Neanderthals (UK: /niˈændərˌtɑːl/, also US: /neɪ-, -ˈɑːn-,
-ˌtɔːl, -ˌθɔːl/), more rarely known as Neandertals,[a]
were archaic humans that became extinct about 40,000 years
ago. 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
which is characterised by tools made out of stone flakes.
Neanderthals are considered either a distinct species, Homo
neanderthalensis, or more rarely a subspecies of Homo
sapiens (H. s. neanderthalensis). A 2010 report found that
Neanderthals share 99.7% of their DNA and are
hence much more closely related than to their closest non-human
relative, the chimpanzee (98.8%). Another report in 2017 concluded
that 1.8 to 2.6 percent of the genes of non-Africans come from
Neanderthals. Compared to modern humans,
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, extending
to 1,736 cm3 (105.9 cu in) in the male Amud 1. This
is within range of anatomically modern humans, whose cranial capacity
range from 1,230 to 1,880 cm3 (75 to 115 cu in).
Males stood 164 to 168 cm (65 to 66 in) and females 152 to
156 cm (60 to 61 in) tall.
Neanderthal genome project
Neanderthal genome project revealed in 2010 that, through
Neanderthals may have contributed to the
DNA of modern
humans, likely between 50,000 and 60,000 years ago.
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 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
the ancestors of
East Asians only.
4.1 Timeline of research
5 Habitat and range
7.1 Question of art and adornment
8.2 Interbreeding with modern humans
9.1 Climate change
9.2 Coexistence with modern humans
10 Interbreeding hypotheses
10.1 Pre-2010 interbreeding hypotheses
10.2 Interbreeding hypotheses since 2010
11.1 Notable European Neanderthals
11.2 Notable Southwest Asian Neanderthals
11.3 Notable Central Asian Neanderthal
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 In popular culture
13 See also
16 Further reading
17 External links
Neanderthals are named after one of the first sites where their
fossils were discovered in the 19th century in the Neandertal in
Erkrath, Germany.[b] Thal is an older spelling of the German word Tal
(with the same pronunciation), which means "valley" (cognate with
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". 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
Neanderthal fossil was distinct enough from humans to warrant
a separate genus. Nevertheless, King's name had priority over the
proposal put forward in 1866 by Ernst Haeckel,
Homo stupidus. The
practice of referring to "the Neanderthals" and "a Neanderthal"
emerged in the popular literature of the 1920s. 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/). 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/), although scientists typically use the /t/
as in German.
Anatomical comparison of skulls of
Homo sapiens (left) and Homo
(in Cleveland Museum of Natural History)
Features compared are the braincase shape, forehead, browridge, nasal
bone, projection, cheek bone angulation, chin and occipital contour
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.
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."
During the early 20th century, the prevailing view was heavily
Arthur Keith and Marcellin Boule, who wrote the first
scientific description of a nearly complete
Senior members of their respective national paleontological institutes
and among the most eminent paleoanthropologists of their time,
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.
During the 1930s, scholars Ernst Mayr,
George Gaylord Simpson and
Theodosius Dobzhansky reinterpreted the existing fossil record and
came to different conclusions.
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
See also: Accretion model of
(stage 1, 450 ka, male)
(stage 2, 350 ka, male)
(stage 3, 130 ka, female)
(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.
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
Levant and reached Georgia (fossils of Dmanisi). Hominins had
reached China by 1.7 Ma and Iberia (Spain) by 1.4 Ma. 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
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. 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
population that stayed in Africa would have evolved, perhaps through
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. As of
1998[update], there is a fossil gap in Europe between 300 and 243 ka
(MIS 8); no hominin has ever been dated to this period.
Conventionally, therefore, European hominins younger than 243,000
years old are called Neanderthals.
The quality of the fossil record greatly increases from 130,000 years
ago onwards. Specimens younger than this date make up the bulk of
Neanderthal skeletons and were the first whose anatomy was
comprehensively studied. They are known as typical or Classic
Neanderthals. 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).
Neanderthal fossils were first discovered in 1829 in the
(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 (Feldhofer Cave), located in the
Neandertal in Erkrath,
Germany (about 12 km (7 mi) east of
Düsseldorf), in August 1856, three years before Charles Darwin's On
the Origin of Species was published.
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.
Timeline of research
Engis 2, child (1829)
Gibraltar 1, female (1848)
Neanderthal 1, male (upper skull 1856, left-cheek 2000)
Spy 2 skull, sex unclear (1886)
Krapina 3, female (1899)
1829: A damaged skull of a
Engis 2, is discovered
in Engis, Netherlands (now Belgium).
1848: A female
Gibraltar 1, is found in Forbes'
Quarry, Gibraltar, but its importance is not recognised.
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
Homo neanderthalensis. He then changed his mind on placing it in
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."
1880: The mandible of a
Neanderthal child is discovered in a secure
Šipka cave, in the Austro-Hungarian Empire (now the Czech
Republic), 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.
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, said by the excavators to
be a burial, a claim still heatedly contested. For
historical reasons it remains the most famous Neanderthal
Marcellin Boule publishes his now discredited influential study
Neanderthal skeletal morphology based on
La Chapelle-aux-Saints 1.
Neanderthal skeletons are excavated in
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.
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
would show that the adult males were genetically related, but the
adult females were not, suggesting female exogamy.
1997: Matthias Krings et al. are the first to amplify Neanderthal
DNA (mtDNA) using a specimen from Feldhofer grotto in
the Neander valley.
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.
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.
Max Planck Institute for Evolutionary Anthropology
Max Planck Institute for Evolutionary Anthropology and
associated institutions launch the
Neanderthal genome project
Neanderthal genome project to
Neanderthal nuclear genome.
2010: 1–4% of the
DNA of living non-African humans are found by the
Max Planck Institute to likely come from Neanderthals, a
result confirmed in 2012, and refined to 1.5–2.1% in
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.
2014: Some researchers express worry that almost all theories for the
Neanderthals' extinction assume that there is something modern humans
Neanderthals did not.
2014: The most comprehensive dating ever of
Neanderthal bones and
tools from hundreds of sites in Europe is carried out. It suggested
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
reached the continent.
Habitat and range
Southwest Asian Neanderthals
Southwest Asian Neanderthals and List of
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
Neanderthal remains are found in Western, Central, Eastern,
and Mediterranean Europe, as well as Southwest, Central, and
Northern Asia up to the
Altai Mountains in Siberia. No
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), although it
is difficult to assess the limits of their northern range because
glacial advances destroy most human remains, the Bontnewydd tooth
Middle Palaeolithic artifacts have been found up to
60° N on the Russian plains.
There likely never were more than 70,000
Neanderthals at any given
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, 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.
Evidence suggests they were much stronger than modern humans, with
particularly strong arms and hands, while they were
comparable in height; based on 45 long bones from at most 14 males and
Neanderthal males averaged 164 to 168 cm (65 to
66 in) and females 152 to 156 cm (60 to 61 in)
tall. 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. A 2007 genetic study suggested some
have had red hair and blond hair, along with a light skin tone.
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
In 2017, researchers using 3D reconstructions of nasal cavities and
Computational Fluid Dynamics
Computational Fluid Dynamics techniques have found that Neanderthals
and modern humans both adapted their noses (independently and in a
convergent way) to help breathe in cold and dry conditions.
The large nose seen in Neanderthals, as well as
affected the shape of the skull and the muscle attachments, and gave
them a weaker bite force than in modern humans.
In The Spread of Modern Humans in Europe (2002) John F. Hoffecker,
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
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.
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 drainage of the dural venous
sinuses (low pressure blood vessels that run between the meninges and
skull leading down through the skull) in the occipital lobe region of
Neanderthal brains appears more asymmetric than other hominid
brains. 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. They had almost the same degree of encephalisation
(i.e. brain-to-body-size ratio) as modern humans.
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
Levallois point – Beuzeville, France
See also: Pleistocene human diet
Neanderthals made stone tools, used fire, and were hunters.
The consensus on their behaviour ends there. It had actually long been
Neanderthals were hunters or scavengers, but 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, and fed on red deer,
reindeer, ibex, wild boar, aurochs and on occasion mammoth,
straight-tusked elephant and rhinoceros. They
appear to have occasionally used vegetables as fall-back
food, revealed in the 2000s and 2010s by isotope analysis of
their teeth and study of their coprolites (fossilised
faeces). Dental analysis of specimens from
Spy, Belgium and
El Sidrón, Spain in 2017 suggested that 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".
The size and distribution of
Neanderthal sites, along with genetic
Neanderthals lived in much smaller and more
sparsely distributed groups than anatomically-modern Homo
sapiens. 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
DNA 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".
The bones of the El Sidrón group show signs of defleshing, suggesting
that they were victims of cannibalism. 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.
Neanderthals deliberately buried their dead, and if they
did, whether such burials had any symbolic meaning,:158–60 are
heavily contested. 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, and this has in
turn been heavily criticised.
Question of art and adornment
Whether Neandethals created art and used adornments, which would
indicate a capability for complex symbolic thought, remains
unresolved. A 2010 paper on radiocarbon dates cast doubt on the
Châtelperronian beads with Neanderthals, and Paul
Mellars considered the evidence for symbolic behavior to have been
refuted. This conclusion, however, is controversial, and others
Jean-Jacques Hublin and colleagues have re-dated material
associated with the
Châtelperronian artefacts and used proteomic
evidence to restate the challenged association with Neanderthals.
A very large number of other claims of
Neanderthal art, adornment, and
structures have been made. These are often taken by the media as
Neanderthals were capable of symbolic thought, or
were "mental equals" to anatomically modern humans. As
evidence of symbolism, none of them are widely accepted, although
the same is true for
Middle Palaeolithic anatomically modern humans.
Among many others:
Flower pollen on the body of pre-
Shanidar 4, Iraq, had in
1975 been argued to be a flower burial. Once popular, this theory
is no longer accepted.
Pigmented shells from Murcia, Spain, were argued in 2009 to be
Neanderthal make-up containers.
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 took to mean
Neanderthals wore bird feathers as
Deep scratches were found in 2012 on a cave floor underlying
Neanderthal layer in Gorham's Cave, Gibraltar, which some have
controversially interpreted as art.
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.
In 2015, a study argued that a number of 130,000-year-old eagle talons
found in a cache near Krapina, Croatia along with
had been modified to be used as jewellery.
All of these appeared only in single locations. Yet in 2018, using
uranium-thorium dating methods, red painted symbols comprising a
scalariform (ladder shape), a negative hand stencil, and red lines and
dots on the cave walls of three Spanish caves 700 km
(430 mi) apart were dated to at least 64,000 years old. If
the dating is correct, they were painted before the time anatomically
modern humans are thought to have arrived in Europe.
John D. Hawks argues these findings demonstrate
Neanderthals were capable of symbolic behavior previously thought to
be unique to modern humans.
Neanderthal genome project
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
In 1997, geneticists were able to extract a short sequence of
Neanderthal bones. The extraction of mt
DNA from a second specimen
was reported in 2000, and showed no sign of modern human descent from
In July 2006, the
Max Planck Institute for Evolutionary Anthropology
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
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. From
DNA analysis estimates, the two shared a common ancestor about
500,000 years ago. An article 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. A 2007 study
pushes the point of divergence back to around 800,000 years ago.
Edward Rubin of the
Lawrence Berkeley National Laboratory
Lawrence Berkeley National Laboratory states
recent genome testing of
Neanderthals suggests human and Neanderthal
DNA are some 99.5% to nearly 99.9% identical.
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
Interbreeding with modern humans
On November 16, 2006,
Lawrence Berkeley National Laboratory
Lawrence Berkeley National Laboratory issued a
press release suggesting
Neanderthals and ancient humans probably did
not interbreed. Edward M. Rubin, director of the U.S. Department
Lawrence Berkeley National Laboratory
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
Their results show the genomes of modern humans and
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
In 2008 Richard E. Green et al. from
Max Planck Institute for
Evolutionary Anthropology in Leipzig, Germany, published the full
DNA (mtDNA) and suggested
Neanderthals had a long-term effective population size smaller than
that of modern humans." In the same publication, it was disclosed
Svante Pääbo that in the previous work at the Max Planck
Institute, "Contamination was indeed an issue," and they eventually
realised that 11% of their sample was modern human DNA.
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
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.
However, an analysis of a first draft of the
Neanderthal genome by the
same team released in May 2010 indicates interbreeding may have
occurred. "Those of us who live outside Africa carry a little
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]. 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.
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 colonising 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 colonisation process.
It is suggested that 20 percent of
DNA survived in modern
humans, notably expressed in the skin, hair and diseases of modern
people.[unreliable source?] Modern human genes involved in
making keratin—the protein found in skin, hair, and nails—have
specially high levels of
Neanderthal DNA. For example, around 66%
East Asians contain the
Neanderthal skin gene, while 70% of
Europeans possess 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. Eight
DNA comes from an unknown group of archaic
humans, tantalising hints of unknown groups from Asia and Africa that
left genes in
Denisovans and modern humans, respectively.
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.
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
Neanderthals than other Africans were, because of ancient
genetic divisions within Africa. 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
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.
Among the genes shown to differ between present-day humans and
Neanderthals were RPTN, SPAG17, CAN15, TTF1,
FOXP2 and PCD16.
Specifically, a visualisation map of the reference modern-human
containing the genome regions with high degree of similarity or with
novelty according to a
Neanderthal of 50k has been built by Pratas
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
Genetics that the relatively greater quantity of
DNA in the genomes of individuals of East Asian
descent (than those of European descent) cannot be explained by
differences in selection. 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. 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."
In a subsequent interview, Dr. Lohmueller did note that these findings
go against the commonly-held perception that
Neanderthals were mostly
localised 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."[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
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
East Asians interbred with
Neanderthals just a little bit
more."[better source needed]
Studies published in March 2016 suggest that modern humans bred with
hominins, including Neanderthals, on multiple occasions. 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. This could explain why no modern man had to date been
found with a
Neanderthal Y chromosome. 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.
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.
In 2016 researchers reported that they had found
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.
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. The reconstructed DNA
methylation map allowed researchers to assess gene activity levels
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
HOX cluster of genes were behind many of the morphological
Neanderthals and modern humans, including shorter
limbs, curved bones and more.
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
Neanderthal survival in the peninsula after 42,000 years
ago is almost non-existent.
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. The exact nature of
biological and cultural interaction between
Neanderthals and other
human groups is contested.
Possible scenarios for the extinction of the
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.
Jared Diamond has suggested a
scenario of violent conflict and displacement.
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
1,600 generations ago.
Neanderthal range in light grey
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
Neanderthals died out in Europe between 41,000 and
39,000 years ago, apparently coinciding with the start of a very cold
period. 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
Coexistence with modern humans
Approximate reconstruction of a
Neanderthal skeleton and artistic
interpretation of the
La Ferrassie 1
La Ferrassie 1
Neanderthal man from the National
Museum of Nature and Science
In November 2011 tests conducted at the Oxford
Unit in England on what were previously thought to be
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. 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. Nonetheless, the exact
nature of biological and cultural interaction between
other human groups has been contested.
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
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. 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
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
Cro-Magnon population by interbreeding. This would be counter to
strict versions of the recent African origin theory, since it would
imply that at least part of the genome of Europeans would descend from
Pre-2010 interbreeding hypotheses
Until the early 1950s, most scholars believed
Neanderthals were not in
the ancestry of living humans.:232–234 Nevertheless,
Thomas H. Huxley in 1904 saw among Frisians the presence of what he
believed to be Neanderthaloid skeletal and cranial characteristics as
an evolutionary development from
Neanderthal rather than as a result
of interbreeding, saying that "the blond long-heads may exhibit one of
the lines of evolution of the men of the Neanderthaloid type," yet he
raised the possibility that the Frisians alternatively "may be the
result of the admixture of the blond long-heads with
thus separating "blond" from "Neanderthaloid."
Hans Peder Steensby proposed interbreeding in 1907 in the article Race
studies in Denmark. He strongly emphasised that all living humans are
of mixed origins. He held that this would best fit observations,
and challenged the widespread idea that
Neanderthals were ape-like or
inferior. Basing his argument primarily on cranial data, he noted that
the Danes, like the Frisians and the Dutch, exhibit some
Neanderthaloid characteristics, and felt it was reasonable to "assume
something was inherited" and that
Neanderthals "are among our
Carleton Stevens Coon in 1962 found it likely, based upon evidence
from cranial data and material culture, that
Neanderthal and Upper
Paleolithic peoples either interbred or that the newcomers reworked
Neanderthal implements "into their own kind of tools."
Christopher Thomas Cairney in 1989 went further, laying out a
rationale for hybridisation and adding a broader discussion of
physical characteristics as well as commentary on interbreeding and
its importance to adaptive European phenotypes. Cairney specifically
discussed the "intermixture of racial elements" and
By the early 2000s, the majority of scholars supported the Out of
Africa hypothesis, according to which anatomically modern
humans left Africa about 50,000 years ago and replaced Neanderthals
with little or no interbreeding. Yet some scholars still argued for
hybridisation with Neanderthals. The most vocal proponent of the
hybridisation hypothesis was
Erik Trinkaus of Washington
University. Trinkaus claimed various fossils as products of
hybridised populations, including the child of Lagar Velho, a skeleton
found at Lagar Velho in Portugal and the Peștera
Muierii in Romania.
Interbreeding hypotheses since 2010
In 2010, geneticists announced that interbreeding had likely taken
place, a result confirmed in
2012.[page needed] The genomes of all non-Africans
include portions that are of
Neanderthal origin, a share
estimated in 2014 to 1.5–2.1%. This
DNA is absent in Sub-Saharan
Yoruba people and San subjects).
Ötzi the iceman,
Europe's oldest preserved mummy, was found to possess an even higher
Neanderthal ancestry. The two percent of
DNA 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.
2012 genetic studies seem to suggest that modern humans may have mated
with "at least two groups" of archaic humans:
Denisovans. Some researchers suggest admixture of 3.4–7.9% in
modern humans of non-African ancestry, rejecting the hypothesis of
ancestral population structure. 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. 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.
While some modern human nuclear
DNA has been linked to the extinct
Neanderthals, no mitochondrial
Neanderthal origin has been
detected, 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.
Notable European Neanderthals
La Ferrassie 1, skull cast
Le Moustier 1 in 1909
Le Moustier 1 in 2011
Shanidar 1, skull cast
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 fossilised 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
La Ferrassie 1: A fossilised 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
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
Mousterian tool techno-complex is named after its
Notable Southwest Asian Neanderthals
Remains of more than 70
Southwest Asian Neanderthals
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
Shanidar microfauna and whose burrowing activity can be
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:
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
Notable Central Asian Neanderthal
Teshik-Tash 1: An 8–11 year old skeleton discovered in
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.
This section describes bones with
Neanderthal traits in chronological
H. heidelbergensis traits
> 350 ka: Sima de los Huesos c. 500:350 ka
350–200 ka: Pontnewydd 225 ka ago.
200–135 ka: Atapuerca, Vértesszőlős, Ehringsdorf, Casal
de'Pazzi, Biache, La Chaise, Montmaurin, Prince, Lazaret,
H. neanderthalensis fossils
130–50 ka: Krapina, Saccopastore skulls, Malarnaud, Altamura,
Gánovce, Denisova, Okladnikov, Pech de l'Azé, Tabun
Shanidar 1 to 9
La Ferrassie 1
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, 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), Arcy-sur-Cure,
Châtelperron, Figueira Brava, Mezmaiskaya (41±1 cal ka),
Zafarraya, Vindija, Velika Pećina.
H. s. sapiens with traits reminiscent of Neanderthals
Pestera cu Oase
Pestera cu Oase 35 ka,
Mladeč 31 ka, Pestera
Muierii 30 ka (n/s),
Lapedo Child 24.5 ka.
In popular culture
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, emphasising hairiness and rough, dark complexion.
Early human migrations
Neanderthals in Gibraltar
Prehistoric Autopsy (2012 BBC documentary)
Dawn of Humanity (2015 PBS film)
^ 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 established by King in 1864, Homo
neanderthalensis. 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.
^ The valley is named after Joachim Neander, whose Greek-style last
name had been changed by his grandfather from "Neumann" ("new
^ 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 18th and
19th centuries.) Tal became standardized with the German spelling
reform of 1901, thus the German name Neandertal for both the valley
^ 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.
^ 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.
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
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.
^ Romeo, Luigi (1979). Ecce Homo!:A Lexicon of Man. Amsterdam: John
Benjamins Publishing Company. p. 92. ISBN 9027220069.
^ 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.
Neanderthal in ODE". Oxford Dictionaries.
^ ""Neanderthal" in Random House Dictionary (US) & Collins
Dictionary (UK)". Dictionary.com.
^ "Neanderthal". Wiley-Blackwell Encyclopedia of
Chichester, West Sussex: Wiley-Blackwell. 2013.
^ a b c King, William (Jan 1864). "The Reputed Fossil Man of the
Neanderthal" (PDF). The Quarterly Journal of Science. 1: 96.
^ "Neandertal oder Neanderthal? – Was ist denn nun richtig?".
Neanderthal museum. Retrieved February 1, 2017. Heute
sollten Ortsbezeichnungen das „Neandertal“ ohne „h“
bezeichnen. Alle Namen, die sich auf den prähistorischen Menschen
beziehen, führen das „h“. [Today one should write for place names
"Neandertal" without an "h". All names related to the prehistoric
humans keep the "h".]
^ 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
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)
^ 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)
^ 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
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)
^ 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:
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^ McKie, Robin (June 2, 2013). "Why did the
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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
^ a b c d "New dates rewrite
Neanderthal story". BBC News.
^ Shaw, Ian; Jameson, Robert, eds. (1999). A Dictionary of
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Neanderthal genome sequenced:
DNA signatures found in
present-day Europeans and Asians, but not in Africans, ScienceDaily
^ "Biologists Sequence a New
Vindija Cave in
Croatia". SciTechDaily. 6 October 2017. Retrieved 11 March 2018.
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Archived from the original on June 17, 2006. : Includes
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