The first recognised Neanderthal fossil,Neanderthal 1, was discovered in 1856 in theNeander Valley, Germany. At first, Neanderthal 1 was considered to be one of thelower races in accord withhistorical race concepts. As more fossils were discovered through the early 20th century, Neanderthals were characterised as a unique species of underdeveloped human, in particular byMarcellin Boule. By the mid-twentieth century, it was believed thathuman evolution progressed from an ape-like ancestor through a "Neanderthal phase" to modern humans. This gave way to the"Out of Africa" theory in the 1970s. With the sequencing ofNeanderthal genetics first in 2010, it was discovered that Neanderthalsinterbred with modern humans.
Neanderthal anatomy is characterised by a long and low skull, a heavy and rounded brow ridge (supraorbital torus), anoccipital bun (bony projection) at the back of the skull, strong teeth and jaws, a wide chest, and short limbs. These traits gradually became more frequent through theMiddle Pleistocene of Europe, possibly due tonatural selection in a cold climate, as well asgenetic drift when populations collapsed duringglacial periods. Neanderthals would also have been effective sprinters. Neanderthal specimens vary in height from 147.5 to 177 cm (4 ft 10 in to 5 ft 10 in), with average male dimensions estimated at 165 cm (5 ft 5 in) and 75 kg (165 lb). While Neanderthal brain volume and ratio to body size averaged higher than any living human population — 1,640 cc (100 cu in) for males and 1,460 cc (89 cu in) for females — their brain organisation differed from modern humans in areas related to cognition and language, which could explain the comparative simplicity ofNeanderthal behaviour to Cro-Magnons in the archaeological record.
Neanderthals maintained a low population and sufferedinbreeding depression, which may have impeded their ability to progress technologically. They producedMousterianstone tools (aMiddle Palaeolithicindustry) and possibly wore blankets andponchos. They maintained and might havecreated fire. 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 andanimal attacks. Examples ofPalaeolithic 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 buried their dead, but there is no clear indication that they believed inlife after death.
Neanderthals are named after theNeander Valley in which the first identified specimen was found. The valley was spelledNeanderthal and the species was spelledNeanderthaler in German until thespelling reform of 1901.[b] The spellingNeandertal for the species is occasionally seen in English, even in scientific publications, but the scientific name,H. neanderthalensis, is always spelled withth according to theprinciple of priority. The vernacular name of the species in German is alwaysNeandertaler ("inhabitant of the Neander Valley"), whereasNeandertal always refers to the valley.[c][10] The valley itself was named after the late 17th century German theologian and hymn writerJoachim Neander, who often visited the area.[9] His grandfather, a musician, had changed the family name from the original GermanNeumann "new man" (cf. "Newman") to the Graeco-Roman formNeander (deriving with Greekἀνήρănḗr "man"), following the fashion of the time.[11]
Neanderthal 1, thetype 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".[16] The binomial nameHomo neanderthalensis was first proposed by Irish geologistWilliam King in a paper read to the 33rdBritish 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.[17][18][19] 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[d]] conceptions".[20]
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 prehistorianPhilippe-Charles Schmerling in theGrottes 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.[21] In 1848,Gibraltar 1 fromForbes' 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.[22]
FollowingCharles Darwin's 1859On the Origin of Species, Fuhlrott and Schaaffhausen argued that Neanderthal 1 represents a primitivelower human form, aligning more closely with non-human apes as well asNegroids,Eskimos, andAboriginal Australians (which were variably classified as separate species orsubspecies of human at the time).[24][20][25][26] The uniqueness of Neanderthal Man met opposition namely from the pathologistRudolf Virchow, who argued against defining new species based on only a single find. In 1872, Virchow erroneously interpreted Neanderthal characteristics as evidence ofsenility, disease, and malformation instead of archaicness,[27] which stalled Neanderthal research until the end of the century.[24][25]
By the early 20th century, numerous other Neanderthal discoveries were made, establishingH. 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 anatomistGustav Albert Schwalbe. This hypothesis was opposed by French palaeontologistMarcellin Boule, who authored several publications starting in 1908 describing the French Neanderthal specimenLa 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.[24][28][29][30][31]
Boule suggested two different lineages existed in Ice Age Europe: a more evolved one descending from the BritishPiltdown Man (a hoax) to the FrenchGrimaldi Man (aCro-Magnon) which would culminate with modern Europeans; and a less evolved dead-end lineage leading from the GermanHeidelberg 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 asJava Man andPeking 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 theOld World. Some specimens were described as "progressive" Neanderthals which would evolve into some local subspecies ofH. sapiens (polycentricism), while the "classic" Neanderthals of the Western EuropeanWürm glaciation would not.[32]
In the 1970s, with the formulation ofcladistics 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 toH. neanderthalensis.[33] 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").[34] In 2010, the first mapping of the Neanderthalgenome demonstrated that there was at least someinterbreeding between archaic and modern humans.[35] Subsequent genetic studies continue to raise questions on how Neanderthals should be classified relative to modern humans.[36]
Neanderthals can be classified as a unique species asH. neanderthalensis, though some authors argue expanding the definition ofH. sapiens to include other ancient humans, with combinations such asH. sapiens neanderthalensis (splitters and lumpers). The latter opinion has generally been justified usingNeanderthal genetics, as well as inferences on the complexity ofNeanderthal behaviour based on the archaeological record. While there seems to have been some genetic contact between these two groups, there are potential indicators ofhybrid incompatibility,[f] which if true could justify species distinction. The crux of the issue lies in the vagueness of the term "species" (thespecies problem).[36][38][39]
Among identified archaic humans, Neanderthals are most closely related toDenisovans based onnuclear DNA (nDNA) analyses. Denisovans are an enigmatic group ofLate Pleistocene humans only recognisable by a genetic signature rather than anatomical landmarks.[40] Likely due to more recent interbreeding episodes, themitochondrial DNA (mtDNA, passed down maternally)[41] andY-chromosome DNA (passed down paternally)[40] are more similar between Neanderthals and modern humans than between Neanderthals and Denisovans. Similarly, 430,000 year old fossils from theSima de los Huesos are more closely related to Neanderthals in their nDNA, but their mtDNA aligns more closely with Denisovans.[42]
Typical Neanderthal skull traits appear in the European fossil record near the beginning of theMiddle Pleistocene, in specimens usually classified asH. 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 (namelyInuit groups) are often used as modern Neanderthal analogues to study "hyper-arctic" adaptations. Additionally,glacial periods may have forced populations into smallrefugia, reducinggenetic diversity, leading to the development of other typical Neanderthal traits throughgenetic drift orpleiotropy.[33] The 120,000 to 140,000-year-old IsraeliNesher Ramla remains may represent one such source population which would recolonise Europe following thePenultimate Glacial Period.[44]
The occurrence of typical Neanderthal traits in the Middle Pleistocene was highly variable even among individuals of the same population.[33] The speed of Neanderthalization may have also been impeded by gene flow between Western Europe and Africa; this is exemplified by anomalous specimens which lack typical Neanderthal traits, such asCeprano Man.[42] The first recognisable "early Neanderthals" show up in the fossil record by the end ofMarine Isotope Stage 7 (beginning roughly 243,000 years ago) and give way to "classic" or "late Neanderthals" by the end ofMarine Isotope Stage 5e. This spans thePenultimate Glacial Period to theLast Interglacial. Some early Neanderthal teeth fromPayré, France, potentially date to MIS 8, but the dating is uncertain.[1][33]
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.[42][45][46] 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.[47]
Genetic data indicates that Neanderthals, at least after 100,000 years ago, maintained a small population with low genetic diversity, weakeningnatural selection and proliferatingharmful mutations. It is unclear how long European populations suffered this population stress, or to what extent it influenced Neanderthalization.[48]
The Neanderthals were the first human species to permanently occupy Europe.[49] While pre-Neanderthals are mostly identified around Western Europe, classic Neanderthals are recorded across Europe as well asSouthwest[33] and Central Asia, up to theAltai 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.[50]
The southernmost find was recorded atShuqba Cave, Levant;[51] reports of Neanderthals from the North AfricanJebel Irhoud[52] andHaua Fteah[53] have been reidentified asH. sapiens. Their easternmost presence is recorded atDenisova Cave, Siberia85°E; the southeast ChineseMaba Man, a skull, shares several physical attributes with Neanderthals, although these may be the result ofconvergent evolution rather than Neanderthals extending their range to the Pacific Ocean.[54] The northernmost bound is generally accepted to have been55°N, with unambiguous sites known between50–53°N, but this is difficult to assess because glacial advances destroy most human remains.[55][56] Middle Palaeolithic artefacts have been found up to 60°N on the Russian plains,[57][58][59] but these are more likely attributed to modern humans.[60]
Map of Europe 20,000 to 70,000 years ago during theWürm glaciation
It is possible Neanderthal range expanded and contracted as the ice retreated and grew, respectively, to avoidpermafrost areas, residing in certain refuge zones during glacial maxima.[61] Stable environments with mild mean annual temperatures may have been the most suitable Neanderthal habitats.[62]
Like modern humans, Neanderthals probably descended from a very small population with aneffective 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.[63][64] Archaeological evidence suggests that the initial Cro-Magnon population was 10 times higher than Neanderthals.[65]
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.[66][61] 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.[67] Infant mortality was estimated to have been very high for Neanderthals, about 43% in northern Eurasia.[68]
The Neanderthal skull has a flat and broad skullcap, rounded supraorbital torus (the buldge that forms the brow ridges), larger, wideorbits (eye sockets), a broad nose, mid-facialprognathism (the face projects far from thebase of the skull), an "en bombe" (bomb-like) skull shape when viewed from the back, afossa (depression) on the back of the skull below the level of theinion (suprainiac fossa), and anoccipital bun (bony projection) at the back of the skull. Like those of other archaic humans, their jaws lack a truechin.[33]
The Neanderthal braincase averages 1,640 cm3 (100 cu in) for males and 1,460 cm3 (89 cu in) for females,[69] which is significantly larger than the averages for all living populations.[70] The largest Neanderthal brain,Amud 1, was calculated to be 1,736 cm3 (105.9 cu in), one of the largest ever recorded in humans.[71] 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.[72][73][74]
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.[75] A large nose does not necessarily equate to a better sense of smell, and neurologically, because theolfactory bulbs are smaller, Neanderthals may have had a poorer sense of smell andolfactory memory than modern humans.[76]
Thecheek bones are strong, theincisors are large andshovel-shaped, themolars have a swollentooth 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 orattritive foods, or because Neanderthals regularly used the mouth as a third hand.[77]
Neanderthals were generally short and stocky. In a sample of 45 Neanderthallong bones from 14 men and 7 women, the average height was 164 to 168 cm (5 ft 5 in to 5 ft 6 in) for males and 152 to 156 cm (5 ft 0 in to 5 ft 1 in) for females.[78] The fossil record shows that adult Neanderthals varied from about 147.5 to 177 cm (4 ft 10 in to 5 ft 10 in) in height.[79] Average malebody mass index would have been 26.9–28.3 using a size of 164 to 168 cm (5 ft 5 in to 5 ft 6 in) and 76 kg (168 lb).[78][80]
The Neanderthal chest was deep and wide, with a proportionally expansivethoracic cavity, and possibly strongerlung performance. Neanderthals also had relatively morefast-twitch muscle fibres,[81] and much highercaloric demands.[82] The limbs are proportionally short. The body plan has traditionally been explained as a "hyper-arctic" adaptation (Allen's rule).[83][84][85] Stronger lungs, more fast-twitch muscle, and shorter limbs would have also boosted sprinting efficiency.[81][86]
Skin colour seems to have ranged from dark to light. Some Neanderthals had dark or brown hair.[87][88] If red was another possible hair colour, it does not appear to have been a common one.[89]
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.[90] One extreme example isShanidar 1, who shows signs of anamputation of the right arm likely due to anonunion after breaking a bone in adolescence,osteomyelitis (a bone infection) on the leftclavicle, an abnormalgait, vision problems in the left eye, and possible hearing loss[91] (perhapsswimmer's ear).[92] The high trauma rate may be ascribed to a dangerous hunting strategy,[67] or frequent animal attacks.[93]
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 ofbirth defects than in modern humans.[48]
Genetically, Neanderthals may be grouped into three distinct regions (above), dots indicate sampled specimens.[94]
It is difficult to infer Neanderthal group size, but indirect data generally suggests small bands of 10 to 30 individuals.[95] 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.[96] Neanderthals may have been outcompetingcave bears for cave space.[97] Intergroup movement may have been predominantlypatrilocal (male relationships as the basis of groups with females from other groups entering for breeding).[98]
Neanderthals maintained a low population across their range, which may have hindered their ability to maintain long-distance trade routes[99] and to avoidinbreeding.[48] They may have regularly interacted with closely neighbouring communities within a region, but not so often beyond.[100] 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.[94]
While the Cro-Magnons are usually assumed to have generally practisedsexual 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.[101]
Reconstruction of a Neanderthal man butchering a goat at theNeanderthal Museum
Neanderthals were once thought of asscavengers, but are now considered to have beenapex predators.[102] They appear to have eaten predominantly what was abundant within their immediate surroundings.[103] 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.[104][105]
Neanderthals possibly employed a wide range of food preparation techniques. At Cueva del Sidrón, Spain, Neanderthals may have beenroasting andsmoking meat, and used certain plants—such asyarrow andcamomile—for flavouring,[111] although these plants may have instead been used for their medicinal properties.[112][113] AtGorham's Cave, Gibraltar, Neanderthals may have been roastingpinecones to accesspine nuts,[108] and atGruta da Figueira Brava,brown crabs to soften the shell before cracking them open.[114] AtGrotte du Lazaret, France, a total of twenty-three red deer, six ibexes, three aurochs, and oneroe deer appear to have been hunted in a single autumn hunting season, when strong male and female deer herds would group together forrut. It is possible these Neanderthals werecuring and storing all this meat before winter set in.[115] Neanderthals at Neumark-Nord may have beenrendering fat from animal bones to offsetprotein toxicity.[116]
Neanderthals competed with several large carnivores, but also seem to have hunted them down, namelycave lions andwolves,[117] as well as cave andbrown bear both in and out ofhibernation.[118] Neanderthals and other predators may have sometimes avoided competition by pursuing different prey, namely withcave hyenas[104] and wolves (niche differentiation).[119] Neanderthals, nonetheless, were frequently victims ofanimal attacks.[93]
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.[121] Some shells may have been painted.[122] Gibraltarian palaeoanthropologistsClive andGeraldine Finlayson suggested that Neanderthals used various bird parts as artistic media, especially black feathers.[123][124] A 2020 study found evidence of a3-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.[125]
There are several instances of nondescript engravings and scratches on flints, bones, pebbles, and stone slabs — as of 2014[update], 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.[126] Neanderthals may have producedfinger flutings on the walls of La Roche-Cotard over 57,000 years ago.[127]
Neanderthals used ochre, aclay 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.[128]
The 43,000-year-oldDivje Babe flute (a cave bearfemur) from Slovenia has been attributed by some researchers to Neanderthals, though its status as aPalaeolithic flute is heavily disputed. Many researchers consider it to be most likely the product of a carnivorous animal chewing the bone.[129]
Neanderthals manufacturedMiddle Palaeolithicstone tools, and are associated with theMousterianindustry, specifically theLevallois technique. After developing this technology from theAcheulean industry,[130] 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.[61][90] Neanderthals normally collected raw materials from a nearby source, no more than 5 km (3.1 mi).[95] Some communities were also making tools from shells[131] and bone.[132] They may havehafted tips onto spears usingbirch bark tar.[133] European populations have also been manufacturing wood spears, namely the 400,000 year old BritishClacton Spear; 300,000 year old GermanSchöningen spears; and 120,000 year old GermanLehringen Spear,[134] including both likely thrown (Schöningen)[135] and thrusting (Lehringen) types.[136] It has been suggested that Neanderthals likely specifically selected particular wood types (such asEuropean yew in the case of the Clacton and Lehringen spears) for manufacturing spears for their beneficial material properties.[136]
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.[137][138][139] They may have been using fire for cooking, keeping warm, and deterring predators.[140] They were also capable of zoning areas for specific activities, such as for knapping, butchering,hearths, and wood storage.[95] AtAbric 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.[141]
Reconstruction of an elderly Neanderthal man
The only known Neanderthal tools that could have been used to fashion clothes are hidescrapers as no bone sewing-needles andstitching awls have been found as in Cro-Magnon sites. Hide scrapers could have been used to make items similar to blankets orponchos. There is no direct evidence that Neanderthals could make fitted clothes from animal hide.[142][143] Unfitted clothes would have limited range of mobility while dressed, and decreased the time Neanderthals could spend unprotected from the elements away from shelters.[144] Anteriordental 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.[145]
Neanderthals appear to have lived lives of frequent traumatic injury and recovery, indicating the setting ofsplints and dressing of major wounds. By and large, they appear to have avoided severe infections, indicating long-term treatment. Their knowledge ofmedicinal plants was comparable to that of Cro-Magnons.[146]
Stone tools on various Greek islands could indicate early seafaring through the Mediterranean, employing simple reed boats for one-day crossings,[147] but the evidence for such a big claim is limited.[148]
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.[149][150][151] In experiments with modern humans, theLevallois technique can be taught with purelyobservational learning without spoken instruction.[152]
While thehyoid 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.[153] Neanderthals had theFOXP2 gene, which is associated with speech and language development, but not the modern human variant.[154]
Engraved flint from a Neanderthal grave atKiik-Koba, Crimea
Neanderthals, probably uncommonly, buried their dead. This may explain the abundance of fossil remains.[103] The behaviour is not indicative of a religious belief oflife after death because it could also have hadnon-symbolic motivations.[155][156] The dead were buried in simple, shallow graves and pits,[156] 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.[157] Some sites with multiple well-preserved Neanderthal skeletons may representcemeteries.[156]
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 andhollyhock.[158] The medicinal properties of the plants led American archaeologistRalph Solecki to claim that the man buried was some leader, healer, orshaman, and that "the association of flowers with Neanderthals adds a whole new dimension to our knowledge of his humanness, indicating that he had 'soul'".[159] It is also possible the pollen was deposited by a small burrowing rodent after the man's death.[160]
Neanderthals were once thought to have ritually killed and eaten cave bears or other Neanderthals, but the evidence is circumstantial.[155] In 2019, the Finlayson's reported that Neanderthals disproportionately butchered thegolden eagle over anybird of prey orcorvid species, and speculated that Neanderthals viewed the golden eagle as a symbol of power like some recent modern human societies did.[124]
Reconstruction ofOase 2 with around 7.3% Neanderthal DNA (from an ancestor 4–6 generations back)[161]
Hybridisation between Neanderthals and early modern humans had been suggested early on,[162] such as by English anthropologistThomas Huxley in 1890,[163] Danish ethnographerHans Peder Steensby in 1907,[164] and Coon in 1962.[165] In the early 2000s, supposed hybrid specimens were discovered:Lagar Velho 1[166][167][168][169] andMuierii 1.[170] Similar anatomy could also have been caused by adapting to a similar environment rather than interbreeding.[171]
The first Neanderthal genome sequence was published in 2010, and strongly indicated interbreeding between Neanderthals and early modern humans.[35] 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 ago. Interbreeding also occurred in other populations which are not ancestral to any living person.[172] 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 toSub-Saharan Africa).[173] In all, approximately 20% of the Neanderthal genome appears to have survived in the modern humangene pool.[174] This Neanderthal DNA is derived primarily from the children of female modern humans and male Neanderthals.[175][37]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 benon-coding ("junk") DNA with few biological functions.[171] Some Neanderthal-derived genes, nonetheless, may have functional implications related to metabolism, brain function, and skeletal and muscular development.[35][176] Some genes may have helped immigrating modern human populations acclimatise faster, such as genes related toimmune response.[177]
Neanderthals in the Siberian Altai Mountains interbred with the local Denisovan population, and it may have been a common occurrence here.[178] About 17% of the genome of one Altai Denisovan specimen derived from Neanderthals.[179]
The extinction of Neanderthals was part of the broaderLate Pleistocene megafaunal extinction event.[181] Neanderthals were replaced by modern humans, indicated by the near-complete replacement of Middle Palaeolithic Mousterian stone technology with modern humanUpper PalaeolithicAurignacian stone technology across Europe (the Middle-to-Upper Palaeolithic Transition) from 39,000 to 41,000 years ago.[180][182][183] Neanderthals may have persisted in Spain for longer, but the dates of the latest Mousterian and earliest Aurignacian are poorly constrained. InCatalonia andAragón (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.[184] Similar refuge zones have also been proposed on other temperate European peninsulas, namely Italy, the Balkans, andCrimea.[185][186]
Historically, the cause of extinction of Neanderthals and other archaic humans was viewed under an imperialistic guise, with the superior invading modern humansexterminating and replacing the inferior species.[24]
Whensapiens began to expand and spread, he eliminated the other contemporary races [including Neanderthals] just as the white man drove out the Australian aborigines and the North American Indians.
In general, the extinction of Neanderthals is ascribed predominantly tocompetition 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.[188] Assimilation had long been hypothesised with supposed hybrid specimens, and was revitalised with the discovery of archaic human DNA in modern humans.[189] Similarly, theChâtelperronian industry of central France and northern Spain may represent a culture of Neanderthals adopting modern human techniques, viaacculturation.[190][191] Other ambiguous transitional cultures include the ItalianUluzzian industry,[192] and the Central EuropeanSzeletian industry.[193]
Neanderthal extinction has also been ascribed to their low population as well as the resultingmutational meltdown, making them less adaptable to major environmental changes or new diseases introduced by immigrating modern humans.[194] 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 (specificallyHeinrich event 4) or a natural disaster (theCampanian Ignimbrite eruption), Neanderthals may not have been as fast as modern humans in recolonising.[195] TheLaschamp event 39,000 to 42,000 years ago may have increasedultraviolet radiation, disproportionately affecting Neanderthals who lacked protective fitted clothes, and may not have utilised ochre as sunscreen to the extent Cro-Magnons did.[144]
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.[196]
^The German/t/ phoneme was frequently spelledth from the 15th to 19th centuries until theGerman Orthographic Conference of 1901. The German spellingThal ("valley", acognate of Englishdale) changed toTal, and theh was also dropped fromNeandertal for the valley andNeandertaler for the species.[10]
^InMettmann, "Neander Valley", there is a local idiosyncrasy in use of the outdated spellings withth, such as with theNeanderthal Museum (but the name is in English [German would requireNeandertalermuseum]), theNeanderthal station (Bahnhof Neanderthal), and some other rare occasions meant for tourists. Beyond these, city convention is to useth when referring to the species.[10]
^The bones were discovered by workers ofWilhelm Beckershoff andFriedrich Wilhelm Pieper. Initially, the workers threw the bones out as debris, but Beckershoff then told them to store the bones. Pieper asked Fuhlrott to come up to the cave and investigate the bones, which Beckershoff and Pieper believed belonged to acave bear.[9]
^abc"Neandertal oder Neanderthal? Was ist denn nun richtig?" [Neandertal or Neanderthal? So which is actually right?]. Kreisstadt Mettmann. Archived fromthe original on August 9, 2020. RetrievedFebruary 1, 2017.Heute sollten Ortsbezeichnungen das 'Neandertal' ohne 'h' bezeichnen. Alle Namen, die sich auf den prähistorischen Menschen beziehen, führen das 'h'. (Nowadays, place names should refer to the Neander Valley ['Neandertal'] without an 'h'. All names referring to the prehistoric humans have the 'h'.)
^Schaaffhausen, H. (1858). "Zur Kenntnis der ältesten Rassenschädel" [Acknowledging the oldest racial skull].Archiv für Anatomie, Physiologie und Wissenschaftliche Medicin (in German):453–478.
^abSchlager, S.; Wittwer-Backofen, U. (2015). "Images in Paleoanthropology: Facing Our Ancestors". In Henke, W.; Tattersall, I. (eds.).Handbook of Paleoanthropology. Springer-Verlag Berlin Heidelberg. pp. 1019–1027.doi:10.1007/978-3-642-39979-4_70.ISBN978-3-642-39978-7.
^Langdon, J. H. (2016). "Case study 18. Neanderthals in the mirror: imagining our relatives".The science of human evolution: getting it right. Springer.ISBN978-3-319-41584-0.
^Spencer, F.; Smith, F. H. (1981). "The significance of Aleš Hrdlička's 'Neanderthal phase of man': A historical and current assessment".American Journal of Physical Anthropology.56 (4):435–459.doi:10.1002/ajpa.1330560417.
^abcdefHublin, J.-J. (2002). "Climatic Changes, Paleogeography, and the Evolution of the Neandertals". In Akazawa, T.; Aoki, K.; Bar-Yosef, O. (eds.).Neandertals and Modern Humans in Western Asia.doi:10.1007/b109961.ISBN0-306-45924-8.
^Holliday, T. W.; Gautney, J. R.; Friedl, L. (2014). "Right for the Wrong Reasons".Current Anthropology.55 (6):696–724.doi:10.1086/679068.
^abChevy, Elizabeth T.; Huerta-Sánchez, Emilia; Ramachandran, Sohini (August 14, 2023)."Integrating sex-bias into studies of archaic introgression on chromosome X".PLOS Genetics.19 (8) e1010399.doi:10.1371/journal.pgen.1010399.ISSN1553-7404.PMC10449224.PMID37578977.We have shown that the observed low level of archaic coverage on chromosome X could be explained merely by a reduction in the effect of heterosis and sex-biases in the introgression events, without involving a more complex model with hybrid incompatibilities. Our work also suggests that negative selection was likely acting on archaic variants, and provides an appropriate set of null models for evaluating positive selection on introgressed segments on chromosome X.
^Callander, J. (2004). "Dorothy Garrod's excavations in the Late Mousterian of Shukbah Cave in Palestine reconsidered".Proceedings of the Prehistoric Society.70:207–231.doi:10.1017/S0079497X00001171.S2CID191630165.
^Nielsen, T. K.; Benito, B. M.; et al. (2017). "Investigating Neanderthal dispersal above 55°N in Europe during the Last Interglacial Complex".Quaternary International.431:88–103.Bibcode:2017QuInt.431...88N.doi:10.1016/j.quaint.2015.10.039.
^Nielsen, T. K.; Riede, F. (2018). "On research history and Neanderthal occupation at its northern margins".European Journal of Archaeology.21 (4):506–527.doi:10.1017/eaa.2018.12.S2CID165849999.
^Holloway, R. L. (1985). "The poor brain ofHomo sapiens neanderthalensis: see what you please". In Delson, E. (ed.).Ancestors: The hard evidence. Alan R. Liss.ISBN0-471-84376-8.
^Cerqueira, C. C.; Piaxão-Côrtes, V. R.; Zambra, F. M. B.; Hünemeier, T.; Bortolini, M. (2012). "PredictingHomo pigmentation phenotype through genomic data: From neanderthal to James Watson".American Journal of Human Biology.24 (5):705–709.doi:10.1002/ajhb.22263.PMID22411106.S2CID25853632.
^French, Jennifer C. (2024). "Sex, gender, and the division of labour in the European Middle and Upper Palaeolithic".The Routledge Handbook of Gender Archaeology. London: Routledge. pp. 161–174.doi:10.4324/9781003257530-14.ISBN978-1-003-25753-0.
^Valensi, P.; Michel, V.; et al. (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.316:123–139.Bibcode:2013QuInt.316..123V.doi:10.1016/j.quaint.2013.10.034.
^Lycett, S. J.; von Cramon-Taubadel, N. (2013). "A 3D morphometric analysis of surface geometry in Levallois cores: patterns of stability and variability across regions and their implications".Journal of Archaeological Science.40 (3):1508–1517.Bibcode:2013JArSc..40.1508L.doi:10.1016/j.jas.2012.11.005.
^Collard, M.; Tarle, L.; Sandgathe, D.; Allan, A. (2016). "Faunal evidence for a difference in clothing use between Neanderthals and early modern humans in Europe".Journal of Anthropological Archaeology.44:235–246.Bibcode:2016JAnAr..44..235C.doi:10.1016/j.jaa.2016.07.010.hdl:2164/9989.
^Ferentinos, G.; Gkioni, M.; Geraga, M.; Papatheodorou, G. (2012). "Early seafaring activity in the southern Ionian Islands, Mediterranean Sea".Journal of Archaeological Science.39 (7):2167–2176.Bibcode:2011JQS....26..553S.doi:10.1016/j.jas.2012.01.032.
^Sommer, J. D. (1999). "The Shanidar IV 'flower burial': a re-evaluation of Neanderthal burial ritual".Cambridge Archaeological Journal.9 (1):127–129.doi:10.1017/s0959774300015249.S2CID162496872.
^abHigham, T.; Douka, K.; Wood, R.; Ramsey, C. B.; Brock, F.; Basell, L.; Camps, M.; Arrizabalaga, A.; Baena, J.; Barroso-Ruíz, C.; C. Bergman; C. Boitard; P. Boscato; M. Caparrós; N.J. Conard; C. Draily; A. Froment; B. Galván; P. Gambassini; A. Garcia-Moreno; S. Grimaldi; P. Haesaerts; B. Holt; M.-J. Iriarte-Chiapusso; A. Jelinek; J.F. Jordá Pardo; J.-M. Maíllo-Fernández; A. Marom; J. Maroto; M. Menéndez; L. Metz; E. Morin; A. Moroni; F. Negrino; E. Panagopoulou; M. Peresani; S. Pirson; M. de la Rasilla; J. Riel-Salvatore; A. Ronchitelli; D. Santamaria; P. Semal; L. Slimak; J. Soler; N. Soler; A. Villaluenga; R. Pinhasi; R. Jacobi; et al. (2014). "The timing and spatiotemporal patterning of Neanderthal disappearance".Nature.512 (7514):306–309.Bibcode:2014Natur.512..306H.doi:10.1038/nature13621.hdl:1885/75138.PMID25143113.S2CID205239973.We show that the Mousterian [the Neanderthal tool-making tradition] ended by 41,030–39,260 calibrated years BP (at 95.4% probability) across Europe. We also demonstrate that succeeding 'transitional' archaeological industries, one of which has been linked with Neanderthals (Châtelperronian), end at a similar time.
^Higham, T. (2011). "European Middle and Upper Palaeolithic radiocarbon dates are often older than they look: problems with previous dates and some remedies".Antiquity.85 (327):235–249.doi:10.1017/s0003598x00067570.S2CID163207571.Few events of European prehistory are more important than the transition from ancient to modern humans about 40,000 years ago, a period that unfortunately lies near the limit of radiocarbon dating. This paper shows that as many as 70 per cent of the oldest radiocarbon dates in the literature may be too young, due to contamination by modern carbon.
^Straus, Lawrence Guy (2020). "Neanderthal last stand? Thoughts on Iberian refugia in late MIS 3".Journal of Quaternary Science.37 (2):283–290.doi:10.1002/jqs.3252.ISSN1099-1417.
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Papagianni, D.; Morse, M. A. (2013)."Still with us?".Neanderthals rediscovered: how modern science is rewriting their story. Thames and Hudson.ISBN978-0-500-77311-6.
Shipman, P. (2015). "How humans and their dogs drove Neanderthals to extinction".The invaders: how humans and their dogs drove Neanderthals to extinction. Harvard University Press.doi:10.2307/j.ctvjf9zbs.ISBN978-0-674-42538-5.JSTORj.ctvjf9zbs.
Tattersall, I. (2015). "Neanderthals, DNA, and creativity".The strange case of the Rickety Cossack: and other cautionary tales from human evolution. St. Martin's Publishing Group.ISBN978-1-4668-7943-0.
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