Compared to modern humans, Neanderthals were stockier, with shorter legs and bigger bodies. In conformance with Bergmann's rule, as well as Allen's rule, this was likely an adaptation to preserve heat in cold climates. Male and female Neanderthals had cranial capacities averaging 1,600 cm3 (98 cu in) and 1,300 cm3 (79 cu in), respectively, within the range of the values for anatomically modern humans. Average males stood around 164 to 168 cm (5.4 to 5.5 ft) and females 152 to 156 cm (5.0 to 5.1 ft) tall.
Neanderthal 1 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". The binomial name Homo neanderthalensis--extending the name "Neanderthal man" from the individual type specimen to the entire group--was first proposed by the Anglo-Irish geologist William King in a paper read to the British Association in 1863, although in the following year he stated that the specimen was not human and rejected the name. King's name had priority over the proposal put forward in 1866 by Ernst Haeckel, Homo stupidus. Popular English usage of "Neanderthal" as shorthand for "Neanderthal man", as in "the Neanderthals" or "a Neanderthal", emerged in the popular literature of the 1920s.
Since the historical spelling -th- in German represents the phoneme /t/, not the fricative /?/, standard British pronunciation of "Neanderthal" is with /t/ (IPA: /ni:'ænd?rt?:l/). Because of the usual sound represented by the digraph ?th? in English, "Neanderthal" is also pronounced with the voiceless fricative /?/ (as /ni:'ænd?r:l/).
The spelling Neandertal is occasionally seen in English, even in scientific publications.
Since "Neanderthal", or "Neandertal", is a common name, there is no authoritative prescription on its spelling, unlike the spelling of the binominal name H. neanderthalensis, which is predicated by King 1864. 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-1900 orthography.
Ever since the discovery of the Neanderthal fossils, expert opinion has been divided as to whether Neanderthals should be considered a separate species (Homo neanderthalensis) or a subspecies (Homo sapiens neanderthalensis) relative to modern humans.Svante Pääbo (2014) described such "taxonomic wars" as unresolveable in principle, "since there is no definition of species perfectly describing the case." The question depends on the definition of Homo sapiens as a chronospecies, which has also been in flux throughout the 20th century. Authorities preferring classification of Neanderthals as subspecies have introduced the subspecies name Homo sapiens sapiens for the anatomically modern Cro-Magnon population which lived in Europe at the same time as Neanderthals, while authorities preferring classification as separate species use Homo sapiens as equivalent to "anatomically modern humans".
During the early 20th century, a prevailing view of Neanderthals as "simian", influenced by Arthur Keith and Marcellin Boule, tended to exaggerate the anatomical differences between Neanderthals and Cro-Magnons. Beginning in the 1930s, revised reconstructions of Neanderthals increasingly emphasized the similarity rather than differences from modern humans. From the 1940s throughout the 1970s, it was increasingly common to use the subspecies classification of Homo sapiens neanderthalensis vs. Homo sapiens sapiens. The hypothesis of "multiregional origin" of modern man was formulated in the 1980s on such grounds, arguing for the presence of an unbroken succession of fossil sites in both Europe and Asia. Hybridization between Neanderthals and Cro Magnon had been suggested on skeletal and craniological grounds since the early 20th century, and found increasing support in the later 20th century, until Neanderthal admixture was found to be present in modern populations' genomes in the 2010s.
It is largely thought that Homo heidelbergensis was the last common ancestor of Neanderthals, Denisovans, and modern humans after populations became isolated in Europe, Africa, and Asia respectively. The taxonomic distinction between H. heidelbergensis and Neanderthals is mostly due to a fossil gap in Europe between 300,000 and 243,000 years ago (Marine isotope stage 8). "Neanderthals", by conventions, are fossils which date to after this gap. The quality of the fossil record greatly increases from 130,000 years ago onwards. Specimens younger than this date make up the bulk of known Neanderthal skeletons and were the first whose anatomy was comprehensively studied. In morphological studies, the term "classic Neanderthal" may be used in a narrower sense for Neanderthals younger than 71,000 years old (MIS 4 and 3).
Numerous dates for the Neanderthal/human split exist. The date of around 250,000 years ago cites the Florisbad Skull ("H. helmei") as being the ancestral species, and the split is associated with the Levallois technique of making stone tools. The date of 400,000 years ago uses Homo rhodesiensis as the ancestral species. 600,000 years ago claims that H. heidelbergensis only inhabited Europe and directly evolved into Neanderthal, classified as a chronospecies. 800,000 has Homo antecessor as the ancestral species, however, different variations of this model would push the date back to 1 million years ago. DNA studies have yielded various results on divergence time, such as 182-592 kya, 321-553 kya, 475-654 kya, 550-690 kya, 550-765 kya, 520-800 kya, and so forth.
The time of divergence between archaic Homo sapiens and the ancestors of the Neanderthals and the Denisovans was dated at 744,000 years ago, and was likely caused by a population bottleneck of the latter. Neanderthals diverged from Denisovans about 300 generations after that split.
Distribution of Neanderthals and locations of main excavation sites
Early Neanderthals, living before the Eemian interglacial (130 ka), are poorly known and come mostly from European sites. From 130 ka onwards, the quality of the fossil record increases dramatically, recorded from Western, Central, Eastern, and Mediterranean Europe, as well as Southwest, Central, and Northern Asia up to the Altai Mountains in Siberia. The southernmost find was recorded at about 30°N at Shuqba Cave, Levant, and the easternmost as 85°E at Denisova Cave, Siberia.
Sites where "classic Neanderthal" fossils (70-40 kya) have been found. Ice sheets during the Last Glacial Maximum (approx. 25-15 kya) are indicated. These are significantly larger than at 70-40 kya.
The limit of their northern range appears to have been south of
53°N (Bontnewydd, Wales),
although it is difficult to assess because glacial advances destroy most human remains, the Bontnewydd tooth being exceptional. Middle Palaeolithic artefacts have been found up to 60°N on the Russian plains.
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.
Various studies, using mtDNA analysis, yield varying effective populations, such as about 15,000 individuals (corresponding to a total population of roughly 150,000 individuals); 5,000 to 9,000 remaining constant; or 3,000 to 25,000 steadily increasing until 50,000 BCE before declining until extinction. However, all agree on low population, which may have been up to 10 times smaller than contemporary human populations in Western Europe. This 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 150,000 year stagnation in Neanderthal technology.
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.
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. However, it has also been estimated that the age pyramids for Neanderthals and Upper Paleolithic humans were the same. Infant mortality was very high, about 43% in northern Eurasia.
Analysis of the genomic DNA from three locations suggests that about 120,000 years ago there were genetically distinct Neanderthal populations in Western Europe and Siberia. Later specimens show that the western population not only still populated Europe through 40,000 years ago, but also had spread east, being present at the Siberian locale by 90,000 years ago.
Comparison of faces of early European Homo sapiens (left) and Homo neanderthalensis (right) based on forensic facial reconstructions exhibited at the Neanderthal Museum.
Neanderthal anatomy differed from modern humans in that they had a more robust build and distinctive morphological features, especially on the cranium, which gradually accumulated more derived aspects as it was described by Marcellin Boule, particularly in certain isolated geographic regions. These include shorter limb proportions, a wider, barrel-shaped rib cage, a reduced chin, sloping forehead, and a large nose, being at the modern human higher end in both width and length,[b] and started somewhat higher on the face than in modern humans. The Neanderthal skull is typically more elongated and less globular than that of anatomically modern humans, and features a notable occipital bun. Inherited Neanderthal DNA variants may subtly influence the skull shape of living people.
Neanderthals were much stronger than modern humans, with particularly strong arms and hands, while they were comparable in height; based on 45 long bones from 14 males and 7 females, three different methods of estimating height produced averages for Neanderthal males from 164 to 168 cm (65 to 66 in) and 152 to 156 cm (60 to 61 in) for females. Samples of 26 specimens found an average weight of 77.6 kg (171 lb) for males and 66.4 kg (146 lb) for females.
Neanderthals had a large cranial capacity, which at 1,600 cm3 (98 cu in) is larger on average than that of modern humans. One study has found that drainage of the dural venous sinuses (low pressure blood vessels that run between the meninges and skull leading down through the skull) in the occipital lobe region of Neanderthal brains appears more asymmetric than other hominid brains. Three-dimensional computer-assisted reconstructions of Neanderthal infants based on fossils from Russia and Syria indicated that Neanderthal and modern human brains were the same size at birth, but that by adulthood, the Neanderthal brain was larger than the modern human brain. They had almost the same degree of encephalisation (i.e. brain-to-body-size ratio) as modern humans.
Adaptations to the cold
Three-dimensional reconstructions of nasal cavities and computational fluid dynamics techniques have found that Neanderthals and modern humans both adapted their noses (independently and in a convergent way) to help breathe in cold and dry conditions. The large nose seen in Neanderthals, as well as Homo heidelbergensis, affected the shape of the skull and the muscle attachments, and gave them a weaker bite force than in modern humans. Larger eye sockets and larger areas of the brain devoted to vision suggest that their eyesight may have been better than that of modern humans. Dental remains from two Italian sites indicate that Neanderthal dental features had evolved by around 450,000 years ago during the Middle Pleistocene epoch.
Two Neanderthal specimens from Italy and Spain were found to have an allele of the melanocortin 1 receptor (MC1R) with reduced activity. This receptor plays a role in mammalian pigmentation, and the activity of the novel allele in Neanderthals was found to be reduced sufficiently to allow for visibly lighter pigment expression. Although not found in the small European sample studied by Lalueza et al., a larger study found that the derived variant was present at 70% frequency in Taiwanese Aborigines, 50% frequency in Cheyenne Native Americans, 30% frequency in Han Chinese, and 5% frequency in Europeans. It is therefore unclear whether this loss-of-function variant is responsible for any other traits other than lightening the skin (such as red or blonde hair). This allele was not found in the Croatian or Altai Neanderthal specimens subjected to whole-genome sequencing, nor have the MC1R variants known to cause red hair in modern humans, though the Altai specimen was polymorphic for another variant MC1R allele of unknown effect. Genomic analysis of three Croatian specimens for the alleles of numerous genes that affect pigment in modern humans showed the Neanderthals to have more of the alleles that produce dark pigment in modern humans than those producing reduced pigmentation. Based on this they concluded these Neanderthals had darker hair, skin and eye coloration than modern Europeans. Skin pigmentation prediction for archaic humans is controversial, as there are no living samples in which to evaluate the effect of SNP variants, and, with the tested samples coming from a single Neanderthal population, they may not be representative of the diversity across Neanderthals' full geographic range.
The overall shorter limbs and in general more stout body proportions of Neanderthals may have been an adaptation to colder climates. In comparison to modern humans, Neanderthals were more suited for sprinting and pouncing activities rather than endurance running, which would have been adaptive in the forests and woodlands that seem to have been their preferred environment. Genomic evidence possibly points to a higher proportion of fast-twitch muscle fiber in the Neanderthal. Evidence suggests that Neanderthals walked upright much like modern humans.
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.
Maximum lifespan, and the timing of adulthood, menopause, and gestation were most likely very similar to modern humans. However, it has been hypothesized that Neanderthals matured faster than modern humans based on the growth rates of teeth and tooth enamel, though this is not backed by age biomarkers.
Neanderthals suffered a high rate of traumatic injury, with by some estimates 79% of specimens showing evidence of healed major trauma. It was thus theorized that Neanderthals employed a risky hunting strategy. However, rates of cranial trauma are not significantly different between Neanderthal and middle paleolithic anatomically modern human samples.
Artist's reconstruction of a Neanderthal man and child
Neanderthals likely lived in much smaller and more sparsely distributed groups than Upper Paleolithic humans. 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". Small parties composed no more of perhaps three members may have went off on days-long hunting trips, venturing off on average 10-12 km (6.2-7.5 mi) from base camp, though possibly as much as 25 km (16 mi). Bands moved between certain caves depending on the season, and likely returned to the same locations generation after generation.
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. In humans, 20 or more 'bands' often create cultural ties and self-identify as a single people, but population density was likely too low for this to occur in Neanderthals. 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. 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, 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. It has been proposed Neanderthals would have had to have maintained strong alliances to cope with leaner times and enemies as is exhibited in the low-density hunter gatherer societies of the Western Desert of Australia. 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 or mating networks could have formed, collectively encompassing 13,000 km2 (5,000 sq mi), with each band claiming 1,200-2,800 km2 (460-1,080 sq mi).
It is sometimes suggested, since they were hunters of 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 were involved in hunting, instead of just men hunting and women and children foraging. This is used as an explanation as to why contemporary humans achieved higher populations, as they could better exploit resources than Neanderthals. However, in modern human societies, the higher the meat dependency, the higher the division of labour.
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. Like human societies, social hierarchies, if they did form, would have been most common in areas with abundant resources, such as Vézère Valley.
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. Upon reaching adolescence, and individual would join in hunting large game. 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.
Neanderthal diet may have varied significantly region to region, with some communities subsisting primarily on a plant-based diet. Edible plant remains are recorded from several caves. For example, study looking at Neanderthals from Sidrón Cave in Spain, based on dental tarter, found a meatless diet of mushrooms, pine nuts, and moss, indicating they were forest foragers. Neanderthal faecal matter from El Salt, Spain, dated to 50,000 years BCE-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). The low-calorie plant diet suggests the minimum daily caloric intake was low. 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. Remnants from the Israeli Amud Cave indicates a diet of figs, palm tree fruits, and various cereals and edible grasses. 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.
It is possible Neanderthals could employ a wide range of cooking techniques such as roasting, and may have been able to heat up or boil soup, stew, or animal stock. 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 died of starvation. These methods would have increased protein consumption to make up for a lack of carbohydrates. At Sidrón Cave, Neanderthals likely cooked and possibly smoked food, as well as used certain plants-such as yarrow and camomile-to flavor food, 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. They likely lacked any method of storing food, and had to eat whatever they hunted or foraged immediately.
There are several examples of Neanderthals practising cannibalism. The first undisputed example came from Gran Dolina, and other examples were found at Sidrón Cave,Zafarraya in Spain; and the French Moula-Guercy Cave,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 morrow, 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.
Proposed Neanderthal jewelry: white-tailed eagle claw with striations at the Neanderthal site of Krapina, Croatia, circa 130,000 BP.
Nonetheless, a large number of other claims of Neanderthal art, adornment, and structures have been made. These are often taken by the media as showing Neanderthals were capable of symbolic thought, or were "mental equals" to anatomically modern humans. As evidence of symbolism, none of them are widely accepted, although the same is true for Middle Palaeolithic anatomically modern humans. Among many others:
Incision-decorated raven bones from the Zaskalnaya VI (Kolosovskaya) Neanderthal site, Crimea, Micoquian industry dated to between cal. 43,000 and 38,000 BP. 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.
Flower pollen on the body of pre-Neanderthal Shanidar 4, Iraq, had in 1975 been argued to be a flower burial, but the pollen could have also been deposited by natural events.
Bird bones were argued to show evidence for feather plucking in a 2012 study examining 1,699 ancient sites across Eurasia, which the authors controversially took to mean Neanderthals wore bird feathers as personal adornments.
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. The authors claim artificial lighting would have been required as this part of the cave is beyond the reach of daylight.
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.
In 2018, red painted symbols comprising hand stencils, ladder-shaped figured, prints, dots, discs, lines, and representations of animals on the cave walls of three Spanish caves 700 km (430 mi) apart-La Pasiega, Maltravieso, and Doña Trinidad-were dated to be older than, respectively, 64,800; 66,700; and for one painting 65,500 years ago. If the dating is correct, they were painted before the time anatomically modern humans are thought to have arrived in Europe, and demonstrate Neanderthals were capable of symbolic behaviour.
In 2018, perforated seashell beads and pigments that are at least 115,000 years old were found in Cueva de los Aviones in southeastern Spain.
Neanderthals made stone tools, used fire, and were hunters. This is the extent of the consensus on their behaviour. It had long been debated whether Neanderthals were hunters or scavengers, but the discovery of the pre-Neanderthal Schöningen wooden spears in Germany helped settle the debate in favour of hunting. Neanderthals are associated with the Mousterian stone tool industry. The Levallois technique they adopted maximizes the cutting surface with the least amount of raw material. As a difficult-to-learn process, the technique may have been directly taught generation to generation rather than via purely observational learning. 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.
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.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. 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.
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. A study from 2019 showed that birch tar production can instead be a very simple process - merely involving the burning of birch bark near smooth vertical surfaces in open air conditions.
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. They probably did not need tailored clothings as humans due to their anatomical adaptations to the cold, though this may not have made such an impactful difference. Nonetheless, Neanderthals probably covered up to 70-80% of their body, and, depending on the frigidness, the hands and feet; though contemporary humans would have covered 80-90%. 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.
Remains of Middle Paleolithic stone tools on Greek islands indicate early seafaring of 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. Other Mediterranean islands include Sardinia, Melos, Alonnisos, and it is possible they crossed the Strait of Gibraltar. Their ability to engineer these boats and navigate through open waters speaks to their advanced cognitive and technical skills.
Claims that Neanderthals held funerals for their dead with symbolic meaning,:158-60 are heavily contested. 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. 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. 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. The medicinal properties of the plants led American archaeologist Ralph Solecki to claim that the man buried was some leader, healer, or shaman. However, it is also possible the pollen was deposited by a small rodent after the man's death. 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.
It was once asserted that the bones of the short-faced bear in some European caves were arranged in a particular 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 'peculiar' arrangement of bones could also be well-explained by natural causes, 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.
It was also once thought Neanderthals hunted, killed, and cannabalised other Neanderthals, and used the skull as the focus of some ritual. It is now thought that isolated damaged skull remains-which were put forward as evidence of the skull cult-were the result of predation by cave hyenas or simple decay. Nonetheless, Neanderthals are known to have practiced cannibalism, but there is unsubstantial evidence to suggest ritual defleshing.
An alternative proposal regarding the fate of Neanderthals is that rather than being replaced by modern humans and going extinct, Neanderthals were absorbed into the Cro-Magnon population by interbreeding. This would be counter to strict versions of theory of recent African origin of modern humans, since it would imply that at least part of the genome of Europeans would descend from Neanderthals. Evidence of interbreeding has been detected in genomes of ancient and modern humans, though this could have resulted from occasional interbreeding and introgression, rather than population absorption.
Pre-2010 interbreeding hypotheses
Until the early 1950s, most scholars thought Neanderthals were not in the ancestry of living humans.:232-34 Nevertheless, in 1904 Thomas H. Huxley saw among Frisians the presence of what he suspected to be Neanderthaloid skeletal and cranial characteristics as an evolutionary development from Neanderthal rather than as a result of interbreeding, saying that "the blond long-heads may exhibit one of the lines of evolution of the men of the Neanderthaloid type," yet he raised the possibility that the Frisians alternatively "may be the result of the admixture of the blond long-heads with Neanderthal men," thus separating "blond" from "Neanderthaloid."
Hans Peder Steensby proposed interbreeding in 1907 in the article Race studies in Denmark. He strongly emphasised that all living humans are of mixed origins. He held that this would best fit observations, and challenged the widespread idea that Neanderthals were ape-like or inferior. Basing his argument primarily on cranial data, he noted that the Danes, like the Frisians and the Dutch, exhibit some Neanderthaloid characteristics, and felt it was reasonable to "assume something was inherited" and that Neanderthals "are among our ancestors."
Carleton Stevens Coon in 1962 found it likely, based upon evidence from cranial data and material culture, that Neanderthal and Upper Paleolithic peoples either interbred or that the newcomers reworked Neanderthal implements "into their own kind of tools." Christopher Thomas Cairney in 1989 went further, laying out a rationale for hybridisation and adding a broader discussion of physical characteristics as well as commentary on interbreeding and its importance to adaptive European phenotypes. Cairney specifically discussed the "intermixture of racial elements" and "hybridisation."
In 2010, geneticists announced that interbreeding had likely taken place, a result confirmed in 2012.[page needed] The genomes of all non-Africans (such as Europeans, Asians, and Oceanians such as Papua New Guineans) include portions that are of Neanderthal origin, a share estimated in 2014 to 1.5-2.1%. This DNA is absent in Sub-Saharan Africans (Yoruba people and San subjects).Ötzi the iceman, Europe's oldest preserved mummy, was found to possess an even higher percentage of Neanderthal ancestry. The two percent of Neanderthal DNA in Europeans and Asians is not the same in all Europeans and Asians: in all, approximately 20% of the Neanderthal genome appears to survive in the modern human gene pool.
Genomic studies suggest that modern humans mated with at least two groups of archaic humans: Neanderthals and Denisovans. Some researchers suggest admixture of 3.4-7.9% in modern humans of non-African ancestry, rejecting the hypothesis of ancestral population structure., and others later suggesting Neanderthal admixture of 1.5-2.1%. Detractors have argued and continue to argue that the signal of Neanderthal interbreeding may be due to ancient African substructure, meaning that the similarity is only a remnant of a common ancestor of both Neanderthals and modern humans and not the result of interbreeding.John D. Hawks has argued that the genetic similarity to Neanderthals may indeed be the result of both structure and interbreeding, as opposed to just one or the other.
An approximately 40,000 year old anatomically-modern human skeleton from Pe?tera cu Oase, Romania, was found in 2015 to have a much larger proportion of DNA matching the Neanderthal genome than seen in humans of today, and this was estimated to have resulted from an interbreeding event as few as four generations earlier. However, this hybrid Romanian population does not appear to have made a substantial contribution to the genomes of later Europeans.
While some modern human nuclear DNA has been linked to the extinct Neanderthals, no mitochondrial DNA of Neanderthal origin has been detected, which in primates is almost always maternally transmitted. This observation has prompted the hypothesis that whereas female humans interbreeding with male Neanderthals were able to generate fertile offspring, the progeny of female Neanderthals who mated with male humans were either rare, absent or sterile.
Eastern Neanderthals from the Altai show evidence of an introgression from modern humans not seen in western Neanderthals. This contribution to their genome derived from a modern human population that diverged from most other modern humans about 120 kya and expanded from Africa into Eurasia, but which was later largely replaced by a second expansion of modern humans out of Africa after 75,000 years ago that gave rise to modern Eurasians, although 2% of the genome of New Guineans derives from this earlier dispersal. Kuhlwilm et al. argue that the admixture between this early modern human group, modern Eurasians, and Neanderthals took place in Southern Arabia or the Levant and that the introgressed Siberian Neanderthals had spread there from the Middle East.
Interbreeding with Denisovans
Sequencing of the genome of a Denisovan, a distinct but related archaic hominin, from the Denisova cave in the Siberian Altai region has shown that 17% of its genome represents Neanderthal DNA. This Neanderthal DNA present in the Denisovan genome more closely resembled that found in the genome of a 120,000-year-old Neanderthal bone found in the same cave than that of Neanderthals from the Vindija cave in Croatia or the Mezmaiskaya cave in the Caucasus, suggesting that the gene flow came from a local interbreeding. However, the complete genome sequencing of DNA from a 90,000-year-old bone fragment, Denisova 11, showed it to have belonged to a Denisovan-Neanderthal hybrid whose father was a typical Denisovan with the Altai Neanderthal component dating to an interbreeding more than 300 generations earlier, but the specimen's mother was a Neanderthal belonging to a population more closely related to the Vindija Neanderthal than to the sequenced Altai Neanderthal genome. This suggests mobility or turnover among the distinct Neanderthal populations.
The mtDNA phylogeny of the Neanderthals, Denisovans, and modern humans produces a puzzling picture. Based on mtDNA sequences of Neanderthals from the Vindija and Altai sites, the most recent common ancestor of modern humans and Neanderthals lived approximately 440,000 years ago, while that between Neanderthals and Denisovans is far back as 830,000 years. This contrasts with nuclear genome analysis, in which the Neanderthals and Denisovans are sister groups. The mtDNA of the 400,000-year-old Sima de los Huesos sample, thought to be ancestral to Western European Neanderthals, is much closer to Denisovan mtDNA. Paabo et al. suggest several alternative explanations for this, including genetic substructure within the populations, introgression of mtDNA from one group to another, or incorrect taxonomic placement of the Sima de los Huesos hominins. Peyrégne et al. again suggested genetic substructure or introgression as possible explanations for an approximately 124,000-year-old German Neanderthal specimen with mtDNA that diverged from that of other Neanderthal specimens (except for Sima de Huesos) about 270,000 years ago, while its genomic DNA was consistent with divergence less than 150,000 years ago.
According to a 2014 study by Thomas Higham and colleagues of organic samples from European sites, Neanderthals died out in Europe between 41,000 and 39,000 years ago.[d] New dating in Iberia, where Neanderthal dates as late as 24,000 years had been reported before, now suggests evidence of Neanderthal survival in the peninsula after 42,000 years ago is almost non-existent.
mtDNA-based simulation of modern human expansion in Europe starting 1,600 generations ago. Neanderthal range in light grey
Anatomically modern humans arrived in Mediterranean Europe between 45,000 and 43,000 years ago, so the two different human populations shared Europe for several thousand years. The exact nature of biological and cultural interaction between Neanderthals and other human groups is contested.
Possible scenarios for the extinction of the Neanderthals are:
Neanderthals were a separate species from modern humans, and became extinct (because of climate change or interaction with modern humans) and were replaced by modern humans moving into their habitat between 45,000 and 40,000 years ago.Jared Diamond has suggested a scenario of violent conflict and displacement.
Neanderthals were a contemporary subspecies that bred with modern humans and disappeared through absorption (interbreeding theory).
About 55,000 years ago, the climate began to fluctuate wildly from extreme cold conditions to mild cold and back in a matter of decades. Neanderthal bodies were well-suited for survival in a cold climate--their stocky chests and limbs stored body heat better than the Cro-Magnons. Neanderthals died out in Europe between 41,000 and 39,000 years ago, coinciding with the start of a very cold period.
Raw material sourcing and the examination of faunal remains found in the southern Caucasus suggest that modern humans may have had a survival advantage, being able to use social networks to acquire resources from a greater area. In both the Late Middle Palaeolithic and Early Upper Palaeolithic more than 95% of stone artifacts were drawn from local material, suggesting Neanderthals restricted themselves to more local sources.
Coexistence with modern humans
The exact nature of biological and cultural interaction between Neanderthals and other human groups has been contested. The dating of the period of overlap has been repeatedly revised. In 2011, this period of coexistence was extended when what were previously thought to be Neanderthal baby teeth, unearthed in 1964 from the Grotta del Cavallo in Italy, were identified as the oldest modern human remains discovered anywhere in Europe, dating from between 43,000 and 45,000 years ago. At that time, Neanderthal samples from Iberia were thought to have dated as recently as 24,000 years ago, but a 2014 study of Neanderthal bones and tools led to a redating of these samples, and indicated that Neanderthals died out in Europe between 41,000 and 39,000 years ago, with the two different human populations sharing Europe only for about 5,000 years. However, a 2019 reanalysis of skull fragments from southern Greece assumed to have belonged to Neanderthals concluded that they belonged to a modern human, and dated them to 210,000 years ago, vastly extending the period of potential overlap in Europe between the two groups.
Neanderthals inhabited that continent long before the arrival of modern humans. These modern humans may have introduced a disease that contributed to the extinction of Neanderthals, and that may be added to other recent explanations for their extinction. When Neanderthal ancestors left Africa potentially as early as over 800,000 years ago they adapted to the pathogens in their European environment, unlike modern humans, who adapted to African pathogens. This transcontinental movement is known as the Out of Africa model. If contact between humans and Neanderthals occurred in Europe and Asia the first contact may have been devastating to the Neanderthal population, because they would have had little, if any, immunity to the African pathogens. More recent historical events in Eurasia and the Americas show a similar pattern, whereby the unintentional introduction of viral or bacterial pathogens to unprepared populations has led to mass mortality and local population extinction. The most well-known example of this is the arrival of Christopher Columbus to the New World, which brought and introduced foreign diseases when he and his crew arrived to a native population who had no immunity.
1856: Limestone miners discover the Neanderthal-type specimen, Neanderthal 1, in Neandertal, western Prussia (now Germany).
1864: William King is the first to recognise Neanderthal 1 as belonging to a separate species, for which he gives the scientific name Homo neanderthalensis. He then changed his mind on placing it in the genus Homo, arguing that the upper skull was different enough to warrant a separate genus since, to him, it had likely been "incapable of moral and theistic conceptions."
1880: The mandible of a Neanderthal child is discovered in a secure context in ?ipka cave, in the Austro-Hungarian Empire (now the Czech Republic), associated with cultural debris, including hearths, Mousterian tools, and bones of extinct animals.
1886: Two well-preserved Neanderthal skeletons are found at Spy, Belgium, making the hypothesis that Neanderthal 1 was only a diseased modern human difficult to sustain.
1899: Sand excavation workers find hundreds of fragmentary Neanderthal remains representing at least 12 and likely as much as 70 individuals on a hill in Krapina, in the Austro-Hungarian Empire (now Croatia).
Neanderthal 1: The first human bones recognised as showing a non-modern anatomy. Discovered in 1856 in a limestone quarry at the Feldhofer grotto in Neanderthal, Western Prussia (now Germany), they consist of a skull cap, the two femora, the three right arm bones, two left arm bones, the ilium, and fragments of a scapula and ribs.
La Chapelle-aux-Saints 1: Called the Old Man, a fossilised skeleton discovered in La Chapelle-aux-Saints, France, by A. and J. Bouyssonie, and L. Bardon in 1908. Characteristics include a low vaulted cranium and large browridge typical of Neanderthals. Estimated to be about 60,000 years old, the specimen was severely arthritic and had lost all his teeth long before death, leading some to suggest he was cared for by others.
La Ferrassie 1: A fossilised skull discovered in La Ferrassie, France, by R. Capitan in 1909. It is estimated to be 70,000 years old. Its characteristics include a large occipital bun, low-vaulted cranium and heavily worn teeth.
Le Moustier 1: One of the rare nearly complete Neanderthal skeletons to be discovered, it was excavated by a German team in 1908, at Peyzac-le-Moustier, France. Sold to a Berlin museum, the post cranial skeleton was bombed and mostly destroyed in 1945, and parts of the mid face were lost sometime after then. The skull, estimated to be less than 45,000 years old, includes a large nasal cavity and a less developed brow ridge and occipital bun than seen in other Neanderthals. The Mousterian tool techno-complex is named after its discovery site.
Shanidar 1 to 10: Eight Neanderthals and two pre-Neanderthals (Shanidar 2 and 4) were discovered in the Zagros Mountains in Iraqi Kurdistan. One of the skeletons, Shanidar 4, was once thought to have been buried with flowers, a theory no longer accepted. To Paul B. Pettitt the "deliberate placement of flowers has now been convincingly eliminated", since "[a] recent examination of the microfauna from the strata into which the grave was cut suggests that the pollen was deposited by the burrowing rodent Meriones tersicus, which is common in the Shanidar microfauna and whose burrowing activity can be observed today".
Amud 1: A male adult Neanderthal, dated to roughly 55,000 BP, and one of several found in a cave at Nahal Amud, Israel. At 178 cm (70 in), it is the tallest known Neanderthal. It also has the largest cranial capacity of all extinct hominins: 1,736 cm3.
Kebara 2: A male adult post-cranial skeleton, dated to roughly 60,000 BP, that was discovered in 1983 in Kebara Cave, Israel. It has been studied extensively, for its hyoid, ribcage, and pelvis are much better preserved than in all other Neanderthal specimens.
Notable Central Asian Neanderthal
Teshik-Tash 1: The skeleton of an 8-11-year-old child discovered in Uzbekistan by Okladnikov in 1938. This is the only fairly complete skeleton discovered to the east of Iraq. Okladnikov claimed it was a deliberate burial, but this is debated.
Neanderthals have been portrayed in popular culture including appearances in literature, visual media and comedy. Early 20th century artistic interpretations often presented Neanderthals as beastly creatures, emphasising hairiness and a rough, dark complexion.
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.
^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.
^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). Homo sapiens sapiens 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.
^Higham et al did not study samples from sites outside Europe and they stated that further work was required to rule out later survival at Gorhams Cave, Gibraltar.
^ abcT. Higham; K. Douka; R. Wood; C.B. Ramsey; F. Brock; L. Basell; M. Camps; A. Arrizabalaga; J. Baena; C. Barroso-Ruíz; 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. PMID25143113. 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.
^T. Higham (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 40,000 years ago, a period that unfortunately lies near the limit of radiocarbon dating. This paper shows that as many as 70 per cent of the oldest radiocarbon dates in the literature may be too young, due to contamination by modern carbon.(subscription required)
^ abB. Galván; C.M. Hernández; C. Mallol; N. Mercier; A. Sistiaga; V. Soler (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. PMID25016565.(subscription required)
^McKie, Robin (June 2, 2013). "Why did the Neanderthals die out?". The Guardian. Retrieved 2017. "It was once thought we appeared in Europe around 35,000 years ago and that we coexisted with Neanderthals for thousands of years after that. They may have hung on in pockets - including caves in Gibraltar - until 28,000 years ago [said Chris Stringer]" Previous research on Neanderthal sites which suggested that they were more recent than 40,000 years old appears to be wrong," said Stringer. "That is a key finding that will be discussed at the conference."[...] However, scientists have set out to get round these problems. At Oxford University, scientists led by Tom Higham have developed new methods to remove contamination and have been able to make much more precise radiocarbon dating for this period.
^Holloway, R.L. (1985). "The poor brain of Homo sapiens neanderthalensis: see what you please". In Delson, E. (ed.). Ancestors: The hard evidence. New York: Alan R. Liss. ISBN978-0471843764.; extending to 1,736 cm3 (105.9 cu in) in the male Amud 1.
^Amano, H.; Kikuchi, T.; Morita, Y.; Kondo, O.; Suzuki, Hiromasa; et al. (August 2015). "Virtual Reconstruction of the Neanderthal Amud 1 Cranium". American Journal of Physical Anthropology. 158 (2): 185-97. doi:10.1002/ajpa.22777. hdl:10261/123419. PMID26249757.
^ abHelmuth H (1998). "Body height, body mass and surface area of the Neanderthals". Zeitschrift für Morphologie und Anthropologie. 82 (1): 1-12. PMID9850627.
^ abcdefGreen, 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.; Prufer, 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.; Paabo, S. (May 6, 2010). "A Draft Sequence of the Neandertal Genome". Science. 328 (5979): 710-722. Bibcode:2010Sci...328..710G. doi:10.1126/science.1188021. PMC5100745. PMID20448178.
^Murray, John; Nasheuer, Heinz Peter; Seoighe, Cathal; McCormack, Grace P.; Williams, D. Michael & Harper, David 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, JSTOR10.3318/ijes.2015.33.1(subscription required)
^"Neanderthal". Wiley-Blackwell Encyclopedia of Human Evolution. Chichester, West Sussex: Wiley-Blackwell. 2013.
^"Neandertal oder Neanderthal? - Was ist denn nun richtig?" [Neanderthal or Neanderthal? - What is right?]. mettmann.de. Neanderthal museum. Retrieved 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'.]
^ abD. 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. PMID9614635.
^B. Vandermeersch; M.D. Garralda (2011). S. Condemi; G.-C. Weniger (eds.). "Continuity or Discontinuity in the Peopling of Europe: One Hundred and Fifty Years of Neanderthal Study". Vertebrate Paleobiology and Paleoanthropology. Springer Netherlands: 113-25. doi:10.1007/978-94-007-0492-3_10. Cite journal requires |journal= (help)
^ abRogers, 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.
^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. PMID15288521.
Slimak, L., et al. (2012). "Response to 'Comment on Late Mousterian Persistence near the Arctic Circle'" Science, 335 (6065), 167 doi:10.1126/science.1210211
Zwyns, N., et al. (2012). "Comment on 'Late Mousterian Persistence near the Arctic Circle'" Science, 335 (6065), 167 doi:10.1126/science.1209908
^ abFabrizio Mafessoni, Kay Prüfer, "Better support for a small effective population size of Neandertals and a long shared history of Neandertals and Denisovans", PNAS 114 (48), November 28, 2017, E10256-E10257, doi:10.1073/pnas.1716918114.
^ abcdefgBocquet-Appel, J.; Degioanni, A. (2013). "Neanderthal Demographic Estimates". Current Anthropology. 54: 202-214. doi:10.1086/673725.
^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. PMID15689531.
^Silberman, Neil. The Oxford Companion to Archaeology, p. 455 (Oxford University Press 2012): "[I]t is with the Neanderthals that we see the full achievement, for the first time, of the degree of encephalization (brain to body size ratio) that characterizes modern humans."
^Abramiuk, Marc. The Foundations of Cognitive Archaeology, p. 199 (MIT Press 2012): "the encephalization quotient was slightly smaller".
^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. (November 30, 2007). "A Melanocortin 1 Receptor Allele Suggests Varying Pigmentation Among Neanderthals". Science. 318 (5855): 1453-1455. Bibcode:2007Sci...318.1453L. doi:10.1126/science.1147417. PMID17962522.
^Ding, Qiliang; Hu, Ya; Xu, Shuhua; Wang, Chuan-Chao; Li, Hui; Zhang, Ruyue; Yan, Shi; Wang, Jiucun; Jin, Li (August 2014). "Neanderthal Origin of the Haplotypes Carrying the Functional Variant Val92Met in the MC1R in Modern Humans". Molecular Biology and Evolution. 31 (8): 1994-2003. doi:10.1093/molbev/msu180. PMID24916031.
^Stewart, J.R.; García-Rodríguez, O.; Knul, M.V.; Sewell, L.; Montgomery, H.; Thomas, M.G.; Diekmann, Y. (August 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.
^Churchill, Steven E.; Franciscus, Robert G.; McKean-Peraza, Hilary A.; Daniel, Julie A.; Warren, Brittany R. (August 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. PMID19615713.
^Bocherens, Hervé; Drucker, Dorothée G.; Billiou, Daniel; Patou-Mathis, Marylène; Vandermeersch, Bernard (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. PMID15869783.
^ abLev, 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.
^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.
^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. PMID21187393.
^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. doi:10.1073/pnas.0609662104.
^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. PMID29472483.
^Lycett, Stephen J.; von Cramon-Taubadel, Noreen (March 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.
^Boëda, Eric; Geneste, J. M.; Griggo, C.; Mercier, N.; Muhesen, S.; Reyss, J. L.; Taha, A.; Valladas, H. (January 2, 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.
^ abcKedar, Yafit; Barkai, Ran (2019). "The Significance of Air Circulation and Hearth Location at Paleolithic Cave Sites". Open Quaternary. 5. doi:10.5334/oq.52.
^ abCollard, Mark; Tarle, Lia; Sandgathe, Dennis; Allan, Alexander (December 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.
^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.
^ abRendu, William; Beauval, Cédric; Crevecoeur, Isabelle; Bayle, Priscilla; Balzeau, Antoine; Bismuth, Thierry; Bourguignon, Laurence; Delfour, Géraldine; Faivre, Jean-Philippe; Lacrampe-Cuyaubère, François; Muth, Xavier; Pasty, Sylvain; Semal, Patrick; Tavormina, Carlotta; Todisco, Dominique; Turq, Alain; Maureille, Bruno (May 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.
^ abGargett, R.H. (1989). "Grave Shortcomings: The Evidence for Neandertal Burial". Current Anthropology. 30 (2): 157-90. doi:10.1086/203725.
^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. PMID10375476.
^ abDibble, 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.
^Steensby, H. P. (1907). "Racestudier i Danmark" [Race Studies in Denmark] (PDF). Geographical Journal (in Danish). Royal Library, Denmark. Retrieved 2017.
^Coon, Carleton Stevens (1962). The Origin of races. New York: Knopf. pp. 548-49.
^Liu, Prugnolle et al. (2006). "Currently available genetic and archaeological evidence is supportive of a recent single origin of modern humans in East Africa. However, this is where the consensus on human settlement history ends, and considerable uncertainty clouds any more detailed aspect of human colonization history."
^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.
^Lowery, Robert K.; Uribe, Gabriel; Jimenez, Eric B.; Weiss, Mark A.; Herrera, Kristian J.; Regueiro, Maria; Herrera, Rene J. (November 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. PMID23872234.
^ abFinlayson, C.; Carrión, J. S. (April 2007). "Rapid ecological turnover and its impact on Neanderthal and other human populations". Trends in Ecology & Evolution. 22 (4): 213-22. doi:10.1016/j.tree.2007.02.001. PMID17300854.
^Genovés, S. (1954). "The problem of the sex of certain fossil hominids, with special reference to the Neandertal skeletons from Spy". The Journal of the Royal Anthropological Institute of Great Britain and Ireland. 84 (1/2): 131-44. doi:10.2307/2844006. JSTOR2844006.
^Green, Richard E.; Krause, Johannes; Ptak, Susan E.; Briggs, Adrian W.; Ronan, Michael T.; Simons, Jan F.; Du, Lei; Egholm, Michael; Rothberg, Jonathan M.; Paunovic, Maja; Pääbo, Svante (November 2006). "Analysis of one million base pairs of Neanderthal DNA". Nature. 444 (7117): 330-336. Bibcode:2006Natur.444..330G. doi:10.1038/nature05336. PMID17108958.
^Duveau, Jérémy; Berillon, Gilles; Verna, Christine; Laisné, Gilles; Cliquet, Dominique (September 24, 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): 19409-19414. doi:10.1073/pnas.1901789116. PMID31501334.
^The Neanderthal Dead, exploring mortuary variability in middle paleolithic eurasia. Paul B. Pettitt (2002)
^Rink, W. Jack; Schwarcz, H.P.; Lee, H.K.; Rees-Jones, J.; Rabinovich, R.; Hovers, E. (August 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): 701-17. doi:10.1002/gea.1017.
^Valladas, Hélène; Merciera, N.; Frogeta, L.; Hoversb, E.; Joronc, J.L.; Kimbeld, W.H.; Rak, Y. (March 1999). "TL Dates for the Neanderthal Site of the Amud Cave, Israel". Journal of Archaeological Science. 26 (3): 259-68. doi:10.1006/jasc.1998.0334.
^Wood, R. E.; Higham, T. F. G.; de Torres, T.; Tisnérate-Laborde, N.; Valladas, H.; Ortiz, J. E.; Lalueza-Fox, C.; Sánchez-Moral, S.; Cañaveras, J. C.; Rosas, A.; Santamaría, D.; de la Rasilla, M. (March 20, 2012). "A new date for the Neanderthals from El Sidrón Cave (Asturias, Northern Spain)". Archaeometry. 55 (1): 148-58. doi:10.1111/j.1475-4754.2012.00671.x. hdl:10261/79592.
Boë, Louis-Jean; Heim, Jean-Louis; Honda, Kiyoshi; Maeda, Shinji (2002). "The potential Neandertal vowel space was as large as that of modern humans". Journal of Phonetics. 30 (3): 465-84. doi:10.1006/jpho.2002.0170.
Lieberman, Philip (October 2007). "Current views on Neanderthal speech capabilities: A reply to Boe et al. (2002)". Journal of Phonetics. 35 (4): 552-63. doi:10.1016/j.wocn.2005.07.002.