Alexander Platt |author3=Matthew E.B. Hansen |author4=Shaohua Fan |author5=Michael A. McQuillan |author6=Thomas Nyambo |author7=Sununguko Wata Mpoloka |author8=Gaonyadiwe George Mokone |author9=Gurja Belay |author10=Charles Fokunang |author11=Alfred K. Njamnshi |author12=Sarah A. Tishkoff |title=Diverse African genomes reveal selection on ancient modern human introgressions in Neanderthals |journal=Current Biology |date=2023 |volume=33 |issue=22 |pages=4905–4916.e5 |doi=10.1016/j.cub.2023.09.066 |pmid=37837965 |issn=0960-9822|pmc=10841429 }}</ref> An individual whose ancestry lies beyond sub-Saharan Africa may carry about 2% of Neanderthal DNA. Sub-Saharan Africans can carry Neanderthal DNA presumably descending from back migration (the interbreeding population having migrated back to [[Sub-Saharan Africa]]).<ref name="Chen Wolf Fu Li Akey">{{cite journal |first1=L. |last1=Chen |first2=A. B. |last2=Wolf |first3=W. |last3=Fu |first4=J. M. |last4=Akey |year=2020 |title=Identifying and Interpreting Apparent Neanderthal Ancestry in African Individuals |journal=Cell |volume=180 |issue=4 |pages=677–687.e16 |doi=10.1016/j.cell.2020.01.012 |pmid=32004458 |s2cid=210955842|doi-access=free |pmc=12805117 }}</ref> In all, approximately 20% of the Neanderthal genome appears to have survived in the modern human [[gene pool]].<ref name=vernot2014>{{cite journal |title=Resurrecting surviving Neandertal lineages from modern human genomes |journal=Science |volume=343 |issue=6174 |pages=1017–1021 |year=2014 |bibcode=2014Sci...343.1017V |last1=Vernot |first1=B. |last2=Akey |first2=J. M. |doi=10.1126/science.1245938 |pmid=24476670 |s2cid=23003860|doi-access=free }}</ref> This Neanderthal DNA is derived primarily from the children of female modern humans and male Neanderthals.<ref>{{cite journal |last1=Platt |first1=Alexander |last2=Harris |first2=Daniel N. |last3=Tishkoff |first3=Sarah A. |title=Interbreeding between Neanderthals and modern humans was strongly sex biased |journal=Science |year= 2026 |volume=391 |issue=6788 |pages=922–925 |doi=10.1126/science.aea6774 |pmid=41747031 |bibcode=2026Sci...391..922P }}</ref><ref name="Reilly2022">{{Cite journal |last1=Reilly |first1=Patrick F. |last2=Tjahjadi |first2=Audrey |last3=Miller |first3=Samantha L. |last4=Akey |first4=Joshua M. |last5=Tucci |first5=Serena |date=September 2022 |title=The contribution of Neanderthal introgression to modern human traits |journal=Current Biology |language=en |volume=32 |issue=18 |pages=R970–R983 |doi=10.1016/j.cub.2022.08.027 |pmc=9741939 |pmid=36167050|bibcode=2022CBio...32.R970R }}</ref><ref name="Chevy Huerta-Sánchez Ramachandran 2023 p. e1010399">{{cite journal | last1=Chevy | first1=Elizabeth T. | last2=Huerta-Sánchez | first2=Emilia | last3=Ramachandran | first3=Sohini | title=Integrating sex-bias into studies of archaic introgression on chromosome X | journal=PLOS Genetics | volume=19 | issue=8 | date=August 14, 2023 | issn=1553-7404 | pmid=37578977 | pmc=10449224 | doi=10.1371/journal.pgen.1010399 | doi-access=free | article-number=e1010399 |quote= We have shown that the observed low level of archaic coverage on chromosome X could be explained merely by a reduction in the effect of heterosis and sex-biases in the introgression events, without involving a more complex model with hybrid incompatibilities. Our work also suggests that negative selection was likely acting on archaic variants, and provides an appropriate set of null models for evaluating positive selection on introgressed segments on chromosome X.}}</ref> Due to their low population and proliferation of deleterious mutations, many Neanderthal genes were probably selected out of the modern human gene pool ([[negative selection (natural selection)|negative selection]]). Similarly, a large portion of surviving introgression appears to be [[non-coding DNA|non-coding]] ("junk") DNA with few biological functions.{{sfn|Reich|2018}} Some Neanderthal-derived genes, nonetheless, may have functional implications related to metabolism, brain function, and skeletal and muscular development.<ref name=green/><ref name=Dolgova2018>{{cite journal |first1=O. |last1=Dolgova |first2=O. |last2=Lao |year=2018 |title=Evolutionary and medical consequences of archaic introgression into modern human genomes |journal=Genes |volume=9 |issue=7 |page=358 |doi=10.3390/genes9070358 |pmc=6070777 |pmid=30022013 |doi-access=free}}</ref> Some genes may have helped immigrating modern human populations acclimatise faster, such as genes related to [[immune response]].<ref name=Nedelec2016>{{cite journal |first1=Y. |last1=Nédélec |first2=J. |last2=Sanz |first3=G. |last3=Baharian |display-authors=et al. |year=2016 |title=Genetic ancestry and natural selection drive population differences in immune responses to pathogens |journal=Cell |volume=167 |issue=3 |pages=657–669 |doi=10.1016/j.cell.2016.09.025 |pmid=27768889 |doi-access=free}}</ref>

The genetic evidence suggests that the interbreeding mainly took place between Neanderthal men and modern human women.<ref name="Papini 2020 p. 619">{{cite book | last=Papini | first=M. | title=Comparative Psychology: Evolution and Development of Brain and Behavior, 3rd Edition | publisher=Taylor & Francis | year=2020 | isbn=978-1-000-17770-1 | url=https://books.google.com/books?id=hvj3DwAAQBAJ&pg=PT619 | access-date=2024-03-28 | page=619 |quote="There is evidence of somewhat selective interbreeding. mtDNA from Neanderthals is absent in modern humans. Because mtDNA is carried only by females, interbreeding may have occurred mainly between Neanderthal males and modern females (Krings, Stone, Schmitz, Krainitzki, Stoneking, & Pääbo, 1997). "}}</ref> This is based on the absence of Neanderthal mtDNA, as well as the relatively low level of Neanderthal admixture in the modern human [[X chromosome]].<ref name=Serre2004>{{cite journal |doi=10.1371/journal.pbio.0020057 |title=No evidence of Neandertal mtDNA contribution to early modern humans |year=2004 |author-last1=Serre |author-first1=D. |author-last2=Langaney |author-first2=A. |author-last3=Chech |author-first3=M. |author-last4=Teschler-Nicola |author-first4=M. |author-last5=Paunovic |author-first5=M. |author-last6=Mennecier |author-first6=P. |author-last7=Hofreiter |author-first7=M. |author-last8=Possnert |author-first8=G. |author-last9=Pääbo |author-first9=S. |author-link9=Svante Pääbo |journal=PLOS Biology |volume=2 |issue=3 |article-number=e57 |pmid=15024415 |pmc=368159 |doi-access=free }}</ref><ref name="KringsStone1997">{{cite journal |last1=Krings |first1=M. |last2=Stone |first2=A. |last3=Schmitz |first3=R. W. |last4=Krainitzki |first4=H. |last5=Stoneking |first5=M. |last6=Pääbo |first6=S. |author-link6=Svante Pääbo|title=Neandertal DNA sequences and the origin of modern humans |journal=Cell |year= 1997 |volume=90 |issue=1 |pages=19–30 |doi=10.1016/s0092-8674(00)80310-4 |pmid=9230299 |hdl=11858/00-001M-0000-0025-0960-8 |s2cid=13581775 |hdl-access=free}}</ref><ref name=luo2018>{{cite journal |last1=Luo |first1=S. |last2=Valencia |first2=C. A. |last3=Zhang |first3=J. |last4=Lee |first4=N.-C. |last5=Slone |first5=J. |last6=Gui |first6=B. |last7=Wang |first7=X.|last8=Li |first8=Z.|last9=Dell |first9=S. |last10=Brown |first10=J. |last11=Chen |first11=S. M.|last12=Chien |first12=Y.-H. |last13=Hwu |first13=W.-L. |last14=Fan |first14=P.-C.|last15=Wong |first15=L.-J. |last16=Atwal |first16=P. S. |last17=Huang |first17=T.|title=Biparental inheritance of mitochondrial DNA in humans |journal=Proceedings of the National Academy of Sciences |year=2018 |volume=115 |issue=51 |pages=13039–13044 |doi=10.1073/pnas.1810946115 |pmid=30478036 |pmc=6304937|bibcode=2018PNAS..11513039L |doi-access=free }}</ref> According to [[Svante Pääbo]], it is not clear that modern humans were socially dominant over Neanderthals, which may explain why the interbreeding occurred primarily between Neanderthal males and modern human females.<ref name="Paabo 2014 p. 19">{{cite book | last=Paabo | first=S. | title=Neanderthal Man: In Search of Lost Genomes | publisher=Basic Books | year=2014 | isbn=978-0-465-02083-6 | url=https://books.google.com/books?id=cguNAgAAQBAJ&pg=PA19 | access-date=2024-03-28 | page=19}}</ref> A 2026 study confirmed the evidence of sex bias, reporting that [[sexual selection]] (i.e., a preference for male Neanderthals and modern human females) was the primary driver of the asymmetric mating patterns, rather than sex biases in demography or migration.<ref>{{cite journal |last1= Platt|first1= Alexander|last2= Harris|first2= Daniel N.|last3= Tishkoff|first3= Sarah A.|title=Interbreeding between Neanderthals and modern humans was strongly sex biased |journal= Science|date=February 2026 |volume=391 |issue=6788 |pages= 922–925|doi=10.1126/science.aea6774 |pmid= 41747031|bibcode= 2026Sci...391..922P|url=https://www.science.org/doi/abs/10.1126/science.aea6774 |language=en}}</ref>

Neanderthals in the Siberian Altai Mountains interbred with the local Denisovan population, and it may have been a common occurrence here.<ref>{{cite journal |author=Warren, Matthew |title=Mum's a Neanderthal, dad's a Denisovan: First discovery of an ancient-human hybrid |journal=Nature News |volume=560 |issue=7719 |pages=417–418 |year=2018 |doi= 10.1038/d41586-018-06004-0 |pmid=30135540 |bibcode=2018Natur.560..417W |doi-access=free}}</ref> About 17% of the genome of one Altai Denisovan specimen derived from Neanderthals.<ref name=Pennisi>{{cite journal |doi=10.1126/science.340.6134.799 |pmid=23687020 |title=More genomes from Denisova Cave show mixing of early human groups |journal=Science |volume=340 |issue=6134 |page=799 |year=2013 |last1=Pennisi |first1=E. |author-link=Elizabeth Pennisi |bibcode=2013Sci...340..799P}}</ref>

Even before genetic evidence had confirmed this admixture, hybridisation between Neanderthals and early modern humans had been proposed early on,<ref name="McFarland">{{cite book |url=http://www.electricscotland.com/webclans/cairney/14.htm|title=Clans and families of Ireland and Scotland, an ethnography of the Gael |last1=Cairney |first1=C. T. |year=1989 |publisher=McFarland |isbn=0-89950-362-4 |page=14}}</ref> such as by English anthropologist [[Thomas Huxley]] in 1890,<ref>{{cite journal |last=Huxley |first=T. |author-link=Thomas Huxley |year=1891 |title=The Aryan question and pre-historic man |url=http://aleph0.clarku.edu/huxley/CE7/Aryan.html |journal=The Popular Science Monthly |volume=38 |pages=512–516}}</ref> Danish ethnographer [[Hans Peder Steensby]] in 1907,<ref>{{cite journal |first=H. P. |last=Steensby |title=Racestudier i Danmark |trans-title=Race Studies in Denmark |language=da |url=http://img.kb.dk/tidsskriftdk/pdf/gto/gto_0019-PDF/gto_0019_67206.pdf |journal=Geographical Journal |publisher=Geografisk Tidsskrift |volume=9 |year=1907}}</ref> and Coon in 1962.<ref>{{cite journal |last=Coon |first=C. S. |author-link=Carleton Stevens Coon |title=The origin of races |journal=Science |year=1962 |volume=140 |issue=3563 |publisher=Knopf |pages=548–549 |doi=10.1126/science.140.3563.208 |pmid=14022816 |url=https://archive.org/details/B-001-001-289/page/n631}}</ref> In the early 2000s, supposed hybrid specimens were discovered: [[Lagar Velho 1]]<ref name="Tattersall">{{cite journal |doi=10.1073/pnas.96.13.7117 |title=Hominids and hybrids: The place of Neanderthals in human evolution |year=1999 |last1=Tattersall |first1=I. |author1-link=Ian Tattersall |last2=Schwartz |first2=J. H. |journal=Proceedings of the National Academy of Sciences |volume=96 |issue=13 |pages=7117–19 |jstor=48019 |bibcode=1999PNAS...96.7117T |pmid=10377375 |pmc=33580|doi-access=free }}</ref><ref name="DuarteMauricio1999">{{cite journal |last1=Duarte |first1=C. |last2=Maurício |first2=J. |last3=Pettitt |first3=P. B. |last4=Souto |first4=P. |last5=Trinkaus |first5=E.|author-link5=Erik Trinkaus|last6=van der Plicht |first6=H. |last7=Zilhão |first7=J. |title=The early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern human emergence in Iberia |journal=Proceedings of the National Academy of Sciences of the United States of America |year=1999 |volume=96 |issue=13 |pages=7604–7609 |doi=10.1073/pnas.96.13.7604 |pmid=10377462 |pmc=22133 |bibcode=1999PNAS...96.7604D|doi-access=free }}</ref><ref name="speciation">{{cite journal |last1=Hublin |first1=J. J. |year=2009 |title=The origin of Neandertals |journal=Proceedings of the National Academy of Sciences |volume=106 |issue=38 |pages=16022–16027 |bibcode=2009PNAS..10616022H |doi=10.1073/pnas.0904119106 |jstor=40485013|pmc=2752594 |pmid=19805257|doi-access=free }}</ref><ref>{{Cite journal |last1=Harvati |first1=K. |last2=Frost |first2=S. R. |last3=McNulty |first3=K. P. |year=2004 |title=Neanderthal taxonomy reconsidered: implications of 3D primate models of intra- and interspecific differences |journal=Proceedings of the National Academy of Sciences |volume=101 |issue=5 |pages=1147–52 |bibcode=2004PNAS..101.1147H |doi=10.1073/pnas.0308085100 |pmc=337021 |pmid=14745010|doi-access=free }}</ref> and [[Peștera Muierilor|Muierii 1]].<ref>{{cite journal |doi=10.1073/pnas.0608443103 |title=Early modern humans from the Peștera Muierii, Baia de Fier, Romania |year=2006 |last1=Soficaru |first1=A. |last2=Dobos |first2=A. |last3=Trinkaus |first3=E. |author-link3=Erik Trinkaus |journal=Proceedings of the National Academy of Sciences |volume=103 |issue=46 |pages=17196–17201 |jstor=30052409 |bibcode=2006PNAS..10317196S |pmid=17085588 |pmc=1859909|doi-access=free }}</ref> These proposals were based on shared anatomy, but modern geneticists have cautioned that some shared anatomy might have been acquired by adaptation to a similar environment, rather than admixture alone.{{sfn|Reich|2018}}

==Extinction== {{Main|Neanderthal extinction}}

[[File:Protoaurignacian Fumane Cave retouched bladelets Falcucci PlosOne 2017.png|upright=1.5|thumb|The Neanderthal [[Mousterian]] culture was replaced by modern human [[Aurignacian]] culture (above, [[Protoaurignacian]] bladelets).<ref name="higham2014"/>]] The extinction of Neanderthals was part of the broader [[Late Pleistocene extinctions|Late Pleistocene megafaunal extinction event]].<ref>{{cite journal |last1=Hortolà |first1=Policarp |last2=Martínez-Navarro |first2=Bienvenido |title=The Quaternary megafaunal extinction and the fate of Neanderthals: An integrative working hypothesis |journal=Quaternary International |date=May 2013 |volume=295 |pages=69–72 |doi=10.1016/j.quaint.2012.02.037 |bibcode=2013QuInt.295...69H }}</ref> Neanderthals were replaced by modern humans, indicated by the near-complete replacement of Middle Palaeolithic Mousterian stone technology with modern human [[Upper Palaeolithic]] [[Aurignacian]] stone technology across Europe (the Middle-to-Upper Palaeolithic Transition) from 39,000 to 41,000 years ago.<ref name="higham2014">{{cite journal |last1=Higham |first1=T. |last2=Douka |first2=K. |last3=Wood |first3=R. |last4=Ramsey |first4=C. B. |last5=Brock |first5=F. |last6=Basell |first6=L. |last7=Camps |first7=M. |last8=Arrizabalaga |first8=A. |last9=Baena |first9=J. |last10=Barroso-Ruíz |first10=C. |author11=C. Bergman |author12=C. Boitard |author13=P. Boscato |author14=M. Caparrós |author15=N.J. Conard |display-authors=etal |year=2014 |title=The timing and spatiotemporal patterning of Neanderthal disappearance |journal=[[Nature (journal)|Nature]] |volume=512 |issue=7514 |pages=306–309 |bibcode=2014Natur.512..306H |doi=10.1038/nature13621 |pmid=25143113 |s2cid=205239973 |hdl-access=free |author16=C. Draily |author17=A. Froment |author18=B. Galván |author19=P. Gambassini |author20=A. Garcia-Moreno |author21=S. Grimaldi |author22=P. Haesaerts |author23=B. Holt |author24=M.-J. Iriarte-Chiapusso |author25=A. Jelinek |author26=J.F. Jordá Pardo |author27=J.-M. Maíllo-Fernández |author28=A. Marom |author29=J. Maroto |author30=M. Menéndez |author31=L. Metz |author32=E. Morin |author33=A. Moroni |author34=F. Negrino |author35=E. Panagopoulou |author36=M. Peresani |author37=S. Pirson |author38=M. de la Rasilla |author39=J. Riel-Salvatore |author40=A. Ronchitelli |author41=D. Santamaria |author42=P. Semal |author43=L. Slimak |author44=J. Soler |author45=N. Soler |author46=A. Villaluenga |author47=R. Pinhasi |author48=R. Jacobi |quotation=We show that the Mousterian [the Neanderthal tool-making tradition] ended by 41,030–39,260 calibrated years BP (at 95.4% probability) across Europe. We also demonstrate that succeeding 'transitional' archaeological industries, one of which has been linked with Neanderthals (Châtelperronian), end at a similar time. |hdl=1885/75138}}</ref><ref name=higham2011>{{cite journal |first=T. |last=Higham |year=2011 |title=European Middle and Upper Palaeolithic radiocarbon dates are often older than they look: problems with previous dates and some remedies |journal=Antiquity |volume=85 |issue=327 |pages=235–249 |doi=10.1017/s0003598x00067570 |s2cid=163207571 |quotation=Few events of European prehistory are more important than the transition from ancient to modern humans about 40,000 years ago, a period that unfortunately lies near the limit of radiocarbon dating. This paper shows that as many as 70 per cent of the oldest radiocarbon dates in the literature may be too young, due to contamination by modern carbon.}}</ref><ref>{{Cite journal |last1=Agusti |first1=J. |last2=Rubio-Campillo |first2=X. |year=2017 |title=Were Neanderthals responsible for their own extinction? |journal=[[Quaternary International]] |volume=431 |pages=232–237 |bibcode=2017QuInt.431..232A |doi=10.1016/j.quaint.2016.02.017}}</ref> Neanderthals may have persisted in Spain for longer, but the dates of the latest Mousterian and earliest Aurignacian are poorly constrained. In [[Catalonia]] and [[Aragón]] (northern Spain), the Mousterian may have survived to about 39,000 years ago, and in southern Spain and Gibraltar potentially 32,000 to 35,000 years ago.<ref>{{cite journal|first1=Lawrence Guy|last1=Straus|year=2020|title=Neanderthal last stand? Thoughts on Iberian refugia in late MIS 3|journal=Journal of Quaternary Science|issn=1099-1417|pages=283–290|volume=37|issue=2|doi=10.1002/jqs.3252}}</ref> Similar refuge zones have also been proposed on other temperate European peninsulas, namely Italy, the Balkans, and [[Crimea]].<ref>{{cite journal|first1=Nuno|last1=Bicho|first2=Milena|last2=Carvalho|title=Peninsular southern Europe refugia during the Middle Palaeolithic: an introduction|journal=Journal of Quaternary Science|year=2022 |issn=1099-1417|pages=133–135|volume=37|issue=2|doi=10.1002/jqs.3410|doi-access=free|bibcode=2022JQS....37..133B }}</ref><ref>{{cite journal|first1=E. M.|last1=Pigott|first2=T.|last2=Uthmeier|first3=V.|last3=Chabai|first4=T. F. G.|last4=Higham|title=The Late Middle and Early Upper Palaeolithic in Crimea (Ukraine)—A Review of the Neanderthal Refugium Hypothesis|journal=Journal of Paleolithic Archaeology|year=2024|issn=2520-8217|page=27|volume=7|issue=1|doi=10.1007/s41982-024-00194-y|doi-access=free|bibcode=2024JPalA...7...27P }}</ref>

Historically, the cause of extinction of Neanderthals and other archaic humans was viewed under an imperialistic guise, with the superior invading modern humans [[prehistoric warfare|exterminating]] and replacing the inferior species.<ref name=drell2000/>

{{blockquote|text=When ''sapiens'' began to expand and spread, he eliminated the other contemporary races [including Neanderthals] just as the white man drove out the Australian aborigines and the North American Indians.|author=[[Ernst Mayr]], 1950<ref name=Mayr1950>{{cite journal |last=Mayr |first=E. |author-link=Ernst Mayr |year=1950 |title=Taxonomic categories in fossil hominids |journal=Cold Spring Harbor Symposia on Quantitative Biology |volume=15 |pages=109–118 |doi=10.1101/SQB.1950.015.01.013|pmid=14942702 }}</ref>}}

In general, the extinction of Neanderthals is ascribed predominantly to [[competitive exclusion principle|competition]] with modern humans. The success of modern humans over Neanderthals is usually attributed to a higher birth rate and population, facilitated by better long-distance mobility and more complex technologies and subsistence strategies. Some Neanderthal populations may have also been assimilated into modern human populations rather than being ecologically outcompeted.<ref name=Timmermann2020>{{cite journal|first1=Axel|last1=Timmermann|title=Quantifying the potential causes of Neanderthal extinction: Abrupt climate change versus competition and interbreeding|journal=Quaternary Science Reviews|year=2020|issn=0277-3791|article-number=106331|volume=238|doi=10.1016/j.quascirev.2020.106331|doi-access=free|bibcode=2020QSRv..23806331T }}</ref> Assimilation had long been hypothesised with supposed hybrid specimens, and was revitalised with the discovery of archaic human DNA in modern humans.<ref name=Villa2014>{{cite journal |first1=P. |last1=Villa |first2=W. |last2=Roebroeks |year=2014 |title=Neandertal demise: an archaeological analysis of the modern human superiority complex |journal=PLOS ONE |volume=9 |issue=4 |article-number=e96424 |doi=10.1371/journal.pone.0096424 |pmc=4005592 |pmid=24789039 |bibcode=2014PLoSO...996424V|doi-access=free }}</ref> Similarly, the [[Châtelperronian]] industry of central France and northern Spain may represent a culture of Neanderthals adopting modern human techniques, via [[acculturation]].<ref>{{cite journal|first1=I.|last1=Djakovic|first2=M.|last2=Roussel|first3=M.|last3=Soressi|year=2024|title=Stone Tools in Shifting Sands: Past, Present, and Future Perspectives on the Châtelperronian Stone Tool Industry|journal=Journal of Paleolithic Archaeology|volume=7|issue=1 |article-number=29 |doi=10.1007/s41982-024-00193-z|doi-access=free|bibcode=2024JPalA...7...29D }}</ref><ref>{{cite journal |first1=M. |last1=Roussel |first2=M. |last2=Soressi |first3=J.-J. |last3=Hublin |year=2016 |title=The Châtelperronian conundrum: blade and bladelet lithic technologies from Quinçay, France |journal=Journal of Human Evolution |volume=95 |pages=13–32 |doi=10.1016/j.jhevol.2016.02.003 |pmid=27260172|bibcode=2016JHumE..95...13R }}</ref> Other ambiguous transitional cultures include the Italian [[Uluzzian]] industry,<ref>{{cite journal |first1=P. |last1=Villa |first2=L. |last2=Pollarolo |first3=J. |last3=Conforti |display-authors=et al. |year=2018 |title=From Neandertals to modern humans: new data on the Uluzzian |journal=PLOS ONE |volume=13 |issue=5 |article-number=e0196786 |doi=10.1371/journal.pone.0196786 |pmc=5942857 |pmid=29742147 |bibcode=2018PLoSO..1396786V|doi-access=free }}</ref> and the Central European [[Szeletian]] industry.<ref name="Hoffecker2009">{{cite journal |last=Hoffecker |first=J. F. |year=2009 |title=The spread of modern humans in Europe |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=106 |issue=38 |pages=16040–16045 |bibcode=2009PNAS..10616040H |doi=10.1073/pnas.0903446106 |pmc=2752585 |pmid=19571003 |doi-access=free}}</ref>

Neanderthal extinction has also been ascribed to their low population as well as the resulting [[mutational meltdown]], making them less adaptable to major environmental changes or new diseases introduced by immigrating modern humans.<ref>{{cite journal |first1=A. |last1=Degioanni |first2=C. |last2=Bonenfant |first3=S. |last3=Cabut |first4=S. |last4=Condemi |year=2019 |title=Living on the edge: Was demographic weakness the cause of Neanderthal demise? |journal=PLOS ONE |volume=14 |issue=5 |article-number=e0216742 |doi=10.1371/journal.pone.0216742 |pmc=6541251 |pmid=31141515 |bibcode=2019PLoSO..1416742D |doi-access=free}}</ref> It is unclear if climatic degradation would have severely impacted Neanderthals given how many glacial periods they persisted through in Europe. If areas were depopulated of Neanderthals as a consequence of climate change (specifically [[Heinrich event]] 4) or a natural disaster (the [[Campanian Ignimbrite eruption]]), Neanderthals may not have been as fast as modern humans in recolonising.<ref name="Staubwasser">{{cite journal |last1=Staubwasser |first1=M. |last2=Drăgușin |first2=V. |last3=Onac |first3=B. P. |year=2018 |title=Impact of climate change on the transition of Neanderthals to modern humans in Europe |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=115 |issue=37 |pages=9116–9121 |bibcode=2018PNAS..115.9116S |doi=10.1073/pnas.1808647115 |pmc=6140518 |pmid=30150388 |doi-access=free}}</ref> The [[Laschamp event]] 39,000 to 42,000 years ago may have increased [[ultraviolet]] radiation, disproportionately affecting Neanderthals who lacked protective fitted clothes, and may not have utilised ochre as sunscreen to the extent Cro-Magnons did.<ref name=Mukhopadhyay2025/>

==In popular culture== {{Main|Neanderthals in popular culture}}

[[File:BlackTerror1636.jpg|thumb|Cavemen in ''[[wikisource: The Black Terror/16|The Black Terror #16]]'' (1946)|alt=3 panels, a brawny man standing to the left and a child to the right on the grass in front of a cave. The man is holding a hammer in his right hand and has only 2 teeth visible, the child a spear and dragging a cat behind him, and both are dressed in wraps around their waist with a strap around one shoulder. The man says, "Well, son, you did very well on your first hunting trip...now you get your second lesson in surviving!" The child says, "Me got food, what else do me have to do?" to which the man responds, "You got to learn to start fire and cook food!" The third panel shows the child looking at a small tipi pyre, the man walks away and says, "There is wood, now you light! I can't tell how—you must find out for yourself!" and the child says to himself, "Jumping mammoths. Me gotta figure out all alone! I know, will use flint stone to make sparks!"]]

Neanderthals have been portrayed in popular culture including appearances in literature, visual media and comedy. The "[[caveman]]" [[archetype]] often mocks Neanderthals and depicts them as primitive, hunchbacked, knuckle-dragging, club-wielding, grunting, nonsocial characters driven solely by animal instinct. "Neanderthal" can also be used as an insult.{{sfn|Papagianni|Morse|2013}}

In literature, they are sometimes depicted as brutish or monstrous, such as in [[H. G. Wells]]' ''The Grisly Folk'' and [[Elizabeth Marshall Thomas]]' ''The Animal Wife'', but sometimes with a civilised but unfamiliar culture, as in [[William Golding]]'s ''[[The Inheritors (Golding novel)|The Inheritors]]'', [[Björn Kurtén]]'s ''[[Dance of the Tiger]]'', and [[Jean M. Auel]]'s ''[[Clan of the Cave Bear]]'' and her ''[[Earth's Children]]'' series.<ref name=drell2000/>

==See also== * {{annotated link|Denisovan}} * [[Early human migrations]] * {{annotated link|Cro-Magnon}} * {{annotated link|Homo floresiensis|''Homo floresiensis''}} * {{annotated link|Homo luzonensis|''Homo luzonensis''}} * {{annotated link|Homo naledi|''Homo naledi''}} * {{annotated link|Timeline of human evolution}}

== Footnotes == <templatestyles src="Reflist/styles.css" /><div class="reflist reflist-lower-alpha"><references group="lower-alpha" /></div>

== References == <references />

==Sources== *{{cite book|last=Broodbank |first=Cyprian |title=The Making of the Middle Sea: A History of the Mediterranean from the Beginning to the Emergence of the Classical World |publisher=Thames & Hudson |location =London |year=2013|isbn=978-0-500-29208-2}} * {{cite book |first1=K. |last1=Brown |first2=D. A. |last2=Fa |first3=G. |last3=Finlayson |author3-link=Geraldine Finlayson |first4=C. |last4=Finlayson |author4-link=Clive Finlayson |year=2011 |chapter=Small game and marine resource exploitation by Neanderthals: the evidence from Gibraltar |title=Trekking the shore: changing coastlines and the antiquity of coastal settlement |series=Interdisciplinary contributions to archaeology |publisher=Springer |isbn=978-1-4419-8218-6 |chapter-url=https://www.researchgate.net/publication/227198441 |ref={{harvid |Brown |2011}}}} * {{cite book |first=C. |last=Finlayson |author-link=Clive Finlayson |year=2019 |title=The smart Neanderthal: bird catching, cave art, and the cognitive revolution |publisher=Oxford University Press |isbn=978-0-19-251812-5 |url={{google books |plainurl=yes |id=7hqJDwAAQBAJ}}}} *{{cite book|last=French |first=Jennifer |title=Palaeolithic Europe: A Demographic and Social Prehistory |publisher=Cambridge University Press |year=2021|isbn=978-1-108-49206-5}} *{{cite book |chapter-url=https://books.google.com/books?id=9As7CwAAQBAJ&pg=PT171 |first1=D. |last1=Papagianni |first2=M. A. |last2=Morse |year=2013 |chapter=Still with us? |title=Neanderthals rediscovered: how modern science is rewriting their story |publisher=Thames and Hudson |isbn=978-0-500-77311-6}} * {{cite book |first=D. |last=Reich |author-link=David Reich (geneticist) |year=2018 |chapter=Encounters with Neanderthals |title=Who we are and how we got here: ancient DNA and the new science of the human past |publisher=Oxford University Press |isbn=978-0-19-882125-0 |url={{google books |plainurl=yes |id=uLNSDwAAQBAJ |page=25}}}} * {{cite book |first=P. |last=Shipman |year=2015 |title=The invaders: how humans and their dogs drove Neanderthals to extinction |publisher=Harvard University Press |jstor=j.ctvjf9zbs |isbn=978-0-674-42538-5 |chapter=How humans and their dogs drove Neanderthals to extinction|doi=10.2307/j.ctvjf9zbs }} * {{cite book |first=I. |last=Tattersall |author-link=Ian Tattersall |year=2015 |chapter=Neanderthals, DNA, and creativity |title=The strange case of the Rickety Cossack: and other cautionary tales from human evolution |publisher=St. Martin's Publishing Group |isbn=978-1-4668-7943-0}}

==Further reading== *{{cite journal| last= Hunt|first=Chris |display-authors=etal| journal= Journal of Archaeological Science|title=Shanidar et ses fleurs? Reflections on the palynology of the Neanderthal 'Flower Burial' hypothesis|volume= 159|number= |date=August 28, 2023|article-number=105822 |doi=10.1016/j.jas.2023.105822 |bibcode=2023JArSc.159j5822H |s2cid=261325698 |issn= |doi-access=free}} *{{cite book|last1=Pettitt|first1=Paul|last2=White|first2=Mark|title=The British Palaeolithic: hominin societies at the edge of the Pleistocene world|date=2012|publisher=Routledge|location=London|isbn=978-0-415-67454-6}} * {{cite book |last1=Romagnoli |first1=Francesca |last2=Rivals |first2=Florent |last3=Benazzi |first3=Stefano |title=Updating Neanderthals: Understanding Behavioural Complexity in the Late Middle Palaeolithic |date=2022 |publisher=Academic Press |isbn=978-0-12-821429-9 |url=https://books.google.com/books?id=sTJcEAAAQBAJ&pg=PA78 |language=en}} * {{cite book |last1=Stringer |first1=C. |author-link=Chris Stringer |last2=Gamble |first2=C. |year=1993 |title=In search of the Neanderthals |url=https://archive.org/details/insearchofneande00stri |publisher=Thames and Hudson |isbn=0-500-05070-8 |ref={{harvid |Stringer |1993}}}} *{{cite book |first = Rebecca Wragg |last = Sykes | title=[[Kindred: Neanderthal Life, Love, Death and Art]] |year=2020 |publisher=Bloomsbury Sigma |location=London |isbn= 978-1-4729-3749-0}}

==External links== {{Commons category|Homo neanderthalensis}} {{Wikibooks|Introduction to Paleoanthropology}} {{Wikispecies|Homo neanderthalensis}} {{NIE Poster|Neanderthal Man}} {{Wikinews|Neanderthals 'knew what they were doing': Archæologist Dr Naomi Martisius discusses her findings about Neanderthals' behaviour with Wikinews}} * {{cite web | title = ''Homo neanderthalensis'' | publisher = The Smithsonian Institution | url = http://humanorigins.si.edu/evidence/human-fossils/species/homo-neanderthalensis| date = February 14, 2010 }} * [http://humanorigins.si.edu/evidence/human-evolution-timeline-interactive Human Timeline (Interactive)] – [[Smithsonian Institution|Smithsonian]], [[National Museum of Natural History]] (August 2016). * {{cite web |title=Neanderthal DNA |publisher=International Society of Genetic Genealogy |url=http://www.isogg.org/neanderthaldna.htm |archive-url=https://web.archive.org/web/20060617204513/http://www.isogg.org/neanderthaldna.htm |archive-date=June 17, 2006 }}: Includes Neanderthal mtDNA sequences * [https://www.ncbi.nlm.nih.gov/search/all/?term=homo%20sapiens%20neanderthalensis GenBank records for ''H. s. neanderthalensis''] maintained by the [[National Center for Biotechnology Information]] (NCBI) * {{cite web |last1=Alex |first1=Bridget |title=What's Behind the Evolution of Neanderthal Portraits |url=https://www.sapiens.org/archaeology/neanderthal-art-paleolithic-archaeology/ |website=SAPIENS |date=February 21, 2024}} *[https://theclimatechronicles.com/ The Climate Chronicles], explores the impact of Pleistocene climate change on Neanderthals and other hominins. {{Prehistoric technology}} {{Human Evolution}} {{Homo neanderthalensis|state=expanded}} {{#invoke:Portal bar|main|Evolutionary biology|Science}} {{#invoke:Taxonbar|taxonbar|from=Q40171}} {{#invoke:Authority control|authorityControl}}

[[Category:Neanderthals| ]] [[Category:Fossil taxa described in 1864]] [[Category:Stone Age Asia]] [[Category:Stone Age Europe]] [[Category:Extinct apex predators]]