Neanderthal#Language

{{See also|Human taxonomy}}

File:Neanderthal excavation site.JPG 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 and Jürgen Thissen.}}|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]]

Neanderthals are named after the 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 spelling reform of 1901.{{efn|The German spelling Thal ("valley") was current until 1901 but has been Tal since then. (The German noun is cognate with English dale.) The German /t/ phoneme was frequently spelled th from the 15th to 19th centuries, but the spelling Tal became standardised in 1901 and the old spellings of the German names Neanderthal for the valley and Neanderthaler for the species were both changed to the spellings without h.}} 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Neandertal always refers to the valley.{{efn|In Mettmann, "Neander Valley", there is a local idiosyncrasy in use of the outdated spellings with th, such as with the Neanderthal Museum (but the name is in English [German would require Neandertalermuseum]), the Neanderthal station (Bahnhof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.{{cite web |url=https://web.archive.org/web/20200809042411/https://www.mettmann.de/neandertal/schreibweise.php |title=Neandertal oder Neanderthal? Was ist denn nun richtig? |publisher=Kreisstadt Mettmann |trans-title=Neandertal or Neanderthal? So which is actually right? |access-date=February 1, 2017 |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'.)}}}} The valley itself was named after the late 17th century German theologian and hymn writer Joachim Neander, who often visited the area. His grandfather, a musician, had changed the family name from the original German Neumann ("new man") to the Graeco-Roman form Neander, following the fashion of the time.{{cite EB1911 |wstitle=Neander, Joachim |volume=19 |pages=320–321}}

Neanderthal can be pronounced using the {{IPA|/t/}} (as in {{IPAc-en|n|i|ˈ|æ|n|d|ər|t|ɑː|l}}){{cite web |url=https://www.collinsdictionary.com/dictionary/english/neanderthal |title=Neanderthal |website=Collins English Dictionary |access-date=February 18, 2020}} or the standard English pronunciation of th with the fricative /θ/ (as {{IPAc-en|n|i|ˈ|æ|n|d|ər|θ|ɔː|l}}).{{cite web |url=https://www.merriam-webster.com/dictionary/Neanderthal |title=Neanderthal |website=Merriam-Webster Dictionary |access-date=February 18, 2020}}{{cite web |url=https://www.ahdictionary.com/word/search.html?q=Neanderthal |title=Neanderthal |website=American Heritage Dictionary |access-date=February 18, 2020}} The latter pronunciation, nevertheless, has no basis in the original German word which is pronounced always with a t regardless of the historical spelling.{{cite magazine|first=B.|last=Alex|year=2016|title=Is It Neander-TAL or Neander-THAL?|magazine=Discover|access-date=March 3, 2025|url=https://www.discovermagazine.com/planet-earth/is-it-neander-tal-or-neander-thal}}

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".{{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, UK |pages=302, 473}} The binomial name Homo neanderthalensis—extending the name "Neanderthal man" from the individual specimen to the entire species, and formally recognising it as distinct from humans—was first proposed by Irish geologist William King in a paper read to the 33rd British Science Association in 1863.{{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}}{{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.|s2cid=131804686 }}{{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 |s2cid=224796921}} 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 [theistic{{efn|King made a typo and said "theositic".}}] conceptions".{{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}}

File:Calotte crânienne, type de l'espèce Homo neanderthalensis, vallée de Néander.jpg 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]]

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 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.{{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}} 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.{{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 }}

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.{{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 }}}} 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 (hip bone), part of the right shoulder blade, and pieces of the ribs.{{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}}

File:Guide leaflet (1901) (14767192135).jpg'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) Heidelberg Man

7) Neanderthal Man

8) Cro-Magnon Man

9) Australian Black-fellow (pejorative term for Aboriginal Australians)

10) Hottentot (pejorative term for South African pastoralists)

11) Chinese

12) American Caucasian

]]

Following Charles Darwin's 1859 On the Origin of Species, Fuhlrott and Schaaffhausen argued that Neanderthal 1 represents a primitive lower human form, aligning more closely with non-human apes as well as Negroids, Eskimos, and Aboriginal Australians (which were variably classified as separate species or subspecies of human at the time).{{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}}{{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}} 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 senility, disease, and malformation instead of archaicness,{{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}} which stalled Neanderthal research until the end of the century.{{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 |s2cid=54616107 }}

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 notably 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.{{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}}{{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 |s2cid=164060946}}{{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}}{{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 |s2cid=145778787 |url=http://blog.wbkolleg.unibe.ch/wp-content/uploads/Sommer2006_MirrorMirrorNeanderthals.pdf}}

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 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. Several specimens around the Old World were classified as "progressive" Neanderthals which would eventually evolve into some local subspecies of H. sapiens (polycentricism), or in Europe into either the modern European subspecies or the "classic" Neanderthals.{{cite journal|first=F.|last=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}}

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. At around the same time, the "Out of Asia" hypothesis was overturned by the "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").{{cite journal|last=Holliday|first=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}} In 2010, the first mapping of the Neanderthal genome demonstrated that there was at least some interbreeding between archaic and modern humans.{{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}} Subsequent genetic studies continue to raise questions on how Neanderthals should be classified relative to modern humans.

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 (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).{{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 | page=e1010399}}}} which if true could justify species distinction. The crux of the issue lies in the vagueness of the term "species" (the species problem).{{cite journal|first=A.|last=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|doi=10.1093/evolinnean/kzae033|doi-access=free}}{{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}}{{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 |s2cid=15005225 |url=https://digitalcommons.wayne.edu/cgi/viewcontent.cgi?article=1316&context=humbiol}}

Among identified archaic humans, Neanderthals are most closely related to Denisovans based on nuclear DNA (nDNA) analyses. Denisovans are an enigmatic group of Late Pleistocene humans only recognisable by a genetic signature.{{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. |date=September 25, 2020 |title=The evolutionary history of Neanderthal and Denisovan Y chromosomes |url=https://www.science.org/doi/10.1126/science.abb6460 |journal=Science |language=en |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 |issn=0036-8075|hdl-access=free }} Likely due to more recent interbreeding episodes, the mitochondrial DNA (mtDNA, passed down maternally){{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 |page=16046 |year=2017 |doi=10.1038/ncomms16046| pmid=28675384 | pmc=5500885 | bibcode=2017NatCo...816046P}} and Y-chromosome DNA (passed down paternally){{cite journal|first1=Martin|last1=Petr|first2=Mateja|last2=Hajdinjak|first3=Qiaomei|last3=Fu|first4=Elena|last4=Essel|title=The evolutionary history of Neanderthal and Denisovan Y chromosomes|url=https://www.science.org/doi/10.1126/science.abb6460|journal=Science|year=2020|pages=1653–1656|volume=369|issue=6511|doi=10.1126/science.abb6460|first5=Hélène|last5=Rougier|first6=Isabelle|last6=Crevecoeur|first7=Patrick|last7=Semal|first8=Liubov V.|last8=Golovanova|first9=Vladimir B.|last9=Doronichev|first10=Carles|last10=Lalueza-Fox|first11=Marco|last11=de la Rasilla|first12=Antonio|last12=Rosas|first13=Michael V.|last13=Shunkov|first14=Maxim B.|last14=Kozlikin|first15=Anatoli P.|last15=Derevianko|first16=Benjamin|last16=Vernot|first17=Matthias|last17=Meyer|first18=Janet|last18=Kelso|hdl=21.11116/0000-0007-11C2-A|hdl-access=free}} 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 placed as ancestors to Neanderthals using nDNA, but their mtDNA aligns more closely with Denisovans.{{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 |s2cid=4467094}}

A 2021 phylogeny of some Middle Pleistocene and Neanderthal fossils using tip dating:{{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=English |volume=2 |issue=3 |doi=10.1016/j.xinn.2021.100130 |issn=2666-6758 |pmc=8454562 |pmid=34557770 |last13=Stringer |first13=Chris |last12=Grün |first12=Rainer |page=100130 |bibcode=2021Innov...200130N}}

{{Clade|label1=Homo|{{Clade

|1=H. habilis

|2={{clade

|1=H. erectus

|2={{clade

|1=Tighennif

|2={{clade

|1={{clade

|1=Saldanha

|2={{clade

|1={{clade

|1=Petralona

|2=Kabwe 1

}}

|2={{clade

|1=Ceprano

|2=Bodo

}}

}}

}}

|2={{clade

|1={{clade

|1=Mauer 1

|2=Arago

}}

|2={{clade

|1={{clade

|1=Narmada

|2=Maba

}}

|2={{clade

|1=Ndutu

|2={{clade

|label1=Neanderthal clade

|1={{clade

|1=Sima de los Huesos

|2={{clade

|1=Saccopastore

|2={{clade

|1={{clade

|1={{clade

|1=Tabun

|2=Gibraltar 1

}}

|2={{clade

|1=Spy

|2=Neanderthal 1

}}

}}

|2={{clade

|1={{clade

|1=Amud 1

|2=Saint-Césaire

}}

|2={{clade

|1=Shanidar

|2={{clade

|1=La Chapelle-aux-Saints 1

|2=La Ferrassie 1

}}

}}

}}

}}

}}

}}

|2=H. sapiens

}}

}}

}}

}}

}}

}}

}}

}}

}}

File:Skull5 complete right Sima de los Huesos.jpg") from Sima de los Huesos, Spain]]

Typical Neanderthal skull traits appear in the European fossil record near the beginning of the Middle Pleistocene, in specimens usually classified as 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 periods may have forced populations into small refugia, reducing genetic diversity, leading to the development of other typical Neanderthal traits through genetic drift or pleiotropy. The 120,000 to 140,000-year-old Israeli Nesher Ramla remains may represent one such source population which would recolonise Europe following the Penultimate Glacial Period.{{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}}

The occurrence of typical Neanderthal traits in the Middle Pleistocene was highly variable even among individuals of the same population. The speed of Neanderthalization may have also been impeded by gene flow between Western Europe and Africa, exemplified by anomalous specimens which lack typical Neanderthal traits, such as Ceprano Man.{{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 |s2cid=4467094}} The "classical Neanderthal" anatomy appears by the Last Interglacial (Eemian).

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 even earlier.{{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}}{{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 }} 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.{{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 |page=eaay5483 |doi=10.1126/sciadv.aay5483 |pmid=32128408 |pmc=7032934 |bibcode=2020SciA....6.5483R}}

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 mutations. It is unclear how long European populations suffered this population stress, or to what extent it influenced Neanderthalization.{{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 |page=20130374 |doi=10.1098/rstb.2013.0374 |pmc=4275882 |pmid=25487326}}

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

File:Tabun 1 NMNH.jpg, 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 Southwest 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.{{cite journal |first1=J. |last1=Serangeli |first2=M. |last2=Bolus |url=http://www.quartaer.eu/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 |access-date=October 11, 2022 |archive-date=February 29, 2020 |archive-url=https://web.archive.org/web/20200229163117/http://www.quartaer.eu/pdfs/2008/2008_serangeli.pdf |url-status=dead }}

The southernmost find was recorded at Shuqba Cave, Levant;{{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 |s2cid=191630165}} reports of Neanderthals from the North African Jebel Irhoud{{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 }} and Haua Fteah{{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 }} have been reidentified as H. sapiens. Their easternmost presence is recorded at Denisova Cave, Siberia 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.{{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}} The northernmost bound is generally accepted to have been 55°N, with unambiguous sites known between 50–53°N, but this is difficult to assess because glacial advances destroy most human remains.{{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}}{{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 |s2cid=165849999}} Middle Palaeolithic artefacts have been found up to 60°N on the Russian plains,{{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 }}{{cite journal |first1=L. |last1=Slimak |first2=J. I. |last2=Svendsen |first3=J. |last3=Mangerud |first4=H. |last4=Plisson |year=2011 |title=Late Mousterian persistence near the Arctic Circle |journal=Science |volume=332 |issue=6031 |pages=841–845 |doi=10.1126/science.1203866 |pmid=21566192 |jstor=29784275 |bibcode=2011Sci...332..841S |s2cid=24688365}}{{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 |pages=167 |doi=10.1126/science.1210211 |pmid=22246757 |bibcode=2012Sci...335..167S |doi-access=free}} but these are more likely attributed to modern humans.{{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}}

File:Weichsel-Würm-Glaciation.png 70–20,000 years ago]]

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. Stable environments with mild mean annual temperatures may have been the most suitable Neanderthal habitats.{{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 |url=https://www.cambridge.org/core/product/identifier/S0033589423000327/type/journal_article |journal=Quaternary Research |language=en |volume=116 |pages=108–132 |doi=10.1017/qua.2023.32 |bibcode=2023QuRes.116..108P |issn=0033-5894 |access-date=February 7, 2024|hdl=2164/21236 |hdl-access=free }}

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.{{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 |pages=e1006340 |doi=10.1371/journal.pgen.1006340 |pmid=27824859 |pmc=5100956 |doi-access=free }}{{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 }} Archaeological evidence suggests that the initial Cro-Magnon population was 10 times higher than Neanderthals.{{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 |s2cid=28256970}}

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.{{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 }}{{cite journal |first1=J. |last1=Bocquet-Appel |first2=A. |last2=Degioanni |year=2013 |title=Neanderthal demographic estimates |journal=Current Anthropology |volume=54 |pages=202–214 |doi=10.1086/673725 |s2cid=85090309}} 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.{{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 }} Infant mortality was estimated to have been very high for Neanderthals, about 43% in northern Eurasia.{{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 |s2cid=43859422}}

{{clear}}

{{Neanderthal map}}

{{Main|Neanderthal anatomy}}

File:Neanderthal cranial anatomy.jpg

The Neanderthal skull has a flat and broad skullcap, rounded supraorbital torus (the brow ridges), high 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, and an occipital bun at the back of the skull.{{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=978-0-306-45924-5}}

The Neanderthal braincase averages {{cvt|1,640|cm3}} for males and {{cvt|1,460|cm3}} for females,{{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=978-0-471-84376-4}} which is significantly larger than the averages for all living populations.{{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 |url=http://syslearn.oregonstate.edu/instruction/anth/smith/TimeMach1984.pdf |journal=Current Anthropology |volume=12 |issue=3 |pages=301–30 |doi=10.1086/203138 |s2cid=86147507}} The largest Neanderthal brain, Amud 1, was calculated to be {{cvt|1,736|cm3}}, one of the largest ever recorded in humans.{{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 |s2cid=36974955 |url=https://www.zora.uzh.ch/id/eprint/120401/6/AmudPaper.pdf}} Neanderthal brain organisation differs in areas related to cognition and language, which may be implicated in the comparative simplicity of Neanderthal behaviour compared to Cro-Magnons in the archaeological record.{{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 }}{{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 |s2cid=9755048 |url=http://nrs.harvard.edu/urn-3:HUL.InstRepos:3716473 |doi-access=free }}{{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 }}

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.{{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 }} A large nose does not necessarily equate to a better sense of smell, and neurologically, because the olfactory bulbs are smaller, Neanderthals may have had a poorer sense of smell and olfactory memory than modern humans.{{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 }}

The cheek bones are strong, the incisors are large and shovel-shaped, the 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 attritive foods, or because Neanderthals regularly used the mouth as a third hand.{{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 }}

File:Neanderthal and homo sapiens sapiens.jpg]]

Neanderthals were generally short and stocky. In a sample of 45 Neanderthal long bones 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. The fossil record shows that adult Neanderthals varied from about {{cvt|147.5|to|177|cm|ftin|0}} in height.{{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 }} Average male body mass index would have been 26.9–28.2 (overweight) using a size of {{cvt|164 to 168|cm|ftin|0}} and {{cvt|76|kg}}.{{Cite journal |last=Helmuth |first=H. |title=Body height, body mass and surface area of the Neanderthals |journal=Zeitschrift für Morphologie und Anthropologie |volume=82 |issue=1 |pages=1–12 |year=1998 |pmid=9850627 |jstor=25757530}}{{cite journal |last1=Froehle |first1=A. W. |last2=Churchill |first2=S. E. |year=2009 |title=Energetic competition between Neandertals and anatomically modern humans |journal=PaleoAnthropology |pages=96–116 |url=http://www.paleoanthro.org/journal/content/PA20090096.pdf}}

The Neanderthal chest was deep and wide, with a proportionally expansive thoracic cavity, and possibly stronger lung performance. Neanderthals also had relatively more fast-twitch muscle fibres,{{cite journal|first=M.|last=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|page=11762|doi=10.1038/s41598-022-15344-x|doi-access=free|pmc=9273616|pmid=35817835}} and much higher caloric demands.{{cite journal |first1=A. W. |last1=Froehle |first2=S. E. |last2=Churchill |year=2009 |title=Energetic competition between Neanderthals and anatomically modern humans |journal=PaleoAnthropology |pages=96–116 |url=http://paleoanthro.reedd.webfactional.com/static/journal/content/PA20090096.pdf}} The limbs are proportionally short. The body plan has traditionally been explained as a "hyper-arctic" adaptation (Allen's rule).{{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}}{{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}}{{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 }} Stronger lungs, more fast-twitch muscle, and shorter limbs would have also boosted sprinting efficiency.{{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 |s2cid=133980969 |url=https://www.researchgate.net/publication/329811193}}

Skin colour seems to have ranged from dark to light. Some Neanderthals had dark or brown hair.{{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 }}{{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}} If red was another possible hair colour, it does not appear to have been a common one.{{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}}

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.{{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}} 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{{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 |page=e0186684 |doi=10.1371/journal.pone.0186684 |pmc=5650169 |pmid=29053746 |bibcode=2017PLoSO..1286684T|doi-access=free }} (perhaps swimmer's ear).{{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 |page=e0220464 |doi=10.1371/journal.pone.0220464 |pmid=31412053 |pmc=6693685 |bibcode=2019PLoSO..1420464T|doi-access=free }} The high trauma rate may be ascribed to a dangerous hunting strategy,{{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 }} or frequent animal attacks.{{cite journal |url=https://www.researchgate.net/publication/276353034 |first1=E. |last1=Camarós |first2=M. |last2=Cueto |first3=C. |last3=Lorenzo |first4=V. |last4=Villaverde |year=2016 |title=Large carnivore attacks on hominins during the Pleistocene: a forensic approach with a Neanderthal example |journal=Archaeological and Anthropological Sciences |volume=8 |issue=3 |pages=635–646 |doi=10.1007/s12520-015-0248-1 |bibcode=2016ArAnS...8..635C |hdl=10550/54275 |s2cid=82001651 |hdl-access=free}}

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 defects than in modern humans.{{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 |page=20130374 |doi=10.1098/rstb.2013.0374 |pmc=4275882 |pmid=25487326}}

{{Main|Neanderthal behavior}}

File:Neanderthal genetic subgroups.png

It is difficult to infer Neanderthal group size, but indirect data generally suggests small bands of 10 to 30 individuals.{{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}} 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 over a century.{{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}} Neanderthals may have been outcompeting cave bears for cave space.{{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}} Intergroup movement may have been predominantly female-driven, with at least some groups practicing patrilocal residency (the woman moves out of her group to live with her mate).{{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 }}

Neanderthals maintained a low population across their range, which may have hindered their ability to maintain long-distance trade routes{{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 |page=20130168 |doi=10.1098/rspb.2013.0168 |pmc=3619466 |pmid=23486442}} and avoid inbreeding.{{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 |page=20130374 |doi=10.1098/rstb.2013.0374 |pmc=4275882 |pmid=25487326}} They may have regularly interacted with closely neighbouring communities within a region, but not as often beyond.{{cite journal |url=https://www.academia.edu/4182063 |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 }} 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.{{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 |pages=e5151 |doi=10.1371/journal.pone.0005151 |pmc=2664900 |pmid=19367332 |bibcode=2009PLoSO...4.5151F|doi-access=free }}

While Cro-Magnons are usually assumed to have practised sexual division of labour with men hunting and women gathering like in recent hunter-gatherer societies, it is unclear to what extent this can 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, on the other hand, could indicate men and women typically carried different items with their mouths, maybe during tasks not related to subsistence. The women at El Sidrón Cave, Spain, may have been eating more seeds and nuts than the men. The lack of a strict sexual division in Neanderthals has usually been linked to their small population and group size, falling short of the demographic threshold where task specialisation becomes feasible — which may also explain the comparative simplicity of Neanderthal material culture.{{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}}

{{See also|Pleistocene human diet}}

File:Homo sapiens neanderthalensis-Jäger.jpg]]

Neanderthals were once thought of as scavengers, but are now considered to have been apex predators.{{cite journal |last=Jaouen |first=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 |volume=116 |issue=11 |pages=4928–4933 |doi=10.1073/pnas.1814087116 |pmid=30782806 |pmc=6421459|bibcode=2019PNAS..116.4928J |doi-access=free }} They appear to have eaten predominantly what was abundant within their immediate surroundings.{{sfn|Tattersall|2015}} Cro-Magnons, in contrast, seem to have used more complex food extraction strategies and generally had a more diverse diet.{{cite journal |first1=S. |last1=El Zaatari |first2=F. E. |last2=Grine |first3=P. S. |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 |page=e0153277 |doi=10.1371/journal.pone.0153277 |pmc=4847867 |pmid=27119336 |bibcode=2016PLoSO..1153277E|doi-access=free }}

In many European sites, prey items include reindeer, horse, aurochs, and steppe bison. Neanderthals in Southwest Asia more commonly hunted mountain gazelle, Persian fallow deer, wild goat, and camels. They may have less frequently taken down larger Pleistocene megafauna whenever locally abundant, such as woolly mammoth and woolly rhinoceros.{{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}} At the 125,000 year old Neumark-Nord site, Germany, there is evidence of regular hunting of straight-tusked elephants maybe every 5 to 6 years.{{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 |pages=eadd8186 |doi=10.1126/sciadv.add8186 |issn=2375-2548 |pmc=9891704 |pmid=36724231 |bibcode=2023SciA....9D8186G }} 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.{{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 }}{{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}}

Neanderthals possibly employed a wide range of food preparation techniques. At Cueva del Sidrón, Spain, Neanderthals may have been roasting and smoking meat, and used certain plants—such as yarrow and camomile—for flavouring,{{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 |volume=89 |issue=344 |pages=464–471 |doi=10.15184/aqy.2014.7 |s2cid=86646905 |url=https://www.researchgate.net/publication/283861536}} although these plants may have instead been used for their medicinal properties.{{cite journal |first1=S. |last1=Buckley |first2=K. |last2=Hardy |first3=M. |last3=Huffman |year=2013 |title=Neanderthal self-medication in context |journal=Antiquity |volume=87 |issue=337 |pages=873–878 |doi=10.1017/S0003598X00049528 |s2cid=160563162 |url=https://www.researchgate.net/publication/256374897|doi-access=free }}{{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 |url=https://www.researchgate.net/publication/229160372 |bibcode=2012NW.....99..617H |s2cid=10925552}} At Gorham's Cave, Gibraltar, Neanderthals may have been roasting pinecones to access pine nuts,{{sfn|Brown|2011}} and at Gruta da Figueira Brava, brown crabs to soften the shell before cracking them open.{{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}} 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. It is possible these Neanderthals were curing and storing all this meat before winter set in.{{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}}

Neanderthals competed with several large carnivores, but also seem to have hunted them down, namely cave lions and wolves,{{sfn|Shipman|2015|loc=pp. 120–143}} as well as cave and brown bear both in and out of hibernation.{{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}} Neanderthals and other predators may have sometimes avoided competition by pursuing different prey, namely with cave hyenas{{cite book |author-first=G. L. |author-last=Dusseldorp |title=Zooarchaeology and Modern Human Origins |chapter=Neanderthals and Cave Hyenas: Co-existence, Competition or Conflict? |year=2013 |pages=191–208 |chapter-url=https://link.springer.com/content/pdf/10.1007%2F978-94-007-6766-9_12.pdf |doi=10.1007/978-94-007-6766-9_12 |series=Vertebrate paleobiology and paleoanthropology |publisher=Springer Science+Business Media Dordrecht |editor1-first=J. L. |editor1-last=Clark |editor2-first=J. D. |editor2-last=Speth |isbn=978-94-007-6765-2}} and wolves (niche differentiation).{{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 |date=October 2013 |title=Middle Pleistocene ecology and Neanderthal subsistence: Insights from stable isotope analyses in Payre (Ardèche, southeastern France) |url=https://linkinghub.elsevier.com/retrieve/pii/S0047248413001590 |journal=Journal of Human Evolution |language=en |volume=65 |issue=4 |pages=363–373 |doi=10.1016/j.jhevol.2013.06.013 |pmid=23920410}} Neanderthals, nonetheless, were frequently victims of animal attacks.

There are multiple instances of Neanderthals practicing cannibalism, but it may have only been done in times of extreme food shortages, as in some cases in recorded human history.{{cite journal |author-first1=J. |author-last1=Yravedra |author-first2=M. |author-last2=Yustos |year=2015 |title=Cannibalism in the Neanderthal world: an exhaustive revision |journal=Journal of Taphonomy |volume=13 |issue=1 |pages=33–52 |url=https://www.researchgate.net/publication/316915426}}

{{See also|Prehistoric art}}

File:Neandertal Jewelry (from PLoS).jpg talon jewellery from 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.{{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 |url=https://www.researchgate.net/publication/230703161}} Some shells may have been painted.{{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 |pages=eaar5255 |doi=10.1126/sciadv.aar5255 |pmid=29507889 |pmc=5833998 |bibcode=2018SciA....4.5255H}} Gibraltarian palaeoanthropologists Clive and Geraldine Finlayson suggested that Neanderthals used various bird parts as artistic media, especially black feathers.{{sfn|Finlayson|2019|loc=pp. 129–132}} A 2020 study found evidence of a 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.{{Cite journal |last=Hardy |first=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=2020-04-09 |title=Direct evidence of Neanderthal fibre technology and its cognitive and behavioral implications |url=https://www.nature.com/articles/s41598-020-61839-w |journal=Scientific Reports |language=en |volume=10 |issue=1 |doi=10.1038/s41598-020-61839-w |issn=2045-2322 |pmc=7145842 |pmid=32273518}}

There are several instances of nondescript engravings and scratches on flints, bones, pebbles, and stone slabs — as of 2014, 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.{{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 |page=e0195049 |doi=10.1371/journal.pone.0195049 |pmid=29718916 |pmc=5931501 |bibcode=2018PLoSO..1395049M|doi-access=free }} Neanderthals may have produced finger flutings on the walls of La Roche-Cotard over 57,000 years ago.{{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|page=e0286568|doi=10.1371/journal.pone.0286568|doi-access=free|pmid=37343032 |bibcode=2023PLoSO..1886568M }}

Neanderthals used ochre, a clay earth pigment. While modern humans have used this for decorative or symbolic colouration, it has also been used as medicine, hide tanning agent, food preservative, and insect repellent.{{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.}}

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 Palaeolithic flute is heavily disputed. Many researchers consider it to be most likely the product of a carnivorous animal chewing the bone.{{cite journal | doi=10.1111/j.1468-0092.2006.00264.x | title=Mousterian Musicianship? The Case of the Divje Babe I Bone | year=2006 | last1=Morley | first1=Iain | journal=Oxford Journal of Archaeology | volume=25 | issue=4 | pages=317–333 |url=https://www.researchgate.net/publication/227587744 |access-date=May 30, 2024 }}

File:Mousterian point.png point|alt=A thin, black triangular rock]]

Neanderthals manufactured Middle Palaeolithic stone tools, and are associated with the Mousterian industry, specifically the Levallois technique. After developing this technology from the Acheulean industry,{{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 }} 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.{{cite journal |first1=J. |last1=Bocquet-Appel |first2=A. |last2=Degioanni |year=2013 |title=Neanderthal demographic estimates |journal=Current Anthropology |volume=54 |pages=202–214 |doi=10.1086/673725 |s2cid=85090309}}{{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}} Neanderthals normally collected raw materials from a nearby source, no more than {{cvt|5|km}}. Some communities were also making tools from shells{{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 |page=e0226690 |doi=10.1371/journal.pone.0226690 |pmid=31940356 |pmc=6961883 |bibcode=2020PLoSO..1526690V|doi-access=free }} and bone.{{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}} They may have hafted tips onto spears using birch bark tar.{{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 |page=e0213473 |doi=10.1371/journal.pone.0213473 |pmc=6586293 |pmid=31220106 |bibcode=2019PLoSO..1413473D|doi-access=free }} European populations have also been manufacturing yew wood thrusting spears for close-range combat, namely the 400,000 year old British Clacton Spear; 300,000 year old German Schöningen spears; and 120,000 year old German Lehringen Spear.{{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}}

Many Neanderthal sites have evidence of fire, some for extended periods of time, though it is unclear whether they were capable of starting fire or simply scavenged from naturally occurring wildfires.{{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 |pages=10065 |doi=10.1038/s41598-018-28342-9 |pmid=30026576 |pmc=6053370 |issn=2045-2322 |bibcode=2018NatSR...810065S}}{{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}}{{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}} They may have been using fire for cooking, keeping warm, and deterring predators.{{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 | pages=e0292075 |pmid=37819902 | pmc=10566745 |bibcode=2023PLoSO..1892075A }} They were also capable of zoning areas for specific activities, such as for knapping, butchering, hearths, and wood storage. At Abric Romaní rock shelter, Spain, Neanderthas 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.{{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}}

The only known Neanderthal tools that could have been used to fashion clothes are hide scrapers as no bone sewing-needles and stitching awls have been found as in Cro-Magnon sites. Hide scrapers could have been used to make items similar to blankets or ponchos. There is no direct evidence that Neanderthals could make fitted clothes from animal hide.{{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 |hdl=2164/9989 |hdl-access=free}}{{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 }} Unfitted clothes would have limited range of mobility while dressed, and decreased the time Neanderthals could spend unprotected from the elements away from shelters.{{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 |doi=10.1126/sciadv.adq7275 |pmid=40238891 |doi-access=free|pmc=12002135 }}

Neanderthals appear to have lived lives of frequent traumatic injury and recovery, indicating the setting of splints and dressing of major wounds. By and large, they appear to have avoided severe infections, indicating long-term treatment. The quality of medical care may have ensured their survival as a species for so long. Their knowledge of medicinal plants was comparable to that of Cro-Magnons.{{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}}

Stone tools on various Greek islands could indicate early seafaring through the Mediterranean, employing simple reed boats for one-day crossings,{{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}} but the evidence for such a big claim is limited.{{sfn|Broodbank|2013|pp=107-108}}

{{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.{{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 |url=https://pure.mpg.de/rest/items/item_2521815_7/component/file_2538918/content |hdl=21.11116/0000-0000-1667-4 |s2cid=54391128 |hdl-access=free}}{{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 }}{{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}} In experiments with modern humans, the Levallois technique can be taught with purely observational learning without spoken instruction.{{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}}

Anatomically, the Neanderthal hyoid bone (which supports the tongue) is almost identical to that in modern humans, but this does not provide insight of the entire vocal tract.{{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}} Neanderthals had the FOXP2 gene, which is associated with speech and language development, but not the modern human variant.{{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 |pages=22157 |bibcode=2016NatSR...622157M |doi=10.1038/srep22157 |pmc=4766443 |pmid=26912479}}

{{See also|Paleolithic religion|History of religion}}

File:Neandertal scratches.jpg, 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 life after death because it could also have had non-symbolic motivations. The dead were buried in simple, shallow graves and pits,{{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}} 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.{{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}} Some sites with multiple well-preserved Neanderthal skeletons may represent cemeteries.

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, centaury, ragwort, grape hyacinth, joint pine and hollyhock.{{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}} The medicinal properties of the plants led American archaeologist Ralph Solecki to claim that the man buried was some 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{{'"}}.{{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}} It is also possible the pollen was deposited by a small rodent after the man's death.{{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 }}

Neanderthals were once thought to have ritually killed and eaten cave bears or other Neanderthals, but the evidence is circumstantial.{{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}} The Finlayson's speculate that Neanderthals viewed the golden eagle as a symbol of power.{{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 }}

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

File:Homo sapiens sapiens, Oase, Rumänien (Daniela Hitzemann).jpg with around 7.3% Neanderthal DNA (from an ancestor 4–6 generations back){{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}}|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]]

Hybridisation between Neanderthals and early modern humans had been suggested early on,{{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=978-0-89950-362-2 |page=14}} such as by English anthropologist Thomas Huxley in 1890,{{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}} Danish ethnographer Hans Peder Steensby in 1907,{{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}} and Coon in 1962.{{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}} In the early 2000s, supposed hybrid specimens were discovered: Lagar Velho 1{{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 }}{{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 }}{{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 }}{{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 }} and Muierii 1.{{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 }} Similar anatomy could also have been caused by adapting to a similar environment rather than interbreeding.{{sfn|Reich|2018}}

The first Neanderthal genome sequence was published in 2010, and strongly indicated interbreeding between Neanderthals and early modern humans. 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. Interbreeding also occurred in other populations which are not ancestral to any living person.{{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 |url=https://www.sciencedirect.com/science/article/pii/S0960982223013155 |access-date=8 May 2024 |issn=0960-9822|pmc=10841429 }} 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 modern human migration between Eurasia and Africa.{{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 }} In all, approximately 20% of the Neanderthal genome appears to have survived in the modern human gene pool.{{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 }} This Neanderthal DNA may descend primarily from the children of female modern humans and male Neanderthals.{{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 }}{{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 | page=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.}}

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). Similarly, a large portion of surviving introgression appears to be non-coding ("junk") DNA with few biological functions.{{sfn|Reich|2018}} Some Neanderthal-derived genes, nonetheless, may have functional (though not necessarily positive) implications related to metabolism, brain function, and skeletal and muscular development.{{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}} Some genes may have helped immigrating modern humans populations acclimatise faster, such as genes related to immune response.{{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}}

Neanderthals in the Siberian Altai Mountains interbred with the local Denisovan population, and it may have been a common occurrence here.{{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}} About 17% of the genome of one Altai Denisovan specimen derived from Neanderthals.{{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}}

{{Main|Neanderthal extinction}}

File:Protoaurignacian Fumane Cave retouched bladelets Falcucci PlosOne 2017.png culture was replaced by modern human Aurignacian culture (above, Protoaurignacian bladelets).]]

The extinction of Neanderthals was part of the broader Late Pleistocene megafaunal extinction event.{{Cite journal |last1=Hortolà |first1=Policarp |last2=Martínez-Navarro |first2=Bienvenido |date=May 8, 2013 |title=The Quaternary megafaunal extinction and the fate of Neanderthals: An integrative working hypothesis |url=https://www.sciencedirect.com/science/article/pii/S1040618212001188 |journal=Quaternary International |series=East meets West: First settlements and human evolution in Eurasia |volume=295 |pages=69–72 |doi=10.1016/j.quaint.2012.02.037 |bibcode=2013QuInt.295...69H |issn=1040-6182 |access-date=February 5, 2024 |via=Elsevier Science Direct}} 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.{{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 |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}}{{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.}}{{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}} 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.{{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}} Similar refuge zones have also been proposed on other temperate European peninsulas, namely Italy, the Balkans, and Crimea.{{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}}{{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|pages=27|volume=7|issue=1|doi=10.1007/s41982-024-00194-y|doi-access=free}}

Historically, the cause of extinction of Neanderthals and other archaic humans was viewed under an imperialistic guise, with the superior invading modern humans exterminating and replacing the inferior species.

{{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{{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 |issue=0 |pages=109–118 |doi=10.1101/SQB.1950.015.01.013}}}}

In general, the extinction of Neanderthals is ascribed to predominantly competition with modern humans. The success of modern humans over Neanderthals is usually attributed to a higher birth rate and population, 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.{{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|pages=106331|volume=238|doi=10.1016/j.quascirev.2020.106331|doi-access=free}} Assimilation had long been hypothesised with supposed hybrid specimens, and was revitalised with the discovery of archaic human DNA in modern humans.{{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 |page=e96424 |doi=10.1371/journal.pone.0096424 |pmc=4005592 |pmid=24789039 |bibcode=2014PLoSO...996424V|doi-access=free }} Similarly, the Châtelperronian industry of central France and northern Spain may represent a culture of Neanderthals adopting modern human techniques, via acculturation.{{cite journal|first=I.|last=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=29|doi=10.1007/s41982-024-00193-z|doi-access=free}}{{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 }} Other ambiguous transitional cultures include the Italian Uluzzian industry,{{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 |page=e0196786 |doi=10.1371/journal.pone.0196786 |pmc=5942857 |pmid=29742147 |bibcode=2018PLoSO..1396786V|doi-access=free }} and the Central European Szeletian industry.{{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}}

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.{{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 |at=e0216742 |doi=10.1371/journal.pone.0216742 |pmc=6541251 |pmid=31141515 |bibcode=2019PLoSO..1416742D |doi-access=free}} 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.{{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}} 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.

{{Main|Neanderthals in popular culture}}

File:BlackTerror1636.jpg (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, Björn Kurtén's Dance of the Tiger, and Jean M. Auel's Clan of the Cave Bear and her Earth's Children series.

• {{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}}

• {{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, UK |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 |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 |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 |location =Cambridge, UK |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|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=978-0415674546}}
• {{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 |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=978-0-500-05070-5 |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, UK |isbn= 978-1-4729-3749-0}}
• {{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}}

• {{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|page=105822 |doi=10.1016/j.jas.2023.105822 |bibcode=2023JArSc.159j5822H |s2cid=261325698 |issn= |doi-access=free}}

{{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, 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

|url-status=dead

|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}}

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Category:Fossil taxa described in 1864

Category:Stone Age Asia

Category:Stone Age Europe