Neanderthals (/niˈændərtɑːl, n-, -θɔːl/;[7] or Neandertals, German: Neandertaler [neˈ(ʔ)andɐtaːlɐ];[8] Homo neanderthalensis or Homo sapiens neanderthalensis),[9] are an extinct species or subspecies of archaic humans who lived in Eurasia until about 40,000 years ago (40 kya [thousand years ago]).[10][11][12][13] They probably went extinct due to competition with or extermination by immigrating modern humans[14][15][16] or due to great climatic change,[17][18][19] disease,[20][21] or a combination of these factors.[19]

Neanderthal
Temporal range: MiddleLate Pleistocene 0.43/0.25–0.04 Ma
Slightly angled head-on view of a Neanderthal skeleton, stepping forward with the left leg
An approximate reconstruction of a Neanderthal skeleton. The central rib cage, including the sternum, and parts of the pelvis are from modern humans.
Scientific classification edit
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Haplorhini
Infraorder: Simiiformes
Family: Hominidae
Subfamily: Homininae
Tribe: Hominini
Genus: Homo
Species:
H. neanderthalensis
Binomial name
Homo neanderthalensis
King, 1864
Stretching across all of Portugal, Spain, Switzerland, Italy, England, southern Germany and Austria, all of Czech Republic, Hungary, Romania, Croatia, Montenegro, the Peloponnesian Peninsula, Crimean peninsula, Black Sea–Caspian Steppe west of the Caucasus, southern Turkey, northern Syria, the Levant, northern Iraq spilling over into Iran, the east end of Uzbekistan, and just northeast of Kazakhstan in Russia
Known Neanderthal range in Europe (blue), Southwest Asia (orange), Uzbekistan (green), and the Altai Mountains (violet).
Synonyms[6]

It is unclear when Neanderthals split from modern humans; DNA studies have produced results ranging from 182 kya[22] to more than 800 kya.[23] The time of divergence of Neanderthals from their ancestor H. heidelbergensis is also unclear. The oldest potential Neanderthal bones are dated to 430 kya, but the classification is uncertain.[24] Neanderthals are known from numerous fossils, especially from after 130 kya.[25] The type specimen, Neanderthal 1, was found in 1856 in the German Neander Valley. After much debate over their validity, Neanderthals were depicted as being primitive, stupid, and brutish for much of the early 20th century. Though knowledge and perception of them has markedly changed since then in the scientific community, the image of the unevolved caveman archetype remains prevalent in popular culture.[26][27]

Compared to modern humans, Neanderthals were stockier, with somewhat shorter limbs and a larger chest and nose. These features are often explained as adaptations to conserve heat in a cold climate, but (aside from body fat storage)[28] they are more likely products of genetic drift[29] and adaptations for sprinting in the warmer, forested landscape that Neanderthals often inhabited.[30] The braincases of Neanderthal men and women averaged about 1,600 cm3 (98 cu in) and 1,300 cm3 (79 cu in) respectively,[31][32][33] which is within the range of the values for modern humans. Average Neanderthal men stood around 165 cm (5 ft 5 in) and women 153 cm (5 ft) tall, similar to contemporary humans.[34]

Neanderthal technology is thought to have been quite sophisticated. It includes the Mousterian stone tool industry[35][36] and the abilities to create fire[37][38] and build cave hearths,[39][40] make the adhesive birch bark tar,[41] craft simple clothes similar to blankets and ponchos,[42] go seafaring through the Mediterranean,[43][44] make use of medicinal plants[45][46][47] as well as treat severe injuries,[48] and use various cooking techniques such as roasting[49] and smoking.[50] Neanderthals made use of a wide array of food, mainly hoofed mammals,[51] but also other megafauna,[52][53] plants,[54][55][56] small mammals, birds, and marine resources.[57] Though they were likely apex predators, they still competed with cave bears, cave lions, cave hyaenas, and other large predators.[58]:120–143 Several examples of Upper Paleolithic art have been controversially attributed to Neanderthals, most famously Spanish cave paintings contentiously[59] dated to before 65 kya[60][61] and the Divje Babe Flute.[62] Some claims of religious beliefs have been made.[63] Neanderthals were capable of speech, though it is unclear how complex their language was.[64][65]

The total population remained low, and they lived in small groups, interacting rarely with outsiders.[39][66] This led to the accumulation of harmful genes and inbreeding.[66] Neanderthals also lived in a high-stress environment with high trauma rates, and about 80% died before the age of 40.[67] Evidence for interbreeding between Neanderthals and anatomically modern humans was presented in the 2010 Neanderthal genome project's draft report.[68][69][70] It possibly occurred 316–219 kya[71] and more likely 100 kya and again after 65 kya.[72] Around 1–4% of all non-Subsaharan African genomes (Eurasians, Oceanians, Native Americans, and North Africans) derive from Neanderthals,[68][73][74] and about 20% of the Neanderthal genome survives today,[75] though many of the inherited genes may have been detrimental and selected out.[76]

TaxonomyEdit

EtymologyEdit

 
What remains of Kleine Feldhofer Grotte where Neanderthal 1 was discovered

Neanderthals are named after the site from which they were first identified, the Neander Valley, at the time in the Rhine Province of the Kingdom of Prussia (now in North Rhine-Westphalia, Germany). The valley itself was named for Joachim Neander, Neander being the Hellenized form of the surname Neumann ("new man").[77][78][79]

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".[80] The binomial name Homo neanderthalensis—extending the name "Neanderthal man" from the individual type specimen to the entire group, and formally recognizing 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.[81][82][83] However, in 1864, he recommended the genus name also be distinct from modern humans as he compared the Neanderthal braincase to that of a chimpanzee and argued they were "incapable of moral and theositic conceptions".[84]

 
Ernst Haeckel's Primate family tree showing H. stupidus (Neanderthal) as the ancestor to H. sapiens[1]

Since the historical spelling -th- in German represents the phoneme /t/, not the usual pronunciation of th with the fricative /θ/, standard British pronunciation of "Neanderthal" is with /t/ (IPA: /niːˈændərtɑːl/).[85][86][87][88][89] However, in English, "Neanderthal" is also pronounced with the usual th sound (as /niːˈændərθɔːl/).[90]

The German spelling of Thal ("valley") was current until 1901; it is now spelt Tal.[a][91][92] The spelling Neandertal is occasionally seen in English, even in scientific publications. Since Neanderthal and Neandertal are common names, there is no authoritative prescription on its spelling, unlike the spelling of the binominal name, H. neanderthalensis, which is predicated by King, 1864.[93] The common name in German is invariably Neandertaler (lit. "of the valley of Neander"), not Neandertal, but the spelling of the name of the Neander Valley itself (Neandertal vs. Neanderthal) has been affected by the species name, the names of the Neanderthal Museum and of Neanderthal station persisting with pre-20th-century spelling.[92]

Research historyEdit

The first Neanderthal remains—Engis 2—were discovered in 1829 by Dutch naturalist Philippe-Charles Schmerling in the Schmerling Caves, but he thought it was an ancient skull of an anatomically modern human.[94] In 1848, Gibraltar 1 from Forbes' Quarry was presented to the Gibraltar Scientific Society by their Secretary Lieutenant Edmund Henry Réné Flint, but was also thought to be a modern human skull.[95] In 1856, local schoolteacher Johann Carl Fuhlrott recognized bones from Feldhofer Cave in Neander Valley—Neanderthal 1 (the holotype specimen)—as distinct from modern humans, and gave it 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.[96][84] Following Charles Darwin's On the Origin of Species, Fuhlrott and Schaaffhausen argued the bones represented an ancient modern human form;[27][97][98][84] Schaaffhausen, a social Darwinist, believed that humans linearly progressed from savage to civilised, and so concluded that Neanderthals were barbarous cave-dwellers.[27] Fuhlrott and Schaaffhausen met opposition namely from the prolific 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,[99] which stalled Neanderthal research until the end of the century.[27][97]

 
Le Moustier by Charles R. Knight, 1920

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. French palaeontologist Marcellin Boule authored several publications, among the first to establish palaeontology as a science, detailing the specimen, but reconstructed it as ape-like and only remotely related to modern humans.[100][27] He fueled the popular image of Neanderthals as barbarous, slouching, club-wielding primitives; this image was reproduced for several decades and popularised in science fiction works, such as the 1911 The Quest for Fire by J.-H. Rosny aîné and the 1927 The Grisly Folk by H. G. Wells where they are depicted as monsters.[27] In 1911, Scottish anthropologist Arthur Keith reconstructed La Chapelle-aux-Saints 1 as an immediate precursor to modern humans, sitting next to a fire, producing tools, wearing a necklace, and having a more humanlike posture, but this failed to garner much scientific rapport, and Keith later abandoned his thesis in 1915.[27][101][97]

 
Neanderthal in a business suit at the Neanderthal Museum, Germany

By the middle of the century, based on several recent findings of other archaic humans (such as Homo erectus), the scientific community began to rework its understanding of Neanderthals. Ideas such as Neanderthal behaviour, intelligence, and culture were being discussed, and a more humanlike image of them emerged. In 1939, American anthropologist Carlton Coon reconstructed a Neanderthal in a modern business suit and hat to emphasize that they would be, more or less, indistinguishable from modern humans had they survived into the present. William Golding's 1955 novel The Inheritors depicts Neanderthals as much more emotional and civilised.[27][26] However, Boule's image continued to influence works until the 1960s. In modern day, Neanderthal reconstructions are often very humanlike.[97]

Hybridization between Neanderthals and early modern humans had been suggested early on,[102] such as by English anthropologist Thomas Huxley in 1890,[103] ethnographer Hans Peder Steensby in 1907,[104] and Coon in 1962.[105] In the early 2000s, several researchers argued in favour of admixture based on supposed hybrid specimens such as Lagar Velho 1[106][107][108][109] and Muierii 1.[110] Neanderthal admixture was found to be present in modern populations in 2010 with the mapping of the first Neanderthal genome sequence.[68]

ClassificationEdit

Homo sapiens

Denisovan from Denisova Cave

Denisovan from Baishiya Karst Cave

Neanderthal from Denisova Cave

Neanderthal from Sidrón Cave

Neanderthal from Vindija Cave

Phylogeny based on comparison of ancient proteomes and genomes with those of modern species.[111]

Neanderthals are hominids in the genus Homo, humans, and generally classified as a distinct species, H. neanderthalensis, though sometimes as a subspecies of modern human as H. sapiens neanderthalensis. This would necessitate the classification of modern humans as H. s. sapiens.[9]

A large part of the controversy stems from the vagueness of the term "species", as it is generally used to distinguish two genetically isolated populations, but admixture between modern humans and Neanderthals is known to have occurred.[9][112] However, the absence of Neanderthal-derived patrilineal Y-chromosome and matrilineal mitochondrial DNA (mtDNA) in modern humans, along with the underrepresentation of Neanderthal X chromosome DNA, could imply reduced fertility or frequent sterility of some hybrid crosses,[70][113][114][115] representing a partial biological reproductive barrier between the groups.[70]

In 2014, geneticist Svante Pääbo described such "taxonomic wars" as unresolveable, "since there is no definition of species perfectly describing the case".[9]

Neanderthals are thought to have been more closely related to Denisovans than to modern humans. Likewise, Neanderthals and Denisovans share a more recent last common ancestor (LCA) based on nuclear DNA (nDNA) than to humans. However, Neanderthals and modern humans share a more recent LCA based on mtDNA. This likely resulted from an interbreeding event subsequent to the Neanderthal/Denisovan split which introduced another mtDNA line. This involved either introgression coming from an unknown archaic human into Denisovans,[111][116][117][118][119] or introgression from an earlier unidentified modern human wave from Africa into Neanderthals.[120]

EvolutionEdit

Stage 1: early pre-Neanderthal, possibly H. erectus, (Tautavel Man, 450 kya)
Stage 2: archaic Neanderthal, possibly H. heidelbergensis (Miguelón, 430 kya)
Stage 3: early Neanderthal (Saccopastore I, 130 kya)
Stage 4: classic European Neanderthal (La Chapelle-aux-Saints 1, 50 kya)
The accretion model[121]

It is largely thought that H. heidelbergensis was the last common ancestor of Neanderthals, Denisovans, and modern humans after populations became isolated in Europe, Asia, and Africa respectively.[122] The taxonomic distinction between H. heidelbergensis and Neanderthals is mostly due to a fossil gap in Europe between 300 and 243 thousand years ago (kya) during marine isotope stage 8. "Neanderthals", by conventions, are fossils which date to after this gap.[121][52][22] However, 430,000 year (ka) old bones at Sima de los Huesos could represent early Neanderthals or a closely related group,[24] and the 400 ka old Aroeira 3 could represent a transitional phase. Basal and derived morphs could have lived concurrently.[123] The quality of the fossil record greatly increases from 130 kya onwards,[124] and specimens from this period make up the bulk of known Neanderthal skeletons.[125][126] Dental remains from the Italian Visogliano and Fontana Ranuccio sites indicate that Neanderthal dental features had evolved by around 450–430 kya during the Middle Pleistocene.[127]

There are two main hypotheses regarding the evolution of Neanderthals following the Neanderthal/human split: two-step and accretion. The former argues a single major environmental event—such as the Saale glaciation—caused European H. heidelbergensis to rapidly increase body size and robustness, as well as undergo an enlengthenment of the head, which then led to other changes in skull anatomy.[108] However, Neanderthal anatomy was most likely not driven by adapting to cold weather.[30] The latter holds that Neanderthals slowly evolved over time from the ancestral H. heidelbergensis, divided into 4 stages: early-pre-Neanderthals (MIS 12), pre-Neanderthals (MIS 119), early Neanderthals (MIS 7–5), and classic Neanderthals (MIS 4–3).[121]

Numerous dates for the Neanderthal/human split exist. The date of around 250 kya cites the Florisbad Skull ("H. helmei") as being the last common ancestor (LCA), and the split is associated with the Levallois technique of making stone tools. The date of about 400 kya uses H. heidelbergensis as the LCA. 600 kya says that "H. rhodesiensis" was the LCA, which split off into modern human lineage and a Neanderthal/H. heidelbergensis lineage.[128] 800 kya has H. antecessor as the LCA; however, different variations of this model would push the date back to 1 million years ago.[128][24] DNA studies have yielded various results on divergence time, such as 592–182 kya,[22] 553–321 kya,[129] 654–475 kya,[128] 690–550 kya,[77] 765–550 kya,[24][118] 800–520 kya,[130] before 800 kya,[23] and so forth.

Neanderthals and Denisovans are more closely related to each other than they are to modern humans, meaning the Neanderthal/Denisovan split occurred after their split with modern humans.[131][118][24][117] Using a mutation rate of 1x10-9 or 0.5x10-9 per base pair (bp) per year, the Neanderthal/Denisovan split occurred around either 236–190 kya or 473–381 kya respectively.[118] Using 1.1x10-8 per generation with a new generation every 29 years, the time is 744 kya. Using 5x10-10 nucleotide site per year, it is 644 kya. Using the latter dates, the split had likely already occurred by the time hominins spread out across Europe, and unique Neanderthal features had begun evolving by 600–500 kya.[117]

DemographicsEdit

RangeEdit

 
One of the most southernmost Neanderthal remains from Tabun Cave, Israel (120–50 kya) at the Israel Museum

Early Neanderthals, living before the Eemian interglacial (130 kya), are poorly known and come mostly from European sites. From 130 kya onwards, the quality of the fossil record increases dramatically. These later Neanderthals are recorded from Western, Central, Eastern, and Mediterranean Europe,[25] as well as Southwest, Central, and Northern Asia up to the Altai Mountains in southern Siberia.[132]

The southernmost find was recorded at Shuqba Cave, Levant;[133] reports of Neanderthals from the North African Jebel Irhoud[134] and Haua Fteah[135] 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, though these may be the result of convergent evolution rather than Neanderthals extending their range to the Pacific Ocean.[136] The limit of their northern bound appears to have been 53°N (Bontnewydd, Wales),[137] although it is difficult to assess because glacial advances destroy most human remains; Middle Palaeolithic artefacts have been found up to 60°N on the Russian plains,[138][139][140] but these are more likely attributed to modern humans.[141] A 2017 study claimed the presence of Homo at the 130 ka Californian Cerutti Mastodon site in North America,[142] but this is highly unlikely.[143][144][145]

It is unknown how the rapidly fluctuation climate of the last glacial period (Dansgaard–Oeschger events) impacted Neanderthals, as warming periods would produce more favorable temperatures, but encourage forest growth and deter megafauna; whereas frigid periods would produce the opposite.[66] However, Neanderthals may have preferred a forested landscape.[30] Populations may have peaked in cold but not extreme intervals, such as marine isotope stages 8 and 6 (respectively 300 and 191 kya). It is possible their range expanded and contracted as the ice retreated and grew respectively to avoid permafrost areas, residing in certain refuge zones.[66]

 
Distribution of Neanderthals and locations of main excavation sites

PopulationEdit

Like modern humans, Neanderthals probably descended from a very small population with an effective population—the number of individuals who can bear children—of 3,000 to 12,000 approximately. However, Neanderthals maintained this very low population, living in small, isolated, inbred groups.[146][117] Various studies, using mtDNA analysis, yield varying effective populations,[66] such as about 1,000 to 5,000;[146] 5,000 to 9,000 remaining constant;[147] or 3,000 to 25,000 steadily increasing until 50,000 BCE before declining until extinction.[148] However, all agree on low population,[66] which may have been up to 10 times smaller than contemporary human populations in Western Europe.[149] Estimates giving a total population in the tens of thousands[117] are contested.[146] A consistently low population may be explained in the context of the "Boserupian Trap": a population's carrying capacity is limited by the amount of food it can obtain, which in turn is limited by its technology. Innovation increases with population, but if the population is too low, innovation will not occur very rapidly and the population will remain low. This is consistent with the apparent 150 ka stagnation in Neanderthal lithic technology.[66]

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.[67] However, it has also been estimated that the age pyramids for Neanderthals and contemporary modern humans were the same.[66] Infant mortality was estimated to have been very high for Neanderthals, about 43% in northern Eurasia.[150]

AnatomyEdit

BuildEdit

Comparisons of a human (left) and a Neanderthal (right) skull at the Cleveland Museum of Natural History
Neanderthal skull features

Neanderthals had more robust and stockier builds than modern humans,[34] though still maintained an upright posture;[158] wider and barrel-shaped rib cages; wider pelvises;[159][52] and proportionally shorter forearms and forelegs.[160][30]

Based on 45 Neanderthal long bones from 14 men and 7 women, the average height was 164 to 168 cm (5 ft 5 in) for males and 152 to 156 cm (5 ft) for females. For comparison, the average height of 16 Upper Paleolithic and Mesolithic modern humans was 168.1 cm (5 ft 6 in) for males and 152.5 cm (5 ft) for females.[34] For Neanderthal weight, samples of 26 specimens found an average of 77.6 kg (171 lb) for males and 66.4 kg (146 lb) for females.[161] Using 76 kg (168 lb), the body mass index for Neanderthal males was calculated to be 26.9–28.2, which in modern humans correlates to being overweight. This indicates a very stocky build.[34] 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.[28]

The neck vertebrae of Neanderthals are longer and thicker than those of modern humans, lending to stability, possibly due to different head shape and size.[162] Though the Neanderthal thorax (where the ribcage is) was similar in size to modern humans, the longer and straighter ribs would have equated to a widened mid-lower thorax and stronger breathing in the lower thorax, which are indicative of a larger diaphragm and possibly greater lung capacity.[159][163][164] The lung capacity of Kebara 2 was estimated to have been 9.04 L (2.39 US gal). The Neanderthal chest was also more pronounced (expanded antero-posteriorly). The sacrum (where the pelvis connects to the spine) was more vertically inclined, and was placed lower in relation to the pelvis, causing the spine to be less curved (exhibit less lordosis) and to fold in on itself somewhat. Such modifications to the spine would have enhanced side-to-side (mediolateral) flexion, better supporting the wider lower thorax. This condition may be normal for Homo, with the condition of a narrower thorax in modern humans being a unique characteristic.[159]

Body proportions were usually cited as being "hyperarctic" as adaptations to the cold, because they are similar to those of human populations which developed in cold climates[165]—the Neanderthal build is most similar to that of Eskimos among modern humans[166]—and shorter limbs equates to higher retention of body heat.[160][165][167] However, Neanderthals from more temperate climates—such as Iberia—still retain the "hyperarctic" physique.[168] Further, the increasing evidence that Neanderthals preferred warmer wooded areas instead of open mammoth steppe and cold climate suggests to the contrary, and DNA analysis indicates a higher proportion of fast-twitch muscle fibers in the Neanderthals than modern humans. It is possible their body proportions and greater muscle mass were adaptations to sprinting as opposed to the endurance-oriented modern human physique,[30] as persistence hunting may only be effective in hot climates where the hunter can run prey to the point of heat exhaustion (hyperthermia). They had longer heel bones, reducing their ability for endurance running.[169] Their shorter limbs would have reduced moment arm at the limbs, which allowed for greater rotational force at the wrists and ankles without extra exertion of the rotating muscles at the elbows and knees by increasing the speed at which the muscles contract, allowing for faster acceleration.[160][170]

FaceEdit

 
Le Moustier Neanderthal facial reconstitution at the Neues Museum, Berlin

Neanderthals had a reduced chin, sloping forehead, and large nose, which also started somewhat higher on the face than in modern humans. The Neanderthal skull is typically more elongated and less globular than that of modern humans, and features an occipital bun,[171] or "chignon", a protrusion on the back of the skull, though it is within the range of variation for humans who have it. It is caused by the cranial base and temporal bones being placed higher and more towards the front of the skull, and a flatter skullcap.[172] They also had larger eyes likely to adapt to the low-light environment.[173]

The large Neanderthal nose and paranasal sinuses were once thought to warm air as it entered the lungs and retain moisture ("nasal radiator hypothesis"),[174] but the bone structure does not indicate any adaptations to cold climate, and sinuses are generally reduced in cold-adapted creatures. More likely, the large nose was caused by genetic drift; also, the sinuses are not grossly large, and are comparable in size to those of modern humans.[29][175][174]

Neanderthals featured a protrusion of the jaw (prognathism), which was once cited as a response to a large bite force evidenced by heavy wearing of Neanderthal front teeth (the "anterior dental loading hypothesis"), but similar wearing trends are seen in contemporary humans. It could also have evolved to fit larger teeth in the jaw, which would better resist wear and abrasion,[176][174] and the increased wear on the front teeth compared to the back teeth probably stem from repetitive use. Neanderthal dental wear patterns are most similar to those of modern Inuit. The bite force of Neanderthals and modern humans is now thought to be about the same,[174] about 285 N (64 lbf) and 255 N (57 lbf) in modern human males and females, respectively.[177]

BrainEdit

 
Bust at the Hall of Human Origins, Washington, D.C.

The Neanderthal braincase averages 1,600 cm3 (98 in3) for males and 1,300 cm3 (79 in3) for females,[31][32][33] within the possible range of modern humans,[178] which is, on average, 1,270 cm3 (78 in3) for males and 1,130 cm3 (69 in3) for females.[179] The largest Neanderthal brain, Amud 1, was calculated to be 1,736 cm3 (105.9 in3), one of the largest ever recorded in hominids.[180] However, both Neanderthal and human infants measure about 400 cm3 (24 in3), and either Neanderthal brain development sped up or modern human development slowed down from the last common ancestor.[181]

In Neanderthals, the occipital lobe—operating vision—was much larger than in modern humans, and, similarly, they had larger eyes, probably as an adaptation to lower light conditions in Europe. More brain tissue was devoted to bodily maintenance and control, and, consequently, the cognitive areas of the brain were proportionally smaller than in modern humans,[173] including the cerebellum (operating muscle memory, and possibly language, attention, working memory, social abilities, and thought), the parietal lobes (visuospatial function and episodic memory), the temporal lobes (language comprehension and associations with emotions), the orbitofrontal cortex (decision making), and the olfactory bulb (sense of smell).[182] A 2011 study looking at the brain asymmetry of 20 Neanderthals to predict handedness found 85% to be right-hand dominant and the remaining 15% left-handed, whereas modern humans are 52% right-dominant, 12% left-dominant, and 36% ambidextrous.[183]

Hair and skin colourEdit

 
Reconstruction of a Croatian Neanderthal at the Zagros Paleolithic Museum

The lack of sunlight most likely led to the proliferation of lighter skin in Neanderthals,[184] though modern Europeans did not evolve light skin until the Holocene.[185] Genetically, BNC2 was present in Neanderthals, which is associated with light skin colour; however, a second variation of BNC2 was also present, which is associated with darker skin colour in the UK Biobank.[184] It is likely Neanderthal skin colour varied from region to region. The DNA of three Croatian Neanderthals shows they had darker hair, skin, and eye colour than modern Europeans.[186]

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 5 known variants in modern humans of the gene which cause loss-of-function and are associated with light skin and hair colour, and another unknown variant in Neanderthals (the R307G variant) which could be associated with pale skin and red hair. The R307G variant was identified in a Neanderthal from Monti Lessini, Italy, and possibly Sidrón Cave, Spain.[187] However, like in modern humans, red was probably not a very common hair colour because the variant is not present in many other sequenced Neanderthals.[184]

MetabolismEdit

The Neanderthal physical activity level (PAL) was assumed to be a very high 650 counts per minute per day (CPM/d), in comparison to 200 CPM/d in modern Siberian hunter-gatherers. Average body fat percentage (BFP) was estimated to be 25%, though it may have been 13% for males and 20% for females in more temperate areas. Using these measurements and average height and weight, the daily total energy expenditure (TEE)—the amount of calories consumed in one day—was estimated to be 3,454–4,019 and 3,828–4,483 kcal for males with high and low BFPs respectively, and 3,115–3,538 and 3,258–3,710 kcal for females. However, if the PAL was reduced to that of modern Siberian hunter-gatherers, the TEE becomes 2,959–3,524 and 3,333–3,988 kcal for males, and 2,620–3,043 and 2,764–3,215 kcal for females. This is comparable with the upper end of energetic demands of modern hunter gatherers, and the latter estimates are most similar to the Siberian Yakuts, which contradicts earlier estimates of vastly higher energetic demands in Neanderthals than modern humans. Further, some Neanderthal populations are thought to have had a predominantly low-calorie plant diet, which suggests the minimum daily caloric intake was also low.[188]

 
Reconstruction of a Neanderthal woman[189]

Maximum lifespan, and the timing of adulthood, menopause, and gestation were most likely very similar to modern humans.[66] However, it has been hypothesized that Neanderthals matured faster than modern humans based on the growth rates of teeth and tooth enamel,[190][191] though this is not backed by age biomarkers.[67]

Genetically, Neanderthal-derived alleles near ASB1 and EXOC6 are associated with being an evening person, narcolepsy, and day-time napping.[184]

PathologyEdit

InjuriesEdit

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;[192] and an estimated 80% succumbed to their injuries and died before reaching 40. It was thus theorized that Neanderthals employed a risky hunting strategy.[67] 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[193] (perhaps swimmer's ear).[194]

However, trauma patterns between Neanderthals and contemporary humans are about the same,[195] and rates of cranial trauma are not significantly different between Neanderthals and Middle Paleolithic modern humans,[196] which indicate that they got hurt doing the same things. Therefore, interpersonal violence or large carnivores were more likely the source of severe trauma. A 2016 study looking at 124 Neanderthal specimens found that about 36% were victims of bear attacks, 21% big cat attacks, and 17% wolf attacks (totalling 92 positive animal attack cases, 74%). There were no cases of hyaena attacks, though hyaenas still nonetheless probably killed Neanderthals, at least opportunistically;[195] however, hyaenas and Neanderthals may have actively avoided each other.[51] Such intense predation probably stemmed from common confrontations due to competition over food and cave space, and from Neanderthals hunting these carnivores.[195]

Other ailmentsEdit

Likely due to advanced age (60s or 70s), La Chapelle-aux-Saints 1 had signs of Baastrup's disease, affecting the spine, and osteoarthritis.[158] Shanidar 1, who likely died at about 40 or 50, was diagnosed with the most ancient case of diffuse idiopathic skeletal hyperostosis (DISH), a degenerative disease which can restrict movement.[197]

Low population caused a low genetic diversity and probably inbreeding, which reduced the population's ability to filter out harmful mutations (inbreeding depression). However, 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 humans.[198] It is known, however, that the 13 inhabitants of Sidrón Cave collectively exhibited 17 different birth defects likely due to inbreeding.[199]

Neanderthals were likely subject to several infectious diseases and parasites. Modern humans likely transmitted diseases to them; one possible candidate is the stomach bacteria Helicobacter pylori.[200] A Neanderthal at Sidrón Cave shows evidence of a gastrointestinal Enterocytozoon bieneusi infection.[47] The leg bones of 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.[201] Neanderthals had a lower cavity rate than modern humans, despite some populations consuming typically cavity-causing foods in great quantity, which could indicate a lack of cavity-causing oral bacteria, namely Streptococcus mutans.[202]

Two 250 ka old Neanderthal children from Payré, France present the earliest known cases of lead exposure of any hominin. 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 25 km (16 mi) of the site.[203]

CultureEdit

Social structureEdit

Group dynamicsEdit

 
Artist's reconstruction of a Neanderthal man and child

Neanderthals likely lived in more sparsely distributed groups than contemporary modern humans,[66] but group size is thought to have averaged 10 to 30 individuals, similar to modern hunter-gatherers.[39] Reliable evidence of Neanderthal group composition comes from Sidrón Cave and the footprints at Le Rozel:[151] the former shows 7 adults, 3 adolescents, 2 juveniles, and an infant;[204] whereas the latter, based on footprint size, shows a group of 10 to 13 members where juveniles and adolescents made up 90%.[151]

A Neanderthal child's teeth analysed in 2018 showed it was weaned after 2.5 years, similar to modern hunter gatherers, and was born in the spring, which is consistent with modern humans and other mammals whose birth cycles coincide with environmental cycles.[203] Indicated from various ailments resulting from high stress at a low age, such as stunted growth, it was hypothesized that children of both sexes were put to work directly after weaning;[150] and that, upon reaching adolescence, an individual may have been expected to join in hunting large and dangerous game.[67] However, the bone trauma is comparable to modern Inuits, and their childhood was more likely similar to contemporary modern humans.[205] Further, such stunting more likely resulted from harsh winters and bouts of low food resources.[203]

Sites showing evidence of no more than three individuals may have represented nuclear families or temporary camping sites for special task groups (such as a hunting party).[39] Bands likely moved between certain caves depending on the season, returning to the same locations generation after generation.[206] Cave bears may have greatly competed with Neanderthals for cave space, and there is a decline in cave bear populations starting 50 kya onwards (though their extinction is attributed to modern humans).[207][208]

Inter-group relationsEdit

A self-sustaining population which avoids inbreeding consists of about 450–500 individuals, which would necessitate these bands to interact with 8–53 other bands, but more likely the more conservative estimate given low population density.[39] Analysis of the mtDNA of the Neanderthals of Sidrón Cave showed that the adult three men belonged to the same maternal lineage, while the three adult women belonged to different ones. This suggests a patrilocal residence (that a women moved out of her group to live with her husband).[209] However, the DNA of a Neanderthal from Denisova Cave shows that her parents were half-siblings,[118] and the inhabitants of Sidrón Cave were likely highly inbred.[199] Neanderthal groups probably rarely exchanged mates given the abundance of harmful genes.[117]

Considering most Neanderthal artefacts were sourced no more than 5 km (3.1 mi) from the main settlement, it is unlikely these bands interacted very often.[39] However, a skeleton from La Roche à Pierrot, France, showed a healed fracture on top of the skull apparently caused by a deep blade wound,[210] and another from Shanadir Cave was found to have a rib lesion characteristic of projectile weapon injuries, which could be considered as evidence for conflict.[211]

A few Neanderthal artefacts in a settlement could have originated 20, 30, 100, and 300 km (12.5, 18.5, 60, and 185 mi) away. It is possible that macro-bands formed, collectively encompassing 13,000 km2 (5,000 sq mi), with each band claiming 1,200–2,800 km2 (460–1,080 sq mi), maintaining strong alliances for mating networks or to cope with leaner times and enemies as is exhibited in the low-density hunter gatherer societies of the Western Desert of Australia.[39] There is also an apparent cemetery of six or seven individuals at La Ferrassie, which, in modern humans, is typically used as evidence of a corporate group which maintained a distinct social identity and controlled some resource, trading, manufacturing, and so on. La Ferrassie is also located in one of the richest animal-migration routes of Pleistocene Europe.[39] However, mapping of the Neanderthal brain indicates they may not have had the cognitive function required for complex social interactions and trade.[173]

Social hierarchyEdit

It is sometimes suggested, since they were hunters of challenging big game and lived in small groups, there was no sexual division of labour as seen in human societies. That is, men, women, and children all had to be involved in hunting, instead of men hunting with women and children foraging. However, in modern human societies, the higher the meat dependency, the higher the division of labour.[39] Further, tooth-wearing patterns in Neanderthal men and women suggest they commonly used their teeth for carrying items, but men exhibit more wearing on the upper teeth, and women the lower, suggesting some cultural differences in tasks.[212]

 
Skeleton and restoration model of La Ferrassie 1 at the National Museum of Nature and Science, Tokyo

It is controversially proposed that some Neanderthals wore decorative clothing or jewellery—such as a leopard skin or raptor feathers—to display elevated status in the group. The small number of Neanderthal graves found could be explained as only high-ranking members receiving an elaborate burial, as is the case for some contemporary human societies.[39] It has also been suggested that elderly Neanderthals were given special burial rites for lasting so long given the high mortality rates.[67] Alternatively, many more Neanderthals may have received burials, but the graves were infiltrated and destroyed by bears.[213] Given that 20 graves of Neanderthals aged under 4 have been found—over a third of all known graves—deceased children may have received greater care during burial than other age demographics.[205]

A study looking at Neanderthal skeletons recovered from several natural rock shelters shows that, although they were recorded as bearing several trauma-related injuries, none of them had significant trauma to the legs that would debilitate movement. This might indicate that individuals who could not keep up with the group while moving from cave to cave were left behind. The high mortality rate indicates grandparents were rare. These all could indicate a culture based on the idea that self-worth derives from contributing food to the group; a debilitating injury would remove this self-worth and result in near-immediate death.[67] However, there are several examples of individuals with debilitating injuries being nursed for several years, and, lacking much contact with neighbouring groups, they may have focused extra attention onto group member wellbeing.[205][48] Such an altruistic strategy may have ensured their survival as a species for so long, especially given the high trauma rates.[48]

FoodEdit

Hunting and gatheringEdit

 
Red deer, presumably the preferred game of Neanderthals

Neanderthals were once thought of as scavengers, but are now considered to have been apex predators.[214][215] Living in a forested environment, Neanderthals were likely ambush hunters, getting close to and attacking their target—a prime adult—in a short burst of speed, thrusting in a spear at close quarters.[216][30] Younger or wounded animals may have been hunted using traps, projectiles, or pursuit.[216] Nonetheless, they were able to adapt to a variety of habitats.[57][217] In 1980, it was hypothesised that two piles of mammoth skulls at La Cotte de St Brelade, Channel Islands, at the base of a gulley was evidence of mammoth drive hunting (causing them to stampede off a ledge),[218] but this is contested.[217]

They appear to have fed predominantly on hoofed mammals, namely red deer and reindeer as these two were the most abundant game,[51] but also on other Pleistocene megafauna such as ibex, wild boar, aurochs, mammoth, straight-tusked elephant, and woolly rhinoceros.[52][53][219] There is evidence of directed cave and brown bear hunting both in and out of hibernation, as well as butchering.[220] Analysis of Neanderthal bone collagen from the Croatian Vindija Cave shows nearly all of their protein needs derived from animal meat.[53] Some caves show evidence of regular rabbit and tortoise consumption. At Gibraltar sites, there are remains of 143 different bird species, many ground-dwelling such as the common quail, corn crake, woodlark, and crested lark.[57] Neanderthals also exploited marine resources on the Iberian, Italian, and Peloponnesian Peninsulas, where they waded or dived for shellfish,[57][221][222] and, at Vanguard Cave, Gibraltar, consumed large marine creatures such as the Mediterranean monk seal, the short-beaked common dolphin, the common bottlenose dolphin, Atlantic bluefin tuna, sea breem, and the purple sea urchin.[57][223] They also fished for freshwater fish in some areas, such as (though not limited to) Castelcivita Cave, Italy, for trout, chub, and eel.[222]

Some communities subsisted primarily on a plant-based diet.[54] For example, Neanderthals from Sidrón Cave in Spain, based on dental tarter, likely had a meatless diet of mushrooms, pine nuts, and moss, indicating they were forest foragers.[47] Edible plant and mushroom remains are recorded from several caves.[55] Remnants from the Israeli Amud Cave indicates a diet of figs, palm tree fruits, and various cereals and edible grasses.[56] Several bone traumas in the leg joints could possibly suggest habitual squatting, which, if the case, was likely done while gathering food.[224]

Dental tartar from Spy Cave indicates the inhabitants had a meat-heavy diet including woolly rhinoceros and mouflon sheep, while also regularly consuming mushrooms.[47] Neanderthal faecal matter from El Salt, Spain, dated to 50 kya—the oldest human faecal matter remains recorded—show elevated coprostanol levels (digested cholesterol indicating a meat-heavy diet) and elevated stigmastanol (deriving from plant matter).[225] Evidence of cooked food plants—mainly legumes and, to a far lesser extent, acorns—were discovered in the Israeli Kebara Cave, with its inhabitants possibly gathering plants in spring and fall and hunting in all seasons except fall, though the cave was probably abandoned in late summer to early fall.[46] At the Iraqi Shanidar Cave, Neanderthals collected plants with various harvest seasons, indicating they scheduled returns to the area to harvest certain plants, and that they had complex food-gathering behaviors for both meat and plants.[226]

Food preparationEdit

Neanderthals probably could employ a wide range of cooking techniques, such as roasting, and they may have been able to heat up or boil soup, stew, or animal stock.[49] The abundance of animal bone fragments at settlements may indicate the making of fat stocks from boiling bone marrow, possibly taken from animals that had already died of starvation. These methods would have substantially increased protein consumption, which was a major component of their nutrition to compensate for low carbohydrate intake.[49][227] Neanderthal tooth size had a decreasing trend after 100 kya, which could indicate an increased dependence on cooking or the advent of boiling, a technique that would soften food.[228]

 
Yarrow growing in Spain

At Sidrón Cave, Neanderthals likely cooked and possibly smoked food,[50] as well as used certain plants—such as yarrow and camomile—as flavouring,[49] though these plants may have instead been used for their medicinal properties. Further, the former hypothesis would assume a degree of cuisine complexity which lacks proper evidence.[45] However, flavouring food may not require greatly enhanced humanlike cognitive or imaginative abilities, as Japanese macaques are known to dunk sweet potatoes and wheat in the ocean before eating them to give them a salty taste.[229] At Gorham's Cave, Gibraltar, Neanderthals may have been roasting pinecones to access pine nuts.[57]

The archaeological record shows they commonly used animal hide and birch bark, and it is possible they used them to make cooking containers, though this is based largely on circumstantial evidence as neither fossilise well.[228] It is possible the Neanderthals at Kebara Cave used the shells of the spur-thighed tortoise as containers.[230]

After a big kill, the animal would probably have been butchered on the spot as it is unlikely they hauled back big kills intact to a settlement nor the entire group (including pregnant women and infants) followed the hunting party to eat at the site of the kill. To prevent spoilage during transport, one practical method would have been to construct scaffolds to hang and dry the meat, which would also reduce overall weight by a third.[231] Though there is no evidence that they could store food,[39] it seems likely they had the ability considering the great quantities of meat and fat which could have been gathered from typical prey items (namely mammoths).[232]

CompetitionEdit

 
Cave hyaena skeleton

Competition from large Ice Age predators was rather high. Cave lions likely targeted horses, large deer and wild cattle; and leopards primarily reindeer and roe deer; which heavily overlapped with Neanderthal diet. To defend a kill against such ferocious predators, Neanderthals may have engaged in a group display of yelling, arm waving, or stone throwing; or quickly gathered meat and abandoned the kill. However, at Spy Cave, the remains of wolves, cave lions, and cave bears—which were all major predators of the time—indicate Neanderthals hunted their competitors to some extent.[58]:120–143

Neanderthals and cave hyaenas may have exemplified niche differentiation, and actively avoided competing with each other. Though they both mainly targeted the same groups of creatures—deer, horses, and cattle—Neanderthals mainly hunted the former and cave hyaenas the latter two. Further, animal remains from Neanderthal caves indicate they preferred to hunt prime individuals, whereas cave hyaenas hunted weaker or younger prey and had a greater proportion of carnivore remains than herbivore remains. In fact, cave hyaena caves are distinguished from Neanderthal caves by the higher abundance of carnivore remains.[51] Nonetheless, cave hyaenas likely stole food and leftovers from Neanderthal campsites, and scavenged on dead Neanderthal bodies.[233]

CannibalismEdit

There are several instances of Neanderthals practising cannibalism across their range.[152][234] The first undisputed example came from Gran Dolina in 1999,[235] and other examples were found at Sidrón Cave,[236] Zafarraya in Spain; and the French Moula-Guercy Cave,[237] Les Pradelles, and La Quina. For the five cannabalised Neanderthals at the Goyet Caves in Belgium, there is evidence that the upper limbs were disarticulated, the lower limbs defleshed and also smashed (likely to extract bone marrow), the chest cavity disemboweled, and the jaw dismembered. There is also evidence that the butchers used some bones to retouch their tools. The processing of Neanderthal meat at Goyet Caves is similar to how they processed horse and reindeer.[152][234]

These cannibalistic tendencies have been explained as either ritual defleshing, pre-burial defleshing (to prevent scavengers or foul smell), for nutritional value, or as an act of war. Due to a small number of cases, and the higher number of cut marks seen on cannibalized individuals than animals (indicating inexperience), cannibalism was probably not a very common practice, and it may have only been done in times of extreme food shortages as in some cases in recorded human history.[234]

ArtEdit

White-tailed eagle claw with striations[238]
The scratched floor of Gorham's Cave

A large number of claims of Neanderthal art, adornment, and structures have been made. If accurate, these claims would show that Neanderthals were capable of symbolic thought or were cognitively comparable to modern humans. However, many of these are ambiguously attributed as the dating interlaps with modern human presence in Europe.[60][61] Claims are:

  • Purported Neanderthal bone flute fragments made of bear long bones are reported from Potok Cave, Slovenia, in the 1920s; and Istállóskö Cave, Hungary,[239] and Mokriška Jama Cave, Slovenia, in 1985; but these are now attributed to modern human activities.[240][241] The 1995 140–42 ka Divje Babe Flute from Slovenia has been attributed to Neanderthals, and Canadian musicologist Robert Fink said the original flute had either a diatonic or pentatonic musical scale.[242] Alternatively, it has also been claimed that the holes were made by a scavenging hyaena as there are a lack of cut marks stemming from whittling,[241] but it is highly unlikely the punctures were made by teeth, and cut marks are not always present on bone flutes.[62]
  • Excavated from 1949–1963 from the French Grotte du Renne, Châtelperronian beads made from animal teeth, shells, and ivory were found associated with Neanderthal bones, but the dating is uncertain and the beads may have been made by modern humans and simply redeposited with Neanderthal remains.[243][244][245][246]
  • In 1975, flower pollen on the body of the Iraqi Shanidar 4 was argued to be evidence of a flower burial,[247] but the pollen could have also been deposited by natural events.[248][249]
  • Discovered in 1975, the so-called Mask of la Roche-Cotard, a mostly flat piece of flint with a bone pushed through a hole on the midsection—dated to 32, 40, or 75 kya[250]—has been purported to resemble the upper half of a face, with the bone representing eyes.[251][252] It is contested whether it represents a face, or if it even counts as art.[253]
  • In a 2012 study examining 1,699 ancient sites across Eurasia, raptor and corvid bones, species not typically consumed by any human society, were argued to show evidence of feather plucking, specifically of the large flight feathers. The authors used this as evidence of bird feathers being worn as personal adornment.[254]
  • In 2012, deep scratches on the floor of Gorham's Cave, Gibraltar, were discovered, dated to older than 39 kya, which the discoverers have interpreted as Neanderthal abstract art.[255][256] The scratches could have also been produced by a bear.[213]
  • In 2015, a study argued that a number of 130 ka eagle talons found in a cache near Krapina, Croatia, associated with Neanderthal bones had been modified to be used as jewellery.[238][257] A similar 39 ka Spanish imperial eagle talon necklace was reported in 2019 at Cova Foradà in Spain, though from the contentious Châtelperronian layer.[258]
  • In 2017, incision-decorated raven bones from the Zaskalnaya VI (Kolosovskaya) Neanderthal site, Crimea, Micoquian industry dated to 43–38 kya were reported. Given there are 17 of these objects at seven different sites in the area, and the notches on all of them are more-or-less equidistant to each other, they are very unlikely to have originated from simple butchering.[259]
  • In 2018, red painted symbols comprising hand stencils, a ladder-shape figure, dots, discs, lines, and representations of animals on the cave walls of several caves across Spain 700 km (430 mi) apart, including La Pasiega, Cave of El Castillo, and Doña Trinidad–were dated to be older than 66 kya. This is at least 20 ka prior to the arrival of anatomically modern humans in western Europe.[60][61][260] However, the dating, and thus, its attribution to Neanderthals, is contested.[59]
  • In 2018, 3 perforated seashell beads (as if they were worn as jewellery) that are at least 115 ka old were found in Cave of Los Aviones in southeastern Spain. One has traces of red pigment. A fourth unperforated shell has multiple colours, which the author asserted was a palette. However, the perforations were naturally occurring, and the shells may have been picked up for no specific reason.[261]
  • In 2018, an engraved flint flake was found in the grave a Neanderthal child in Crimea, Ukraine.[262]

TechnologyEdit

Despite the apparent 150 ka stagnation in Neanderthal innovation,[66] there is evidence that Neanderthal technology was more sophisticated than was previously thought.[64] However, the high frequency of potentially debilitating injuries could have prevented very complex technologies from emerging, as a major injury would have impeded an expert's ability to effectively teach a novice.[192]

Tool manufacturingEdit

Levallois point from Beuzeville, France
Levallois technique

Neanderthals made stone tools, and are associated with the Mousterian industry.[35] The Mousterian is also associated with North African H. sapiens as early as 315 kya[263] and was found in Northern China about 47–37 kya.[264] The Levallois technique they adopted (also not unique to Neanderthals) maximized the cutting surface with the least amount of flint. As a difficult-to-learn process, the technique may have been directly taught generation to generation rather than via purely observational learning.[36] There is some debate if they had long-ranged weapons.[265] A wound on the neck of an African wild ass was likely inflicted by a Levallois point javelin,[266] and bone trauma consistent with habitual throwing has been reported in Neanderthals.[265][267]

Neanderthals may also be associated with the Châtelperronian 45–40 kya in central France and northern Spain, borrowing tool-making techniques from immigrating modern humans, namely crafting bone tools using grinding and polishing rather than the percussion technique used with stone,[268][269][270][271] though Neanderthal attribution of these artefacts is contested.[272]

The Neanderthals in 10 coastal sites in Italy (namely Grotta del Cavallo and Grotta dei Moscerini) and Kalamakia Cave, Greece, have been known to craft scrapers using smooth clam shells, and possibly hafted them to a wooden handle. They probably chose this clam species because it has the most durable shell. At Grotta dei Moscerini, about 24% of the shells were gathered alive from the seafloor, meaning these Neanderthals had to wade or dive into shallow water to collect them. At Grotta di Santa Lucia, Italy, in the Campanian volcanic arc, Neanderthals collected the porous volcanic pumice, which, in contemporary humans, was probably used for polishing points and needles. The pumices are associated with shell tools.[222]

At the Italian Poggetti Vecchi site, there is evidence they used fire to process boxwood branches to make digging sticks, a common implement in hunter-gatherer societies.[273]

Fire and constructionEdit

Neanderthal were able to create fire,[37][274][38] and utilise complex spatial organisation, indicating equatable intelligence levels with contemporary humans. Certain areas in a settlement were used for specific activities, such as for knapping, butchering, hearths, and wood storage. Many Neanderthal sites lack evidence for such activity likely due to natural degradation of the area, such as by bear infiltration after abandonment of the settlement.[213]

In a number of caves, evidence of hearths has been detected. Neanderthals likely considered air circulation when making hearths as a lack of proper ventilation for a single hearth can render a cave uninhabitable in several minutes.[40] Abric Romaní rock shelter indicates 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.[39][40] At Cueva de Bolomor, with hearths lined up against the wall, the smoke flowed upwards to the ceiling, and led to outside the cave. In Grotte du Lazaret, smoke was probably naturally ventilated during the winter as the interior cave temperature was greater than the outside temperature; likewise, the cave was likely only inhabited in the winter.[40]

 
The ring structures in Bruniquel Cave, France

In 1990, two 176 ka ring structures made of broken stalagmite pieces, several metres wide, were discovered in a large chamber more than 300 m (980 ft) from the entrance within Bruniquel Cave, France. One ring was 6.7 m × 4.5 m (22 ft × 15 ft) with stalagmite pieces averaging 34.4 cm (13.5 in) in length, and the other 2.2 m × 2.1 m (7.2 ft × 6.9 ft) with pieces averaging 29.5 cm (11.6 in). There were also 4 other piles of stalagmite pieces for a total of 112 m (367 ft) or 2.2 t (2.4 short tons) worth of stalagmite pieces. A team of Neanderthals was likely necessary to construct these, and the structures may have been used as light sources, but the chamber's actual purpose is uncertain. Building complex structures so deep in a cave is unprecedented in the archaeological record, and indicates sophisticated lighting and construction technology, and great comfortability in subterranean environments.[275]

Bark tarEdit

Neanderthal produced the adhesive birch bark tar, perhaps using plant-based resins for hafting.[276] It was long believed that birch bark tar required a complex recipe to be followed, and that it thus showed complex cognitive skills and cultural transmission. However, a 2019 study showed it can be made simply by burning birch bark on smooth vertical surfaces, such as a flat, inclined rock.[41]

ClothesEdit

Neanderthals were likely able to survive in a similar range of temperatures as modern humans while sleeping; about 32 °C (90 °F) while naked in the open and windspeed 5.4 km/h (3.4 mph), or 27–28 °C (81–82 °F) while naked in an enclosed space. Since ambient temperature were markedly lower than this—averaging during the Last Interglacial 17.4 °C (63.3 °F) in July and 1 °C (34 °F) in January and dropping as a low as −30 °C (−22 °F) on the coldest days—they may have required tailored clothing capable of preventing airflow to the skin. Especially during extended periods of travelling (such as a hunting trip), tailored footwear completely enwrapping the feet may have been necessary.[231]

However, as opposed to the bone sewing-needles and stitching awls of contemporary modern humans, the only known Neanderthal tools that could have been used to fashion clothes are hide scrapers, which could have made items similar to blankets or ponchos.[42][277] Nonetheless, Neanderthals would have needed to cover up most of their body. Contemporary humans would have covered 80–90%.[277][278]

Since human/Neanderthal admixture is known to have occurred in the Middle East, and no modern body louse species descends from Neanderthals (body lice only inhabit clothed individuals), it is possible Neanderthals (and/or humans) in hotter climates did not wear clothes, or Neanderthal lice were highly specialised.[278]

SeafaringEdit

Remains of Middle Paleolithic stone tools on Greek islands indicate early seafaring by Neanderthals in the Ionian Sea possibly starting as far back as 200–150 kya. The oldest stone artefacts from Crete date to 130–107 kya, Cephalonia 125 kya, and Zakynthos 110–35 kya. Here, they likely employed simple reed boats and made one-day crossings back and forth.[43] Other Mediterranean islands include Sardinia, Melos, Alonnisos,[44] and Naxos (though Naxos may have been connected to land),[279] and it is possible they crossed the Strait of Gibraltar.[44] Their ability to engineer these boats and navigate through open waters speaks to their advanced cognitive and technical skills.[44][279]

MedicineEdit

They appear to have had some knowledge of medicine comparable to contemporary humans.[48] An individual at Sidrón Cave seems to have been self-medicating a dental abscess using poplar—which contains salicylic acid, the active ingredient in aspirin—and there were also traces of the antibiotic-producing Penicillium.[47] They may have also used yarrow and camomile, and their bitter taste—which should act as a deterrent as it could indicate poison—means it was likely a deliberate act.[45] In Kebara Cave, plant remains which have historically been used for their medicinal properties were found, including the common grape vine, the pistachios of the Persian turpentine tree, ervil seeds, and oak acorns.[46]

Given high trauma rates and evidence of healing, Neanderthals appear to have been well-equipped at handling severe injuries. Well-healed fractures on many bones indicate the setting of primitive splints. Individuals with severe head and rib traumas (which would have caused massive blood loss) indicates they had some manner of dressing major wounds, and bandages could have been made from animal skin. By-and-large, they appear to have avoided severe infections, indicating good long-term treatment of such wounds.[48]

LanguageEdit

 
Reconstruction of the Kebara 2 skeleton at the Natural History Museum, London

It was once thought Neanderthals were anatomically unable to produce quantal vowels, which are present in all modern human languages, because they had a large mouth and thus lacked the necessity for a descended larynx to fit the entire tongue inside the mouth.[280][281] However, this is not a safe assumption to make,[282] and the modern human vocal apparatus and thus vocal repertoire were likely already present in the ancestral H. heidelbergensis.[65] The 1983 discovery of a Neanderthal hyoid bone—used in speech production in humans—in Kebara 2 which is almost identical to that of humans suggests Neanderthals were capable of speech.[65] The prevailing hypothesis for a long time was that speech spontaneously developed very recently in modern humans,[64] and some argued that the hyoid could have a different usage in Neanderthals, as it is simply used in tongue movement, including while chewing.[280][281]

The degree of language complexity is difficult to establish, but given that Neanderthals achieved some technical and cultural complexity, and interbred with humans, it is reasonable to assume they were at least fairly articulate, comparable to modern humans. A somewhat complex language—possibly using syntax—was likely necessary to survive in their harsh environment, with Neanderthals needing to communicate about topics such as locations, hunting and gathering, and tool-making techniques.[64][283][284]

Neuroscientist Andrey Vyshedskiy argued that Neanderthals lacked mental synthesis, the behaviorally modern human imaginative ability to craft effectively infinite ideas using a finite amount of words. This is a hallmark of behavioural modernity, which he believed spontaneously appeared by about 70 kya (the "Upper Paleolithic Revolution").[285] However, behavioural modernity is regarded as a process initiated as early as 400 kya,[286] and was possibly also exhibited in Neanderthals.[287][284]

The FOXP2 gene in modern humans is important in speech and language development. FOXP2 was present in Neanderthals,[288] but not the gene's modern human variant.[289] Neurologically, Neanderthals had an expanded Broca's area—operating the formulation of sentences, and speech comprehension—but 11 out of 48 genes which encode for language brainwaves had different methylation patterns between Neanderthals and modern humans. This indicates a stronger ability in modern humans than in Neanderthals to express language.[290]

ReligionEdit

FuneralsEdit

 
Reconstruction of the grave of La Chapelle-aux-Saints 1 at the Musée de La Chapelle-aux-Saints

Claims that Neanderthals held funerals for their dead with symbolic meaning,[124]:158–60 are heavily contested.[291][292][293] Even if the burial was intentional, it is not indicative of a religious belief of life after death, as such burial could have been the result of great emotion[294] or to prevent scavenging.[285]

The debate on Neanderthal funerals has been active since the 1908 discovery of La Chapelle-aux-Saints 1 in a small, nonnatural hole in a cave in southwestern France, very controversially attributed to have been buried in a symbolic fashion.[295][291][296] Another grave at Shanidar Cave was associated with the pollen of several flowers that may have been in bloom at the time of deposition—yarrow, Centaurea, ragwort, grape hyacinth, joint pine, and hollyhock.[297] The medicinal properties of the plants led American archaeologist Ralph Solecki to claim that the man buried was some leader, healer, or shaman.[298] However, it is also possible the pollen was deposited by a small rodent after the man's death.[299]

The graves of children and infants, especially, are associated with grave goods such as artefacts and bones. The grave of a newborn from La Ferrassie was found with three flint scrapers, and infant from Dederiyeh Cave, Syria, was found with a triangular flint placed on its chest. A 10 month old from Amud Cave was associated with a red deer mandible, likely purposefully placed there given other animal remains are now reduced to fragments. Teshik-Tash 1 from Uzbekistan was associated with a circle of ibex horns and a limestone slab argued to have supported the head.[205] A child from Kiik-Koba, Crimea, Ukraine had a flint flake with some purposeful engraving on it, likely requiring a great deal of skill, and possibly made with some symbolic significance.[262] Nonetheless, these contentiously constitute evidence of symbolic meaning as the grave goods' significance and worth are unclear.[205]

CultsEdit

It was once argued that the bones of the cave bear, particularly the skull, in some European caves were arranged in a specific order, indicating an ancient bear cult that killed bears and then ceremoniously arranged the bones. This would be consistent with bear-related rituals of modern human Arctic hunter gatherers, but the alleged peculiarity of the arrangement could also be well-explained by natural causes,[63][294] and bias could be introduced as the existence of a bear cult would conform with the idea that totemism was the earliest religion, leading to undue extrapolation of evidence.[300]

It was also once thought that Neanderthals hunted, killed, and cannibalised other Neanderthals to use the skull as the focus of some ritual.[234] In 1962, Italian palaeontologist Alberto Blanc believed a skull from Grotta Guattari, Italy, had evidence of a swift blow to the head—indicative of ritual murder—and a precise and deliberate incising at the base to access the brain. He compared it to the victims of headhunters in Malaysia and Borneo,[301] putting it forward as evidence of a skull cult.[294] However, it is now thought to have been a result of cave hyaena predation.[302] Though Neanderthals are known to have practiced cannibalism, there is unsubstantial evidence to suggest ritual defleshing.[152]

InterbreedingEdit

Interbreeding with modern humansEdit

 
Reconstruction of the upper Paleolithic human Oase 2 with around 7.3% Neanderthal DNA (from an ancestor 4–6 generations back)[303]

The first Neanderthal genome sequence was published in 2010, and strongly indicated interbreeding between Neanderthals and early modern humans.[68][304][305][73] The genomes of all non-sub-Saharan populations contain Neanderthal DNA.[68][306][70][74] Various estimates exist for the proportion: 1–4% in modern Eurasians,[68] 3.4–7.9%,[307] and 1.8–2.4% in modern Europeans and 2.3–2.6% in modern East Asians.[308] Such low percentages indicate infrequent interbreeding.[309] Of the inherited Neanderthal genome, 25% in modern Europeans and 32% in modern East Asians may be related to viral immunity.[310] In all, approximately 20% of the Neanderthal genome appears to have survived in the modern human gene pool.[75]

However, due to their small population and resulting reduced effectivity of natural selection, Neanderthals accumulated several weakly harmful mutations, which were introduced to and slowly selected out of the much larger human population; the initial hybridised population may have experienced up to a 94% reduction in fitness compared to contemporary humans. By this measure, Neanderthals may have substantially increased in fitness.[76] A 2017 study focusing on archaic genes in Turkey found associations with coeliac disease, malaria severity, and Costello syndrome.[311] Nonetheless, some genes may have helped modern human Europeans adapt to the climate; the Val92Met variant of the MC1R gene, the most frequent variant associated with red hair and light skin, may descend from Neanderthals[312] though this is contested as the variant was rare in Neanderthals,[184] and light skin in modern humans did not become prevalent until the Holocene.[185]

According to linkage disequilibrium mapping, the last Neanderthal gene flow into the modern human genome occurred 86–37 kya, but most likely 65–47 kya.[313] However, the approximately 40 ka old modern human Oase 2 was found, in 2015, to have had 6–9% (point estimate 7.3%) Neanderthal DNA, indicating a Neanderthal ancestor up to four to six generations earlier,[303] but this hybrid Romanian population does not appear to have made a substantial contribution to the genomes of later Europeans.[303]

In 2016, the DNA of Neanderthals from Denisova Cave shows evidence of interbreeding 100 kya, and interbreeding with an earlier dispersal of H. sapiens may have occurred as early as 120 kya in places such as the Levant.[72] The earliest H. sapiens remains outside of Africa occur at Misliya Cave 194–177 kya, and Skhul and Qafzeh 120–90 kya.[314] However, the Neanderthals of the German Hohlenstein-Stadel Cave have deeply divergent mtDNA compared to more recent Neanderthals, possibly due to introgression of human mtDNA between 316–219 kya, or simply because they were genetically isolated.[71] Whatever the case, these first interbreeding events have not left any trace in modern human genomes.[315]

Due to the absence of Neanderthal-derived mtDNA (which is passed on from mother to child) in modern populations,[115][316][77] it has been suggested that the progeny of Neanderthal females who mated with modern human males were either rare, absent, or sterile—that is to say, admixture stems from the progeny of Neanderthal males with modern human females.[317][316][113][69] Due to the lack of Neanderthal-derived Y-chromosomes in modern humans (which is passed on from father to son), it has also been suggested that the hybrids that contributed ancestry to modern populations were predominantly females, or the Neanderthal Y-chromosome was not compatible with H. sapiens and became extinct.[318]

Detractors of the interbreeding model argue that the genetic similarity is only a remnant of a common ancestor instead of interbreeding,[319] though this is unlikely as it fails to explain why sub-Saharan Africans do not have Neanderthal DNA.[305] Anthropologist John D. Hawks has argued that the genetic similarity to Neanderthals may be the result of both common ancestry and interbreeding, as opposed to just one or the other.[320]

Interbreeding with DenisovansEdit

 
Chris Stringer's Homo family tree. The horizontal axis represents geographic location, and the vertical time in millions of years ago.[b]

Though nDNA confirms that Neanderthals and Denisovans are more closely related to each other than they are to modern humans, Neanderthals and modern humans share a more recent maternally-transmitted mtDNA common ancestor, possibly due to interbreeding between Denisovans and some unknown human species. The 400 kya Neanderthal-like humans from Sima de los Huesos in northern Spain, looking at mtDNA, are more closely related to Denisovans than Neanderthals. Several Neanderthal-like fossils in Eurasia from a similar time period are often grouped into H. heidelbergensis, of which some may be relict populations of earlier humans, which could have interbred with Denisovans.[322] This is also used to explain an approximately 124 ka old German Neanderthal specimen with mtDNA that diverged from other Neanderthals (except for Sima de los Huesos) about 270 kya, while its genomic DNA indicated divergence less than 150 kya.[71]

Sequencing of the genome of a Denisovan from Denisova Cave has shown that 17% of its genome derives from Neanderthals.[116] This Neanderthal DNA more closely resembled that of a 120 ka old Neanderthal bone from the same cave than that of Neanderthals from Vindija Cave in Croatia or Mezmaiskaya Cave in the Caucasus, suggesting that interbreeding was local.[118]

For the 90 ka old Denisova 11, it was found that her father was a Denisovan related to more recent inhabitants of the region, and her mother a Neanderthal related to more recent European Neanderthals at Vindija Cave, Croatia. The discovery of a first generation hybrid indicates interbreeding was very common between these species, and Neanderthal migration across Eurasia likely occurred sometime after 120 kya.[323]

ExtinctionEdit

 
Map emphasizing the Ebro river in northern Spain

Neanderthals are thought to have died out between 41 and 39 kya.[324][10][11][12][13] Though some Neanderthals in Gibraltar were dated to later than this—such as Zafarraya (30 kya)[325] and Gorham's Cave (28 kya)[326]—prompting a hypothesis of some Iberian refuge with the Ebro River providing a geographical barrier, these dates are likely incorrect as they were based on ambiguous artefacts instead of direct dating.[13] A claim of Neanderthals surviving in a polar refuge in the Ural Mountains[139] is loosely supported by Mousterian stone tools dating to 34–31 kya at a time when modern humans may not yet have colonised the northern reaches of Europe;[141] however, modern human remains are known from the northern Siberian Mamontovaya Kurya site dating to 40 kya.[327]

Whatever the cause of their extinction, Neanderthals were replaced by modern humans, indicated by near full replacement of Mousterian stone technology with modern human Aurignacian stone technology by 38 kya in the fossil record.[16] Modern human remains dating to 45–43 kya have been found in Italy[328] and Britain,[329] and this migration of modern humans replaced Neanderthals.[10] A 2019 reanalysis of 210 ka old skull fragments from the Greek Apidima Cave assumed to have belonged to a Neanderthal concluded that they belonged to a modern human, and DNA evidence indicates H. sapiens contact with Neanderthals and admixture as early as 100 kya, meaning there were multiple human immigration events into Europe. A Neanderthal skull dating to 170 kya from Apidima Cave indicates H. sapiens were replaced by Neanderthals until their return about 40 kya.[330]

Modern humansEdit

 
Replacement of Neanderthals by early modern humans.

ExterminationEdit

In 1911, Boule was the first person to suggest modern humans forcefully took Europe from the Neanderthals due to their inherent inferiority[100] ; this erroneous[39][26][27] sentiment echoed for decades. In 1992, Jared Diamond in his book The Third Chimpanzee said competitive replacement often occurs in human history when a more technologically advanced culture (modern humans) meets a less advanced culture (Neanderthals).[15] However, there is a notable lack of Neanderthal/human violence in the fossil record. To explain this, in 1999, military historian Azar Gat suggested that, instead of an overt genocide, modern humans used hostile demonstrations to scare Neanderthals off of favourable living grounds, but still massacred them when these demonstrations were ineffective. Though peaceful interactions occurred, he maintained that aggressive ones were predominant, which led to their extinction.[331]

The first evidence of Neanderthal/human contact was discovered in 2009 with a mandible belonging to a young Neanderthal from the modern-human-inhabited (indicated by Aurignacian technology) Les Rois Cave, France. The mandible shows cut marks which are similar to the butchery seen in the reindeer bones also in the cave, which could indicate human predation of Neanderthals, human skinning of Neanderthal heads (as trophies), or a pre-burial ritual involving tooth extraction. The latter is better supported as other Aurignacian sites have evidence of post-mortem tooth extraction (but for modern humans), perhaps to use as jewellery.[332]

Shanidar 3 died from complications from a stab wound, likely originating from a light-weight, long-range projectile, a technology that possibly only H. sapiens had, which implies Neanderthal/modern human violence.[333] However, the Lower Paleolithic Schöningen spears[265] and Neanderthal trauma consistent with habitual throwing could indicate they were familiar with ranged weapons.[265][267]

CompetitionEdit

Though modern human expansion and Neanderthal contraction are correlated, this may instead due to the competitive exclusion principle with modern humans outcompeting and outperforming Neanderthals rather than exterminating them.[14] However, largely human-free tropical Asia was colonised by modern humans by 60 kya, meaning European colonisation was, for some reason, delayed, and, though colder climate may have influenced immigration speed, it is possible the presence of Neanderthal settlements inhibited modern human expansion for some time.[334]

Though Neanderthals likely exhibited modern behaviour, discrediting arguments of plain modern human superiority, the spread of grasslands and open steppe would have favoured humans (which evolved in a grassland environment) over Neanderthals (forest environment);[16] modern humans could push into colder areas with bigger game wearing their fitted clothes, which were more effective at insulating than Neanderthal ponchos;[335] and raw material and animal remain sourcing in the southern Caucasus suggest that modern humans were able to use extensive social networks to acquire resources from a greater area in leaner times, whereas Neanderthals likely restricted themselves to more local sources since most of their stone artefacts were drawn from within 5 km (3.1 mi).[39][336] This climate change may have depopulated several regions of Neanderthals, like previous cold spikes, but these areas were instead repopulated by immigrating humans, leading to Neanderthal extinction.[337]

Anthropologist Pat Shipman suggested that domestication of the dog could have played a role in Neanderthals' extinction, or rather, a symbiosis with wolves long preceding domestication. She claims that modern humans, about 50–45 kya, evolved the whites of the eyes to allow for more effective non-verbal communications with wolves, and this gave modern humans an advantage in hunting. She also claims that Neanderthals did not have very prominent whites of the eyes, like the rest of the animal kingdom.[58]:214–226

Climate changeEdit

It has also been proposed that climate change was the primary driver, as Neanderthals extinction coincided with the start of a very cold period, and their low population left them vulnerable to any environmental change, with even a small drop in survival or fertility rates possibly quickly leading to their extinction.[338] Their ultimate extinction coincides with Heinrich event 4, a period of intense cold and dry climate causing their preferred forest landscape to give way to steppeland, and future Heinrich events are also associated with massive cultural turnovers where European human populations collapsed.[17][18]

 
Graphic of the ash cloud from the Campanian Ignimbrite Eruption

Extinction also coincides with the Campanian Ignimbrite Eruption in Italy around 40 kya, which caused a 2–4°C cooling event for a year and acid rain for several years. By that time, Neanderthal populations may have already been dwindling from other factors, and the eruption could have led to their final demise.[19][339]

DiseaseEdit

Modern humans may have introduced African diseases to Neanderthals, contributing to their extinction. Lacking immunity, compounded by an already low population, first contact may have been devastating to the Neanderthal population. Low genetic diversity may have also rendered fewer Neanderthals naturally immune to these new diseases.[21]

Low population and inbreeding depression may have caused maladaptive birth defects, which could have contributed to their decline.[199]

In late-20th-century New Guinea, due to ritual cannibalism, the population was decimated by transmissible spongiform encephalopathies (kuru), a highly virulent disease spread by ingestion of prions found in brain tissue. However, individuals with the 129 variant of the PRNP gene were naturally immune to the prions. Because there were some cases of cannibalism in Neanderthals, it is possible that a kuru-like disease was shared between modern humans and Neanderthals, and the Neanderthals had very few naturally immune members due to their already small population, and were very quickly killed off.[20][340][341]

In popular cultureEdit

 
Cavemen in The Black Terror #16 (1946)

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, anti-social characters driven solely by animal instinct. "Neanderthal" can also be used as an insult.[26]

In literature, they are sometimes depicted as brutish or monstrous, such as in H. G. Well's The Grisly Folk and Elisabeth Thomas' The Animal Wife, but also civilised, such as 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.[27]

See alsoEdit

NotesEdit

  1. ^ The German noun is cognate with English dale. The German /t/ phoneme was frequently spelled th throughout the 15th to 19th centuries; Tal became standardized with the German spelling reform of 1901, thus the German name Neandertal for both the valley and species.
  2. ^ Homo floresiensis originated in an unknown location from unknown ancestors and reached remote parts of Indonesia. Homo erectus spread from Africa to western Asia, then east Asia and Indonesia; its presence in Europe is uncertain, but it gave rise to Homo antecessor, found in Spain. Homo heidelbergensis originated from Homo erectus in an unknown location and dispersed across Africa, southern Asia and southern Europe (other scientists interpret fossils, here named heidelbergensis, as late erectus). Humans spread from Africa to western Asia and then to Europe and southern Asia, eventually reaching Australia and the Americas. In addition to Neanderthals and Denisovans, a third gene flow of archaic Africa origin is indicated at the right.[321]

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