Neanderthal

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 th in Neanderthal can be pronounced as (hence ) following the German convention or Anglicized as fricative /θ/ (hence ), as per the standard English pronunciation of th.

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 was first proposed by Irish geologist William King in a paper read to the 33rd British Science Association in 1863. He extended the name "Neanderthal man" from the individual specimen to the entire species and formally recognised it as distinct from modern humans. 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

1. Notharctus

2. Propliopithecus

3. Dryopithecus

4. Java Man

5. Piltdown Man

6. Heidelberg Man

7. Neanderthal Man

8. Cro-Magnon Man

9. Australian Black-fellow (pejorative term for Aboriginal Australians)

10. Hottentot (pejorative term for the Southern African Khoisan)

11. Chinese

12. American Caucasian ]] 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 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. The definition of "Neanderthal" expanded to include several anatomically variable specimens around the Old World. Some specimens were described as "progressive" Neanderthals which would evolve into some local subspecies of H. sapiens (polycentricism), while the "classic" Neanderthals of the Western European Würm glaciation would not.

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. 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 rather than anatomical landmarks. 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 in their 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 occurrence of typical Neanderthal traits in the Middle Pleistocene was highly variable even among individuals of the same population. 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

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.

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.

Compared to Cro-Magnons, 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

Main article: Neanderthal anatomy

Skull

The Neanderthal skull has a flat and broad skullcap, rounded supraorbital torus (the buldge that forms the brow ridges), larger, wide 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, a fossa (depression) on the back of the skull below the level of the inion (suprainiac fossa), and an occipital bun (bony projection) at the back of the skull. Like those of other archaic humans, their jaws lack a true chin.

The Neanderthal braincase averages 1,640 cm3 for males and 1,460 cm3 for females, which is significantly larger than the averages for all living populations. The largest Neanderthal brain, Amud 1, was calculated to be 1,736 cm3, 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 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 cheek bones are strong, the incisors are large and shovel-shaped, the 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 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 164 to 168 cm for males and 152 to 156 cm for females. The fossil record shows that adult Neanderthals varied from about 147.5 to in height. Average male body mass index would have been 26.9–28.3 using a size of 164 to 168 cm and 76 kg.

The Neanderthal chest was deep and wide, with a proportionally expansive thoracic cavity, and possibly stronger lung performance. Neanderthals also had relatively more fast-twitch muscle fibres, and much higher 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

Main article: Neanderthal behavior

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 more than a century. Neanderthals may have been outcompeting cave bears for cave space. Intergroup movement may have been predominantly patrilocal (male relationships as the basis of groups with females from other groups entering for breeding).

Neanderthals maintained a low population across their range, which may have hindered their ability to maintain long-distance trade routes and to avoid inbreeding. They may have regularly interacted with closely neighbouring communities within a region, but not so 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 the Cro-Magnons are usually assumed to have generally practised sexual division of labour with men hunting and women gathering such as in the preponderance of recent hunter-gatherer societies, it is unclear to what extent this may be applied to Neanderthals. Both Neanderthal men and women have similar traumatic injury patterns, which might imply that both genders were involved in hunting. Dental wearing patterns among Neanderthals, on the other hand, could indicate men and women typically carried different items with their mouths, perhaps not related to tasks related to subsistence however. The women at El Sidrón Cave, Spain, may have been eating more seeds and nuts than the men. The lack of distinctive task specialization in Neanderthals has usually been linked to their small population and group size, falling short of the demographic threshold where task specialization 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 maintained a more diverse diet even in settings where certain foods would have been harder to procure; for example, Neanderthals living in forests ate about the same proportion of foodplants as Cro-Magnons, but Neanderthals on open steppe (where foodplants are harder to find) ate far less foodplants.

In many European sites, prey items include reindeer, horse, aurochs, and steppe bison. Neanderthals in Southwest Asia more commonly hunted mountain gazelle, Persian fallow deer, wild goat, and camels. 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 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, 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. It is possible these Neanderthals were curing and storing all this meat before winter set in. Neanderthals at Neumark-Nord may have been rendering fat from animal bones to offset protein toxicity.

Neanderthals competed with several large carnivores, but also seem to have hunted them down, namely cave lions and 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, though it may have only been done in times of extreme food shortages, as in some 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 and Geraldine Finlayson suggested that Neanderthals used various bird parts as artistic media, especially black feathers.

There are several instances of nondescript engravings and scratches on flints, bones, pebbles, and stone slabs — , 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. It is unclear if this constitutes evidence of artmaking because, while modern humans have used red ochre for decorative or symbolic colouration, they have also used ochre 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 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 5 km. and bone. They may have 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, including both likely thrown (Schöningen) and thrusting (Lehringen) types. 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.

The only known Neanderthal tools that could have been used to fashion clothes are hide 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. Anterior dental microwear of Neanderthals living in open environments is similar to that of the modern Ipiutak and Nunavut people, who are known to use their anterior teeth for clamping while preparing hides, suggesting that Neanderthals may have engaged in similar behaviour.

Neanderthals appear to have lived lives of frequent traumatic injury and recovery, indicating the setting of splints and dressing of major wounds. By and large, they appear to have avoided severe infections, indicating long-term treatment. Their knowledge of 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.

While the hyoid bone (a bone that supports the tongue) is almost identical to that of modern humans, this does not provide insight into 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 life after death because it could also have had non-symbolic motivations. 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, 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 burrowing 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. In 2019, the Finlayson's reported that Neanderthals disproportionately butchered the golden eagle over any bird of prey or corvid species, and speculated that Neanderthals viewed the golden eagle as a symbol of power like some recent modern human societies did.

Interbreeding{{anchor|Interbreeding}}

Main article: Interbreeding between archaic and modern humans, Neanderthal genetics

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 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. 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 back migration (the interbreeding population migrated back to Sub-Saharan Africa). In all, approximately 20% of the Neanderthal genome appears to have survived in the modern human gene pool. This Neanderthal DNA is derived 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). Similarly, a large portion of surviving introgression appears to be 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

Main article: Neanderthal extinction

The extinction of Neanderthals was part of the broader 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 exterminating and replacing the inferior species.

In general, the extinction of Neanderthals is ascribed predominantly to competition with modern humans. The success of modern humans over Neanderthals is usually attributed to a higher birth rate and population, facilitated by 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

Main article: Neanderthals 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, Björn Kurtén's Dance of the Tiger, and Jean M. Auel's Clan of the Cave Bear and her Earth's Children series.

Footnotes

References

Sources

References

1. Haeckel, E.. (1895). "Systematische Phylogenie: Wirbelthiere". G. Reimer.
2. Schwalbe, G.. (1906). "Studien zur Vorgeschichte des Menschen". Stuttgart, E. Nägele.
3. Klaatsch, H.. (1909). "Preuves que l{{'}}''Homo Mousteriensis Hauseri'' appartient au type de Neandertal". L'Homme Préhistorique.
4. Romeo, L.. (1979). "Ecce Homo!: a lexicon of man". John Benjamins Publishing Company.
5. (1939). "The stone age of Mount Carmel. The fossil human remains from the Levalloisso-Mousterian". Clarenden Press.
6. (2013). "Evolutionary history of the Primates". Academic Press.
7. {{Cite LPD. 3
8. "Neandertal oder Neanderthal? Was ist denn nun richtig?". Kreisstadt Mettmann.
9. "Neanderthal".
10. Alex, B.. (2016). "Is It Neander-TAL or Neander-THAL?".
11. "Neanderthal".
12. "Neanderthal".
13. Vogt, K. C.. (1864). "Lectures on man: his place in creation, and in the history of the earth". Longman, Green, Longman and Roberts.
14. (1864). "On the Neanderthal skull, or reasons for believing it to belong to the Clydian Period and to a species different from that represented by man". Report of the British Association for the Advancement of Science, Notices and Abstracts, Newcastle-upon-Tyne, 1863.
15. (2015). "The contribution of William King to the early development of palaeoanthropology". Irish Journal of Earth Sciences.
16. Winner, A. K.. (1964). "Terminology". Current Anthropology.
17. King, W.. (1864). "The reputed fossil man of the Neanderthal". Quarterly Journal of Science.
18. Schmerling, P.. (1834). "Recherches sur les ossemens fossiles découverts dans les cavernes de la province de Liége". P. J. Collardin.
19. Menez, A.. (2018). "Custodian of the Gibraltar skull: the history of the Gibraltar Scientific Society". Earth Sciences History.
20. (2002). "The Neandertal type site revisited: interdisciplinary investigations of skeletal remains from the Neander Valley, Germany". Proceedings of the National Academy of Sciences.
21. Schaaffhausen, H.. (1858). "Zur Kenntnis der ältesten Rassenschädel". Archiv für Anatomie, Physiologie und Wissenschaftliche Medicin.
22. (2015). "Handbook of Paleoanthropology". Springer-Verlag Berlin Heidelberg.
23. Fuhlrott, J. C.. (1859). "Menschliche Überreste aus einer Felsengrotte des Düsselthales". Verh Naturhist Ver Preuss Rheinl.
24. Virchow, R.. (1872). "Untersuchung des Neanderthal-Schädels". Verh Berl Anthrop Ges.
25. Drell, J. R. R.. (2000). "Neanderthals: a history of interpretation". Oxford Journal of Archaeology.
26. Van Reybrouck, D.. (2002). "Boule's error: on the social context of scientific knowledge". Antiquity.
27. Langdon, J. H.. (2016). "The science of human evolution: getting it right". Springer.
28. Sommer, M.. (2006). "Mirror, mirror on the wall: Neanderthal as image and 'distortion' in early 20th-century French science and press". Social Studies of Science.
29. (1981). "The significance of Aleš Hrdlička's 'Neanderthal phase of man': A historical and current assessment". American Journal of Physical Anthropology.
30. (2014). "Right for the Wrong Reasons". Current Anthropology.
31. (2010). "A draft sequence of the Neandertal genome". Science.
32. (2024). "''Homo sapiens'', Neanderthals and speciation complexity in palaeoanthropology". Evolutionary Journal of the Linnean Society.
33. (2014). "Neanderthal man: in search of lost genomes". Basic Books.
34. (2011). "Drafting human ancestry: What does the Neanderthal genome tell us about hominid evolution? Commentary on Green et al. (2010)". Human Biology.
35. (2020). "The evolutionary history of Neanderthal and Denisovan Y chromosomes". Science.
36. (2017). "Deeply divergent archaic mitochondrial genome provides lower time boundary for African gene flow into Neanderthals". Nature Communications.
37. (2016). "Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins". Nature.
38. (2021). "Massive cranium from Harbin in northeastern China establishes a new Middle Pleistocene human lineage". The Innovation.
39. (2021). "Neanderthal cranial remains from Baume Moula-Guercy (Soyons, Ardèche, France)". American Journal of Physical Anthropology.
40. Prüfer, K.. (2014). "The complete genome sequence of a Neanderthal from the Altai Mountains". Nature.
41. (2015). "Nuclear and mitochondrial DNA sequences from two Denisovan individuals". Proceedings of the National Academy of Sciences.
42. (2020). "Neanderthal-Denisovan ancestors interbred with a distantly related hominin". Science Advances.
43. Callander, J.. (2004). "Dorothy Garrod's excavations in the Late Mousterian of Shukbah Cave in Palestine reconsidered". Proceedings of the Prehistoric Society.
44. (2007). "Earliest evidence of modern human life history in North African early ''Homo sapiens''". Proceedings of the National Academy of Sciences of the United States of America.
45. (2014). "The chronostratigraphy of the Haua Fteah cave (Cyrenaica, northeast Libya)". Journal of Human Evolution.
46. (2016). "The endocranial anatomy of Maba 1". American Journal of Physical Anthropology.
47. (2017). "Investigating Neanderthal dispersal above 55°N in Europe during the Last Interglacial Complex". Quaternary International.
48. (2018). "On research history and Neanderthal occupation at its northern margins". European Journal of Archaeology.
49. (2004). "The Pleistocene colonization of northeastern Europe: a report on recent research". Journal of Human Evolution.
50. (2011). "Late Mousterian persistence near the Arctic Circle". Science.
51. Slimak, L.. (2012). "Response to "Comment on Late Mousterian persistence near the Arctic Circle"". Science.
52. Zwyns, N.. (2012). "Comment on Late Mousterian persistence near the Arctic Circle". Science.
53. (July 17, 2023). "Late Pleistocene Neanderthal exploitation of stable and mosaic ecosystems in northern Iberia shown by multi-isotope evidence". [[Quaternary Research]].
54. (2016). "The strength of selection against Neanderthal introgression". PLOS Genetics.
55. (2017). "Better support for a small effective population size of Neandertals and a long shared history of Neandertals and Denisovans". Proceedings of the National Academy of Sciences.
56. (2011). "Tenfold population increase in Western Europe at the Neandertal-to-modern human transition". Science.
57. Trinkaus, E.. (2011). "Late Pleistocene adult mortality patterns and modern human establishment". Proceedings of the National Academy of Sciences.
58. (2013). "Neanderthal demographic estimates". Current Anthropology.
59. Trinkaus, E.. (1995). "Neanderthal mortality patterns". Journal of Archaeological Science.
60. Pettitt, R. B.. (2000). "Neanderthal lifecycles: developmental and social phases in the lives of the last archaics". World Archaeology.
61. Hublin, J.-J.. (2002). "Neandertals and Modern Humans in Western Asia".
62. (1985). "The poor brain of ''Homo sapiens neanderthalensis'': see what you please". Alan R. Liss.
63. (1984). "Brain size, cranial morphology, climate, and time machines". Current Anthropology.
64. (2015). "Virtual reconstruction of the Neanderthal Amud 1 cranium". American Journal of Physical Anthropology.
65. (2015). "Brain Ontogeny and Life History in Pleistocene Hominins". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences.
66. (2008). "Middle Cranial Fossa Anatomy and the Origin of Modern Humans". The Anatomical Record.
67. (2010). "Brain Development after Birth Differs between Neanderthals and Modern Humans". Current Biology.
68. (2017). "Nasal airflow simulations suggest convergent adaptation in Neanderthals and modern humans". Proceedings of the National Academy of Sciences.
69. (2011). "Evolution of the Base of the Brain in Highly Encephalized Human Species". Nature Communications.
70. (2012). "Tooth wear, Neanderthal facial morphology and the anterior dental loading hypothesis". Journal of Human Evolution.
71. (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.
72. Helmuth, H.. (1998). "Body height, body mass and surface area of the Neanderthals". Zeitschrift für Morphologie und Anthropologie.
73. (2009). "Energetic competition between Neandertals and anatomically modern humans". PaleoAnthropology.
74. (2022). "Variation in human 3D trunk shape and its functional implications in hominin evolution". Scientific Reports.
75. (2009). "Energetic competition between Neanderthals and anatomically modern humans". PaleoAnthropology.
76. Holliday, T. W.. (1997). "Postcranial evidence of cold adaptation in European Neandertals". American Journal of Physical Anthropology.
77. Trinkaus, E.. (1981). "Aspects of human evolution". Taylor and Francis Ltd..
78. Weaver, T. D.. (2009). "The meaning of Neandertal skeletal morphology". Proceedings of the National Academy of Sciences.
79. (2019). "Palaeoecological and genetic evidence for Neanderthal power locomotion as an adaptation to a woodland environment". Quaternary Science Reviews.
80. (2007). "A melanocortin 1 receptor allele suggests varying pigmentation among Neanderthals". Science.
81. (2012). "Predicting ''Homo'' pigmentation phenotype through genomic data: From neanderthal to James Watson". American Journal of Human Biology.
82. (2017). "The contribution of Neanderthals to phenotypic variation in modern humans". The American Journal of Human Genetics.
83. (2017). "The effect of trauma on Neanderthal culture: A mathematical analysis". Homo.
84. (2017). "External auditory exostoses and hearing loss in the Shanidar 1 Neandertal". PLOS ONE.
85. (2019). "External auditory exostoses among western Eurasian late Middle and Late Pleistocene humans". PLOS ONE.
86. (2016). "Large carnivore attacks on hominins during the Pleistocene: a forensic approach with a Neanderthal example". Archaeological and Anthropological Sciences.
87. Hayden, B.. (2012). "Neandertal social structure?". Oxford Journal of Archaeology.
88. Farizy, C.. (1994). "Spatial patterning of Middle Paleolithic sites". Journal of Anthropological Archaeology.
89. (2010). "Withering away—25,000 years of genetic decline preceded cave bear extinction". Molecular Biology and Evolution.
90. (2011). "Genetic evidence for patrilocal mating behavior among Neandertal groups". Proceedings of the National Academy of Sciences.
91. (2013). "New insights into differences in brain organization between Neanderthals and anatomically modern humans". Proceedings of the Royal Society B.
92. (2015). "Almost 20 years of Neanderthal palaeogenetics: adaptation, admixture, diversity, demography and extinction". Philosophical Transactions of the Royal Society B.
93. Ruebens, K.. (2013). "Regional behaviour among late Neanderthal groups in Western Europe: A comparative assessment of late Middle Palaeolithic bifacial tool variability". Journal of Human Evolution.
94. (2009). "Genetic evidence of geographical groups among Neanderthals". PLOS ONE.
95. (2024). "The Routledge Handbook of Gender Archaeology". Routledge.
96. Jaouen, K.. (2019). "Exceptionally high δ15N values in collagen single amino acids confirm Neandertals as high-trophic level carnivores". Proceedings of the National Academy of Sciences.
97. (2016). "Neandertal versus modern human dietary responses to climatic fluctuations". PLOS ONE.
98. (February 3, 2023). "Hunting and processing of straight-tusked elephants 125.000 years ago: Implications for Neanderthal behavior". [[Science Advances]].
99. (2018). "Dental calculus indicates widespread plant use within the stable Neanderthal dietary niche". Journal of Human Evolution.
100. (2016). "Evidence for the paleoethnobotany of the Neanderthal: a review of the literature". Scientifica.
101. (2015). "Flavouring food: the contribution of chimpanzee behaviour to the understanding of Neanderthal calculus composition and plant use in Neanderthal diets". Antiquity.
102. (2013). "Neanderthal self-medication in context". Antiquity.
103. (2012). "Neanderthal medics? Evidence for food, cooking, and medicinal plants entrapped in dental calculus". The Science of Nature.
104. (2024). "Neanderthal brown crab recipes: A combined approach using experimental, archaeological and ethnographic evidence". Historical Biology.
105. (2013). "New data on human behavior from a 160,000 year old Acheulean occupation level at Lazaret cave, south-east France: An archaeozoological approach". Quaternary International.
106. (2025). "Large-scale processing of within-bone nutrients by Neanderthals, 125,000 years ago". Science Advances.
107. (2018). "Bears and humans, a Neanderthal tale. Reconstructing uncommon behaviors from zooarchaeological evidence in southern Europe". [[Journal of Archaeological Science]].
108. (2013). "Zooarchaeology and Modern Human Origins". Springer Science+Business Media Dordrecht.
109. (2013). "Middle Pleistocene ecology and Neanderthal subsistence: Insights from stable isotope analyses in Payre (Ardèche, southeastern France)". [[Journal of Human Evolution]].
110. (2015). "Cannibalism in the Neanderthal world: an exhaustive revision". Journal of Taphonomy.
111. (2012). "Non utilitarian objects in the Palaeolithic: emergence of the sense of precious?". Archaeology, Ethnology & Anthropology of Eurasia.
112. (2018). "Symbolic use of marine shells and mineral pigments by Iberian Neandertals 115,000 years ago". Science Advances.
113. (April 9, 2020). "Direct evidence of Neanderthal fibre technology and its cognitive and behavioral implications". Scientific Reports.
114. (2018). "Assessing the significance of Palaeolithic engraved cortexes. A case study from the Mousterian site of Kiik-Koba, Crimea". PLOS ONE.
115. (2023). "The earliest unambiguous Neanderthal engravings on cave walls: La Roche-Cotard, Loire Valley, France". PLOS ONE.
116. (2012). "Use of red ochre by early Neandertals". [[Proceedings of the National Academy of Sciences of the United States of America]].
117. (2006). "Mousterian Musicianship? The Case of the Divje Babe I Bone". Oxford Journal of Archaeology.
118. (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.
119. (2020). "Neandertals on the beach: use of marine resources at Grotta dei Moscerini (Latium, Italy)". PLOS ONE.
120. (2020). "Non-destructive ZooMS identification reveals strategic bone tool raw material selection by Neandertals". Scientific Reports.
121. (2019). "Hafting of Middle Paleolithic tools in Latium (central Italy): new data from Fossellone and Sant'Agostino caves". PLOS ONE.
122. Allington-Jones, L. (2015). "The Clacton Spear: The Last One Hundred Years". Archaeological Journal.
123. (December 2015). "New insights on the wooden weapons from the Paleolithic site of Schöningen". Journal of Human Evolution.
124. Milks, A.  (2020) ''[https://sticks-and-stones.blog/wp-content/uploads/2022/11/milks-2021.pdf 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. {{ISBN. 978-94-93194-35-9
125. (2018). "Neandertal fire-making technology inferred from microwear analysis". Scientific Reports.
126. (2019). "Geochemical evidence for the control of fire by Middle Palaeolithic hominins". Scientific Reports.
127. (2016). "Selection and Use of Manganese Dioxide by Neanderthals". Scientific Reports.
128. (2023). "Formation processes, fire use, and patterns of human occupation across the Middle Palaeolithic (MIS 5a-5b) of Gruta da Oliveira (Almonda karst system, Torres Novas, Portugal)". PLOS ONE.
129. (2016). "Faunal evidence for a difference in clothing use between Neanderthals and early modern humans in Europe". Journal of Anthropological Archaeology.
130. Wales, N.. (2012). "Modeling Neanderthal clothing using ethnographic analogues". Journal of Human Evolution.
131. (2025). "Wandering of the auroral oval 41,000 Years Ago". [[Science Advances]].
132. (April 2017). "Anterior dental microwear textures show habitat-driven variability in Neandertal behavior". [[Journal of Human Evolution]].
133. (2019). "Living to fight another day: The ecological and evolutionary significance of Neanderthal healthcare". Quaternary Science Reviews.
134. (2012). "Early seafaring activity in the southern Ionian Islands, Mediterranean Sea". Journal of Archaeological Science.
135. (2018). "Neanderthal language revisited: not only us". Current Opinion in Behavioral Sciences.
136. Johansson, S.. (2015). "Language abilities in Neanderthals". Annual Review of Linguistics.
137. (2018). "Were Neanderthals rational? A stoic approach". Humanities.
138. (1997). "Transmission of tool-making through verbal and non-verbal communication: Preliminary experiments in Levallois flake production". [[Journal of Anthropological Sciences]].
139. (1991). "What sayeth thou Neanderthal? A look at the evolution of their vocal tract and speech". American Journal of Physical Anthropology.
140. (2016). "The evolutionary history of genes involved in spoken and written language: beyond FOXP2". Scientific Reports.
141. (2017). "The Oxford Handbook of Archaeology". Oxford University Press.
142. (2014). "The Cradle of Thought: Growth, Learning, Play and Attachment in Neanderthal Children". Oxford Journal of Archaeology.
143. Leroi-Gourhan, A.. (1975). "The flowers found with Shanidar IV, a Neanderthal burial in Iraq". Science.
144. Solecki, R. S.. (1975). "Shanidar IV: a Neanderthal flower burial in northern Iraq". Science.
145. Sommer, J. D.. (1999). "The Shanidar IV 'flower burial': a re-evaluation of Neanderthal burial ritual". Cambridge Archaeological Journal.
146. Wunn, I.. (2001). "Cave bear worship in the Paleolithic". Cadernos do Laboratorio Xeolóxico de Laxe.
147. (2019). "Neanderthals and the cult of the sun bird". Quaternary Science Reviews.
148. (2015). "An early modern human from Romania with a recent Neanderthal ancestor". Nature.
149. (1989). "Clans and families of Ireland and Scotland, an ethnography of the Gael". McFarland.
150. Huxley, T.. (1891). "The Aryan question and pre-historic man". The Popular Science Monthly.
151. Steensby, H. P.. (1907). "Racestudier i Danmark". Geografisk Tidsskrift.
152. Coon, C. S.. (1962). "The origin of races". Knopf.
153. (1999). "Hominids and hybrids: The place of Neanderthals in human evolution". Proceedings of the National Academy of Sciences.
154. (1999). "The early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern human emergence in Iberia". Proceedings of the National Academy of Sciences of the United States of America.
155. (2009). "The origin of Neandertals". Proceedings of the National Academy of Sciences.
156. (2004). "Neanderthal taxonomy reconsidered: implications of 3D primate models of intra- and interspecific differences". Proceedings of the National Academy of Sciences.
157. (2006). "Early modern humans from the Peștera Muierii, Baia de Fier, Romania". Proceedings of the National Academy of Sciences.
158. (2023). "Diverse African genomes reveal selection on ancient modern human introgressions in Neanderthals". Current Biology.
159. (2020). "Identifying and Interpreting Apparent Neanderthal Ancestry in African Individuals". Cell.
160. (2014). "Resurrecting surviving Neandertal lineages from modern human genomes". Science.
161. (September 2022). "The contribution of Neanderthal introgression to modern human traits". Current Biology.
162. (August 14, 2023). "Integrating sex-bias into studies of archaic introgression on chromosome X". PLOS Genetics.
163. (2016). "Genetic ancestry and natural selection drive population differences in immune responses to pathogens". Cell.
164. Warren, Matthew. (2018). "Mum's a Neanderthal, dad's a Denisovan: First discovery of an ancient-human hybrid". Nature News.
165. (2013). "More genomes from Denisova Cave show mixing of early human groups". Science.
166. (May 8, 2013). "The Quaternary megafaunal extinction and the fate of Neanderthals: An integrative working hypothesis". [[Quaternary International]].
167. (2014). "The timing and spatiotemporal patterning of Neanderthal disappearance". [[Nature (journal).
168. Higham, T.. (2011). "European Middle and Upper Palaeolithic radiocarbon dates are often older than they look: problems with previous dates and some remedies". Antiquity.
169. (2017). "Were Neanderthals responsible for their own extinction?". [[Quaternary International]].
170. (2020). "Neanderthal last stand? Thoughts on Iberian refugia in late MIS 3". Journal of Quaternary Science.
171. (2022). "Peninsular southern Europe refugia during the Middle Palaeolithic: an introduction". Journal of Quaternary Science.
172. (2024). "The Late Middle and Early Upper Palaeolithic in Crimea (Ukraine)—A Review of the Neanderthal Refugium Hypothesis". Journal of Paleolithic Archaeology.
173. Mayr, E.. (1950). "Taxonomic categories in fossil hominids". Cold Spring Harbor Symposia on Quantitative Biology.
174. (2020). "Quantifying the potential causes of Neanderthal extinction: Abrupt climate change versus competition and interbreeding". Quaternary Science Reviews.
175. (2014). "Neandertal demise: an archaeological analysis of the modern human superiority complex". PLOS ONE.
176. (2024). "Stone Tools in Shifting Sands: Past, Present, and Future Perspectives on the Châtelperronian Stone Tool Industry". Journal of Paleolithic Archaeology.
177. (2016). "The Châtelperronian conundrum: blade and bladelet lithic technologies from Quinçay, France". Journal of Human Evolution.
178. (2018). "From Neandertals to modern humans: new data on the Uluzzian". PLOS ONE.
179. Hoffecker, J. F.. (2009). "The spread of modern humans in Europe". [[Proceedings of the National Academy of Sciences of the United States of America]].
180. (2019). "Living on the edge: Was demographic weakness the cause of Neanderthal demise?". PLOS ONE.
181. (2018). "Impact of climate change on the transition of Neanderthals to modern humans in Europe". [[Proceedings of the National Academy of Sciences of the United States of America]].