Neandertals

Neanderthals ( ; ''Homo neanderthalensis'' or sometimes ''H. sapiens neanderthalensis'') are an extinct group of archaic humans who inhabited Europe and Western and Central Asia during the Middle to Late Pleistocene. Neanderthal extinction occurred roughly 40,000 years ago with the immigration of modern humans (Cro-Magnons), but Neanderthals in Gibraltar may have persisted for thousands of years longer.

The first recognised Neanderthal fossil, Neanderthal 1, was discovered in 1856 in the Neander Valley, Germany. At first, Neanderthal 1 was considered to be one of the lower races in accord with historical race concepts. As more fossils were discovered through the early 20th century, Neanderthals became characterised most especially by Marcellin Boule as a unique species of underdeveloped human. By the mid-20th century, human evolution was described as progressing from an apelike ancestor, through a "Neanderthal phase", ending in modern humans. This gave way to the "Out of Africa" theory in the 1970s. With the sequencing of Neanderthal genetics first in 2010, it was discovered that Neanderthals interbred with modern humans.

Neanderthal anatomy is characterised by a long and low skull, a heavy and rounded brow ridge (supraorbital torus), an occipital bun at the back of the skull, strong teeth and jaws, a wide chest, and short limbs. These traits gradually became more frequent through the Middle Pleistocene of Europe, possibly due to natural selection in a cold climate, as well as genetic drift when populations collapsed during glacial periods. Neanderthals would have also been effective sprinters. Neanderthal specimens vary in height from , with average male dimensions of maybe and . While Neanderthal brain volume averaged higher than any living population — for males and for females — their brain organisation differed from modern humans in areas related to cognition and language, which could explain the comparative simplicity of Neanderthal behaviour compared to Cro-Magnons in the archaeological record.

Neanderthals maintained a low population and suffered inbreeding depression, which may have impeded their ability to progress technologically. They produced Mousterian stone tools (a Middle Palaeolithic industry), maintained and might have created fire, and possibly wore blankets and ponchos. They predominantly ate whatever was abundant close to home, usually big game as well as plants and mushrooms. Neanderthals were frequently victims of major physical traumas and animal attacks. Examples of Palaeolithic art have been inconclusively attributed to Neanderthals, namely possible ornaments made from bird claws and feathers; collections of unusual objects including crystals and fossils; and engravings. Neanderthals uncommonly buried their dead, but this is not indicative of a religious belief of an afterlife.

Taxonomy

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. 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. The valley itself was named after the late 17th century German theologian and hymn writer Joachim Neander, who often visited the area. His grandfather, a musician, had changed the family name from the original German ''Neumann'' ("new man") to the Graeco-Roman form ''Neander'', following the fashion of the time.

''Neanderthal'' can be pronounced using the (as in ) or the standard English pronunciation of ''th'' with the fricative / θ/ (as ). The latter pronunciation, nevertheless, has no basis in the original German word which is pronounced always with a ''t'' regardless of the historical spelling.

Neanderthal 1, the type specimen, was known as the "Neanderthal cranium" or "Neanderthal skull" in anthropological literature, and the individual reconstructed on the basis of the skull was occasionally called "the Neanderthal man". The binomial name ''Homo neanderthalensis''—extending the name "Neanderthal man" from the individual specimen to the entire species, and formally recognising it as distinct from humans—was first proposed by Irish geologist William King in a paper read to the 33rd British Science Association in 1863. However, in 1864, he recommended that Neanderthals and modern humans be classified in different genera as he compared the Neanderthal braincase to that of a chimpanzee and argued that they were "incapable of moral and theistic] conceptions".

Discovery

A number of Neanderthal fossils had been discovered before their antiquity was fully understood. The first Neanderthal remains— Engis 2 (a skull)—were discovered in 1829 by Dutch/Belgian prehistorian Philippe-Charles Schmerling in the Grottes d'Engis, Belgium. He concluded that these "poorly developed" human remains must have been buried at the same time and by the same causes as the co-existing remains of extinct animal species. In 1848, Gibraltar 1 from Forbes' Quarry was presented to the Gibraltar Scientific Society by their Secretary Lieutenant Edmund Henry Réné Flint, but was thought to be a modern human skull.

In 1856, local schoolteacher Johann Carl Fuhlrott recognised bones from Kleine Feldhofer Grotte in Neander Valley—Neanderthal 1—as distinct from modern humans, and gave them to German anthropologist Hermann Schaaffhausen to study in 1857. It comprised the cranium, thigh bones, right arm, left humerus and ulna, left ilium (hip bone), part of the right shoulder blade, and pieces of the ribs.

Research history

Following Charles Darwin's 1859 ''On the Origin of Species'', Fuhlrott and Schaaffhausen argued that Neanderthal 1 represents a primitive lower human form, aligning more closely with non-human apes as well as Negroids, Eskimos, and Aboriginal Australians (which were variably classified as separate species or subspecies of human at the time). The uniqueness of Neanderthal Man met opposition namely from the pathologist Rudolf Virchow, who argued against defining new species based on only a single find. In 1872, Virchow erroneously interpreted Neanderthal characteristics as evidence of senility, disease, and malformation instead of archaicness, which stalled Neanderthal research until the end of the century.

By the early 20th century, numerous other Neanderthal discoveries were made, establishing ''H. neanderthalensis'' as a legitimate species. At first, many palaeontologists considered Neanderthals to be an intermediary phase between modern humans and more apelike ancestors, as suggested by German anatomist Gustav Albert Schwalbe. This hypothesis was notably opposed by French palaeontologist Marcellin Boule, who authored several publications starting in 1908 describing the French Neanderthal specimen La Chapelle-aux-Saints 1 ("The Old Man") as a slouching, ape-like creature distantly related to modern man. Boule's ideas would define discussions of Neanderthals for some time.

Boule suggested two different lineages existed in Ice Age Europe: a more evolved one descending from the British Piltdown Man (a hoax) to the French Grimaldi Man (a Cro-Magnon) which would culminate with modern Europeans; and a less evolved dead-end lineage leading from the German Heidelberg Man to Neanderthal Man. As the focus of human origins shifted from Europe to East Asia ("Out of Asia" hypothesis) by the 1930s and 40s with discoveries such as Java Man and Peking Man (as well as the marginalisation of Piltdown Man), the question of a "Neanderthal phase" in human evolution once again became a topic of discussion. Several specimens around the Old World were classified as "progressive" Neanderthals which would eventually evolve into some local subspecies of ''H. sapiens'' ( polycentricism), or in Europe into either the modern European subspecies or the "classic" Neanderthals.

In the 1970s, with the formulation of cladistics and the consequent refinement of the anatomical definitions of species, this "global morphological pattern" fell apart. The "Neanderthaloids" of Africa and East Asia were reclassified as distant relatives to ''H. neanderthalensis''. At around the same time, the "Out of Asia" hypothesis was overturned by the "Out of Africa" hypothesis, which posited that all modern humans share a fully modern common ancestor (monogenism). There were two main schools of thought: modern humans competitively replaced all other archaic humans ("Replacement"), or extensively interbred with them while dispersing throughout the world ("Regional Continuity"). In 2010, the first mapping of the Neanderthal genome demonstrated that there was at least some interbreeding between archaic and modern humans. Subsequent genetic studies continue to raise questions on how Neanderthals should be classified relative to modern humans.

Classification

Neanderthals can be classified as a unique species as ''H. neanderthalensis'', though some authors argue expanding the definition of ''H. sapiens'' to include other ancient humans, with combinations such as ''H. sapiens neanderthalensis'' ( splitters and lumpers). The latter opinion has generally been justified using Neanderthal genetics, as well as inferences on the complexity of Neanderthal behaviour based on the archaeological record. While there seems to have been some genetic contact between these two groups, there are potential indicators of hybrid incompatibility, which if true could justify species distinction. The crux of the issue lies in the vagueness of the term "species" (the species problem).

Among identified archaic humans, Neanderthals are most closely related to Denisovans based on nuclear DNA (nDNA) analyses. Denisovans are an enigmatic group of Late Pleistocene humans only recognisable by a genetic signature. Likely due to more recent interbreeding episodes, the mitochondrial DNA (mtDNA, passed down maternally) and Y-chromosome DNA (passed down paternally) are more similar between Neanderthals and modern humans than between Neanderthals and Denisovans. Similarly, 430,000 year old fossils from the Sima de los Huesos are more closely related to Neanderthals using nDNA, but their mtDNA aligns more closely with Denisovans.

A 2021 phylogeny of some Middle Pleistocene and Neanderthal fossils using tip dating:

Evolution

Typical Neanderthal skull traits appear in the European fossil record near the beginning of the Middle Pleistocene, in specimens usually classified as '' H. heidelbergensis''. These "pre-Neanderthals" seem to have gradually accreted these traits ("Neanderthalization") as populations adapted to the cold environment, evolving a "hyper-arctic" physique. Circumpolar peoples (namely Inuit groups) are often used as modern Neanderthal analogues to study "hyper-arctic" adaptations. Additionally, glacial periods may have forced populations into small refugia, reducing genetic diversity, leading to the development of other typical Neanderthal traits through genetic drift or pleiotropy. The 120,000 to 140,000-year-old Israeli Nesher Ramla remains may represent one such source population which would recolonise Europe following the Penultimate Glacial Period.

The occurrence of typical Neanderthal traits in the Middle Pleistocene was highly variable even among individuals of the same population. The speed of Neanderthalization may have also been impeded by gene flow between Western Europe and Africa, exemplified by anomalous specimens which lack typical Neanderthal traits, such as Ceprano Man. The first recognisable "early Neanderthals" show up in the fossil record by the end of Marine Isotope Stage 7 (beginning roughly 243,000 years ago) and give way to "classic" or "late Neanderthals" by the end of Marine Isotope Stage 5e. This spans the Penultimate Glacial Period to the Last Interglacial. Some early Neanderthal teeth from Payré, France, potentially date to MIS 8, but the dating is uncertain.

Genetic data usually estimates that Neanderthals diverged from modern humans sometime during the early Middle Pleistocene. Neanderthals and Denisovans are more closely related to each other than they are to modern humans, meaning the Neanderthal/Denisovan split occurred sometime later. 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 million years ago.

Genetic data indicates that Neanderthals, at least after 100,000 years ago, maintained a small population with low genetic diversity, weakening natural selection and proliferating harmful mutations. It is unclear how long European populations suffered this population stress, or to what extent it influenced Neanderthalization.

Demographics

Range

The Neanderthals were the first human species to permanently occupy Europe. While pre-Neanderthals are mostly identified around Western Europe, classic Neanderthals are recorded across Europe as well as Southwest and Central Asia, up to the Altai Mountains in southern Siberia. Pre- and early Neanderthals 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 (without leaving Europe). Nonetheless, southwestern France has the highest density of sites for pre- and classic Neanderthals.

The southernmost find was recorded at Shuqba Cave, Levant; reports of Neanderthals from the North African Jebel Irhoud and Haua Fteah have been reidentified as ''H. sapiens''. Their easternmost presence is recorded at Denisova Cave, Siberia 85°E; the southeast Chinese Maba Man, a skull, shares several physical attributes with Neanderthals, although these may be the result of convergent evolution rather than Neanderthals extending their range to the Pacific Ocean. The northernmost bound is generally accepted to have been 55°N, with unambiguous sites known between 50– 53°N, but this is difficult to assess because glacial advances destroy most human remains. Middle Palaeolithic artefacts have been found up to 60°N on the Russian plains, but these are more likely attributed to modern humans.

It is possible Neanderthal range expanded and contracted as the ice retreated and grew, respectively, to avoid permafrost areas, residing in certain refuge zones during glacial maxima. Stable environments with mild mean annual temperatures may have been the most suitable Neanderthal habitats.

Population

Like modern humans, Neanderthals probably descended from a very small population with an effective population—the number of individuals who can bear or father children—of 3,000 to 12,000 approximately. Neanderthals maintained this low population, proliferating weakly harmful genes due to the reduced effectivity of natural selection. Archaeological evidence suggests that the initial Cro-Magnon population was 10 times higher than Neanderthals.

Neanderthals may have been at a demographic disadvantage due to a lower fertility rate, a higher infant mortality rate, or a combination of the two. 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. Infant mortality was estimated to have been very high for Neanderthals, about 43% in northern Eurasia.

Anatomy

Skull

The Neanderthal skull has a flat and broad skullcap, rounded supraorbital torus (the brow ridges), high orbits (eye sockets), a broad nose, mid-facial prognathism (the face projects far from the base of the skull), an "en bombe" (bomb-like) skull shape when viewed from the back, and an occipital bun at the back of the skull.

The Neanderthal braincase averages for males and for females, which is significantly larger than the averages for all living populations. The largest Neanderthal brain, Amud 1, was calculated to be , one of the largest ever recorded in humans. Neanderthal brain organisation differs in areas related to cognition and language, which may be implicated in the comparative simplicity of Neanderthal behaviour compared to Cro-Magnons in the archaeological record.

Neanderthals had large and wide noses, probably an adaptation to warm greater quantities of cold air to fuel their assumed heightened metabolism and activity levels. A large nose does not necessarily equate to a better sense of smell, and neurologically, because the olfactory bulbs are smaller, Neanderthals may have had a poorer sense of smell and olfactory memory than modern humans.

The zygomatic bone, cheek bones are strong, the incisors are large and shovel-shaped incisors, shovel-shaped, the molar (tooth), molars have a swollen tooth pulp (taurodontism), and there is a gap behind the molars (retromolar space). These dental traits are usually interpreted as a response to habitual heavy loading of the front teeth, either to process mechanically challenging or dental attrition, attritive foods, or because Neanderthals regularly used the mouth as a third hand.

Build

Neanderthals were generally short and stocky. In a sample of 45 Neanderthal long bones from 14 men and 7 women, the average height was for males and for females. The fossil record shows that adult Neanderthals varied from about in height. Average male body mass index would have been 26.9–28.2 using a size of and .

The Neanderthal chest was deep and wide, with a proportionally expansive thoracic cavity, and possibly stronger lung performance. Neanderthals also had relatively more skeletal muscle#Force generation, fast-twitch muscle fibres, and much higher calorie, caloric demands. The limbs are proportionally short. The body plan has traditionally been explained as a "hyper-arctic" adaptation (Allen's rule). Stronger lungs, more fast-twitch muscle, and shorter limbs would have also boosted sprinting efficiency.

Skin colour seems to have ranged from dark to light. Some Neanderthals had dark or brown hair. If red was another possible hair colour, it does not appear to have been a common one.

Pathology

Neanderthals suffered a high rate of traumatic injury, with an estimated 79–94% of specimens showing evidence of healed major trauma, of which 37–52% were severely injured, and 13–19% injured before reaching adulthood. One extreme example is Shanidar 1, who shows signs of an amputation of the right arm likely due to a nonunion after breaking a bone in adolescence, osteomyelitis (a bone infection) on the left clavicle, an abnormal gait, vision problems in the left eye, and possible hearing loss (perhaps swimmer's ear). The high trauma rate may be ascribed to a dangerous hunting strategy, or frequent animal attacks.

Low population caused a low genetic diversity and probably inbreeding, which reduced the population's ability to filter out harmful mutations (inbreeding depression). It is unknown how this affected a single Neanderthal's genetic burden and, thus, if this caused a higher rate of birth defects than in modern humans.

Culture

Social structure

It is difficult to infer Neanderthal group size, but indirect data generally suggests small bands of 10 to 30 individuals. Bands likely moved between certain caves depending on the season, indicated by remains of seasonal materials, such as certain foods. They returned to the same locations generation after generation, and some sites may have been used for over a century. Neanderthals may have been outcompeting cave bears for cave space. Intergroup movement may have been predominantly female-driven, with at least some groups practicing patrilocal residence, patrilocal residency (the woman moves out of her group to live with her mate).

Neanderthals maintained a low population across their range, which may have hindered their ability to maintain long-distance trade routes and avoid inbreeding. They may have regularly interacted with closely neighbouring communities within a region, but not as often beyond. 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.

While Cro-Magnons are usually assumed to have practised sexual division of labour with men hunting and women gathering like in recent hunter-gatherer societies, it is unclear to what extent this can be applied to Neanderthals. Both Neanderthal men and women have similar traumatic injury patterns, which might imply that both sexes were involved in hunting. Dental wearing patterns, on the other hand, could indicate men and women typically carried different items with their mouths, maybe during tasks not related to subsistence. The women at El Sidrón Cave, Spain, may have been eating more seeds and nuts than the men. The lack of a strict sexual division in Neanderthals has usually been linked to their small population and group size, falling short of the demographic threshold where task specialisation becomes feasible — which may also explain the comparative simplicity of Neanderthal material culture.

Food

Neanderthals were once thought of as scavengers, but are now considered to have been apex predators. They appear to have eaten predominantly what was abundant within their immediate surroundings. Cro-Magnons, in contrast, seem to have used more complex food extraction strategies and generally had a more diverse diet.

In many European sites, prey items include reindeer, wild horse, horse, aurochs, and steppe bison. Neanderthals in Southwest Asia more commonly hunted mountain gazelle, Persian fallow deer, wild goat, and camels. They may have less frequently taken down larger Pleistocene megafauna whenever locally abundant, such as woolly mammoth and woolly rhinoceros. At the 125,000 year old Neumark-Nord site, Germany, there is evidence of regular hunting of straight-tusked elephants maybe every 5 to 6 years. Some waterside communities ate fish and shellfish; and at Vanguard Cave, Gibraltar, dolphin and Mediterranean monk seal. Neanderthals also hunted small game, and some caves show evidence of regular rabbit and tortoise consumption. At Gibraltar sites, there are butchered remains of 143 different bird species, many ground-dwelling such as the common quail, corn crake, woodlark, and crested lark. Neanderthals also consumed a variety of plants and mushrooms across their range — at Kebara Cave, Israel, over 50 species of seeds, nuts, fruits, and cereals.

Neanderthals possibly employed a wide range of food preparation techniques. At Cueva del Sidrón, Spain, Neanderthals may have been roasting and smoking (cooking), smoking meat, and used certain plants—such as yarrow and camomile—for flavouring, although these plants may have instead been used for their medicinal properties. At Gorham's Cave, Gibraltar, Neanderthals may have been roasting pinecones to access pine nuts, and at Gruta da Figueira Brava, Cancer pagurus, brown crabs to soften the shell before cracking them open. 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 (mammalian reproduction), rut. It is possible these Neanderthals were curing (food preservation), curing and storing all this meat before winter set in.

Neanderthals competed with several large carnivores, but also seem to have hunted them down, namely Panthera spelaea, cave lions and wolf, wolves, as well as cave and brown bear both in and out of hibernation. Neanderthals and other predators may have sometimes avoided competition by pursuing different prey, namely with cave hyenas and wolves (niche differentiation). Neanderthals, nonetheless, were frequently victims of animal attacks.

There are multiple instances of Neanderthals practicing cannibalism, but it may have only been done in times of extreme food shortages, as in some List of incidents of cannibalism, cases in recorded human history.

The arts

Neanderthals collected non-functional, uniquely-shaped objects, namely shells, fossils, and gems. It is unclear if these objects were simply picked up for their aesthetic qualities, or if some symbolic significance was applied to them. Some shells may have been painted. Gibraltarian palaeoanthropologists Clive Finlayson, Clive and Geraldine Finlayson suggested that Neanderthals used various bird parts as artistic media, especially black feathers. A 2020 study found evidence of a yarn weight, 3-ply cord fragment made from conifer inner-bark fibres at Abri du Maras, France, which can be used to knit light items, such as strings for hanging beads. 115,000-year-old perforated shell beads from Cueva Antón were possibly strung together to make a necklace.

There are several instances of nondescript engravings and scratches on flints, bones, pebbles, and stone slabs — as of 2014, 63 purported engravings have been reported from 27 different European and Middle Eastern Lower-to-Middle Palaeolithic sites. It is debated if these were made with symbolic intent. Neanderthals may have produced finger flutings on the walls of La Roche-Cotard over 57,000 years ago.

Neanderthals used ochre, a clay earth pigment. While modern humans have used this for decorative or symbolic colouration, it has also been used as medicine, hide tanning agent, food preservative, and insect repellent.

The 43,000-year-old Divje Babe flute (a cave bear femur) from Slovenia has been attributed by some researchers to Neanderthals, though its status as a Paleolithic flute, Palaeolithic flute is heavily disputed. Many researchers consider it to be most likely the product of a carnivorous animal chewing the bone.

Technology

Neanderthals manufactured Middle Palaeolithic stone tools, and are associated with the Mousterian industry, specifically the Levallois technique. After developing this technology from the Acheulean industry, there is a 150,000 year stagnation in Neanderthal stone tool innovation. Stalled technological growth may have followed from their low population, impeding complex ideas from being spread across their range or passed down generationally. Neanderthals normally collected raw materials from a nearby source, no more than . Some communities were also making tools from shells and bone. They may have hafting, hafted tips onto spears using birch bark tar. European populations have also been manufacturing wood spears, namely the 400,000 year old British Clacton Spear; 300,000 year old German Schöningen spears; and 120,000 year old German Lehringen spear, Lehringen Spear, including both likely thrown (Schöningen) and thrusting (Lehringen) types.Milks, A.  (2020)
Yew wood, would you? An exploration of the selection of wood for Pleistocene spears
'' In: Berihuete-Azorin, M., Martin Seijo, M., Lopez-Bulto, O. and Pique, R. (eds.) The Missing Woodland Resources: Archaeobotanical studies of the use of plant raw materials. Advances in Archaeobotany, 6 (6). Barkhuis Publishing, Groningen, pp. 5-22. It has been suggested that Neanderthals likely specifically selected particular wood types (such as European yew in the case of the Clacton and Lehringen spears) for manufacturing spears for their beneficial material properties.

Many Neanderthal 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. They may have been using fire for cooking, keeping warm, and deterring predators. They were also capable of zoning areas for specific activities, such as for knapping, butchering, hearths, and wood storage. At Capellades, Abric Romaní rock shelter, Spain, Neanderthals may have maintained 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.

The only known Neanderthal tools that could have been used to fashion clothes are hide scraper (archaeology), scrapers as no bone sewing-needles and stitching awls have been found as in Cro-Magnon sites. Hide scrapers could have been used to make items similar to blankets or ponchos. There is no direct evidence that Neanderthals could make fitted clothes from animal hide. Unfitted clothes would have limited range of mobility while dressed, and decreased the time Neanderthals could spend unprotected from the elements away from shelters.

Neanderthals appear to have lived lives of frequent traumatic injury and recovery, indicating the setting of splint (medicine), splints and dressing of major wounds. By and large, they appear to have avoided severe infections, indicating long-term treatment. The quality of medical care may have ensured their survival as a species for so long. Their knowledge of medicinal plants#Prehistoric times, medicinal plants was comparable to that of Cro-Magnons.

Stone tools on various Greek islands could indicate early seafaring through the Mediterranean, employing simple reed boats for one-day crossings, but the evidence for such a big claim is limited.

Language

It is unclear if Neanderthals had the capacity for complex language, but some researchers have argued that Neanderthals required complex communications to discuss locations, hunting and gathering, and tool-making techniques in order to survive in their harsh environment. In experiments with modern humans, the Levallois technique can be taught with purely observational learning without spoken instruction.

Anatomically, the Neanderthal hyoid bone (which supports the tongue) is almost identical to that in modern humans, but this does not provide insight of the entire vocal tract. Neanderthals had the FOXP2 gene, which is associated with speech and language development, but not the modern human variant.

Burials and religion

Neanderthals, probably uncommonly, buried their dead. This may explain the abundance of fossil remains. The behaviour is not indicative of a religious belief of afterlife, life after death because it could also have had non-symbolic motivations. The dead were buried in simple, shallow graves and pits, but special care seems to have been given to child graves. The graves of children and infants, especially, are associated with grave goods such as artefacts and bones. Some sites with multiple well-preserved Neanderthal skeletons may represent cemeteries.

One grave in Shanidar Cave, Iraq, was associated with the pollen of several flowers that may have been in bloom at the time of deposition—yarrow, Centaurea, centaury, 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, and that "the association of flowers with Neanderthals adds a whole new dimension to our knowledge of his humanness, indicating that he had 'soul. It is also possible the pollen was deposited by a small rodent after the man's death.

Neanderthals were once thought to have ritually killed and eaten cave bears or other Neanderthals, but the evidence is circumstantial. The Finlayson's speculate that Neanderthals viewed the golden eagle as a symbol of power.

Interbreeding

Hybridisation between Neanderthals and early modern humans had been suggested early on, such as by English anthropologist Thomas Huxley in 1890, Danish ethnographer Hans Peder Steensby in 1907, and Coon in 1962. In the early 2000s, supposed hybrid specimens were discovered: Lagar Velho 1 and Peștera Muierilor, Muierii 1. Similar anatomy could also have been caused by adapting to a similar environment rather than interbreeding.

The first Neanderthal genome sequence was published in 2010, and strongly indicated interbreeding between Neanderthals and early modern humans. Neanderthal-derived genes descend from at least 2 interbreeding episodes outside of Africa: one about 250,000 years ago, and another 40,000 to 54,000 years. Interbreeding also occurred in other populations which are not ancestral to any living person. An individual whose ancestry lies beyond sub-Saharan Africa may carry about 2% of Neanderthal DNA. Sub-Saharan Africans can carry Neanderthal DNA, presumably descending from modern human migration between Eurasia and Africa. In all, approximately 20% of the Neanderthal genome appears to have survived in the modern human gene pool. This Neanderthal DNA may descend primarily from the children of female modern humans and male Neanderthals.

Due to their low population and proliferation of deleterious mutations, many Neanderthal genes were probably selected out of the modern human gene pool (negative selection (natural selection), negative selection). Similarly, a large portion of surviving introgression appears to be non-coding DNA, non-coding ("junk") DNA with few biological functions. Some Neanderthal-derived genes, nonetheless, may have functional implications related to metabolism, brain function, and skeletal and muscular development. Some genes may have helped immigrating modern human populations acclimatise faster, such as genes related to immune response.

Neanderthals in the Siberian Altai Mountains interbred with the local Denisovan population, and it may have been a common occurrence here. About 17% of the genome of one Altai Denisovan specimen derived from Neanderthals.

Extinction

The extinction of Neanderthals was part of the broader Late Pleistocene extinctions, Late Pleistocene megafaunal extinction event. Neanderthals were replaced by modern humans, indicated by the near-complete replacement of Middle Palaeolithic Mousterian stone technology with modern human Upper Palaeolithic Aurignacian stone technology across Europe (the Middle-to-Upper Palaeolithic Transition) from 39,000 to 41,000 years ago. Neanderthals may have persisted in Spain for longer, but the dates of the latest Mousterian and earliest Aurignacian are poorly constrained. In Catalonia and Aragón (northern Spain), the Mousterian may have survived to about 39,000 years ago, and in southern Spain and Gibraltar potentially 32,000 to 35,000 years ago. Similar refuge zones have also been proposed on other temperate European peninsulas, namely Italy, the Balkans, and Crimea.

Historically, the cause of extinction of Neanderthals and other archaic humans was viewed under an imperialistic guise, with the superior invading modern humans prehistoric warfare, exterminating and replacing the inferior species.

In general, the extinction of Neanderthals is ascribed predominantly to competitive exclusion principle, competition with modern humans. The success of modern humans over Neanderthals is usually attributed to a higher birth rate and population, better long-distance mobility, and more complex technologies and subsistence strategies. Some Neanderthal populations may have also been assimilated into modern human populations rather than being ecologically outcompeted. Assimilation had long been hypothesised with supposed hybrid specimens, and was revitalised with the discovery of archaic human DNA in modern humans. Similarly, the Châtelperronian industry of central France and northern Spain may represent a culture of Neanderthals adopting modern human techniques, via acculturation. Other ambiguous transitional cultures include the Italian Uluzzian industry, and the Central European Szeletian industry.

Neanderthal extinction has also been ascribed to their low population as well as the resulting mutational meltdown, making them less adaptable to major environmental changes or new diseases introduced by immigrating modern humans. It is unclear if climatic degradation would have severely impacted Neanderthals given how many glacial periods they persisted through in Europe. If areas were depopulated of Neanderthals as a consequence of climate change (specifically Heinrich event 4) or a natural disaster (the Campanian Ignimbrite eruption), Neanderthals may not have been as fast as modern humans in recolonising. The Laschamp event 39,000 to 42,000 years ago may have increased ultraviolet radiation, disproportionately affecting Neanderthals who lacked protective fitted clothes, and may not have utilised ochre as sunscreen to the extent Cro-Magnons did.

In popular culture

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.

In literature, they are sometimes depicted as brutish or monstrous, such as in H. G. Wells' ''The Grisly Folk'' and Elizabeth Marshall Thomas' ''The Animal Wife'', but sometimes with a civilised but unfamiliar culture, as in William Golding's The Inheritors (Golding novel), The Inheritors, Björn Kurtén's ''Dance of the Tiger'', and Jean M. Auel's ''Clan of the Cave Bear'' and her ''Earth's Children'' series.

See also

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* Early human migrations
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Footnotes

References

Sources

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Further reading

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External links

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Human Timeline (Interactive)
– Smithsonian Institution, Smithsonian, National Museum of Natural History (August 2016).
* : Includes Neanderthal mtDNA sequences

GenBank records for ''H. s. neanderthalensis''
maintained by the National Center for Biotechnology Information (NCBI)
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The Climate Chronicles
explores the impact of Pleistocene climate change on Neanderthals and other hominins.

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Neanderthals,
Fossil taxa described in 1864
Stone Age Asia
Stone Age Europe