Neanderthal

Neanderthal Temporal range: Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
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
Domain:	Eukaryota
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
Known Neanderthal range in Europe (blue), Southwest Asia (orange), Uzbekistan (green), and the Altai Mountains (violet)
Synonyms[6]
Homo H. stupidus Haeckel, 1895[1] H. europaeus primigenius Wilser, 1898 H. primigenius Schwalbe, 1906[2] H. antiquus Adloff, 1908 H. transprimigenius mousteriensis Farrer, 1908 H. mousteriensis hauseri Klaatsch 1909[3][4] H. priscus Krause , 1909 H. chapellensis von Buttel-Reepen, 1911 H. calpicus Keith, 1911 H. acheulensis moustieri Wiegers, 1915 H. lemousteriensis Wiegers, 1915 H. naulettensis Baudouin, 1916 H. sapiens neanderthalensis Kleinshmidt, 1922 H. heringsdorfensis Werthe, 1928 H. galilensis Joleaud, 1931 H. primigenius galilaeensis Sklerj, 1937 H. kiikobiensis Bontsch-Osmolovskii, 1940 H. sapiens krapinensis Campbell, 1962 H. erectus mapaensis Kurth, 1965 Palaeoanthropus P. neanderthalensis Keith, 1939[5] P. heidelbergensis Keith, 1939[5] P. ehringsdorfensis Paterson, 1940[5] P. krapinensis Sergi, 1911[5] P. palestinensis Keith, 1939[5] P. europaeus Sergi, 1910 Protanthropus P. atavus Haeckel, 1895 P. tabunensis Bonarelli, 1944 Acanthropus A. neanderthalensis Arldt, 1915 A. primigenius Abel, 1920 A. neanderthalensis Dawkins, 1926

Neanderthals (

The reasons for Neanderthal extinction are disputed.[16]^[17] Theories for their extinction include demographic factors such as small population size and inbreeding, competitive replacement,^[18] interbreeding and assimilation with modern humans,^[19] change of climate,^[20][21][22] disease,^[23][24] or a combination of these factors.^[22]

For much of the early 20th century, European researchers depicted Neanderthals as primitive, unintelligent and brutish. Although knowledge and perception of them has markedly changed since then in the scientific community, the image of the unevolved

Compared with modern humans, Neanderthals had a more robust build and proportionally shorter limbs. Researchers often explain these features as adaptations to conserve heat in a cold climate, but they may also have been adaptations for sprinting in the warmer, forested landscape that Neanderthals often inhabited.^[66] They had cold-specific adaptations, such as specialised body-fat storage^[67] and an enlarged nose to warm air^[68] (although the nose could have been caused by genetic drift^[69]). Average Neanderthal men stood around 165 cm (5 ft 5 in) and women 153 cm (5 ft 0 in) tall, similar to pre-industrial modern Europeans.^[70] The braincases of Neanderthal men and women averaged about 1,600 cm³ (98 cu in) and 1,300 cm³ (79 cu in), respectively,^[71][72][73] which is considerably larger than the modern human average (1,260 cm³ (77 cu in) and 1,130 cm³ (69 cu in), respectively).^[74] The Neanderthal skull was more elongated and the brain had smaller parietal lobes^[75][76][77] and cerebellum,^[78][79] but larger temporal, occipital and orbitofrontal regions.^[80][81]

The total population of Neanderthals remained low, proliferating weakly harmful gene variants^[82] and precluding effective long-distance networks. Despite this, there is evidence of regional cultures and regular communication between communities.^[83][84] They may have frequented caves and moved between them seasonally.^[85] Neanderthals lived in a high-stress environment with high trauma rates, and about 80% died before the age of 40.^[86]

The 2010

Etymology

Neanderthals are named after the Neander Valley in which the first identified specimen was found. The valley was spelled Neanderthal and the species was spelled Neanderthaler in German until the spelling reform of 1901.^[b] The spelling Neandertal for the species is occasionally seen in English, even in scientific publications, but the scientific name, H. neanderthalensis, is always spelled with th according to the principle of priority. The vernacular name of the species in German is always Neandertaler ("inhabitant of the Neander Valley"), whereas Neandertal always refers to the valley.^[c] The valley itself was named after the late 17th century German theologian and hymn writer Joachim Neander, who often visited the area.^[101] His name in turn means 'new man', being a learned Graecisation of the German surname Neumann.

Neanderthal can be pronounced using the /t/ (as in /niˈændərtɑːl/)^[104] or the standard English pronunciation of th with the fricative /θ/ (as /niˈændərθɔːl/).^[105][106]

H. heidelbergensis (Miguelón, 430,000 years ago)

Stage 3: early Neanderthal (Saccopastore I, 130,000 years ago)

Stage 4: classic European Neanderthal (La Chapelle-aux-Saints 1, 50,000 years ago)

The accretion model^[142]

Neanderthals and Denisovans are more closely related to each other than they are to modern humans, meaning the Neanderthal/Denisovan split occurred after their split with modern humans.[14]^{[92][138][158]} Assuming a mutation rate of 1 × 10⁻⁹ or 0.5 × 10⁻⁹ per base pair (bp) per year, the Neanderthal/Denisovan split occurred around either 236–190,000 or 473–381,000 years ago, respectively.^[92] Using 1.1 × 10⁻⁸ per generation with a new generation every 29 years, the time is 744,000 years ago. Using 5 × 10⁻¹⁰ nucleotide sites per year, it is 616,000 years ago. 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 600–500,000 years ago.^[138] Before splitting, Neanderthal/Denisovans (or "Neandersovans") migrating out of Africa into Europe apparently interbred with an unidentified "superarchaic" human species who were already present there; these superarchaics were the descendants of a very early migration out of Africa around 1.9 mya.^[159]

Genetic evidence suggests that following their split from Denisovans, Neanderthals experienced gene flow (around 3% of their genome) from the lineage leading to modern humans prior to the expansion of modern humans outside of Africa during the Last Glacial Period, with this interbreeding suggested to have taken place around 200-300,000 years ago.^[141]

Demographics

Range

Pre- and early Neanderthals, living before the Eemian interglacial (130,000 years ago), are poorly known and come mostly from Western European sites. From 130,000 years ago onwards, the quality of the fossil record increases dramatically with classic Neanderthals, who are recorded from Western, Central, Eastern and Mediterranean Europe,^[15] as well as Southwest, Central and Northern Asia up to the Altai Mountains in southern Siberia. Pre- and early Neanderthals, on the other hand, seem to have continuously occupied only France, Spain and Italy, although some appear to have moved out of this "core-area" to form temporary settlements eastward (although without leaving Europe). Nonetheless, southwestern France has the highest density of sites for pre-, early and classic Neanderthals.^[160] The Neanderthals were the first human species to permanently occupy Europe as the continent was only sporadically occupied by earlier humans.^[161]

The southernmost find was recorded at

Population

Like modern humans, Neanderthals probably descended from a very small population with an

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Locations of Neanderthal finds in Europe and the Levant.
View references

Show map of Europe

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Locations of Neanderthal finds in Eurasia (note, part of Spain is cut off)
View references

Show map of Asia

Anatomy

Build

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

Neanderthal skull features

Neanderthals had more robust and stockier builds than typical modern humans,^[70] wider and barrel-shaped rib cages; wider pelvises;^[25][184] and proportionally shorter forearms and forelegs.^[66][185]

Based on 45 Neanderthal

The

Neanderthals had less developed chins, sloping foreheads, and longer, broader, more projecting noses. The Neanderthal skull is typically more elongated, but also wider, and less globular than that of most modern humans, and features much more of an

The Neanderthal face is characterized by mid-facial prognathism, where the zygomatic arches are positioned in a rearward location relative to modern humans, while their maxillary bones and nasal bones are positioned in a more forward direction, by comparison.^[202] Neanderthal eyeballs are larger than those of modern humans. One study proposed that this was due to Neanderthals having enhanced visual abilities, at the expense of neocortical and social development.^[203] However, this study was rejected by other researchers who concluded that eyeball size does not offer any evidence for the cognitive abilities of Neanderthal or modern humans.^[204]

The projected Neanderthal nose and

Brain

The Neanderthal braincase averages 1,640 cm³ (100 cu in) for males and1,460 cm³ (89 cu in) for females,^[72][73] which is significantly larger than the averages for all groups of extant humans;^[74] for example, modern European males average 1,362 cm³ (83.1 cu in) and females 1,201 cm³ (73.3 cu in).^[210] For 28 modern human specimens from 190,000 to 25,000 years ago, the average was about 1,478 cm³ (90.2 cu in) disregarding sex, and modern human brain size is suggested to have decreased since the Upper Palaeolithic.^[211] The largest Neanderthal brain, Amud 1, was calculated to be 1,736 cm³ (105.9 cu in), one of the largest ever recorded in hominids.^[73] Both Neanderthal and human infants measure about 400 cm³ (24 cu in).^[212]

When viewed from the rear, the Neanderthal braincase has lower, wider, rounder appearance than in anatomically modern humans. This characteristic shape is referred to as "en bombe" (bomb-like), and is unique to Neanderthals, with all other hominid species (including most modern humans) generally having narrow and relatively upright cranial vaults, when viewed from behind.

Maximum natural lifespan and the timing of adulthood, menopause and gestation were most likely very similar to modern humans.^[176] However, it has been hypothesised, based on the growth rates of teeth and tooth enamel,^[228][229] that Neanderthals matured faster than modern humans, although this is not backed up by age biomarkers.^[86] The main differences in maturation are the atlas bone in the neck as well as the middle thoracic vertebrae fused about 2 years later in Neanderthals than in modern humans, but this was more likely caused by a difference in anatomy rather than growth rate.^[230][231]

Generally, models on Neanderthal caloric requirements report significantly higher intakes than those of modern humans because they typically assume Neanderthals had higher basal metabolic rates (BMRs) due to higher muscle mass, faster growth rate and greater body heat production against the cold;^{[232][233][234]} and higher daily physical activity levels (PALs) due to greater daily travelling distances while foraging.^[233][234] However, using a high BMR and PAL, American archaeologist Bryan Hockett estimated that a pregnant Neanderthal would have consumed 5,500 calories per day, which would have necessitated a heavy reliance on big game meat; such a diet would have caused numerous deficiencies or nutrient poisonings, so he concluded that these are poorly warranted assumptions to make.^[234]

Neanderthals may have been more active during dimmer light conditions rather than broad daylight because they lived in regions with reduced daytime hours in the winter, hunted large game (such predators typically hunt at night to enhance ambush tactics), and had large eyes and visual processing neural centres. Genetically,

Low population caused a low

Neanderthals were subject to several infectious diseases and parasites. Modern humans likely transmitted diseases to them; one possible candidate is the stomach bacteria Helicobacter pylori.^[246] The modern human papillomavirus variant 16A may descend from Neanderthal introgression.^[247] A Neanderthal at Cueva del Sidrón, Spain, shows evidence of a gastrointestinal Enterocytozoon bieneusi infection.^[248] The leg bones of the French La Ferrassie 1 feature lesions that are consistent with periostitis—inflammation of the tissue enveloping the bone—likely a result of hypertrophic osteoarthropathy, which is primarily caused by a chest infection or lung cancer.^[249] Neanderthals had a lower cavity rate than modern humans, despite some populations consuming typically cavity-causing foods in great quantity, which could indicate a lack of cavity-causing oral bacteria, namely Streptococcus mutans.^[250]

Two 250,000-year-old Neanderthaloid children from

Social structure

Group dynamics

Neanderthals likely lived in more sparsely distributed groups than contemporary modern humans,^[176] but group size is thought to have averaged 10 to 30 individuals, similar to modern hunter-gatherers.^[31] Reliable evidence of Neanderthal group composition comes from Cueva del Sidrón, Spain, and the footprints at Le Rozel, France:^[188] the former shows 7 adults, 3 adolescents, 2 juveniles and an infant;^[252] whereas the latter, based on footprint size, shows a group of 10 to 13 members where juveniles and adolescents made up 90%.^[188]

A Neanderthal child's teeth analysed in 2018 showed it was weaned after 2.5 years, similar to modern hunter gatherers, and was born in the spring, which is consistent with modern humans and other mammals whose birth cycles coincide with environmental cycles.^[251] Indicated from various ailments resulting from high stress at a low age, such as stunted growth, British archaeologist Paul Pettitt hypothesised that children of both sexes were put to work directly after weaning;^[183] and Trinkaus said that, upon reaching adolescence, an individual may have been expected to join in hunting large and dangerous game.^[86] However, the bone trauma is comparable to modern Inuit, which could suggest a similar childhood between Neanderthals and contemporary modern humans.^[253] Further, such stunting may have also resulted from harsh winters and bouts of low food resources.^[251]

Sites showing evidence of no more than three individuals may have represented

In 2022, remains of the first-known Neanderthal family (six adults and five children) were excavated from Chagyrskaya Cave in the Altai Mountains of southern Siberia in Russia. The family, which included a father, a daughter, and what appear to be cousins, most likely died together, presumably from starvation.[261]^[262]

Inter-group relations

Canadian

Considering most Neanderthal artifacts were sourced no more than 5 km (3.1 mi) from the main settlement, Hayden considered it unlikely these bands interacted very often,[31] and mapping of the Neanderthal brain and their small group size and population density could indicate that they had a reduced ability for inter-group interaction and trade.^[203] However, 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. Based on this, Hayden also speculated that macro-bands formed which functioned much like those of the low-density hunter-gatherer societies of the Western Desert of Australia. Macro-bands collectively encompass 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.^[31] Similarly, British anthropologist Eiluned Pearce and Cypriot archaeologist Theodora Moutsiou speculated that Neanderthals were possibly capable of forming geographically expansive ethnolinguistic tribes encompassing upwards of 800 people, based on the transport of obsidian up to 300 km (190 mi) from the source compared to trends seen in obsidian transfer distance and tribe size in modern hunter-gatherers. However, according to their model Neanderthals would not have been as efficient at maintaining long-distance networks as modern humans, probably due to a significantly lower population.^[264] Hayden noted an apparent cemetery of six or seven individuals at La Ferrassie, France, which, in modern humans, is typically used as evidence of a corporate group which maintained a distinct social identity and controlled some resource, trading, manufacturing and so on. La Ferrassie is also located in one of the richest animal-migration routes of Pleistocene Europe.^[31]

Genetic analysis indicates there were at least three distinct geographical groups—Western Europe, the Mediterranean coast, and east of the Caucasus—with some migration among these regions.^[84] Post-Eemian Western European Mousterian lithics can also be broadly grouped into three distinct macro-regions: Acheulean-tradition Mousterian in the southwest, Micoquien in the northeast, and Mousterian with bifacial tools (MBT) in between the former two. MBT may actually represent the interactions and fusion of the two different cultures.^[83] Southern Neanderthals exhibit regional anatomical differences from northern counterparts: a less protrusive jaw, a shorter gap behind the molars, and a vertically higher jawbone.^[265] These all instead suggest Neanderthal communities regularly interacted with neighbouring communities within a region, but not as often beyond.^[83]

Nonetheless, over long periods of time, there is evidence of large-scale cross-continental migration. Early specimens from

There is also evidence of inter-group conflict: a skeleton from La Roche à Pierrot, France, showing a healed fracture on top of the skull apparently caused by a deep blade wound,[268] and another from Shanidar Cave, Iraq, found to have a rib lesion characteristic of projectile weapon injuries.^[269]

Social hierarchy

It is sometimes suggested that, since they were hunters of challenging big game and lived in small groups, there was no sexual division of labour as seen in modern hunter-gatherer societies. That is, men, women and children all had to be involved in hunting, instead of men hunting with women and children foraging. However, with modern hunter-gatherers, the higher the meat dependency, the higher the division of labour.^[31] Further, tooth-wearing patterns in Neanderthal men and women suggest they commonly used their teeth for carrying items, but men exhibit more wearing on the upper teeth, and women the lower, suggesting some cultural differences in tasks.^[270]

It is controversially proposed that some Neanderthals wore decorative clothing or jewellery—such as a leopard skin or

Hunting and gathering

Neanderthals were once thought of as

Edible plant and mushroom remains are recorded from several caves.[49] Neanderthals from Cueva del Sidrón, Spain, based on dental tartar, likely had a meatless diet of mushrooms, pine nuts and moss, indicating they were forest foragers.^[248] Remnants from Amud Cave, Israel, indicates a diet of figs, palm tree fruits and various cereals and edible grasses.^[50] Several bone traumas in the leg joints could possibly suggest habitual squatting, which, if the case, was likely done while gathering food.^[300] Dental tartar from Grotte de Spy, Belgium, indicates the inhabitants had a meat-heavy diet including woolly rhinoceros and mouflon sheep, while also regularly consuming mushrooms.^[248] Neanderthal faecal matter from El Salt, Spain, dated to 50,000 years ago—the oldest human faecal matter remains recorded—show a diet mainly of meat but with a significant component of plants.^[301] Evidence of cooked plant foods—mainly legumes and, to a far lesser extent, acorns—was discovered at the Kebara Cave site in Israel, with its inhabitants possibly gathering plants in spring and fall and hunting in all seasons except fall, although the cave was probably abandoned in late summer to early fall.^[40] At Shanidar Cave, Iraq, 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 behaviours for both meat and plants.^[48]

Food preparation

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.^[44] The abundance of animal bone fragments at settlements may indicate the making of fat stocks from boiling bone marrow, possibly taken from animals that had already died of starvation. These methods would have substantially increased fat consumption, which was a major nutritional requirement of communities with low carbohydrate and high protein intake.^[44][302] Neanderthal tooth size had a decreasing trend after 100,000 years ago, which could indicate an increased dependence on cooking or the advent of boiling, a technique that would have softened food.^[303]

At Cueva del Sidrón, Spain, Neanderthals likely cooked and possibly

At Grotte du Lazaret, France, a total of twenty-three red deer, six ibexes, three aurochs, and one roe deer appear to have been hunted in a single autumn hunting season, when strong male and female deer herds would group together for rut. The entire carcasses seem to have been transported to the cave and then butchered. Because this is such a large amount of food to consume before spoilage, it is possible these Neanderthals were curing and preserving it before winter set in. At 160,000 years old, it is the oldest potential evidence of food storage.^[43] The great quantities of meat and fat which could have been gathered in general from typical prey items (namely mammoths) could also indicate food storage capability.^[304] With shellfish, Neanderthals needed to eat, cook, or in some manner preserve them soon after collection, as shellfish spoils very quickly. At Cueva de los Aviones, Spain, the remains of edible, algae eating shellfish associated with the alga Jania rubens could indicate that, like some modern hunter gatherer societies, harvested shellfish were held in water-soaked algae to keep them alive and fresh until consumption.^[305]

Competition

Competition from large Ice Age predators was rather high. Cave lions likely targeted horses, large deer and wild cattle; and leopards primarily reindeer and roe deer; which heavily overlapped with Neanderthal diet. To defend a kill against such ferocious predators, Neanderthals may have engaged in a group display of yelling, arm waving, or stone throwing; or quickly gathered meat and abandoned the kill. However, at Grotte de Spy, Belgium, 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.^[52]

Neanderthals and

There are several instances of Neanderthals practising

Personal adornment

Neanderthals used ochre, a

Neanderthals collected uniquely shaped objects and are suggested to have modified them into pendants, such as a fossil

Gibraltarian palaeoanthropologists

As of 2014, 63 purported engravings have been reported from 27 different European and Middle Eastern Lower-to-Middle Palaeolithic sites, of which 20 are on flint cortexes from 11 sites, 7 are on slabs from 7 sites, and 36 are on pebbles from 13 sites. It is debated whether or not these were made with symbolic intent.

Despite the apparent 150,000-year stagnation in Neanderthal lithic innovation,[176] there is evidence that Neanderthal technology was more sophisticated than was previously thought.^[64] However, the high frequency of potentially debilitating injuries could have prevented very complex technologies from emerging, as a major injury would have impeded an expert's ability to effectively teach a novice.^[236]

Stone tools

Mousterian projectile point

Levallois technique

Neanderthals made stone tools, and are associated with the Mousterian industry.^[27] The Mousterian is also associated with North African H. sapiens as early as 315,000 years ago^[341] and was found in Northern China about 47–37,000 years ago in caves such as Jinsitai or Tongtiandong.^[342] It evolved around 300,000 years ago with the Levallois technique which developed directly from the preceding Acheulean industry (invented by H. erectus about 1.8 mya). Levallois made it easier to control flake shape and size, and as a difficult-to-learn and unintuitive process, the Levallois technique may have been directly taught generation to generation rather than via purely observational learning.^[28]

There are distinct regional variants of the Mousterian industry, such as: the

There is some debate if Neanderthals had long-ranged weapons.[345]^[346] A wound on the neck of an African wild ass from Umm el Tlel, Syria, was likely inflicted by a heavy Levallois-point javelin,^[347] and bone trauma consistent with habitual throwing has been reported in Neanderthals.^[345][346] Some spear tips from Abri du Maras, France, may have been too fragile to have been used as thrusting spears, possibly suggesting their use as darts.^[298]

Organic tools

The Châtelperronian in central France and northern Spain is a distinct industry from the Mousterian, and is controversially hypothesised to represent a culture of Neanderthals borrowing (or by process of acculturation) tool-making techniques from immigrating modern humans, crafting bone tools and ornaments. In this frame, the makers would have been a transitional culture between the Neanderthal Mousterian and the modern human Aurignacian.^{[348][349][350][351][352]} The opposing viewpoint is that the Châtelperronian was manufactured by modern humans instead.^[353] Abrupt transitions similar to the Mousterian/Châtelperronian could also simply represent natural innovation, like the La Quina–Neronian transition 50,000 years ago featuring technologies generally associated with modern humans such as bladelets and microliths. Other ambiguous transitional cultures include the Italian Uluzzian industry,^[354] and the Balkan Szeletian industry.^[355]

Before immigration, the only evidence of Neanderthal bone tools are animal rib lissoirs—which are rubbed against hide to make it more supple or waterproof—although this could also be evidence for modern humans immigrating earlier than expected. In 2013, two 51,400- to 41,100-year-old deer rib lissoirs were reported from Pech-de-l'Azé and the nearby Abri Peyrony in France.^{[350][100][100]} In 2020, five more lissoirs made of aurochs or bison ribs were reported from Abri Peyrony, with one dating to about 51,400 years ago and the other four to 47,700–41,100 years ago. This indicates the technology was in use in this region for a long time. Since reindeer remains were the most abundant, the use of less abundant bovine ribs may indicate a specific preference for bovine ribs. Potential lissoirs have also been reported from Grosse Grotte, Germany (made of mammoth), and Grottes des Canalettes, France (red deer).^[356]

The Neanderthals in 10 coastal sites in Italy (namely

At Abri du Maras, France, twisted fibres and a 3-ply inner-bark-fibre cord fragment associated with Neanderthals show that they produced string and cordage, but it is unclear how widespread this technology was because the materials used to make them (such as animal hair, hide, sinew, or plant fibres) are biodegradable and preserve very poorly. This technology could indicate at least a basic knowledge of weaving and knotting, which would have made possible the production of nets, containers, packaging, baskets, carrying devices, ties, straps, harnesses, clothes, shoes, beds, bedding, mats, flooring, roofing, walls and snares, and would have been important in hafting, fishing and seafaring. Dating to 52–41,000 years ago, the cord fragment is the oldest direct evidence of fibre technology, although 115,000-year-old perforated shell beads from Cueva Antón possibly strung together to make a necklace are the oldest indirect evidence.^[36][298] In 2020, British archaeologist Rebecca Wragg Sykes expressed cautious support for the genuineness of the find, but pointed out that the string would have been so weak that it would have had limited functions. One possibility is as a thread for attaching or stringing small objects.^[357]

The archaeological record shows that Neanderthals commonly used animal hide and birch bark, and may have used them to make cooking containers, although this is based largely on

Many Mousterian sites have evidence of fire, some for extended periods of time, though it is unclear whether they were capable of starting fire or simply scavenged from naturally occurring wildfires. Indirect evidence of fire-starting ability includes pyrite residue on a couple of dozen bifaces from late Mousterian (c. 50,000 years ago) northwestern France (which could indicate they were used as percussion fire starters), and collection of manganese dioxide by late Neanderthals which can lower the combustion temperature of wood.^{[29][30][360]} They were also capable of zoning areas for specific activities, such as for knapping, butchering, hearths and wood storage. Many Neanderthal sites lack evidence for such activity perhaps due to natural degradation of the area over tens of thousands of years, such as by bear infiltration after abandonment of the settlement.^[271]

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, Spain, 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.^[31][32] At Cueva de Bolomor, Spain, with hearths lined up against the wall, the smoke flowed upwards to the ceiling, and led to outside the cave. In Grotte du Lazaret, France, smoke was probably naturally ventilated during the winter as the interior cave temperature was greater than the outside temperature; likewise, the cave was likely only inhabited in the winter.^[32]

In 1990, two 176,000-year-old ring structures, several metres wide, made of broken stalagmite pieces, were discovered in a large chamber more than 300 m (980 ft) from the entrance within Grotte de Bruniquel, France. One ring was 6.7 m × 4.5 m (22 ft × 15 ft) with stalagmite pieces averaging 34.4 cm (13.5 in) in length, and the other 2.2 m × 2.1 m (7.2 ft × 6.9 ft) with pieces averaging 29.5 cm (11.6 in). There were also four other piles of stalagmite pieces for a total of 112 m (367 ft) or 2.2 t (2.4 short tons) worth of stalagmite pieces. Evidence of the use of fire and burnt bones also suggest human activity. A team of Neanderthals was likely necessary to construct the structure, but the chamber's actual purpose is uncertain. Building complex structures so deep in a cave is unprecedented in the archaeological record, and indicates sophisticated lighting and construction technology, and great familiarity with subterranean environments.^[361]

The 44,000-year-old Moldova I open-air site, Ukraine, shows evidence of a 7 m × 10 m (23 ft × 33 ft) ring-shaped dwelling made out of mammoth bones meant for long-term habitation by several Neanderthals, which would have taken a long time to build. It appears to have contained hearths, cooking areas and a flint workshop, and there are traces of woodworking. Upper Palaeolithic modern humans in the Russian plains are thought to have also made housing structures out of mammoth bones.^[85]

Birch tar

Neanderthal produced the adhesive

Neanderthals were likely able to survive in a similar range of temperatures to modern humans while sleeping: about 32 °C (90 °F) while naked in the open and windspeed 5.4 km/h (3.4 mph), or 27–28 °C (81–82 °F) while naked in an enclosed space. Since ambient temperatures were markedly lower than this—averaging, during the Eemian interglacial, 17.4 °C (63.3 °F) in July and 1 °C (34 °F) in January and dropping to as a low as −30 °C (−22 °F) on the coldest days—Danish physicist Bent Sørensen hypothesised that Neanderthals required tailored clothing capable of preventing airflow to the skin. Especially during extended periods of travelling (such as a hunting trip), tailored footwear completely enwrapping the feet may have been necessary.^[364]

Since human/Neanderthal admixture is known to have occurred in the Middle East, and no modern body louse species descends from their Neanderthal counterparts (body lice only inhabit clothed individuals), it is possible Neanderthals (and/or humans) in hotter climates did not wear clothes, or Neanderthal lice were highly specialised.^[366]

Seafaring

Remains of Middle Palaeolithic stone tools on Greek islands indicate early seafaring by Neanderthals in the

Given their dangerous hunting and extensive skeletal evidence of healing, Neanderthals appear to have lived lives of frequent traumatic injury and recovery. Well-healed fractures on many bones indicate the setting of splints. Individuals with severe head and rib traumas (which would have caused massive blood loss) indicate they had some manner of dressing major wounds, such as bandages made from animal skin. By and large, they appear to have avoided severe infections, indicating good long-term treatment of such wounds.^[42]

Their knowledge of

The degree of language complexity is difficult to establish, but given that Neanderthals achieved some technical and cultural complexity, and interbred with humans, it is reasonable to assume they were at least fairly articulate, comparable to modern humans. A somewhat complex language—possibly using syntax—was likely necessary to survive in their harsh environment, with Neanderthals needing to communicate about topics such as locations, hunting and gathering, and tool-making techniques.^{[64][368][369]} The FOXP2 gene in modern humans is associated with speech and language development. FOXP2 was present in Neanderthals,^[370] but not the gene's modern human variant.^[371] Neurologically, Neanderthals had an expanded Broca's area—operating the formulation of sentences, and speech comprehension, but out of a group of 48 genes believed to affect the neural substrate of language, 11 had different methylation patterns between Neanderthals and modern humans. This could indicate a stronger ability in modern humans than in Neanderthals to express language.^[372]

In 1971, cognitive scientist

It is debated if behavioural modernity is a recent and uniquely modern human innovation, or if Neanderthals also possessed it.^{[55][369][379][380]}

Religion

Funerals

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

Although Neanderthals did bury their dead, at least occasionally—which may explain the abundance of fossil remains^[53]—the behavior is not indicative of a religious belief of life after death because it could also have had non-symbolic motivations, such as great emotion^[381] or the prevention of scavenging.^[382]

Estimates made regarding the number of known Neanderthal burials range from thirty-six to sixty.^{[383][384][385][386]} The oldest confirmed burials do not seem to occur before approximately 70,000 years ago.^[387] The small number of recorded Neanderthal burials implies that the activity was not particularly common. The setting of inhumation in Neanderthal culture largely consisted of simple, shallow graves and pits.^[388] Sites such as La Ferrassie in France or Shanidar in Iraq may imply the existence of mortuary centers or cemeteries in Neanderthal culture due to the number of individuals found buried at them.^[388]

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

The graves of children and infants, especially, are associated with grave goods such as artefacts and bones. The grave of a newborn from La Ferrassie, France, was found with three flint scrapers, and an infant from Dederiyeh [de] Cave, Syria, was found with a triangular flint placed on its chest. A 10-month-old from Amud Cave, Israel, was associated with a red deer mandible, likely purposefully placed there given other animal remains are now reduced to fragments. Teshik-Tash 1 from Uzbekistan was associated with a circle of ibex horns, and a limestone slab argued to have supported the head.^[253] A child from Kiik-Koba, Crimea, Ukraine, had a flint flake with some purposeful engraving on it, likely requiring a great deal of skill.^[58] Nonetheless, these contentiously constitute evidence of symbolic meaning as the grave goods' significance and worth are unclear.^[253]

Cults

It was once argued that the bones of the cave bear, particularly the skull, in some European caves were arranged in a specific order, indicating an ancient

It was also once thought that Neanderthals ritually hunted, killed and cannibalised other Neanderthals and used the skull as the focus of some ceremony.[309] 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,^[396] putting it forward as evidence of a skull cult.^[381] However, it is now thought to have been a result of cave hyaena scavengery.^[397] Although Neanderthals are known to have practiced cannibalism, there is unsubstantial evidence to suggest ritual defleshing.^[308]

In 2019, Gibraltarian palaeoanthropologists Stewart, Geraldine and Clive Finlayson and Spanish archaeologist Francisco Guzmán speculated that the golden eagle had iconic value to Neanderthals, as exemplified in some modern human societies because they reported that golden eagle bones had a conspicuously high rate of evidence of modification compared to the bones of other birds. They then proposed some "Cult of the Sun Bird" where the golden eagle was a symbol of power.^[55][322] There is evidence from Krapina, Croatia, from wear use and even remnants of string, that suggests that raptor talons were worn as personal ornaments.^[398]

Interbreeding

Interbreeding with modern humans

The first Neanderthal genome sequence was published in 2010, and strongly indicated interbreeding between Neanderthals and early modern humans.^{[87][400][401][402]} The genomes of all studied modern populations contain Neanderthal DNA.^{[87][89][403][404][405]} Various estimates exist for the proportion, such as 1–4%^[87] or 3.4–7.9% in modern Eurasians,^[406] or 1.8–2.4% in modern Europeans and 2.3–2.6% in modern East Asians.^[407] Pre-agricultural Europeans appear to have had similar, or slightly higher,^[405] percentages to modern East Asians, and the numbers may have decreased in the former due to dilution with a group of people which had split off before Neanderthal introgression.^[100] Typically, studies have reported finding no significant levels of Neanderthal DNA in Sub-Saharan Africans, but a 2020 study detected 0.3-0.5% in the genomes of five African sample populations, likely the result of Eurasians back-migrating and interbreeding with Africans, as well as human-to-neanderthal gene flow from dispersals of Homo sapiens preceding the larger Out-of-Africa migration, and also showed more equal Neanderthal DNA percentages for European and Asian populations.^[405] Such low percentages of Neanderthal DNA in all present day populations indicate infrequent past interbreeding,^[408] unless interbreeding was more common with a different population of modern humans which did not contribute to the present day gene pool.^[100] Of the inherited Neanderthal genome, 25% in modern Europeans and 32% in modern East Asians may be related to viral immunity.^[409] In all, approximately 20% of the Neanderthal genome appears to have survived in the modern human gene pool.^[94]

Neanderthal mtDNA (which is passed on from mother to child) is absent in modern humans.[136]^[155][419] This is evidence that interbreeding occurred mainly between Neanderthal males and modern human females.^[420] According to Svante Pääbo, it is not clear that modern humans were socially dominant over Neanderthals, which may explain why the interbreeding occurred primarily between Neanderthal males and modern human females.^[421] Furthermore, even if Neanderthal women and modern human males had interbred, Neanderthal mtDNA lineages may have gone extinct if women who carried them only gave birth to sons.^[421]

Due to the lack of Neanderthal-derived Y-chromosomes in modern humans (which is passed on from father to son), it has also been suggested that the hybrids that contributed ancestry to modern populations were predominantly females, or the Neanderthal Y-chromosome was not compatible with H. sapiens and became extinct.^[100][422]

According to

Detractors of the interbreeding model argue that the genetic similarity is only a remnant of a common ancestor instead of interbreeding,[427] although this is unlikely as it fails to explain why sub-Saharan Africans do not have Neanderthal DNA.^[401]

In December 2023, scientists reported that

Although nDNA confirms that Neanderthals and Denisovans are more closely related to each other than they are to modern humans, Neanderthals and modern humans share a more recent maternally-transmitted mtDNA common ancestor, possibly due to interbreeding between Denisovans and some unknown human species. The 400,000-year-old Neanderthal-like humans from Sima de los Huesos in northern Spain, looking at mtDNA, are more closely related to Denisovans than Neanderthals. Several Neanderthal-like fossils in Eurasia from a similar time period are often grouped into H. heidelbergensis, of which some may be relict populations of earlier humans, which could have interbred with Denisovans.^[430] 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 270,000 years ago, while its genomic DNA indicated divergence less than 150,000 years ago.^[90]

Sequencing of the genome of a Denisovan from Denisova Cave has shown that 17% of its genome derives from Neanderthals.^[93] 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, Croatia, or Mezmaiskaya Cave in the Caucasus, suggesting that interbreeding was local.^[92]

For the 90,000-year-old

Transition

The extinction of Neanderthals was part of the broader

Some Neanderthals in Gibraltar were dated to much later than this—such as Zafarraya (30,000 years ago)^[441] and Gorham's Cave (28,000 years ago)^[442]—which may be inaccurate as they were based on ambiguous artefacts instead of direct dating.^[11] A claim of Neanderthals surviving in a polar refuge in the Ural Mountains^[169] is loosely supported by Mousterian stone tools dating to 34,000 years ago from the northern Siberian Byzovaya site at a time when modern humans may not yet have colonised the northern reaches of Europe;^[171] however, modern human remains are known from the nearby Mamontovaya Kurya site dating to 40,000 years ago.^[443] Indirect dating of Neanderthals remains from Mezmaiskaya Cave reported a date of about 30,000 years ago, but direct dating instead yielded 39,700 ±1,100 years ago, more in line with trends exhibited in the rest of Europe.^[10]

The earliest indication of Upper Palaeolithic modern human immigration into Europe is the Balkan Bohunician industry beginning 48,000 years ago, likely deriving from the Levantine Emiran industry,^[355] and the earliest bones in Europe date to roughly 45–43,000 years ago in Bulgaria,^[444] Italy,^[445] and Britain.^[446] This wave of modern humans replaced Neanderthals.^[8] However, Neanderthals and H. sapiens have a much longer contact history. DNA evidence indicates H. sapiens contact with Neanderthals and admixture as early as 120–100,000 years ago. A 2019 reanalysis of 210,000-year-old skull fragments from the Greek Apidima Cave assumed to have belonged to a Neanderthal concluded that they belonged to a modern human, and a Neanderthal skull dating to 170,000 years ago from the cave indicates H. sapiens were replaced by Neanderthals until returning about 40,000 years ago.^[447] This identification was refuted by a 2020 study.^[448] Archaeological evidence suggests that Neanderthals displaced modern humans in the Near East around 100,000 years ago until about 60–50,000 years ago.^[100]

Cause

Modern humans

Historically, modern human technology was viewed as vastly superior to that of Neanderthals, with more efficient weaponry and subsistence strategies, and Neanderthals simply went extinct because they could not compete.^[19]

The discovery of Neanderthal/modern human introgression has caused the resurgence of the multiregional hypothesis, wherein the present day genetic makeup of all humans is the result of complex genetic contact among several different populations of humans dispersed across the world. By this model, Neanderthals and other recent archaic humans were simply assimilated into the modern human genome – that is, they were effectively bred out into extinction.^[19] Modern humans coexisted with Neanderthals in Europe for around 3,000 to 5,000 years.^[449]

Climate change

Their ultimate extinction coincides with Heinrich event 4, a period of intense seasonality; later Heinrich events are also associated with massive cultural turnovers when European human populations collapsed.^[20][21] This climate change may have depopulated several regions of Neanderthals, like previous cold spikes, but these areas were instead repopulated by immigrating humans, leading to Neanderthal extinction.^[450] In southern Iberia, there is evidence that Neanderthal populations declined during H4 and the associated proliferation of Artemisia-dominated desert-steppes.^[451]

It has also been proposed that climate change was the primary driver, as 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.

Modern humans may have introduced African diseases to Neanderthals, contributing to their extinction. A lack of immunity, compounded by an already low population, was potentially devastating to the Neanderthal population, and low genetic diversity could have also rendered fewer Neanderthals naturally immune to these new diseases ("differential pathogen resistance" hypothesis). However, compared to modern humans, Neanderthals had a similar or higher genetic diversity for 12 major histocompatibility complex (MHC) genes associated with the adaptive immune system, casting doubt on this model.^[24]

Low population and inbreeding depression may have caused maladaptive birth defects, which could have contributed to their decline (mutational meltdown).^[243]

In late-20th-century New Guinea, due to cannibalistic funerary practices, the

Neanderthals have been portrayed in popular culture including appearances in literature, visual media and comedy. The "caveman" archetype often mocks Neanderthals and depicts them as primitive, hunchbacked, knuckle-dragging, club-wielding, grunting, nonsocial characters driven solely by animal instinct. "Neanderthal" can also be used as an insult.^[25]

In literature, they are sometimes depicted as brutish or monstrous, such as in

• Denisovan – Asian archaic human
• Early human migrations
• Early European modern humans
   – Earliest anatomically modern humans in EuropePages displaying short descriptions of redirect targets
• Homo floresiensis – Archaic human from Flores, Indonesia
• Homo luzonensis – Archaic human from Luzon, Philippines
• Homo naledi – South African archaic human species
• Human timeline

Footnotes

References