text:
Neanderthal
From Wikipedia, the free encyclopedia
Jump to: navigation, search
Neanderthal
Temporal range: Middle–Late Pleistocene 0.250–0.040 Ma
PreЄ
Є
O
S
D
C
P
T
J
K
Pg
N
↓
The Neanderthal skull of La Chapelle-aux-Saints 1
An approximate reconstruction of a Neanderthal skeleton. (The central ribcage, including the sternum, and parts of the pelvis are from modern humans.)
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Haplorhini
Infraorder: Simiiformes
Family: Hominidae
Genus: Homo
Species: H. neanderthalensis or Homo sapiens
Binomial name
Homo neanderthalensis or Homo sapiens
King, 1864
Known Neanderthal range in Europe (blue), Southwest Asia (orange), Uzbekistan (green), and the Altai mountains (violet).
Synonyms
Homo mousteriensis[1]
Palaeoanthropus neanderthalensis[2]
Neanderthals (UK: /niˈændərˌtɑːl/, also US: /neɪ-, -ˈɑːn-, -ˌtɔːl, -ˌθɔːl/),[3][4] more rarely known as Neandertals,[a] were archaic humans that became extinct about 40,000 years ago.[8][9][10][11][12][13] They seem to have appeared in Europe and later expanded into Southwest, Central, and Northern Asia. There, they left hundreds of stone tool assemblages. Almost all of those younger than 160,000 years are of the so-called Mousterian techno-complex, which is characterised by tools made out of stone flakes.[14]
Neanderthals are considered either a distinct species, Homo neanderthalensis,[15][16][17] or more rarely[18] a subspecies of Homo sapiens (H. s. neanderthalensis).[19][20] Modern humans and Neanderthals share 99.7% of their DNA[21] and are hence much more closely related than to their closest non-human relative, the chimpanzee (98.8%). Compared to modern humans, Neanderthals were stockier, with shorter legs and a bigger body. In conformance with Bergmann's rule, this likely was a Darwinian adaptation to preserve heat in cold climates. Male and female Neanderthals had cranial capacities averaging 1,600 cm3 (98 cu in) and 1,300 cm3 (79 cu in) respectively, [22][23] extending to 1,736 cm3 (105.9 cu in) in the male Amud 1.[24] 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.[25]
The Neanderthal genome project revealed in 2010 that, through interbreeding, Neanderthals contributed to the DNA of modern humans, likely between 50,000 and 60,000 years ago.[26][27][28][29] Today, this is apparent in the genome of most people living outside sub-Saharan Africa, as well as in some sub-Saharan Africans. Subsequent studies suggested there may have been three episodes of interbreeding. The first would have occurred soon after non-African modern humans left Africa. The second would have occurred after the ancestral Melanesians had branched off — these people seem to have thereafter bred with Denisovans. The third would have involved Neanderthals and the ancestors of East Asians only.[30][31][32]
Contents
[hide]
1 Name
2 Classification
3 Origins
4 Discovery
4.1 Timeline of research
5 Habitat and range
6 Anatomy
7 Behavior
7.1 Claims of art and adornment
8 Genome
8.1 Background
8.2 Interbreeding with modern humans
8.3 Epigenetics
9 Extinction
9.1 Climate change
9.2 Coexistence with modern humans
10 Interbreeding hypotheses
11 Specimens
11.1 Notable European Neanderthals
11.2 Notable Southwest Asian Neanderthals
11.3 Notable Central Asian Neanderthal
11.4 Chronology
11.4.1 Mixed with H. heidelbergensis traits
11.4.2 H. neanderthalensis fossils
11.4.3 H. s. sapiens with traits reminiscent of Neanderthals
12 Popular culture
13 See also
14 Notes
15 References
16 Further reading
17 External links
Name[edit]
Human timeline
view • discuss • edit
-10 —
–
-9 —
–
-8 —
–
-7 —
–
-6 —
–
-5 —
–
-4 —
–
-3 —
–
-2 —
–
-1 —
–
0 —
Human-like
apes
Nakalipithecus
Ouranopithecus
Sahelanthropus
Orrorin
Ardipithecus
Australopithecus
Homo habilis
Homo erectus
Neanderthal
Homo sapiens
←
Earlier apes
←
Separation from gorillas
←
Possibly bipedal
←
Separation from chimpanzees
←
Earliest bipedal
←
Earliest stone tools
←
Earliest exit from Africa
←
Earliest fire use
←
Earliest in Europe
←
Earliest cooking
←
Earliest clothes
←
Modern speech
←
Modern humans

P
l
e
i
s
t
o
c
e
n
e

P
l
i
o
c
e
n
e

M
i
o
c
e
n
e

H

o

m

i

n

i

d

s
Axis scale: millions of years ago.
Also see: Life timeline and Nature timeline
Human development
0.2 Mya H. sapiens
0.5 Mya Neanderthal
1.0 Mya H. erectus
2.0 Mya H. habilis
3.0 Mya Australopithecus
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][34][35]
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".[36] 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.[6] Nevertheless, King's name had priority over the proposal put forward in 1866 by Ernst Haeckel, Homo stupidus.[34] The practice of referring to "the Neanderthals" and "a Neanderthal" emerged in the popular literature of the 1920s.[37]
The German pronunciation of 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/).[38][39][40] 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/),[41] although scientists typically use the /t/ as in German.[42][43]
Classification[edit]
Boule (1912)'s now discredited interpretation of Neanderthal morphology (left), compared to an anatomically modern human
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.[44][45] 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."[46]
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.[47] Senior members of their respective national paleontological institutes and among the most eminent paleoanthropologists of their time,[48][49] both men argued that the primitive traits of this skeleton meant it could not be a direct ancestor of Homo sapiens. This idea was reflected in Boule (1912)'s reconstruction of the Chapelle-aux-Saints 1 Neanderthal, now believed inaccurate, in which the skeleton was mounted in a crooked pose with bowed legs.[50]
During the 1930s scholars Ernst Mayr, George Gaylord Simpson and Theodosius Dobzhansky reinterpreted the existing fossil record and came to different conclusions.[51] Neanderthal man was classified as Homo sapiens neanderthalensis – an early subspecies contrasted with what was now called Homo sapiens sapiens. The unbroken succession of fossil sites of both groups in Europe was considered evidence of a slow, 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.[52]
Origins[edit]
See also: Accretion model of Neanderthal origins
Early pre-Neanderthal
(stage 1, 450 ka, male)
Pre-Neanderthal
(stage 2, 350 ka, male)
Early Neanderthal
(stage 3, 130 ka, female)
Classic Neanderthal
(stage 4, male, 50 ka)
The evolution of Neanderthals according to the accretion model. Note the larger nose and rounder eyes of the Classic Neanderthal.
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[53] and Iberia (Spain) by 1.4 Ma.[54] The discoverers of fragmented bones in Spain (Iberia) 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.[55]
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.[56][33][57][58][59][60][61][62][63] 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 or earlier.
Neanderthal traits are present in Homo heidelbergensis specimens beginning between 600,000 and 350,000 years ago.[64][65][66][67] As of 1998, there is a fossil gap in Europe between 300 and 243 ka (MIS 8); no hominin has ever been dated to this period.[68] Conventionally, therefore, European hominins younger than 243,000 years old are called Neanderthals.[68][69]
The quality of the fossil record greatly increases from 130,000 years ago onwards.[70] 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 typical or Classic Neanderthals.[71][72] In morphological studies, the latter term may also be used in a narrower sense for Neanderthals younger than 71,000 years old (MIS 4 and 3).[68]
Discovery[edit]
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.[73]
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.[74]
Timeline of research[edit]
Engis 2, child (1829)
Gibraltar 1, female (1848)
Neanderthal 1, male (upper skull 1856, left-cheek 2000)
Spy 2 skull, sex unclear[75] (1886)
Krapina 3, female (1899)
1829: A damaged skull of a Neanderthal child, Engis 2, is discovered in Engis, Netherlands (now Belgium).
1848: A female Neanderthal skull, Gibraltar 1, is found in Forbes' Quarry, Gibraltar, but its importance is not recognised.
1856: Limestone miners discover the Neanderthal type specimen, Neanderthal 1, in Neandertal, western Prussia (now Germany).
1864: William King is the first to recognise Neanderthal 1 as belonging to a separate species, for which he gives the scientific name Homo neanderthalensis. He then changed his mind on placing it in the Homo genus, arguing that the upper skull was different enough to warrant a separate genus since, to him, it had likely been "incapable of moral and theistic conceptions."[6]
1880: The mandible of a Neanderthal child is discovered in a secure context in Šipka cave, in the Austro-Hungarian Empire (now Czechia), associated with cultural debris, including hearths, Mousterian tools, and bones of extinct animals.
1886: Two well-preserved Neanderthal skeletons are found at Spy, Belgium, making the hypothesis that Neanderthal 1 was only a diseased modern human difficult to sustain.[76]
1899: Sand excavation workers find hundreds of fragmentary Neanderthal remains representing at least 12 and likely as much as 70 individuals on a hill in Krapina, in the Austro-Hungarian Empire (now Croatia).
1908: A very well preserved Neanderthal, La Chapelle-aux-Saints 1, is found in its eponymous site in France,[77] said by the excavators to be a burial, a claim still heatedly contested.[78][79][80] For historical reasons it remains the most famous Neanderthal skeleton.[70]:15
1912: Marcellin Boule publishes his now discredited influential study of Neanderthal skeletal morphology based on La Chapelle-aux-Saints 1.
1953–1957: Ten Neanderthal skeletons are excavated in Shanidar Cave, Iraqi Kurdistan, by Ralph Solecki and colleagues.
1975: Erik Trinkaus's study of Neanderthal feet strongly argues that Neanderthals walked like modern humans.
1981: The site of Bontnewydd, Wales yielded an early Neanderthal tooth, the most north-western Neanderthal remain ever.
1987: Israeli Neanderthal Kebara 2 is dated (by TL and ESR) to 60,000 BP, thus later than the Israeli anatomically modern humans dated to 90,000 and 80,000 BP at Qafzeh and Skhul.
1994: The site of Sidrón Cave, Spain, is discovered where the remains of 12 males and females were to be found. Mitochondrial DNA studies would show that the adult males were genetically related, but the adult females were not, suggesting female exogamy.[81][82]
1997: Matthias Krings et al. are the first to amplify Neanderthal mitochondrial DNA (mtDNA) using a specimen from Feldhofer grotto in the Neander valley.[33]
1997–2000: A small part of the skull of Neanderthal 1, the Neanderthal type fossil discovered in 1856, is found in a dump of limestone mining debris near the original discovery site.[83]
1998: The body of a c. 24,000 year old early Upper Paleolithic child is discovered at Abrigo do Lagar Velho, Portugal, and is described as presenting a mosaic of anatomically modern human and more archaic features, reminiscent of Neanderthals.[45]
2005: The Max Planck Institute for Evolutionary Anthropology and associated institutions launch the Neanderthal genome project to sequence the Neanderthal nuclear genome.[84]
2010: 1–4% of the DNA of living non-African humans are found by the Max Planck Institute to likely come from Neanderthals,[27][85] a result confirmed in 2012,[86][87] and refined to 1.5–2.1% in 2014.[56]
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.[88]
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.[89]
2014: The most comprehensive dating ever of Neanderthal bones and tools from hundreds of sites in Europe is carried out. It suggested that European Neanderthals died out between 41,000 and 39,000 years ago, which coincides with the start of a very cold period in Europe and is no more than 5,000 years after anatomically-modern Homo sapiens reached the continent.[13][90]
Habitat and range[edit]
Further information: Southwest Asian Neanderthals and List of Neanderthal sites
Sites where typical Neanderthal fossils have been found
Early Neanderthals, living before the Eemian interglacial (130 ka), are poorly known and come mostly from European sites. From 130 ka onwards, the quality of the fossil record increases dramatically. From then on, Neanderthal remains are found in Western, Central, Eastern, and Mediterranean Europe,[91] as well as Southwest, Central, and Northern Asia up to the Altai Mountains in Siberia. No Neanderthal has ever been found outside Western Eurasia, namely neither to the south of Jerusalem (Shuqba), nor further east than Kazakhstan (Denisova, Russia), nor to the north of Wales (Bontnewydd),[92][93] although it is difficult to assess the limits of their northern range because glacial advances destroy most human remains, the Bontnewydd tooth being exceptional. Middle Palaeolithic artifacts have been found up to 60° N on the Russian plains.[94]
There likely never were more than 70,000 Neanderthals at any given time.[95]
Anatomy[edit]
Main article: Neanderthal anatomy
Neanderthal anatomy differed from modern humans in that they had a more robust build and distinctive morphological features, especially on the cranium, which gradually accumulated more derived aspects as it was described by Marcellin Boule,[96] particularly in certain isolated geographic regions. These include shorter limb proportions, a wider, barrel-shaped rib cage, a reduced chin, and a large nose, being at the modern human higher end in both width and length,[d] and started somewhat higher on the face than in modern humans.[69] Evidence suggests they were much stronger than modern humans, with particularly strong arms and hands,[97][98] 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.[25] 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.[99] A 2007 genetic study suggested some Neanderthals may have had red hair and blond hair, along with a light skin tone.[100]
Gregory Cochran and Henry Harpending, in the book The 10,000 Year Explosion, investigated whether it is accurate to depict Neanderthals as having hair patterns 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."[101]
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.[102]
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.[103]
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.[104] 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.[105] They had almost the same degree of encephalization (i.e. brain to body size ratio) as modern humans.[106][107]
The Neanderthal skeleton suggests they consumed 100 to 350 kcal (420 to 1,460 kJ) more per day than male modern humans of 68.5 kg (151 lb) and females of 59.2 kg (131 lb).[99]
Behavior[edit]
Levallois point – Beuzeville, France
Main article: Neanderthal behavior
Neanderthals made stone tools,[108] used fire,[109] and were hunters. The consensus on their behaviour ends there.
It had actually long been debated whether Neanderthals were hunters or scavengers.[69][further explanation needed] The discovery of the pre-Neanderthal Schöningen wooden spears in Germany helped settle the debate in favour of hunting. Most available evidence suggests they were apex predators,[110][111] and fed on red deer, reindeer, ibex, wild boar, aurochs and on occasion mammoth, straight-tusked elephant and rhinoceros.[69][112][113][114] They also appear to have occasionally used vegetables as fall-back food,[111][115] which was revealed in the 2000s and 2010s by isotope analysis of their teeth and the study of their coprolites (fossilized feces).[113][116] Dental analysis of specimens from Spy, Belgium and El Sidrón, Spain[117] 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".[118]
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.[119][120] The bones of twelve Neanderthals were discovered at El Sidrón cave in northwestern 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.[121]
The bones of the El Sidrón group show signs of defleshing, suggesting that they were victims of cannibalism.[121] 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, suggesting interpersonal violence.[122]
Claims that Neanderthals deliberately buried their dead, and if they did, whether such burials had any symbolic meaning,[70]:158–60 are heavily contested.[79][123][80] 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,[88] and this has in turn been heavily criticised.[78]
Claims of art and adornment[edit]
Upon Higham et al.'s (2010)[124] publication of new radiocarbon dates shedding doubt on the association of Châtelperronian beads with Neanderthals, Paul Mellars wrote that “the single most impressive and hitherto widely cited pillar of evidence for the presence of complex ‘symbolic’ behavior among the late Neanderthal populations in Europe has now effectively collapsed”.[125] This conclusion, however, is controversial, and others such as Jean-Jacques Hublin and colleagues have re-dated material associated with the Châtelperronian artefacts[126] and used proteomic evidence to restate the challenged association with Neanderthals.[127]
There exists a very large number of other claims of Neanderthal art, adornment, and structures. These are often taken by the media as showing Neanderthals were capable of symbolic thought,[128][129] or were “mental equals” to anatomically modern humans.[130][131] As evidence of symbolism, none of them are widely accepted,[132] although the same is true for Middle Palaeolithic anatomically modern humans. Among many others:
Flower pollen on the body of pre-Neanderthal Shanidar 4, Iraq, had in 1975 been argued to be a flower burial.[133] Once popular, this theory is no longer accepted.[134][135]
Pigmented shells from Murcia, Spain, were argued in 2009 to be Neanderthal make-up containers.[136]
Bird bones were argued to show evidence for feather plucking in a 2012 study examining 1,699 ancient sites across Eurasia, which the authors controversially[137] took to mean Neanderthals wore bird feathers as personal adornments.[138]
Deep scratches were found in 2012 on a cave floor underlying Neanderthal layer in Gorham's Cave, Gibraltar, which some have controversially[139] interpreted as art.[140]
Two 176,000-year-old stalagmite ring structures, several metres wide, were reported in 2016 more than 300 metres from the entrance within Bruniquel Cave, France. The authors claim artificial lighting would have been required as this part of the cave is beyond the reach of daylight and that the structures had been made by early Neanderthals, the only humans in Europe at this time.[141]
All of these claimed manifestations of symbolic thought are one-offs and have not been found at more than one location.
Genome[edit]
Further information: Neanderthal genome project
Background[edit]
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.[59]
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[84] 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.[145] A 2007 study pushes the point of divergence back to around 800,000 years ago.[146]
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.[147][148]

Geneticists first sequenced the entire genome of a Neanderthal in 2013 by extracting it from the phalanx bone of a 50,000-year-old Siberian Neanderthal.[149][150][56]
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.[151] 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.[152]
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."[152]
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."[60] 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.
Geneticist at the Max Planck Institute for Evolutionary Anthropology extracting ancient 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.[27][85] "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 [later refined to 1.5 to 2.1 percent].[56] 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.[85]
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.[85]
It is suggested that 20 percent of Neanderthal DNA survived in modern humans, notably expressed in the skin, hair, and diseases of modern people.[155][156][unreliable source?] Modern human genes involved in making keratin – the protein found in skin, hair, and nails – have specially high levels of Neanderthal DNA.[156] 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.[156][155] 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][100]
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.[85] 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.[159][160][161][162]
Among the genes shown to differ between present-day humans and Neanderthals were RPTN, SPAG17, CAN15, TTF1, FOXP2 and PCD16.[85]
Specifically, a visualization map of the reference modern-human containing the genome regions with high degree of similarity or with novelty according to a Neanderthal of 50k [56] has been built by Pratas et al.[163]
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][better source needed] 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][better source needed]
But John D. Hawks, Professor of Anthropology at the University of Wisconsin writes "that in comparison of East Asian samples (Japanese, Han Chinese in Beijing, and Han Chinese originating in South China) and European samples (Tuscans, British, Finn and CEU samples, along with a handful of Spanish), Europeans average a bit more Neanderthal than Asians. The within-population differences between individuals are large, and constitute noise as far as our comparisons between populations are concerned. At present, we can take as a hypothesis that Europeans have more Neandertal ancestry than Asians. If this is true, we can further guess that Europeans may have mixed with Neanderthals as they moved into Europe, constituting a second process of population mixture beyond that shared by European and Asian ancestors".[167]
Studies published in March 2016 suggest that modern humans bred with hominins, including Neanderthals, on multiple occasions.[168] 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.[169] This could explain why no modern man had to date been found with a Neanderthal Y chromosome.[170] Melanesians and Australoid populations show evidence of only one interbreeding event, possibly about 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.[168]
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.[171]
Epigenetics[edit]
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.[168][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]
Extinction[edit]
Main article: Neanderthal extinction
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.[e] 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]
Anatomically 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.[8][173] The exact nature of biological and cultural interaction between Neanderthals and other human groups is contested.[174]
Possible scenarios for the extinction of the Neanderthals are:
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.[175] Jared Diamond has suggested a scenario of violent conflict and displacement.[176]
Neanderthals were a contemporary subspecies that bred with modern humans and disappeared through absorption (interbreeding theory).
Volcanic catastrophe: see Campanian Ignimbrite Eruption
mtDNA-based simulation of modern human expansion in Europe starting 1600 generations ago. Neanderthal range in light grey[177]
Climate change[edit]
About 55,000 years ago, the climate began to fluctuate wildly from extreme cold conditions to mild cold and back in a matter of decades. Neanderthal bodies were well-suited for survival in a cold climate – their stocky chests and limbs stored body heat better than the Cro-Magnons. Neanderthals died out in Europe between 41,000 and 39,000 years ago, apparently coinciding with the start of a very cold period.[13][178] Raw material sourcing and the examination of faunal remains by Adler et al. (2006) in the southern Caucasus 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 restricted themselves to more local sources.[179]
Coexistence with modern humans[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.[180] 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.[174]
Modern humans co-existed with them in Europe starting around 45,000 years ago and perhaps even earlier. Neanderthals inhabited that continent long 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.[181] 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.[182]
Interbreeding hypotheses[edit]
Main article: Archaic human admixture with modern humans
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.[f]
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.[184] 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.[185] Trinkaus claims various fossils as products of hybridized populations, including the child of Lagar Velho, a skeleton found at Lagar Velho in Portugal.[186][187][188] 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.[189]
Genetic research has asserted that some admixture took place.[190] The genomes of all non-Africans include portions that are of Neanderthal origin,[191][192] 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 1.5 to 2.1 percent[56] of the DNA in Europeans and Asians (French, Chinese and Papua probands) is non-modern, and shared with ancient Neanderthal DNA. This DNA is absent in Sub-Saharan Africans (Yoruba people and San probands).[85] Ötzi the iceman, Europe's oldest preserved mummy, was found to possess an even higher percentage of Neanderthal ancestry.[193] The two percent of Neanderthal DNA that lives on in Europeans and Asians is not the same in all Europeans and Asians: In all, approximately 20% of the Neanderthal genome appears to survive in the modern human gene pool.[194]
2012 genetic studies seem to suggest that modern humans may have mated with "at least two groups" of archaic humans: Neanderthals and Denisovans.[195] Some researchers suggest admixture of 3.4–7.9% in modern humans of non-African ancestry, rejecting the hypothesis of ancestral population structure.[196] 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.[161][197] 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.[198]
While some modern human nuclear DNA has been linked to the extinct Neanderthals, no mitochondrial DNA of Neanderthal origin has been detected,[33] 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.[199] However, some researchers have argued that there is evidence of possible interbreeding between female Neanderthals and male modern humans.[171][200][201]
Specimens[edit]
Notable European Neanderthals[edit]
La Ferrassie 1, skull cast
Le Moustier 1 in 1909
Le Moustier 1 in 2011
Shanidar 1, skull cast
Kebara 2
Teshik-Tash 1
Remains of more than 300 European Neanderthals have been found. For the most important, see List of human evolution fossils.
Neanderthal 1: The first human bones recognised as showing a non-modern anatomy. Discovered in 1856 in a limestone quarry at the Feldhofer grotto in Neanderthal, Germany, they consist of a skull cap, the two femora, the three right arm bones, two left arm bones, the 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 long before death, leading some to suggest he was cared for by others.
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 1: One of the rare nearly complete Neanderthal skeletons to be discovered, it was excavated by a German team in 1908, at Peyzac-le-Moustier, France. Sold to a Berlin museum, the post cranial skeleton was bombed and mostly destroyed in 1945, and parts of the mid face were lost sometime after then. The skull, estimated to be less than 45,000 years old, includes a large nasal cavity and a less developed brow ridge and occipital bun than seen in other Neanderthals. The Mousterian tool techno-complex is named after its discovery site.
Notable Southwest Asian Neanderthals[edit]
Remains of more than 70 Southwest Asian Neanderthals have been found. For a complete list see List of Southwest Asian Neanderthals.
Shanidar 1 to 10: Eight Neanderthals and two pre-Neanderthals (Shanidar 2 and 4) were discovered in the Zagros Mountains in Iraqi Kurdistan. One of the skeletons, Shanidar 4, was once thought to have been buried with flowers, a theory no longer accepted. To Paul B. Pettitt the "deliberate placement of flowers has now been convincingly eliminated", since "[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".[202]
Amud 1: A male adult Neanderthal, dated to roughly 55,000 BP, and one of several found in a cave at Nahal Amud, Israel. At 178 cm (70 in), it is the tallest known Neanderthal. It also has the largest cranial capacity of all extinct hominins: 1,736 cm3.[69][203]
Kebara 2: A male adult post-cranial skeleton, dated to roughly 60,000 BP, that was discovered in 1983 in Kebara Cave, Israel. It has been studied extensively, for its hyoid, ribcage, and pelvis are much better preserved than in all other Neanderthal specimens.
Notable Central Asian Neanderthal[edit]
Teshik-Tash 1: An 8–11 year old skeleton discovered in Uzbekistan by Okladnikov in 1938. This is the only fairly complete skeleton discovered to the east of Iraq. Okladnikov claimed it was a deliberate burial, but this is debated.
Chronology[edit]
This section describes bones with Neanderthal traits in chronological order.
Mixed with H. heidelbergensis traits[edit]
> 350 ka: Sima de los Huesos c. 500:350 ka ago[65][204]
350–200 ka: Pontnewydd 225 ka ago.
200–135 ka: Atapuerca,[205] Vértesszőlős, Ehringsdorf, Casal de'Pazzi, Biache, La Chaise, Montmaurin, Prince, Lazaret, Fontéchevade
H. neanderthalensis fossils[edit]
130–50 ka: Krapina, Saccopastore skulls, Malarnaud, Altamura, Gánovce, Denisova, Okladnikov, Pech de l'Azé, Tabun 120–7002100000000000000♠100±5 ka,[206] Shanidar 1 to 9 80–60 ka, La Ferrassie 1 70 ka, Kebara 60 ka, Régourdou, Mt. Circeo, Combe Grenal, Erd 50 ka, La Chapelle-aux Saints 1 60 ka, Amud I 7001530000000000000♠53±8 ka,[207][208] Teshik-Tash.
In radiocarbon range, > 50 ka: Le Moustier, Feldhofer, La Quina, l'Hortus, Kulna, Šipka, Saint Césaire, Bacho Kiro, El Castillo, Bañolas, El Sidrón (48±3 cal ka),[209] Arcy-sur-Cure, Châtelperron, Figueira Brava, Mezmaiskaya (41±1 cal ka),[10] Zafarraya, Vindija, Velika Pećina.
H. s. sapiens with traits reminiscent of Neanderthals[edit]
< 35 Pestera cu Oase 35 ka, Mladeč 31 ka, Pestera Muierii 30 ka (n/s),[210] Lapedo Child 24.5 ka.
Popular culture[edit]
Main article: Neanderthals in popular culture
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.[211]
See also[edit]
Dawn of Humanity (2015 PBS film)
Denisova hominin
Early human migrations
Homo heidelbergensis
Human evolution
Human timeline
Life timeline
List of Neanderthal sites
Neanderthal behaviour
Neanderthals in popular culture
Neanderthals of Gibraltar
Neanderthals of the Furninha cave, Portugal
Prehistoric Autopsy (2012 BBC documentary)
Notes[edit]
Jump up ^ 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[5] established by King in 1864, Homo neanderthalensis.[6] 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 (lit. "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.[7]
Jump up ^ The valley is named after Joachim Neander, whose Greek-style last name had been changed by his grandfather from "Neumann" ("new man").[33]
Jump up ^ 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.
Jump up ^ There are modern humans with noses as wide as those of Neanderthals and modern humans with similar nose lengths, but none with both Neanderthal nose width and nose length.
Jump up ^ 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.[8]
Jump up ^ 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.[183]
References[edit]
Jump up ^ Romeo, Luigi (1979). Ecce Homo!:A Lexicon of Man. Amsterdam: John Benjamins Publishing Company. p. 92. ISBN 9027220069.
Jump up ^ Camp, C. L; Allison, H. J.; Nichols, R. H. (1964). Bibliography of Fossil Vertebrates 1954–1958. New York: The Geological Society of America, Inc. p. 556. ISBN 9780813710921.
Jump up ^ "Neanderthal in ODE". Oxford Dictionaries.
Jump up ^ ""Neanderthal" in Random House Dictionary (US) & Collins Dictionary (UK)". Dictionary.com.
Jump up ^ "Neanderthal". Wiley-Blackwell Encyclopedia of Human Evolution. Chichester, West Sussex: Wiley-Blackwell. 2013.
^ Jump up to: a b c King, William (Jan 1864). "The Reputed Fossil Man of the Neanderthal" (PDF). The Quarterly Journal of Science. 1: 96.
Jump up ^ "Neandertal oder Neanderthal? – Was ist denn nun richtig?". mettmann.de. 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".]
^ Jump up to: a b c 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): 306–09. Bibcode:2014Natur.512..306H. 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. CS1 maint: Uses authors parameter (link)
Jump up ^ T. Higham (2011). "European Middle and Upper Palaeolithic radiocarbon dates are often older than they look: problems with previous dates and some remedies" (PDF). Antiquity. 85 (327): 235–49. 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. CS1 maint: Uses authors parameter (link)
^ Jump up to: 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 (21): 8611–16. Bibcode:2011PNAS..108.8611P. 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. CS1 maint: Uses authors parameter (link)
^ Jump up to: 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" (PDF). Journal of Human Evolution. 75: 16–27. doi:10.1016/j.jhevol.2014.06.002. PMID 25016565. CS1 maint: Uses authors parameter (link)
Jump up ^ 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.
^ Jump up to: a b c d "New dates rewrite Neanderthal story". BBC News.
Jump up ^ Shaw, Ian; Jameson, Robert, eds. (1999). A Dictionary of Archaeology. Blackwell. p. 408. ISBN 0-631-17423-0. Retrieved August 1, 2016.
Jump up ^ Hublin, J. J. (2009). "The origin of Neandertals". Proceedings of the National Academy of Sciences. 106 (38): 16022–27. Bibcode:2009PNAS..10616022H. doi:10.1073/pnas.0904119106. JSTOR 40485013. PMC 2752594  . PMID 19805257.
Jump up ^ 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 (5): 1147–52. Bibcode:2004PNAS..101.1147H. doi:10.1073/pnas.0308085100. PMC 337021  . PMID 14745010.
Jump up ^ "Scientists Identify Neanderthal Genes in Modern Human DNA". Sci-News.com. January 30, 2014. Retrieved October 29, 2015.
Jump up ^ "Homo neanderthalensis King, 1864". Wiley-Blackwell Encyclopedia of Human Evolution. Chichester, West Sussex: Wiley-Blackwell. 2013. pp. 328–31.
Jump up ^ Bednarik, R.G. (2012). "U–Th analysis and rock art: a response to Pike et al". Rock Art Research. 29 (2): 244–46.
Jump up ^ 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.
Jump up ^ Complete Neanderthal genome sequenced: DNA signatures found in present-day Europeans and Asians, but not in Africans, ScienceDaily
Jump up ^ 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.
Jump up ^ 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.
Jump up ^ 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 (2): 185–97. doi:10.1002/ajpa.22777.
^ Jump up to: 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.
Jump up ^ 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 (10): e47765. Bibcode:2012PLoSO...747765S. doi:10.1371/journal.pone.0047765. PMC 3474783  . PMID 23082212. Retrieved May 29, 2016.
^ Jump up to: a b c Rincon, Paul (May 6, 2010). "Neanderthal genes 'survive in us'". BBC News. Retrieved May 7, 2010.
Jump up ^ 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): 445–49. Bibcode:2014Natur.514..445F. doi:10.1038/nature13810. PMC 4753769  . PMID 25341783.
Jump up ^ Brahic, Catherine. "Humanity's forgotten return to Africa revealed in DNA", The New Scientist (February 3, 2014).
Jump up ^ S. Sankararaman; S. Mallick; M. Dannemann; K. Prüfer; J. Kelso; N. Patterson; D. Reich (2014). "The landscape of Neandertal ancestry in present-day humans". Nature. 507 (7492). pp. 354–357. doi:10.1038/nature12961.
Jump up ^ Sankararaman, Sriram; Mallick, Swapan; Patterson, Nick; Reich, David (2016). "The Combined Landscape of Denisovan and Neanderthal Ancestry in Present-Day Humans". Current Biology. 26 (9): 1241–47. doi:10.1016/j.cub.2016.03.037. ISSN 0960-9822.
Jump up ^ "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.
^ Jump up to: a b c d Krings, Matthias; Stone, Anne; Schmitz, Ralf W; Krainitzki, Heike; Stoneking, Mark; Pääbo, Svante (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.
^ Jump up to: 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.
Jump up ^ Foley, Tim. TalkOrigins Archive. "Neanderthal or Neandertal?". 2005.
Jump up ^ 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.
Jump up ^ Inter alia, Boys' Life, p. 18. January 1924.
Jump up ^ The Oxford Illustrated Dictionary. Great Britain: Oxford University Press. 1976 [1975]. p. 564. (tahl)
Jump up ^ "Neanderthal adjective – definition in British English Dictionary & Thesaurus". Dictionary.cambridge.org. January 8, 2013. Retrieved January 22, 2013.
Jump up ^ "Oxford Learner's Dictionaries – Find pronunciation, clear meanings and definitions of words". OxfordLearnersDictionaries.com. [dead link]
Jump up ^ "Neanderthal | Define Neanderthal at Dictionary.com". Dictionary.reference.com. Retrieved January 22, 2013.
Jump up ^ Kurtén, Björn (October 10, 1995). Dance of the Tiger: A Novel of the Ice Age. University of California Press. p. xxi. ISBN 0-520-20277-5. Retrieved May 9, 2012.
Jump up ^ Pollet, Carl J. (September 21, 1991). "…And Etymology". Science News. 140 (12): 191. doi:10.2307/3975867. JSTOR 3975867.
Jump up ^ 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): 7117–19. Bibcode:1999PNAS...96.7117T. doi:10.1073/pnas.96.13.7117. JSTOR 48019. PMC 33580  . PMID 10377375.
^ Jump up to: a b Duarte, C.; Mauricio, J.; Pettitt, P. B. (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): 7604–09. Bibcode:1999PNAS...96.7604D. doi:10.1073/pnas.96.13.7604. ISSN 0027-8424. PMID 10377462. Retrieved June 21, 2009.
Jump up ^ Pääbo, Svante (2014). Neanderthal Man: In Search of Lost Genomes. New York: Basic Books. p. 237.
Jump up ^ "L'homme fossile de La Chapelle-aux-Saints – full text: Volume VI (pp. 11–172), Volume VII (pp. 21–56), Volume VIII (pp. 1–70), 1911–13". Royal College of Surgeons of England. Retrieved July 26, 2016.
Jump up ^ "Marcellin Boule – French geologist". Encyclopædia Britannica. Retrieved July 26, 2016.
Jump up ^ "Arthur Keith". Royal Anthropological Institute. Retrieved July 26, 2016.
Jump up ^ "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.
Jump up ^ "Our Neandertal Brethren: Why They Were Not a Separate Species". Scientific American. August 1, 2010. Retrieved July 26, 2016.
Jump up ^ 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.
Jump up ^ Rightmire, G.P. (2001). "Patterns of hominid evolution and dispersal in the Middle Pleistocene". Quaternary International. 75 (1): 77–84. Bibcode:2001QuInt..75...77R. doi:10.1016/S1040-6182(00)00079-3.
Jump up ^ Toro-Moyano, I. (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. PMID 23481345.
Jump up ^ Wayman, Erin (November 26, 2012). "Homo antecessor: Common Ancestor of Humans and Neanderthals?". Smithsonian Institution. Retrieved May 17, 2017.
^ Jump up to: a b c d e f K. Prüfer; et.al. (2014). "The complete genome sequence of a Neanderthal from the Altai Mountains". Nature. 505 (7481): 43–49. Bibcode:2014Natur.505...43P. doi:10.1038/nature12886. PMC 4031459  . PMID 24352235.
Jump up ^ M. Krings (1999). "DNA sequence of the mitochondrial hypervariable region II from the Neandertal type specimen". Proceedings of the National Academy of Sciences USA. 96 (10): 5581–85. Bibcode:1999PNAS...96.5581K. doi:10.1073/pnas.96.10.5581.
Jump up ^ P. Beerli; S.V. Edwards (2002). "When did Neanderthals and modern humans diverge?" (PDF). Evolutionary Anthropology. 11 (S1): 60–63. doi:10.1002/evan.10058.
^ Jump up to: a b Ovchinnikov, Igor V.; Götherström, Anders; Romanova, Galina P.; Kharitonov, Vitaliy M.; Lidén, Kerstin; Goodwin, William (2000). "Molecular analysis of Neanderthal DNA from the northern Caucasus". Nature. 404 (6777): 490–93. Bibcode:2000Natur.404..490O. doi:10.1038/35006625. ISSN 0028-0836. PMID 10761915.
^ Jump up to: a b 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. ISSN 0092-8674. PMC 2602844 Freely accessible. PMID 18692465. doi:10.1016/j.cell.2008.06.021.
Jump up ^ R.E. Green; et. al. (2009). "Targeted Retrieval and Analysis of Five Neandertal mtDNA Genomes" (PDF). Science. 325 (5938): 318–21. Bibcode:2009Sci...325..318B. doi:10.1126/science.1174462.
Jump up ^ 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" (PDF). Journal of Human Evolution. 59 (1): 87–95. doi:10.1016/j.jhevol.2010.04.005. PMID 20510437.
Jump up ^ A. Rieux (2014). "Improved calibration of the human mitochondrial clock using ancient genomes". Molecular Biology and Evolution. 31 (10): 2780–92. doi:10.1093/molbev/msu222. PMC 4166928  . PMID 25100861.
Jump up ^ Cookson, Clive (June 27, 2014). "Palaeontology: How Neanderthals evolved". Financial Times. Retrieved October 28, 2015.
^ Jump up to: a b Bischoff, James L.; Shamp, et.al (2003). "The Sima de los Huesos Hominids Date to Beyond U/Th Equilibrium (>350kyr) and Perhaps to 400–500kyr: New Radiometric Dates" (PDF). Journal of Archaeological Science. 30 (3): 275–80. doi:10.1006/jasc.2002.0834.
Jump up ^ Stringer, Chris (2011). The Origin of our Species. Penguin. pp. 26–29, 202. ISBN 978-0-141-03720-2.
Jump up ^ Johansson, Donald; Edgar, Blake (2006). From Lucy to Language. Simon & Schuster. p. 38. ISBN 978-0-7432-8064-8.
^ Jump up to: a b c D. Dean; J.-J. Hublin; R. Holloway; R. Ziegler (1998). "On the phylogenetic position of the pre-Neandertal specimen from Reilingen, Germany". Journal of Human Evolution. 34 (5). pp. 485–508. doi:10.1006/jhev.1998.0214.
^ Jump up to: a b c d e Papagianni, Dmitra; Morse, Michael (2013). The Neanderthals Rediscovered. Thames & Hudson. ISBN 978-0-500-05177-1.
^ Jump up to: a b c Stringer, C.; Gamble, C. (1993). In Search of the Neanderthals. London: Thames and Hudson. ISBN 978-0500050705.
Jump up ^ B. Vandermeersch; M.D. Garralda (2011). S. Condemi; G.-C. Weniger, eds. "Continuity or Discontinuity in the Peopling of Europe: One Hundred and Fifty Years of Neanderthal Study". Vertebrate Paleobiology and Paleoanthropology. Springer Netherlands. pp. 113–25. doi:10.1007/978-94-007-0492-3_10.
Jump up ^ N.J. Conard; J. Richter, eds. (2011). "2". Neanderthal Lifeways, Subsistence and Technology. Vertebrate Paleobiology and Paleoanthropology. 19. Springer. pp. 7–14. doi:10.1007/978-94-007-0415-2_2. ISBN 978-94-007-0414-5.
Jump up ^ "Homo neanderthalensis". Smithsonian Institution. Archived from the original on May 21, 2009. Retrieved May 18, 2009.
Jump up ^ "New Evidence On The Role Of Climate In Neanderthal Extinction". Science Daily.
Jump up ^ S. Genovés (1954). "The problem of the sex of certain fossil hominids, with special reference to the Neandertal skeletons from Spy". The Journal of the Royal Anthropological Institute of Great Britain and Ireland. 84 (1/2). pp. 131–44.
Jump up ^ "Neanderthal Man". Encyclopedia Britannica (15th ed.). 1982.
Jump up ^ "Neanderthal – Homo neanderthalensis – Details – Key Fossils – La Chapelle-aux-Saints". Encyclopedia of Life. Retrieved July 18, 2016.
^ Jump up to: a b 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. 53: 649–57. doi:10.1016/j.jas.2014.04.019. CS1 maint: Uses authors parameter (link)
^ Jump up to: a b 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.
^ Jump up to: a b Gargett, R.H. (1989). "Grave Shortcomings: The Evidence for Neandertal Burial". Current Anthropology. 30 (2): 157–90. doi:10.1086/203725.
Jump up ^ "Universitat Pompeu Fabra, Barcelona, A paleogenetical study determines the blood group of Neanderthal man" Archived November 22, 2012, at the Wayback Machine.
Jump up ^ Lalueza-Fox, Carles; Rosas, Antonio; Rasilla, Marco de la (2012). "Palaeogenetic research at the El Sidrón Neanderthal site". Annals of Anatomy – Anatomischer Anzeiger. 194: 133–37. doi:10.1016/j.aanat.2011.01.014.
Jump up ^ 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): 13342–47. Bibcode:2002PNAS...9913342S. doi:10.1073/pnas.192464099. PMC 130635  . PMID 12232049.
^ Jump up to: a b Green, Richard E.; Krause, Johannes; Ptak, Susan E.; Briggs, Adrian W.; Ronan, Michael T.; Simons, Jan F.; Du, Lei; Egholm, Michael; Rothberg, Jonathan M.; Paunovic, Maja; Pääbo, Svante (2006). "Analysis of one million base pairs of Neanderthal DNA". Nature. 444 (7117): 330–36. Bibcode:2006Natur.444..330G. doi:10.1038/nature05336. ISSN 0028-0836. PMID 17108958. Retrieved 9 September 2017.
^ Jump up to: 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 5100745  . PMID 20448178.
Jump up ^ Sankararaman, S.; Patterson, N.; Li, H.; Pääbo, S.; Reich, D; Akey, J.M. (2012). "The Date of Interbreeding between Neandertals and Modern Humans". PLoS Genetics. 8 (10): e1002947. doi:10.1371/journal.pgen.1002947. PMC 3464203  . PMID 23055938.
Jump up ^ Yang, M.A.; Malaspinas, A.S.; Durand, E.Y.; Slatkin, M. (2012). "Ancient Structure in Africa Unlikely to Explain Neanderthal and Non-African Genetic Similarity". Molecular Biology and Evolution. 29 (10): 2987–95. doi:10.1093/molbev/mss117. PMC 3457770  . PMID 22513287.
^ Jump up to: 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. Bibcode:2014PNAS..111...81R. doi:10.1073/pnas.1316780110.
Jump up ^ P. Villa, W. Roebroeks (2014). "Neandertal Demise: An Archaeological Analysis of the Modern Human Superiority Complex". PLoS ONE. 9 (4): e96424. Bibcode:2014PLoSO...996424V. doi:10.1371/journal.pone.0096424. PMC 4005592  . PMID 24789039. CS1 maint: Uses authors parameter (link)
Jump up ^ "The Human Lineage by Matt Cartmill, Fred H. Smith". Google Books. Retrieved September 27, 2015.
Jump up ^ "Ancient tooth provides evidence of Neanderthal movement" (Press release). Durham University. February 11, 2008. Retrieved May 18, 2009.
Jump up ^ Wade, Nicholas (October 2, 2007). "Fossil DNA Expands Neanderthal Range". The New York Times. Retrieved May 18, 2009.
Jump up ^ Ravilious, Kate (October 1, 2007). "Neandertals Ranged Much Farther East Than Thought". National Geographic Society. Retrieved May 18, 2009.
Jump up ^ 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.
Jump up ^ O'Neill, Dennis. "Evolution of Modern Humans: Neanderthals", Palomar College, June 10, 2011. Retrieved August 21, 2011.
Jump up ^ Puech, Pierre-François; Puech, Bernard. "L'Homme de Neanderthal par Paul Dardé : L'Homme Primitif" [Neanderthal Man by Paul Dardé: Primitive Man]. Academia.edu (in French). Retrieved July 5, 2017.
Jump up ^ "Science & Nature – Wildfacts – Neanderthal". BBC. Archived from the original on January 15, 2009. Retrieved June 21, 2009.
Jump up ^ "Neanderthal". BBC. Archived from the original on January 15, 2009. Retrieved May 18, 2009.
^ Jump up to: a b Froehle, Andrew W; Churchill, Steven E (2009). "Energetic Competition Between Neandertals and Anatomically Modern Humans" (PDF). PaleoAnthropology: 96–116. Retrieved October 31, 2011.
^ Jump up to: a b Lalueza-Fox, C.; Rompler, H.; Caramelli, D.; Staubert, C.; Catalano, G.; Hughes, D.; Rohland, N.; Pilli, E.; Longo, L.; Condemi, S.; de la Rasilla, M.; Fortea, J.; Rosas, A.; Stoneking, M.; Schoneberg, T.; Bertranpetit, J.; Hofreiter, M. (2007). "A Melanocortin 1 Receptor Allele Suggests Varying Pigmentation Among Neanderthals" (PDF). Science. 318 (5855): 1453–55. Bibcode:2007Sci...318.1453L. doi:10.1126/science.1147417. ISSN 0036-8075. PMID 17962522. Retrieved September 10, 2017. ; see also Rincon, Paul (October 25, 2007). "Neanderthals 'were flame-haired'". BBC News. Retrieved October 25, 2007.
Jump up ^ The 10,000 Year Explosion: How Civilizations Accelerated Human Evolution (2009). Gregory Cochran and Henry Harpending. Basic Books, New York, NY
Jump up ^ 14:186–98. Hoffecker JF (2009) The spread of modern humans in Europe. Proc. Nat. Acad. Sci. 106:16040– References 399 Hoffecker JF, Cleghorn ...
Jump up ^ "Neanderthal brains focused on vision and movement leaving less room for social networking". Science Daily. March 19, 2013.
Jump up ^ SINC Servicio de Información y Noticias Científicas. "El cerebro neandertal era más asimétrico que el del 'Homo sapiens'".
Jump up ^ "Neanderthal Brain Size at Birth Sheds Light on Human Evolution". National Geographic. September 9, 2008. Retrieved September 19, 2009.
Jump up ^ 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."
Jump up ^ Abramiuk, Marc. The Foundations of Cognitive Archaeology, p. 199 (MIT Press 2012): "the encephalization quotient was slightly smaller".
Jump up ^ Moskvitch, Katia (September 24, 2010). "Neanderthals were able to 'develop their own tools'". BBC News. BBC. Retrieved October 1, 2010.
Jump up ^ Heyes, Peter; Anastasakis, Konstantinos; de Jong, Wiebren (2016). "Selection and Use of Manganese Dioxide by Neanderthals". Scientific Reports. 6: 22159. Bibcode:2016NatSR...622159H. doi:10.1038/srep22159. ISSN 2045-2322. PMC 4770591  . PMID 26922901.
Jump up ^ 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.
^ Jump up to: a b Ghosh, Pallab. "Neanderthals cooked and ate vegetables." BBC News. December 27, 2010.
Jump up ^ Lichfield, John (September 30, 2006). "French dig up Neanderthal 'butcher's shop'". The New Zealand Herald.
^ Jump up to: 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): 7663–66. Bibcode:2000pnas...97.7663r. doi:10.1073/pnas.120178997. JSTOR 122870. PMC 16602  . PMID 10852955.
Jump up ^ 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. Bibcode:2011PLoSO...614769F. doi:10.1371/journal.pone.0014769. PMC 3060801  . PMID 21445243.
Jump up ^ 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): 486–91. Bibcode:2011PNAS..108..486H. doi:10.1073/pnas.1016868108. PMC 3021051  . PMID 21187393.
Jump up ^ Webb, Jonathan (June 25, 2014). "Oldest human faeces show Neanderthals ate vegetables". BBC News.
Jump up ^ "Neanderthal behaviour, diet, and disease inferred from ancient DNA in dental calculus : Nature". The Guardian. Retrieved March 9, 2017.
Jump up ^ Nicola Davis (March 8, 2017). "Neanderthal dental tartar reveals plant-based diet – and drugs". The Guardian. Retrieved March 9, 2017.
Jump up ^ Shaw, Kate (July 29, 2011). "Sheer Numbers Gave Early Humans Edge Over Neanderthals". Wired.com.
Jump up ^ Vergano, Dan (April 22, 2014). "Neanderthals Lived in Small, Isolated Populations, Gene Analysis Shows". National Geographic.
^ Jump up to: 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.
Jump up ^ Zollikofer, Christoph; Marcia, Ponce; Leon, De; Vandermeersch, Bernard; Leveque, Francois (2002). "Evidence for Interpersonal Violence in the St. Césaire Neanderthal". PNAS. 99 (9): 6444–48. Bibcode:2002PNAS...99.6444Z. doi:10.1073/pnas.082111899. PMC 122968  . PMID 11972028.
Jump up ^ 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 (1): 27–90. doi:10.1006/jhev.1999.0301. PMID 10375476.
Jump up ^ Higham T, Jacobi R, Julien M, David F, Basell L, Wood R, Davies W, Ramsey CB.C (2010). "Chronology of the Grotte du Renne (France) and implications for the context of ornaments and human remains within the Chatelperronian". Proc Natl Acad Sci USA. doi:10.1073/pnas.1007963107 PMID 20956292
Jump up ^ Mellars P. (2010). "Neanderthal symbolism and ornament manufacture: The bursting of a bubble?" Proc Natl Acad Sci USA doi:10.1073/pnas.1014588107
Jump up ^ J.-J. Hublin; S. Talamo; M. Julien; F. David; N. Connet; P. Bodu; B. Vandermeersch; M.P. Richards. "Radiocarbon dates from the Grotte du Renne and Saint-Césaire support a Neandertal origin for the Châtelperronian". Proceedings of the National Academy of Sciences USA. 109 (46). doi:10.1073/pnas.1212924109.
Jump up ^ F. Welkera; M. Hajdinjak; S. Talamo; K. Jaouen; M. Dannemann; F. David; M. Julien; M. Meyer; J. Kelso; I. Barnes; S. Brace; P. Kamminga; R. Fischer; B.M. Kessler; J.R. Stewart; S. Pääbo; M.J. Collins; J.-J. Hublin. "Palaeoproteomic evidence identifies archaic hominins associated with the Châtelperronian at the Grotte du Renne". Proceedings of the National Academy of Sciences USA. 113 (40). pp. 11, 162–11, 167. doi:10.1073/pnas.1605834113.
Jump up ^ "Neanderthals were not inferior to modern humans, study finds". ScienceDaily. 30 April 2014.
Jump up ^ E. de Lazaro (18 January 2017). "Neanderthals Capable of Incorporating Symbolic Objects into Their Culture, Discovery Suggests". Sci News.
Jump up ^ C.Q. Choi (8 January 2010). "Heavy Brows, High Art?: Newly Unearthed Painted Shells Show Neandertals Were Homo sapiens' Mental Equals". Scientific American.
Jump up ^ I. Sample (30 April 2014). "Neanderthals were not less intelligent than modern humans, scientists find". The Guardian.
Jump up ^ N. Branan (2010). "Neandertal Symbolism: Evidence Suggests a Biological Basis for Symbolic Thought". Scientific American.
Jump up ^ R. S. Solecki (1975). "Shanidar IV, a Neanderthal Flower Burial in Northern Iraq, Science". 190 (4217): 880–81.
Jump up ^ D.J. Sommer (1999). "The Shanidar IV 'Flower Burial': a Re-evaluation of Neanderthal Burial Ritual". Cambridge Archaeological Journal. 9 (1): 127–29.
Jump up ^ Paul B. Pettitt (2002). "The Neanderthal Dead, exploring mortuary variability in Middle Paleolithic Eurasia". Before Farming. 1 (4).
Jump up ^ "Neanderthal 'make-up' discovered". BBC News. 9 January 2010. Retrieved 20 May 2010.
Jump up ^ "Did Neanderthals use feathers for fashion?". New Scientist. Retrieved 16 June 2017.
Jump up ^ Finlayson, Clive; Brown, Kimberly; Blasco, Ruth; Rosell, Jordi; Negro, Juan José; Bortolotti, Gary R; Finlayson, Geraldine; Sánchez Marco, Antonio; Giles Pacheco, Francisco; Rodríguez Vidal, Joaquín; Carrión, José S; Fa, Darren A; Rodríguez Llanes, José M (17 September 2012). "Birds of a Feather: Neanderthal Exploitation of Raptors and Corvids". Plos One. 7 (9): e45927. Bibcode:2012PLoSO...745927F. doi:10.1371/journal.pone.0045927.
Jump up ^ E. Callaway (2014). "Neanderthals made some of Europe's oldest art". Nature News. doi:10.1038/nature.2014.15805.
Jump up ^ P. Rincon (1 September 2014). "Neanderthal 'artwork' found in Gibraltar cave". BBC.
Jump up ^ 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 (2 June 2016) [online 25 May 2016]. "Early Neanderthal Constructions deep in Bruniquel Cave in Southwestern France". Nature. 534 (7605): 111–14. Bibcode:2016Natur.534..111J. doi:10.1038/nature18291. ISSN 0028-0836. PMID 27251286.
Jump up ^ Brown, Cynthia Stokes. Big History. New York, NY: The New Press, 2008. Print.
Jump up ^ Moulson, Geir; Associated Press (July 20, 2006). "Neanderthal genome project launches". MSNBC. Retrieved August 22, 2006.
Jump up ^ 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)."
Jump up ^ Wade, Nicholas (November 15, 2006). "New Machine Sheds Light on DNA of Neanderthals". The New York Times. Retrieved May 18, 2009.
Jump up ^ 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.
Jump up ^ "Neanderthal bone gives DNA clues". CNN. Associated Press. November 16, 2006. Archived from the original on November 18, 2006. Retrieved May 18, 2009.
Jump up ^ Than, Ker; LiveScience (November 15, 2006). "Scientists decode Neanderthal genes". MSNBC. Retrieved May 18, 2009.
Jump up ^ "Studies find Neanderthal genes in modern humans".
Jump up ^ Zimmer, Carl (December 18, 2013). "Toe Fossil Provides Complete Neanderthal Genome". New York Times. Retrieved December 18, 2013.
Jump up ^ "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.
^ Jump up to: 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.
^ Jump up to: a b Elizabeth Pennisi (2009). "Tales of a Prehistoric Human Genome". Science. 323 (5916): 866–71. doi:10.1126/science.323.5916.866. PMID 19213888.
Jump up ^ 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 2725275  . PMID 19661919.
^ Jump up to: a b "Surprise! 20 Percent of Neanderthal Genome Lives On in Modern Humans, Scientists Find". National Geographic. Retrieved October 7, 2016.
^ Jump up to: a b c "New studies reveal 20 Percent of Neanderthal genome lives on in modern humans". Retrieved October 7, 2016.
Jump up ^ "Ancient DNA and Neanderthals". The Smithsonian Institution's Human Origins Program.
Jump up ^ "Science/Nature – Neanderthals 'were flame-haired'". BBC News.
Jump up ^ Neanderthals did not interbreed with humans, scientists find Archived April 19, 2014, at the Wayback Machine.. Telegraph. Retrieved May 24, 2014.
Jump up ^ Press Association (February 4, 2013). "Neanderthals 'unlikely to have interbred with human ancestors'". The Guardian. Retrieved May 24, 2014.
^ Jump up to: 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.
Jump up ^ Neanderthal breeding idea doubted. BBC. Retrieved March 2, 2016.
Jump up ^ Pratas, D; Hosseini, M; Silva, R; Pinho, A; Ferreira, P (June 20–23, 2017). "Visualization of Distinct DNA Regions of the Modern Human Relatively to a Neanderthal Genome". Iberian Conference on Pattern Recognition and Image Analysis. Springer. Lecture Notes in Computer Science. 10255: 235–42. doi:10.1007/978-3-319-58838-4_26. ISBN 978-3-319-58837-7.
^ Jump up to: a b Vernot, Benjamin; Akey, Joshua M (2015). "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. PMID 25683122.
Jump up ^ Kim, BY; Lohmueller, KE (2015). "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 4375557  . PMID 25683122.
^ Jump up to: a b [1]. The Daily Mail. Retrieved January 26, 2016.
Jump up ^ [2]
^ Jump up to: a b c Zimmer, Carl (March 17, 2016). "Humans Interbred With Hominins on Multiple Occasions, Study Finds". The New York Times. Retrieved March 17, 2016.
Jump up ^ 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): 728–34. doi:10.1016/j.ajhg.2016.02.023. PMC 4833433  . PMID 27058445. Retrieved April 12, 2016.
Jump up ^ "DNA points to Neanderthal breeding barrier". BBC News, Science & Environment. April 8, 2016. Retrieved April 12, 2016.
^ Jump up to: a b Sample, Ian (February 17, 2016). "Oldest known case of Neanderthal-human sex revealed by DNA test". the Guardian. Retrieved May 10, 2016.
^ Jump up to: 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): 523–27. Bibcode:2014Sci...344..523G. doi:10.1126/science.1250368. PMID 24786081.
Jump up ^ 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)
^ Jump up to: 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. CS1 maint: Uses authors parameter (link)
Jump up ^ "First genocide of human beings occurred 30,000 years ago". Pravda. October 24, 2007. Retrieved May 18, 2009.
Jump up ^ 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.
Jump up ^ 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 532389  . PMID 15562317.
Jump up ^ Wong, Kate (August 1, 2009). "The Mysterious Downfall of the Neandertals". Scientific American.
Jump up ^ 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.
Jump up ^ Wilford, John Noble (November 2, 2011). "Fossil Teeth Put Humans in Europe Earlier Than Thought". New York Times. Retrieved August 27, 2014.
Jump up ^ Wolff, H. "Result Filters." National Center for Biotechnology Information. U.S. National Library of Medicine, July 2010. Web. October 22, 2014.
Jump up ^ McKie, Robin (March 1, 2015). "How hunting with wolves helped humans outsmart the Neanderthals". The Guardian. Retrieved May 5, 2016.
Jump up ^ Stringer, Chris (2012). "Evolution: What makes a modern human". Nature. 485 (7396): 33–35. Bibcode:2012Natur.485...33S. doi:10.1038/485033a. PMID 22552077.
Jump up ^ Steensby, H. P. (1907). "Racestudier i Danmark" [Race Studies In Denmark] (PDF). Geographical Journal (in Danish). Royal Library, Denmark. Retrieved July 6, 2017.
Jump up ^ 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.
Jump up ^ Foley, Jim (July 31, 2000). "The Lagar Velho 1 Skeleton". Fossil Hominids FAQ. TalkOrigins Archive. Retrieved July 6, 2017.
Jump up ^ Sample, Ian (September 13, 2006). "Life on the edge: was a Gibraltar cave last outpost of the lost neanderthal?". The Guardian. Retrieved July 6, 2017.
Jump up ^ "Not a lasting last for the Neandertals". john hawks weblog. September 13, 2006. Retrieved July 6, 2017.
Jump up ^ 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 1859909  . PMID 17085588.
Jump up ^ Wade, Nicholas (July 26, 2012). "Genetic Data and Fossil Evidence Tell Differing Tales of Human Origins". The New York Times. Retrieved May 5, 2015.
Jump up ^ 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.
Jump up ^ Viegas, Jennifer (July 18, 2011). "All Non-Africans Part Neanderthal, Genetics Confirm". DNews.
Jump up ^ "Neandertal ancestry "Iced"". john hawks weblog. August 15, 2012. Retrieved July 6, 2017.
Jump up ^ "Resurrecting Surviving Neandertal Lineages from Modern Human Genomes". Science. January 29, 2014. Retrieved February 2, 2014.
Jump up ^ Mitchell, Alanna (January 30, 2012). "DNA Turning Human Story Into a Tell-All". NYTimes. Retrieved January 31, 2012.
Jump up ^ 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:1307.8263  . Bibcode:2013arXiv1307.8263L.
Jump up ^ Jha, Alok (August 14, 2012). "Study casts doubt on human-Neanderthal interbreeding theory". The Guardian. Retrieved February 19, 2015.
Jump up ^ 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.
Jump up ^ Mason, Paul H.; Short, Roger V. (2011). "Neanderthal-human Hybrids". Hypothesis. 9: e1. doi:10.5779/hypothesis.v9i1.215.
Jump up ^ Viegas, Jennifer (March 27, 2013). "First Love Child of Human, Neanderthal Found". Discovery.
Jump up ^ 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. Bibcode:2013PLoSO...859781C. doi:10.1371/journal.pone.0059781. PMC 3609795  . PMID 23544098.
Jump up ^ The Neanderthal Dead, exploring mortuary variability in middle paleolithic eurasia. Paul B. Pettitt (2002)
Jump up ^ "Homo neanderthalensis – The Neanderthals". Australian Museum. Retrieved June 26, 2014.
Jump up ^ 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.
Jump up ^ Kreger, C. David. "Homo neanderthalensis". ArchaeologyInfo.com. Retrieved May 16, 2009.
Jump up ^ 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): 252–55. Bibcode:1993Natur.363..252M. doi:10.1038/363252a0. ISSN 0028-0836. PMID 8387643.
Jump up ^ 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): 701–17. doi:10.1002/gea.1017.
Jump up ^ 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): 259–68. doi:10.1006/jasc.1998.0334.
Jump up ^ 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): 148–58. doi:10.1111/j.1475-4754.2012.00671.x. CS1 maint: Uses authors parameter (link)
Jump up ^ Hayes, Jacqui (November 2, 2006). "Humans and Neanderthals interbred". Cosmos. Archived from the original on February 22, 2009. Retrieved May 17, 2009.
Jump up ^ Neanderthal image by Kupka, based on Boule, 1909, in Humanity's Journeys Dr. Kathryn Denning, 2005. Retrieved March 17, 2012.
Journals
Boë, Louis-Jean; Heim, Jean-Louis; Honda, Kiyoshi; Maeda, Shinji (2002). "The potential Neandertal vowel space was as large as that of modern humans". Journal of Phonetics. 30 (3): 465–84. doi:10.1006/jpho.2002.0170.
Lieberman, Philip (October 2007). "Current views on Neanderthal speech capabilities: A reply to Boe et al. (2002)". Journal of Phonetics. 35 (4): 552–63. doi:10.1016/j.wocn.2005.07.002.
Serre, David; Langaney, André; Chech, Mario; Teschler-Nicola, Maria; Paunovic, Maja; Mennecier, Philippe; Hofreiter, Michael; Possnert, Göran; Pääbo, Svante (2004). "No Evidence of Neandertal mtDNA Contribution to Early Modern Humans". PLoS Biology. 2 (3): e57. doi:10.1371/journal.pbio.0020057. PMC 368159  . PMID 15024415.
Wild, Eva M.; Teschler-Nicola, Maria; Kutschera, Walter; Steier, Peter; Trinkaus, Erik; Wanek, Wolfgang (2005). "Direct dating of Early Upper Palaeolithic human remains from Mladeč". Nature. 435 (7040): 332–35. Bibcode:2005Natur.435..332W. doi:10.1038/nature03585. PMID 15902255.
Zilhão, João; Davis, Simon J. M.; Duarte, Cidália; Soares, António M. M.; Steier, Peter; Wild, Eva (2010). Hawks, John, ed. "Pego do Diabo (Loures, Portugal): Dating the Emergence of Anatomical Modernity in Westernmost Eurasia". PLoS ONE. 5 (1): e8880. Bibcode:2010PLoSO...5.8880Z. doi:10.1371/journal.pone.0008880. PMC 2811729  . PMID 20111705. Lay summary – ScienceDaily (January 27, 2010).
Bibliography
Derev'anko, Anatoliy P.; Powers, William Roger; Shimkin, Demitri Boris (1998). The Paleolithic of Siberia: new discoveries and interpretations. Novosibirsk: Institute of Anthropology and Ethnography. ISBN 978-0-252-02052-0. OCLC 36461622.
Lunine, Jonathan I. (2013). Earth: Evolution of a Habitable World. Cambridge University Press. 327. ISBN 978-0-521-85001-8.
Further reading[edit]
Sankararaman, Sriram; Mallick, Swapan; Dannemann, Michael; Prüfer, Kay; Kelso, Janet; Patterson, Nick; Reich, David (2014). "The genomic landscape of Neanderthal ancestry in present-day humans". Nature. 507 (7492): 354–57. Bibcode:2014Natur.507..354S. doi:10.1038/nature12961. PMC 4072735  . PMID 24476815.
Vattathil, S.; Akey, J.M. (2015). "Small amounts of archaic admixture provide big insights into human history". Cell. 163 (2): 281–84. doi:10.1016/j.cell.2015.09.042. PMID 26451479.
External links[edit]
Wikimedia Commons has media related to Homo neanderthalensis.
Wikibooks has a book on the topic of: Introduction to Paleoanthropology
Wikispecies has information related to: Homo neanderthalensis
Wikisource has the text of the 1905 New International Encyclopedia article Neanderthal Man.
Kreger, C. David (June 30, 2000). "Homo neanderthalensis". ArchaeologyInfo.com. Retrieved May 23, 2009.
O'Neil, Dennis (May 12, 2009). "Evolution of Modern Humans: Neandertals". Retrieved May 23, 2009.
New Portuguese skull may be an early relative of Neandertals – article by Ann Gibbons at Science, March 13, 2017.
In Neanderthal DNA, Signs of a Mysterious Human Migration – article by Carl Zimmer, NY Times, July 4, 2017
"Homo neanderthalensis". The Smithsonian Institution.
"Neanderthal DNA". International Society of Genetic Genealogy. Archived from the original on June 17, 2006. : Includes Neanderthal mtDNA sequences
Panoramio – The Neandertal foot prints' (photo of ≈25K years old fossilized footprints discovered in 1970 on volcanic layers near Demirkopru Dam Reservoir, Manisa, Turkey)
Did better mothering defeat the Neanderthals?
My Great-great-great Grandfather's a Neanderthal
Ancient tryst fortified human immune system
Neanderthal-human hybridisation hypothesis
Neanderthal hybridization and Haldane's rule
Neanderthal Studies Professional Online Service (NESPOS) fossil overview
Human Timeline (Interactive) – Smithsonian, National Museum of Natural History (August 2016).
[show]
v
t
e
Prehistoric technology
Prehistory
timeline
outline
Stone Age
subdivisions
New Stone Age
synoptic table
Technology
history
[show]
Tools
Farming
Neolithic Revolution
founder crops
New World crops
Ard / plough
Celt
Digging stick
Domestication
Goad
Irrigation
Secondary products
Sickle
Terracing
Food processing
Fire
Basket
Cooking
Earth oven
Granaries
Grinding slab
Ground stone
Hearth
Aşıklı Höyük
Qesem Cave
Manos
Metate
Mortar and pestle
Pottery
Quern-stone
Storage pit
Hunting
Arrow
Boomerang
throwing stick
Bow and arrow
history
Nets
Spear
Spear-thrower
baton
harpoon
woomera
Schöningen Spears
Projectile points
Arrowhead
Bare Island
Cascade
Clovis
Cresswell
Cumberland
Eden
Folsom
Lamoka
Manis Site
Plano
Transverse arrowhead
Systems
Game drive system
Buffalo jump
Toolmaking
Earliest toolmaking
Oldowan
Acheulean
Mousterian
Clovis culture
Cupstone
Fire hardening
Gravettian culture
Hafting
Hand axe
Grooves
Langdale axe industry
Levallois technique
Lithic core
Lithic reduction
analysis
debitage
flake
Lithic technology
Magdalenian culture
Metallurgy
Microblade technology
Mining
Prepared-core technique
Solutrean industry
Striking platform
Tool stone
Uniface
Yubetsu technique
Other tools
Adze
Awl
bone
Axe
Bannerstone
Blade
prismatic
Bone tool
Bow drill
Burin
Canoe
Oar
Pesse canoe
Chopper
tool
Cleaver
Denticulate tool
Fire plough
Fire-saw
Hammerstone
Knife
Microlith
Quern-stone
Racloir
Rope
Scraper
side
Stone tool
Tally stick
Weapons
Wheel
illustration
[show]
Architecture
Ceremonial
Göbekli Tepe
Kiva
Standing stones
megalith
row
Stonehenge
Pyramid
Dwellings
Neolithic architecture
British megalith architecture
Nordic megalith architecture
Burdei
Cave
Cliff dwelling
Dugout
Hut
Quiggly hole
Jacal
Longhouse
Mud brick
Mehrgarh
Neolithic long house
Pit-house
Pueblitos
Pueblo
Rock shelter
Blombos Cave
Abri de la Madeleine
Sibudu Cave
Stone roof
Roundhouse
Stilt house
Alp pile dwellings
Wattle and daub
Water management
Check dam
Cistern
Flush toilet
Reservoir
Water well
Other architecture
Archaeological features
Broch
Burnt mound
fulacht fiadh
Causewayed enclosure
Tor enclosure
Circular enclosure
Goseck
Cursus
Henge
Thornborough
Oldest buildings
Megalithic architectural elements
Midden
Timber circle
Timber trackway
Sweet Track
[show]
Arts and culture
Material goods
Baskets
Beadwork
Beds
Chalcolithic
Clothing/textiles
timeline
Cosmetics
Glue
Hides
shoes
Ötzi
Jewelry
amber use
Mirrors
Pottery
Cardium
Grooved ware
Linear
Jōmon
Unstan ware
Sewing needle
Weaving
Wine
Winery
wine press
PrehistArt
Art of the Upper Paleolithic
Art of the Middle Paleolithic
Blombos Cave
List of Stone Age art
Bird stone
Bradshaw rock paintings
Cairn
Carved Stone Balls
Cave paintings
painting
pigment
Cup and ring mark
Geoglyph
Golden hats
Guardian stones
Megalithic art
Petroform
Petroglyph
Petrosomatoglyph
Pictogram
Rock art
Stone carving
Sculpture
Statue menhir
Stone circle
list
British Isles and Brittany
Venus figurines
Burial
Burial mounds
Bowl barrow
Round barrow
Mound Builders culture
U.S. sites
Chamber tomb
Severn-Cotswold
Cist
Dartmoor kistvaens
Clava cairn
Court tomb
Cremation
Dolmen
Great dolmen
Funeral pyre
Gallery grave
transepted
wedge-shaped
Grave goods
Jar burial
Long barrow
unchambered
Grønsalen
Megalithic tomb
Mummy
Passage grave
Rectangular dolmen
Ring cairn
Simple dolmen
Stone box grave
Tor cairn
Tumulus
Unchambered long cairn
Other cultural
Astronomy
sites
lunar calendar
Behavioral modernity
Origin of language
Prehistoric medicine
trepanning
Evolutionary musicology
music archaeology
Prehistoric music
Alligator drum
flutes
Divje Babe flute
gudi
Prehistoric numerals
Origin of religion
Paleolithic religion
Prehistoric religion
Spiritual drug use
Prehistoric warfare
Symbols
symbolism
[show]
v
t
e
Human evolution
Taxonomy
(Hominins)
Orrorin
Sahelanthropus
Last common ancestors
Chimpanzee–human
Gorilla–human
Orangutan–human
Gibbon–human
Australopithecines
Ardipithecus
Kenyanthropus
Australopithecus
A. afarensis
A. africanus
A. anamensis
A. bahrelghazali
A. deyiremeda
A. garhi
A. sediba
Paranthropus
P. aethiopicus
P. boisei
P. robustus
Humans and
proto-humans
(Homo)
Proto-humans
H. ergaster
H. floresiensis
H. gautengensis
H. habilis
H. rudolfensis
H. tsaichangensis
Homo erectus
H. e. erectus
H. e. georgicus
H. e. lantianensis
H. e. nankinensis
H. e. palaeojavanicus
H. e. pekinensis
H. e. soloensis
H. e. tautavelensis
H. e. yuanmouensis
Archaic humans
Callao Man
?
Denisovan
H. antecessor
H. cepranensis
H. heidelbergensis
H. helmei
H. naledi
H. neanderthalensis
H. rhodesiensis
Modern humans
Homo sapiens
Cro-Magnon
H. s. idaltu
H. s. sapiens (anatomically modern human)
Manot people
Red Deer Cave people
?
Models
General models
Hunting
Gathering
Endurance running
Aquatic ape
Sexual selection
Self-domestication
Specific models
Diet
Cooking
Expensive tissue
Shore-based
Drunken monkey
Behavior
Killer ape
Cooperative eye
Life history
Grandmother
Patriarch
Topics
Bipedalism
Skeleton
Muscles
Skin color
Hair
Thermoregulation
Color vision
Speech
Language
Intelligence
Gender roles
Origin of modern humans
Recent African origin
Multiregional origin
Behavioral modernity
Early migrations
Timelines
Human evolution
Human prehistory
Others
Theorists
Books
Fossils
Evolutionary anthropology
Evolutionary biology portal
[hide]
v
t
e
Neanderthal
Ancestors
Homo habilis → Homo erectus (→ Homo antecessor)? → Homo heidelbergensis
Biology
Origins (Accretion model)
Anatomy
Behaviour
Admixture with H. sapiens
Extinction
Tool traditions
Mousterian
Châtelperronian (controversial)
Regional variants
Gibraltar Neanderthals
Southwest Asian Neanderthals
Notable Specimens
Early (250–130 ka)
Ehringsdorf 9
Altamura 1
Typical
(130–50 ka)
La Ferrassie 1 (site)
La Chapelle-aux-Saints 1 (site)
La Quina 5 and 18
Krijn
Scladina
Saccopastore 1 and 2
Altamura man
Mt. Circeo 1
Krapina remains
Shanidar 1-4
Teshik-Tash 1
Kebara 2 (site)
Tabun C1
Amud 1 and 7 (site)
OR-1
In radiocarbon
range (50–40 ka)
Neanderthal 1 (type
site)
Spy 1 and 2
Engis 2
Sidrón remains
Gibraltar 1 and 2
Saint-Césaire 1
Le Moustier
Fontéchevade
Mezmaiskaya 1
Fossils with claimed
Neanderthal traits
H. heidelbergensis
Bontnewydd tooth
Atapuerca
Samu
Azykhantrop
Homo sapiens
Peștera cu Oase
Lapedo child
Contemporary species
Homo erectus
Denisovan
Homo floresiensis
Homo naledi
Homo sapiens
Research
Excavation sites
Neanderthal Museum
Neanderthal genome project (In Search of Lost Genomes)
Popular culture
The Quest for Fire (film)
The Grisly Folk
The Neanderthal Man
The Inheritors
The Ugly Little Boy
Neanderthal Planet
Dance of the Tiger
Earth's Children (film)
Neanderthal
The Neanderthal Parallax
Documentaries
Prehistoric Autopsy
Reliable sourcing guide for Palaeolithic articles
Taxon identifiers
Wd: Q40171
EoL: 4454114
Fossilworks: 83087
GBIF: 4827584
Authority control
GND: 4171356-4
NDL: 00568063
Anthropology portal
Evolutionary biology portal
Science portal
Retrieved from "https://en.wikipedia.org/w/index.php?title=Neanderthal&oldid=810130950"
Categories:
Neanderthals
Fossil taxa described in 1864
Stone Age Europe
Stone Age Asia
Pleistocene primates
Tool-using mammals
Hidden categories:
CS1 maint: Uses authors parameter
All articles with dead external links
Articles with dead external links from July 2016
Webarchive template wayback links
CS1 French-language sources (fr)
Pages containing links to subscription-only content
CS1 Danish-language sources (da)
Articles with dead external links from October 2017
Use mdy dates from May 2017
Articles with 'species' microformats
Wikipedia articles needing clarification from April 2017
All articles lacking reliable references
Articles lacking reliable references from October 2017
Articles lacking reliable references from August 2017
Wikipedia articles with GND identifiers
Navigation menu
Personal tools
Not logged in
Talk
Contributions
Create account
Log in
Namespaces
Article
Talk
Variants
Views
Read
Edit
View history
More
Search
Navigation
Main page
Contents
Featured content
Current events
Random article
Donate to Wikipedia
Wikipedia store
Interaction
Help
About Wikipedia
Community portal
Recent changes
Contact page
Tools
What links here
Related changes
Upload file
Special pages
Permanent link
Page information
Wikidata item
Cite this page
Print/export
Create a book
Download as PDF
Printable version
In other projects
Wikimedia Commons
Wikispecies
Languages
Afrikaans
Alemannisch
العربية
Asturianu
Azərbaycanca
تۆرکجه
বাংলা
Беларуская
Български
Boarisch
Bosanski
Brezhoneg
Català
Чӑвашла
Čeština
Cymraeg
Dansk
Deutsch
Eesti
Ελληνικά
Español
Esperanto
Euskara
فارسی
Français
Frysk
Gaeilge
Galego
한국어
Հայերեն
हिन्दी
Hrvatski
Bahasa Indonesia
Interlingua
Íslenska
Italiano
עברית
ქართული
Қазақша
Kiswahili
Кыргызча
ລາວ
Latina
Latviešu
Lëtzebuergesch
Lietuvių
Limburgs
Magyar
Македонски
മലയാളം
Bahasa Melayu
Nederlands
日本語
Norsk
Norsk nynorsk
Occitan
Oʻzbekcha/ўзбекча
ਪੰਜਾਬੀ
پنجابی
پښتو
Plattdüütsch
Polski
Português
Română
Русский
Саха тыла
Scots
Sicilianu
Simple English
Slovenčina
Slovenščina
Српски / srpski
Srpskohrvatski / српскохрватски
Suomi
Svenska
Tagalog
தமிழ்
Татарча/tatarça
తెలుగు
ไทย
Türkçe
Українська
اردو
Vepsän kel’
Tiếng Việt
Winaray
Yorùbá
粵語
Zazaki
中文
Edit links
This page was last edited on 13 November 2017, at 14:32.
Text is available under the Creative Commons Attribution-ShareAlike License; additional terms may apply. By using this site, you agree to the Terms of Use and Privacy Policy. Wikipedia® is a registered trademark of the Wikimedia Foundation, Inc., a non-profit organization.
Privacy policy
About Wikipedia
Disclaimers
Contact Wikipedia
Developers
Cookie statement
Mobile view
Enable previews

key: html

type: local

value: div

attrs: id class

values: mw-page-base noprint

values: mw-head-base noprint

value: a

attrs: id class lang

values: firstHeading firstHeading en

values: siteSub noprint

values: jump-to-nav mw-jump

attrs: href title

values: /wiki/Megaannum mw-redirect Megaannum

value: span

value: small

values: position:absolute; height:100%; text-align:center; background-color:rgb(129,170,114); left:36.892307692308px; width:18.818461538462px;

values: position:absolute; height:100%; text-align:center; background-color:rgb(0,169,138); left:55.710769230769px; width:14.08px;

values: position:absolute; height:100%; text-align:center; background-color:rgb(166,223,197); left:69.790769230769px; width:8.3261538461539px;

values: position:absolute; height:100%; text-align:center; background-color:rgb(221,150,81); left:78.116923076923px; width:20.409230769231px;

values: position:absolute; height:100%; text-align:center; background-color:rgb(63,174,173); left:98.526153846154px; width:20.307692307692px;

values: position:absolute; height:100%; text-align:center; background-color:rgb(247,88,60); left:118.83384615385px; width:15.907015384615px;

values: position:absolute; height:100%; text-align:center; background-color:rgb(153,78,150); left:134.74086153846px; width:17.126830769231px;

values: position:absolute; height:100%; text-align:center; background-color:rgb(0,187,231); left:151.86769230769px; width:19.055384615385px;

values: position:absolute; height:100%; text-align:center; background-color:rgb(111,200,107); left:170.92307692308px; width:26.738461538462px;

values: position:absolute; height:100%; text-align:center; background-color:rgb(254,161,99); left:197.66153846154px; width:14.543692307692px;

values: position:absolute; height:100%; text-align:center; background-color:rgb(254,221,45); left:212.20523076923px; width:6.9215384615385px;

values: position:absolute; height:100%; background-color:#666; width:1px; left:219px

values: margin:0 auto; line-height:0; clear:both; width:220px; padding:0px; height:8px; overflow:visible; background-color:transparent; position:relative; top:-4px; z-index:100; position:absolute; left:219.91538461538px; font-size:50% position:relative; left:-0.42em

values: margin: 4px auto 0; clear:both; width:220px; padding:0px; height:18px; overflow:visible; border:1px #666; border-style:solid none; position:relative; z-index:0; font-size:13px;

values: text-align: center; background-color: rgb(235,235,210)

attrs: alt src width height srcset data-file-width data-file-height

attrs: alt src width height data-file-width data-file-height

values: font-size:smaller; float:right; padding-right:0.4em; margin-left:-3em;

values: min-width:15em; text-align: center; background-color: rgb(235,235,210)

values: text-align: left; width: 200px; font-size: 100%

values: Archaic human admixture with modern humans

values:
/wiki/East_Asians
mw-redirect
East Asians

values: background-color:#77bb77; font-weight:bold; padding:0.2em; text-align:center; margin:0.6em 0 1.2em 0

values: You can edit this timeline. Please use the preview button before saving.

values: float:right; position:absolute; right:-1px; top:36em;

values: float:right; position:absolute; right:-1px; top:34.2em;

values: float:right; position:absolute; right:-1px; top:32.4em;

values: float:right; position:absolute; right:-1px; top:30.6em;

values: float:right; position:absolute; right:-1px; top:28.8em;

values: float:right; position:absolute; right:-1px; top:27em;

values: float:right; position:absolute; right:-1px; top:25.2em;

values: float:right; position:absolute; right:-1px; top:23.4em;

values: float:right; position:absolute; right:-1px; top:21.6em;

values: float:right; position:absolute; right:-1px; top:19.8em;

values: float:right; position:absolute; right:-1px; top:18em;

values: float:right; position:absolute; right:-1px; top:16.2em;

values: float:right; position:absolute; right:-1px; top:14.4em;

values: float:right; position:absolute; right:-1px; top:12.6em;

values: float:right; position:absolute; right:-1px; top:10.8em;

values: float:right; position:absolute; right:-1px; top:9em;

values: float:right; position:absolute; right:-1px; top:7.2em;

values: float:right; position:absolute; right:-1px; top:5.4em;

values: float:right; position:absolute; right:-1px; top:3.6em;

values: float:right; position:absolute; right:-1px; top:1.8em;

values: float:right; position:absolute; right:-1px; top:0em;

values: width:3.1em; position:relative; top:-0.82em; float:left; height:36em; padding:0px

values: width:1px; float:left; height:36em; padding:0; background-color:#242020

values: position:relative; align:center; vertical-align:middle; bottom:-12.96em;

values: font-size:100%; background-color:#ffa500; border-style: none; position:absolute; text-align:center; margin-top:10.08em; height:25.92em; margin-left:0em; width:10em; font-size:100%; position:relative; top:-6.16em; left:-0.3em;

values: position:relative; align:center; vertical-align:middle; bottom:-0.18em;

values: font-size:100%; background-color:#ffa500; border-style: none; position:absolute; text-align:center; margin-top:34.92em; height:0.36000000000001em; margin-left:1em; width:9em; font-size:100%; position:relative; top:-0.86em; left:-0.5em;

values: font-size:100%; background-color:#ffa500; border-style: none; position:absolute; text-align:center; margin-top:32.04em; height:0.36em; margin-left:1em; width:9em; font-size:100%; position:relative; top:-0.86em; left:-0.3em;

values: font-size:100%; background-color:#ffa500; border-style: none; position:absolute; text-align:center; margin-top:24.84em; height:0.36em; margin-left:1em; width:9em; font-size:100%; position:relative; top:-0.86em; left:-0.3em;

values: font-size:100%; background-color:#ffa500; border-style: none; position:absolute; text-align:center; margin-top:21.24em; height:0.36em; margin-left:1em; width:9em; font-size:100%; position:relative; top:-0.86em; left:-2em;

values: font-size:100%; background-color:#ffa500; border-style: none; position:absolute; text-align:center; margin-top:15.48em; height:0.36em; margin-left:1em; width:9em; font-size:100%; position:relative; top:-0.86em; left:-1em;

values: position:relative; align:center; vertical-align:middle; bottom:-4.32em;

values: font-size:100%; background-color:#ffa500; border-style: none; position:absolute; text-align:center; margin-top:4.32em; height:8.64em; margin-left:0em; width:10em; font-size:100%; position:relative; top:3.14em; left:0.4em;

values: position:relative; align:center; vertical-align:middle; bottom:-2.34em;

values: font-size:100%; background-color:#ffb732; border-style: none; position:absolute; text-align:center; margin-top:5.4em; height:4.68em; margin-left:1em; width:9em; font-size:100%; position:relative; top:-0.56em; left:0.4em;

values: position:relative; align:center; vertical-align:middle; bottom:-3.177em;

values: font-size:100%; background-color:#ffc966; border-style: none; position:absolute; text-align:center; margin-top:0.126em; height:6.354em; margin-left:2em; width:8em; font-size:100%; position:relative; top:0.14em; left:0.1em;

values: font-size:100%; background-color:#ffc966; border-style: none; position:absolute; text-align:center; margin-top:4.32em; height:1.08em; margin-left:1em; width:1em; font-size:100%; position:relative; top:-0.86em; left:0.0em;

values: position:relative; align:center; vertical-align:middle; bottom:-1.008em;

values: font-size:100%; background-color:#ffdb99; border-style: none; position:absolute; text-align:center; margin-top:0.144em; height:2.016em; margin-left:1em; width:9em; font-size:100%; position:relative; top:-0.56em; left:0.4em;

values: position:relative; align:center; vertical-align:middle; bottom:-0.351em;

values: font-size:100%; background-color:#ffff00; border-style: none; position:absolute; text-align:center; margin-top:0em; height:0.702em; margin-left:2em; width:8em; font-size:100%; position:relative; top:-0.86em; left:0.0em;

values: font-size:100%; background-color:#ffff00; border-style: none; position:absolute; text-align:center; margin-top:0em; height:0.126em; margin-left:0em; width:1em; font-size:100%; position:relative; top:-0.86em; left:0.0em;

values: font-size:100%; background-color:#ffff00; border-style: none; position:absolute; text-align:center; margin-top:0em; height:0.144em; margin-left:1em; width:1em; font-size:100%; position:relative; top:-0.86em; left:0.0em;

values: font-size:100%; width:100%; height:36em; padding:0px; float:left; width:10em; border:none; background-color:#ffc966;

values: width:auto; font-size:100%; position:absolute; text-align:center; margin-top:18em; text-size:100%; float:left; position:relative; text-align:left; top:-0.75em; left:0em; width:7em;

values: position:relative; text-size:100%; top:-0.86em; left:0em; float:left;

values: position:relative; width:auto; z-index:10; font-size:90%; color:black; vertical-align:middle; line-height:100%; bottom:0;

values: text-size:100%; float:left; position:relative; text-align:left; top:-0.75em; left:0em; width:7em; line-height:9pt

values: width:auto; font-size:100%; position:absolute; text-align:center; margin-top:36em;

values: width:auto; font-size:100%; position:absolute; text-align:center; margin-top:32.4em;

values: width:auto; font-size:100%; position:absolute; text-align:center; margin-top:25.2em;

values: width:auto; font-size:100%; position:absolute; text-align:center; margin-top:20.88em;

values: Human skeletal changes due to bipedalism

values: text-size:100%; float:left; position:relative; text-align:left; top:-0.75em; left:0em; width:7em; line height:9pt

values: width:auto; font-size:100%; position:absolute; text-align:center; margin-top:14.58em;

values: width:auto; font-size:100%; position:absolute; text-align:center; margin-top:11.88em;

values: width:auto; font-size:100%; position:absolute; text-align:center; margin-top:6.48em;

values: width:auto; font-size:100%; position:absolute; text-align:center; margin-top:5.4em;

values: width:auto; font-size:100%; position:absolute; text-align:center; margin-top:3.96em;

values: width:auto; font-size:100%; position:absolute; text-align:center; margin-top:2.844em;

values: width:auto; font-size:100%; position:absolute; text-align:center; margin-top:1.8em;

values: width:auto; font-size:100%; position:absolute; text-align:center; margin-top:0.9em;

values: width:auto; font-size:100%; position:absolute; text-align:center; margin-top:0.18em;

values: width:auto; font-size:100%; position:absolute; text-align:center; margin-top:0em; text-size:100%; float:left; position:relative; text-align:left; top:-0.75em; left:-13em; width:7em;

values: width:auto; font-size:100%; position:absolute; text-align:center; margin-top:12.6em; text-size:100%; float:left; position:relative; text-align:left; top:-0.75em; left:-13em; width:7em;

values: width:auto; font-size:100%; position:absolute; text-align:center; margin-top:24.3em; text-size:100%; float:left; position:relative; text-align:left; top:-0.75em; left:-13em; width:7em;

values: width:auto; font-size:100%; position:absolute; text-align:center; margin-top:15.12em; text-size:100%; float:left; position:relative; text-align:left; top:-0.75em; left:-4.20em; width:7em;

values: margin-left:3.1em; border-style:none; float:left; clear:both;

values: margin:0 0.5em; float:left; padding:0 auto; border-style:none; clear:both; text-align:center; width:20.5em

values: float: right; text-align: left; padding:0.5em 0 0 0; border-style:solid; float:right;clear:right; padding:0 0.5em; margin:0.3em 0 0.8em 1.4em; overflow: hidden;

values:
/wiki/Type_specimen
mw-redirect
Type specimen

values: Representation in the International Phonetic Alphabet (IPA)

attrs: alt src width height class srcset data-file-width data-file-height

values:
/wiki/Out_of_Africa_I
mw-redirect
Out of Africa I

values:
/wiki/Archaic_Homo_sapiens
mw-redirect
Archaic Homo sapiens

values:
/wiki/Marine_Isotope_Stage
mw-redirect
Marine Isotope Stage

values:
/wiki/%C5%A0ipka_(cave)
mw-redirect
Šipka (cave)

values:
/wiki/Electron_spin_resonance
mw-redirect
Electron spin resonance

values: Max Planck Institute for Evolutionary Anthropology

values:
/wiki/Southwest_Asian_Neanderthals
mw-redirect
Southwest Asian Neanderthals

values:
/wiki/Eemian_interglacial
mw-redirect
Eemian interglacial

values:
/wiki/Human_skull
mw-redirect
Human skull

values:
/wiki/Cranial_capacity
mw-redirect
Cranial capacity

values: Detail the scavenger vs. hunter debate. (April 2017)

values:
/wiki/Sch%C3%B6ningen_spears
mw-redirect
Schöningen spears

values:
/wiki/El_Sidr%C3%B3n
mw-redirect
El Sidrón

values: Edit section: Claims of art and adornment

values:
/wiki/Middle_Palaeolithic
mw-redirect
Middle Palaeolithic

values:
/wiki/Neanderthal_genome
mw-redirect
Neanderthal genome

values: Edit section: Interbreeding with modern humans

values:
/wiki/Max_Planck_Institute
mw-redirect
Max Planck Institute

values: The material near this tag may rely on an unreliable source. (October 2017)

values:
/wiki/Biliary_cirrhosis
mw-redirect
Biliary cirrhosis

values: Daily Mail has a reputation as an occasional fabricator. Wikipedia:Reliable_sources/Noticeboard/Archive_220#Daily_Mail_RfC (August 2017)

values: noprint Inline-Template noprint Template-Fact

values:
/wiki/HOX_cluster
mw-redirect
HOX cluster

values:
/wiki/Neanderthal_admixture
mw-redirect
Neanderthal admixture

values:
/wiki/Campanian_Ignimbrite_Eruption
mw-redirect
Campanian Ignimbrite Eruption

values: Edit section: Coexistence with modern humans

values:
/wiki/Domestication_of_the_dog
mw-redirect
Domestication of the dog

values:
/wiki/Recent_African_Origin
mw-redirect
Recent African Origin

values:
/wiki/Washington_University
mw-redirect
Washington University

values:
/wiki/Lagar_Velho
mw-redirect
Lagar Velho

values:
/wiki/Pe%C8%99tera_Muierii
mw-redirect
Peștera Muierii

values:
/wiki/Modern_humans
mw-redirect
Modern humans

values: Edit section: Notable European Neanderthals

values: Edit section: Notable Southwest Asian Neanderthals

values:
/wiki/Paul_B._Pettitt
mw-redirect
Paul B. Pettitt

values: Edit section: Notable Central Asian Neanderthal

values: Edit section: Mixed with H. heidelbergensis traits

values: Pontnewydd (Bontnewydd-Llanelwy) Paleaolithic site

values: Edit section: H. neanderthalensis fossils

values:
/wiki/La_Chapelle-aux_Saints_1
mw-redirect
La Chapelle-aux Saints 1

values: Edit section: H. s. sapiens with traits reminiscent of Neanderthals

values:
/wiki/Pestera_cu_Oase
mw-redirect
Pestera cu Oase

values:
/wiki/Lapedo_Child
mw-redirect
Lapedo Child

values:
/wiki/Dawn_of_Humanity_(film)
mw-redirect
Dawn of Humanity (film)

values:
/wiki/Denisova_hominin
mw-redirect
Denisova hominin

values:
/wiki/Neanderthal_behaviour
mw-redirect
Neanderthal behaviour

values:
/wiki/Neanderthals_of_Gibraltar
mw-redirect
Neanderthals of Gibraltar

values: -moz-column-width: 30em; -webkit-column-width: 30em; column-width: 30em;

values:
reflist mw-references-wrap
list-style-type: lower-alpha;

values:
nofollow
external text
//doi.org/10.1038%2Fnature13621

values:
/wiki/PubMed_Identifier
mw-redirect
PubMed Identifier

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/25143113

values: Category:CS1 maint: Uses authors parameter

values:
nofollow
external text
//doi.org/10.1017%2Fs0003598x00067570

values:
nofollow
external text
//doi.org/10.1073%2Fpnas.1018938108

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/25016565

values:
nofollow
external text
//doi.org/10.1073%2Fpnas.0904119106

values:
nofollow
external text
//www.jstor.org/stable/40485013

attrs: alt src title width height srcset data-file-width data-file-height

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/19805257

values:
nofollow
external text
//doi.org/10.1073%2Fpnas.0308085100

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/14745010

values:
nofollow
external text
//doi.org/10.1002%2Fajpa.22777

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/9850627

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/23082212

values:
nofollow
external text
//doi.org/10.1038%2Fnature13810

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/25341783

values:
/wiki/The_New_Scientist
mw-redirect
The New Scientist

values:
nofollow
external text
//doi.org/10.1038%2Fnature12961

values:
nofollow
external text
//doi.org/10.1016%2Fj.cub.2016.03.037

values:
nofollow
external text
//www.worldcat.org/issn/0960-9822

values:
nofollow
external text
//www.worldcat.org/issn/0092-8674

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/9230299

values:
nofollow
external text
//doi.org/10.1086%2F401978

values:
nofollow
external text
//www.jstor.org/stable/2816956

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/13506025

values:
nofollow
external text
//doi.org/10.2307%2F3975867

values:
nofollow
external text
//www.jstor.org/stable/3975867

values:
nofollow
external text
//doi.org/10.1073%2Fpnas.96.13.7117

values:
nofollow
external text
//www.jstor.org/stable/48019

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/10377375

values:
nofollow
external text
//doi.org/10.1073%2Fpnas.96.13.7604

values:
nofollow
external text
//www.worldcat.org/issn/0027-8424

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/10377462

values: American Journal of Physical Anthropology

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/10766948

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/23481345

values:
nofollow
external text
//doi.org/10.1038%2Fnature12886

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/24352235

values:
nofollow
external text
//doi.org/10.1073%2Fpnas.96.10.5581

values:
nofollow
external text
//doi.org/10.1002%2Fevan.10058

values:
nofollow
external text
//doi.org/10.1038%2F35006625

values:
nofollow
external text
//www.worldcat.org/issn/0028-0836

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/10761915

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/18692465

values:
nofollow
external text
//doi.org/10.1126%2Fscience.1174462

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/20510437

values:
nofollow
external text
//doi.org/10.1093%2Fmolbev%2Fmsu222

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/25100861

values:
nofollow
external text
//doi.org/10.1006%2Fjasc.2002.0834

values:
nofollow
external text
//doi.org/10.1006%2Fjhev.1998.0214

values:
nofollow
external text
http://eol.org/pages/4454114/details

values:
nofollow
external text
//doi.org/10.1016%2Fj.jas.2014.04.019

values:
nofollow
external text
//doi.org/10.1016%2Fj.jas.2016.02.006

values:
nofollow
external text
//doi.org/10.1086%2F203725

values:
nofollow
external text
//doi.org/10.1016%2Fj.aanat.2011.01.014

values:
nofollow
external text
//doi.org/10.1073%2Fpnas.192464099

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/12232049

values:
nofollow
external text
//doi.org/10.1038%2Fnature05336

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/17108958

values:
nofollow
external text
//doi.org/10.1126%2Fscience.1188021

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/20448178

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/23055938

values:
nofollow
external text
//doi.org/10.1093%2Fmolbev%2Fmss117

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/22513287

values:
nofollow
external text
//doi.org/10.1073%2Fpnas.1316780110

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/24789039

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/15288521

values:
nofollow
external text
//doi.org/10.1126%2Fscience.1147417

values:
nofollow
external text
//www.worldcat.org/issn/0036-8075

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/17962522

values:
nofollow
external text
//doi.org/10.1038%2Fsrep22159

values:
nofollow
external text
//www.worldcat.org/issn/2045-2322

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/26922901

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/15869783

values:
nofollow
external text
//doi.org/10.1073%2Fpnas.120178997

values:
nofollow
external text
//www.jstor.org/stable/122870

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/10852955

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/21445243

values:
nofollow
external text
//doi.org/10.1073%2Fpnas.1016868108

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/21187393

values:
nofollow
external text
//doi.org/10.1073%2Fpnas.082111899

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/11972028

values:
nofollow
external text
//doi.org/10.1006%2Fjhev.1999.0301

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/10375476

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/20956292

values:
nofollow
external text
//doi.org/10.1073%2Fpnas.1212924109

values:
nofollow
external text
//doi.org/10.1073%2Fpnas.1605834113

values:
nofollow
external text
//doi.org/10.1038%2Fnature.2014.15805

values:
nofollow
external text
//doi.org/10.1038%2Fnature18291

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/27251286

values:
nofollow
http://www.msnbc.msn.com/id/13955661/
external text

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/17510332

values:
nofollow
http://www.msnbc.msn.com/id/15732243/
external text

values:
/wiki/New_York_Times
mw-redirect
New York Times

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/19213888

values:
nofollow
external text
//doi.org/10.1038%2Femboj.2009.222

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/19661919

values:
nofollow
external text
//doi.org/10.1016%2Fj.gene.2013.06.005

values:
nofollow
external text
//www.worldcat.org/issn/0378-1119

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/23872234

values:
nofollow
external text
//doi.org/10.1016%2Fj.ajhg.2015.01.006

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/25683122

values:
nofollow
external text
//doi.org/10.1016%2Fj.ajhg.2014.12.029

values:
nofollow
external text
//doi.org/10.1016%2Fj.ajhg.2016.02.023

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/27058445

values:
nofollow
external text
//doi.org/10.1126%2Fscience.1250368

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/24786081

values:
/wiki/Gen._Eng._Biotechnol._News
mw-redirect
Gen. Eng. Biotechnol. News

values: font-size:0.95em; font-size: 90%; color: #555

values:
nofollow
external text
//doi.org/10.1016%2Fj.tree.2007.02.001

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/17300854

values:
nofollow
external text
//www.worldcat.org/oclc/60088352

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/15562317

values:
nofollow
external text
//doi.org/10.1086%2F432455

values:
nofollow
external text
//doi.org/10.1038%2F485033a

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/22552077

values:
nofollow
external text
http://www.cosmosmagazine.com/node/814

values:
nofollow
external text
//doi.org/10.1073%2Fpnas.0608443103

values:
nofollow
external text
//www.jstor.org/stable/30052409

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/17085588

values:
nofollow
external text
//doi.org/10.1093%2Fmolbev%2Fmsr024

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/21266489

values:
nofollow
external text
//arxiv.org/abs/1307.8263

values:
nofollow
external text
//doi.org/10.5779%2Fhypothesis.v9i1.215

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/23544098

values:
nofollow
external text
//doi.org/10.1006%2Fjhev.1997.0133

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/9300343

values:
nofollow
external text
//doi.org/10.1038%2F363252a0

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/8387643

values:
nofollow
external text
//doi.org/10.1002%2Fgea.1017

values:
nofollow
external text
//doi.org/10.1006%2Fjasc.1998.0334

values:
nofollow
external text
//doi.org/10.1006%2Fjpho.2002.0170

values:
nofollow
external text
//doi.org/10.1016%2Fj.wocn.2005.07.002

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/15024415

values:
nofollow
external text
//doi.org/10.1038%2Fnature03585

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/15902255

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/20111705

values: Institute of Anthropology and Ethnography

values:
nofollow
external text
//www.worldcat.org/oclc/36461622

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/24476815

values:
nofollow
external text
//doi.org/10.1016%2Fj.cell.2015.09.042

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/26451479

values: s:The New International Encyclopædia/Neanderthal Man

values:
nofollow
http://www.sciencemag.org/
external text

values:
nofollow
external text
http://www.isogg.org/neanderthaldna.htm

values:
nofollow
external text
http://www.panoramio.com/photo/322489

values:
/wiki/Demirkopru_Dam
mw-redirect
Demirkopru Dam

values: Synoptic table of the principal old world prehistoric cultures

values: Use of animals during the Gravettian period

values: nowraplinks collapsible collapsed navbox-subgroup

values: Prehistoric pile dwellings around the Alps

values: Neolithic circular enclosures in Central Europe

values: Timeline of clothing and textiles technology

values: Stone circles in the British Isles and Brittany

values: List of burial mounds in the United States

values:
/wiki/Megalithic_tomb
mw-redirect
Megalithic tomb

values: List of archaeoastronomical sites by country

values: Entheogenic drugs and the archaeological record

values: nowraplinks hlist collapsible autocollapse navbox-inner

values: nowraplinks hlist collapsible collapsed navbox-inner

values:
/wiki/Ehringsdorf_skull
mw-redirect
Ehringsdorf skull

values:
/wiki/Altamura_man
mw-redirect
Altamura man

values:
/wiki/Krapina_Neanderthal_Site
mw-redirect
Krapina Neanderthal Site

values:
/wiki/H._heidelbergensis
mw-redirect
H. heidelbergensis

values: Neanderthal Man: In Search of Lost Genomes

values:
/wiki/The_Grisly_Folk
mw-redirect
The Grisly Folk

values: Wikipedia:Reliable sources for Palaeolithic articles

values: nowraplinks hlist collapsible expanded navbox-inner

values: navigation navbox Neanderthal padding:3px

values:
row
navbox-group
width:1%;"letter-spacing: .1px;"

values:
nofollow
external text
http://eol.org/pages/4454114

values: Global Biodiversity Information Facility

values:
nofollow
external text
http://www.gbif.org/species/4827584

values:
nofollow
external text
https://d-nb.info/gnd/4171356-4

values:
noprint metadata navbox
navigation
Portals
font-weight:bold;padding:0.4em 2em

values: Category:All articles with dead external links

values: Category:Articles with dead external links from July 2016

values: Category:Webarchive template wayback links

values: Category:CS1 French-language sources (fr)

values: Category:Pages containing links to subscription-only content

values: Category:CS1 Danish-language sources (da)

values: Category:Articles with dead external links from October 2017

values: Category:Articles with 'species' microformats

values: Category:Wikipedia articles needing clarification from April 2017

values: Category:All articles lacking reliable references

values: Category:Articles lacking reliable references from October 2017

values: Category:Articles lacking reliable references from August 2017

values: Category:Wikipedia articles with GND identifiers

values: Discussion about edits from this IP address [ctrl-alt-n]

values: A list of edits made from this IP address [ctrl-alt-y]

values: You are encouraged to create an account and log in; however, it is not mandatory

values: You're encouraged to log in; however, it's not mandatory. [ctrl-alt-o]

values:
/wiki/Neanderthal
View the content page [ctrl-alt-c]
c

values: Discussion about the content page [ctrl-alt-t]

values: p-namespaces navigation vectorTabs p-namespaces-label

values:
p-variants
navigation
vectorMenu emptyPortlet
p-variants-label

values: Past revisions of this page [ctrl-alt-h]

values: p-views navigation vectorTabs p-views-label

values:
p-cactions
navigation
vectorMenu emptyPortlet
p-cactions-label

values:
mw-wiki-logo
/wiki/Main_Page
Visit the main page

values:
/wiki/Main_Page
Visit the main page [ctrl-alt-z]
z

values: Featured content – the best of Wikipedia

values: Find background information on current events

values: Guidance on how to use and edit Wikipedia

values: About the project, what you can do, where to find things

values: A list of recent changes in the wiki [ctrl-alt-r]

values: portal navigation p-interaction p-interaction-label

values: List of all English Wikipedia pages containing links to this page [ctrl-alt-j]

values:
uls-settings-trigger
Language settings
0
button
true

values: نياندرتال – Arabic

values: নিয়ানডার্থাল – Bangla

values: Неандэрталец – Belarusian

values: Неандерталец – Bulgarian

values: Неандерталь çынни – Chuvash

values: interlanguage-link interwiki-de badge-Q17437796 badge-featuredarticle

values: Νεάντερταλ – Greek

values: نئاندرتال – Persian

values: interlanguage-link interwiki-fr badge-Q17437796 badge-featuredarticle

values: 네안데르탈인 – Korean

values: Նեանդերթալյան մարդ – Armenian

values: निअंडरथल मानव – Hindi

values: האדם הניאנדרטלי – Hebrew

values: interlanguage-link interwiki-he badge-Q17437796 badge-featuredarticle

values: ნეანდერტალელი – Georgian

values: Неандерталдықтар – Kazakh

values: Неандерталь адамдары – Kyrgyz

values: ນີແອນເດີທອນ – Lao

values: interlanguage-link interwiki-hu badge-Q17437796 badge-featuredarticle

values: Неандерталец – Macedonian

values: നിയാന്തർത്താൽ മനുഷ്യൻ – Malayalam

values: ネアンデルタール人 – Japanese

values: ਨੀਐਂਡਰਥਾਲ – Punjabi

values: نيانډرټال – Pashto

values: Неандерталец – Russian

values: Палеоантроп – Sakha

values: Неандерталац – Serbian

values: நியண்டர்தால் மனிதன் – Tamil

values: నియాండర్తల్ – Telugu

values: นีแอนเดอร์ทาล – Thai

values: Неандерталець – Ukrainian

values: نینڈرتهال – Urdu

values: /static/images/wikimedia-button-1.5x.png 1.5x, /static/images/wikimedia-button-2x.png 2x

values: /static/images/poweredby_mediawiki_132x47.png 1.5x, /static/images/poweredby_mediawiki_176x62.png 2x

values: mediawiki ltr sitedir-ltr mw-hide-empty-elt ns-0 ns-subject page-Neanderthal rootpage-Neanderthal vector-nav-directionality skin-vector action-view