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For other uses, see Neanderthal (disambiguation).
Temporal range: MiddleLate Pleistocene 0.250–0.024 Ma
Homo sapiens neanderthalensis.jpg
The Neanderthal skull of La Chapelle-aux-Saints 1
Mounted Neanderthal skeleton at American Museum of Natural History
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Haplorhini
Family: Hominidae
Genus: Homo
Species: H. neanderthalensis
Binomial name
Homo neanderthalensis
King, 1864
Range of Homo neanderthalensis.png

Homo mousteriensis[1]
Homo sapiens neanderthalensis
Palaeoanthropus neanderthalensis[2]

Neanderthals, or more rarely Neandertals,[a] (UK /niˈændərˌtɑːl/, us also /n/-, -/ˈɑːndər/-, -/ˌtɔːl/, -/ˌθɔːl/;[6][7] named for the Neandertal region in Germany) were a species or subspecies of archaic humans in the genus Homo that became extinct about 24,000 years ago.[8][9][10][11] Neanderthals and modern humans share 99.7% of their DNA[12] and are hence closely related.[13][14] (By comparison, both modern humans and Neanderthals share 98.8% of their DNA with their closest non-human living relatives, the chimpanzees.)[12] Neanderthals left bones and stone tools in Eurasia, from Western Europe to Central and Northern Asia. Due to accumulating genetic and fossil evidence suggesting Neanderthals evolved in Europe separately from modern humans in Africa for over 600,000 years, paleontologists generally classify them as a distinct species, Homo neanderthalensis. From the 1950s to the early 1980s, however, Neanderthals were widely considered a subspecies of Homo sapiens (H. s. neanderthalensis) and a minority of scholars still hold this view.[15][16][17]

Several cultural assemblages have been linked to the Neanderthals in Europe. The earliest, the Mousterian stone tool culture, dates to about 160,000 years ago.[18] Late Mousterian artifacts were found in Gorham's Cave on the south-facing coast of Gibraltar.[19][20]

Compared to Homo sapiens, Neanderthals had a lower surface-to-volume ratio, with shorter legs and a bigger body, in conformance with Bergmann's rule, as an energy-loss reduction adaptation to life in a high-latitude (i.e. seasonally cold) climate. Male Neanderthals had cranial capacities averaging 1600 cm3, females 1300 cm3,[21][22] extending to 1736 cm3 in Amud 1.[23] This is notably larger than the 1250–1400 cm3 typical of modern humans. Males stood 164–168 cm (65–66 in) and females 152–156 cm (60–61 in) tall.[24]

The Neanderthal genome project published papers in 2010 and 2014 stating that Neanderthals contributed to the DNA of modern humans, including most humans outside sub-Saharan Africa, as well as a few populations in sub-Saharan Africa, through interbreeding, likely between 50,000 and 60,000 years ago.[25][26][27][28] Recent studies also show that a few Neanderthals began mating with ancestors of modern humans long before the large "out of Africa migration" of present day non-Africans, as early as 100,000 years ago.[29] In 2016, research indicated that there were three distinct episodes of interbreeding between modern humans and Neanderthals: the first encounter involved the ancestors of non-African modern humans, probably soon after leaving Africa; the second, after the ancestral Melanesian group had branched off (and subsequently had a unique episode of interbreeding with Denisovans); and the third, involving the ancestors of East Asians only.[30]

Claims that Neanderthals deliberately buried their dead, and if they did, whether such burials had any symbolic meaning,[31] are heavily contested.[32][33][34] The debate on deliberate Neanderthal burials has been active since the 1908 discovery of the well-preserved Chapelle-aux-Saints 1 skeleton in a small hole in a cave in southwestern France. In this controversy's most recent installment, a team of French researchers reinvestigated the Chapelle-aux-Saints cave and in January 2014 reasserted the century-old claim that the 1908 Neanderthal specimen had been deliberately buried,[35] and this has in turn been heavily criticised.[36]

In 2013, scientists sequenced the entire genome of a Neanderthal for the first time. The genome was extracted from the phalanx bone of a 50,000-year-old Neanderthal found in a Siberian cave.[37][38][39] In 2016, elaborate constructions of rings of broken stalagmites made by early Neanderthals around 176,000 years ago were discovered 336 m (1,102 ft) inside Bruniquel Cave in southwestern France. This would have required a more advanced social structure than previously known for Neanderthals.[40]


The species is named after one of the first sites where its fossils were discovered in the 19th century, about 12 km (7 mi) east of Düsseldorf, Germany, in the Feldhofer Cave, located in the Düssel River's Neander valley.[b] Thal is an older spelling of the German word Tal (with the same pronunciation), which means "valley" (cognate with English dale).[c][42][43]


The German pronunciation of Neanderthaler or Neandertaler is [neˈandɐˌtʰaːlɐ] in the International Phonetic Alphabet. In British English, "Neanderthal" is pronounced with the /t/ as in German, but different vowels (IPA: /niːˈændərtɑːl/).[46][47][48] In layman's American English, "Neanderthal" is pronounced with a /θ/ (the voiceless th as in thin) and /ɔ/ instead of the longer British /aː/ (IPA: /niːˈændərθɔːl/),[49] although scientists typically use the /t/ as in German.[50][51]


Scientists to this day debate over whether Neanderthals should be classified as a distinct species - Homo neanderthalensis - or as Homo sapiens neanderthalensis, the latter placing Neanderthals as a subspecies of H. sapiens.[52][53] During the early 20th century the prevailing view was heavily influenced by Arthur Keith and Marcellin Boule, who wrote the first scientific description of a nearly complete Neanderthal skeleton.[54] Senior members of their respective national paleontological institutes and among the most eminent paleoanthropologists of their time,[55][56] both men argued that this "primitive" Neanderthal could not be a direct ancestor of modern man. This idea was reflected in an erroneous and inaccurate reconstruction of the Neanderthal findings of La Chapelle-aux-Saints, mounted in a crooked pose with a deformed and heavily curved spine and legs buckled.[57]

During the 1930s scholars Ernst Mayr, George Gaylord Simpson and Theodosius Dobzhansky reinterpreted the existing fossil record and came to different conclusions.[58] Neanderthal man was classified as Homo sapiens neanderthalensis - an early subspecies contrasted with what was now called Homo sapiens sapiens. The obviously unbroken succession of fossil sites of both subspecies in Europe was considered evidence that there was a slow and gradual evolutionary transition from Neanderthals to modern humans. Contextual interpretations of similar excavation sites in Asia lead to the hypothesis of multiregional origin of modern man in the 1980s.[59]

Current views[edit]

Current scientific ideas hold that both species evolved from a common African ancestor: Homo erectus. Fossil and tool finds support the idea that during the first migration wave around 2 million years ago, small groups of Homo erectus left Africa via the Levant and the Black Sea area into Georgia (fossils of Dmanisi) and possibly via north-west Africa towards southern Spain. The 1.2-million-year-old fragmented finds in Spain are referred to as Homo antecessor by their discoverers, and considered to be the ancestors of Neanderthal. This interpretation, however, is controversial.

Some 600,000 years ago a second propagation wave of African Homo erectus took place. Skulls from that period found in Spain for example, suggest a brain volume of between 1100 and 1450 cm3. The brain volume of fossils from the first propagation wave is however estimated only to be slightly above 1000 ccm. Homo erectus that arrived in Europe in the second wave of colonization developed into Neanderthals through the intermediate Homo heidelbergensis. Since about 200,000 years ago in Africa the so-called early anatomically modern humans descended from the local Homo erectus through the intermediate Homo rhodesiensis. Evidence from mitochondrial DNA studies has been interpreted as evidence that Neanderthals were not a subspecies of H. sapiens.[60] Others, for example University of Cambridge Professor Paul Mellars, say "no evidence has been found of cultural interaction".[61]

The period when the Neanderthal lineage separated from the lineage that led to modern man remains debated. In 2010 a period between 440,000 and 270,000 years B.P. was calculated by the molecular clock method. However, the reliability of molecular clocks is questionable as dating determined by stratigraphic methods varies considerably from those determined using the molecular clock. In 2012 recalculation of the mutation rates revealed evidence of a much earlier separation: between 800,000 and 400,000 years ago.


The first representatives of the genus Homo with proto-Neanderthal[62] traits are believed to have existed in Eurasia as early as 350,000–600,000 years ago[63] with the first "true Neanderthals" appearing between 200,000 and 250,000 years ago.[64] The exact date of their extinction had been disputed but in 2014, a team led by Thomas Higham of the University of Oxford used an improved radiocarbon dating technique on material from 40 archaeological sites to show that Neanderthals died out in Europe between 41,000 and 39,000 years ago, with the last group disappearing from southern Spain 24,000 years ago.[8][9] Radiocarbon dating performed on charcoal in Gorham's Cave in Gibraltar in 2006 suggests that Neanderthals lived there 24,000 to 28,000 years ago. This coincides with the start of a very cold period in Europe and is 5,000 to 18,000 years after Homo sapiens reached the continent.[65]




Neanderthal skulls were first discovered in the Engis Caves (the partial skull dubbed Engis 2) (fr), in what is now Belgium (1829) by Philippe-Charles Schmerling and in Forbes' Quarry, Gibraltar, dubbed Gibraltar 1 (1848), both prior to the type specimen discovery in a limestone quarry of the Neander Valley in Erkrath near Düsseldorf in August 1856, three years before Charles Darwin's On the Origin of Species was published.[70]


To date, the bones of over 400 Neanderthals have been found.[71]

Timeline of research[edit]

Neanderthal fossils
Skull, found in 1886 in Spy, Belgium
Frontal bone of a neanderthal child from the cave of La Garigüela
Skull from La Chapelle aux Saints
Semi-frontal view of a neanderthal skull from Gibraltar
  • 1829: The Engis 2 Neanderthal skulls were discovered in Engis, in present-day Belgium.
  • 1905: During the excavation in Krapina more than 5,000 items were found, of which 874 residue of human origin, including bones of prehistoric man and animals, artifacts.
  • 1908: A nearly complete Neanderthal skeleton was discovered in La Chapelle-aux-Saints, France in association with Mousterian tools and bones of extinct animals.[72]
  • 1926: Skull fragments of Gibraltar 2, a four-year-old Neanderthal girl, discovered by Dorothy Garrod.
  • 1981: The Bontnewydd Palaeolithic site, Wales yields the most north-western site in Eurasia.
  • 1987: Thermoluminescence results from Israeli fossils dates Kebara 2 to 60,000 BP and humans at Qafzeh to 90,000 BP. These dates were confirmed by electron spin resonance (ESR) dates for Qafzeh (90,000 BP) and Es Skhul (80,000 BP).
  • 1994: Neanderthal remains inadvertently uncovered inside the Sidrón Cave in Piloña municipality, Asturias, northwestern Spain. Since then the remains of at least 12 individuals: three men, three adolescent boys, three women, and three infants have been found. In 2009 Neanderthal ancient mtDNA was partially sequenced in HVR region for three distinct Neanderthals there.[73][74]
  • 1997–2000: During new excavations in the Neandertal, additional bone fragments are found, some of which fit the fragments found in 1856, thus pinpointing the exact location of the original find. The exact location had previously been unknown, as the site of the find (the „Kleine Feldhofer Grotte“) was destroyed by limestone mining.[76]
  • 2000: Igor Ovchinnikov et al. retrieved mitochondrial DNA of a Neanderthal infant from Mezmaiskaya Cave in the Caucasus.[78]
  • 2005: The Max Planck Institute for Evolutionary Anthropology launched the Neanderthal genome project in co-operation with Connecticut-based 454 Life Sciences and the Institute of Quaternary Paleontology and Geology in Zagreb, Croatia.[79] In 2009, the Max Planck Institute announced the "first draft" of a complete Neanderthal genome is completed.[80]
  • 2014: Prof Thomas Higham of the University of Oxford performed the most comprehensive dating of Neanderthal bones and tools ever carried out, which demonstrated that Neanderthals died out in Europe between 41,000 and 39,000 years ago—this coincides with the start of a very cold period in Europe and is 5,000 years after Homo sapiens reached the continent.[87][88] Other studies suggest a more recent date, around 24,000 years ago instead.[8][9][10][11]

Habitat and range[edit]

Further information: List of Neanderthal sites
Sites where typical Neanderthal fossils have been found

Early Neanderthals lived in the last glacial period for a span of about 100,000 years. Because of the damaging effects the glacial period had on the Neanderthal sites, not much is known about the early species. Countries where their remains are known include most of Europe south of the line of glaciation, roughly along the 50th parallel north. This includes most of Western Europe, Central Europe, the Carpathians, and the Balkans,[89] some sites in Ukraine and in western Russia, Central and Northern Asia up to the Altai Mountains, and Western Asia from the Levant up to the Indus River. The Bontnewydd Palaeolithic site at Denbighshire, North Wales is the most north-western site of Neanderthal remains and one of the oldest remains in Britain (230,000 years ago). It is estimated that the total Neanderthal population across this habitat range numbered at around 70,000 at its peak.[90]



Separate developments in the human line, in other regions such as Southern Africa, somewhat resembled the Eurasian Neanderthals, but these people were not Neanderthals. One such example is Rhodesian Man (Homo rhodesiensis), who existed long before any classic Eurasian Neanderthals, but had a more modern set of teeth. Some H. rhodesiensis populations appeared to be on the road to modern H. sapiens sapiens. At any rate, the populations in Eurasia underwent increasing "Neanderthalization" over time. Some have suggested that H. rhodesiensis in general was ancestral to both modern humans and Neanderthals, and that at some point the two populations went their separate ways, but this argument presupposes that H. rhodesiensis existed around 600,000 years ago.



Classic Neanderthal fossils have been found over a large area, from northern Germany to Israel and Mediterranean countries like Spain[94] and Italy[95] in the south and from England and Portugal in the west to Uzbekistan in the east. This area probably was not occupied all at the same time. The northern border of their range, in particular, would have contracted frequently with the onset of cold periods. On the other hand, the northern border of their range as represented by fossils may not be the real northern border of the area they occupied, since Middle Palaeolithic-looking artifacts have been found even farther north, up to 60° N, on the Russian plain.[96] Recent evidence has extended the Neanderthal range by about 1,250 miles (2,010 km) east into southern Siberia's Altai Mountains.[97][98]


Main article: Neanderthal anatomy

Neanderthal anatomy differed from modern humans in that they had a more robust build and distinctive morphological features, especially on the cranium, which gradually accumulated more derived aspects as it was described by Marcellin Boule,[99] particularly in certain isolated geographic regions. These include shorter limb proportions, a wider, barrel-shaped rib cage, a reduced chin, and a large nose, which was much larger in both length and width, and started somewhat higher on the face, than in modern humans.[64] Evidence suggests they were much stronger than modern humans, with particularly strong arms and hands,[100][101] while they were comparable in height; based on 45 long bones from at most 14 males and 7 females, Neanderthal males averaged 164–168 cm (65–66 in) and females 152–156 cm (60–61 in) tall.[24] Samples of 26 specimens in 2010 found an average weight of 77.6 kg (171 lb) for males and 66.4 kg (146 lb) for females.[102] A 2007 genetic study suggested some Neanderthals may have had red hair and blond hair, along with a light skin tone.[103]

Gregory Cochran and Henry Harpending in the book "The 10,000 year explosion," investigated whether it is accurate to depict Neanderthals as having hair pattens similar to Homo Sapiens. They concluded that, "We don’t yet know for sure, but it seems likely that, as part of their adaptation to cold, Neanderthals were furry. Chimpanzees have ridges on their finger bones that stem from the way that they clutch their mother’s fur as infants. Modern humans don’t have these ridges, but Neanderthals do." (Cochran, G. & H. Harpending (2009).[full citation needed]

In 'The Spread of Modern Humans in Europe; 2002 John F Hoffecker, writes "Neanderthal sites show no evidence of tools for making tailored clothing. There are only hide scrapers, which might have been used to make blankets or ponchos. This is in contrast to Upper Paleolithic (modern human) sites, which have an abundance of eyed bone needles and bone awls Moreover, microwear analysis of Neanderthal hide scrapers shows that they were used only for the initial phases of hide preparation, and not for the more advanced phases of clothing production ."[full citation needed]


Neanderthals are known for their large cranial capacity, which at 1600 cm3 is larger on average than that of modern humans. One study has found that Neanderthal brains were more asymmetric than other hominid brains.[105] In 2008, a group of scientists produced a study using three-dimensional computer-assisted reconstructions of Neanderthal infants based on fossils found in Russia and Syria. It indicated that Neanderthal and modern human brains were the same size at birth, but that by adulthood, the Neanderthal brain was larger than the modern human brain.[106] They had almost the same degree of encephalization (i.e. brain to body size ratio) as modern humans.[107][108]


Main article: Neanderthal behavior

Neanderthals made advanced tools,[109] apparently kindled fire on demand,[110] may have had a language (the nature of which is debated and likely unknowable), and lived in complex social groups.[111][112][113] The Molodova I archaeological site in eastern Ukraine suggests some Neanderthals built dwellings using animal bones. A building was made of mammoth skulls, jaws, tusks and leg bones, and had 25 hearths inside.[114][115]

Circumstantial evidence suggests Neanderthals may have been building some form of watercraft since the Middle Paleolithic.[116][117] Scientists have speculated that these watercraft may have been similar to dugout canoes, which are among the oldest known boats in the archaeological record.[117] Mousterian stone tools discovered on the southern Ionian Islands suggests that Neanderthals were sailing the Mediterranean Sea as early as 110,000 years ago.[118] Quartz hand-axes, three-sided picks, and stone cleavers from Crete have also been recovered that date back about 170,000 years ago.[119]

It was once thought that Neanderthals lacked the sophistication for hunting, perhaps scavenging meat from carcasses,[64] but increasing evidence suggests they were apex predators,[120][121] capable of bringing down a wide range of prey from red deer, reindeer, ibex and wild boar, to larger animals such as aurochs and even, on occasion, mammoth, straight-tusked elephant and rhinoceros.[64][122] However, while they were largely carnivorous,[123][124] new studies indicate Neanderthals also had cooked vegetables in their diet.[121][125] In 2010, an isotope analysis of Neanderthal teeth found traces of cooked vegetable matter, and more recently a 2014 study of Neanderthal coprolites (fossilized feces) found substantial amounts of plant matter, contradicting the earlier belief they were exclusively (or almost exclusively) carnivorous.[123][126] A dental analysis of specimens from Spy, Belgium and El Sidrón, Spain by an international team, published[127] in 2017 in Nature revealed, that these Neanderthals "had a wide-ranging diet", even claiming the "Neanderthals from El Sidrón showed no evidence of meat eating" at all while living on "a mixture of forest moss, pine nuts and a mushroom known as split gill".[128]

The size and distribution of Neanderthal sites, along with genetic evidence, suggests Neanderthals lived in much smaller and more sparsely distributed groups than their Homo sapiens contemporaries.[129][130] Some experts suggest that this disparity alone was a major contributing factor to their ultimate replacement by Homo sapiens, which may have outnumbered them by as much as 9 to 1 according to some estimates.[129] Their lower population density may have also increased Neanderthal susceptibility to mutations caused by inbreeding.[130]

The bones of twelve Neanderthals were discovered at El Sidrón cave in the Atapuerca Mountains of Spain. They are believed to be a social group which was massacred about 50,000 years ago, and analysis of the mtDNA shows that the three adult males belonged to the same maternal lineage, while the three adult females belonged to different ones. This suggests a social structure where males remained in the same social group and females "married" out.[131]

The bones of the El Sidron group show signs of defleshing, suggesting that they were victims of cannibalism, and as their bones also show signs that they suffered from food shortage, they may have been victims of "survival cannibalism" by another Neanderthal group.[131] The St. Césaire 1 skeleton discovered in 1979 at La Roche à Pierrot, France, showed a healed fracture on top of the skull apparently caused by a deep blade wound. Researchers have taken this as evidence of the presence of interpersonal violence among the Neanderthals.[132]

In 2016, Jacques Jaubert and colleagues reported the discovery of 176,000 year old stalagmite constructions 336 meters deep in Bruniquel Cave near Bruniquel in south-western France. There were two annular (ring) structures, one 6.7 by 4.5 metres, and the other 2.2 by 2.1, with one to four layers of aligned broken stalagmites, and short pieces within the layers to support them. There were also four stack structures, two within the large ring and two outside it. Artificial lighting would have been required as the cave is far beyond the reach of daylight, and 57 of the stalagmite pieces are reddened and 66 blackened by fire. Burnt organic material was also found. The researchers concluded that the constructions must have been made by early Neanderthals, as they were the only human species in the area at the time. The elaborate structures show that the Neanderthals had a more sophisticated level of social organisation than previously thought possible.[40]


Further information: Neanderthal genome project



In 1997, geneticists were able to extract a short sequence of DNA from Neanderthal bones.[133] The extraction of mtDNA from a second specimen was reported in 2000, and showed no sign of modern human descent from Neanderthals.[78]


Svante Pääbo has tested more than 70 Neanderthal specimens. The Neanderthal genome is almost the same size as the human genome and is identical to ours to a level of 99.7% by comparing the accurate order of the nitrogenous bases in the double nucleotide chain.[135] From mtDNA analysis estimates, the two species shared a common ancestor about 500,000 years ago. An article[136] appearing in the journal Nature has calculated the species diverged about 516,000 years ago, whereas fossil records show a time of about 400,000 years ago.[137] A 2007 study pushes the point of divergence back to around 800,000 years ago.[138]

Edward Rubin of the Lawrence Berkeley National Laboratory states recent genome testing of Neanderthals suggests human and Neanderthal DNA are some 99.5% to nearly 99.9% identical.[139][140]

Interbreeding with modern humans[edit]





Scientist at the Max Planck Institute for Evolutionary Anthropology extracting the DNA




It is suggested that 20 percent of Neanderthal DNA survived in modern humans, notably expressed in the skin, hair, and diseases of the modern people.[146] Modern human genes involved in making keratin—the protein found in skin, hair, and nails—have specially high levels of Neanderthal DNA.[146] For example, around 66% of East Asians contain the Neanderthal skin gene, while 70% of Europeans contain the Neanderthal gene which affects skin colour. POU2F3 is found in around 66 percent of East Asians, while the Neanderthal version of BNC2, which affects skin color, among other traits, is found in 70 percent of Europeans. Neanderthal are the variants in genes that affect the risk of several diseases, including lupus, biliary cirrhosis, Crohn's disease, and type 2 diabetes. 8% of Neanderthal DNA comes from unknown species, tantalizing hints of unknown groups from Asia and Africa that left genes in Denisovans and modern humans, respectively.[146] [147] 17% of the genes of Denisovians and Neanderthals are shared. According to a research led by an Oxford scientist, it is possible that the gene causing red hair in modern humans is inherited from the Neanderthals, which is up to 100,000 years old, although a different mutation on Neanderthals was found in the MRC1 gene than on modern humans.[148][149][150][151]





Studies published in March 2016 suggest that modern humans bred with hominins, including Neanderthals, on multiple occasions.[159] Another study in April 2016 found differences between Homo sapiens and Neanderthal Y chromosomes that, they postulated, could cause female Homo Sapiens to miscarry male babies that had Neanderthal fathers.[160] This could explain why no modern man had to date been found with a Neanderthal Y chromosome.[161] Melanesians and Australoid populations show evidence of only one interbreeding event, possibly ~100,000 years ago, occurring in the Middle East, Europeans show a second event, which may also be of Middle Eastern origin, occurring possibly 50,000 years ago, while East Asians show an additional third interbreeding event possibly 30,000 years ago occurring in Siberia. Evidence that Neanderthal genomic material is often found amongst genes of the immune system suggests that some of the interbreeding may have secured resistance to diseases that Neanderthal populations had bred resistance to.[159]




Extinction hypotheses[edit]

According to a 2014 study by Thomas Higham and colleagues of organic samples from European sites, Neanderthals died out in Europe between 41,000 and 39,000 years ago,[d] although other studies suggested around 24,000 years ago instead.[8][9][10][11] Nonetheless, Homo sapiens arrived in Mediterranean Europe between 45,000 and 43,000 years ago, so the two different human populations shared Europe for several thousand years.[165][166] The exact nature of biological and cultural interaction between Neanderthals and other human groups has been contested.[167]

Possible scenarios for the extinction of the Neanderthals are:

  1. Neanderthals were a separate species from modern humans, and became extinct (because of climate change or interaction with modern humans) and were replaced by modern humans moving into their habitat between 45,000 and 40,000 years ago.[168] Jared Diamond has suggested a scenario of violent conflict and displacement.[169]
  3. Volcanic catastrophe: see Campanian Ignimbrite Eruption


Climate change[edit]



Coexistence with Homo sapiens[edit]


In November 2011 tests conducted at the Oxford Radiocarbon Accelerator Unit in England on what were previously thought to be Neanderthal baby teeth, which had been unearthed in 1964 from the Grotta del Cavallo in Italy, were identified as the oldest modern human remains discovered anywhere in Europe, dating from between 43,000 and 45,000 years ago.[174] Given that the 2014 study by Thomas Higham of Neanderthal bones and tools indicates that Neanderthals died out in Europe between 41,000 and 39,000 years ago, the two different human populations shared Europe for as long as 5,000 years.[87] Other studies, however, suggest the Neanderthals became extinct around 24,000 years ago instead.[8][9][10][11] Nonetheless, the exact nature of biological and cultural interaction between Neanderthals and other human groups has been contested.[167]



Interbreeding hypotheses[edit]



Hans Peder Steensby, while strongly emphasising that all modern humans are of mixed origins, proposed the interbreeding hypothesis in 1907, in the article Race studies in Denmark.[178] He held that this would best fit current observations, and attacked the widespread idea that Neanderthals were ape-like or inferior.

The most vocal proponent of the hybridization hypothesis is Erik Trinkaus of Washington University.[179] Trinkaus claims various fossils as products of hybridized populations, including the child of Lagar Velho, a skeleton found at Lagar Velho in Portugal.[180][181] In a 2006 publication co-authored by Trinkaus, the fossils found in 1952 in the cave of Peștera Muierii, Romania, are likewise claimed as descendants of previously hybridized populations.[182]

Genetic research has asserted that some admixture took place.[183] The genomes of all non-Africans include portions that are of Neanderthal origin,[184][185] due to interbreeding between Neanderthals and the ancestors of Eurasians in Northern Africa or the Middle East prior to their spread. Rather than absorption of the Neanderthal population, this gene flow appears to have been of limited duration and limited extent. An estimated 1 to 4 percent of the DNA in Europeans and Asians (French, Chinese and Papua probands) is non-modern, and shared with ancient Neanderthal DNA rather than with Sub-Saharan Africans (Yoruba people and San probands).[81] Ötzi the iceman, Europe's oldest preserved mummy, was found to possess an even higher percentage of Neanderthal ancestry.[186] 2014 findings suggest there may be even more Neanderthal genes in non-African humans than previously expected: approximately 20% of the Neanderthal gene pool was present in a broad sampling of non-African individuals, though each individual's genome was on average only 2% Neanderthal.[187]

2012 genetic studies seem to suggest that modern humans may have mated with "at least two groups" of ancient humans: Neanderthals and Denisovans.[188] Some researchers suggest admixture of 3.4–7.9% in modern humans of non-African ancestry, rejecting the hypothesis of ancestral population structure.[189] Detractors have argued and continue to argue that the signal of Neanderthal interbreeding may be due to ancient African substructure, meaning that the similarity is only a remnant of a common ancestor of both Neanderthals and modern humans and not the result of interbreeding.[154][190] John D. Hawks has argued that the genetic similarity to Neanderthals may indeed be the result of both structure and interbreeding, as opposed to just one or the other.[191]

While some modern human nuclear DNA has been linked to the extinct Neanderthals, no mitochondrial DNA of Neanderthal origin has been detected,[75] which in primates is always maternally transmitted. This observation has prompted the hypothesis that whereas female humans interbreeding with male Neanderthals were able to generate fertile offspring, the progeny of female Neanderthals who mated with male humans were either rare, absent or sterile.[192] However, some researchers have argued that there is evidence of possible interbreeding between female Neanderthals and male modern humans.[162][193][194]


Notable specimens[edit]

Type Specimen, Neanderthal 1
  • La Chapelle-aux-Saints 1: Called the Old Man, a fossilized skull discovered in La Chapelle-aux-Saints, France, by A. and J. Bouyssonie, and L. Bardon in 1908. Characteristics include a low vaulted cranium and large browridge typical of Neanderthals. Estimated to be about 60,000 years old, the specimen was severely arthritic and had lost all his teeth, with evidence of healing. For him to have lived on suggests that someone might have had to process his food for him. This is one possible example of Neanderthal altruism (see also Shanidar I).
A cast of the Ferrassie 1 skull
  • Shanidar Cave: Found in the Zagros Mountains in Iraqi Kurdistan; a total of nine skeletons found believed to have lived in the Middle Paleolithic. One of the nine remains was missing part of its right arm, which is theorized to have been broken off or amputated. The find is also significant because it shows that stone tools were present among this tribe's culture. One of the skeletons was originally thought to have been buried with flowers, signifying that some type of burial ceremony may have occurred. This is no longer considered to be the case, and Paul B. Pettitt has stated that the "deliberate placement of flowers has now been convincingly eliminated", noting that "A recent examination of the microfauna from the strata into which the grave was cut suggests that the pollen was deposited by the burrowing rodent Meriones tersicus, which is common in the Shanidar microfauna and whose burrowing activity can be observed today".[195]


This section describes bones with Neanderthal traits in chronological order.

Mixed with H. heidelbergensis traits[edit]

Typical H. neanderthalensis traits[edit]

Homo sapiens with some neanderthal-like archaic traits[edit]

Popular culture[edit]


Early artistic reconstructions mostly presented Neanderthals as beastly creatures, emphasizing hairiness and rough, dark complexion.[209]

See also[edit]





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Further reading[edit]

  • Sankararaman, Sriram; Mallick, Swapan; Dannemann, Michael; Prüfer, Kay; Kelso, Janet; Patterson, Nick; Reich, David (2014). "The genomic landscape of Neanderthal ancestry in present-day humans". Nature. 507 (7492): 354–357. doi:10.1038/nature12961. 
  • Vattathil, S.; Akey, J.M. (2015). "Small amounts of archaic admixture provide big insights into human history". Cell. 163 (2): 281–284. doi:10.1016/j.cell.2015.09.042. PMID 26451479. 

External links[edit]