From Wikipedia, the free encyclopedia
Jump to: navigation, search
Temporal range: MiddleLate Pleistocene 0.250–0.040 Ma
Homo sapiens neanderthalensis.jpg
The Neanderthal skull of La Chapelle-aux-Saints 1
An approximate reconstruction of a Neanderthal skeleton at the 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 or Homo sapiens neanderthalensis
Binomial name
Homo neanderthalensis or Homo sapiens neanderthalensis
King, 1864
Range of Homo neanderthalensis.png
Range of Homo neanderthalensis. Eastern and northern ranges may extend to include Okladnikov in Altai and Mamotnaia in Ural

Homo mousteriensis[1]
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 went extinct about 40,000 years ago.[8][9][10][11][12][13] Neanderthals and modern humans share 99.7% of their DNA[14] and are hence closely related.[15][16] (By comparison, both modern humans and Neanderthals share 98.8% of their DNA with their closest non-human living relatives, the chimpanzees.)[14] Neanderthals left bones and stone tools in Eurasia, from Western Europe to Central and Northern Asia. Fossil evidence suggests Neanderthals evolved in Europe, separate from modern humans in Africa for more than 400,000 years. They are classified either as a distinct species, Homo neanderthalensis,[17][18][19] or as a subspecies of Homo sapiens (H. s. neanderthalensis).[20][21]

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.[22] Late Mousterian artifacts were found in Gorham's Cave on the south-facing coast of Gibraltar.[23][24]

Compared to modern humans, 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 1,600 cm3 (98 cu in), females 1,300 cm3 (79 cu in),[25][26] extending to 1,736 cm3 (105.9 cu in) in Amud 1.[27] This is notably larger than the 1,250 to 1,400 cm3 (76 to 85 cu in) typical of modern humans. Males stood 164 to 168 cm (65 to 66 in) and females 152 to 156 cm (60 to 61 in) tall.[28]

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.[29][30][31][32] Recent studies also show that some Neanderthals mated with ancestors of modern humans long before the "out of Africa migration" of present day non-Africans, as early as 100,000 years ago.[33] 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.[34]

Claims that Neanderthals deliberately buried their dead, and if they did, whether such burials had any symbolic meaning,[35]:158–160 are heavily contested.[36][37][38] 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,[39] and this has in turn been heavily criticised.[40]

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.[41][42][43] 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.[44]


Neanderthals are named after one of the first sites where their 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][46][47]

Neanderthal 1 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".[48] The binomial name Homo neanderthalensis – extending the name "Neanderthal man" from the individual type specimen to the entire group – was first proposed by the Anglo-Irish geologist William King in 1864, although that same year King changed his mind and thought that the Neanderthal fossil was distinct enough from humans to warrant a separate genus.[4] Nevertheless, King's name had priority over the proposal put forward in 1866 by Ernst Haeckel, Homo stupidus.[46] The practice of referring to "the Neanderthals" and "a Neanderthal" emerged in the popular literature of the 1920s.[49]

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/).[50][51][52] 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/),[53] although scientists typically use the /t/ as in German.[54][55]


Scientists still dispute whether Neanderthals should be classified as a distinct species - Homo neanderthalensis - or as Homo sapiens neanderthalensis, placing Neanderthals as a subspecies of H. sapiens.[56][57] 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.[58] Senior members of their respective national paleontological institutes and among the most eminent paleoanthropologists of their time,[59][60] 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.[61]

During the 1930s scholars Ernst Mayr, George Gaylord Simpson and Theodosius Dobzhansky reinterpreted the existing fossil record and came to different conclusions.[62] 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.[63]

DNA researcher Svante Pääbo referred to the ongoing "taxonomic wars" over whether Neanderthals were a separate species or subspecies as the type of debate that cannot be resolved, "since there is no definition of species perfectly describing the case."[64]


Both Neanderthals and living humans are thought to have evolved from Homo erectus. In the earliest known migration wave into Eurasia dated to 1.81 million years ago (Ma), Homo erectus left Africa most probably via the Levant and reached Georgia (fossils of Dmanisi). Hominins had reached China by 1.7 Ma[65] and Spain by 1.4 Ma.[66] The discoverers of fragmented bones in Spain dated to 1.2 million years, assigned to a new species Homo antecessor, argue these are the remains of the ancestors of Neanderthals and of the older species Homo heidelbergensis, an interpretation rejected by most anthropologists.[67]

A large number of molecular clock genetic studies place the divergence time of the Neanderthal and modern human lineages between 800,000 and 400,000 years ago.[68][69][70][71][72][73][74][75][76] For this reason, most scholars believe Neanderthals descend, via Homo heidelbergensis, from another Homo erectus migration out of Africa that would have occurred in this time frame. Parts of the Homo erectus population that stayed in Africa would have evolved, perhaps through the intermediate Homo rhodesiensis, into early anatomically modern humans by 200,000 years ago.

Neanderthal traits are present in Homo heidelbergensis specimens beginning between 600,000 and 350,000 years ago.[77][78][79][80] Conventionally, European hominins are called Neanderthals from c. 250,000 years ago.[81] Because the quality of the fossil record greatly increases from 130,000 years ago onwards,[35] specimens younger than this date make up the bulk of known Neanderthal skeletons and were the first whose anatomy was comprehensively studied. They are known as Classic Neanderthals.


Neanderthal excavation site (former Kleine Feldhofer Grotte) in Neander Valley, and its position in North Rhine-Westphalia, Germany

Neanderthal fossils were first discovered in 1829 in the Engis caves (the partial skull dubbed Engis 2), in what is now Belgium by Philippe-Charles Schmerling and the Gibraltar 1 skull in 1848 in the Forbes' Quarry, Gibraltar, both prior to the type specimen discovery in a limestone quarry in the Neandertal in Germany in August 1856, three years before Charles Darwin's On the Origin of Species was published.[82]

The type specimen, dubbed Neanderthal 1, consisted of a skull cap, two femora, three bones of the right arm, two of the left arm, parts of the left ilium, fragments of a scapula, and ribs. The workers who recovered the objects originally thought them to be the remains of a cave bear. However, they eventually gave the material to amateur naturalist Johann Carl Fuhlrott, who turned the fossils over to anatomist Hermann Schaaffhausen.

To date, the bones of over 400 Neanderthals have been found.[83]

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.
  • 1848: Neanderthal skull Gibraltar 1 found in Forbes' Quarry, Gibraltar. Called "an ancient human" at the time.
  • 1856: Johann Karl Fuhlrott first recognized the fossil called "Neanderthal 1", discovered in the Neandertal, a valley near Mettmann in what is now North Rhine-Westphalia, Germany.
  • 1864: William King proposed the name Homo neanderthalensis at a meeting of the British Association for the Advancement of Science, but then changed his mind and argued that Neanderthals were different enough from humans to warrant a separate genus, under the assumption that they were likely "incapable of moral and theositic conceptions."[4]
  • 1880: The mandible of a Neanderthal child was found in a secure context and associated with cultural debris, including hearths, Mousterian tools, and bones of extinct animals.
  • 1886: Two nearly perfect skeletons of a man and woman were found at Spy, Belgium at the depth of 16 ft with numerous Mousterian-type implements.
  • 1899: Hundreds of Neanderthal bones were described in stratigraphic position in association with cultural remains and extinct animal bones.
  • 1899: Sand excavation workers found bone fragments on a hill in Krapina, Croatia called Hušnjakovo brdo. Local Franciscan friar Dominik Antolković requested Dragutin Gorjanović-Kramberger to study the remains of bones and teeth that were found there.
  • 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.[84]
  • 1925: Francis Turville-Petre finds the 'Galilee Man' or 'Galilee Skull' in the Zuttiyeh Cave in Wadi Amud in The British Mandate of Palestine (now Israel).
  • 1926: Skull fragments of Gibraltar 2, a four-year-old Neanderthal girl, discovered by Dorothy Garrod.
  • 1953–1957: Ralph Solecki uncovered nine Neanderthal skeletons in Shanidar Cave in the Kurdistan region of northern Iraq.
  • 1975: Erik Trinkaus' study of Neanderthal feet confirmed they walked like modern humans.
  • 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).
  • 1991: ESR dates showed the Tabun Neanderthal was contemporaneous with modern humans from Skhul and Qafzeh.
  • 1993: 127,000-year-old DNA is found on the child of Sclayn, found in Scladina (fr), Belgium.
  • 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.[85][86]
  • 1997: Matthias Krings et al. are the first to amplify Neanderthal mitochondrial DNA (mtDNA) using a specimen from Feldhofer grotto in the Neander valley.[87]
  • 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.[88]
  • 1998: A team led by pre-history archeologist João Zilhão discovered an early Upper Paleolithic human burial in Portugal, at Abrigo do Lagar Velho, which provided evidence of early modern humans from the west of the Iberian Peninsula. The remains, a largely complete skeleton of an approximately 4-year-old child, buried with pierced shell and red ochre, is dated to ca. 24,500 years BP.[89] The cranium, mandible, dentition, and postcrania present a mosaic of European early modern human and more archaic features reminiscent of Neanderthals.[89]
  • 2000: Igor Ovchinnikov et al. retrieved mitochondrial DNA of a Neanderthal infant from Mezmaiskaya Cave in the Caucasus.[90]
  • 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.[91] In 2009, the Max Planck Institute announced the "first draft" of a complete Neanderthal genome is completed.[92]
  • 2010: Comparison of Neanderthal genome with modern humans from Africa and Eurasia shows that 1–4% of modern non-African human genome might come from the Neanderthals.[30][93]
  • 2010: Discovery of Neanderthal tools far away from the influence of anatomically modern humans indicate that Neanderthals might have been able to create and evolve tools on its own, and therefore be more intelligent than previously thought. Furthermore, it was proposed that the Neanderthals might be more closely related to modern humans than previously thought and may in fact be a subspecies of Homo sapiens.[94] Evidence has more recently emerged that these artifacts are probably of H. sapiens sapiens origin.[95]
  • 2012: Charcoal found next to six paintings of seals in Nerja caves, Malaga, Spain, has been dated to between 42,300 and 43,500 years old. The paintings themselves will be dated in 2013, and if their pigment matches the date of the charcoal, they would be the oldest known cave paintings. José Luis Sanchidrián at the University of Cordoba, Spain believes the paintings are more likely to have been painted by Neanderthals than early modern humans.[96]
  • 2013: A jawbone found in Italy had features intermediate between Neanderthals and anatomically modern humans, suggesting it could be a hybrid. The mitochondrial DNA is Neanderthal.[97]
  • 2013: In the midst of a hundred-year old debate over the existence of Neanderthal deliberate burials, a team of French researchers reasserted a heavily contested 1908 claim that the Chapelle-aux-Saints 1 skeleton was deliberately buried.[39]
  • 2014: Some researchers express worry that almost all theories for the Neanderthals' extinction assume that there is something modern humans had that Neanderthals did not.[98]
  • 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 anatomically-modern Homo sapiens reached the continent.[13][99]

Habitat and range[edit]

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 early Neanderthals. 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,[100] 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.[101]

Neanderthal fossils have not been found to date in Africa, but there have been finds close to North Africa, both on Gibraltar and in the Levant. At some Levantine sites, Neanderthal remains date from after the same sites were vacated by modern humans. Mammal fossils of the same time period show cold-adapted animals were present alongside these Neanderthals in this region of the Eastern Mediterranean. This implies Neanderthals were better adapted biologically to cold weather than modern humans and at times displaced them in parts of the Middle East when the climate got cold enough. However this has been disputed recently through studies of their cranial morphology and sinuses. Some scholars have also posited that Neanderthals could have arisen in Asia, then expanded into Europe, making Neanderthals of tropical origin rather than cold adaptive.[102][103]

Anatomically modern humans appear to have been the only human type in the Nile River Valley during these periods, and Neanderthals are not known to have ever lived south-west of present-day Israel. When climate change caused warmer temperatures, the Neanderthal range likewise retreated to the north, along with the cold-adapted species of mammals. Apparently these weather-induced population shifts took place before modern people secured competitive advantages over the Neanderthal, as these shifts in range took place well over ten thousand years before modern people totally replaced the Neanderthal, despite the recent evidence of some successful interbreeding.[102]

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.

To date, no intimate connection has been found between these similar archaic people and the Eurasian Neanderthals, at least during the same time as H. rhodesiensis seems to have lived about 600,000 years ago, long before the time of classic Neanderthals. This said, some researchers think that H. rhodesiensis may have lived much later than this period, depending on the method used to date the fossils, so this question remains open to debate. Some H. rhodesiensis features, like the large brow ridge, may have been caused by convergent evolution.

It appears incorrect, based on present research and known fossil finds, to refer to any fossils outside Eurasia as true Neanderthals. They had a known range that possibly extended as far east as the Altai Mountains, but not farther to the east or south, and apparently not into Africa. At any rate, in North-East Africa the land immediately south of the Neanderthal range was possessed by modern humans Homo sapiens idaltu or Homo sapiens, since at least 160,000 years before the present. 160,000-year-old hominid fossils at Jebel Irhoud in Morocco were previously thought to be Neanderthal, but it is now clear that they are early modern humans.[104]

Classic Neanderthal fossils have been found over a large area, from northern Germany to Israel and Mediterranean countries like Spain[105] and Italy[106] 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.[107] Recent evidence has extended the Neanderthal range by about 1,250 miles (2,010 km) east into southern Siberia's Altai Mountains.[108][109]


Artist interpretation of the head of the Shanidar 1 fossil, a Neanderthal male who lived c. 70,000 years ago (John Gurche 2010)

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,[110] 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.[81] Evidence suggests they were much stronger than modern humans, with particularly strong arms and hands,[111][112] while they were comparable in height; based on 45 long bones from at most 14 males and 7 females, Neanderthal males averaged 164 to 168 cm (65 to 66 in) and females 152 to 156 cm (60 to 61 in) tall.[28] 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.[113] A 2007 genetic study suggested some Neanderthals may have had red hair and blond hair, along with a light skin tone.[114]

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 anatomically modern humans. 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 mothers' fur as infants. Modern humans don’t have these ridges, but Neanderthals do." [115][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.[116]

A 2013 study of Neanderthal skulls suggests that their eyesight may have been better than that of modern humans, owing to larger eye sockets and larger areas of the brain devoted to vision.[117]

Neanderthals are known for their large cranial capacity, which at 1,600 cm3 (98 cu in) is larger on average than that of modern humans. One study has found that Neanderthal brains were more asymmetric than other hominid brains.[118] 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.[119] They had almost the same degree of encephalization (i.e. brain to body size ratio) as modern humans.[120][121]


Neanderthals made stone tools,[122] used fire,[123] and were hunters. The consensus on their behaviour ends here.

It had actually long been debated whether Neanderthals were hunters or scavengers.[81][further explanation needed] Most available evidence suggests they were apex predators,[124][125] and fed on red deer, reindeer, ibex, wild boar, aurochs and on occasion mammoth, straight-tusked elephant and rhinoceros.[81][126] However, while they were largely carnivorous,[127][128] new studies suggest Neanderthals occasionally used vegetables as fall-back food.[125][129] In 2010, an isotope analysis of Neanderthal teeth found traces of cooked vegetable matter, and a 2014 study of Neanderthal coprolites (fossilized feces) found substantial amounts of plant matter.[127][130] Dental analysis of specimens from Spy, Belgium and El Sidrón, Spain [131] in 2017 argued these Neanderthals had a wide-ranging diet, and that those "from El Sidrón showed no evidence of meat eating" at all and seemed to have lived on "a mixture of forest moss, pine nuts and a mushroom known as split gill".[132] The size and distribution of Neanderthal sites, along with genetic evidence, suggests Neanderthals lived in much smaller and more sparsely distributed groups than anatomically-modern Homo sapiens.[133][134] Some experts suggest that this disparity alone was a major contributing factor to their ultimate replacement by Homo sapiens sapiens, which may have outnumbered them by as much as 9 to 1 according to some estimates.[133] Their lower population density may have also increased Neanderthal susceptibility to mutations caused by inbreeding.[134]

The bones of twelve Neanderthals were discovered at El Sidrón cave in the Atapuerca Mountains of Spain. They are believed to have been a group killed and butchered about 50,000 years ago. Analysis of the mtDNA showed 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.[135]

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.[135] 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.[136]

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.[137][138]

Circumstantial evidence suggests Neanderthals may have been building some form of watercraft since the Middle Paleolithic.[139][140][unbalanced opinion] Quartz hand-axes, three-sided picks, and stone cleavers from Crete are controversially claimed to have been recovered on the island in layers dating to 170,000 BP.[141]

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 m (22 by 15 ft), and the other 2.2 by 2.1 m (7.2 by 6.9 ft), 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 controversially concluded that the constructions must have been made by early Neanderthals, as they were the only archaic humans in the area at the time.[44][unbalanced opinion]



Early investigations concentrated on mitochondrial DNA (mtDNA), which, owing to strictly matrilineal inheritance and subsequent vulnerability to genetic drift, is of limited value in evaluating the possibility of interbreeding of Neanderthals with Cro-Magnon people.

In 1997, geneticists were able to extract a short sequence of DNA from Neanderthal bones.[142] The extraction of mtDNA from a second specimen was reported in 2000, and showed no sign of modern human descent from Neanderthals.[90]

In July 2006, the Max Planck Institute for Evolutionary Anthropology and 454 Life Sciences announced that they would sequence the Neanderthal genome over the next two years. This genome was expected to be roughly the size of the human genome, three-billion base pairs, and share most of its genes. It was hoped the comparison would expand understanding of Neanderthals, as well as the evolution of humans and human brains.[143]

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.[144] From mtDNA analysis estimates, the two shared a common ancestor about 500,000 years ago. An article[145] appearing in the journal Nature has calculated they diverged about 516,000 years ago, whereas fossil records show a time of about 400,000 years ago.[146] A 2007 study pushes the point of divergence back to around 800,000 years ago.[147]

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.[148][149]

Interbreeding with modern humans[edit]

On November 16, 2006, Lawrence Berkeley National Laboratory issued a press release suggesting Neanderthals and ancient humans probably did not interbreed.[150] Edward M. Rubin, director of the U.S. Department of Energy's Lawrence Berkeley National Laboratory and the Joint Genome Institute (JGI), sequenced a fraction (0.00002) of genomic nuclear DNA (nDNA) from a 38,000-year-old Vindia Neanderthal femur. They calculated the common ancestor to be about 353,000 years ago, and a complete separation of the ancestors of the groups about 188,000 years ago.[151]

Their results show the genomes of modern humans and Neanderthals are at least 99.5% identical, but despite this genetic similarity, and despite the two groups having coexisted in the same geographic region for thousands of years, Rubin and his team did not find any evidence of any significant interbreeding between the two. Rubin said, "While unable to definitively conclude that interbreeding between the two species of humans did not occur, analysis of the nuclear DNA from the Neanderthal suggests the low likelihood of it having occurred at any appreciable level."[151]

In 2008 Richard E. Green et al. from Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany, published the full sequence of Neanderthal mitochondrial DNA (mtDNA) and suggested "Neanderthals had a long-term effective population size smaller than that of modern humans."[152]

In the same publication, it was disclosed by Svante Pääbo that in the previous work at the Max Planck Institute, "Contamination was indeed an issue," and they eventually realized that 11% of their sample was modern human DNA.[153][154] Since then, more of the preparation work has been done in clean areas and 4-base pair 'tags' have been added to the DNA as soon as it is extracted so the Neanderthal DNA can be identified.

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

With 3 billion nucleotides sequenced, analysis of about ⅓ showed no sign of admixture between modern humans and Neanderthals, according to Pääbo. This concurred with the work of Noonan from two years earlier. The variant of microcephalin common outside Africa, which was suggested to be of Neanderthal origin and responsible for rapid brain growth in humans, was not found in Neanderthals. Nor was the MAPT variant, a very old variant found primarily in Europeans.[153]

However, an analysis of a first draft of the Neanderthal genome by the same team released in May 2010 indicates interbreeding may have occurred.[30][93] "Those of us who live outside Africa carry a little Neanderthal DNA in us," said Pääbo, who led the study. "The proportion of Neanderthal-inherited genetic material is about 1 to 4 percent. It is a small but very real proportion of ancestry in non-Africans today," says Dr. David Reich of Harvard Medical School, who worked on the study. This research compared the genome of the Neanderthals to five modern humans from China, France, sub-Saharan Africa, and Papua New Guinea. The finding is that about 1 to 4 percent of the genes of the humans outside sub-Saharan Africa came from Neanderthals, compared to the baseline defined by the two sub-Saharan Africans.[93]

This indicates a gene flow from Neanderthals to modern humans, i.e., interbreeding between the two populations. Since the three non-African genomes show a similar proportion of Neanderthal sequences, the interbreeding must have occurred early in the migration of modern humans out of Africa, perhaps in the Middle East. No evidence for gene flow in the direction from modern humans to Neanderthals was found. Gene flow from modern humans to Neanderthals would not be expected if contact occurred between a small colonizing population of modern humans and a much larger resident population of Neanderthals. A very limited amount of interbreeding could explain the findings, if it occurred early enough in the colonization process.[93]

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.[155] Modern human genes involved in making keratin—the protein found in skin, hair, and nails—have specially high levels of Neanderthal DNA.[155] 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 an unknown group of archaic humans, tantalizing hints of unknown groups from Asia and Africa that left genes in Denisovans and modern humans, respectively.[155] [156] The genetic variant of the MC1R gene linked to red hair in Neanderthals has not been found in modern humans, hence red hair may be an example of convergent evolution.[157][158][159]

While interbreeding is viewed as the most parsimonious interpretation of the genetic discoveries, the authors point out they cannot conclusively rule out an alternative scenario, in which the source population of non-African modern humans was already more closely related to Neanderthals than other Africans were, because of ancient genetic divisions within Africa.[93] Other studies carried out since the sequencing of the Neanderthal genome have cast doubt on the level of admixture between Neanderthals and modern humans, or even as to whether the groups interbred at all. One study has asserted that the presence of Neanderthal or other archaic human genetic markers can be attributed to shared ancestral traits between the lineages originating from a 500,000-year-old common ancestor.[160][161][162] [163]

Among the genes shown to differ between present-day humans and Neanderthals were RPTN, SPAG17, CAN15, TTF1, FOXP2 and PCD16.[93]

More recent research suggests that Neanderthal–Homo sapiens sapiens interbreeding appears to have occurred asymmetrically among the ancestors of modern-day humans, and that this is a possible rationale for differing frequencies of Neanderthal-specific DNA in the genomes of modern humans. In 2015, researchers Benjamin Vernot and Joshua Akey at the University of Washington conclude in a paper in the American Journal of Human Genetics that the relatively greater quantity of Neanderthal-specific DNA in the genomes of individuals of East Asian descent (than those of European descent) cannot be explained by differences in selection.[164] They further suggest that "two additional demographic models, involving either a second pulse of Neandertal gene flow into the ancestors of East Asians or a dilution of Neandertal lineages in Europeans by admixture with an unknown ancestral population" are parsimonious with their data.[164] Similar conclusions were reached in a paper published in the same publication by researchers Bernard Kim and Kirk Lohmueller at UCLA: "Using simulations of a broad range of models of selection and demography, we have shown that this hypothesis [that the greater proportion of Neandertal ancestry in East Asians than in Europeans is due to the fact that purifying selection is less effective at removing weakly deleterious Neandertal alleles from East Asian populations] cannot account for the higher proportion of Neandertal ancestry in East Asians than in Europeans. Instead, more complex demographic scenarios, most likely involving multiple pulses of Neandertal admixture, are required to explain the data."[165]

In a subsequent interview, Dr. Lohmueller did note that these findings go against the commonly-held perception that Neanderthals were mostly localized to modern-day Europe and western Asia: "It’s very hard to put these findings into spatial context. The key idea is that there would have to have been some additional interbreeding events involving East Asians, but not Europeans. These interbreeding events could have been directly between Neanderthals and East Asians, maybe in some other indirect way."[166] Vernot also noted that "[H]umans have been constantly migrating throughout their history - this makes it hard to say exactly where interactions with Neanderthals occurred. It's possible, for example, that all of the interbreeding with Neanderthals occurred in the Middle East, before the ancestors of modern non-Africans spread out across Eurasia. In the model from the paper, the ancestors of all non-Africans interbred with Neanderthals, and then split up into multiple groups that would later become Europeans, East Asians. Shortly after they split up, the ancestors of East Asians interbred with Neanderthals just a little bit more."[166]

Studies published in March 2016 suggest that modern humans bred with hominins, including Neanderthals, on multiple occasions.[167] Another study in April 2016 found differences between modern human and Neanderthal Y chromosomes that, they postulated, could cause female Homo sapiens sapiens to miscarry male babies that had Neanderthal fathers.[168] This could explain why no modern man had to date been found with a Neanderthal Y chromosome.[169] 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.[167]

In 2016 researchers reported that they had found Human DNA in the genome of a female Neanderthal from the Altai mountains region near the border between Mongolia and Russia. They calculated that the mating must have taken place about 100,000 years ago.[170]


In April 2014, a first glimpse into the epigenetics of the Neanderthal was obtained with the publication of the full DNA methylation of the Neanderthal and the Denisovan.[171][172] The reconstructed DNA methylation map allowed researchers to assess gene activity levels throughout the Neanderthal genome and compare them to modern humans. One of the major findings focused on the limb morphology of Neanderthals. Gokhman et al. found that changes in the activity levels of the HOX cluster of genes were behind many of the morphological differences between Neanderthals and modern humans, including shorter limbs, curved bones and more.[172]


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] New dating in Iberia, where Neanderthal dates as late as 28,000 years had been reported, suggests evidence of Neanderthal survival in the peninsula after 42,000 years ago is almost non-existent.[11]

Anatomicaly modern humans arrived in Mediterranean Europe between 45,000 and 43,000 years ago, so the two different human populations shared Europe for several thousand years.[173][174] The exact nature of biological and cultural interaction between Neanderthals and other human groups is contested.[175]

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.[176] Jared Diamond has suggested a scenario of violent conflict and displacement.[177]
  2. Neanderthals were a contemporary subspecies that bred with modern humans and disappeared through absorption (interbreeding theory).
  3. Volcanic catastrophe: see Campanian Ignimbrite Eruption
mtDNA-based simulation of modern human expansion in Europe starting 1600 generations ago. Neanderthal range in light grey[178]

As Paul Jordan notes: "A natural sympathy for the underdog and the disadvantaged lends a sad poignancy to the fate of the Neanderthal folk, however it came about." Jordan, though, does say that there was perhaps interbreeding to some extent, but that populations that remained totally Neanderthal were probably out-competed and marginalized to extinction by the Aurignacians.[102]

Climate change[edit]

About 55,000 years ago, the weather began to fluctuate wildly from extreme cold conditions to mild cold and back in a matter of a few decades. Neanderthal bodies were well suited for survival in a cold climate—their barrel chests and stocky limbs stored body heat better than the Cro-Magnons. However, the rapid fluctuations of weather caused ecological changes to which the Neanderthals could not adapt; familiar plants and animals would be replaced by completely different ones within a lifetime. Neanderthals' ambush techniques would have failed as grasslands replaced trees. Neanderthals died out in Europe between 41,000 and 39,000 years ago, which may coincide with the start of a very cold period.[13][179] Raw material sourcing and the examination of faunal remains by Adler et al. (2006) in the southern Caucasus region suggest that modern humans may have had a survival advantage during this period, being able to use social networks to acquire resources from a greater area. They found that in both the Late Middle Palaeolithic and Early Upper Palaeolithic more than 95% of stone artifacts were drawn from local material, suggesting Neanderthals were restricted to more local resources. Furthermore, excavations at Ortvale Klde Rockshelter discovered that there was a clear break between the Late Middle Paleolithic and the Early Upper Paleolithic lithic assemblages, which were attributed to Neanderthals and modern humans respectively. This would suggest that modern humans came in and replaced Neanderthals, rather than a slow shift or integration occurring in this region. [180]

Studies on Neanderthal body structures have shown that they needed more energy to survive than any other types of hominid. Their energy needs were up to 100 to 350 kcal (420 to 1,460 kJ) more per day comparing to projected anatomically modern human males weighing 68.5 kg (151 lb) and females 59.2 kg (131 lb).[181] When food became scarce, this difference may have played a major role in the Neanderthals' extinction.[179]

Coexistence with Homo sapiens sapiens[edit]

Approximate reconstruction of a Neanderthal skeleton and artistic interpretation of the La Ferrassie 1 Neanderthal man from the National Museum of Nature and Science

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.[182] 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.[13] Nonetheless, the exact nature of biological and cultural interaction between Neanderthals and other human groups has been contested.[175]

Modern humans co-existed with them in Europe starting around 45,000 years ago and perhaps even earlier. Neanderthals inhabited that continent for a long period of time before the arrival of modern humans. These modern humans may have introduced a disease that contributed to the extinction of Neanderthals, and that may be added to other recent explanations for their extinction. When Neanderthal ancestors left Africa roughly 100,000 years earlier they adapted to the pathogens in their European environment, unlike modern humans who adapted to African pathogens. This transcontinental movement is known as the Out of Africa model. If contact between humans and Neanderthals occurred in Europe and Asia the first contact may have been devastating to the Neanderthal population, because they would have had little if any immunity to the African pathogens. More recent historical events in Eurasia and the Americas show a similar pattern, where the unintentional introduction of viral, or bacterial pathogens to unprepared populations has led to mass mortality and local population extinction.[183] The most well-known example of this is the arrival of Christopher Columbus to the New World, which brought and introduced foreign diseases when he and his crew arrived to a native population who had no immunity.

Anthropologist Pat Shipman, of Pennsylvania State University, suggested that domestication of the dog could have played a role in Neanderthals' extinction.[184]

Interbreeding hypotheses[edit]

Chris Stringer's hypothesis of the family tree of genus Homo, published 2012 in Nature – the horizontal axis represents geographic location, and the vertical axis represents time in millions of years ago.[e]

An alternative to extinction is that Neanderthals were absorbed into the Cro-Magnon population by interbreeding. This would be counter to strict versions of the Recent African Origin, since it would imply that at least part of the genome of Europeans would descend from Neanderthals.

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.[186] 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.[187] Trinkaus claims various fossils as products of hybridized populations, including the child of Lagar Velho, a skeleton found at Lagar Velho in Portugal.[188][189] 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.[190]

Genetic research has asserted that some admixture took place.[191] The genomes of all non-Africans include portions that are of Neanderthal origin,[192][193] 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).[93] Ötzi the iceman, Europe's oldest preserved mummy, was found to possess an even higher percentage of Neanderthal ancestry.[194] 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.[195]

2012 genetic studies seem to suggest that modern humans may have mated with "at least two groups" of ancient humans: Neanderthals and Denisovans.[196] Some researchers suggest admixture of 3.4–7.9% in modern humans of non-African ancestry, rejecting the hypothesis of ancestral population structure.[197] 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.[162][198] 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.[199]

While some modern human nuclear DNA has been linked to the extinct Neanderthals, no mitochondrial DNA of Neanderthal origin has been detected,[87] 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.[200] However, some researchers have argued that there is evidence of possible interbreeding between female Neanderthals and male modern humans.[170][201][202]


Notable specimens[edit]

Type Specimen, Neanderthal 1
  • Neanderthal 1: The first Neanderthal specimen found during mining in August 1856. It was discovered in a limestone quarry at the Feldhofer grotto in Neanderthal, Germany. The find consisted of a skull cap, two femora, three right arm bones, two left arm bones, ilium, and fragments of a scapula and ribs.
  • 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
  • La Ferrassie 1: A fossilized skull discovered in La Ferrassie, France, by R. Capitan in 1909. It is estimated to be 70,000 years old. Its characteristics include a large occipital bun, low-vaulted cranium and heavily worn teeth.
  • Le Moustier: A fossilized skull, discovered in 1909, at the archaeological site in Peyzac-le-Moustier, Dordogne, France. The Mousterian tool culture is named after Le Moustier. The skull, estimated to be less than 45,000 years old, includes a large nasal cavity and a somewhat less developed brow ridge and occipital bun as might be expected in a juvenile.
  • 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".[203]
  • Amud 1: Fossilized remains of an adult Neanderthal, dated to roughly 45,000 years ago, and one of several found in a cave at Nahal Amud, Israel, at least some of which may have been deliberately buried. A particularly notable feature of this find is its cranial capacity, which, at 1,740 cm3, is among the largest known for any hominid, living or extinct.[81][204]


This section describes bones with Neanderthal traits in chronological order.

Mixed with H. heidelbergensis traits[edit]

H. neanderthalensis fossils[edit]

Homo sapiens sapiens with archaic traits reminiscent of Neanderthals[edit]

Popular culture[edit]

Neanderthals have been portrayed in popular culture including appearances in literature, visual media and comedy. Early 20th century artistic interpretations often presented Neanderthals as beastly creatures, emphasizing hairiness and rough, dark complexion.[213]

See also[edit]


  1. ^ The common species name Neanderthal is on occasion written Neandertal, even in scientific publications, under the somewhat mistaken assumption that this common name is taken directly from the German and that it might hence have to follow spelling reforms of that language. (In German Thal, meaning valley, is written Tal since 1901.) In reality, the common species name Neanderthal comes from the binomial scientific name[3] established by King in 1864, Homo neanderthalensis.[4] The binomial name is indeed taken from German but because binomial names are normally unalterable, the binomial still reflects the pre-1901 German spelling and hence so does, for most authors, the common name. The Neandertal region in Germany is in English written without an h. Note that in German the common species name is almost always Neandertaler (litt. "of the valley of Neander") not Neandertal, but in the few instances where the word Neandertal is used to refer not to the place but to the prehistoric humans, as is the case of the Neanderthal Museum, the h is kept for the same reason as in English that it reflects the scientific name.[5]
  2. ^ The valley is named after Joachim Neander, whose Greek-style last name had been changed by his grandfather from "Neumann" ("new man").[45]
  3. ^ Some words beginning with th in older varieties of German were the result of a spelling embellishment that had no connection to English th. (Teil, meaning 'part,' was sometimes spelled Theil in the 18h and 19th centuries.) Tal became standardized with the German spelling reform of 1901, thus the German name Neandertal for both the valley and species/subspecies.
  4. ^ Higham et al did not study samples from sites outside Europe and they stated that further work was required to rule out later survival at Gorhams Cave, Gibraltar.[173]
  5. ^ Homo floresiensis originated in an unknown location from unknown ancestors and reached remote parts of Indonesia. Homo erectus spread from Africa to western Asia, then east Asia and Indonesia; its presence in Europe is uncertain, but it gave rise to Homo antecessor, found in Spain. Homo heidelbergensis originated from Homo erectus in an unknown location and dispersed across Africa, southern Asia and southern Europe (other scientists interpret fossils, here named heidelbergensis, as late erectus). Homo sapiens sapiens spread from Africa to western Asia and then to Europe and southern Asia, eventually reaching Australia and the Americas. In addition to Neanderthals and Denisovans, a third gene flow of archaic Africa origin is indicated at the right.[185]


  1. ^ Romeo, Luigi (1979). Ecce Homo!:A Lexicon of Man. Amsterdam: John Benjamins Publishing Company. p. 92. 
  2. ^ {cite book|last1=Camp|first1=C. L|last2=Allison|first2=H. J.|last3=Nichols|first3=R. H.|title=Bibliography of Fossil Vertebrates 1954-1958|publisher=The Geological Society of America, Inc.|location=New York|year=1964|page=556|url=}}
  3. ^ "Neanderthal". Wiley-Blackwell Encyclopedia of Human Evolution. Chichester, West Sussex: Wiley-Blackwell. 2013. 
  4. ^ a b c King, William (Jan 1864). "The Reputed Fossil Man of the Neanderthal" (PDF). The Quarterly Journal of Science. 1: 96. 
  5. ^ "Neandertal oder Neanderthal? - Was ist denn nun richtig?". Neanderthal museum. Retrieved February 1, 2017. Heute sollten Ortsbezeichnungen das „Neandertal“ ohne „h“ bezeichnen. Alle Namen, die sich auf den prähistorischen Menschen beziehen, führen das „h“. [Today one should write for place names "Neandertal" without an "h". All names related to the prehistoric humans keep the "h".] 
  6. ^ "Neanderthal in ODE". Oxford Dictionaries. 
  7. ^ ""Neanderthal" in Random House Dictionary (US) & Collins Dictionary (UK)". 
  8. ^ T. Higham, K. Douka, R. Wood, C.B. Ramsey, F. Brock, L. Basell, M. Camps, A. Arrizabalaga, J. Baena, C. Barroso-Ruíz, 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 (2014). "The timing and spatiotemporal patterning of Neanderthal disappearance". Nature. 512 (7514): 306–9. doi:10.1038/nature13621. PMID 25143113. 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. (subscription required)
  9. ^ T. Higham (2011). "European Middle and Upper Palaeolithic radiocarbon dates are often older than they look: problems with previous dates and some remedies". Antiquity. 85: 235–249. doi:10.1017/s0003598x00067570. Few events of European prehistory are more important than the transition from ancient to modern humans around 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. (subscription required)
  10. ^ a b R. Pinhasi, T.F.G. Higham, L.V. Golovanova, V.B. Doronichev (2011). "Revised age of late Neanderthal occupation and the end of the Middle Paleolithic in the northern Caucasus". Proceedings of the National Academy of Sciences USA. 108: 8611–8616. doi:10.1073/pnas.1018938108. The direct date of the fossil (39,700 ± 1,100 14C BP) is in good agreement with the probability distribution function, indicating at a high level of probability that Neanderthals did not survive at Mezmaiskaya Cave after 39 ka cal BP. [...] This challenges previous claims for late Neanderthal survival in the northern Caucasus. [...] Our results confirm the lack of reliably dated Neanderthal fossils younger than ~40 ka cal BP in any other region of Western Eurasia, including the Caucasus. (subscription required)
  11. ^ a b B. Galván, C.M. Hernández, C. Mallol, N. Mercier, A. Sistiaga, V. Soler (2014). "New evidence of early Neanderthal disappearance in the Iberian Peninsula". Journal of Human Evolution. 75: 16–27. doi:10.1016/j.jhevol.2014.06.002. (subscription required)
  12. ^ McKie, Robin (June 2, 2013). "Why did the Neanderthals die out?". The Guardian. Retrieved April 6, 2017. "It was once thought we appeared in Europe around 35,000 years ago and that we coexisted with Neanderthals for thousands of years after that. They may have hung on in pockets – including caves in Gibraltar – until 28,000 years ago [said Chris Stringer]" Previous research on Neanderthal sites which suggested that they were more recent than 40,000 years old appears to be wrong," said Stringer. "That is a key finding that will be discussed at the conference."[...] However, scientists have set out to get round these problems. At Oxford University, scientists led by Tom Higham have developed new methods to remove contamination and have been able to make much more precise radiocarbon dating for this period. 
  13. ^ a b c d "BBC News—New dates rewrite Neanderthal story". BBC News. 
  14. ^ a b Complete Neanderthal genome sequenced: DNA signatures found in present-day Europeans and Asians, but not in Africans, ScienceDaily
  15. ^ Colin P.T. Baillie; University of California, Berkeley. "Neandertals: Unique from Humans, or Uniquely Human?" (PDF). 
  16. ^ Smithsonian Institution National Museum of American History. "Ancient DNA and Neanderthals". 
  17. ^ Hublin, J. J. (2009). "The origin of Neandertals". Proceedings of the National Academy of Sciences. 106 (38): 16022–7. Bibcode:2009PNAS..10616022H. doi:10.1073/pnas.0904119106. JSTOR 40485013. PMC 2752594Freely accessible. PMID 19805257. 
  18. ^ Harvati, K.; Frost, S.R.; McNulty, K.P. (2004). "Neanderthal taxonomy reconsidered: implications of 3D primate models of intra- and interspecific differences". Proc. Natl. Acad. Sci. U.S.A. 101: 1147–52. doi:10.1073/pnas.0308085100. PMC 337021Freely accessible. PMID 14745010. 
  19. ^ "Scientists Identify Neanderthal Genes in Modern Human DNA". January 30, 2014. Retrieved October 29, 2015. 
  20. ^ Bodkin-Kowacki, Eva (March 14, 2017). "How a 400,000-year-old skull fragment hints at ancient 'unified humanity'". The Christian Science Monitor. Retrieved May 21, 2017. 
  21. ^ Whigham, Nick; AFP (March 15, 2017). "A 400,000-year-old skull fragment found in Portugal points to mystery people". Retrieved May 21, 2017. 
  22. ^ Shaw, Ian; Jameson, Robert, eds. (1999). A Dictionary of Archaeology. Blackwell. p. 408. ISBN 0-631-17423-0. Retrieved August 1, 2016. 
  23. ^ Finlayson, C; Pacheco, FG; Rodríguez-Vidal, J; Fa, DA; Gutierrez López, JM; Santiago Pérez, A; Finlayson, G; Allue, E; Baena Preysler, J; Cáceres, I; Carrión, JS; Fernández Jalvo, Y; Gleed-Owen, CP; Jimenez Espejo, FJ; López, P; López Sáez, JA; Riquelme Cantal, JA; Sánchez Marco, A; Guzman, FG; Brown, K; Fuentes, N; Valarino, CA; Villalpando, A; Stringer, CB; Martinez Ruiz, F; Sakamoto, T (October 2006). "Late survival of Neanderthals at the southernmost extreme of Europe". Nature. 443 (7113): 850–3. Bibcode:2006Natur.443..850F. doi:10.1038/nature05195. ISSN 0028-0836. PMID 16971951. 
  24. ^ Outside Europe, Mousterian tools were made by both Neanderthals and early modern Homo sapiens. (Donald Johanson & Blake Edgar (2006) From Lucy to Language, Simon & Schuster, p. 272
  25. ^ Stringer, C. (1984). "Human evolution and biological adaptation in the Pleistocene". In Foley, R. Hominid evolution and community ecology. New York: Academic Press. ISBN 978-0122619205. 
  26. ^ Holloway, R.L. (1985). "The poor brain of Homo sapiens neanderthalensis: see what you please". In Delson, E. Ancestors: The hard evidence. New York: Alan R. Liss. ISBN 978-0471843764. 
  27. ^ Amano, H.; Kikuchi, T.; Morita, Y.; Kondo, O.; Suzuki, Hiromasa; et al. (August 2015). "Virtual Reconstruction of the Neanderthal Amud 1 Cranium". American Journal of Physical Anthropology. 158: 185–197. doi:10.1002/ajpa.22777. 
  28. ^ a b Helmuth H (1998). "Body height, body mass and surface area of the Neanderthals". Zeitschrift für Morphologie und Anthropologie. 82 (1): 1–12. PMID 9850627. 
  29. ^ Sánchez-Quinto, F; Botigué, LR; Civit, S; Arenas, C; Avila-Arcos, MC; Bustamante, CD; Comas, D; Lalueza-Fox, C (October 17, 2012). "North African Populations Carry the Signature of Admixture with Neandertals". PLOS ONE. 7: e47765. doi:10.1371/journal.pone.0047765. PMC 3474783Freely accessible. PMID 23082212. Retrieved May 29, 2016. 
  30. ^ a b c Rincon, Paul (May 6, 2010). "Neanderthal genes 'survive in us'". BBC News. Retrieved May 7, 2010. 
  31. ^ Fu, Q; Li, H; Moorjani, P; Jay, F; Slepchenko, SM; Bondarev, AA; Johnson, PL; Aximu-Petri, A; Prüfer, K; de Filippo, C; Meyer, M; Zwyns, N; Salazar-García, DC; Kuzmin, YV; Keates, SG; Kosintsev, PA; Razhev, DI; Richards, MP; Peristov, NV; Lachmann, M; Douka, K; Higham, TF; Slatkin, M; Hublin, JJ; Reich, D; Kelso, J; Viola, TB; Pääbo, S (October 23, 2014). "Genome sequence of a 45,000-year-old modern human from western Siberia". Nature. 514 (7523): 445–449. doi:10.1038/nature13810. PMC 4753769Freely accessible. PMID 25341783. 
  32. ^ Brahic, Catherine. "Humanity's forgotten return to Africa revealed in DNA", The New Scientist (February 3, 2014).
  33. ^ "Neanderthals mated with modern humans much earlier than previously thought, study finds: First genetic evidence of modern human DNA in a Neanderthal individual". ScienceDaily. February 17, 2016. Retrieved March 6, 2016. 
  34. ^ The Combined Landscape of Denisovan and Neanderthal Ancestry in Present-Day Humans, Current Biology, Sankararaman et al., 26, 1241–1247, 2016.
  35. ^ a b Stringer, C.; Gamble, C. (1993). In Search of the Neanderthals. London: Thames and Hudson. ISBN 978-0500050705. 
  36. ^ Rendu W, Beauval C, Crevecoeur I, Bayle P, Balzeau A, Bismuth T, Bourguignon L, Delfour G, Faivre JP, Lacrampe-Cuyaubère F, Muth X, Pasty S, Semal P, Tavormina C, Todisco D, Turq A, Maureille B (2016). "Let the dead speak...comments on Dibble et al.'s reply to "Evidence supporting an intentional burial at La Chapelle-aux-Saints"". Journal of Archaeological Science. 69: 12–20. doi:10.1016/j.jas.2016.02.006. 
  37. ^ Gargett, R.H. (1999). "Middle Palaeolithic burial is not a dead issue: the view from Qafzeh, Saint-Césaire, Kebara, Amud, and Dederiyeh". Journal of Human Evolution. 37: 27–90. doi:10.1006/jhev.1999.0301. 
  38. ^ Gargett, R.H. (1989). "Grave Shortcomings: The Evidence for Neandertal Burial". Current Anthropology. 30 (2): 157–190. doi:10.1086/203725. 
  39. ^ a b Rendu W, Beauval C, Crevecoeur I, Bayle P, Balzeau A, Bismuth T, Bourguignon L, Delfour G, Faivre JP, Lacrampe-Cuyaubère F, Tavormina C, Todisco D, Turq A, Maureille B (January 2014). "Evidence supporting an intentional Neandertal burial at La Chapelle-aux-Saints". Proceedings of the National Academy of Sciences. 111 (1): 81–86. doi:10.1073/pnas.1316780110. 
  40. ^ H. Dibble and V. Aldeias and P. Goldberg and D. Sandgathe and T.E. Steele (2015). "A critical look at evidence from La Chapelle-aux-Saints supporting an intentional burial". Journal of Archaeological Science: 649–657. 
  41. ^ "Studies find Neanderthal genes in modern humans". 
  42. ^ Zimmer, Carl (December 18, 2013). "Toe Fossil Provides Complete Neanderthal Genome". New York Times. Retrieved December 18, 2013. 
  43. ^ Prüfer, Kay; et al. (2014). "The complete genome sequence of a Neanderthal from the Altai Mountains". Nature. 505 (7481): 43–49. doi:10.1038/nature12886. PMC 4031459Freely accessible. PMID 24352235. 
  44. ^ a b Jaubert, Jacques; Verheyden, Sophie; Genty, Dominique; Soulier, Michel; Cheng, Hai; Blamart, Dominique; Burlet, Christian; Camus, Hubert; Delaby, Serge; Deldicque, Damien; Edwards, R. Lawrence; Ferrier, Catherine; Lacrampe-Cuyaubère, François; Lévêque, François; Maksud, Frédéric; Mora, Pascal; Muth, Xavier; Régnier, Édouard; Rouzaud, Jean-Noël; Santos, Frédéric (June 2, 2016) [online May 25, 2016]. "Early Neanderthal Constructions deep in Bruniquel Cave in Southwestern France". Nature. 534 (7605): 111–114. doi:10.1038/nature18291. ISSN 0028-0836. PMID 27251286. 
  45. ^ Kunzig, Robert. "The Year in Science: Human Origins 1997", Discover (magazine) (January 1, 1998) reprinted in Contemporary Readings in Physical Anthropology, p. 145 (Alan Almquist ed., Prentice Hall, 2000)
  46. ^ a b Howell, F. Clark (1957). "The evolutionary significance of variation and varieties of 'Neanderthal' man". The Quarterly Review of Biology. 32 (4): 330–47. doi:10.1086/401978. JSTOR 2816956. PMID 13506025. 
  47. ^ Foley, Tim. TalkOrigins Archive. "Neanderthal or Neandertal?". 2005.
  48. ^ Vogt, Karl C (1864). Lectures on Man: His Place in Creation, and in the History of the Earth. London: Longman, Green, Longman and Roberts. pp. 302, 473. 
  49. ^ Inter alia, Boys' Life, p. 18. January 1924.
  50. ^ The Oxford Illustrated Dictionary. Great Britain: Oxford University Press. 1976 [1975]. p. 564. (tahl) 
  51. ^ "Neanderthal adjective—definition in British English Dictionary & Thesaurus—Cambridge Dictionary Online". January 8, 2013. Retrieved January 22, 2013. 
  52. ^ "Oxford Learner's Dictionaries—Find pronunciation, clear meanings and definitions of words at". [dead link]
  53. ^ "Neanderthal | Define Neanderthal at". Retrieved January 22, 2013. 
  54. ^ Kurtén, Björn (October 10, 1995). Dance of the Tiger: A Novel of the Ice Age. University of California Press. pp. xxi. ISBN 0-520-20277-5. Retrieved May 9, 2012. 
  55. ^ Pollet, Carl J. (September 21, 1991). "…And Etymology". Science News. 140 (12): 191. doi:10.2307/3975867. JSTOR 3975867. 
  56. ^ Tattersall, Ian; Schwartz, Jeffrey H. (1999). "Hominids and hybrids: The place of Neanderthals in human evolution". Proceedings of the National Academy of Sciences. 96 (13): 7117–9. Bibcode:1999PNAS...96.7117T. doi:10.1073/pnas.96.13.7117. JSTOR 48019. PMC 33580Freely accessible. PMID 10377375. 
  57. ^ Duarte, Cidália; Mauricio, João; Pettitt, Paul B.; Souto, Pedro; Trinkaus, Erik; Van Der Plicht, Hans; Zilhao, João (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. 96 (13): 7604–9. Bibcode:1999PNAS...96.7604D. doi:10.1073/pnas.96.13.7604. JSTOR 48106. PMC 22133Freely accessible. PMID 10377462. 
  58. ^ "L'homme fossile de La Chapelle-aux-Saints - full text - Volume VI (p. 11–172), Volume VII (p. 21–56), Volume VIII (p. 1–70), 1911–1913". Royal College of Surgeons of England. Retrieved July 26, 2016. 
  59. ^ "Marcellin Boule - French geologist". Encyclopædia Britannica. Retrieved July 26, 2016. 
  60. ^ "Arthur Keith". Royal Anthropological Institute. Retrieved July 26, 2016. 
  61. ^ "La Chapelle-Aux-Saints - The old man of La Chapelle - The original reconstruction of the 'Old Man of La Chapelle' by scientist Pierre Marcellin Boule led to the reason why popular culture stereotyped Neanderthals as dim-witted brutes for so many years.". Smithsonian Institution. Retrieved July 26, 2016. 
  62. ^ "Our Neandertal Brethren: Why They Were Not a Separate Species". SCIENTIFIC AMERICAN. August 1, 2010. Retrieved July 26, 2016. 
  63. ^ Wolpoff, MH; Hawks, J; Caspari, R (2000). "Multiregional, not multiple origins" (pdf). American Journal of Physical Anthropology. 112 (1): 129–36. doi:10.1002/(SICI)1096-8644(200005)112:1<129::AID-AJPA11>3.0.CO;2-K. PMID 10766948. 
  64. ^ Pääbo, Svante (2014). Neanderthal Man: In Search of Lost Genomes. New York: Basic Books. p. 237. 
  65. ^ Rightmire, G. P. (2001). "Patterns of hominid evolution and dispersal in the Middle Pleistocene". Quaternary International. 75 (1): 77–84. doi:10.1016/S1040-6182(00)00079-3. 
  66. ^ Toro-Moyano, I.; Martínez-Navarro, B.; Agustí, J.; Souday, C.; Bermúdez de Castro, J. M.; Martinón-Torres, M.; Palmqvist, P. (2013). "The oldest human fossil in Europe, from Orce (Spain)". Journal of Human Evolution. 65 (1): 1–9. doi:10.1016/j.jhevol.2013.01.012. 
  67. ^ Wayman, Erin (November 26, 2012). "Homo antecessor: Common Ancestor of Humans and Neanderthals?". Smithsonian Institution. Retrieved May 17, 2017. 
  68. ^ M. Krings; A. Stone; R.W. Schmitz; H. Krainitzki; M. Stoneking; S. Pääbo (1997). "Neandertal DNA sequences and the origin of modern humans". Cell. 90: 19–30. doi:10.1016/S0092-8674(00)80310-4. PMID 9230299. (subscription required)
  69. ^ M. Krings; H. Geisert; R.W. Schmitz; H. Krainitzki; S. Pääbo (1999). "DNA sequence of the mitochondrial hypervariable region II from the Neandertal type specimen". Proceedings of the National Academy of Sciences USA. 96 (10): 5581–5585. doi:10.1073/pnas.96.10.5581. (subscription required)
  70. ^ P. Beerli; S.V. Edwards (2002). "When did Neanderthals and modern humans diverge?". Evolutionary Anthropology. 11 (S1): 60–63. doi:10.1002/evan.10058. (subscription required)
  71. ^ I.V. Ovchinnikov; A. Götherström; G.P. Romanova; V.M. Kharitonov; K. Lidén; W. Goodwin (2002). "Molecular analysis of Neandertal DNA from the northern Caucasus". Nature. 404: 490–493. doi:10.1038/35006625. PMID 10761915. (subscription required)
  72. ^ R.E. Green; A.-S. Malaspinas; J. Krause; A.W. Briggs; P.L.F. Johnson; C. Uhler; M. Meyer; J.M. Good; T. Maricic; U. Stenzel; et al. (2008). "A Complete Neandertal Mitochondrial Genome Sequence Determined by High-Throughput Sequencing". Cell. 134 (3): 416–426. doi:10.1016/j.cell.2008.06.021. PMC 2602844Freely accessible. PMID 18692465. (subscription required)
  73. ^ R.E. Green; A.-S. Malaspinas; J. Krause; A.W. Briggs; P.L.F. Johnson; C. Uhler; M. Meyer; J.M. Good; T. Maricic; U. Stenzel; et al. (2009). "Targeted Retrieval and Analysis of Five Neandertal mtDNA Genomes". Science. 325 (5938): 318–321. doi:10.1126/science.1174462. (subscription required)
  74. ^ P. Endicott; S.Y.W. Ho; C. Stringer (2010). "Using genetic evidence to evaluate four palaeoanthropological hypotheses for the timing of Neanderthal and modern human origins". Journal of Human Evolution. 59 (1): 87–95. doi:10.1016/j.jhevol.2010.04.005. (subscription required)
  75. ^ A. Rieux; A. Eriksson; M. Li; B. Sobkowiak; L.A. Weinert; V. Warmuth; A. Ruiz-Linares; A. Manica; F. Balloux (2014). "Improved calibration of the human mitochondrial clock using ancient genomes". Molecular Biology and Evolution. 31 (10): 2780–2792. doi:10.1093/molbev/msu222. PMC 4166928Freely accessible. PMID 25100861. (subscription required)
  76. ^ K. Prüfer; F. Racimo; N. Patterson; F. Jay; S. Sankararaman; S. Sawyer; A. Heinze; G. Renaud; P.H. Sudmant; C. de Filippo; et al. (2014). "The complete genome sequence of a Neanderthal from the Altai Mountains". Nature. 505: 43–49. doi:10.1038/nature12886. PMC 4031459Freely accessible. PMID 24352235. (subscription required)
  77. ^ Cookson, Clive (June 27, 2014). "Palaeontology: How Neanderthals evolved". Financial Times. Retrieved October 28, 2015. 
  78. ^ Bischoff, James L.; Shamp, Donald D.; Aramburu, Arantza; Arsuaga, Juan Luis; Carbonell, Eudald; Bermudez de Castro, J.M. (2003). "The Sima de los Huesos Hominids Date to Beyond U/Th Equilibrium (>350kyr) and Perhaps to 400–500kyr: New Radiometric Dates". Journal of Archaeological Science. 30 (3): 275–80. doi:10.1006/jasc.2002.0834. 
  79. ^ Stringer, Chris (2011). The Origin of our Species. Penguin. pp. 26–29, 202. ISBN 978-0-141-03720-2. 
  80. ^ Johansson, Donald; Edgar, Blake (2006). From Lucy to Language. Simon & Schuster. p. 38. ISBN 978-0-7432-8064-8. 
  81. ^ a b c d e Papagianni, Dmitra; Morse, Michael (2013). The Neanderthals Rediscovered. Thames & Hudson. ISBN 978-0-500-05177-1. 
  82. ^ "Homo neanderthalensis". Smithsonian Institution. Archived from the original on May 21, 2009. Retrieved May 18, 2009. 
  83. ^ "New Evidence On The Role Of Climate In Neanderthal Extinction". Science Daily. 
  84. ^ "Neanderthal - Homo neanderthalensis - Details - Key Fossils - La Chapelle-aux-Saints". Encyclopedia of Life. Retrieved July 18, 2016. 
  85. ^ "Universitat Pompeu Fabra, Barcelona, A paleogenetical study determines the blood group of Neanderthal man" Archived November 22, 2012, at the Wayback Machine.
  86. ^ "Palaeogenetic research at the El Sidrón Neanderthal site". Annals of Anatomy - Anatomischer Anzeiger. 194: 133–137. doi:10.1016/j.aanat.2011.01.014. 
  87. ^ a b Krings, M; Stone, A; Schmitz, RW; Krainitzki, H; Stoneking, M; Pääbo, S (1997). "Neandertal DNA sequences and the origin of modern humans". Cell. 90 (1): 19–30. doi:10.1016/S0092-8674(00)80310-4. ISSN 0092-8674. PMID 9230299. 
  88. ^ Schmitz, Ralf W; et al. (2002). "The Neandertal type site revisited: Interdisciplinary investigations of skeletal remains from the Neander Valley, Germany". Proceedings of the National Academy of Sciences. 99 (20): 13342–13347. doi:10.1073/pnas.192464099. PMC 130635Freely accessible. PMID 12232049. 
  89. ^ a b Duarte, C; Maurício, J; Pettitt, PB; Souto, P; Trinkaus, E; Van Der Plicht, H; Zilhão, J; et al. (1999). "The early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern human emergence in the Iberian Peninsula". Proceedings of the National Academy of Sciences of the United States of America. PNAS. 96 (13): 7604–7609. Bibcode:1999PNAS...96.7604D. doi:10.1073/pnas.96.13.7604. PMC 22133Freely accessible. PMID 10377462. Retrieved June 21, 2009. 
  90. ^ a b Ovchinnikov, IV; Götherström, A; Romanova, GP; Kharitonov, VM; Lidén, K; Goodwin, W (2000). "Molecular analysis of Neanderthal DNA from the northern Caucasus". Nature. 404 (6777): 490–3. doi:10.1038/35006625. PMID 10761915. 
  91. ^ Green, RE; Krause, J; Ptak, SE; et al. (November 16, 2006). "Analysis of one million base pairs of Neanderthal DNA". Nature. 444: 330–6. doi:10.1038/nature05336. PMID 17108958. Retrieved July 18, 2016. 
  92. ^ Morgan, James (February 12, 2009). "Neanderthals 'distinct from us'". BBC News. Retrieved May 22, 2009. 
  93. ^ a b c d e f g Green, Richard E.; Krause, Johannes; Briggs, Adrian W.; Maricic, Tomislav; Stenzel, Udo; Kircher, Martin; Patterson, Nick; Li, Heng; Zhai, Weiwei; Fritz, Markus Hsi-Yang; Hansen, Nancy F.; Durand, Eric Y.; Malaspinas, Anna-Sapfo; Jensen, Jeffrey D.; Marques-Bonet, Tomas; Alkan, Can; Prüfer, Kay; Meyer, Matthias; Burbano, Hernán A.; Good, Jeffrey M.; Schultz, Rigo; Aximu-Petri, Ayinuer; Butthof, Anne; Höber, Barbara; Höffner, Barbara; Siegemund, Madlen; Weihmann, Antje; Nusbaum, Chad; Lander, Eric S.; Russ, Carsten (2010). "A Draft Sequence of the Neandertal Genome". Science. 328 (5979): 710–22. Bibcode:2010Sci...328..710G. doi:10.1126/science.1188021. PMC 5100745Freely accessible. PMID 20448178. 
  94. ^ "Neanderthals More Intelligent Than Thought". September 22, 2010. Archived from the original on September 25, 2010. Retrieved September 27, 2010. 
  95. ^ Benazzi, S.; Douka, K.; Fornai, C.; Bauer, C. C.; Kullmer, O.; Svoboda, J. Í.; Pap, I.; Mallegni, F.; Bayle, P.; Coquerelle, M.; Condemi, S.; Ronchitelli, A.; Harvati, K.; Weber, G. W. (2011). "Early dispersal of modern humans in Europe and implications for Neanderthal behaviour". Nature. 479 (7374): 525–528. doi:10.1038/Nature10617. PMID 22048311. 
  96. ^ Fergal MacErlean (February 10, 2012). "First Neanderthal cave paintings discovered in Spain". New Scientist. Retrieved February 10, 2012. 
  97. ^ Viegas, Jennifer (March 27, 2013). "First Love Child of Human, Neanderthal Found". Discovery News. Retrieved April 11, 2013. 
  98. ^ P. Villa, W. Roebroeks (2014). "Neandertal Demise: An Archaeological Analysis of the Modern Human Superiority Complex". PLoS ONE. 9 (4): e96424. doi:10.1371/journal.pone.0096424. 
  99. ^ "The Human Lineage by Matt Cartmill, Fred H. Smith". Google Books. Retrieved September 27, 2015. 
  100. ^ "Ancient tooth provides evidence of Neanderthal movement" (Press release). Durham University. February 11, 2008. Retrieved May 18, 2009. 
  101. ^ O'Neill, Dennis. "Evolution of Modern Humans: Neanderthals", Palomar College, June 10, 2011. Retrieved August 21, 2011.
  102. ^ a b c Jordan, P. (2001) Neanderthal: Neanderthal Man and the Story of Human Origins. The History Press ISBN 978-0-7509-2676-8.
  103. ^ Bilsborough, A, Rae, TC (2015) [REVISION OF] Hominoid cranial diversity and adaptation. In: Henke, W., Rothe, H. & Tattersall, I. (eds.) Handbook of Palaeoanthropology, New York: Springer
  104. ^ "Fieldwork—Jebel Irhoud". Max Planck Institute, Department of Human Evolution. 
  105. ^ Arsuaga, J.L; Gracia, A; Martínez, I; Bermúdez de Castro, J.M; Rosas, A; Villaverde, V; Fumanal, M.P (1989). "The human remains from Cova Negra (Valencia, Spain) and their place in European Pleistocene human evolution". Journal of Human Evolution. 19: 55–92. doi:10.1016/0047-2484(89)90023-7. 
  106. ^ Mallegni, F., Piperno, M., and Segre, A (1987). "Human remains of Homo sapiens neanderthalensis from the Pleistocene deposit of Sants Croce Cave, Bisceglie (Apulia), Italy". American Journal of Physical Anthropology. 72 (4): 421–429. doi:10.1002/ajpa.1330720402. PMID 3111268. 
  107. ^ Pavlov P, Roebroeks W, Svendsen JI (2004). "The Pleistocene colonization of northeastern Europe: a report on recent research". Journal of Human Evolution. 47 (1–2): 3–17. doi:10.1016/j.jhevol.2004.05.002. PMID 15288521. 
  108. ^ Wade, Nicholas (October 2, 2007). "Fossil DNA Expands Neanderthal Range". The New York Times. Retrieved May 18, 2009. 
  109. ^ Ravilious, Kate (October 1, 2007). "Neandertals Ranged Much Farther East Than Thought". National Geographic Society. Retrieved May 18, 2009. 
  110. ^ L’Homme de Neanderthal par Paul Dardé : L’Homme Primitif
  111. ^ "Science & Nature—Wildfacts—Neanderthal". BBC. Archived from the original on January 15, 2009. Retrieved June 21, 2009. 
  112. ^ "Neanderthal". BBC. Archived from the original on January 15, 2009. Retrieved May 18, 2009. 
  113. ^ Froehle, Andrew W; Churchill, Steven E (2009). "Energetic Competition Between Neandertals and Anatomically Modern Humans" (PDF). PaleoAnthropology: 96–116. Retrieved October 31, 2011. 
  114. ^ Laleuza-Fox, Carles; Römpler, Holger; et al. (October 25, 2007). "A Melanocortin 1 Receptor Allele Suggests Varying Pigmentation Among Neanderthals". Science. 318 (5855): 1453–5. Bibcode:2007Sci...318.1453L. doi:10.1126/science.1147417. PMID 17962522. ; see also Rincon, Paul (October 25, 2007). "Neanderthals 'were flame-haired'". BBC News. Retrieved October 25, 2007. 
  115. ^ Cochran, G. and Harpending, H., The 10,000 Year Explosion (2009)
  116. ^ 14:186–198. Hoffecker JF (2009) The spread of modern humans in Europe. Proc. Nat. Acad. Sci. 106:16040– References 399 Hoffecker JF, Cleghorn ...
  117. ^ "Neanderthal brains focused on vision and movement leaving less room for social networking". Science Daily. March 19, 2013. 
  118. ^ SINC Servicio de Información y Noticias Científicas. "El cerebro neandertal era más asimétrico que el del 'Homo sapiens'". 
  119. ^ "Neanderthal Brain Size at Birth Sheds Light on Human Evolution". National Geographic. September 9, 2008. Retrieved September 19, 2009. 
  120. ^ Silberman, Neil. The Oxford Companion to Archaeology, p. 455 (Oxford University Press 2012): "[I]t is with the Neanderthals that we see the full achievement, for the first time, of the degree of encephalization (brain to body size ratio) that characterizes modern humans."
  121. ^ Abramiuk, Marc. The Foundations of Cognitive Archaeology, p. 199 (MIT Press 2012): "the encephalization quotient was slightly smaller".
  122. ^ Moskvitch, Katia (September 24, 2010). "Neanderthals were able to 'develop their own tools'". BBC News. BBC. Retrieved October 1, 2010. 
  123. ^ Heyes, Peter; Anastasakis, Konstantinos; de Jong, Wiebren (2016). "Selection and Use of Manganese Dioxide by Neanderthals". Scientific Reports. 6: 22159. doi:10.1038/srep22159. ISSN 2045-2322. PMC 4770591Freely accessible. PMID 26922901. 
  124. ^ Bocherens, Hervé; Drucker, Dorothée G.; Billiou, Daniel; Patou-Mathis, Marylène; Vandermeersch, Bernard (2005). "Isotopic evidence for diet and subsistence pattern of the Saint-Césaire I Neanderthal: Review and use of a multi-source mixing model". Journal of Human Evolution. 49 (1): 71–87. doi:10.1016/j.jhevol.2005.03.003. PMID 15869783. 
  125. ^ a b Ghosh, Pallab. "Neanderthals cooked and ate vegetables." BBC News. December 27, 2010.
  126. ^ Lichfield, John (September 30, 2006). "French dig up Neanderthal 'butcher's shop'". The New Zealand Herald. 
  127. ^ a b Richards, Michael P.; Pettitt, Paul B.; Trinkaus, Erik; Smith, Fred H.; Paunović, Maja; Karavanić, Ivor (2000). "Neanderthal diet at Vindija and Neanderthal predation: The evidence from stable isotopes". Proceedings of the National Academy of Sciences. 97 (13): 7663–6. Bibcode:2000pnas...97.7663r. doi:10.1073/pnas.120178997. JSTOR 122870. PMC 16602Freely accessible. PMID 10852955. 
  128. ^ Fiorenza, Luca; Benazzi, Stefano; Tausch, Jeremy; Kullmer, Ottmar; Bromage, Timothy G.; Schrenk, Friedemann (2011). Rosenberg, Karen, ed. "Molar Macrowear Reveals Neanderthal Eco-Geographic Dietary Variation". PLoS ONE. 6 (3): e14769. doi:10.1371/journal.pone.0014769. PMC 3060801Freely accessible. PMID 21445243. 
  129. ^ Henry, A. G.; Brooks, A. S.; Piperno, D. R. (2010). "Microfossils in calculus demonstrate consumption of plants and cooked foods in Neanderthal diets (Shanidar III, Iraq; Spy I and II, Belgium)". Proceedings of the National Academy of Sciences. 108 (2): 486–491. Bibcode:2011PNAS..108..486H. doi:10.1073/pnas.1016868108. PMC 3021051Freely accessible. PMID 21187393. 
  130. ^ Webb, Jonathan (June 25, 2014). "Oldest human faeces show Neanderthals ate vegetables". BBC News. 
  131. ^ "Neanderthal behaviour, diet, and disease inferred from ancient DNA in dental calculus : Nature". The Guardian. Retrieved March 9, 2017. 
  132. ^ Nicola Davis (March 8, 2017). "Neanderthal dental tartar reveals plant-based diet – and drugs". The Guardian. Retrieved March 9, 2017. 
  133. ^ a b Shaw, Kate (July 29, 2011). "Sheer Numbers Gave Early Humans Edge Over Neanderthals". 
  134. ^ a b Vergano, Dan (April 22, 2014). "Neanderthals Lived in Small, Isolated Populations, Gene Analysis Shows". National Geographic. 
  135. ^ a b Tattersall, Ian (2015). The Strange Case of the Rickety Cossack and other Cautionary Tales from Human Evolution. Palgrave Macmillan. p. 202. ISBN 978-1-137-27889-0. 
  136. ^ Zollikofer, Christoph; Marcia, Ponce; Leon, De; Vandermeersch, Bernard; Leveque, Francois (2002). "Evidence for Interpersonal Violence in the St. Césaire Neanderthal". PNAS. 99 (9): 6444–448. doi:10.1073/pnas.082111899. PMC 122968Freely accessible. PMID 11972028. 
  137. ^ Gray, Richard (December 18, 2011). "Neanderthals built homes with mammoth bones". 
  138. ^ Demay, Laëtitia; Péan, Stéphane; Patou-Mathis, Marylène (2012). "Mammoths used as food and building resources by Neanderthals: Zooarchaeological study applied to layer 4, Molodova I (Ukraine)". Quaternary International. 276–277: 212–226. Bibcode:2012QuInt.276..212D. doi:10.1016/j.quaint.2011.11.019. ISSN 1040-6182. 
  139. ^ Yirka, Bob (March 1, 2012). "Evidence suggests Neanderthals took to boats before modern humans". Retrieved May 5, 2016. 
  140. ^ Marshall, Michael (February 29, 2012). "Neanderthals were ancient mariners". New Scientist. Retrieved May 5, 2016. 
  141. ^ "Neanderthals May Have Sailed to Crete". DNews. December 13, 2012. Retrieved May 5, 2016. 
  142. ^ Brown, Cynthia Stokes. Big History. New York, NY: The New Press, 2008. Print.
  143. ^ Moulson, Geir; Associated Press (July 20, 2006). "Neanderthal genome project launches". MSNBC. Retrieved August 22, 2006. 
  144. ^ Lunine 2013, p. 251: "The Neanderthal genome is about the same size as the human genome, and is identical to ours to a level of 99.7% (this is comparing the ordering of the lettering in the nucleotide bases)."
  145. ^ Green RE, Krause J, Ptak SE, et al. (November 2006). "Analysis of one million base pairs of Neanderthal DNA". Nature. 444 (7117): 330–6. Bibcode:2006Natur.444..330G. doi:10.1038/nature05336. PMID 17108958. 
  146. ^ Wade, Nicholas (November 15, 2006). "New Machine Sheds Light on DNA of Neanderthals". The New York Times. Retrieved May 18, 2009. 
  147. ^ Pennisi, E. (May 2007). "Ancient DNA. No sex please, we're Neandertals". Science. 316 (5827): 967. doi:10.1126/science.316.5827.967a. PMID 17510332. 
  148. ^ "Neanderthal bone gives DNA clues". CNN. Associated Press. November 16, 2006. Archived from the original on November 18, 2006. Retrieved May 18, 2009. 
  149. ^ Than, Ker; LiveScience (November 15, 2006). "Scientists decode Neanderthal genes". MSNBC. Retrieved May 18, 2009. 
  150. ^ "Neanderthal Genome Sequencing Yields Surprising Results And Opens A New Door To Future Studies" (Press release). Lawrence Berkeley National Laboratory. November 16, 2006. Retrieved May 31, 2009. 
  151. ^ a b Hayes, Jacqui (November 15, 2006). "DNA find deepens Neanderthal mystery". Cosmos. Archived from the original on March 26, 2009. Retrieved May 18, 2009. 
  152. ^ Green, RE; Malaspinas, AS; Krause, J; Briggs, Aw; Johnson, PL; Uhler, C; Meyer, M; Good, JM; Maricic, T; Stenzel, U; Prüfer, K; Siebauer, M; Burbano, HA; Ronan, M; Rothberg, JM; Egholm, M; Rudan, P; Brajković, D; Kućan, Z; Gusić, I; Wikström, M; Laakkonen, L; Kelso, J; Slatkin, M; Pääbo, S (2008). "A complete Neandertal mitochondrial genome sequence determined by high-throughput sequencing". Cell. 134 (3): 416–26. doi:10.1016/j.cell.2008.06.021. ISSN 0092-8674. PMC 2602844Freely accessible. PMID 18692465. 
  153. ^ a b Elizabeth Pennisi (2009). "Tales of a Prehistoric Human Genome". Science. 323 (5916): 866–871. doi:10.1126/science.323.5916.866. PMID 19213888. 
  154. ^ Green RE, Briggs AW, Krause J, Prüfer K, Burbano HA, Siebauer M, Lachmann M, Pääbo S (2009). "The Neandertal genome and ancient DNA authenticity". EMBO J. 28 (17): 2494–502. doi:10.1038/emboj.2009.222. PMC 2725275Freely accessible. PMID 19661919. 
  155. ^ a b c "New studies reveal 20 Percent of Neanderthal genome lives on in modern humans". Retrieved October 7, 2016. 
  156. ^ "Surprise! 20 Percent of Neanderthal Genome Lives On in Modern Humans, Scientists Find". National Geographic. Retrieved October 7, 2016. 
  157. ^ "Ancient DNA and Neanderthals - The Smithsonian Institution's Human Origins Program". 
  158. ^ "BBC NEWS - Science/Nature - Neanderthals 'were flame-haired'". 
  159. ^ Lalueza-Fox, Carles; Römpler, Holger; Caramelli, David; Stäubert, Claudia; Catalano, Giulio; Hughes, David; Rohland, Nadin; Pilli, Elena; Longo, Laura; Condemi, Silvana; Rasilla, Marco de la; Fortea, Javier; Rosas, Antonio; Stoneking, Mark; Schöneberg, Torsten; Bertranpetit, Jaume; Hofreiter, Michael (November 30, 2007). "A Melanocortin 1 Receptor Allele Suggests Varying Pigmentation Among Neanderthals". Science. 318 (5855): 1453–1455. doi:10.1126/science.1147417. PMID 17962522 – via 
  160. ^ Neanderthals did not interbreed with humans, scientists find Archived April 19, 2014, at the Wayback Machine.. Telegraph. Retrieved May 24, 2014.
  161. ^ Press Association (February 4, 2013). "Neanderthals 'unlikely to have interbred with human ancestors'". The Guardian. Retrieved May 24, 2014. 
  162. ^ a b Lowery, Robert K.; Uribe, Gabriel; Jimenez, Eric B.; Weiss, Mark A.; Herrera, Kristian J.; Regueiro, Maria; Herrera, Rene J. (2013). "Neanderthal and Denisova genetic affinities with contemporary humans: Introgression versus common ancestral polymorphisms". Gene. 530 (1): 83–94. doi:10.1016/j.gene.2013.06.005. ISSN 0378-1119. PMID 23872234. 
  163. ^ Neanderthal breeding idea doubted. BBC. Retrieved March 2, 2016.
  164. ^ a b "Complex History of Admixture between Modern Humans and Neandertals". American Journal of Human Genetics. 96 (3): 454–61. doi:10.1016/j.ajhg.2015.01.006. 
  165. ^ Kim, BY; Lohmueller, KE. "Selection and Reduced Population Size Cannot Explain Higher Amounts of Neandertal Ancestry in East Asian than in European Human Populations". American Journal of Human Genetics. 96 (3): 448–53. doi:10.1016/j.ajhg.2014.12.029. PMC 4375557Freely accessible. PMID 25683122. 
  166. ^ a b [1]. The Daily Mail. Retrieved January 26, 2016.
  167. ^ a b Zimmer, Carl (March 17, 2016). "Humans Interbred With Hominins on Multiple Occasions, Study Finds". The New York Times. Retrieved March 17, 2016. 
  168. ^ Mendez, Fernando L.; et al. (April 7, 2016). "The Divergence of Neandertal and Modern Human Y Chromosomes" (PDF). The American Journal of Human Genetics. 98 (4): 728–734. doi:10.1016/j.ajhg.2016.02.023. PMC 4833433Freely accessible. PMID 27058445. Retrieved April 12, 2016. 
  169. ^ "DNA points to Neanderthal breeding barrier". BBC News, Science & Environment. April 8, 2016. Retrieved April 12, 2016. 
  170. ^ a b Sample, Ian (February 17, 2016). "Oldest known case of Neanderthal-human sex revealed by DNA test". the Guardian. Retrieved May 10, 2016. 
  171. ^ Zimmer, Carl (March 17, 2016). "Humans Interbred With Hominins on Multiple Occasions, Study Finds". New York Times. Retrieved May 5, 2016. 
  172. ^ a b Gokhman D, Lavi E, Prüfer K, Fraga MF, Riancho JA, Kelso J, Pääbo S, Meshorer E, Carmel L (2014). "Reconstructing the DNA methylation maps of the Neandertal and the Denisovan.". Science. 344 (6183): 523–7. doi:10.1126/science.1250368. PMID 24786081. 
  173. ^ a b Higham, Tom et al. (August 21, 2014). "The timing and spatiotemporal patterning of Neanderthal disappearance". Nature. 512 (512): 306–309. doi:10.1038/nature13621. PMID 25143113. (subscription required)
  174. ^ Stein, Richard A (October 1, 2015). "Copy Number Analysis Starts to Add Up". Genetic Engineering & Biotechnology News (Paper). 35 (17): 20. Neanderthals, which are thought to have come into conteact with modern humans approximately 80,000 years ago, appear to have survived until about 35,000 years ago in some regions of Europe. (subscription required)
  175. ^ a b Finlayson, C., Carrión, J.S. (April 2007). "Rapid ecological turnover and its impact on Neanderthal and other human populations". Trends in Ecology & Evolution (Personal Edition). 22 (4): 213–22. doi:10.1016/j.tree.2007.02.001. PMID 17300854. 
  176. ^ "First genocide of human beings occurred 30,000 years ago". Pravda. October 24, 2007. Retrieved May 18, 2009. 
  177. ^ Diamond, Jared M. (1992). The third chimpanzee: the evolution and future of the human animal. New York City: HarperCollins. p. 52. ISBN 0-06-098403-1. OCLC 60088352. 
  178. ^ Currat, Mathias; Excoffier, Laurent (2004). "Modern Humans Did Not Admix with Neanderthals during Their Range Expansion into Europe". PLoS Biology. 2 (12): e421. doi:10.1371/journal.pbio.0020421. PMC 532389Freely accessible. PMID 15562317. 
  179. ^ a b Wong, Kate (August 1, 2009). "The Mysterious Downfall of the Neandertals". Scientific American. 
  180. ^ Adler, Daniel S.; Bar-Oz, Guy; Belfer-Cohen, Anna; Bar-Yosef, Ofer (2006). "Ahead of the Game: Middle and Upper Palaeolithic Hunting Behaviors in the Southern Caucasus". Current Anthropology. 47 (1): 89–118. doi:10.1086/432455. 
  181. ^ Froehle, Andrew W.; Churchill, Steven E. (2009). "Energetic Competition Between Neandertals and Anatomically Modern Humans" (PDF). PaleoAnthropology: 96–116. 
  182. ^ Wilford, John Noble (November 2, 2011). "Fossil Teeth Put Humans in Europe Earlier Than Thought". New York Times. Retrieved August 27, 2014. 
  183. ^ Wolff, H. "Result Filters." National Center for Biotechnology Information. U.S. National Library of Medicine, July 2010. Web. October 22, 2014.
  184. ^ McKie, Robin (March 1, 2015). "How hunting with wolves helped humans outsmart the Neanderthals". The Guardian. Retrieved May 5, 2016. 
  185. ^ Stringer, Chris (2012). "Evolution: What makes a modern human". Nature. 485 (7396): 33–5. Bibcode:2012Natur.485...33S. doi:10.1038/485033a. PMID 22552077. 
  186. ^[full citation needed]
  187. ^ Dan Jones: The Neanderthal within., New Scientist 193.2007, H. 2593 (March 3), 28–32. Modern Humans, Neanderthals May Have Interbred[dead link]; Humans and Neanderthals interbred Archived February 22, 2009, at the Wayback Machine.
  188. ^ [2]; [,,1871842,00.html[full citation needed]
  189. ^ Not a lasting last for the Neandertals—John Hawks weblog, September 13, 2006[full citation needed]
  190. ^ Soficaru, Andrei; Dobos, Adrian; Trinkaus, Erik (2006). "Early modern humans from the Pestera Muierii, Baia de Fier, Romania". Proceedings of the National Academy of Sciences. 103 (46): 17196–201. Bibcode:2006PNAS..10317196S. doi:10.1073/pnas.0608443103. JSTOR 30052409. PMC 1859909Freely accessible. PMID 17085588. 
  191. ^ Wade, Nicholas (July 26, 2012). "Genetic Data and Fossil Evidence Tell Differing Tales of Human Origins". The New York Times. Retrieved May 5, 2015. 
  192. ^ Yotova, V.; Lefebvre, J.-F.; Moreau, C.; Gbeha, E.; Hovhannesyan, K.; Bourgeois, S.; Bédarida, S.; Azevedo, L.; Amorim, A.; Sarkisian, T.; Avogbe, P. H.; Chabi, N.; Dicko, M. H.; Kou' Santa Amouzou, E. S.; Sanni, A.; Roberts-Thomson, J.; Boettcher, B.; Scott, R. J.; Labuda, D. (2011). "An X-Linked Haplotype of Neandertal Origin is Present Among All Non-African Populations". Molecular Biology and Evolution. 28 (7): 1957–62. doi:10.1093/molbev/msr024. PMID 21266489. 
  193. ^ Viegas, Jennifer (July 18, 2011). "All Non-Africans Part Neanderthal, Genetics Confirm". DNews. 
  194. ^ Neandertal ancestry "Iced" - John Hawks weblog, August 15, 2012[full citation needed]
  195. ^ "Resurrecting Surviving Neandertal Lineages from Modern Human Genomes". Science. January 29, 2014. Retrieved February 2, 2014. 
  196. ^ Mitchell, Alanna (January 30, 2012). "DNA Turning Human Story Into a Tell-All". NYTimes. Retrieved January 31, 2012. 
  197. ^ Lohse, Konrad; Frantz, Laurent A. F. (2013). "Maximum likelihood evidence for Neandertal admixture in Eurasian populations from three genomes". Populations and Evolution. 1307: 8263. arXiv:1307.8263Freely accessible. Bibcode:2013arXiv1307.8263L. 
  198. ^ Jha, Alok (August 14, 2012). "Study casts doubt on human-Neanderthal interbreeding theory". The Guardian. Retrieved February 19, 2015. 
  199. ^ Hawks, John (2013). "Significance of Neandertal and Denisovan Genomes in Human Evolution". Annual Review of Anthropology. 42: 433–49. doi:10.1146/annurev-anthro-092412-155548. 
  200. ^ Mason, Paul H.; Short, Roger V. (2011). "Neanderthal-human Hybrids". Hypothesis. 9: e1. doi:10.5779/hypothesis.v9i1.215. 
  201. ^ Viegas, Jennifer (March 27, 2013). "First Love Child of Human, Neanderthal Found". Discovery. 
  202. ^ Condemi, Silvana; Mounier, Aurélien; Giunti, Paolo; Lari, Martina; Caramelli, David; Longo, Laura (2013). Frayer, David, ed. "Possible Interbreeding in Late Italian Neanderthals? New Data from the Mezzena Jaw (Monti Lessini, Verona, Italy)". PLoS ONE. 8 (3): e59781. doi:10.1371/journal.pone.0059781. PMC 3609795Freely accessible. PMID 23544098. 
  203. ^ The Neanderthal Dead, exploring mortuary variability in middle paleolithic eurasia. Paul B. Pettitt (2002)
  204. ^ "Homo neanderthalensis–The Neanderthals". Australian Museum. Retrieved June 26, 2014. 
  205. ^ Bischoff, J; Shamp, Donald D.; Aramburu, Arantza; Arsuaga, Juan Luis; Carbonell, Eudald; Bermudez De Castro, J.M. (2003). "The Sima de los Huesos Hominids Date to Beyond U/Th Equilibrium (>350kyr) and Perhaps to 400–500kyr: New Radiometric Dates". Journal of Archaeological Science. 30 (3): 275–280. doi:10.1006/jasc.2002.0834. 
  206. ^ Arsuaga JL, Martínez I, Gracia A, Lorenzo C (1997). "The Sima de los Huesos crania (Sierra de Atapuerca, Spain). A comparative study". Journal of Human Evolution. 33 (2–3): 219–81. doi:10.1006/jhev.1997.0133. PMID 9300343. 
  207. ^ Kreger, C. David. "Homo neanderthalensis". Retrieved May 16, 2009. 
  208. ^ Mcdermott, F; Grün, R; Stringer, Cb; Hawkesworth, Cj (May 1993). "Mass-spectrometric U-series dates for Israeli Neanderthal/early modern hominid sites". Nature. 363 (6426): 252–5. Bibcode:1993Natur.363..252M. doi:10.1038/363252a0. ISSN 0028-0836. PMID 8387643. 
  209. ^ Rink, W. Jack; Schwarcz, H.P.; Lee, H.K.; Rees-Jones, J.; Rabinovich, R.; Hovers, E. (August 2002). "Electron spin resonance (ESR) and thermal ionization mass spectrometric (TIMS) 230Th/234U dating of teeth in Middle Paleolithic layers at Amud Cave, Israel". Geoarchaeology. 16 (6): 701–717. doi:10.1002/gea.1017. 
  210. ^ Valladas, Hélène; Merciera, N.; Frogeta, L.; Hoversb, E.; Joronc, J.L.; Kimbeld, W.H.; Rak, Y. (March 1999). "TL Dates for the Neanderthal Site of the Amud Cave, Israel". Journal of Archaeological Science. 26 (3): 259–268. doi:10.1006/jasc.1998.0334. 
  211. ^ R.E. Wood, T.F.G. Higham, T. de Torres, N. Tisnérate-Laborde, H. Valladas, J.E. Ortiz, C. Lalueza-Fox, S. Sánchez-Moral, J.C. Cañaveras, A. Rosas, D. Santamaría, M. de la Rasilla (March 20, 2012). "A new date for the Neanderthals from El Sidrón Cave (Asturias, Northern Spain)". Archaeometry. 55 (1): 148–158. doi:10.1111/j.1475-4754.2012.00671.x. 
  212. ^ Hayes, Jacqui (November 2, 2006). "Humans and Neanderthals interbred". Cosmos. Archived from the original on February 22, 2009. Retrieved May 17, 2009. 
  213. ^ Neanderthal image by Kupka, based on Boule, 1909, in Humanity's Journeys Dr. Kathryn Denning, 2005. Retrieved March 17, 2012.



Further reading[edit]

External links[edit]