Open main menu

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

Neanderthal
Temporal range: MiddleLate Pleistocene 0.43/0.25–0.04 Ma
Neanderthalensis.jpg
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
Range of NeanderthalsAColoured.png
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 humans, with DNA studies ranging from 182 kya[22] to before 800 kya,[23] and the time of divergence from the ancestral Homo heidelbergensis is also vague. The oldest potential Neanderthal bones are dated to 430 kya, but the classification is uncertain.[24] They are known from numerous fossils, especially following 130 kya.[25] The type specimen, Neanderthal 1, was found in 1856 in the German Neander Valley. After much debate over its validity, Neanderthals were depicted as being primitive, stupid, and brutish, for much of the early 20th century. Though their image in the scientific community has markedly changed since, the unevolved caveman archetype remains aloof in popular culture.[26][27]

Compared to modern humans, Neanderthals were stockier with somewhat shorter limbs, and bigger chest and nose. These are often explained as adaptations to conserve heat in a cold climate, but are more likely adaptations for sprinting in the warmer, forested landscape they often inhabited,[28] and products of genetic drift.[29] The braincases of Neanderthal men and women averaged about 1,600 cm3 (98 cu in) and 1,300 cm3 (79 cu in) respectively,[30][31][32] within the range of the values for modern humans. Average Neanderthal men stood around 165 cm (5.5 ft) and women 153 cm (5 ft) tall, similar to contemporary humans.[33]

Neanderthal technology is thought to have been somewhat sophisticated, and include the Mousterian stone tool industry,[34][35] the ability to create fire[36][37] and build cave hearths,[38][39] making the adhesive birch bark tar,[40] crafting simple clothes similar to blankets and ponchos,[41] seafaring through the Mediterranean,[42][43] making use of medicinal plants,[44][45][46] and using various cooking techniques (such as roasting[47] and smoking[48]). Though they were likely apex predators, they still competed with cave bears, cave lions, cave hyaenas, and wolves.[49]:120–143 Several examples of Upper Paleolithic art have been controversially attributed to Neanderthals–most famously Spanish cave paintings contentiously[50] dated to before 65 kya[51][52]–and some claims of religious beliefs have been made.[53] They were capable of speech, though it is unclear how complex their language would have been.[54][55]

They likely lived in small groups, lacked sexual division of labour, and put children to work at a very young age. Neanderthals lived in a high-stress environment with high trauma rates, and about 80% died before the age of 40.[56] They had a low population, leading to the accumulation of harmful genes and inbreeding. Interbreeding between Neanderthals and anatomically modern humans was concluded in the 2010 Neanderthal genome project's draft report,[57][58][59] possibly occurring 316–219 kya,[60] and more likely occurring 100 kya and again after 65 kya.[61] Around 1–4% of non-Subsaharan African genomes (North Africans, Eurasians, Oceanians, and Native Americans) derive from Neanderthals,[57][62][63] and about 20% of the Neanderthal genome survives today,[64] but many of the inherited genes may have been detrimental and selected out.[65]

TaxonomyEdit

EtymologyEdit

 
What remains of Kleine Feldhofer Grotte where Neanderthal 1 was discovered

Neanderthals are named after the site they were first identified in, 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").[66][67][68]

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".[69] 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.[70][71][72] However, in 1864, he recommended the genus name also be distinct from humans as he compared the Neanderthal braincase to that of a chimpanzee and argued it was "incapable of moral and theositic conceptions".[73]

 
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/).[74][75][76][77][78] However, in English, "Neanderthal" is also pronounced with the usual th sound (as /niːˈændərθɔːl/).[79]

The German spelling of "Thal" ("valley") was current until 1901; it is now spelt "Tal".[a][80][81] 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.[82] 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.[81]

Research historyEdit

The first Neanderthal remains ever discovered–Engis 2–was in 1829 by Dutch naturalist Philippe-Charles Schmerling in the Schmerling Caves, but he thought it was an ancient human skull.[83] 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 human skull.[84] In 1856, local schoolteacher Johann Carl Fuhlrott recognized bones from Feldhofer Cave in Neander Valley–Neanderthal 1 (the holotype specimen)–as distinct from 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.[85][73] Following Charles Darwin's On the Origin of Species, Fuhlrott and Schaaffhausen argued the bones represented an ancient human form;[27][86][87][73] Schaaffhausen, a social darwinist, believed that humans linearly progressed from savage to civilised, and so concluded that Neanderthals were barbarous cave-dwellers.[27] However, they 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,[88] which stalled Neanderthal research until the end of the century.[27][86]

By the early 20th century, numerous other Neanderthal discoveries were made, establishing Neanderthal as a legitimate species, the most influential specimen La Chapelle-aux-Saints 1. French palaeontologist Marcellin Boule made 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 humans,[89][27] and 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 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][90][86]

By the middle of the century, based on several recent findings of other human ancestors (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 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.[86]

Hybridization between Neanderthals and early humans had been suggested early on,[91] such as by English anthropologist Thomas Huxley in 1890,[92] ethnographer Hans Peder Steensby in 1907,[93] and Coon in 1962.[94] In the early 2000s, several researchers argued in favour of admixture based on supposed hybrid specimens such as Lagar Velho 1[95][96][97][98] and Muierii 1.[99] Neanderthal admixture was found to be present in modern populations in 2010 with the mapping of the first Neanderthal genome sequence.[57]

ClassificationEdit

Hominina

Chimpanzee

Humans

Denisovan from Denisova Cave

Denisovan from Baishiya Karst Cave

Neanderthal from Denisova Cave

Neanderthal from Sidrón Cave

Neanderthal from Vindija Cave

Phylogeny of Hominina based on comparison of ancient proteomes and genomes with those of modern species.[100]

Neanderthals are hominids in the genus Homo, and generally classified as a distinct species, H. neanderthalensis, though sometimes as a subspecies of 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][101] 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,[59][102][103][104] representing a partial biological reproductive barrier between the groups.[59]

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

Neanderthals were more closely related to Denisovans than to humans based on nuclear DNA. However modern humans and Neanderthals share a more recent common mitochondrial lineage than that of Denisovans. 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,[100][105][106][107][108] or the mtDNA of Neanderthals being replaced by that of a modern human lineage deriving from an early H. sapiens wave from Africa.[109] Older specimens show distinct genetic populations of Neanderthals in different regions, but DNA samples from Mezmaiskaya Cave in the Caucasus[108] and Denisova Cave in the Siberian Altai Mountains[60] show apparent replacement of their local populations by a different population also found in Western European sites.

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[110]

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.[111] 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.[110][112][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 ky old Aroeira 3 could represent a transitional phase. Basal and derived morphs could have lived concurrently.[113] The quality of the fossil record greatly increases from 130,000 years ago (kya) onwards,[114] and make up the bulk of known Neanderthal skeletons.[115][116] 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.[117]

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, robustness, and an enlengthenment of the head, which then led to other changes in skull anatomy.[97] However, Neanderthal anatomy was most likely not driven by adapting to cold weather.[28] 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).[110]

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 a human lineage and a Neanderthal/H. heidelbergensis lineage.[118] 800 kya has H. antecessor as the LCA; however, different variations of this model would push the date back to 1 million years ago.[118][24] DNA studies have yielded various results on divergence time, such as 592–182 kya,[22] 553–321 kya,[119] 654–475 kya,[118] 690–550 kya,[66] 765–550 kya,[24][107] 800–520 kya,[120] before 800 kya,[23] and so forth.

Neanderthals and Denisovans are more closely related to each other than they are to humans, meaning the Neanderthal/Denisovan split occurred after their split with humans.[121][107][24][106] 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.[107] 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.[106]

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, 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.[122]

The southernmost find was recorded at Shuqba Cave, Levant,[123] and the easternmost 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.[124] The limit of their northern bound appears to have been 53°N (Bontnewydd, Wales),[125] 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,[126][127][128] but these are more likely attributed to humans.[129] A 2017 study claimed the presence of Homo at the 130 ka Californian Cerutti Mastodon site,[130] but this is highly unlikely.[131][132][133]

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. Populations may have peaked in cold but not extreme intervals, such as marine isotope stages 8 and 6. It is possible their range expanded and contracted as the ice retreated and grew respectively to avoid permafrost areas, residing in certain refuge zones.[134] However, Neanderthals may have preferred a forested landscape.[28]

 
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.[135][106] Various studies, using mtDNA analysis, yield varying effective populations,[134] such as about 1,000 to 5,000;[135] 5,000 to 9,000 remaining constant;[136] or 3,000 to 25,000 steadily increasing until 50,000 BCE before declining until extinction.[137] However, all agree on low population,[134] which may have been up to 10 times smaller than contemporary human populations in Western Europe.[138] Estimates giving a total population in the tens of thousands[106] are contested.[135] 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 maintain its low population. This is consistent with the apparent 150 kya stagnation in Neanderthal technology.[134]

This low population caused a low genetic diversity, inbreeding, and reduced the population's ability to filter out harmful mutations. 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.[139] It is known, however, that the Neanderthals of Sidrón Cave displayed several birth defects.[140]

Based on tropical human hunter-gatherer societies, adult Neanderthals possibly lived on average 25–40 years. 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.[56] However, it has also been estimated that the age pyramids for Neanderthals and Upper Paleolithic humans were the same.[134] Infant mortality was very high, about 43% in northern Eurasia.[141]

AnatomyEdit

BuildEdit

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

Neanderthals had a more robust and stockier build than humans,[33] though still maintained an upright posture;[149] wider and barrel-shaped rib cage; wider pelvis;[112] and proportionally shorter forearms and forelegs.[150][28]

Based on 45 Neanderthals long bones from 14 men and 7 women, the average height was 164 to 168 cm (5.4 to 5.5 ft) for men and 152 to 156 cm (5 ft) for women. For comparison, the average height of 16 Upper Paleolithic and Mesolithic humans was 168.1 cm (5.5 ft) for men and 152.5 cm (5 ft) for women.[33] For Neanderthal weight, samples of 26 specimens found an average of 77.6 kg (171 lb) for men and 66.4 kg (146 lb) for women.[151] Using 76 kg (168 lb), the body mass index for Neanderthal men was calculated to be 2.69–2.82, which in humans correlates to obesity. This indicates a very stocky build.[33]

Body proportions are usually cited as being "hyperarctic" as adaptations to the cold, as they are similar to those of human populations which developed in cold climates[152]–the Neanderthal build is most similar to Eskimos[153]–and shorter limbs equates to higher retention of body heat,[150][152] but Neanderthals from more temperate climates–such as Iberia–still retain the "hyperarctic" physique.[154] 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 humans. It is possible their body proportions and greater muscle mass were adaptations to sprinting as opposed to the endurance-oriented human physique,[28] as shorter limbs reduce moment arm at the limbs, which allows 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.[150][155]

Several adaptations in the leg joints could possibly suggest habitual squatting, which, if the case, was likely done while gathering food.[156]

FaceEdit

 
Le Moustier Neanderthal facial reconstitution at the Neues Museum, Berlin

Neanderthals had a reduced chin, sloping forehead, and large nose[b] 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 humans, and features an occipital bun,[157] 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.[158] They likely also had larger eyes to adapt to the low-light environment.[159]

The large Neanderthal nose and paranasal sinuses were once thought to warm air as it entered the lungs and retain moisture ("nasal radiator hypothesis"),[160] 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 humans.[29][161][160]

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 better resists wear and abrasion,[162][160] 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 humans is now thought to be about the same,[160] about 285 N (64 lbf) and 255 N (57 lbf) in human men and women, respectively.[163]

BrainEdit

The Neanderthal braincase averages 1,600 cm3 (98 in3) for men and 1,300 cm3 (79 in3) for women,[30][31][32] within the possible range of modern humans,[164] which is, on average, 1,270 cm3 (78 in3) for men and 1,130 cm3 (69 in3) for women.[165] The largest Neanderthal brain, Amud 1, was calculated to be 1,736 cm3 (105.9 in3), one of the largest ever recorded in hominids.[166] However, both Neanderthal and human infants measure about 400 cm3 (24 in3), and either Neanderthal brain development sped up or human development slowed down from the last common ancestor.[167]

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 humans,[159] including the cerebellum–operating muscle memory, and possibly language, attention, working memory, social abilities, and thought–the parietal lobesvisuospatial 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.[168] 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 humans are 52% right-dominant, 12% left-dominant, and 36% ambidextrous.[169]

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. 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.[170] 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,[171] though modern Europeans did not evolve light skin and hair until the Holocene.[172]

The DNA of a Neanderthal from Monti Lessini, Valpolicella, Italy, showed depressed activity of the MC1R gene, which is associated with red or blond hair.[173][174] However, like in humans, red was probably not a very common hair color.[170]

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% in men and 20% in women 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 men with high and low BFPs respectively, and 3,115–3,538 and 3,258–3,710 kcal for women. 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 men, and 2,620–3,043 and 2,764–3,215 kcal for women. 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 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.[175]

 
Reconstruction of a Neanderthal woman.[176]

Maximum lifespan, and the timing of adulthood, menopause, and gestation were most likely very similar to modern humans.[134] However, it has been hypothesized that Neanderthals matured faster than modern humans based on the growth rates of teeth and tooth enamel,[177][178] though this is not backed by age biomarkers.[56]

Genetically, Neanderthal-derived alleles near ASB1 and EXOC6 are associated with being an evening person, narcolepsy, and day-time napping. In a survey, people who had archaic haplotypes near CDH6 more frequently reported feeling unenthusiasm and disinterest. These are consistent with the idea that sunlight affects circadian rhythm and mood.[170]

PathologyEdit

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% before reaching adulthood;[179] and an estimated 80% succumbed to their injuries and died before reaching 40.[56] It was thus theorized that Neanderthals employed a risky hunting strategy, though rates of cranial trauma are not significantly different between Neanderthals and Middle Paleolithic humans.[180] Shanidar 1 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. It is unlikely any of these are what ultimately killed him, however.[181]

Likely due to advanced age (60s or 70s), La Chapelle-aux-Saints 1 had signs of Baastrup's disease, affecting the spine, and osteoarthritis.[149] 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 restricts muscle movement, which likely led to his death.[182]

Low population also led to low genetic diversity and probably inbreeding which could have led to inbreeding depression. The 13 inhabitants of Sidrón Cave collectively exhibited 17 birth defects likely due to this.[140]

Neanderthals were likely subject to several infectious diseases and parasites. Humans likely transmitted diseases to them, one likely candidate the stomach bacteria Helicobacter pylori.[183] A Neanderthal at Sidrón Cave showed evidence of a gastrointestinal Enterocytozoon bieneusi infection.[46] The leg bones of La Ferrassie 1 feature lesions which 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 carcinoma.[184]

CultureEdit

Social structureEdit

Group dynamicsEdit

 
Artist's reconstruction of a Neanderthal man and child

Neanderthals likely lived in much smaller and more sparsely distributed groups than Upper Paleolithic humans.[134] Reliable evidence of Neanderthal group composition comes from Sidrón Cave and the footprints at Le Rozel:[142] the former shows 7 adults, 3 adolescents, 2 juveniles, and an infant;[185] whereas the latter, based on footprint size, shows juveniles and adults making up 80–90% of the group.[142]

Children were likely weaned after 5 years – one year later than humans – probably to increase birth spacing. Indicated from various ailments resulting from high stress at a low age, such as stunted growth, children of both sexes were likely put to work directly after weaning.[141] Upon reaching adolescence, an individual would likely have been expected to join in hunting large game.[56]

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).[38] Bands likely moved between certain caves depending on the season, returning to the same locations generation after generation.[186]

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.[38] Analysis of the mtDNA of the Neanderthals of Sidrón Cave showed that the three men belonged to the same maternal lineage, while the three women belonged to different ones. This suggests that women "married out".[187] However, the DNA of a Neanderthal from Denisova Cave shows that her parents were half-siblings,[107] and the inhabitants of Sidrón Cave were likely highly inbred.[140] Neanderthal groups probably rarely exchanged mates given the abundance of harmful genes.[106]

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.[38] 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,[188] 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.[189] Further, 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.[38] However, mapping of the Neanderthal brain indicates they may not have had the cognitive function required for complex social interactions and trade.[159]

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 just men hunting and women and children foraging which is a more efficient food-collecting system. However, in modern human societies, the higher the meat dependency, the higher the division of labour.[38] 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, implying some cultural differences in tasks.[190]

 
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 jewelry–such as a leopard skin or raptor feathers–to display elevated status in the group. The few 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. An apparent cemetery of six or seven individuals at La Ferrassie may indicate they consciously identified as a single group.[38]

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 which 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. In this hypothesis, elderly Neanderthals were given special burial rites for lasting so long.[56]

FoodEdit

Hunting and gatheringEdit

 
Red deer, the preferred game of Neanderthals

Neanderthals were probably an apex predator,[191] and fed predominantly on deer, namely red deer and reindeer, as they were the most abundant game,[192] but also on ibex, wild boar, aurochs, and less frequently mammoth, straight-tusked elephant and woolly rhinoceros.[112][193][194] Isotope studies of Neanderthals from two French sites showed similar profiles to other carnivores, suggesting that these populations ate fresh meat.[195] Analysis of Neanderthal bone collagen from the Croatian Vindija Cave shows nearly all of their protein needs derived from animal meat.[193] 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.[196][28] Younger or wounded animals may have been hunted using traps, projectiles, or pursuit.[196]

Neanderthal diet may have varied significantly region to region, with some communities subsisting primarily on a plant-based diet.[197] Edible plant remains are recorded from several caves.[198] 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.[46] Remnants from the Israeli Amud Cave indicates a diet of figs, palm tree fruits, and various cereals and edible grasses.[199]

Dental tarter from Spy Cave indicates they had a meat-heavy diet including woolly rhinoceros and mouflon sheep, while also regularly consuming mushrooms.[46] 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).[200] Evidence of cooked food plants–mainly legumes and, to a far lesser extent, acorns–were discovered in the Israeli Kebara Cave, possibly gathering plants in spring and fall and hunting in all seasons except fall, though it was probably abandoned in late summer to early fall.[45] 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.[201]

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.[47] The abundance of animal bone fragments at settlements may indicate the making of fat stocks from boiling bone marrow, possibly taken from animals which 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.[47][202] Neanderthal tooth size had a decreasing trend after 100 kya, which could indicate an increased dependence on cooking or the advent of boiling which would soften food.[203]

 
Yarrow growing in Spain

At Sidrón Cave, Neanderthals likely cooked and possibly smoked food,[48] as well as used certain plants–such as yarrow and camomile–as flavouring,[47] 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.[44] 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.[204]

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.[203] It is possible the Neanderthals at Kebara Cave used the shells of the spur-thighed tortoise as containers.[205]

They likely lacked any method of storing food, and had to eat whatever they hunted or foraged immediately.[38]

CompetitionEdit

 
Cave hyaena skeleton

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. In fact, cave hyaena caves are distinguished from Neanderthal caves by the higher abundance of carnivore remains.[192] Nonetheless, cave hyaenas likely stole food and leftovers from Neanderthal campsites, and scavenged on dead Neanderthal bodies.[206]

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, and that pressure from competition was rather high. Cave lions likely targeted horses, large deer and wild cattle; and the leopard primarily reindeer and roe deer; which heavily overlapped with Neanderthal diet. However, given the ferocity of these creatures and the lack of long-range weapons, confrontations likely would have resulted in a group display of yelling, arm waving, or stone throwing; or quickly gathering meat and abandoning a kill.[49]:120–143

CannibalismEdit

There are several examples of Neanderthals practising cannibalism occurring across their range.[143][207] The first undisputed example came from Gran Dolina in 1999,[208] and other examples were found at Sidrón Cave,[187][209] Zafarraya in Spain; and the French Moula-Guercy Cave,[210] 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, the jaw dismembered, and 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.[143][207]

These cannibalistic tendencies have been explained as either ritual defleshing, for nutritional value, or as an act of war. Due to a small amount of cases, and the higher amount 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 historical cases in human history. It has also been suggested evidence of butchering was caused by pre-burial defleshing to prevent scavenging by wild animals or a foul smell. However, it is not uncommon to find Neanderthal remains eaten by an animal.[207]

ArtEdit

 
Proposed Neanderthal jewelry: white-tailed eagle claw with striations at the Neanderthal site of Krapina, Croatia, circa 130,000 BP.[211]

A large number of claims of Neanderthal art, adornment, and structures have been made which would show Neanderthals were capable of symbolic thought or were cognitively comparable to anatomically modern humans. However, many of these are ambiguously attributed as the dating interlaps with anatomically modern human presence in Europe.[51][52] Among many others:

  • Flower pollen on the body of pre-Neanderthal Shanidar 4, Iraq, had in 1975 been argued to be a flower burial,[212] but the pollen could have also been deposited by natural events.[213][214]
  • In 1975, a mostly flat piece of flint with a bone pushed through a hole on the midsection–dating to 32, 40, or 75 kya[215]–has been purported to resemble the upper half of a face, with the bone representing eyes–the Mask of la Roche-Cotard.[216][217] It is contested whether it represents a face, or if it even constitutes as art.[218]
  • Châtelperronian beads have been attributed to Neanderthals, but the dating is uncertain and the beads may have been made by modern humans.[219][220][221][222]
  • Raptor and corvid bones were argued to show evidence of feather plucking by a 2012 study examining 1,699 ancient sites across Eurasia, which the authors controversially took to mean Neanderthals wore bird feathers as personal adornments.[223]
 
The scratched floor of Gorham's Cave
  • Deep scratches on the floor of Gorham's Cave, Gibraltar, were dated to older than 39 kya in 2012, which some have controversially interpreted as Neanderthal abstract art.[224][225]
  • Two 176,000-year-old stalagmite ring structures, several metres wide, were reported in 2016 more than 300 m (980 ft) from the entrance within Bruniquel Cave, France. Being so far inside the cave, this shows a high degree of proficiency in underground environments in Neanderthals.[226] Other red-painted stalagmites in Spain were dated to 65,500 years ago.[227]
  • In 2015, a study argued that a number of 130,000-year-old eagle talons found in a cache near Krapina, Croatia along with Neanderthal bones, had been modified to be used as jewelry.[211][228] A similar talon necklace was reported in 2019 at Cova Foradà in Spain.[229]
  • 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.[230]
  • In 2018, red painted symbols comprising hand stencils, a ladder-shape figure,[227] dots, discs, lines, and representations of animals on the cave walls of several caves across Spain 700 km (430 mi) apart, including La Pasiega,[227] Cave of Maltravieso,[227] Cave of El Castillo,[231] and Doña Trinidad–were dated to be older than 66,000 years ago.[227] This is at least 20,000 years prior to the arrival of anatomically modern humans in western Europe, and demonstrate Neanderthals were capable of symbolic behavior.[227][51][52][232] However, the dating, and thus its attribution to Neanderthals, is contested.[50]
  • In 2018, perforated seashell beads and pigments that are at least 115 kya were found in Cave of Los Aviones in southeastern Spain.[233]
  • In 2018, an engraved flake flint was found in the grave a Neanderthal child in Crimea, Ukraine.[234]

TechnologyEdit

Despite the apparent 150 kya stagnation in Neanderthal innovation,[134] there is evidence that Neanderthal technology was more sophisticated than was previously thought.[54] However, the high frequency of potentially debilitating injuries would have prevented very complex technologies from emerging, as a major injury would have impeded an expert's ability to effectively teach a novice.[179]

Neanderthals made stone tools, and are associated with the Mousterian industry.[34] The Levallois technique they adopted maximizes the cutting surface with the least amount of raw material (p. As a difficult-to-learn process, the technique may have been directly taught generation to generation rather than via purely observational learning.[35] A Levallois point embedded in the vertebrae of an African wild ass indicated that a javelin had been thrown with a parabolic trajectory to disable the animal.[235] They may also be associated with the Châtelperronian 45–40 kya, borrowing tool-making techniques from immigrating humans,[236] though this is highly controversial.[237]

Neanderthal were able to create fire,[36][238][37] and, 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.[39] 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.[38][39] 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; similarly, the cave was likely only inhabited in the winter.[39]

It was long believed that an adhesive (birch bark tar) made by Neanderthals required to follow a complex recipe, and that it thus showed complex cognitive skills and cultural transmission. However, a 2019 study showed it can be made simply by burning birch bark on smooth vertical surfaces, such as a flat, inclined rock.[40]

As opposed to the bone sewing needles and stitching awls of Upper Paleolithic 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.[41][239] Nonetheless, Neanderthals would have needed to cover up most of their body. Contemporary humans would have covered 80–90%.[239][240] 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 in hotter climates did not wear clothes, or their lice were highly specialised.[240]

Remains of Middle Paleolithic stone tools on Greek islands indicate early seafaring by Neanderthals in the Ionian Sea possibly starting as far back as 150,000–200,000 BCE. The oldest stone artefacts from Crete date to 130,000–107,000 BCE, Cephalonia 125,000 BCE, and Zakynthos 110,000–35,000 BCE. Here, they likely employed simple reed boats and made one-day crossings back and forth.[42] Other Mediterranean islands include Sardinia, Melos, Alonnisos, and it is possible they crossed the Strait of Gibraltar[43] and sailed to Naxos (though Naxos may have been connected to land).[241] Their ability to engineer these boats and navigate through open waters speaks to their advanced cognitive and technical skills.[43][241]

They appear to have had some knowledge of medicine. 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.[46] 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.[44] 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.[45]

LanguageEdit

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

The 1983 discovery of a Neanderthal hyoid bone–used in speech production in humans–in Kebara 2 almost identical to that of humans suggests Neanderthals were capable of speech.[55] The prevailing hypothesis for a long time was that speech spontaneously developed very recently in humans,[54] and some argued that the hyoid could have a different usage in Neanderthals, as it is simply used in tongue movement including while chewing. It was once thought Neanderthals were anatomically unable to produce quantal vowels, which are present in all human languages, because they had a large mouth to house the tongue and lacked necessity for a descended larynx[242][243] but this is not a safe assumption to make,[244] and the modern human vocal apparatus and thus vocal repertoire were likely already present in the ancestral H. heidelbergensis.[55]

The degree of language complexity is difficult to establish, but given Neanderthals achieved some technical and cultural complexity, and interbred with humans, it is reasonable to assume they were at least fairly articulate, comparable to humans. A somewhat complex language–possibly using syntax–was likely necessary to survive in their harsh environment, needing to communicate about topics as locations, hunting and gathering, and tool-making techniques.[54][245][246] However, they may have lacked mental synthesis, the human imaginative ability to craft effectively infinite ideas using a finite amount of words, a hallmark of behavioural modernity which appeared by about 70 kya,[247] though behavioural modernity is now believed to have been a process started about 400 kya,[248] and possibly also exhibited in Neanderthals.[249][246]

Genetically, the FOXP2 gene in humans is identified to be very important in speech and language development. FOXP2 is present in Neanderthals,[250] but not the modern human variant.[251] Neurologically, they 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, indicating a stronger ability in modern humans to express language.[252]

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,[114]:158–60 are heavily contested.[253][254][255] 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[256] or to prevent scavengers.[247]

The debate on Neanderthal funerals has been active since the 1908 discovery of La Chapelle-aux-Saints 1 in a small hole in a cave in southwestern France, very controversially attributed to have been buried in a symbolic fashion.[257][253][258]

Another grave at Shanidar Cave was associated with the pollen of several flowers which may have been in bloom at the time of deposition–yarrow, Centaurea, ragwort, grape hyacinth, joint pine, and hollyhock.[259] The medicinal properties of the plants led American archaeologist Ralph Solecki to claim that the man buried was some leader, healer, or shaman.[260] However, it is also possible the pollen was deposited by a small rodent after the man's death.[261]

The grave of Teshik-Tash 1 from Uzbekistan was associated with a circle of ibex horns, which was asserted by Sir Paul Mellars to indicate a ritualistic burial. However, the abundance of ibex horns in the vicinity casts doubt on this.[262]

The grave of a Neanderthal child from Kiik-Koba from 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.[234]

CultsEdit

It was once asserted 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 which killed bears and then ceremoniously arranged the bones. This would be consistent with bear-related rituals of human Arctic hunter gatherers, but the alleged peculiarity of the arrangement could also be well-explained by natural causes,[53][256] 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.[263]

It was also once thought Neanderthals hunted, killed, and cannabalised other Neanderthals to use the skull as the focus of some ritual.[207] 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,[264] putting it forward as evidence of a skull cult.[256] However, it is now thought to have been a result of cave hyaena predation.[265] Though Neanderthals are known to have practiced cannibalism, there is unsubstantial evidence to suggest ritual defleshing.[143]

InterbreedingEdit

Interbreeding with modern humansEdit

 
Reconstruction of the upper Paleolithic human Oase 2 with around 7.3% Neanderthal DNA (4–6 generations back)[266]

The first Neanderthal genome sequence was published in 2010, and strongly indicated interbreeding between Neanderthals and humans.[57][267][268][62] The genomes of all non-sub-Saharan-Africans contain Neanderthal DNA.[57][269][59][63] Various estimates exist for the proportion: 1–4% in Eurasians,[57] 3.4–7.9%,[270] and 1.8–2.4% in Europeans and 2.3–2.6% in East Asians.[271] Such low percentages indicate infrequent interbreeding.[272] Of the inherited Neanderthal genome, 25% in Europeans and 32% in East Asians may be related to viral immunity.[273] In all, approximately 20% of the Neanderthal genome appears to have survived in the modern human gene pool.[64] However, due to their small population and resulting reduced effectivity of natural selection, Neanderthals mutated several weakly harmful mutations, which were introduced to and slowly selected out of the much larger human population; the initial hybridized 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.[65]

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.[274] However, the approximately 40 ka anatomically-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,[266] but this hybrid Romanian population does not appear to have made a substantial contribution to the genomes of later Europeans.[266]

In 2016, the DNA of Altai Neanderthals showed evidence of interbreeding 100 kya, and interbreeding with a later dispersal of humans may have occurred as late as 120 kya in places such as the Levant.[61] Palaeontologically, the earliest human remains outside of Africa occurs at Misliya Cave 194–177 kya, and Skhul and Qafzeh 120–90 kya.[275] 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.[60] Whatever the case, these first interbreeding event has not left any trace in modern human genomes.[276]

Due to the absence of Neanderthal-derived mtDNA (which is passed on from mother to child) in modern populations,[104][277][66] it has been suggested that the progeny of Neanderthal women who mated with modern human men were either rare, absent, or sterile–that is to say, admixture stems from the progeny of Neanderthal men with modern human women.[278][277][102][58] 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 female, or the Neanderthal Y-chromosome was not compatible with humans and became extinct.[279]

Detractors of the interbreeding model argue that the genetic similarity is only a remnant of a common ancestor instead of interbreeding.[280] 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.[281]

Interbreeding with DenisovansEdit

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

Though nDNA confirms that Neanderthals and Denisovans are more closely related to each other than they are to humans, Neanderthals and humans share a more recent maternally-transmitted mtDNA common ancestor, possibly due to interbreeding between Denisovans and some unknown hominid. The 400 kya Neanderthal-like hominids 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 hominids, which could have interbred with Denisovans.[283] This is also used to explain an approximately 124,000-year-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.[60]

Sequencing of the genome of a Denisovan from Denisova Cave has shown that 17% of its genome derives from Neanderthal.[105] This Neanderthal DNA more closely resembled that of a 120,000-year-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.[107]

For the 90 ky 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 groups, and Neanderthal migration across Eurasia likely occurred sometime after 120 kya.[284]

ExtinctionEdit

 
Map emphasizing the Ebro river in northern Spain

Neanderthals are thought to have died out between 41 and 39 kya.[285][10][11][12][13] Though some Neanderthals in Gibraltar were dated to later than this–such as Zafarraya (30 kya)[286] and Gorham's Cave (28 kya)[287]–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[127] is loosely supported by Mousterian stone tools dating to 34–31 kya at a time when humans may not yet have colonised the northern reaches of Europe;[129] however, human remains are known from the northern Siberian Mamontovaya Kurya site dating to 40 kya.[288]

Whatever the cause of their extinction, Neanderthals were replaced by humans, indicated by near full replacement of Mousterian stone technology with 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[289] and Britain,[290] and this migration of humans replaced Neanderthals.[10] A 2019 reanalysis of 210 ky old skull fragments from the Greek Apidima Cave assumed to have belonged to a Neanderthal concluded that they belonged to a human, and DNA evidence indicates human 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 humans were replaced by Neanderthals until their return about 40 kya.[291]

Competition with modern humansEdit

 
Replacement of Neanderthals by early modern humans.

Boule was the first person to suggest humans forcefully took Europe from the Neanderthals.[89] Human expansion and Neanderthal contraction are correlated, possibly due to the competitive exclusion principle with humans outcompeting Neanderthals.[14] However, largely hominid-free tropical Asia was colonised by 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 human expansion for some time.[292]

Jared Diamond in his book The Third Chimpanzee said competitive replacement often occurs in human history when a more technologically advanced culture (humans) meets a less advanced culture (Neanderthals).[15] Though Neanderthals likely exhibited modern behaviour, discrediting arguments of plain human superiority, the spread of grasslands and open steppe would have made human projectile weapons much more effective over Neanderthal short-range spears which were adapted to a forest environment;[16] humans could push into colder areas with bigger game wearing their fitted clothes, which were more effective at insulating than Neanderthal ponchos;[293] 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, since most of their stone artifacts were drawn from within 5 km (3.1 mi), likely restricted themselves to more local sources.[38][294]

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 humans, about 50–45 kya, evolved the whites of the eyes to allow for more effective non-verbal communications with wolves, and this gave humans an advantage over hunting. She also claims that Neanderthals did not have very prominent whites of the eyes, like the rest of the animal kingdom.[49]:214–226

Shanidar 3 died from complications from a stab wound, likely originating from a light-weight, long-range projectile, a technology only humans had, which implies Neanderthal/human violence.[295]

Climate changeEdit

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.[296] Their ultimate extinction coincides with Heinrich event 4, a period of intense cold and dry climate, and future Heinrich events are also associated with massive cultural turnovers where European human populations collapsed.[17][18] This time was also characterised by the spread of grasslands and open steppe, which Neanderthal short-range technology was unsuited for, impeding their ability to hunt.[16]

 
Graphic of the ash cloud from the Campanian Ignimbrite Eruption

Neanderthal 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 led to their final demise.[19][297]

DiseaseEdit

Humans may have introduced African diseases to Neanderthals which contributed 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.[140]

In New Guinea, due to cannibalistic practices, the population was decimated from transmissible spongiform encephalopathies (kuru) spread by ingestion of contaminated brains or contact with infected tools. A similar disease could have quickly spread through the small Neanderthal populations even through minimal contact with each other, given its high virulence and Neanderthal cannibalistic tendencies.[20]

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 somewhat 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. ^ There are modern humans with noses as wide as those of Neanderthals and modern humans with similar nose lengths, but none with both Neanderthal nose width and nose length.
  3. ^ 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.[282]

ReferencesEdit

  1. ^ a b Haeckel, E. (1895). Systematische Phylogenie: Wirbelthiere (in German). p. 601.
  2. ^ Schwalbe, G. (1906). Studien zur Vorgeschichte des Menschen [Studies on the history of man] (in German). Stuttgart, E. Nägele. doi:10.5962/bhl.title.61918. hdl:2027/uc1.b4298459.
  3. ^ Klaatsch, H. (1909). "Preuves que l'Homo Mousteriensis Hauseri appartient au type de Neandertal" [Evidence that Homo Mousteriensis Hauseri belongs to the Neanderthal type]. L'Homme Préhistorique (in French). 7: 10–16.
  4. ^ Romeo, Luigi (1979). Ecce Homo!:A Lexicon of Man. Amsterdam: John Benjamins Publishing Company. p. 92. ISBN 978-9027220066 – via Google Books (ebook).
  5. ^ a b c d e McCown, T.; Keith, A. (1939). The stone age of Mount Carmel. The fossil human remains from the Levalloisso-Mousterian. 2. Clarenden Press.
  6. ^ Szalay, F. S.; Delson, E. (2013). Evolutionary history of the Primates. Academic Press. p. 508. ISBN 978-1-4832-8925-0.
  7. ^ Wells, J. (2008). Longman Pronunciation Dictionary (3rd ed.). Pearson Longman. ISBN 978-1-4058-8118-0.
  8. ^ Duden - Das Aussprachewörterbuch: Betonung und Aussprache von über 132.000 Wörtern und Namen [Duden - The pronunciation dictionary: emphasis and pronunciation of over 132,000 words and names] (in German) (7th ed.). Bibliographisches Institut GmbH. 2015. p. 625. ISBN 978-3-411-91151-6.
  9. ^ a b c d Pääbo, S. (2014). Neanderthal Man: In Search of Lost Genomes. New York: Basic Books. p. 237.
  10. ^ a b c Higham, T.; Douka, K.; Wood, R.; Ramsey, C. B.; Brock, F.; Basell, L.; Camps, M.; Arrizabalaga, A.; Baena, J.; Barroso-Ruíz, C.; C. Bergman; C. Boitard; P. Boscato; M. Caparrós; N.J. Conard; C. Draily; A. Froment; B. Galván; P. Gambassini; A. Garcia-Moreno; S. Grimaldi; P. Haesaerts; B. Holt; M.-J. Iriarte-Chiapusso; A. Jelinek; J.F. Jordá Pardo; J.-M. Maíllo-Fernández; A. Marom; J. Maroto; M. Menéndez; L. Metz; E. Morin; A. Moroni; F. Negrino; E. Panagopoulou; M. Peresani; S. Pirson; M. de la Rasilla; J. Riel-Salvatore; A. Ronchitelli; D. Santamaria; P. Semal; L. Slimak; J. Soler; N. Soler; A. Villaluenga; R. Pinhasi; R. Jacobi (2014). "The timing and spatiotemporal patterning of Neanderthal disappearance". Nature. 512 (7514): 306–09. Bibcode:2014Natur.512..306H. doi:10.1038/nature13621. PMID 25143113. We show that the Mousterian [the Neanderthal tool-making tradition] ended by 41,030–39,260 calibrated years BP (at 95.4% probability) across Europe. We also demonstrate that succeeding 'transitional' archaeological industries, one of which has been linked with Neanderthals (Châtelperronian), end at a similar time.
  11. ^ a b Higham, T. (2011). "European Middle and Upper Palaeolithic radiocarbon dates are often older than they look: problems with previous dates and some remedies". Antiquity. 85 (327): 235–49. doi:10.1017/s0003598x00067570. Few events of European prehistory are more important than the transition from ancient to modern humans about 40 kya, a period that unfortunately lies near the limit of radiocarbon dating. This paper shows that as many as 70 per cent of the oldest radiocarbon dates in the literature may be too young, due to contamination by modern carbon.
  12. ^ a b c Pinhasi, R.; Higham, T. F. G.; Golovanova, L. V.; Doronichev, V. B. (2011). "Revised age of late Neanderthal occupation and the end of the Middle Paleolithic in the northern Caucasus". Proceedings of the National Academy of Sciences. 108 (21): 8, 611–8, 616. Bibcode:2011PNAS..108.8611P. doi:10.1073/pnas.1018938108. PMC 3102382. PMID 21555570. The direct date of the fossil (39,700 ± 1,100 14C BP) is in good agreement with the probability distribution function, indicating at a high level of probability that Neanderthals did not survive at Mezmaiskaya Cave after 39 kya cal BP. [...] This challenges previous claims for late Neanderthal survival in the northern Caucasus. [...] Our results confirm the lack of reliably dated Neanderthal fossils younger than ≈40 kya cal BP in any other region of Western Eurasia, including the Caucasus.
  13. ^ a b c Galván, B.; Hernández, C. M.; Mallol, C.; Mercier, N.; Sistiaga, A.; Soler, V. (2014). "New evidence of early Neanderthal disappearance in the Iberian Peninsula". Journal of Human Evolution. 75: 16–27. doi:10.1016/j.jhevol.2014.06.002. PMID 25016565.
  14. ^ a b Banks, W. E.; d'Errico, F.; Peterson, A. T.; Kageyama, M.; Sima, A.; Sánchez-Goñi, M. (2008). "Neanderthal extinction by competitive exclusion". PLOS ONE. 3 (12): e3972. Bibcode:2008PLoSO...3.3972B. doi:10.1371/journal.pone.0003972. PMC 2600607. PMID 19107186.
  15. ^ a b Diamond, J. (1992). The Third Chimpanzee: The Evolution and Future of the Human Animal. Harper Collins. pp. 45–52. ISBN 978-0-06-098403-8.
  16. ^ a b c d Finlayson, C.; Carrión, J. S. (2007). "Rapid ecological turnover and its impact on Neanderthal and other human populations". Trends in Ecology and Evolution. 22 (4): 213–222. doi:10.1016/j.tree.2007.02.001. PMID 17300854.
  17. ^ a b Bradtmöller, M.; Pastoors, A.; Weninger, B.; Weninger, G. (2012). "The repeated replacement model – Rapid climate change and population dynamics in Late Pleistocene Europe". Quaternary International. 247: 38–49. Bibcode:2012QuInt.247...38B. doi:10.1016/j.quaint.2010.10.015.
  18. ^ a b Wolf, D.; Kolb, T.; Alcaraz-Castaño, M.; Heinrich, S. (2018). "Climate deteriorations and Neanderthal demise in interior Iberia". Scientific Reports. 8 (1): 7,048. Bibcode:2018NatSR...8.7048W. doi:10.1038/s41598-018-25343-6. PMC 5935692. PMID 29728579.
  19. ^ a b c Black, B. A.; Neely, R. R.; Manga, M. (2015). "Campanian Ignimbrite volcanism, climate, and the final decline of the Neanderthals" (PDF). Geology. 43 (5): 411–414. Bibcode:2015Geo....43..411B. doi:10.1130/G36514.1.
  20. ^ a b Underdown, S. (2008). "A potential role for transmissible spongiform encephalopathies in Neanderthal extinction". Medical Hypotheses. 71 (1): 4–7. doi:10.1016/j.mehy.2007.12.014. PMID 18280671.
  21. ^ a b Sullivan, A. P.; de Manuel, M.; Marques-Bonet, T.; Perry, G. H. (2017). "An evolutionary medicine perspective on Neandertal extinction" (PDF). Journal of Human Evolution. 108: 62–71. doi:10.1016/j.jhevol.2017.03.004. PMID 28622932.
  22. ^ a b c Stringer, C. (2012). "The status of Homo heidelbergensis (Schoetensack 1908)". Evolutionary Anthropology. 21 (3): 101–107. doi:10.1002/evan.21311. PMID 22718477.
  23. ^ a b Gómez-Robles, A. (2019). "Dental evolutionary rates and its implications for the Neanderthal–modern human divergence". Science Advances. 5 (5): eaaw1268. doi:10.1126/sciadv.aaw1268. PMC 6520022. PMID 31106274.
  24. ^ a b c d e Meyer, M.; Arsuaga, J.; de Filippo, C.; Nagel, S. (2016). "Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins". Nature. 531 (7595): 504–507. Bibcode:2016Natur.531..504M. doi:10.1038/nature17405. PMID 26976447.
  25. ^ a b Klein, R. G. (1983). "What Do We Know About Neanderthals and Cro-Magnon Man?". Anthropology. 52 (3): 386–392. JSTOR 41210959.
  26. ^ a b c Papagianni, D.; Morse, M. A. (2015). "Still with us?". Neanderthals Rediscovered: How Modern Science Is Rewriting Their Story. Thames and Hudson. ISBN 978-0-500-77311-6.
  27. ^ a b c d e f g h i Drell, J. R. R. (2000). "Neanderthals: a history of interpretation". Oxford Journal of Archaeology. 19 (1): 1–24. doi:10.1111/1468-0092.00096.
  28. ^ a b c d e f Stewart, J.R.; García-Rodríguez, O.; Knul, M.V.; Sewell, L.; Montgomery, H.; Thomas, M.G.; Diekmann, Y. (2019). "Palaeoecological and genetic evidence for Neanderthal power locomotion as an adaptation to a woodland environment". Quaternary Science Reviews. 217: 310–315. Bibcode:2019QSRv..217..310S. doi:10.1016/j.quascirev.2018.12.023.
  29. ^ a b de Azevedo, S.; González, M. F.; Cintas, C.; Ramallo, V.; Quinto-Sánchez, M.; Márquez, F.; Hünemeier, T.; Paschetta, C.; Ruderman, A.; Navarro, P.; Pazos, B. A.; Silva de Cerqueira, C. C.; Velan, O.; Ramírez-Rozzi, F.; Calvo, N.; Castro, H. G.; Paz, R. R.; González-José, R. (2017). "Nasal airflow simulations suggest convergent adaptation in Neanderthals and modern humans". Proceedings of the National Academy of Sciences. 114 (47): 12, 442–12, 447. doi:10.1073/pnas.1703790114. PMC 5703271. PMID 29087302.
  30. ^ a b Stringer, C. (1984). "Human evolution and biological adaptation in the Pleistocene". In Foley, R. (ed.). Hominid evolution and community ecology. Academic Press. ISBN 978-0-12-261920-5.
  31. ^ a b Holloway, R. L. (1985). "The poor brain of Homo sapiens neanderthalensis: see what you please". In Delson, E. (ed.). Ancestors: The hard evidence. Alan R. Liss. ISBN 978-0-471-84376-4.
  32. ^ a b Amano, H.; Kikuchi, T.; Morita, Y.; Kondo, O.; Suzuki, H.; et al. (2015). "Virtual Reconstruction of the Neanderthal Amud 1 Cranium" (PDF). American Journal of Physical Anthropology. 158 (2): 185–97. doi:10.1002/ajpa.22777. hdl:10261/123419. PMID 26249757.
  33. ^ a b c d Helmuth H (1998). "Body height, body mass and surface area of the Neanderthals". Zeitschrift für Morphologie und Anthropologie. 82 (1): 1–12. PMID 9850627.
  34. ^ a b Shaw, I.; Jameson, R., eds. (1999). A Dictionary of Archaeology. Blackwell. p. 408. ISBN 978-0-631-17423-3.
  35. ^ a b Lycett, S. J.; von Cramon-Taubadel, N. (2013). "A 3D morphometric analysis of surface geometry in Levallois cores: patterns of stability and variability across regions and their implications". Journal of Archaeological Science. 40 (3): 1508–1517. doi:10.1016/j.jas.2012.11.005.
  36. ^ a b Sorensen, A. C.; Claud, E.; Soressi, M. (July 19, 2018). "Neandertal fire-making technology inferred from microwear analysis". Scientific Reports. 8 (1): 1–16. doi:10.1038/s41598-018-28342-9. ISSN 2045-2322.
  37. ^ a b Brittingham, A.; Hren, M. T.; Hartman, G.; Wilkinson, K. N.; Mallol, C.; Gasparyan, B.; Adler, D. S. (2019). "Geochemical Evidence for the Control of Fire by Middle Palaeolithic Hominins". Scientific Reports. 9 (15, 368): 15368. doi:10.1038/s41598-019-51433-0. PMC 6814844. PMID 31653870.
  38. ^ a b c d e f g h i j Hayden, B. (2012). "Neandertal social structure?". Oxford Journal of Archaeology. 31 (1): 1–26. doi:10.1111/j.1468-0092.2011.00376.x.
  39. ^ a b c d Kedar, Yafit; Barkai, Ran (2019). "The Significance of Air Circulation and Hearth Location at Paleolithic Cave Sites". Open Quaternary. 5. doi:10.5334/oq.52.
  40. ^ a b Schmidt, P.; Blessing, M.; Rageot, M.; Iovita, R.; Pfleging, J.; Nickel, K. G.; Righetti, L.; Tennie, C. (2019). "Birch tar production does not prove Neanderthal behavioral complexity". Proceedings of the National Academy of Sciences. 116 (36): 17, 707–17, 711. doi:10.1073/pnas.1911137116. PMC 6731756. PMID 31427508.
  41. ^ a b Hoffecker, J. F. (2009). "The spread of modern humans in Europe". Proceedings of the National Academy of Sciences. 106 (38): 16040–16045. Bibcode:2009PNAS..10616040H. doi:10.1073/pnas.0903446106. PMC 2752585. PMID 19571003.
  42. ^ a b Ferentinos, G.; Gkioni, M.; Geraga, M.; Papatheodorou, G. (2012). "Early seafaring activity in the southern Ionian Islands, Mediterranean Sea". Journal of Archaeological Science. 39 (7): 2167–2176. doi:10.1016/j.jas.2012.01.032.
  43. ^ a b c Strasser, T. F.; Runnels, C.; Wegmann, K. W.; Panagopoulou, E. (2011). "Dating Palaeolithic sites in southwestern Crete, Greece". Journal of Quaternary Science. 26 (5): 553–560. Bibcode:2011JQS....26..553S. doi:10.1016/j.jas.2012.01.032.
  44. ^ a b c Buckley, S.; Hardy, K.; Huffman, M. (2013). "Neanderthal Self-Medication in Context". Antiquity. 87 (337): 873–878. doi:10.1017/S0003598X00049528.
  45. ^ a b c Lev, E.; Kislev, M. E.; Bar-Yosef, O. (2005). "Mousterian vegetal food in Kebara Cave, Mt. Carmel". Journal of Archaeological Science. 32 (3): 475–484. doi:10.1016/j.jas.2004.11.006.
  46. ^ a b c d e Weyrich, Laura S.; Duchene, Sebastian; Soubrier, Julien; Arriola, Luis; Llamas, Bastien; Breen, James; Morris, Alan G.; Alt, Kurt W.; Caramelli, David; Dresely, Veit; Farrell, Milly; Farrer, Andrew G.; Francken, Michael; Gully, Neville; Haak, Wolfgang; Hardy, Karen; Harvati, Katerina; Held, Petra; Holmes, Edward C.; Kaidonis, John; Lalueza-Fox, Carles; de la Rasilla, Marco; Rosas, Antonio; Semal, Patrick; Soltysiak, Arkadiusz; Townsend, Grant; Usai, Donatella; Wahl, Joachim; Huson, Daniel H.; et al. (2017). "Neanderthal behaviour, diet, and disease inferred from ancient DNA in dental calculus". Nature. 544 (7650): 357–361. Bibcode:2017Natur.544..357W. doi:10.1038/nature21674. hdl:10261/152016. PMID 28273061.
  47. ^ a b c d Krief, S.; Daujeard, C.; Moncel, M.; Lamon, N.; Reynolds, V. (2015). "Flavouring food: the contribution of chimpanzee behaviour to the understanding of Neanderthal calculus composition and plant use in Neanderthal diets". Antiquity. 89 (344): 464–471. doi:10.15184/aqy.2014.7.
  48. ^ a b Hardy, K.; Buckley, S.; Collins, M. J.; Estalrrich, A. (2012). "Neanderthal Medics? Evidence for Food, Cooking, and Medicinal Plants Entrapped in Dental Calculus". The Science of Nature. 99 (8): 617–626. Bibcode:2012NW.....99..617H. doi:10.1007/s00114-012-0942-0. PMID 22806252.
  49. ^ a b c Shipman, P. (2015). "How Humans and Their Dogs Drove Neanderthals to Extinction". The invaders : how humans and their dogs drove Neanderthals to extinction. Harvard University Press. ISBN 978-0-674-42538-5. JSTOR j.ctvjf9zbs.
  50. ^ a b Aubert, M.; Brumm, A.; Huntley, J. (2018). "Early dates for 'Neanderthal cave art' may be wrong". Journal of Human Evolution. 125: 215–217. doi:10.1016/j.jhevol.2018.08.004. PMID 30173883.
  51. ^ a b c Pike, A. W.; Hoffmann, D. L.; Pettitt, P. B.; García-Diez, M.; Zilhão, J. (2017). "Dating Palaeolithic cave art: Why U–Th is the way to go" (PDF). Quaternary International. 432: 41–49. Bibcode:2017QuInt.432...41P. doi:10.1016/j.quaint.2015.12.013.
  52. ^ a b c d D. L. Hoffmann; C. D. Standish; M. García-Diez; P. B. Pettitt; J. A. Milton; J. Zilhão; J. J. Alcolea-González; P. Cantalejo-Duarte; H. Collado; R. de Balbín; M. Lorblanchet; J. Ramos-Muñoz; G.-Ch. Weniger; A. W. G. Pike (2018). "U-Th dating of carbonate crusts reveals Neandertal origin of Iberian cave art". Science. 359 (6378): 912–15. Bibcode:2018Sci...359..912H. doi:10.1126/science.aap7778. PMID 29472483.
  53. ^ a b Wunn, I. (2000). "Beginning of Religion". Numen. 47 (4): 417–452. doi:10.1163/156852700511612.
  54. ^ a b c d Dediu, D.; Levinson, S. C. (2018). "Neanderthal language revisited: not only us". Current Opinion in Behavioral Sciences. 21: 49–55. doi:10.1016/j.cobeha.2018.01.001. hdl:21.11116/0000-0000-1667-4.
  55. ^ a b c d D’Anastasio, R.; Wroe, S.; Tuniz, C.; Mancini, L.; Cesana, D. T. (2013). "Micro-Biomechanics of the Kebara 2 hyoid and its implications for speech in Neanderthals". PLOS ONE. 8 (12): e82261. Bibcode:2013PLoSO...882261D. doi:10.1371/journal.pone.0082261. PMC 3867335. PMID 24367509.
  56. ^ a b c d e f Trinkaus, E. (1995). "Neanderthal mortality patterns". Journal of Archaeological Science. 22 (1): 121–142. doi:10.1016/S0305-4403(95)80170-7.
  57. ^ a b c d e f Green, R. E.; Krause, J.; Briggs, A. W.; Maricic, T.; Stenzel, U.; Kircher, M.; Patterson, N.; Li, H.; Zhai, W.; Fritz, M. H. Y.; Hansen, N. F.; Durand, E. Y.; Malaspinas, A. S.; Jensen, J. D.; Marques-Bonet, T.; Alkan, C.; Prüfer, K.; Meyer, M.; Burbano, H. A.; Good, J. M.; Schultz, R.; Aximu-Petri, A.; Butthof, A.; Hober, B.; Hoffner, B.; Siegemund, M.; Weihmann, A.; Nusbaum, C.; Lander, E. S.; Russ, C.; Novod, N.; Affourtit, J.; Egholm, M.; Verna, C.; Rudan, P.; Brajkovic, D.; Kucan, Z.; Gusic, I.; Doronichev, V. B.; Golovanova, L. V.; Lalueza-Fox, C.; de la Rasilla, M.; Fortea, J.; Rosas, A.; Schmitz, R. W.; Johnson, P. L. F.; Eichler, E. E.; Falush, D.; Birney, E.; Mullikin, J. C.; Slatkin, M.; Nielsen, R.; Kelso, J.; Lachmann, M.; Reich, D.; Pääbo, S. (2010). "A Draft Sequence of the Neandertal Genome". Science. 328 (5979): 710–722. Bibcode:2010Sci...328..710G. doi:10.1126/science.1188021. PMC 5100745. PMID 20448178.
  58. ^ a b Sankararaman, S.; Mallick, S.; Patterson, N.; Reich, D. (2016). "The Combined Landscape of Denisovan and Neanderthal Ancestry in Present-Day Humans". Current Biology. 26 (9): 1, 241–1, 247. doi:10.1016/j.cub.2016.03.037. PMC 4864120. PMID 27032491.
  59. ^ a b c d Sankararaman, S.; Mallick, S.; Dannemann, M.; Prüfer, K.; Kelso, J.; Pääbo, S.; Patterson, N.; Reich, D. (2014). "The genomic landscape of Neanderthal ancestry in present-day humans". Nature. 507 (7492): 354–357. Bibcode:2014Natur.507..354S. doi:10.1038/nature12961. PMC 4072735. PMID 24476815.
  60. ^ a b c d Peyrégne, Stéphane; et al. (2019). "Nuclear DNA from two early Neandertals reveals 80 ka of genetic continuity in Europe". Science Advances. 5 (6): eaaw5873. Bibcode:2019SciA....5.5873P. doi:10.1126/sciadv.aaw5873. PMC 6594762. PMID 31249872.
  61. ^ a b Kuhlwilm, M. (2016). "Ancient gene flow from early modern humans into Eastern Neanderthals". Nature. 530 (7591): 429–433. Bibcode:2016Natur.530..429K. doi:10.1038/nature16544. PMC 4933530. PMID 26886800.
  62. ^ a b Sánchez-Quinto, F.; Botigué, L. R.; Civit, S.; Arenas, C.; Ávila-Arcos, M. C.; Bustamante, C. D.; Comas, D.; Lalueza-Fox, C.; Caramelli, D. (2012). "North African Populations Carry the Signature of Admixture with Neandertals". PLoS ONE. 7 (10): e47765. Bibcode:2012PLoSO...747765S. doi:10.1371/journal.pone.0047765. PMC 3474783. PMID 23082212.
  63. ^ a b Fu, Q.; Li, H.; Moorjani, P.; Jay, F.; Slepchenko, S. M.; Bondarev, A. A.; Johnson, P. L. F.; Aximu-Petri, A.; Prüfer, K.; de Filippo, C.; Meyer, M.; Zwyns, Ni.; Salazar-García, D. C.; Kuzmin, Y. V.; Keates, S. G.; Kosintsev, P. A.; Razhev, D. I.; Richards, M. P.; Peristov, N. V.; Lachmann, M.; Douka, K.; Higham, T. F. G.; Slatkin, M.; Hublin, J.-J.; Reich, D.; Kelso, J.; Viola, T. B.; Pääbo, S. (2014). "Genome sequence of a 45,000-year-old modern human from western Siberia". Nature. 514 (7523): 445–449. Bibcode:2014Natur.514..445F. doi:10.1038/nature13810. PMC 4753769. PMID 25341783.
  64. ^ a b Vernot, B.; Akey, J. M. (2014). "Resurrecting Surviving Neandertal Lineages from Modern Human Genomes". Science. 343 (6174): 1, 017–1, 021. Bibcode:2014Sci...343.1017V. doi:10.1126/science.1245938. PMID 24476670.
  65. ^ a b Juric, I.; Aeschbacher, S.; Coop, G. (2016). "The Strength of Selection against Neanderthal Introgression". PLoS Genetics. 12 (11): e1006340. doi:10.1371/journal.pgen.1006340. PMC 5100956. PMID 27824859.
  66. ^ a b c Krings, M.; Stone, A.; Schmitz, R. W.; Krainitzki, H.; Stoneking, M.; Pääbo, S. (1997). "Neandertal DNA Sequences and the Origin of Modern Humans". Cell. 90 (1): 19–30. doi:10.1016/s0092-8674(00)80310-4. PMID 9230299.
  67. ^ Krings, M.; Geisert, H.; Schmitz, R. W.; Krainitzki, H.; Pääbo, S. (1999). "DNA sequence of the mitochondrial hypervariable region II from the Neandertal type specimen". Proceedings of the National Academy of Sciences. 96 (10): 5581–5585. Bibcode:1999PNAS...96.5581K. doi:10.1073/pnas.96.10.5581. PMC 21903. PMID 10318927.
  68. ^ Beerli, P.; Edwards, S. V. (2003). "When did Neanderthals and modern humans diverge?". Evolutionary Anthropology: Issues, News, and Reviews. 11 (S1): 60–63. doi:10.1002/evan.10058.
  69. ^ Vogt, K. C. (1864). Lectures on Man: His Place in Creation, and in the History of the Earth. London: Longman, Green, Longman and Roberts. pp. 302, 473.
  70. ^ King, W. (1864). "On the Neanderthal Skull, or reasons for believing it to belong to the Clydian Period and to a species different from that represented by man". Report of the British Association for the Advancement of Science, Notices and Abstracts, Newcastle-upon-Tyne, 1863: 81–82 – via Biodiversity Heritage Library.
  71. ^ Murray, J.; Nasheuer, H. P.; Seoighe, C.; McCormack, G. P.; Williams, D. M.; Harper, D. A. T. (2015), "The Contribution of William King to the Early Development of Palaeoanthropology", Irish Journal of Earth Sciences, 33: 1–16, doi:10.3318/ijes.2015.33.1, JSTOR 10.3318/ijes.2015.33.1
  72. ^ Winner, A. K. (1964). "Terminology". Current Anthropology. 5 (2): 119–122. doi:10.1086/200469. JSTOR 2739959.
  73. ^ a b c King, W. (1864). "The reputed fossil man of the Neanderthal" (PDF). Quarterly Journal of Science. 1: 96.
  74. ^ The Oxford Illustrated Dictionary. Oxford University Press. 1976 [1975]. p. 564.
  75. ^ "Neanderthal adjective – definition in British English Dictionary & Thesaurus". Cambridge Dictionaries Online. Cambridge University Press. January 8, 2013. Archived from the original on July 8, 2012. Retrieved January 22, 2013.
  76. ^ "Neanderthal—meaning in the Cambridge English Dictionary". Cambridge Dictionary. Retrieved September 19, 2019.
  77. ^ "Neanderthal". Oxford Learner's Dictionaries. Retrieved October 29, 2019.
  78. ^ Pollet, C. J. (1991). "...And Etymology". Science News. 140 (12): 191. doi:10.2307/3975867. JSTOR 3975867.
  79. ^ "Neanderthal". Dictionary.com. Retrieved January 22, 2013.
  80. ^ a b Howell, F. Clark (1957). "The evolutionary significance of variation and varieties of 'Neanderthal' man". The Quarterly Review of Biology. 32 (4): 330–47. doi:10.1086/401978. JSTOR 2816956. PMID 13506025.
  81. ^ a b "Neandertal oder Neanderthal? Was ist denn nun richtig" [Neandertal or Neanderthal? Which is right?]. Kreisstadt Mettmann. Retrieved February 1, 2017. Heute sollten Ortsbezeichnungen das 'Neandertal' ohne 'h' bezeichnen. Alle Namen, die sich auf den prähistorischen Menschen beziehen, führen das 'h'. [Today one should write for place names 'Neandertal' without an 'h'. All names related to the prehistoric humans keep the 'h'.]
  82. ^ "Neanderthal". Wiley-Blackwell Encyclopedia of Human Evolution. Chichester, West Sussex: Wiley-Blackwell. 2013.
  83. ^ Schmerling, P. (1834). Recherches sur les ossemens fossiles découverts dans les cavernes de la province de Liége [Research on the fossil specimens discovered in the caves of Liège]. P. J. Collardin. pp. 30–32.
  84. ^ Menez, A. (2018). "Custodian of the Gibraltar skull: the history of the Gibraltar Scientific Society". Earth Sciences History. 37 (1): 34–62. doi:10.17704/1944-6178-37.1.34.
  85. ^ Schaaffhausen, H. (1858). "Zur Kenntnis der ältesten Rassenschädel". Archiv für Anatomie, Physiologie und Wissenschaftliche Medicin: 453–478.
  86. ^ a b c d Schlager, S.; Wittwer-Backofen, U. (2015). "Images in paleonthropology: facing our ancestors". In Henke, W.; Tattersall, I. (eds.). Handbook of Paleoanthropology. Springer-Verlag Berlin Heidelberg. pp. 1, 019–1, 027. doi:10.1007/978-3-642-39979-4_70. ISBN 978-3-642-39978-7.
  87. ^ Fuhlrott, J. C. (1859). "Menschliche Überreste aus einer Felsengrotte des Düsselthales" (PDF). Verh Naturhist Ver Preuss Rheinl (in German). 16: 131–153.
  88. ^ Virchow, R. (1872). "Untersuchung des Neanderthal-Schädels" [Examinations on the Neandertal skull]. Verh Berl Anthrop Ges. 4: 157–165.
  89. ^ a b Boule, M. (1911). L'homme fossile de La Chapelle-aux-Saints [Fossil man from La Chapelle-aux-Saints] (in French). Masson. pp. 1–62.
  90. ^ Sommer, M. (2006). "Mirror, Mirror on the Wall: Neanderthal as Image and 'Distortion' in Early 20th-Century French Science and Press" (PDF). Social Studies of Science. 36 (2): 207–240. doi:10.1177/0306312706054527.
  91. ^ Cairney, C. T. (1989). Clans and Families of Ireland and Scotland, an Ethnography of the Gael. McFarland. p. 14. ISBN 978-0899503622.
  92. ^ Huxley, T. (1890). "The Aryan Question and Pre-Historic Man". Collected Essays: Volume VII, Man's Place in Nature.
  93. ^ Steensby, H. P. (1907). "Racestudier i Danmark" [Race Studies in Denmark] (PDF). Geographical Journal (in Danish). Geografisk Tidsskrift. 9.
  94. ^ Coon, C. S. (1962). The Origin of races. Knopf. pp. 548–549.
  95. ^ Tattersall, Ian; Schwartz, Jeffrey H. (1999). "Hominids and hybrids: The place of Neanderthals in human evolution". Proceedings of the National Academy of Sciences. 96 (13): 7117–19. Bibcode:1999PNAS...96.7117T. doi:10.1073/pnas.96.13.7117. JSTOR 48019. PMC 33580. PMID 10377375.
  96. ^ Duarte, C.; Maurício, J.; Pettitt, P. B.; Souto, P.; Trinkaus, E.; van der Plicht, H.; Zilhão, J. (1999). "The early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern human emergence in Iberia". Proceedings of the National Academy of Sciences of the United States of America. 96 (13): 7604–7609. Bibcode:1999PNAS...96.7604D. doi:10.1073/pnas.96.13.7604. PMC 22133. PMID 10377462.
  97. ^ a b Hublin, J. J. (2009). "The origin of Neandertals". Proceedings of the National Academy of Sciences. 106 (38): 16022–27. Bibcode:2009PNAS..10616022H. doi:10.1073/pnas.0904119106. JSTOR 40485013. PMC 2752594. PMID 19805257.
  98. ^ Harvati, K.; Frost, S. R.; McNulty, K. P. (2004). "Neanderthal taxonomy reconsidered: implications of 3D primate models of intra- and interspecific differences". Proc. Natl. Acad. Sci. USA. 101 (5): 1147–52. Bibcode:2004PNAS..101.1147H. doi:10.1073/pnas.0308085100. PMC 337021. PMID 14745010.
  99. ^ Soficaru, Andrei; Dobos, Adrian; Trinkaus, Erik (2006). "Early modern humans from the Pestera Muierii, Baia de Fier, Romania". Proceedings of the National Academy of Sciences. 103 (46): 17196–201. Bibcode:2006PNAS..10317196S. doi:10.1073/pnas.0608443103. JSTOR 30052409. PMC 1859909. PMID 17085588.
  100. ^ a b Chen, F.; Welker, F.; Shen, C. (2019). "A late Middle Pleistocene Denisovan mandible from the Tibetan Plateau". Nature. 569 (7, 756): 409–412. Bibcode:2019Natur.569..409C. doi:10.1038/s41586-019-1139-x. PMID 31043746.
  101. ^ Hofreiter, M. (2011). "Drafting Human Ancestry: What Does the Neanderthal Genome Tell Us about Hominid Evolution? Commentary on Green et al. (2010)". Human Biology. 83 (1): 1–11. doi:10.3378/027.083.0101. PMID 21453001.
  102. ^ a b Currat, M.; Excoffier, L. (2004). "Modern Humans Did Not Admix with Neanderthals during Their Range Expansion into Europe". PLoS Biology. 2 (12): e421. doi:10.1371/journal.pbio.0020421. PMC 532389. PMID 15562317.
  103. ^ Mendez, Fernando L.; Poznik, G. David; Castellano, Sergi; Bustamante, Carlos D. (2016). "The Divergence of Neandertal and Modern Human Y Chromosomes". American journal of Human Genetics. 98 (4): P728–734. doi:10.1016/j.ajhg.2016.02.023.
  104. ^ a b Serre, D.; Langaney, A.; Chech, M.; Teschler-Nicola, M.; Paunovic, M.; Mennecier, P.; Hofreiter, M.; Possnert, G.; Pääbo, S. (2004). "No Evidence of Neandertal mtDNA Contribution to Early Modern Humans". PLoS Biology. 2 (3): e57. doi:10.1371/journal.pbio.0020057. PMC 368159. PMID 15024415.
  105. ^ a b Pennisi, E (2013). "More Genomes from Denisova Cave Show Mixing of Early Human Groups". Science. 340 (6134): 799. Bibcode:2013Sci...340..799P. doi:10.1126/science.340.6134.799. PMID 23687020.
  106. ^ a b c d e f Rogers, A. R.; Bohlender, R. J.; Huff, C. D. (2017). "Early history of Neanderthals and Denisovans". Proceedings of the National Academy of Sciences. 114 (37): 9, 859–9, 863. doi:10.1073/pnas.1706426114. PMC 5604018. PMID 28784789.
  107. ^ a b c d e f Prüfer, K.; et al. (2014). "The complete genome sequence of a Neanderthal from the Altai Mountains". Nature. 505 (7481): 43–49. Bibcode:2014Natur.505...43P. doi:10.1038/nature12886. PMC 4031459. PMID 24352235.
  108. ^ a b Hajdinjak, M.; Fu, Q.; Hübner, A. (2018). "Reconstructing the genetic history of late Neanderthals". Nature. 555 (7, 698): 652–656. Bibcode:2018Natur.555..652H. doi:10.1038/nature26151. PMC 6485383. PMID 29562232.
  109. ^ Posth, Cosimo; Wißing, Christoph; Kitagawa, Keiko; et al. (2017). "Deeply divergent archaic mitochondrial genome provides lower time boundary for African gene flow into Neanderthals". Nature Communications. 8: 16046. Bibcode:2017NatCo...816046P. doi:10.1038/ncomms16046. PMC 5500885. PMID 28675384.
  110. ^ a b c D. Dean; J.-J. Hublin; R. Holloway; R. Ziegler (1998). "On the phylogenetic position of the pre-Neandertal specimen from Reilingen, Germany". Journal of Human Evolution. 34 (5): 485–508. doi:10.1006/jhev.1998.0214. PMID 9614635.
  111. ^ Ko, K. W. (2016). "Hominin interbreeding and the evolution of human variation". Journal of Biological Research-Thessaloniki. 23: 17. doi:10.1186/s40709-016-0054-7. PMC 4947341. PMID 27429943.
  112. ^ a b c Papagianni, D.; Morse, M. (2013). The Neanderthals Rediscovered. Thames & Hudson. ISBN 978-0-500-05177-1.
  113. ^ Daura, J.; Sanz, M.; Arsuaga, J. L.; Hoffman, D. L. (2017). "New Middle Pleistocene hominin cranium from Gruta da Aroeira (Portugal)". Proceedings of the National Academy of Sciences. 114 (13): 3, 397–3, 402. doi:10.1073/pnas.1619040114. PMC 5380066. PMID 28289213.
  114. ^ a b Stringer, C.; Gamble, C. (1993). In Search of the Neanderthals. Thames and Hudson. ISBN 978-0-500-05070-5.
  115. ^ B. Vandermeersch; M.D. Garralda (2011). "Neanderthal Geographical and Chronological Variation". In S. Condemi; G.-C. Weniger (eds.). Continuity and Discontinuity in the Peopling of Europe. Vertebrate Paleobiology and Paleoanthropology. Springer Netherlands. pp. 113–25. doi:10.1007/978-94-007-0492-3_10. ISBN 978-94-007-0491-6.
  116. ^ N.J. Conard; J. Richter, eds. (2011). "2". Neanderthal Lifeways, Subsistence and Technology. Vertebrate Paleobiology and Paleoanthropology. 19. Springer. pp. 7–14. doi:10.1007/978-94-007-0415-2_2. ISBN 978-9400704145.
  117. ^ Zanolli, Clément; Martinón-Torres, María; Bernardini, Federico; Boschian, Giovanni; Coppa, Alfredo; Dreossi, Diego; Mancini, Lucia; Martínez de Pinillos, Marina; Martín-Francés, Laura; Bermúdez de Castro, José María; Tozzi, Carlo; Tuniz, Claudio; Macchiarelli, Roberto (2018). "The Middle Pleistocene (MIS 12) human dental remains from Fontana Ranuccio (Latium) and Visogliano (Friuli-Venezia Giulia), Italy. A comparative high resolution endostructural assessment". PLOS ONE. 13 (10): e0189773. Bibcode:2018PLoSO..1389773Z. doi:10.1371/journal.pone.0189773. PMC 6169847. PMID 30281595.
  118. ^ a b c Endicott, P.; Ho, S. Y. W.; Stringer, C. (2010). "Using genetic evidence to evaluate four palaeoanthropological hypotheses for the timing of Neanderthal and modern human origins" (PDF). Journal of Human Evolution. 59 (1): 87–95. doi:10.1016/j.jhevol.2010.04.005. PMID 20510437.
  119. ^ Briggs, A. W.; Good, J. M.; Green, R. E. (2009). "Targeted retrieval and analysis of five Neandertal mtDNA genomes" (PDF). Science. 325 (5, 938): 318–321. Bibcode:2009Sci...325..318B. doi:10.1126/science.1174462. PMID 19608918.
  120. ^ Green, R. E.; Malaspinas, A. S.; Krause, J.; Briggs, A. W. (2008). "A complete Neandertal mitochondrial genome sequence determined by high-throughput sequencing". Cell. 134 (3): 416–426. doi:10.1016/j.cell.2008.06.021. PMC 2602844. PMID 18692465.
  121. ^ Sawyer, S.; Renaud, G.; Viola, B.; Hublin, J. J. (2015). "Nuclear and mitochondrial DNA sequences from two Denisovan individuals". Proceedings of the National Academy of Sciences. 112 (51): 15, 696–15, 700. Bibcode:2015PNAS..11215696S. doi:10.1073/pnas.1519905112. PMC 4697428. PMID 26630009.
  122. ^ a b Serangeli, J.; Bolus, M. (2008). "Out of Europe - The dispersal of a successful European hominin form" (PDF). Quartär. 55: 83–98.
  123. ^ Callander, J. (2004). "Dorothy Garrod's Excavations in the Late Mousterian of Shukbah Cave in Palestine Reconsidered". Proceedings of the Prehistoric Society. 70: 207–231. doi:10.1017/S0079497X00001171.
  124. ^ Wu, X.-J.; Bruner, E. (2016). "The endocranial anatomy of Maba 1". American Journal of Physical Anthropology. 160 (4): 633–643. doi:10.1002/ajpa.22974. PMID 26972814.
  125. ^ "The oldest people in Wales - Neanderthal teeth from Pontnewydd Cave". National Museum Wales. Retrieved October 28, 2019.
  126. ^ Pavlov P, Roebroeks W, Svendsen JI (2004). "The Pleistocene colonization of northeastern Europe: a report on recent research". Journal of Human Evolution. 47 (1–2): 3–17. doi:10.1016/j.jhevol.2004.05.002. PMID 15288521.
  127. ^ a b Slimak, L.; Svendsen, J. I.; Mangerud, J.; Plisson, H. (2011). "Late Mousterian persistence near the Arctic Circle". Science. 332 (6, 031): 841–845. Bibcode:2011Sci...332..841S. doi:10.1126/science.1203866. JSTOR 29784275. PMID 21566192.
  128. ^ Slimak, L. (2012). "Response to 'Comment on Late Mousterian Persistence near the Arctic Circle". Science. 335 (6065): 167. Bibcode:2012Sci...335..167S. doi:10.1126/science.1210211.
  129. ^ a b Zwyns, N. (2012). "Comment on Late Mousterian Persistence near the Arctic Circle". Science. 335 (6065): 167. Bibcode:2012Sci...335..167Z. doi:10.1126/science.1209908.
  130. ^ Holan, S. R.; Deméré, T. A.; Fisher, D. C.; Fullager, R.; Paces, J. B.; Jefferson, G. T. (2017). "A 130,000-year-old archaeological site in southern California, USA". Nature. 544 (7651): 479–483. doi:10.1038/nature22065.
  131. ^ Sutton, M. Q.; Parkinson, J.; Rosen, M. D. (2019). "Observations Regarding the Cerutti Mastodon". PaleoAmerica. 5 (1). doi:10.1080/20555563.2019.1589409.
  132. ^ Eren, M. I.; Bebber, M. R. (2019). "The Cerutti Mastodon site and experimental archaeology's quiet coming of age". Antiquity. 93 (369): 796–797. doi:10.15184/aqy.2019.50.
  133. ^ Ferrel, P. M. (2019). "The Cerutti Mastodon Site Reinterpreted with Reference to Freeway Construction Plans and Methods". PaleoAmerica. 5 (1): 1–7. doi:10.1080/20555563.2019.1589663.
  134. ^ a b c d e f g h Bocquet-Appel, J.; Degioanni, A. (2013). "Neanderthal Demographic Estimates". Current Anthropology. 54: 202–214. doi:10.1086/673725.
  135. ^ a b c Mafessoni, F.; Prüfer, K. (2017). "Better support for a small effective population size of Neandertals and a long shared history of Neandertals and Denisovans". Proceedings of the National Academy of Sciences. 114 (48): 10, 256–10, 257. doi:10.1073/pnas.1716918114.
  136. ^ Lalueza-Fox, C.; Sampietro, M. L.; Caramelli, D.; Puder, Y. (2013). "Neandertal evolutionary genetics: mitochondrial DNA data from the iberian peninsula". Molecular Biology and Evolution. 22 (4): 1077–1081. doi:10.1093/molbev/msi094. PMID 15689531.
  137. ^ Fabre, V.; Condemi, S.; Degioanni, A. (2009). "Genetic Evidence of Geographical Groups among Neanderthals". PLOS ONE. 4 (4): e5151. Bibcode:2009PLoSO...4.5151F. doi:10.1371/journal.pone.0005151. PMC 2664900. PMID 19367332.
  138. ^ Mellars, P.; French, J. C. (2011). "Tenfold population increase in Western Europe at the Neandertal-to-modern human transition". Science. 333 (6042): 623–627. Bibcode:2011Sci...333..623M. doi:10.1126/science.1206930. PMID 21798948.
  139. ^ Sánchez-Quinto, F.; Lalueza-Fox, C. (2015). "Almost 20 years of Neanderthal palaeogenetics: adaptation, admixture, diversity, demography and extinction". Philosophical Transactions of the Royal Society B. 370 (1, 660): 20130374. doi:10.1098/rstb.2013.0374. PMC 4275882. PMID 25487326.
  140. ^ a b c d Ríos, L.; Kivell, T. L.; Lalueza-Fox, C.; Estalrrich, A. (2019). "Skeletal Anomalies in The Neandertal Family of El Sidrón (Spain) Support A Role of Inbreeding in Neandertal Extinction". Scientific Reports. 9 (1, 697): 1697. Bibcode:2019NatSR...9.1697R. doi:10.1038/s41598-019-38571-1. PMC 6368597. PMID 30737446.
  141. ^ a b Pettitt, R. B. (2000). "Neanderthal Lifecycles: Developmental and Social Phases in the Lives of the Last Archaics". World Archaeology. 31 (3): 351–366. doi:10.1080/00438240009696926. JSTOR 125106. PMID 16475295.
  142. ^ a b c Duveau, J.; Berillon, G.; Verna, C.; Laisné, G.; Cliquet, D. (2019). "The composition of a Neandertal social group revealed by the hominin footprints at Le Rozel (Normandy, France)". Proceedings of the National Academy of Sciences. 116 (39): 19, 409–19, 414. doi:10.1073/pnas.1901789116. PMC 6765299. PMID 31501334.
  143. ^ a b c d Rougier, H.; Crevecoeur, I.; Beauval, C. (2016). "Neandertal cannibalism and Neandertal bones used as tools in Northern Europe". Scientific Reports. 6 (29, 005): 29005. Bibcode:2016NatSR...629005R. doi:10.1038/srep29005. PMC 4933918. PMID 27381450.
  144. ^ McDermott, F.; Grün, R.; Stringer, C. B.; Hawkesworth, C. J. (1993). "Mass-spectrometric U-series dates for Israeli Neanderthal/early modern hominid sites". Nature. 363 (6426): 252–255. Bibcode:1993Natur.363..252M. doi:10.1038/363252a0. PMID 8387643.
  145. ^ Rink, W. Jack; Schwarcz, H. P.; Lee, H. K.; Rees-Jones, J.; Rabinovich, R.; Hovers, E. (2002). "Electron spin resonance (ESR) and thermal ionization mass spectrometric (TIMS) 230Th/234U dating of teeth in Middle Paleolithic layers at Amud Cave, Israel". Geoarchaeology. 16 (6): 701–717. doi:10.1002/gea.1017.
  146. ^ Valladas, H.; Merciera, N.; Frogeta, L.; Hoversb, E.; Joronc, J.L.; Kimbeld, W. H.; Rak, Y. (1999). "TL Dates for the Neanderthal Site of the Amud Cave, Israel". Journal of Archaeological Science. 26 (3): 259–268. doi:10.1006/jasc.1998.0334.
  147. ^ Bischoff, James L.; Shamp; et al. (2003). "The Sima de los Huesos Hominids Date to Beyond U/Th Equilibrium (>350kyr) and Perhaps to 400–500kyr: New Radiometric Dates". Journal of Archaeological Science. 30 (3): 275–80. doi:10.1006/jasc.2002.0834.
  148. ^ Arsuaga, J. L.; Martínez, I.; Gracia, A.; Lorenzo, C. (1997). "The Sima de los Huesos crania (Sierra de Atapuerca, Spain). A comparative study". Journal of Human Evolution. 33 (2–3): 219–281. doi:10.1006/jhev.1997.0133. PMID 9300343.
  149. ^ a b Haeusler, M.; Trinkaus, E.; Fornai, C.; Müller, J.; Bonneau, N.; Boeni, T.; Frater, N. (2019). "Morphology, pathology, and the vertebral posture of the La Chapelle-aux-Saints Neandertal". Proceedings of the National Academy of Sciences. 116 (11): 4, 923–4, 927. doi:10.1073/pnas.1820745116. PMC 6421410. PMID 30804177.
  150. ^ a b c Trinkaus, E. (1981). "Neanderthal limb proportions and cold adaptation". In Stringer, C. B. (ed.). Aspects of human evolution. Taylor and Francis Ltd.
  151. ^ Froehle, A. W.; Churchill, S. E. (2009). "Energetic Competition Between Neandertals and Anatomically Modern Humans" (PDF). PaleoAnthropology: 96–116.
  152. ^ a b Weaver, T. D. (2009). "The meaning of Neandertal skeletal morphology". Proceedings of the National Academy of Sciences. 106 (38): 16, 028–16, 033. doi:10.1073/pnas.0903864106. PMC 2752516. PMID 19805258.
  153. ^ Holliday, T. W. (1997). "Postcranial evidence of cold adaptation in European Neandertals". American Journal of Physiological Anthropology. 104 (2): 245–258. doi:10.1002/(SICI)1096-8644(199710)104:2<245::AID-AJPA10>3.0.CO;2-#. PMID 9386830.
  154. ^ Walker, M. J.; Ortega, J.; Parmová, K.; López, M. V.; Trinkaus, E. (2011). "Morphology, body proportions, and postcranial hypertrophy of a female Neandertal from the Sima de las Palomas, southeastern Spain". Proceedings of the National Academy of Sciences. 108 (25): 10, 087–10, 091. Bibcode:2011PNAS..10810087W. doi:10.1073/pnas.1107318108. PMC 3121844. PMID 21646528.
  155. ^ Lee, S. S. M.; Piazza, S. J. (2009). "Built for speed: musculoskeletal structure and sprinting ability" (PDF). Journal of Experimental Biology. 212 (22): 3, 700–3, 707. doi:10.1242/jeb.031096. PMID 19880732.
  156. ^ Trinkaus, E. (1975). "Squatting among the neandertals: A problem in the behavioral interpretation of skeletal morphology". Journal of Archaeological Science. 2 (4): 327–351. doi:10.1016/0305-4403(75)90005-9.
  157. ^ Gunz, P.; Tilot, A. K.; Wittfeld, K.; Teumer, A.; Shapland, C. Y.; van Erp, T. G. M.; Dannemann, M.; Vernot, B.; Neubauer, S.; Guadalupe, T.; Fernández, G.; Brunner, H. G.; Enard, W.; Fallon, J.; Hosten, N.; Völker, U.; Profico, A.; Di Vincenzo, F.; Manzi, G.; Kelso, J.; St. Pourcain, B.; Hublin, J.-J.; Franke, B.; Pääbo, S.; Macciardi, F.; Grabe, H. J.; Fisher, S. E. (2019). "Neandertal Introgression Sheds Light on Modern Human Endocranial Globularity". Current Biology. 29 (1): 120–127.e5. doi:10.1016/j.cub.2018.10.065. PMC 6380688. PMID 30554901.
  158. ^ Gunz, P.; Harvati, K. (2007). "The Neanderthal "chignon": Variation, integration, and homology". Journal of Human Evolution. 52 (3): 262–274. doi:10.1016/j.jhevol.2006.08.010. PMID 17097133.
  159. ^ a b c Pearce, E.; Stringer, C.; Dunbar, R. I. M. (2013). "New insights into differences in brain organization between Neanderthals and anatomically modern humans". Proceedings of the Royal Society B. 280 (1, 758): 20130168. doi:10.1098/rspb.2013.0168. PMC 3619466. PMID 23486442.
  160. ^ a b c d Clement, A. F.; Hillson, S. W.; Aiello, L. C. (2012). "Tooth wear, Neanderthal facial morphology and the anterior dental loading hypothesis". Journal of Human Evolution. 62 (3): 367–376. doi:10.1016/j.jhevol.2011.11.014. PMID 22341317.
  161. ^ Rae TC, Koppe T, Stringer CB (2011). "The Neanderthal face is not cold adapted". Journal of Human Evolution. 60 (2): 234–239. doi:10.1016/j.jhevol.2010.10.003. PMID 21183202.
  162. ^ O'Connor, C. F.; Franciscus, R. G.; Holton, N. E. (2005). "Bite force production capability and efficiency in Neandertals and modern humans". American Journal of Physical Anthropology. 127 (2): 129–151. doi:10.1002/ajpa.20025. PMID 15558614.
  163. ^ Takaki, P.; Vieira, M.; Bommarito, S. (2014). "Maximum bite force analysis in different age groups". International Archives of Otorhinolaryngology. 18 (3): 272–276. doi:10.1055/s-0034-1374647. PMC 4297017. PMID 25992105.
  164. ^ Beals, Kenneth; Smith, Courtland; Dodd, Stephen (1984). "Brain Size, Cranial Morphology, Climate, and Time Machines" (PDF). Current Anthropology. 12 (3): 301–30. doi:10.1086/203138.
  165. ^ Allen, J. S.; Damasio, H.; Grabowski, T. J. (2002). "Normal neuroanatomical variation in the human brain: an MRI-volumetric study". Anatomical Journal of Physical Anthropology. 118 (4): 341–358. doi:10.1002/ajpa.10092. PMID 12124914.
  166. ^ Amano, H.; Kikuchi, T.; Morita, Y.; Kondo, O.; Suzuki, H.; Ponce de Leon, M. S.; Zollikofer, C.P.E.; Bastir, M.; Stringer, C.; Ogihara, N. (2015). "Virtual Reconstruction of the Neanderthal Amud 1 Cranium" (PDF). American Journal of Physical Anthropology. 158 (2): 185–197. doi:10.1002/ajpa.22777. hdl:10261/123419. PMID 26249757.
  167. ^ Ponce de León, M. S.; Golovanova, L.; Doronichev, V. (2008). "Neanderthal brain size at birth provides insights into the evolution of human life history". Proceedings of the National Academy of Sciences. 105 (37): 13, 764–13, 768. doi:10.1073/pnas.0803917105. PMC 2533682. PMID 18779579.
  168. ^ Kochiyama, T.; Ogihara, N.; Tanabe, H. C. (2018). "Reconstructing the Neanderthal brain using computational anatomy". Scientific Reports. 8 (6296): 6296. Bibcode:2018NatSR...8.6296K. doi:10.1038/s41598-018-24331-0. PMC 5919901. PMID 29700382.
  169. ^ Peña-Melián, A.; Rosas, A.; García-Tabernero, A.; Bastir, M.; de la Rasilla, M. (2011). "Paleoneurology of Two New Neandertal Occipitals from El Sidrón (Asturias, Spain) in the Context of Homo Endocranial Evolution". The Anatomical Record. 294 (8): 1, 370–1, 381. doi:10.1002/ar.21427. hdl:10261/123675. PMID 21714107.
  170. ^ a b c Dannemann, M.; Kelso, J. (2017). "The Contribution of Neanderthals to Phenotypic Variation in Modern Humans". The American Journal of Human Genetics. 101 (4): 578–589. doi:10.1016/j.ajhg.2017.09.010. PMC 5630192. PMID 28985494.
  171. ^ Cerqueira, C. C.; Piaxão-Côrtes, V. R.; Zambra, F. M. B.; Hünemeier, T.; Bortolini, M. (2012). "Predicting homo pigmentation phenotype through genomic data: From neanderthal to James Watson". American Journal of Human Biology. 24 (5): 705–709. doi:10.1002/ajhb.22263. PMID 22411106.
  172. ^ Allentoft, M. E.; Sikora, M. (2015). "Population genomics of Bronze Age Eurasia". Nature. 522 (7, 555): 167–172. Bibcode:2015Natur.522..167A. doi:10.1038/nature14507. PMID 26062507.
  173. ^ Lalueza-Fox, C.; Rompler, H.; Caramelli, D.; Staubert, C.; Catalano, G.; Hughes, D.; Rohland, N.; Pilli, E.; Longo, L.; Condemi, S.; de la Rasilla, M.; Fortea, J.; Rosas, A.; Stoneking, M.; Schoneberg, T.; Bertranpetit, J.; Hofreiter, M. (2007). "A Melanocortin 1 Receptor Allele Suggests Varying Pigmentation Among Neanderthals". Science. 318 (5855): 1, 453–1, 455. Bibcode:2007Sci...318.1453L. doi:10.1126/science.1147417. PMID 17962522.
  174. ^ Ding, Q.; Hu, Y.; Xu, S.; Wang, C.-C.; Li, H.; Zhang, R.; Yan, S.; Wang, J.; Jin, L. (2014). "Neanderthal Origin of the Haplotypes Carrying the Functional Variant Val92Met in the MC1R in Modern Humans". Molecular Biology and Evolution. 31 (8): 1, 994–2, 003. doi:10.1093/molbev/msu180. PMID 24916031.
  175. ^ Venner, S. J. (2018). A New Estimate for Neanderthal Energy Expenditure (MA). CUNY Academic Works.
  176. ^ "Cro-Magnons Conquered Europe, but Left Neanderthals Alone". PLoS Biology. 2 (12): e449. 2004. doi:10.1371/journal.pbio.0020449. PMC 532398.
  177. ^ Smith, T. M.; Tafforeau, P.; Reid, D. J. (2010). "Dental evidence for ontogenetic differences between modern humans and Neanderthals". Proceedings of the National Academy of Sciences. 107 (49): 20, 923–20, 928. Bibcode:2010PNAS..10720923S. doi:10.1073/pnas.1010906107. PMC 3000267. PMID 21078988.
  178. ^ Guatelli-Steinberg, D. (2009). "Recent studies of dental development in Neandertals: Implications for Neandertal life histories". Evolutionary Biology. 18 (1): 9–20. doi:10.1002/evan.20190.
  179. ^ a b Nakahashi, W. (2017). "The effect of trauma on Neanderthal culture: A mathematical analysis". Homo. 68 (2): 83–100. doi:10.1016/j.jchb.2017.02.001. PMID 28238406.
  180. ^ Beier, J.; Anthes, N.; Wahl, J.; Harvati, K. (2018). "Similar cranial trauma prevalence among Neanderthals and Upper Palaeolithic modern humans". Nature. 563 (7, 733): 686–690. Bibcode:2018Natur.563..686B. doi:10.1038/s41586-018-0696-8. PMID 30429606.
  181. ^ Trinkaus, E.; Villotte, S (2017). "External auditory exostoses and hearing loss in the Shanidar 1 Neandertal". PLOS ONE. 12 (10): e0186684. Bibcode:2017PLoSO..1286684T. doi:10.1371/journal.pone.0186684. PMC 5650169. PMID 29053746.
  182. ^ Crubézy, E.; Trinkaus, E. (1992). "Shanidar 1: a case of hyperostotic disease (DISH) in the middle Paleolithic". American Journal of Physical Anthropology. 89 (4): 411–420. doi:10.1002/ajpa.1330890402. PMID 1463085.
  183. ^ Houldcroft, C. J.; Underdown, S. J. (2016). "Neanderthal genomics suggests a pleistocene time frame for the first epidemiologic transition". American Journal of Physical Anthropology. 160 (3): 379–388. doi:10.1002/ajpa.22985. PMID 27063929.
  184. ^ Fennel, K. J.; Trinkaus, E. (1997). "Bilateral Femoral and Tibial Periostitis in the La Ferrassie 1 Neanderthal". Journal of Archaeological Science. 24 (11): 985–995. doi:10.1006/jasc.1996.0176.
  185. ^ Rosas, A.; Estalrrich, A.; García-Vargas, S.; García-Tabernero, A.; Huguet, R.; Lalueza-Fox, C.; de la Rasilla, M. (2013). "Identification of Neandertal individuals in fragmentary fossil assemblages by means of tooth associations: The case of El Sidrón (Asturias, Spain)". Comptes Rendus Palevol. 12 (5): 279–291. doi:10.1016/j.crpv.2013.06.003.
  186. ^ Farizy, C. (1994). "Spatial patterning of Middle Paleolithic sites". Journal of Anthropological Archaeology. 13 (2): 153–160. doi:10.1006/jaar.1994.1010.
  187. ^ a b Tattersall, Ian (2015). The Strange Case of the Rickety Cossack and other Cautionary Tales from Human Evolution. Palgrave Macmillan. p. 202. ISBN 978-1-137-27889-0.
  188. ^ Zollikofer, C. P. E.; Ponce de Leon, M. S.; Vandermeersch, B.; Leveque, F. (2002). "Evidence for interpersonal violence in the St. Cesaire Neanderthal". Proceedings of the National Academy of Sciences. 99 (9): 6444–6448. Bibcode:2002PNAS...99.6444Z. doi:10.1073/pnas.082111899. PMC 122968. PMID 11972028.
  189. ^ Churchill, S. E.; Franciscus, R. G.; McKean-Peraza, H. A.; Daniel, J. A.; Warren, B. R. (2009). "Shanidar 3 Neandertal rib puncture wound and paleolithic weaponry". Journal of Human Evolution. 57 (2): 163–178. doi:10.1016/j.jhevol.2009.05.010. PMID 19615713.
  190. ^ Estalrrich, A.; Rosas, A. (2015). "Division of labor by sex and age in Neandertals: an approach through the study of activity-related dental wear". Journal of Human Evolution. 80: 51–63. doi:10.1016/j.jhevol.2014.07.007. PMID 25681013.
  191. ^ Bocherens, H.; Drucker, D. G.; Billiou, D.; Patou-Mathis, M.; Vandermeersch, B. (2005). "Isotopic evidence for diet and subsistence pattern of the Saint-Césaire I Neanderthal: Review and use of a multi-source mixing model". Journal of Human Evolution. 49 (1): 71–87. doi:10.1016/j.jhevol.2005.03.003. PMID 15869783.
  192. ^ a b Dusseldorp, G. L. (2013). "Neanderthals and Cave Hyenas: Co-existence, Competition or Conflict?" (PDF). In Clark, J. L.; Speth, J. D. (eds.). Zooarchaeology and Modern Human Origins. Vertebrate Paleobiology and Paleoanthropology. Springer Science+Business Media Dordrecht. pp. 191–208. doi:10.1007/978-94-007-6766-9_12. ISBN 978-94-007-6765-2.
  193. ^ a b Richards, M. P.; Pettitt, P. B.; Trinkaus, E.; Smith, F. H.; Paunović, M.; Karavanić, I. (2000). "Neanderthal diet at Vindija and Neanderthal predation: The evidence from stable isotopes". Proceedings of the National Academy of Sciences. 97 (13): 7663–66. Bibcode:2000pnas...97.7663r. doi:10.1073/pnas.120178997. JSTOR 122870. PMC 16602. PMID 10852955.
  194. ^ Fiorenza, Luca; Benazzi, Stefano; Tausch, Jeremy; Kullmer, Ottmar; Bromage, Timothy G.; Schrenk, Friedemann (2011). Rosenberg, Karen (ed.). "Molar Macrowear Reveals Neanderthal Eco-Geographic Dietary Variation". PLOS ONE. 6 (3): e14769. Bibcode:2011PLoSO...614769F. doi:10.1371/journal.pone.0014769. PMC 3060801. PMID 21445243.
  195. ^ Jaouen, Klervia; et al. (2019). "Exceptionally high δ15N values in collagen single amino acids confirm Neandertals as high-trophic level carnivores". Proceedings of the National Academy of Sciences. 116 (11): 4, 928–4, 933. doi:10.1073/pnas.1814087116. PMC 6421459. PMID 30782806.
  196. ^ a b Marín, J.; Saladié, P.; Rodríguez-Hidalgo, A.; Carbonell, E. (2017). "Neanderthal hunting strategies inferred from mortality profiles within the Abric Romaní sequence". PLOS ONE. 12 (11): e0186970. Bibcode:2017PLoSO..1286970M. doi:10.1371/journal.pone.0186970. PMC 5699840. PMID 29166384.
  197. ^ Henry, A. G.; Brooks, A. S.; Piperno, D. R. (2010). "Microfossils in calculus demonstrate consumption of plants and cooked foods in Neanderthal diets (Shanidar III, Iraq; Spy I and II, Belgium)". Proceedings of the National Academy of Sciences. 108 (2): 486–91. Bibcode:2011PNAS..108..486H. doi:10.1073/pnas.1016868108. PMC 3021051. PMID 21187393.
  198. ^ Shipley, G. P.; Kindscher, K. (2016). "Evidence for the Paleoethnobotany of the Neanderthal: A Review of the Literature". Scientifica. 2016: 1–12. doi:10.1155/2016/8927654. PMC 5098096. PMID 27843675.
  199. ^ Madella, M.; Jones, M. K.; Goldberg, P.; Goren, Y.; Hovers, E. (2002). "The Exploitation of Plant Resources by Neanderthals in Amud Cave (Israel): The Evidence from Phytolith Studies". Journal of Archaeological Science. 29 (7): 703–719. doi:10.1006/jasc.2001.0743.
  200. ^ Sistiaga, Ainara; et al. (2014). "The Neanderthal meal: A new perspective using faecal biomarkers". PLOS ONE. 9 (6): e101045. Bibcode:2014PLoSO...9j1045S. doi:10.1371/journal.pone.0101045. PMC 4071062. PMID 24963925.
  201. ^ Henry, A. G.; Brooks, A. S.; Piperno, D. R. (2011). "Microfossils in calculus demonstrate consumption of plants and cooked foods in Neanderthal diets (Shanidar III, Iraq; Spy I and II, Belgium)". Proceedings of the National Academy of Sciences. 108 (2): 486–491. Bibcode:2011PNAS..108..486H. doi:10.1073/pnas.1016868108. PMID 21187393.
  202. ^ Daugeard, C. (2008). "Exploitation du milieu animal par les Néandertaliens dans le Sud-Est de la France" [Exploitation of the environment by Neanderthals in the southeast of France]. Bulletin de la Société préhistorique française (in French). 106 (4): 818–819.
  203. ^ a b Speth, J. D. (2015). "When did humans learn to boil?" (PDF). PaleoAnthropology: 54–67. doi:10.4207/PA.2015.ART96 (inactive November 7, 2019).
  204. ^ Schofield, D. P.; McGrew, W. C.; Takahashi, A.; Hirata, S. (2018). "Cumulative culture in nonhumans: overlooked findings from Japanese monkeys?". Primates. Primates (59): 113–122. doi:10.1007/s10329-017-0642-7.
  205. ^ Speth, J. D.; Tchernov, E. (2002). "Middle Paleolithic Tortoise Use at Kebara Cave (Israel)". Journal of Archaeological Science. 29 (5): 471–483. doi:10.1006/jasc.2001.0740.
  206. ^ Diedrich, C. G. (2010). "The Crocuta crocuta spelaea (Goldfuss 1823) population and its prey from the Late Pleistocene Teufelskammer Cave hyena den besides the famous Paleolithic Neandertal Cave (NRW, NW Germany)". Historical Biology. 23 (2–3): 237–270. doi:10.1080/08912963.2010.530348.
  207. ^ a b c d Yravedra, J.; Yustos, M. (2015). "Cannibalism in the Neanderthal World: An Exhaustive Revision". Journal of Taphonomy. 13 (1): 33–52.
  208. ^ Fernández-Jalvo, Y.; Carlos Díez, J.; Cáceres, I.; Rosell, J. (1999). "Human cannibalism in the Early Pleistocene of Europe (Gran Dolina, Sierra de Atapuerca, Burgos, Spain)". Journal of Human Evolution. 37 (3–4): 591–562. doi:10.1006/jhev.1999.0324. PMID 10497001.
  209. ^ Rosas, A.; Bastir, M.; Martínez-Maza, C. (2006). "Paleobiology and comparative morphology of a late Neandertal sample from El Sidrón, Asturias, Spain". Proceedings of the National Academy of Sciences. 103 (51): 19, 266–19, 271. Bibcode:2006PNAS..10319266R. doi:10.1073/pnas.0609662104. PMC 1748215. PMID 17164326.
  210. ^ Defleur, A.; White, T.; Valensi, P.; Slimak, L. (1999). "Neanderthal cannibalism at Moula-Guercy, Ardèche, France". Science. 286 (5, 437): 128–131. doi:10.1126/science.286.5437.128. PMID 10506562.
  211. ^ a b Radovčić, D.; Sršen, A. O.; Radovčić, J.; Frayer, D. W.; Petraglia, M. D. (2015). "Evidence for Neandertal Jewelry: Modified White-Tailed Eagle Claws at Krapina". PLOS ONE. 10 (3): e0119802. Bibcode:2015PLoSO..1019802R. doi:10.1371/journal.pone.0119802. PMC 4356571. PMID 25760648.
  212. ^ R. S. Solecki (1975). "Shanidar IV, a Neanderthal Flower Burial in Northern Iraq". Science. 190 (4217): 880–81. Bibcode:1975Sci...190..880S. doi:10.1126/science.190.4217.880.
  213. ^ D.J. Sommer (1999). "The Shanidar IV 'Flower Burial': a Re-evaluation of Neanderthal Burial Ritual". Cambridge Archaeological Journal. 9 (1): 127–29. doi:10.1017/s0959774300015249.
  214. ^ Pettitt, P. B. (2002). "The Neanderthal Dead, exploring mortuary variability in Middle Paleolithic Eurasia". Before Farming. 1 (4).
  215. ^ Marquet, J.; Lorblanchet, M.; Oberlin, C.; Thamo-Bozso, E.; Aubry, T. (2016). "New dating of the "mask" of La Roche-Cotard (Langeais, Indre-et-Loire, France)". Paleo Revue d'Archéologie Préhistorique. 27: 253–263.
  216. ^ Marquet, J.-C.; Lorblanchet, M. (2003). "A Neanderthal face? The proto-figurine from La Roche-Cotard, Langeais (Indre-et-Loire, France)". Antiquity. 77 (298): 661–670. doi:10.1017/s0003598x00061627. ISSN 0003-598X.
  217. ^ Pettitt, P. B. (2003). "Is this the infancy of art? Or the art of an infant? A possible Neanderthal face from La Roche-Cotard, France" (PDF). Before Farming. 11 (3). Archived from the original (PDF) on October 2, 2011.
  218. ^ "But is it art?". Science. 302 (5, 652): 1890a–1890. 2003. doi:10.1126/science.302.5652.1890a.
  219. ^ Higham, T.; Jacobi, R.; Julien, M.; David, F.; Basell, L.; Wood, R.; Davies, W.; Ramsey, C. B. (2010). "Chronology of the Grotte du Renne (France) and implications for the context of ornaments and human remains within the Chatelperronian". Proceedings of the National Academy of Sciences. 107 (47): 20234–20239. doi:10.1073/pnas.1007963107. PMC 2996711. PMID 20956292.
  220. ^ Mellars, P. (2010). "Neanderthal symbolism and ornament manufacture: The bursting of a bubble?". Proceedings of the National Academy of Sciences. 107 (47): 20147–20148. Bibcode:2010PNAS..10720147M. doi:10.1073/pnas.1014588107. PMC 2996706. PMID 21078972.
  221. ^ J.-J. Hublin; S. Talamo; M. Julien; F. David; N. Connet; P. Bodu; B. Vandermeersch; M.P. Richards (2012). "Radiocarbon dates from the Grotte du Renne and Saint-Césaire support a Neandertal origin for the Châtelperronian". Proceedings of the National Academy of Sciences USA. 109 (46): 18743–18748. Bibcode:2012PNAS..10918743H. doi:10.1073/pnas.1212924109. PMC 3503158. PMID 23112183.
  222. ^ F. Welkera; M. Hajdinjak; S. Talamo; K. Jaouen; M. Dannemann; F. David; M. Julien; M. Meyer; J. Kelso; I. Barnes; S. Brace; P. Kamminga; R. Fischer; B.M. Kessler; J.R. Stewart; S. Pääbo; M.J. Collins; J.-J. Hublin (2016). "Palaeoproteomic evidence identifies archaic hominins associated with the Châtelperronian at the Grotte du Renne". Proceedings of the National Academy of Sciences USA. 113 (40): 11162–67. doi:10.1073/pnas.1605834113. PMC 5056053. PMID 27638212.
  223. ^ Finlayson, C.; Brown, K.; Blasco, R.; Rosell, J.; Negro, J. José; Bortolotti, G. R.; Finlayson, G.; Sánchez Marco, A.; Giles Pacheco, F.; Rodríguez Vidal, J.; Carrión, J. S.; Fa, D. A.; Rodríguez Llanes, J. M. (2012). "Birds of a Feather: Neanderthal Exploitation of Raptors and Corvids". PLOS ONE. 7 (9): e45927. Bibcode:2012PLoSO...745927F. doi:10.1371/journal.pone.0045927. PMC 3444460. PMID 23029321.
  224. ^ E. Callaway (2014). "Neanderthals made some of Europe's oldest art". Nature News. doi:10.1038/nature.2014.15805.
  225. ^ Rodríguez-Vidal, J.; d'Errico, F.; Pacheco, F. G. (2014). "A rock engraving made by Neanderthals in Gibraltar". Proceedings of the National Academy of Sciences. 111 (37): 13, 301–13, 306. Bibcode:2014PNAS..11113301R. doi:10.1073/pnas.1411529111. PMC 4169962. PMID 25197076.
  226. ^ Jaubert, J.; Verheyden, S.; Genty, D.; Soulier, M.; Cheng, H.; Blamart, D.; Burlet, C.; Camus, H.; Delaby, S.; Deldicque, D.; Edwards, R. L.; Ferrier, C.; Lacrampe-Cuyaubère, F.; Lévêque, F.; Maksud, F.; Mora, P.; Muth, X.; Régnier, É.; Rouzaud, J.-N.; Santos, F. (2016). "Early Neanderthal constructions deep in Bruniquel Cave in southwestern France". Nature. 534 (7605): 111–114. Bibcode:2016Natur.534..111J. doi:10.1038/nature18291. PMID 27251286.
  227. ^ a b c d e f It's Official: Neanderthals Created Art. Chris Standish and Alistair Pike. Sapiens.org. 22 May 2018.
  228. ^ Callaway, E. (2015). "Neanderthals wore eagle talons as jewellery". Nature. doi:10.1038/nature.2015.17095.
  229. ^ Eagle Talon Jewelry Suggests Neanderthals Were Capable of Human-Like Thought. Megan Gannon, Smithsonian Magazine. 1 November 2019.
  230. ^ d’Errico, F.; Tsvelykh, A. (2017). "A decorated raven bone from the Zaskalnaya VI (Kolosovskaya) Neanderthal site, Crimea". PLOS ONE. 12 (3): e0173435. Bibcode:2017PLoSO..1273435M. doi:10.1371/journal.pone.0173435. PMC 5371307. PMID 28355292.
  231. ^ World's Oldest Cave Art Found—Made by Neanderthals?. Ker Than. National Geographic News. Published June 14, 2012.
  232. ^ Marris, E. (2018). "Neanderthal artists made oldest-known cave paintings". Nature News. doi:10.1038/d41586-018-02357-8.
  233. ^ Hoffman, D. L.; Angelucci, D. E.; Villaverde, V.; Zapata, Z.; Zilhão, J. (2018). "Symbolic use of marine shells and mineral pigments by Iberian Neandertals 115,000 years ago". Science Advances. 4 (2): eaar5255. Bibcode:2018SciA....4.5255H. doi:10.1126/sciadv.aar5255. PMC 5833998. PMID 29507889.
  234. ^ a b Majkić, A.; d’Errico, F.; Stepanchuk, V. (2018). "Assessing the significance of Palaeolithic engraved cortexes. A case study from the Mousterian site of Kiik-Koba, Crimea". PLOS ONE. 13 (5): e0195049. Bibcode:2018PLoSO..1395049M. doi:10.1371/journal.pone.0195049. PMC 5931501. PMID 29718916.
  235. ^ Boëda, Eric; Geneste, J. M.; Griggo, C.; Mercier, N.; Muhesen, S.; Reyss, J. L.; Taha, A.; Valladas, H. (2015). "A Levallois point embedded in the vertebra of a wild ass (Equus africanus): hafting, projectiles and Mousterian hunting weapons". Antiquity. 73 (280): 394–402. doi:10.1017/S0003598X00088335.
  236. ^ Roussel, M.; Soressi, M.; Hublin, J.-J. (2016). "The Châtelperronian conundrum: Blade and bladelet lithic technologies from Quinçay, France". Journal of Human Evolution. 95: 13–32. doi:10.1016/j.jhevol.2016.02.003. PMID 27260172.
  237. ^ Gravina, B.; Bachellerie, F.; Caux, S. (2018). "No Reliable Evidence for a Neanderthal-Châtelperronian Association at La Roche-à-Pierrot, Saint-Césaire". Scientific Reports. 8 (1): 15134. Bibcode:2018NatSR...815134G. doi:10.1038/s41598-018-33084-9. PMC 6181958. PMID 30310091.
  238. ^ Heyes, P. J.; Anastasakis, K.; de Jong, W.; van Hoesel, A.; Roebroeks, W.; Soressi, M. (2016). "Selection and Use of Manganese Dioxide by Neanderthals". Scientific Reports. 6 (1): 22,159. Bibcode:2016NatSR...622159H. doi:10.1038/srep22159. PMC 4770591. PMID 26922901.
  239. ^ a b Collard, M.; Tarle, L.; Sandgathe, D.; Allan, A. (2016). "Faunal evidence for a difference in clothing use between Neanderthals and early modern humans in Europe". Journal of Anthropological Archaeology. 44: 235–246. doi:10.1016/j.jaa.2016.07.010.
  240. ^ a b Wales, N. (2012). "Modeling Neanderthal clothing using ethnographic analogues". Journal of Human Evolution. 63 (6): 781–795. doi:10.1016/j.jhevol.2012.08.006. PMID 23084621.
  241. ^ a b Carter, T.; Contreras, D. A.; Holcomb, J.; Mihailović, D. D. (2019). "Earliest occupation of the Central Aegean (Naxos), Greece: Implications for hominin and Homo sapiens' behavior and dispersals". Science Advances. 5 (10): eaax0997. doi:10.1126/sciadv.aax0997. PMID 31663021.
  242. ^ Lieberman, P. (1992). "On Neanderthal Speech and Neanderthal Extinction". Current Anthropology. 33 (4): 409–410. doi:10.1086/204092.
  243. ^ Lieberman, P. (2007). "Current views on Neanderthal speech capabilities: A reply to Boe et al. (2002)". Journal of Phonetics. 35 (4): 552–563. doi:10.1016/j.wocn.2005.07.002.
  244. ^ Boë, L.-J.; Heim, J.-L.; Honda, K.; Maeda, S. (2002). "The potential Neandertal vowel space was as large as that of modern humans". Journal of Phonetics. 30 (3): 465–484. doi:10.1006/jpho.2002.0170.
  245. ^ Johansson, S. (2015). "Language abilities in Neanderthals". Annual Review of Linguistics. 1: 311–322. doi:10.1146/annurev-linguist-030514-124945.
  246. ^ a b Whiting, K.; Konstantakos, L.; Sadler, G.; Gill, C. (2018). "Were Neanderthals Rational? A Stoic Approach". Humanities. 7 (2): 39. doi:10.3390/h7020039.
  247. ^ a b Vyshedskiy, A. (2017). "Language evolution to revolution: from a slowly developing finite communication system with many words to infinite modern language". bioRxiv. 5: e38546. doi:10.1101/166520.
  248. ^ Kissel, M.; Fuentes, A. (2018). "'Behavioral modernity' as a process, not an event, in the human niche". Time and Mind. 11 (2): 163–183. doi:10.1080/1751696X.2018.1469230.
  249. ^ Sterelny, K. (2011). "From hominins to humans: how sapiens became behaviourally modern". Philosophical Transactions of the Royal Society B. 366 (1, 566): 809–822. doi:10.1098/rstb.2010.0301. PMC 3048993. PMID 21320896.
  250. ^ Krause, J.; Lalueza-Fox, C.; Orlando, L. (2007). "The derived FOXP2 variant of modern humans was shared with Neandertals". Current Biology. 17 (21): 1, 908–1, 912. doi:10.1016/j.cub.2007.10.008. PMID 17949978.
  251. ^ Mozzi, A.; Forni, D.; Clerici, M.; Pozzoli, U.; Mascheretti, S. (2016). "The evolutionary history of genes involved in spoken and written language: beyond FOXP2". Scientific Reports. 6: 22157. Bibcode:2016NatSR...622157M. doi:10.1038/srep22157. PMC 4766443. PMID 26912479.
  252. ^ Murphy, E.; Benítez-Burraco, A. (2017). "Paleo-oscillomics: inferring aspects of Neanderthal language abilities from gene regulation of neural oscillations" (PDF). Journal of Anthropological Sciences. 96 (96): 111–124. doi:10.4436/JASS.96010. PMID 30566085.
  253. ^ a b Rendu, W.; Beauval, C.; Crevecoeur, I.; Bayle, P.; Balzeau, A.; Bismuth, T.; Bourguignon, L.; Delfour, G.; Faivre, J.-P.; Lacrampe-Cuyaubère, F.; Muth, X.; Pasty, S.; Semal, P.; Tavormina, C.; Todisco, D.; Turq, A.; Maureille, B. (2016). "Let the dead speak…comments on Dibble et al.'s reply to 'Evidence supporting an intentional burial at La Chapelle-aux-Saints'". Journal of Archaeological Science. 69: 12–20. doi:10.1016/j.jas.2016.02.006.
  254. ^ Gargett, R. H. (1989). "Grave Shortcomings: The Evidence for Neandertal Burial". Current Anthropology. 30 (2): 157–190. doi:10.1086/203725.
  255. ^ Gargett, R. H. (1999). "Middle Palaeolithic burial is not a dead issue: the view from Qafzeh, Saint-Césaire, Kebara, Amud, and Dederiyeh". Journal of Human Evolution. 37 (1): 27–90. doi:10.1006/jhev.1999.0301. PMID 10375476.
  256. ^ a b c Wunn, I. (2001). "Cave bear worship in the Paleolithic" (PDF). Cadernos do Laboratorio Xeolóxico de Laxe. 26: 457–463.
  257. ^ Rendu, W.; Beauval, C.; Crevecoeur, I.; Bayle, P.; Balzeau, A.; Bismuth, T.; Bourguignon, L.; Delfour, G.; Faivre, J. P.; Lacrampe-Cuyaubère, F.; Tavormina, C.; Todisco, D.; Turq, A.; Maureille, B. (2014). "Evidence supporting an intentional Neandertal burial at La Chapelle-aux-Saints". Proceedings of the National Academy of Sciences. 111 (1): 81–86. Bibcode:2014PNAS..111...81R. doi:10.1073/pnas.1316780110. PMC 3890882. PMID 24344286.
  258. ^ Dibble, H.; Aldeias, V.; Goldberg, P.; Sandgathe, D.; Steele, T. E. (2015). "A critical look at evidence from La Chapelle-aux-Saints supporting an intentional burial". Journal of Archaeological Science. 53: 649–57. doi:10.1016/j.jas.2014.04.019.
  259. ^ Leroi Gourhan, A. (1975). "The flowers found with Shanidar IV, a Neanderthal burial in Iraq". Science. 190 (4214): 562–564. Bibcode:1975Sci...190..562L. doi:10.1126/science.190.4214.562.
  260. ^ Solecki, R. S. (1975). "Shanidar IV: a Neanderthal flower burial in northern Iraq". Science. 190 (4217): 880–881. Bibcode:1975Sci...190..880S. doi:10.1126/science.190.4217.880.
  261. ^ Sommer, J. D. (1999). "The Shanidar IV 'flower burial': a re-evaluation of Neanderthal burial ritual". Cambridge Archaeological Journal. 9 (1): 127–129. doi:10.1017/s0959774300015249.
  262. ^ Winzeler, Robert L. (2007). Anthropology and Religion: What We Know, Think, and Question. Altamira Press. p. 51. ISBN 978-0759110465.
  263. ^ Carroll, M. P. (1986). "The bear cult that wasn't: A study in the psychohistory of anthropology". Journal of Psychoanalytic Anthropology. 19 (1): 19–34.
  264. ^ Blanc, A. C. (1939). "Some evidence for the early ideologies of man". In Washburn, S. L. (ed.). Social life of early man. Routledge. pp. 124–126. ISBN 978-1-136-54361-6.
  265. ^ White, T. D.; Toth, N.; Chase, P. G. (1991). "The Question of Ritual Cannibalism at Grotta Guattari". Current Anthropology. 32 (7): 118–138. doi:10.1086/203931. JSTOR 2743640.
  266. ^ a b c Qiaomei Fu; Mateja Hajdinjak; Oana Teodora Moldovan; Silviu Constantin; Swapan Mallick; Pontus Skoglund; Nick Patterson; Nadin Rohland; Iosif Lazaridis; Birgit Nickel; Bence Viola; Kay Prüfer; Matthias Meyer; Janet Kelso; David Reich; Svante Pääbo (2015). "An early modern human from Romania with a recent Neanderthal ancestor". Nature. 524 (7564): 216–219. Bibcode:2015Natur.524..216F. doi:10.1038/nature14558. PMC 4537386. PMID 26098372. [1]
  267. ^ Sankararaman, S.; Patterson, N.; Li, H.; Pääbo, S.; Reich, D; Akey, J. M. (2012). "The Date of Interbreeding between Neandertals and Modern Humans". PLoS Genetics. 8 (10): e1002947. arXiv:1208.2238. Bibcode:2012arXiv1208.2238S. doi:10.1371/journal.pgen.1002947. PMC 3464203. PMID 23055938.
  268. ^ Yang, M. A.; Malaspinas, A. S.; Durand, E. Y.; Slatkin, M. (2012). "Ancient Structure in Africa Unlikely to Explain Neanderthal and Non-African Genetic Similarity". Molecular Biology and Evolution. 29 (10): 2, 987–2, 995. doi:10.1093/molbev/mss117. PMC 3457770. PMID 22513287.
  269. ^ Yotova, V.; Lefebvre, J.-F.; Moreau, C.; Gbeha, E.; Hovhannesyan, K.; Bourgeois, S.; Bédarida, S.; Azevedo, L.; Amorim, A.; Sarkisian, T.; Avogbe, P. H.; Chabi, N.; Dicko, M. H.; Kou' Santa Amouzou, E. S.; Sanni, A.; Roberts-Thomson, J.; Boettcher, B.; Scott, R. J.; Labuda, D. (2011). "An X-Linked Haplotype of Neandertal Origin is Present Among All Non-African Populations". Molecular Biology and Evolution. 28 (7): 1957–62. doi:10.1093/molbev/msr024. PMID 21266489.
  270. ^ Lohse, Konrad; Frantz, Laurent A. F. (2013). "Maximum likelihood evidence for Neandertal admixture in Eurasian populations from three genomes". Populations and Evolution. 1307: 8263. arXiv:1307.8263. Bibcode:2013arXiv1307.8263L.
  271. ^ Prüfer, K.; de Filippo, C.; Grote, S.; Mafessoni, F.; Korlević, P.; Hajdinjak, M.; et al. (2017). "A high-coverage Neandertal genome from Vindija Cave in Croatia". Science. 358 (6363): 655–58. Bibcode:2017Sci...358..655P. doi:10.1126/science.aao1887. PMC 6185897. PMID 28982794.
  272. ^ Pääbo, S. (2015). "The diverse origins of the human gene pool". Nature Reviews Genetics. 16 (6): 313–314. doi:10.1038/nrg3954. PMID 25982166.
  273. ^ Enard, D.; Petrov, D. A. (2018). "Evidence that RNA viruses drove of adaptive introgression between Neanderthals and modern humans". Cell. 175 (2): 360–371. doi:10.1016/j.cell.2018.08.034. PMC 6176737. PMID 30290142.
  274. ^ Sankararaman, S.; Patterson, N.; Li, H.; Pääbo, S.; Reich, D. (2012). "The Date of Interbreeding between Neandertals and Modern Humans". PLoS Genetics. 8 (10): e1002947. PMC 3464203. PMID 23055938.
  275. ^ Hershkovitz, I.; Weber, G. W.; Quam, R.; Duval, M.; Grün, R. (2018). "The earliest modern humans outside Africa". Science. 359 (6, 374): 459. Bibcode:2018Sci...359..456H. doi:10.1126/science.aap8369. PMID 29371468.
  276. ^ Pagani, L. (2016). "Genomic analyses inform on migration events during the peopling of Eurasia". Nature. 538 (7624): 238–242. Bibcode:2016Natur.538..238P. doi:10.1038/nature19792. PMC 5164938. PMID 27654910.
  277. ^ a b Luo, S.; Valencia, C. A.; Zhang, J.; Lee, N.-C.; Slone, J.; Gui, B.; Wang, X.; Li, Z.; Dell, S.; Brown, J.; Chen, S. M.; Chien, Y.-H.; Hwu, W.-L.; Fan, P.-C.; Wong, L.-J.; Atwal, P. S.; Huang, T. (2018). "Biparental Inheritance of Mitochondrial DNA in Humans". Proceedings of the National Academy of Sciences. 115 (51): 13039–13044. doi:10.1073/pnas.1810946115. PMC 6304937. PMID 30478036.
  278. ^ Mason, P. H.; Short, R. V. (2011). "Neanderthal-human Hybrids". Hypothesis. 9: e1. doi:10.5779/hypothesis.v9i1.215.
  279. ^ Mendez, F. L.; Poznik, G. D.; Castellano, S.; Bustamante, C. D. (2016). "The Divergence of Neandertal and Modern Human Y Chromosomes". American Journal of Human Genetics. 98 (4): 728–734. doi:10.1016/j.ajhg.2016.02.023. PMC 4833433. PMID 27058445.
  280. ^ Lowery, Robert K.; Uribe, Gabriel; Jimenez, Eric B.; Weiss, Mark A.; Herrera, Kristian J.; Regueiro, Maria; Herrera, Rene J. (2013). "Neanderthal and Denisova genetic affinities with contemporary humans: Introgression versus common ancestral polymorphisms". Gene. 530 (1): 83–94. doi:10.1016/j.gene.2013.06.005. PMID 23872234.
  281. ^ Hawks, J. (2013). "Significance of Neandertal and Denisovan Genomes in Human Evolution". Annual Review of Anthropology. 42: 433–49. doi:10.1146/annurev-anthro-092412-155548.
  282. ^ Stringer, Chris (2012). "Evolution: What makes a modern human". Nature. 485 (7396): 33–35. Bibcode:2012Natur.485...33S. doi:10.1038/485033a. PMID 22552077.
  283. ^ Pääbo, S. (2014). "A mitochondrial genome sequence of a hominin from Sima de los Huesos" (PDF). Nature. 505 (7483): 403–406. Bibcode:2014Natur.505..403M. doi:10.1038/nature12788. PMID 24305051.
  284. ^ Warren, Matthew (2018). "Mum's a Neanderthal, Dad's a Denisovan: First discovery of an ancient-human hybrid". Nature. 560 (7719): 417–418. Bibcode:2018Natur.560..417W. doi:10.1038/d41586-018-06004-0. PMID 30135540.
  285. ^ Agusti, J.; Rubio-Campillo, X. (2017). "Were Neanderthals responsible for their own extinction?". Quaternary International. 431: 232–237. Bibcode:2017QuInt.431..232A. doi:10.1016/j.quaint.2016.02.017.
  286. ^ Fontugne, M.; Reyss, J. L.; Ruis, C. B.; Lara, P. M. (1995). "The Mousterian site of Zafarraya (Granada, Spain): dating and implications on the palaeolithic peopling processes of Western Europe". Comptes Rendus de l'Académie des Sciences. 321 (10): 931–937.
  287. ^ Finlayson, C.; Pacheco, F. G. (2006). "Late survival of Neanderthals at the southernmost extreme of Europe". Nature. 443 (7, 113): 850–853. Bibcode:2006Natur.443..850F. doi:10.1038/nature05195. PMID 16971951.
  288. ^ Pavlov, P.; Svendsen, J. I.; Indrelid, S. (2001). "Human presence in the European Arctic nearly 40,000 years ago". Nature. 413 (6, 851): 64–67. Bibcode:2001Natur.413...64P. doi:10.1038/35092552. PMID 11544525.
  289. ^ Benazzi, S.; Douka, K.; Fornai, C.; Bauer, C.C.; Kullmer, O.; Svoboda, J.Í.; Pap, I.; Mallegni, F.; Bayle, P.; Coquerelle, M.; Condemi, S.; Ronchitelli, A.; Harvati, K.; Weber, G.W. (2011). "Early dispersal of modern humans in Europe and implications for Neanderthal behaviour". Nature. 479 (7374): 525–8. Bibcode:2011Natur.479..525B. doi:10.1038/nature10617. PMID 22048311.
  290. ^ Higham, T.; Compton, T.; Stringer, C.; Jacobi, R.; Shapiro, B.; Trinkaus, E.; Chandler, B.; Gröning, F.; Collins, C.; Hillson, S.; o’Higgins, P.; Fitzgerald, C.; Fagan, M. (2011). "The earliest evidence for anatomically modern humans in northwestern Europe". Nature. 479 (7374): 521–4. Bibcode:2011Natur.479..521H. doi:10.1038/nature10484. PMID 22048314.
  291. ^ Harvati, Katerina; et al. (2019). "Apidima Cave fossils provide earliest evidence of Homo sapiens in Eurasia". Nature. 571 (7766): 500–504. doi:10.1038/s41586-019-1376-z. PMID 31292546.
  292. ^ Hublin, J. (2017). "The last Neanderthal". Proceedings of the National Academy of Sciences. 40 (114): 10, 520–10, 522. doi:10.1073/pnas.1714533114. PMC 5635937. PMID 28973864.
  293. ^ Roebroeks, Wil (2006). "The human colonisation of Europe: where are we?". Journal of Quaternary Science. 21 (5): 425–435. Bibcode:2006JQS....21..425R. doi:10.1002/jqs.1044.
  294. ^ Adler, Daniel S.; Bar-Oz, Guy; Belfer-Cohen, Anna; Bar-Yosef, Ofer (2006). "Ahead of the Game: Middle and Upper Palaeolithic Hunting Behaviors in the Southern Caucasus". Current Anthropology. 47 (1): 89–118. doi:10.1086/432455. JSTOR 10.1086/432455.
  295. ^ Churchill, S. E.; Franciscus, R. G.; McKean-Peraza, H. A.; Daniel, J. A.; Warren, B. R. (2009). "Shanidar 3 Neandertal rib puncture wound and paleolithic weaponry". Journal of Human Evolution. 57 (2): 163–178. doi:10.1016/j.jhevol.2009.05.010. PMID 19615713.
  296. ^ Degioanni, A.; Bonenfant, C.; Cabut, S.; Condemi, S. (2019). "Living on the edge: Was demographic weakness the cause of Neanderthal demise?". PLOS ONE. 14 (5): e0216742. Bibcode:2019PLoSO..1416742D. doi:10.1371/journal.pone.0216742. PMC 6541251. PMID 31141515.
  297. ^ Marti, A.; Folch, A.; Costa, A.; Engwell, S. (2016). "Reconstructing the plinian and co-ignimbrite sources of large volcanic eruptions: A novel approach for the Campanian Ignimbrite". Scientific Reports. 6 (21, 220): 1–11. Bibcode:2016NatSR...621220M. doi:10.1038/srep21220. PMC 4756320. PMID 26883449.

Further readingEdit

External linksEdit