Neanderthal anatomy#Distinguishing physical traits

File:Evidence as to Man's Place in Nature page 179.jpg's 1863 illustration of an Aboriginal Australian skull overlain by Neanderthal 1, emphasising their similarity]]

When the first Neanderthal fossil, Neanderthal 1 (a skullcap), was discovered in 1856, initial reactions characterised it as belonging to a brutish, savage race of man. The low braincase and flattened forehead were often cited as evidence of its primitiveness, as classic markers of the lower races, in accord with historical race concepts. Consequently, the Neanderthal skull was often anatomically compared to most notably Aboriginal Australians, who were considered the most primitive race alive.{{cite journal|last=Madison|first=P.|year=2021|title=Brutish Neanderthals: History of a merciless characterization|journal=Evolutionary Anthropology: Issues, News, and Reviews|doi=10.1002/evan.21918}} German pathologist Rudolf Virchow 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. The most influential specimen was La Chapelle-aux-Saints 1 ("The Old Man") from La Chapelle-aux-Saints, France. French palaeontologist Marcellin Boule described him as a hairy, slouching, ape-like creature; a reconstruction which would endure until around the middle of the 20th century. By this point, several fossils from across the Old World were classified as "Neanderthaloid", a type of transitional fossil in human evolution halfway between Homo erectus and Homo sapiens. The modern anatomical definition of "Neanderthal" was formalised with the popularisation of cladistics in the late 1970s.{{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}}

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| image1 = Sapiens neanderthal comparison.jpg

| alt1 = A human skull on the left facing a reconstructed Neanderthal skull on the right, emphasising the difference in braincase shape (more cranial length in Neanderthal), shorter forehead ratio, more defined brow ridge, larger nasal bone projection, pinned-back cheekbone angulation, straighter angled chin, and an occipital bun

| caption1 = Comparisons of a modern Eurasian male example (left) and a Neanderthal (right) skull reconstruction at the Cleveland Museum of Natural History

| image2 = Neanderthal cranial anatomy.jpg

| caption2 = Neanderthal skull features

| alt2 = Front and side view diagram of Neanderthal skull reconstruction emphasising large circular orbits, straightened chin, projecting nasal bridge, large brow ridge, receded forehead, long topped braincase, occipital bun, fossa, and a large gap behind the third molar

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The Neanderthal skull is distinguished namely by 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. The occipital bun, or "chignon", is within the range of variation for modern humans who have it. In Neanderthals, it is caused by the high and anterior (more forward) positions of the cranial base and temporal bones, in combination with a flatter skullcap.{{cite journal |first1=P. |last1=Gunz |first2=K. |last2=Harvati |year=2007 |title=The Neanderthal "chignon": variation, integration, and homology |journal=Journal of Human Evolution |volume=52 |issue=3 |pages=262–274 |doi=10.1016/j.jhevol.2006.08.010 |pmid=17097133|bibcode=2007JHumE..52..262G }}

In Neanderthals, the zygomatic arches (cheekbones) are positioned in a rearward location relative to modern humans, while the maxilla (upper jaw) and nasal bones are positioned in a more forward direction.{{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|p=75}} The front (anterior) teeth are characterised by their large size, strong and bulging tooth roots, and tooth wearing (especially on the lower front teeth). The upper incisors are shovel-shaped. There is a large retromolar space (gap behind the molars). Especially in Europe, Neanderthals had a high frequency of taurodontism, a condition where the molars are bulkier due to an enlarged pulp (tooth core). In modern populations the trait has an incidence rate of about 5%, but a weaker form of taurodontism (hypotaurodontism) is somewhat common in some European H. heidelbergensis populations, especially at the Sima de los Huesos site.{{cite journal |first1=S. |last1=Benazzi |first2=H. H. |last2=Nguyen |first3=O. |last3=Kullmer |first4=J.-J. |last4=Hublin |year=2014 |title=Exploring the biomechanics of taurodontism |journal=Journal of Anatomy |volume=226 |issue=2 |pages=180–188 |doi=10.1111/joa.12260 |pmid=25407030 |pmc=4304574}}

These observations are typically explained as a response to habitual heavy loading of the front teeth (anterior dental loading hypothesis), either to process mechanically challenging or attritive foods, or because they regularly used the mouth as a third hand.{{efn|Neanderthals typically present a similar tooth wearing pattern to Greenland Inuit groups, who use their teeth to soften seal hide among other tasks.}} The robusticity of the front teeth and the rest of the face has sometimes been ascribed to the production of a high bite force on the front teeth, but biomechanical studies have typically concluded that Neanderthals could produce similar or even smaller bite forces than modern humans.{{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 }}{{cite journal |first1=C. F. |last1=O'Connor |first2=R. G. |last2=Franciscus |first3=N. E. |last3=Holton |year=2005 |title=Bite force production capability and efficiency in Neandertals and modern humans |journal=American Journal of Physical Anthropology |volume=127 |issue=2 |pages=129–151 |doi=10.1002/ajpa.20025 |pmid=15558614}}

File:Spy Skull.jpg, Belgium]]

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}}{{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}} which is significantly larger than the averages for all groups of recent humans;{{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}} for example, recent European males can average about {{cvt|1,350|cm3}} and females {{cvt|1,200|cm3}}.{{cite journal |first1=J. S. |last1=Allen |first2=H. |last2=Damasio |first3=T. J. |last3=Grabowski |year=2002 |title=Normal neuroanatomical variation in the human brain: an MRI-volumetric study |journal=American Journal of Physical Anthropology |volume=118 |issue=4 |pages=341–358 |doi=10.1002/ajpa.10092 |pmid=12124914 |s2cid=21705705 }} Modern human brain size seems to have decreased since the Upper Palaeolithic, with a sample of 28 modern human specimens from 190,000 to 25,000 years ago averaging about {{cvt|1,478|cm3}} disregarding sex.{{cite journal |first1=A. |last1=Balzeau |first2=D. |last2=Grimaud-Hervé |first3=F. |last3=Detroit |first4=R. L. |last4=Holloway |author4-link=Ralph Holloway |year=2013 |title=First description of the Cro-Magnon 1 endocast and study of brain variation and evolution in anatomically modern Homo sapiens |journal=Bulletins et Mémoires de la Société d'Anthropologie de Paris |volume=25 |issue=1–2 |pages=11–12 |doi=10.1007/s13219-012-0069-z |s2cid=14675512 |url=https://www.researchgate.net/publication/257799835}} The largest Neanderthal brain, Amud 1, was calculated to be {{cvt|1,736|cm3}}, one of the largest ever recorded in humans. Both Neanderthal and modern human infants measure about {{cvt|400|cm3}}.{{cite journal |first1=M. S. |last1=Ponce de León |first2=L. |last2=Golovanova |first3=V. |last3=Doronichev |year=2008 |title=Neanderthal brain size at birth provides insights into the evolution of human life history |journal=Proceedings of the National Academy of Sciences |volume=105 |issue=37 |pages=13764–13768 |doi=10.1073/pnas.0803917105|pmid=18779579|pmc=2533682|doi-access=free}}

The Neanderthal brain had different growth and development rates than modern humans, especially in the orbitofrontal cortex (associated with decision making), parietal and temporal lobes (language processing and memory), and the cerebellum (motor functions). All of these regions are proportionally smaller in Neanderthals, and diverge in growth pattern at what would be a critical period in modern human neurological development.{{cite book |last1=Bruner |first1=Emiliano |last2=Ogihara |first2=Naomichi |last3=Tanabe |first3=Hiroki C. |title=Digital Endocasts: From Skulls to Brains |date=December 28, 2017 |publisher=Springer |isbn=978-4-431-56582-6 |page=25 |url=https://books.google.com/books?id=Z9FEDwAAQBAJ&pg=PA25 |language=en}}{{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 }} Altogether, such differences, while slight, may underlie the differences in Cro-Magnon and Neanderthal behaviour including sociality, technological innovation, and artistic output in the archaeological record.{{cite book |last1=Coolidge |first1=Frederick L. |last2=Wynn |first2=Thomas |last3=Overmann |first3=Karenleigh A. |chapter=The Expert Neandertal Mind and Brain, Revisited |editor1-last=Wynn |editor1-first=Thomas |editor2-last=Overmann |editor2-first=Karenleigh A. |editor3-last=Coolidge |editor3-first=Frederick L. |title=The Oxford Handbook of Cognitive Archaeology |isbn=9780192895950 |pages=131–149 |publisher=Oxford University Press |year=2024}}

File:Drachenlochmuseum Neandertaler2.jpeg]]

The neck vertebrae of Neanderthals are thicker from the front to the rear and transversely than those of (most) modern humans. This may have improved stability of the larger head.{{cite journal |first1=A. |last1=Gómez-Olivencia |first2=E. |last2=Been |first3=J. L. |last3=Arsuaga |first4=J. T. |last4=Stock |year=2013 |title=The Neandertal vertebral column 1: the cervical spine |journal=Journal of Human Evolution |volume=64 |issue=6 |pages=604–630 |doi=10.1016/j.jhevol.2013.02.008 |pmid=23541382|bibcode=2013JHumE..64..608G }}

The Neanderthal chest was deep and wide, with a proportionally expansive thoracic cavity, and possibly better lung performance. This was caused by longer, straighter ribs, and more dorsally (towards the head) orientated transverse processes on the vertebrae (where the rib connects to the spine). The latter caused the vertebrae to protrude farther into the ribcage (invagination).{{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}} To support the wider lower thorax, the sacrum (where the pelvis connects to the spine) was more vertically inclined, and the pelvis was wider.{{cite journal |first1=A. |last1=Gómez-Olivencia |first2=A. |last2=Barash |first3=D. |last3=García-Martínez |display-authors=et al. |year=2018 |title=3D virtual reconstruction of the Kebara 2 Neandertal thorax |journal=Nature Communications |volume=9 |issue=4387 |pages=4387 |doi=10.1038/s41467-018-06803-z |pmc=6207772 |pmid=30377294 |bibcode=2018NatCo...9.4387G}} This also caused the lumbar vertebrae (lower spine) to be less curved (hypolordosis, "flatback"), except for possibly the 5th lumbar vertebra.{{cite journal|first=D.|last=García-Martínez|first2=S.|last2=Martelli|first3=N.|last3=Torres-Tamayo|first4=J. M.|last4=Jiménez-Arenas|first5=A. |last5=González Martín|first6=M.|last6=Campo|first7=O.|last7=Cambra-Moo|first8=S.|last8=Lois-Zlolniski|first9=S.|last9=Nalla|first10=J. A. S.|last10=Gimeno|first11=M.|last11=Bastir|year=2020|title=Sexual dimorphism in the vertebral wedging of the human lumbar vertebrae and its importance as a comparative framework for understanding the wedging pattern of Neanderthals|journal=Quaternary International|volume=566–567|pages=224–232|doi=10.1016/j.quaint.2020.05.054|hdl=10486/715654|hdl-access=free}} Though the skeleton of ancient Homo is poorly documented, because H. ergaster specimens over 1 million years old (namely Turkana Boy) seem to have had a similar physiology, it is possible a wide thorax is normal for Homo. The narrower thorax of modern humans may be a unique adaptation associated with the adoption of endurance running.{{cite journal|first=M.|last=Bastir|first2=D.|last2=García-Martínez|first3=N.|last3=Torres-Tamayo|first4=C. A.|last4=Palancar|first5=B.|last5=Beyer|first6=A.|last6=Barash|first7=C.|last7=Villa|first8=J. A.|last8=Sanchis-Gimeno|first9=A.|last9=Riesco-López|first10=S.|last10=Nalla|first11=I.|last11=Torres-Sánchez|first12=F.|last12=García-Río|first13=E.|last13=Been|first14=A.|last14=Gómez-Olivencia|first15=M.|last15=Haeusler|first16=S. A.|last16=Williams|first17=F.|last17=Spoor|year=2020|title=Rib cage anatomy in Homo erectus suggests a recent evolutionary origin of modern human body shape|journal=Nature Ecology & Evolution|volume=4|pages=1178–1187|doi=10.1038/s41559-020-1240-4}}

The limbs are proportionally short, which has traditionally been explained as a "hyper-arctic" adaptation (Allen's rule). A similar trend is also observed in modern Inuit and Siberian Yupik populations.{{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 }}{{cite journal |first=I. |last=De Groote |year=2011 |title=The Neanderthal lower arm |journal=Journal of Human Evolution |volume=61 |issue=4 |pages=396–410 |doi=10.1016/j.jhevol.2011.05.007 |pmid=21762953|bibcode=2011JHumE..61..396D }} Neanderthals in more temperate climates—such as Iberia—still retain the "hyperarctic" physique.{{cite journal |first1=M. J. |last1=Walker |first2=J. |last2=Ortega |first3=K. |last3=Parmová |first4=M. V. |last4=López |first5=E. |last5=Trinkaus |author-link5=Erik Trinkaus |year=2011 |title=Morphology, body proportions, and postcranial hypertrophy of a Neandertal female from the Sima de las Palomas, southeastern Spain |journal=Proceedings of the National Academy of Sciences |volume=108 |issue=25 |pages=10,087–10,091 |doi=10.1073/pnas.1107318108 |pmc=3121844 |pmid=21646528 |bibcode=2011PNAS..10810087W|doi-access=free }} The shorter limbs, combined with evidence of a stronger respiratory system and a faster metabolism fueling more fast-twitch muscle fibres, could altogether also be explained as adaptations for sprinting. Neanderthals were probably not as adept at endurance running as modern humans given the expanded thorax and long heel bones (causing a long moment arm at the Achilles' tendon, possibly more conducive for acceleration than distance).{{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}}{{cite journal |first1=D. |last1=Raichlen |first2=H. |last2=Armstrong |first3=D. E. |last3=Lieberman |year=2011 |title=Calcaneus length determines running economy: Implications for endurance running performance in modern humans and Neandertals |journal=Journal of Human Evolution |volume=60 |issue=3 |pages=299–308 |doi=10.1016/j.jhevol.2010.11.002 |pmid=21269660|bibcode=2011JHumE..60..299R }}

Neanderthals had proportionately shorter thumbs and index fingers, but this does not seem to have negatively impacted dexterity. Bone trauma indicates Neanderthals habitually made use of a strong, squeezing grip.{{cite journal|first=A.|last=Bardo|first2=M.-H.|last2=Moncel|first3=C. J.|last3=Dunmore|first4=T. L.|last4=Kivell|first5=E.|last5=Pouydebat|first6=R.|last6=Cornette|year=2020|title=The implications of thumb movements for Neanderthal and modern human manipulation|journal=Scientific Reports|volume=10|issue=19323|doi=10.1038/s41598-020-75694-2|hdl=21.11116/0000-0007-A185-C|hdl-access=free}} Like modern humans, Neanderthals seem to have exhibited handedness, generally preferring the right hand.{{cite book|last=Uomini|first=N. T.|year=2011|chapter=Handedness in Neanderthals|title=Neanderthal Lifeways, Subsistence and Technology|series=Vertebrate Paleobiology and Paleoanthropology|publisher=Springer, Dordrecht|doi=10.1007/978-94-007-0415-2_14|isbn=978-94-007-3525-5}}

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 }}

File:Neanderthal and homo sapiens sapiens.jpg]]

For comparison, the average height of 20 male and 10 female Cro-Magnons is, respectively, {{cvt|176.2|cm|ftin|0}} and {{cvt|162.9|cm|ftin|0}}, although this decreases by {{cvt|10|cm|0}} nearer the end of the Upper Palaeolithic based on 21 males and 15 females.{{cite journal |first1=V. |last1=Formicola |first2=M. |last2=Giannecchini |year=1998 |title=Evolutionary trends of stature in Upper Paleolithic and Mesolithic Europe |journal=Journal of Human Evolution |volume=36 |issue=3 |pages=325 |pmid=10074386 |doi=10.1006/jhev.1998.0270}} The average height in the year 1900 was {{cvt|163|cm|ftin|0}} and {{cvt|152.7|cm|ftin|0}}, respectively.{{cite journal |first1=M. |last1=Roser |author1-link=Max Roser |first2=C. |last2=Appel |first3=H. |last3=Ritchie |author3-link=Hannah Ritchie |year=2013 |title=Human height |journal=Our World in Data |url=https://ourworldindata.org/human-height |access-date=June 16, 2020}}

For Neanderthal weight, a sample of 26 specimens found an average of {{cvt|77.6|kg|lbs}} for males and {{cvt|66.4|kg|lbs}} for females.{{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}} Using {{cvt|76|kg|lbs}}, the body mass index for Neanderthal males was calculated to be 26.9–28.3, which in modern humans correlates to being overweight. This indicates a very robust build.{{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}} The Neanderthal LEPR gene concerned with storing fat and body heat production is similar to that of the woolly mammoth, and so was likely an adaptation for cold climate.{{cite journal |first1=M. |last1=Kislev |first2=R. |last2=Barkai |year=2018 |title=Neanderthal and woolly mammoth molecular resemblance |journal=Human Biology |volume=90 |issue=2 |pages=115–128 |doi=10.13110/humanbiology.90.2.03 |pmid=33951886 |s2cid=106401104}}

The orbits (eye sockets) of Neanderthals, and possibly also the eyeballs, were bigger than those of Cro-Magnons and modern humans. It is unclear how this could relate to vision. In modern humans, eye socket size does not correlate with eyeball size; their volumes instead seem to be most influenced by the frontal lobe pushing into the area. In Neanderthals, the separation of the orbits from the frontal lobe is about midway what is seen in H. heidelbergensis and modern humans.{{cite journal|first=A. S.|last=Pereira-Pedro|first2=M.|last2=Masters|first3=E.|last3=Bruner|year=2017|title=Shape analysis of spatial relationships between orbito-ocular and endocranial structures in modern humans and fossil hominids|journal=Journal of Anatomy|volume=231|issue=6|pages=947–960|doi=10.1111/joa.12693|doi-access=free|pmc=5696126}} Neurologically, Neanderthals had a proportionally large occipital lobe compared to Cro-Magnons, associated with visual processing.{{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}}{{cite journal |first1=T. |last1=Kochiyama |first2=N. |last2=Ogihara |first3=H. C. |last3=Tanabe |year=2018 |title=Reconstructing the Neanderthal brain using computational anatomy |journal=Scientific Reports |volume=8 |issue=6296 |pages=6296 |doi=10.1038/s41598-018-24331-0 |pmc=5919901 |pmid=29700382 |bibcode=2018NatSR...8.6296K}} Genetically, colour blindness (which may enhance mesopic vision in low-light conditions) is typically correlated with high-latitude populations, and the Neanderthals from Vindija Cave, Croatia, had some substitutions in the Opsin genes which could have influenced colour vision.

The functional implications of these traits are inconclusive, but altogether, they could indicate a preference for activity in dimmer light conditions. This may have been advantageous in northerly latitudes where daylight hours are much shorter especially in winter, and low-light or nighttime hunting would enhance ambush tactics when pursuing large game.{{cite journal |first1=J. S. |last1=Taylor |first2=T. E. |last2=Reimchen |year=2016 |title=Opsin gene repertoires in northern archaic hominids |journal=Genome |volume=59 |issue=8 |pages=541–549 |doi=10.1139/gen-2015-0164 |pmid=27463216 |hdl=1807/73317 |hdl-access=free}} Neanderthal-derived alleles near ASB1 and EXOC6 are associated with being an evening person, narcolepsy and day-time napping.{{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=578–589 |doi=10.1016/j.ajhg.2017.09.010 |pmid=28985494 |pmc=5630192}}

File:Homo sapiens neanderthalensis.jpg displaying a large piriform aperture (nose hole)]]

The Neanderthal piriform aperture (nose hole) and nasal cavity are much wider and bigger than in modern humans. Assuming the nasal passages took up the entire nasal cavity, this has normally been explained as an adaptation to warm greater quantities of cold air to fuel their assumed heightened metabolism and activity levels compared to modern humans.{{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 }}

In modern humans, high-latitude populations typically instead have taller, narrower piriform apertures and noses, which decrease air intake per breath, but improve turbulence within the nose to moisten and warm air. Since Neanderthals had a much longer skull and face, and consequently probably longer nasal passages, air may have been warmed enough after simply crossing such a distance to reach the lungs, not requiring the additional nasal turbulence.{{cite journal|first=M.|last=Bastir|year=2019|title=Big choanae, larger face: Scaling patterns between cranial airways in modern humans and African apes and their significance in Middle and Late Pleistocene hominin facial evolution|journal=Bulletins et Mémoires de la Société d'Anthropologie de Paris|volume=31|pages=5–13|doi=10.3166/bmsap-2019-0055|doi-access=free|issn=1777-5469|hdl=10261/223991|hdl-access=free}}

The longer, straighter ribs and implied wider mid-lower thorax (where the ribcage is) similarly indicate stronger breathing in the lower thorax, a larger diaphragm, and possibly greater average lung capacity.{{cite journal |first1=D. |last1=García-Martínez |first2=M. |last2=Bastir |first3=R. |last3=Huguet |year=2017 |title=The costal remains of the El Sidrón Neanderthal site (Asturias, northern Spain) and their importance for understanding Neanderthal thorax morphology |journal=Journal of Human Evolution |volume=111 |pages=85–101 |doi=10.1016/j.jhevol.2017.06.003 |pmid=28874276 |bibcode=2017JHumE.111...85G |hdl=10261/158977 |s2cid=4634345 |hdl-access=free}}{{cite journal |first1=A. |last1=Gómez-Olivencia |first2=T. |last2=Holliday |first3=M. |last3=Madelaine |year=2018 |title=The costal skeleton of the Regourdou 1 Neandertal |journal=Journal of Human Evolution |volume=130 |pages=151–171 |doi=10.1016/j.jhevol.2017.12.005 |pmid=29496322 |doi-access=free}} The lung capacity of Kebara 2 was estimated to have been {{cvt|9.04|L|gal}}, compared to the average human capacity of {{cvt|6|L|gal}} for males and {{cvt|4.7|L|gal}} for females.

{{See also|Origin of speech}}

In 1971, cognitive scientist Philip Lieberman attempted to reconstruct the Neanderthal vocal tract and concluded that it was similar to that of a modern human newborn. His reconstruction of the Neanderthal vocal tract left Neanderthals incapable of producing a range of speech sounds, due to the large size of the mouth and the small size of the pharyngeal cavity, thus no need for a descended larynx{{efn|The lack of a descended larynx does not necessarily equate to a reduced vowel capacity.{{cite journal |doi=10.1006/jpho.2002.0170 |title=The potential Neandertal vowel space was as large as that of modern humans |year=2002 |last1=Boë |first1=L.-J. |last2=Heim |first2=J.-L. |last3=Honda |first3=K. |last4=Maeda |first4=S. |journal=Journal of Phonetics |volume=30 |issue=3 |pages=465–484 |s2cid=16685699 }}}} to fit the entire tongue inside the mouth. He concluded that they were anatomically unable to produce the sounds /a/, /i/, /u/, /ɔ/, /g/, and /k/ and thus lacked the capacity for articulate speech, though were still able to speak at a level higher than non-human primates.{{cite journal |first1=P. |last1=Lieberman |author-link=Philip Lieberman |first2=E. S. |last2=Crelin |year=1971 |title=On the speech of Neanderthal man |journal=Linguistic Inquiry |volume=2 |issue=2 |pages=203–222 |doi=10.1515/9783110819434-007 |jstor=4177625}}{{cite journal |last=Lieberman |first=P. |author-link=Philip Lieberman |year=1992 |title=On Neanderthal speech and Neanderthal extinction |journal=Current Anthropology |volume=33 |issue=4 |pages=409–410 |doi=10.1086/204092 |s2cid=144005974}}{{Cite journal|doi=10.1016/j.wocn.2005.07.002 |title=Current views on Neanderthal speech capabilities: A reply to Boe et al. (2002) |year= 2007 |first=P.|last=Lieberman |author-link=Philip Lieberman |journal=Journal of Phonetics |volume=35 |issue=4 |pages=552–563}}

Lieberman's findings have subsequently been debated, especially in light of newer fossil discoveries and re-analyses. The 1983 discovery of a Neanderthal hyoid bone—used in speech production in humans—in Kebara 2 is almost identical to that of humans, which could suggest Neanderthals were capable of speech. Still, it is not possible to accurately reconstruct the entire vocal tract with just the hyoid.{{cite journal|author1=J.T. Laitman|author2=B. Johansson|year=1990|title=The Kebara hyoid: What can it tell us about the evolution of the hominid vocal tract|journal=American Journal of Physical Anthropology|volume=81|number=2|page=254|doi=10.1002/ajpa.1330810204}}{{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}} The Sima de los Huesos fossils, which may be a "pre-Neanderthal" population, also had humanlike hyoid and ear bones.{{cite journal |first1=R. |last1=D’Anastasio |first2=S. |last2=Wroe |first3=C. |last3=Tuniz |first4=L. |last4=Mancini |first5=D. T. |last5=Cesana |year=2013 |title=Micro-biomechanics of the Kebara 2 hyoid and its implications for speech in Neanderthals |journal=PLOS ONE |volume=8 |issue=12 |page=e82261 |doi=10.1371/journal.pone.0082261 |pmc=3867335 |pmid=24367509 |bibcode=2013PLoSO...882261D|doi-access=free }} Some studies can reconstruct the Neanderthal vocal apparatus as comparable to that of modern humans, but soft-tissue reconstructions are problematic in general.{{cite journal |first1=L.J. |last1=Boë |first2=J.L. |last2=Heim |first3=K. |last3=Honda |first4=S. |last4=Maeda |first5=P. |last5=Badin |first6=C. |last6=Abry |year=2007 |title=The vocal tract of newborn humans and Neanderthals: Acoustic capabilities and consequences for the debate on the origin of language. A reply to Lieberman |journal= Journal of Phonetics |volume=35 |issue=4 |pages=564–581 |doi=10.1016/j.wocn.2007.06.006}}

File:Homo sapiens neanderthalensis (Fundort Gibraltar).jpg]]

The lack of sunlight may have led to the proliferation of lighter skin in Neanderthals. Genetically, Neanderthals could carry two different variations of BNC2, which in modern populations are associated with lighter or darker skin colour in the UK Biobank. DNA analysis of three Neanderthal females from southeastern Europe indicates that they had brown eyes and dark skin colour.{{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}}

In modern humans, skin and hair colour is regulated by the melanocyte-stimulating hormone—which increases the proportion of eumelanin (black pigment) to phaeomelanin (red pigment)—which is encoded by the MC1R gene. There are five known variants in modern humans of the gene which cause loss-of-function and are associated with light skin and hair colour. None of the variants associated with red hair have been found in Neanderthals, but they carried an additional variant, R307G, which could be associated with pale skin and red hair. The R307G variant was identified in a Neanderthal from Monti Lessini, Italy, and possibly Cueva del Sidrón, Spain.{{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 }}

Generally, models on Neanderthal caloric requirements report significantly higher intakes than those of modern humans. They typically assume that Neanderthals had higher basal metabolic rates (BMRs) due to higher muscle mass, faster growth rate, and greater body heat production against the cold.{{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}} Higher daily physical activity levels (PALs) are also usually assumed, caused by less efficient foraging techniques than Cro-Magnons, requiring Neanderthals to travel farther and expend more energy daily.{{cite journal |first1=J. J. |last1=Snodgrass |first2=W. R. |last2=Leonard |year=2009 |title=Neandertal energetics revisited: insights into population dynamics and life history evolution |journal=PaleoAnthropology |doi=10.4207/PA.2009.ART31 |url=https://pdfs.semanticscholar.org/f1f6/789a820d1e5dc20e9f7eed18b46900177829.pdf |archive-url=https://web.archive.org/web/20200801154540/https://pdfs.semanticscholar.org/f1f6/789a820d1e5dc20e9f7eed18b46900177829.pdf |url-status=dead |archive-date=August 1, 2020 |pages=220–237 |s2cid=16044193}}{{cite journal |first=B. |last=Hockett |year=2012 |title=The consequences of Middle Paleolithic diets on pregnant Neanderthal women |journal=Quaternary International |volume=264 |pages=78–82 |doi=10.1016/j.quaint.2011.07.002 |bibcode=2012QuInt.264...78H}}

File:Moulage pech de l'azé MHN 1502680.jpg]]

Maximum natural lifespan and the timing of adulthood, menopause, and gestation were most likely similar to modern humans.{{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}}

Based on the growth rates of teeth and tooth enamel, Neanderthals may have matured faster than modern humans.{{cite journal |first1=T. M. |last1=Smith |first2=P. |last2=Tafforeau |first3=D. J. |last3=Reid |year=2010 |title=Dental evidence for ontogenetic differences between modern humans and Neanderthals |journal=Proceedings of the National Academy of Sciences |volume=107 |issue=49 |pages=20923–20928 |doi=10.1073/pnas.1010906107 |pmc=3000267 |pmid=21078988 |bibcode=2010PNAS..10720923S|doi-access=free }}{{cite journal |first=D. |last=Guatelli-Steinberg |year=2009 |title=Recent studies of dental development in Neandertals: Implications for Neandertal life histories |journal=Evolutionary Biology |volume=18 |issue=1 |pages=9–20 |doi=10.1002/evan.20190 |s2cid=84273699 |doi-access=free}} The enamel is thinner in Neanderthals, which may have stemmed from a lower long-period line periodicity and a faster extension rate, which resulted in faster tooth crown formation times.{{cite journal | vauthors = Smith TM, Tafforeau P, Reid DJ, Pouech J, Lazzari V, Zermeno JP, Guatelli-Steinberg D, Olejniczak AJ, Hoffman A, Radovcic J, Makaremi M, Toussaint M, Stringer C, Hublin JJ | display-authors = 6 | title = Dental evidence for ontogenetic differences between modern humans and Neanderthals | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 107 | issue = 49 | pages = 20923–20928 | date = December 2010 | pmid = 21078988 | pmc = 3000267 | doi = 10.1073/pnas.1010906107 | bibcode = 2010PNAS..10720923S | doi-access = free }} Skeletally, the main differences in maturation are the atlas bone in the neck as well as the middle thoracic vertebrae fused about 2 years later in Neanderthals than in modern humans, but this was more likely caused by a difference in anatomy rather than growth rate.{{cite journal |first1=A. |last1=Rosas |first2=L. |last2=Ríos |first3=A. |last3=Estalrrich |display-authors=et al. |year=2017 |title=The growth pattern of Neandertals, reconstructed from a juvenile skeleton from El Sidrón (Spain) |journal=Science |volume=357 |issue=6537 |pages=1282–1287 |doi=10.1126/science.aan6463 |pmid=28935804 |bibcode=2017Sci...357.1282R |doi-access=free}}{{cite journal |first=J. M. |last=DeSilva |year=2018 |title=Comment on 'The growth pattern of Neandertals, reconstructed from a juvenile skeleton from El Sidrón (Spain)' |journal=Science |volume=359 |issue=6380 |pages=eaar3611 |doi=10.1126/science.aar3611 |pmid=29590012 |doi-access=free}}

In a sample of 669 Neanderthal tooth crowns, 75% suffered some degree of enamel hypoplasia, an indicator of developmental stress perhaps caused by recurrent starvation at a young age while the teeth were forming. Comparing the rate to what is seen in recent Tikiġaġmiut, Neanderthals experienced stresses lasting from two weeks to up to three months.{{cite journal | vauthors = Guatelli-Steinberg D, Larsen CS, Hutchinson DL | title = Prevalence and the duration of linear enamel hypoplasia: a comparative study of Neandertals and Inuit foragers | journal = Journal of Human Evolution | volume = 47 | issue = 1–2 | pages = 65–84 | year = 2004 | pmid = 15288524 | doi = 10.1016/j.jhevol.2004.05.004 | bibcode = 2004JHumE..47...65G }}{{cite journal | vauthors = Barrett CK, Guatelli-Steinberg D, Sciulli PW | title = Revisiting dental fluctuating asymmetry in neandertals and modern humans | journal = American Journal of Physical Anthropology | volume = 149 | issue = 2 | pages = 193–204 | date = October 2012 | pmid = 22791408 | doi = 10.1002/ajpa.22107 }}

File:Shanidar I skull and skeleton, c. 60,000 to 45,00o BCE. Iraq Museum.jpg]]

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 }}

In 1995, Trinkaus estimated that about 80% succumbed to their injuries and died before reaching 40, and thus theorised that Neanderthals employed a risky hunting strategy ("rodeo rider" hypothesis).{{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 }} Compared to contemporary modern humans, the rates of cranial trauma are not significantly different (although Neanderthals seem to have had a higher mortality risk),{{cite journal |last1=Beier |first1=J. |last2=Anthes |first2=N. |last3=Wahl |first3=J. |last4=Harvati |first4=K. |title=Similar cranial trauma prevalence among Neanderthals and Upper Palaeolithic modern humans |journal=Nature |year=2018 |volume=563 |issue=7733 |pages=686–690 |doi=10.1038/s41586-018-0696-8 |pmid=30429606 |bibcode=2018Natur.563..686B |s2cid=53306963 |url=https://zenodo.org/record/3569613}} there are few specimens of both Cro-Magnons and Neanderthals who died after the age of 40,{{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 }} and there are overall similar injury patterns between them. In 2012, Trinkaus concluded that Neanderthals may have additionally been injuring themselves in the same way as contemporary humans, such as by interpersonal violence.{{cite journal |first=E. |last=Trinkaus |author-link=Erik Trinkaus |year=2012 |title=Neandertals, early modern humans, and rodeo riders |journal=Journal of Archaeology |volume=39 |issue=2 |pages=3691–3693 |doi=10.1016/j.jas.2012.05.039|bibcode=2012JArSc..39.3691T }}

A 2016 study looking at 124 Neanderthal specimens argued that high trauma rates were instead caused by animal attacks, and found that about 36% of the sample were victims of bear attacks, 21% big cat attacks, and 17% wolf attacks (totalling 92 positive cases, 74%). There were no cases of hyena attacks, although hyenas still nonetheless probably attacked Neanderthals, at least opportunistically. Such intense predation probably stemmed from common confrontations due to competition over food and cave space, and from Neanderthals hunting these carnivores.{{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}}

Some examples of Neanderthal congenital disorders include:

• The 13 inhabitants of Sidrón Cave collectively exhibited 17 different birth defects likely due to inbreeding or recessive disorders{{efn|They exhibited: narrowing of the nasal cavity, retained deciduous canine, clefts of the atlas, hypoplasia of the axis, clefting of the twelfth thoracic vertebra, either a small thoracic rib or a lumbar rib, os centrale carpi and bipartite scaphoid bone, tripartite kneecap, abnormally-shaped tarsal bones, and cuboid-navicular coalition.}}{{cite journal |first1=L. |last1=Ríos |first2=T. L. |last2=Kivell |first3=C. |last3=Lalueza-Fox |first4=A. |last4=Estalrrich |year=2019 |title=Skeletal anomalies in the Neandertal family of El Sidrón (Spain) support a role of inbreeding in Neandertal extinction |journal=Scientific Reports |volume=9 |issue=1 |page=1697 |doi=10.1038/s41598-019-38571-1 |pmc=6368597 |pmid=30737446 |bibcode=2019NatSR...9.1697R}}
• Likely due to advanced age (60s or 70s), La Chapelle-aux-Saints 1 had signs of Baastrup's disease, affecting the spine, and osteoarthritis{{cite journal |first1=M. |last1=Haeusler |first2=E. |last2=Trinkaus |author2-link=Erik Trinkaus |first3=C. |last3=Fornai |first4=J. |last4=Müller |first5=N. |last5=Bonneau |first6=T. |last6=Boeni |first7=N. |last7=Frater |year=2019 |title=Morphology, pathology, and the vertebral posture of the La Chapelle-aux-Saints Neandertal |journal=Proceedings of the National Academy of Sciences |volume=116 |issue=11 |pages=4923–4927 |doi=10.1073/pnas.1820745116 |pmid=30804177 |pmc=6421410|bibcode=2019PNAS..116.4923H |doi-access=free }}
• Shanidar 1, who likely died at about 30 or 40, was diagnosed with the most ancient case of diffuse idiopathic skeletal hyperostosis (DISH), a degenerative disease which can restrict movement, which, if correct, would indicate a moderately high incidence rate for older Neanderthals{{cite journal |first1=E. |last1=Crubézy |first2=E. |last2=Trinkaus |author2-link=Erik Trinkaus |year=1992 |title=Shanidar 1: a case of hyperostotic disease (DISH) in the middle Paleolithic |journal=American Journal of Physical Anthropology |volume=89 |issue=4 |pages=411–420 |doi=10.1002/ajpa.1330890402 |pmid=1463085}}

File:La Ferrassie 1 MdlH 1 2018-10-20.jpg (above) may have had periostitis in his femora and tibiae.|alt=A mostly complete skeleton laid out against a black background horizontally]]

Neanderthals were subject to several infectious diseases and parasites. Modern humans likely transmitted diseases to them; one possible candidate is the stomach bacteria Helicobacter pylori.{{cite journal |first1=C. J. |last1=Houldcroft |first2=S. J. |last2=Underdown |year=2016 |title=Neanderthal genomics suggests a pleistocene time frame for the first epidemiologic transition |journal=American Journal of Physical Anthropology |volume=160 |issue=3 |pages=379–388 |doi=10.1002/ajpa.22985 |pmid=27063929 |url=http://discovery.ucl.ac.uk/1476201/3/Houldcroft_Neanderthal_disease_AJPA-2015-00253.R1_100216.pdf}} The modern human papillomavirus variant 16A may descend from Neanderthal introgression.{{cite journal |first1=V. N. |last1=Pimenoff |first2=C. M. |last2=de Oliveira |first3=I. G. |last3=Bravo |year=2017 |title=Transmission between archaic and modern human ancestors during the evolution of the oncogenic human papillomavirus 16 |journal=Molecular Biology and Evolution |volume=34 |issue=1 |pages=4–19 |doi=10.1093/molbev/msw214 |pmid=28025273 |pmc=5854117}} A Neanderthal at Cueva del Sidrón, Spain, shows evidence of a gastrointestinal Enterocytozoon bieneusi infection.{{cite journal |title=Neanderthal behaviour, diet, and disease inferred from ancient DNA in dental calculus |volume=544 |issue=7650 |pages=357–361 |journal=Nature |doi=10.1038/nature21674 |pmid=28273061 |year=2017 |last1=Weyrich |first1=L. S. |last2=Duchene |first2=S. |last3=Soubrier |first3=J.|display-authors=et al. |bibcode=2017Natur.544..357W |hdl=10261/152016 |s2cid=4457717 |url=https://radar.brookes.ac.uk/radar/items/d6689cb0-38fa-4d3e-98e4-c8be2bd0263e/1}} The leg bones of the French La Ferrassie 1 feature lesions that are consistent with periostitis—inflammation of the tissue enveloping the bone—likely a result of hypertrophic osteoarthropathy, which is primarily caused by a chest infection or lung cancer.{{cite journal |first1=K. J. |last1=Fennel |first2=E. |last2=Trinkaus |author-link2=Erik Trinkaus |year=1997 |title=Bilateral femoral and tibial periostitis in the La Ferrassie 1 Neanderthal |journal=Journal of Archaeological Science |volume=24 |issue=11 |pages=985–995 |doi=10.1006/jasc.1996.0176|bibcode=1997JArSc..24..985F }} La Chapelle-aux-Saints 1 has lesions down the length of his spine consistent with brucellosis, probably infected with Brucella abortus while butchering a carcass or eating raw meat.{{cite journal|first=B.|last=Rothschild|first2=M.|last2=Haeusler|year=2021|title=Possible vertebral brucellosis infection in a Neanderthal|journal=Scientific Reports|volume=11|issue=19846|doi=10.1038/s41598-021-99289-7|pmc=8494896}}

Neanderthals had a lower cavity rate than modern humans, despite some populations consuming typically cavity-causing foods in great quantity. This could indicate a lack of cavity-causing oral bacteria, namely Streptococcus mutans.{{cite journal |first=A. |last=Soltysiak |year=2012 |title=Comment: low dental caries rate in Neandertals: the result of diet or the oral flora composition? |journal=Homo |volume=63 |issue=2 |pages=110–113 |doi=10.1016/j.jchb.2012.02.001 |pmid=22409830}}

In Neanderthals, the Eustachian tubes (which connect the middle ear to the throat) are flat. In modern humans, as an infant grows, the Eustachian tubes become angled to improve drainage of the middle ear and prevent bacterial infection. The flatness of the Eustachian tubes throughout life may have made Neanderthals more prone to developing ear infections.{{cite journal | vauthors = Pagano AS, Márquez S, Laitman JT | title = Reconstructing the Neanderthal Eustachian Tube: New Insights on Disease Susceptibility, Fitness Cost, and Extinction | journal = Anatomical Record | volume = 302 | issue = 12 | pages = 2109–2125 | date = December 2019 | pmid = 31472033 | doi = 10.1002/ar.24248 | s2cid = 201731755 | doi-access = free }} A 2019 study found that 48% of their 77 Neanderthal skull sample size presented bony growths consistent with swimmer's ear (inflammation of the ear canal).

Two 250,000-year-old Neanderthaloid children from Payré, France, present the earliest known cases of lead exposure. They were exposed on two distinct occasions either by eating or drinking contaminated food or water, or inhaling lead-laced smoke from a fire. There are two lead mines within {{cvt|25|km|mi}} of the site.{{cite journal |first1=T. A. |last1=Smith |first2=C. |last2=Austin |first3=D. R. |last3=Green |display-authors=et al. |year=2018 |title=Wintertime stress, nursing, and lead exposure in Neanderthal children |journal=Science Advances |volume=4 |issue=10 |page=eaau9483 |doi=10.1126/sciadv.aau9483 |pmid=30402544 |pmc=6209393 |bibcode=2018SciA....4.9483S}}

• Human anatomy
• Human biology
• Neanderthal behavior
• Neanderthal genetics

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• {{Commons category inline|Homo neanderthalensis}}
• {{Wikibooks inline|Introduction to Paleoanthropology}}
• {{wikispecies inline|Homo neanderthalensis}}

{{Human Evolution}}

{{Anatomy}}

{{Homo neanderthalensis|state=expanded}}

{{DEFAULTSORT:Neanderthal Anatomy}}

Anatomy

Category:Ancient DNA (human)

Category:Human anatomy