Human timeline view • discuss • edit -10 — – -9 — –
-8 — – -7 — – -6 — – -5 — – -4 — – -3 — – -2
— – -1 — – 0 — Human-like
Australopithecus HOMO HABILIS HOMO ERECTUS
NEANDERTHAL HOMO SAPIENS ← Earlier apes
← Possibly bipedal ← Earliest bipedal ←
Earliest stone tools ← Earliest exit
from Africa ← Earliest fire use ← Earliest cooking
← Earliest clothes ←
Axis scale : millions of years .
Also see: Life timeline and Nature timeline
NEANDERTHALS, or more rarely NEANDERTALS, (UK :
/niˈændərˌtɑːl/ , also US : /neɪ-, -ˈɑːn-, -ˌtɔːl,
-ˌθɔːl/ ; named for the
Neandertal region in
Germany ) were a
species or subspecies of archaic humans in the genus
Homo that became
extinct about 40,000 years ago.
Neanderthals and modern humans
share 99.7% of their
DNA and are hence closely related. (By
comparison, both modern humans and
Neanderthals share 98.8% of their
DNA with their closest non-human living relatives, the chimpanzees .)
Neanderthals left bones and stone tools in
Eurasia , from Western
Europe to Central and
Northern Asia .
Fossil evidence suggests
Neanderthals evolved in Europe, separate from modern humans in Africa
for more than 400,000 years. They are considered either a distinct
Homo neanderthalensis, or more rarely as a subspecies of
Homo sapiens (H. s. neanderthalensis).
Hundreds of lithic assemblages were created by
Neanderthals in Europe
and in Western Asia . Almost all of them are of the so-called
Mousterian techno-complex , which begins c. 160,000 years ago when the
makers started making fewer hand axes , and instead started to make
tools out of flakes .
Compared to modern humans ,
Neanderthals had a lower
surface-to-volume ratio, with shorter legs and a bigger body, in
conformance with Bergmann\'s rule , as an energy-loss reduction
adaptation to life in a high-latitude (i.e. seasonally cold) climate.
Neanderthals had cranial capacities averaging 1,600 cm3 (98 cu
in), females 1,300 cm3 (79 cu in), extending to 1,736 cm3 (105.9 cu
Amud 1 . 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.
Neanderthal genome project
Neanderthal genome project published papers in 2010 and 2014
Neanderthals contributed to the
DNA of modern humans ,
including most humans outside sub-Saharan Africa , as well as a few
populations in sub-Saharan Africa, through interbreeding , likely
between 50,000 and 60,000 years ago. Recent studies also show that
Neanderthals mated with ancestors of modern humans long before
the "out of Africa migration" of present-day non-Africans, as early as
100,000 years ago. In 2016, research indicated that there were three
distinct episodes of interbreeding between modern humans and
Neanderthals: the first encounter involved the ancestors of
non-African modern humans, probably soon after leaving Africa; the
second, after the ancestral
Melanesian group had branched off (and
subsequently had a unique episode of interbreeding with
and the third, involving the ancestors of
East Asians only.
Geneticists first sequenced the entire genome of a
2013 by extracting it from the phalanx bone of a 50,000-year-old
* 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
* 8.1 Background
* 8.2 Interbreeding with modern humans
* 9 Extinction
* 9.1 Climate change
* 9.2 Coexistence with modern humans
* 10 Interbreeding hypotheses
* 11 Specimens
* 11.1 Notable European
* 11.2 Notable
Southwest Asian Neanderthals
* 11.3 Notable Central Asian
* 11.4.1 Mixed with
H. heidelbergensis traits
* 11.4.2 H. neanderthalensis fossils
Homo sapiens sapiens with archaic traits reminiscent of
* 12 Popular culture
* 13 See also
* 14 Notes
* 15 References
* 16 Further reading
* 17 External links
Neanderthals are named after one of the first sites where their
fossils were discovered in the 19th century, about 12 km (7 mi) east
Düsseldorf , Germany, in the Feldhofer
Cave , located in the
Düssel River's Neander valley. Thal is an older spelling of the
German word Tal (with the same pronunciation), which means "valley"
(cognate with English dale).
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 Neanderthaler or Neandertaler is in the
International Phonetic Alphabet
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.
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
Neanderthals as a subspecies of H. sapiens.
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 1980s.
Neanderthals and living humans are thought to have evolved from
Homo erectus . In the earliest known migration wave into
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 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
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
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
, into early anatomically modern humans by 200,000 years ago or
Neanderthal traits are present in
Homo heidelbergensis specimens
beginning between 600,000 and 350,000 years ago. Conventionally,
European hominins are called
Neanderthals from c. 250,000 years ago.
Because the quality of the fossil record greatly increases from
130,000 years ago onwards, specimens younger than this date make up
the bulk of known
Neanderthal skeletons and were the first whose
anatomy was comprehensively studied. They are known as Classic
Neanderthal excavation site (former Kleine Feldhofer
Grotte ) in
Neander Valley , and its position in North
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 in the
Germany in August
1856, three years before
Charles Darwin 's On the Origin of Species
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
Johann Carl Fuhlrott , who turned the fossils over to
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,
La Chapelle-aux-Saints 1 , male (1908)
* 1829: A damaged skull of a
Engis 2 , is
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 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 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.
* 1899: Sand excavation workers find hundreds of fragmentary
Neanderthal remains representing dozens of 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 skeleton. :15
Marcellin Boule publishes his now discredited influential
Neanderthal skeletal morphology based on La
Chapelle-aux-Saints 1 .
* 1953–1957: Ten
Neanderthal skeletons are excavated in Shanidar
Iraqi Kurdistan , by
Ralph Solecki and colleagues.
Erik Trinkaus 's study of
Neanderthal feet strongly argues
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
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
* 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 .
* 1997: Matthias Krings et al. are the first to amplify Neanderthal
DNA ) 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
Abrigo do Lagar Velho , Portugal, and is
described as presenting a mosaic of anatomically modern human and more
archaic features, reminiscent of Neanderthals.
* 2005: The
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
Max Planck Institute to likely come from Neanderthals, a result
confirmed in 2012, and refined to 1.5–2.1% in 2014.
* 2013: In the midst of a hundred-year old debate over the existence
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 had that
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 and List of
Neanderthal sites Sites where typical
Neanderthal fossils have
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 being
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, 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 7 females,
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
Neanderthals may 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 pattens similar to anatomically modern humans. They
concluded that, "We don’t yet know for sure, but it seems likely
that, as part of their adaptation to cold,
Neanderthals were furry.
Chimpanzees have ridges on their finger bones that stem from the way
that they clutch their mothers' fur as infants.
Modern humans don’t
have these ridges, but
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
Neanderthal brains were 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
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
encephalization (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 (131 lb).
France Main article:
Neanderthals made stone tools, used fire, and were hunters. The
consensus on their behaviour ends here.
It had actually long been debated whether
Neanderthals were hunters
or scavengers. 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 . However, while they were largely carnivorous , new
Neanderthals occasionally used vegetables as fall-back
food. In 2010, an isotope analysis of
Neanderthal teeth found traces
of cooked vegetable matter, and a 2014 study of
(fossilized feces) found substantial amounts of plant matter. Dental
analysis of specimens from Spy , Belgium and
El Sidrón ,
2017 argued these
Neanderthals had a wide-ranging diet, and that those
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
The bones of twelve
Neanderthals were discovered at
El Sidrón cave in
Atapuerca Mountains of 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 Sidron 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, :158–160 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.
CLAIMS OF ART AND ADORNMENT
Upon Higham et al.'s (2010) publication of new radiocarbon dates
shedding doubt on the association of
Châtelperronian beads with
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
in Europe has now effectively collapsed”. This conclusion, however,
is controversial, and others such as
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.
There exists a very large number of other claims of
adornment, and structures. 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
Middle Palaeolithic anatomically modern humans. Among many
* 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 personal adornments.
* 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
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.
Neanderthal genome project
Neanderthal genome project
Early investigations concentrated on mitochondrial
which, owing to strictly matrilineal inheritance and subsequent
vulnerability to genetic drift , is of limited value in evaluating the
possibility of interbreeding of
In 1997, geneticists were able to extract a short sequence of DNA
Neanderthal bones. The extraction of mt
DNA from a second
specimen was reported in 2000, and showed no sign of modern human
descent from Neanderthals.
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 mtDNA
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 states
recent genome testing of
Neanderthals suggests human and Neanderthal
DNA are some 99.5% to nearly 99.9% identical.
INTERBREEDING WITH MODERN HUMANS
On November 16, 2006,
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 of
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 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. 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
be identified. Geneticist at the
Max Planck Institute for
Evolutionary Anthropology extracting ancient
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. It is a small but very real proportion of ancestry in
non-Africans today," says Dr. David Reich of
Harvard Medical School
Harvard Medical School ,
who worked on the study. This research compared the genome of the
Neanderthals to five modern humans from China, France, sub-Saharan
Africa, and Papua New Guinea. The finding is that about 1 to 4 percent
of the genes of the humans outside sub-Saharan Africa came from
Neanderthals, compared to the baseline defined by the two sub-Saharan
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
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.
It is suggested that 20 percent of
DNA survived in modern
humans, notably expressed in the skin, hair, and diseases of modern
people. 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% 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
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.
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
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
RPTN , SPAG17 , CAN15 ,
FOXP2 and PCD16 .
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
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 cannot account for the higher
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
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." Vernot also noted that "umans have
been constantly migrating throughout their history—this makes it
hard to say exactly where interactions with
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."
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
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. 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
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 1600 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
Neanderthals restricted themselves to more local
COEXISTENCE WITH MODERN HUMANS
Approximate reconstruction of a
Neanderthal skeleton and
artistic interpretation of the
La Ferrassie 1
Neanderthal man from the
National Museum of Nature and Science
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. Given that the 2014 study by Thomas
Neanderthal bones and tools indicates that
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
Neanderthals and 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 for a long period of time before the arrival of modern
humans. These modern humans may have introduced a disease that
contributed to the extinction of Neanderthals, and that may be added
to other recent explanations for their extinction. When Neanderthal
ancestors left Africa roughly 100,000 years earlier they adapted to
the pathogens in their European environment, unlike modern humans who
adapted to African pathogens. This transcontinental movement is known
as the Out of Africa model . If contact between humans and
Neanderthals occurred in Europe and Asia the first contact may have
been devastating to the
Neanderthal population, because they would
have had little if any immunity to the African pathogens. More recent
historical events in
Eurasia and the Americas show a similar pattern,
where the unintentional introduction of viral, or bacterial pathogens
to unprepared populations has led to mass mortality and local
population extinction. The most well-known example of this is the
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
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.
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 , since it would imply
that at least part of the genome of Europeans would descend from
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. 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
Washington University . Trinkaus claims various fossils
as products of hybridized populations, including the child of Lagar
Velho , a skeleton found at
Lagar Velho in
Portugal . In a 2006
publication co-authored by Trinkaus, the fossils found in 1952 in the
Peștera Muierii ,
Romania , are likewise claimed as
descendants of previously hybridized populations.
Genetic research has asserted that some admixture took place. The
genomes of all non-Africans include portions that are of Neanderthal
origin, due to interbreeding between
Neanderthals and the ancestors
of Eurasians in Northern Africa or the Middle East prior to their
spread. Rather than absorption of the
Neanderthal population, this
gene flow appears to have been of limited duration and limited extent.
An estimated 1 to 4 percent of the
DNA in Europeans and Asians
(French, Chinese and Papua probands) is non-modern, and shared with
DNA rather than with Sub-Saharan Africans (Yoruba
people and San probands).
Ötzi the iceman , Europe's oldest
preserved mummy, was found to possess an even higher percentage of
Neanderthal ancestry. 2014 findings suggest there may be even more
Neanderthal genes in non-African humans than previously expected:
approximately 20% of the
Neanderthal gene pool was present in a broad
sampling of non-African individuals, though each individual's genome
was on average only 2% Neanderthal.
2012 genetic studies seem to suggest that modern humans may have
mated with "at least two groups" of ancient 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. However, some researchers
have argued that there is evidence of possible interbreeding between
Neanderthals and male modern humans.
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
List of human evolution fossils .
Neanderthal 1 : The first
Neanderthal specimen found during mining
in August 1856. It was discovered in a limestone quarry at the
Feldhofer grotto in Neanderthal, Germany. The find consisted of a
skull cap, two femora, three right arm bones, two left arm bones,
ilium, and fragments of a scapula and ribs.
La Chapelle-aux-Saints 1 : Called the Old Man, a fossilized skull
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
Mousterian tool techno-complex is named after its
NOTABLE SOUTHWEST ASIAN NEANDERTHALS
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 " 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 45,000 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.
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
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.
This section describes bones with
Neanderthal traits in chronological
Mixed With H. Heidelbergensis Traits
* > 350 ka: Sima de los Huesos c. 500:350 ka ago
* 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:
Saccopastore skulls , Malarnaud, Altamura ,
Gánovce, Denisova , Okladnikov , Pech de l'Azé, Tabun
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
* In radiocarbon range, > 50 ka:
Le Moustier , Feldhofer , La Quina,
l'Hortus, Kulna ,
Šipka , Saint Césaire, Bacho Kiro , El Castillo ,
Bañolas, El Sidron (48±3 cal ka),
Arcy-sur-Cure , Châtelperron,
Figueira Brava, Mezmaiskaya (41±1 cal ka), Zafarraya, Vindija ,
Homo Sapiens Sapiens With Archaic Traits Reminiscent Of Neanderthals
* < 35
Pestera cu Oase 35 ka,
Mladeč 31 ka, Pestera Muierii 30 ka
Lapedo Child 24.5 ka.
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.
* Dawn of Humanity (2015 PBS film)
Early human migrations
* Life timeline
List of Neanderthal sites
Neanderthals in popular culture
Prehistoric Autopsy (2012 BBC documentary)
* ^ 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
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
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.
* ^ There are modern humans with noses as wide as those of
Neanderthals and modern humans with similar nose lengths, but none
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
spread from Africa to western Asia, then east Asia and Indonesia; its
presence in Europe is uncertain, but it gave rise to
Homo antecessor ,
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
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.
* ^ Camp, C. L; Allison, H. J.; Nichols, R. H. (1964). Bibliography
Fossil Vertebrates 1954–1958. New York: The Geological Society of
America, Inc. p. 556.
* ^ "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“.
* ^ "
Neanderthal in ODE". Oxford Dictionaries.
* ^ ""Neanderthal" in Random House Dictionary (US) & Collins
Dictionary (UK)". Dictionary.com.
* ^ 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
Neanderthal disappearance". Nature. 512 (7514): 306–9.
PMID 25143113 . doi :10.1038/nature13621 . We show that the Mousterian
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 )(subscription required)
* ^ T. Higham (2011). "European Middle and Upper Palaeolithic
radiocarbon dates are often older than they look: problems with
previous dates and some remedies". Antiquity. 85: 235–249. doi
:10.1017/s0003598x00067570 . Few events of European prehistory are
more important than the transition from ancient to modern humans
around 40 000 years ago, a period that unfortunately lies near the
limit of radiocarbon dating. This paper shows that as many as 70 per
cent of the oldest radiocarbon dates in the literature may be too
young, due to contamination by modern carbon. CS1 maint: Uses authors
parameter (link )(subscription required)
* ^ A B R. Pinhasi, T.F.G. Higham, L.V. Golovanova, V.B. Doronichev
(2011). "Revised age of late
Neanderthal occupation and the end of the
Middle Paleolithic in the northern Caucasus". Proceedings of the
National Academy of Sciences USA. 108: 8611–8616. doi
:10.1073/pnas.1018938108 . The direct date of the fossil (39,700 ±
1,100 14C BP) is in good agreement with the probability distribution
function, indicating at a high level of probability that Neanderthals
did not survive at Mezmaiskaya
Cave after 39 ka cal BP. This
challenges previous claims for late
Neanderthal survival in the
northern Caucasus. Our results confirm the lack of reliably dated
Neanderthal fossils younger than ≈40 ka cal BP in any other region
of Western Eurasia, including the Caucasus. CS1 maint: Uses authors
parameter (link )(subscription required)
* ^ A B B. Galván, C.M. Hernández, C. Mallol, N. Mercier, A.
Sistiaga, V. Soler (2014). "New evidence of early Neanderthal
disappearance in the Iberian Peninsula". Journal of
75: 16–27. doi :10.1016/j.jhevol.2014.06.002 . CS1 maint: Uses
authors parameter (link )(subscription required)
* ^ McKie, Robin (June 2, 2013). "Why did the
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
" 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.
* ^ A B C D "BBC News—New dates rewrite
Neanderthal story". BBC
* ^ A B Complete
Neanderthal genome sequenced:
DNA signatures found
in present-day Europeans and Asians, but not in Africans, ScienceDaily
* ^ Colin P.T. Baillie;
University of California, Berkeley .
"Neandertals: Unique from Humans, or Uniquely Human?" (PDF).
National Museum of American History .
DNA and Neanderthals". si.edu.
* ^ Hublin, J. J. (2009). "The origin of Neandertals" . Proceedings
of the National Academy of Sciences. 106 (38): 16022–7. Bibcode
JSTOR 40485013 . PMC 2752594 . PMID 19805257
. doi :10.1073/pnas.0904119106 .
* ^ Harvati, K.; Frost, S.R.; McNulty, K.P. (2004). "Neanderthal
taxonomy reconsidered: implications of 3D primate models of intra- and
interspecific differences" . Proc. Natl. Acad. Sci. U.S.A. 101:
1147–52. PMC 337021 . PMID 14745010 . doi
* ^ "Scientists Identify
Genes in Modern
Sci-News.com. January 30, 2014. Retrieved October 29, 2015.
* ^ "
Homo neanderthalensis King, 1864". Wiley-Blackwell
Human Evolution. Chichester, West Sussex:
Wiley-Blackwell. 2013. pp. 328–331.
* ^ Bednarik, R.G. (2012). "U–Th analysis and rock art: a
response to Pike et al.". Rock Art Research. 29 (2): 244–246.
* ^ 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.
* ^ Shaw, Ian; Jameson, Robert, eds. (1999). A Dictionary of
Archaeology. Blackwell. p. 408. ISBN 0-631-17423-0 . Retrieved August
* ^ 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 .
* ^ Holloway, R.L. (1985). "The poor brain of
neanderthalensis: see what you please". In Delson, E. Ancestors: The
hard evidence. New York: Alan R. Liss. ISBN 978-0471843764 .
* ^ Amano, H.; Kikuchi, T.; Morita, Y.; Kondo, O.; Suzuki,
Hiromasa; et al. (August 2015). "Virtual Reconstruction of the
Amud 1 Cranium". American Journal of Physical
Anthropology. 158: 185–197. doi :10.1002/ajpa.22777 .
* ^ 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 .
* ^ Sánchez-Quinto, F; Botigué, LR; Civit, S; Arenas, C;
Avila-Arcos, MC; Bustamante, CD; Comas, D; Lalueza-Fox, C (October 17,
2012). "North African Populations Carry the Signature of Admixture
with Neandertals". PLOS ONE. 7: e47765. PMC 3474783 . PMID 23082212
. doi :10.1371/journal.pone.0047765 . Retrieved May 29, 2016.
* ^ A B C Rincon, Paul (May 6, 2010). "
Neanderthal genes \'survive
in us\'". BBC News. Retrieved May 7, 2010.
* ^ Fu, Q; Li, H; Moorjani, P; Jay, F; Slepchenko, SM; Bondarev,
AA; Johnson, PL; Aximu-Petri, A; Prüfer, K; de Filippo, C; Meyer, M;
Zwyns, N; Salazar-García, DC; Kuzmin, YV; Keates, SG; Kosintsev, PA;
Razhev, DI; Richards, MP; Peristov, NV; Lachmann, M; Douka, K; Higham,
TF; Slatkin, M; Hublin, JJ; Reich, D; Kelso, J; Viola, TB; Pääbo, S
(October 23, 2014). "
Genome sequence of a 45,000-year-old modern human
from western Siberia". Nature. 514 (7523): 445–449. PMC 4753769 .
PMID 25341783 . doi :10.1038/nature13810 .
* ^ Brahic, Catherine. "Humanity\'s forgotten return to Africa
revealed in DNA",
The New Scientist (February 3, 2014).
* ^ "
Neanderthals mated with modern humans much earlier than
previously thought, study finds: First genetic evidence of modern
DNA in a
Neanderthal individual". ScienceDaily. February 17,
2016. Retrieved March 6, 2016.
* ^ The Combined Landscape of
Neanderthal Ancestry in
Present-Day Humans, Current Biology, Sankararaman et al., 26,
* ^ "Studies find
Neanderthal genes in modern humans".
* ^ Zimmer, Carl (December 18, 2013). "Toe
Fossil Provides Complete
New York Times
New York Times . Retrieved December 18, 2013.
* ^ A B C K. Prüfer; F. Racimo; N. Patterson; F. Jay; S.
Sankararaman; S. Sawyer; A. Heinze; G. Renaud; P.H. Sudmant; C. de
Filippo; et al. (2014). "The complete genome sequence of a Neanderthal
from the Altai Mountains" . Nature. 505: 43–49. PMC 4031459 .
PMID 24352235 . doi :10.1038/nature12886 . (subscription required)
* ^ Kunzig, Robert . "The Year in Science:
Human Origins 1997",
Discover (magazine) (January 1, 1998) reprinted in Contemporary
Readings in Physical Anthropology, p. 145 (Alan Almquist ed., Prentice
* ^ 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.
JSTOR 2816956 . PMID 13506025 . doi
* ^ Foley, Tim.
TalkOrigins Archive . "
Neanderthal or Neandertal?".
* ^ 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.
* ^ Inter alia, Boys' Life, p. 18. January 1924.
* ^ The Oxford Illustrated Dictionary. Great Britain: Oxford
University Press. 1976 . p. 564. (tahl)
* ^ "
Neanderthal adjective—definition in British English
Dictionary & Thesaurus—Cambridge Dictionary Online".
Dictionary.cambridge.org. January 8, 2013. Retrieved January 22, 2013.
* ^ "Oxford Learner\'s Dictionaries—Find pronunciation, clear
meanings and definitions of words at OxfordLearnersDictionaries.com".
* ^ "
Neanderthal at Dictionary.com".
Dictionary.reference.com. Retrieved January 22, 2013.
* ^ Kurtén, Björn (October 10, 1995). Dance of the Tiger: A Novel
of the Ice Age. University of California Press. pp. xxi. ISBN
0-520-20277-5 . Retrieved May 9, 2012.
* ^ Pollet, Carl J. (September 21, 1991). "…And Etymology".
Science News. 140 (12): 191.
JSTOR 3975867 . doi :10.2307/3975867 .
* ^ Tattersall, Ian; Schwartz, Jeffrey H. (1999). "Hominids and
hybrids: The place of
Neanderthals in human evolution" . Proceedings
of the National Academy of Sciences. 96 (13): 7117–9. Bibcode
JSTOR 48019 . PMC 33580 . PMID 10377375 . doi
* ^ Duarte, Cidália; Mauricio, João; Pettitt, Paul B.; Souto,
Pedro; Trinkaus, Erik; Van Der Plicht, Hans; Zilhao, João (1999).
Upper Paleolithic human skeleton from the Abrigo do Lagar
Velho (Portugal) and modern human emergence in Iberia" . Proceedings
of the National Academy of Sciences. 96 (13): 7604–9. Bibcode
JSTOR 48106 . PMC 22133 . PMID 10377462 . doi
* ^ Pääbo, Svante (2014).
Neanderthal Man: In Search of Lost
Genomes. New York: Basic Books. p. 237.
* ^ "L\'homme fossile de La Chapelle-aux-Saints—full
text—Volume VI (p. 11–172), Volume VII (p. 21–56), Volume VIII
(p. 1–70), 1911–1913". Royal College of Surgeons of England.
Retrieved July 26, 2016.
* ^ "Marcellin Boule—French geologist". Encyclopædia Britannica.
Retrieved July 26, 2016.
* ^ "Arthur Keith". Royal Anthropological Institute. Retrieved July
* ^ "La Chapelle-Aux-Saints—The old man of La Chapelle - The
original reconstruction of the ‘Old Man of La Chapelle’ by
Marcellin Boule led to the reason why popular culture
Neanderthals as dim-witted brutes for so many years.".
Smithsonian Institution. Retrieved July 26, 2016.
* ^ "Our
Neandertal Brethren: Why They Were Not a Separate
Species". SCIENTIFIC AMERICAN. August 1, 2010. Retrieved July 26,
* ^ Wolpoff, MH; Hawks, J ; Caspari, R (2000). "Multiregional, not
multiple origins" (pdf).
American Journal of Physical Anthropology .
112 (1): 129–36. PMID 10766948 . doi
* ^ Rightmire, G. P. (2001). "Patterns of hominid evolution and
dispersal in the Middle Pleistocene". Quaternary International. 75
(1): 77–84. doi :10.1016/S1040-6182(00)00079-3 .
* ^ Toro-Moyano, I.; Martínez-Navarro, B.; Agustí, J.; Souday,
C.; Bermúdez de Castro, J. M.; Martinón-Torres, M.; Palmqvist, P.
(2013). "The oldest human fossil in Europe, from Orce (Spain)".
Human Evolution. 65 (1): 1–9. doi
* ^ Wayman, Erin (November 26, 2012). "
Homo antecessor: Common
Ancestor of Humans and Neanderthals?". Smithsonian Institution.
Retrieved May 17, 2017.
* ^ M. Krings; A. Stone; R.W. Schmitz; H. Krainitzki; M. Stoneking;
S. Pääbo (1997). "
DNA sequences and the origin of modern
humans". Cell. 90: 19–30. PMID 9230299 . doi
:10.1016/S0092-8674(00)80310-4 . (subscription required)
* ^ M. Krings; H. Geisert; R.W. Schmitz; H. Krainitzki; S. Pääbo
DNA sequence of the mitochondrial hypervariable region II
Neandertal type specimen". Proceedings of the National
Academy of Sciences USA. 96 (10): 5581–5585. doi
:10.1073/pnas.96.10.5581 . (subscription required)
* ^ P. Beerli; S.V. Edwards (2002). "When did
modern humans diverge?". Evolutionary Anthropology. 11 (S1): 60–63.
doi :10.1002/evan.10058 . (subscription required)
* ^ I.V. Ovchinnikov; A. Götherström; G.P. Romanova; V.M.
Kharitonov; K. Lidén; W. Goodwin (2002). "Molecular analysis of
DNA from the northern Caucasus". Nature. 404: 490–493.
PMID 10761915 . doi :10.1038/35006625 . (subscription required)
* ^ R.E. Green; A.-S. Malaspinas; J. Krause ; A.W. Briggs; P.L.F.
Johnson; C. Uhler; M. Meyer; J.M. Good; T. Maricic; U. Stenzel; et al.
(2008). "A Complete
Determined by High-Throughput Sequencing" . Cell. 134 (3): 416–426.
PMC 2602844 . PMID 18692465 . doi :10.1016/j.cell.2008.06.021 .
* ^ R.E. Green; A.-S. Malaspinas; J. Krause; A.W. Briggs; P.L.F.
Johnson; C. Uhler; M. Meyer; J.M. Good; T. Maricic; U. Stenzel; et al.
(2009). "Targeted Retrieval and Analysis of Five
Genomes". Science. 325 (5938): 318–321. doi :10.1126/science.1174462
. (subscription required)
* ^ P. Endicott; S.Y.W. Ho; C. Stringer (2010). "Using genetic
evidence to evaluate four palaeoanthropological hypotheses for the
Neanderthal and modern human origins". Journal of Human
Evolution. 59 (1): 87–95. doi :10.1016/j.jhevol.2010.04.005 .
* ^ A. Rieux; A. Eriksson; M. Li; B. Sobkowiak; L.A. Weinert; V.
Warmuth; A. Ruiz-Linares; A. Manica; F. Balloux (2014). "Improved
calibration of the human mitochondrial clock using ancient genomes" .
Molecular Biology and Evolution. 31 (10): 2780–2792. PMC 4166928
. PMID 25100861 . doi :10.1093/molbev/msu222 . (subscription
* ^ Cookson, Clive (June 27, 2014). "Palaeontology: How
Neanderthals evolved". Financial Times. Retrieved October 28, 2015.
* ^ Bischoff, James L.; Shamp, Donald D.; Aramburu, Arantza;
Arsuaga, Juan Luis; Carbonell, Eudald; Bermudez de Castro, J.M.
(2003). "The Sima de los Huesos Hominids Date to Beyond U/Th
Equilibrium (>350kyr) and Perhaps to 400–500kyr: New Radiometric
Dates". Journal of Archaeological Science. 30 (3): 275–80. doi
* ^ Stringer, Chris (2011). The Origin of our Species. Penguin. pp.
26–29, 202. ISBN 978-0-141-03720-2 .
* ^ Johansson, Donald; Edgar, Blake (2006). From Lucy to Language.
Simon & Schuster. p. 38. ISBN 978-0-7432-8064-8 .
* ^ A B C D E Papagianni, Dmitra; Morse, Michael (2013). The
Neanderthals Rediscovered. Thames & Hudson. ISBN 978-0-500-05177-1 .
* ^ A B C Stringer, C.; Gamble, C. (1993). In Search of the
Neanderthals. London: Thames and Hudson. ISBN 978-0500050705 .
* ^ "
Smithsonian Institution . Archived
from the original on May 21, 2009. Retrieved May 18, 2009.
* ^ "New Evidence On The Role Of Climate In Neanderthal
Extinction". Science Daily.
* ^ "
Neanderthal Man". Encyclopedia Britannica (15th ed.). 1982.
* ^ "Neanderthal—
Fossils—La Chapelle-aux-Saints". Encyclopedia of Life. Retrieved
July 18, 2016.
* ^ 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: 649–657. CS1 maint: Uses authors parameter
* ^ 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 .
* ^ A B Gargett, R.H. (1989). "Grave Shortcomings: The Evidence for
Neandertal Burial". Current Anthropology. 30 (2): 157–190. doi
* ^ "Universitat Pompeu Fabra, Barcelona, A paleogenetical study
determines the blood group of
Neanderthal man" Archived November 22,
2012, at the
Wayback Machine .
* ^ "Palaeogenetic research at the
Annals of Anatomy—Anatomischer Anzeiger. 194: 133–137. doi
* ^ A B Krings, M; Stone, A; Schmitz, RW; Krainitzki, H; Stoneking,
M; Pääbo, S (1997). "
DNA sequences and the origin of
modern humans". Cell. 90 (1): 19–30. ISSN 0092-8674 . PMID 9230299 .
doi :10.1016/S0092-8674(00)80310-4 .
* ^ Schmitz, Ralf W; et al. (2002). "The
Neandertal type site
revisited: Interdisciplinary investigations of skeletal remains from
the Neander Valley, Germany" . Proceedings of the National Academy of
Sciences. 99 (20): 13342–13347. PMC 130635 . PMID 12232049 . doi
* ^ Duarte, C; Maurício, J; Pettitt, PB; Souto, P; Trinkaus, E;
Van Der Plicht, H; Zilhão, J; et al. (1999). "The early Upper
Paleolithic human skeleton from the
Abrigo do Lagar Velho
Abrigo do Lagar Velho (Portugal)
and modern human emergence in the Iberian Peninsula". Proceedings of
the National Academy of Sciences of the United States of America. PNAS
. 96 (13): 7604–7609.
Bibcode :1999PNAS...96.7604D. PMC 22133 .
PMID 10377462 . doi :10.1073/pnas.96.13.7604 . Retrieved June 21,
* ^ Green, RE; Krause, J; Ptak, SE; et al. (November 16, 2006).
"Analysis of one million base pairs of
Neanderthal DNA". Nature. 444:
330–6. PMID 17108958 . doi :10.1038/nature05336 . Retrieved July 18,
2016. CS1 maint: Explicit use of et al. (link )
* ^ 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.
PMC 5100745 . PMID 20448178 . doi :10.1126/science.1188021 .
* ^ 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. PMC 3464203 .
PMID 23055938 . doi :10.1371/journal.pgen.1002947 .
* ^ Yang, M.A.; Malaspinas, A.S.; Durand, E.Y.; Slatkin, M. (2012).
"Ancient Structure in Africa Unlikely to Explain
Non-African Genetic Similarity" . Molecular Biology and Evolution. 29
(10): 2987–2995. PMC 3457770 . PMID 22513287 . doi
* ^ A B Rendu W, Beauval C, Crevecoeur I, Bayle P, Balzeau A,
Bismuth T, Bourguignon L, Delfour G, Faivre JP, Lacrampe-Cuyaubère F,
Tavormina C, Todisco D, Turq A, Maureille B (January 2014). "Evidence
supporting an intentional
Neandertal burial at La
Chapelle-aux-Saints". Proceedings of the National Academy of Sciences.
111 (1): 81–86. doi :10.1073/pnas.1316780110 .
* ^ P. Villa, W. Roebroeks (2014). "
Neandertal Demise: An
Archaeological Analysis of the Modern
Human Superiority Complex". PLoS
ONE. 9 (4): e96424. doi :10.1371/journal.pone.0096424 . CS1 maint:
Uses authors parameter (link )
* ^ "The
Human Lineage by Matt Cartmill, Fred H. Smith". Google
Books. Retrieved September 27, 2015.
* ^ "Ancient tooth provides evidence of
Durham University . February 11, 2008. Retrieved May
* ^ Wade, Nicholas (October 2, 2007). "
Neanderthal Range". The
New York Times
New York Times . Retrieved May 18, 2009.
* ^ Ravilious, Kate (October 1, 2007). "Neandertals Ranged Much
Farther East Than Thought".
National Geographic Society
National Geographic Society . Retrieved
May 18, 2009.
* ^ Pavlov P, Roebroeks W, Svendsen JI (2004). "The Pleistocene
colonization of northeastern Europe: a report on recent research".
Human Evolution. 47 (1–2): 3–17. PMID 15288521 . doi
* ^ O'Neill, Dennis. "Evolution of Modern Humans: Neanderthals",
Palomar College, June 10, 2011. Retrieved August 21, 2011.
* ^ Puech, Pierre-François; Puech, Bernard. "L’Homme de
Neanderthal par Paul Dardé : L’Homme Primitif" .
French). Retrieved July 5, 2017.
* ^ "Science & Nature—Wildfacts—Neanderthal". BBC. Archived
from the original on January 15, 2009. Retrieved June 21, 2009.
* ^ "Neanderthal". BBC. Archived from the original on January 15,
2009. Retrieved May 18, 2009.
* ^ Froehle, Andrew W; Churchill, Steven E (2009). "Energetic
Competition Between Neandertals and Anatomically Modern Humans" (PDF).
PaleoAnthropology: 96–116. Retrieved October 31, 2011.
* ^ Laleuza-Fox, Carles; Römpler, Holger; et al. (October 25,
2007). "A Melanocortin 1 Receptor Allele Suggests Varying Pigmentation
Among Neanderthals". Science . 318 (5855): 1453–5. Bibcode
:2007Sci...318.1453L. PMID 17962522 . doi :10.1126/science.1147417 . ;
see also Rincon, Paul (October 25, 2007). "
flame-haired\'". BBC News. Retrieved October 25, 2007.
* ^ The 10,000 Year Explosion: How Civilizations Accelerated Human
Evolution (2009). Gregory Cochran and Henry Harpending. Basic Books,
New York, NY, USA}}
* ^ 14:186–198. Hoffecker JF (2009) The spread of modern humans
in Europe. Proc. Nat. Acad. Sci. 106:16040– References 399 Hoffecker
JF, Cleghorn ...
* ^ "
Neanderthal brains focused on vision and movement leaving less
room for social networking". Science Daily. March 19, 2013.
* ^ SINC Servicio de Información y Noticias Científicas. "El
cerebro neandertal era más asimétrico que el del \'
* ^ "
Neanderthal Brain Size at Birth Sheds Light on Human
Evolution". National Geographic. September 9, 2008. Retrieved
September 19, 2009.
* ^ Silberman, Neil. The Oxford Companion to Archaeology, p. 455
(Oxford University Press 2012): "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
* ^ Abramiuk, Marc. The Foundations of Cognitive Archaeology, p.
199 (MIT Press 2012): "the encephalization quotient was slightly
* ^ Froehle, Andrew W.; Churchill, Steven E. (2009). "Energetic
Competition Between Neandertals and Anatomically Modern Humans" (PDF).
* ^ Moskvitch, Katia (September 24, 2010). "
Neanderthals were able
to \'develop their own tools\'".
BBC News . BBC. Retrieved October 1,
* ^ Heyes, Peter; Anastasakis, Konstantinos; de Jong, Wiebren
(2016). "Selection and Use of Manganese Dioxide by Neanderthals" .
Scientific Reports. 6: 22159. ISSN 2045-2322 . PMC 4770591 . PMID
26922901 . doi :10.1038/srep22159 .
* ^ 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
Human Evolution. 49 (1): 71–87. PMID 15869783 . doi
* ^ A B Ghosh, Pallab. "
Neanderthals cooked and ate vegetables."
BBC News. December 27, 2010.
* ^ Lichfield, John (September 30, 2006). "French dig up
Neanderthal \'butcher\'s shop\'". The New Zealand Herald.
* ^ 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):
JSTOR 122870 . PMC 16602 .
PMID 10852955 . doi :10.1073/pnas.120178997 .
* ^ 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. PMC 3060801 . PMID 21445243 .
doi :10.1371/journal.pone.0014769 .
* ^ Henry, A. G.; Brooks, A. S.; Piperno, D. R. (2010).
"Microfossils in calculus demonstrate consumption of plants and cooked
Neanderthal diets (
Shanidar III, Iraq; Spy I and II,
Belgium)" . Proceedings of the National Academy of Sciences. 108 (2):
Bibcode :2011PNAS..108..486H. PMC 3021051 . PMID
21187393 . doi :10.1073/pnas.1016868108 .
* ^ Webb, Jonathan (June 25, 2014). "Oldest human faeces show
Neanderthals ate vegetables". BBC News.
* ^ "
Neanderthal behaviour, diet, and disease inferred from ancient
DNA in dental calculus : Nature". The Guardian. Retrieved March 9,
* ^ Nicola Davis (March 8, 2017). "
Neanderthal dental tartar
reveals plant-based diet – and drugs". The Guardian. Retrieved March
* ^ Shaw, Kate (July 29, 2011). "Sheer Numbers Gave Early Humans
Edge Over Neanderthals". Wired.com.
* ^ Vergano, Dan (April 22, 2014). "
Neanderthals Lived in Small,
Isolated Populations, Gene Analysis Shows". National Geographic.
* ^ 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 .
* ^ Zollikofer, Christoph; Marcia, Ponce; Leon, De; Vandermeersch,
Bernard; Leveque, Francois (2002). "Evidence for Interpersonal
Violence in the St. Césaire Neanderthal" . PNAS. 99 (9): 6444–448.
PMC 122968 . PMID 11972028 . doi :10.1073/pnas.082111899 .
* ^ Gargett, R.H. (1999). "
Middle Palaeolithic burial is not a dead
issue: the view from Qafzeh, Saint-Césaire, Kebara, Amud, and
Dederiyeh". Journal of
Human Evolution. 37: 27–90. doi
* ^ 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 U S A. doi
:10.1073/pnas.1007963107 PMID 20956292
* ^ Mellars P. (2010).
Neanderthal symbolism and ornament
manufacture: The bursting of a bubble? Proc Natl Acad Sci U S A. doi
* ^ 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 .
* ^ 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 .
* ^ "
Neanderthals were not inferior to modern humans, study finds".
ScienceDaily. 30 April 2014.
* ^ E. de Lazaro (18 January 2017). "
Neanderthals Capable of
Incorporating Symbolic Objects into Their Culture, Discovery
Suggests". Sci News.
* ^ C.Q. Choi (8 January 2010). "Heavy Brows, High Art?: Newly
Unearthed Painted Shells Show Neandertals Were
Homo sapiens\' Mental
Equals". Scientific American.
* ^ I. Sample (30 April 2014). "
Neanderthals were not less
intelligent than modern humans, scientists find". The Guardian.
* ^ N. Branan (2010). "
Neandertal Symbolism: Evidence Suggests a
Biological Basis for Symbolic Thought". Scientific American.
* ^ R. S. Solecki (1975). "
Shanidar IV, a
Neanderthal Flower Burial
in Northern Iraq, Science". 190 (4217): 880–881.
* ^ D.J. Sommer (1999). "The
Shanidar IV 'Flower Burial': a
Neanderthal Burial Ritual". Cambridge Archaeological
Journal. 9 (1): 127–129.
Paul B. Pettitt (2002). "The
Neanderthal Dead, exploring
mortuary variability in Middle Paleolithic Eurasia". Before Farming. 1
* ^ "
Neanderthal \'make-up\' discovered". BBC News. 9 January 2010.
Retrieved 20 May 2010.
* ^ "Did
Neanderthals use feathers for fashion?". New Scientist.
Retrieved 16 June 2017.
* ^ "Birds of a Feather:
Neanderthal Exploitation of Raptors and
Corvids". PLOS ONE. 7: e45927. 17 September 2012. doi
* ^ E. Callaway (2014). "
Neanderthals made some of Europe\'s oldest
art". Nature News. doi :10.1038/nature.2014.15805 .
* ^ P. Rincon (1 September 2014). "
Neanderthal \'artwork\' found in
Gibraltar cave". BBC.
* ^ 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) .
Neanderthal Constructions deep in
Bruniquel Cave in
Southwestern France". Nature. 534 (7605): 111–114. Bibcode
:2016Natur.534..111J. ISSN 0028-0836 . PMID 27251286 . doi
* ^ Brown, Cynthia Stokes. Big History. New York, NY: The New
Press, 2008. Print.
* ^ Ovchinnikov, IV; Götherström, A; Romanova, GP; Kharitonov,
VM; Lidén, K; Goodwin, W (2000). "Molecular analysis of Neanderthal
DNA from the northern Caucasus". Nature. 404 (6777): 490–3. PMID
10761915 . doi :10.1038/35006625 .
* ^ Moulson, Geir;
Associated Press (July 20, 2006). "Neanderthal
genome project launches".
MSNBC . Retrieved August 22, 2006.
* ^ 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
* ^ Green RE, Krause J, Ptak SE, et al. (November 2006). "Analysis
of one million base pairs of
Neanderthal DNA". Nature. 444 (7117):
Bibcode :2006Natur.444..330G. PMID 17108958 . doi
* ^ Wade, Nicholas (November 15, 2006). "New Machine Sheds Light on
DNA of Neanderthals". The
New York Times
New York Times . Retrieved May 18, 2009.
* ^ Pennisi, E. (May 2007). "Ancient DNA. No sex please, we're
Neandertals". Science. 316 (5827): 967. PMID 17510332 . doi
* ^ "
Neanderthal bone gives
DNA clues". CNN. Associated Press.
November 16, 2006. Archived from the original on November 18, 2006.
Retrieved May 18, 2009.
* ^ Than, Ker;
LiveScience (November 15, 2006). "Scientists decode
MSNBC . Retrieved May 18, 2009.
* ^ "
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.
* ^ 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.
* ^ 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
. PMID 18692465 . doi :10.1016/j.cell.2008.06.021 .
* ^ A B Elizabeth Pennisi (2009). "Tales of a Prehistoric Human
Genome". Science. 323 (5916): 866–871. PMID 19213888 . doi
* ^ 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. PMC 2725275 . PMID
19661919 . doi :10.1038/emboj.2009.222 .
* ^ A B C "New studies reveal 20 Percent of
lives on in modern humans". Retrieved October 7, 2016.
* ^ "Surprise! 20 Percent of
Genome Lives On in Modern
Humans, Scientists Find". National Geographic. Retrieved October 7,
* ^ "Ancient
Neanderthals - The Smithsonian Institution\'s
Human Origins Program".
* ^ "BBC NEWS - Science/Nature -
* ^ Lalueza-Fox, Carles; Römpler, Holger; Caramelli, David;
Stäubert, Claudia; Catalano, Giulio; Hughes, David; Rohland, Nadin;
Pilli, Elena; Longo, Laura; Condemi, Silvana; Rasilla, Marco de la;
Fortea, Javier; Rosas, Antonio; Stoneking, Mark; Schöneberg, Torsten;
Bertranpetit, Jaume; Hofreiter, Michael (November 30, 2007). "A
Melanocortin 1 Receptor Allele Suggests Varying Pigmentation Among
Neanderthals". Science. 318 (5855): 1453–1455. PMID 17962522 . doi
:10.1126/science.1147417 – via science.sciencemag.org.
Neanderthals did not interbreed with humans, scientists find
Archived April 19, 2014, at the
Wayback Machine .. Telegraph.
Retrieved May 24, 2014.
* ^ Press Association (February 4, 2013). "
to have interbred with human ancestors\'". The Guardian. Retrieved May
* ^ A B Lowery, Robert K.; Uribe, Gabriel; Jimenez, Eric B.; Weiss,
Mark A.; Herrera, Kristian J.; Regueiro, Maria; Herrera, Rene J.
Neanderthal and Denisova genetic affinities with contemporary
humans: Introgression versus common ancestral polymorphisms". Gene.
530 (1): 83–94. ISSN 0378-1119 . PMID 23872234 . doi
Neanderthal breeding idea doubted. BBC. Retrieved March 2,
* ^ A B "Complex History of Admixture between Modern Humans and
Neandertals". American Journal of
Human Genetics. 96 (3): 454–61.
doi :10.1016/j.ajhg.2015.01.006 .
* ^ Kim, BY; Lohmueller, KE. "Selection and Reduced Population Size
Cannot Explain Higher Amounts of
Neandertal Ancestry in East Asian
than in European
Human Populations". American Journal of Human
Genetics. 96 (3): 448–53. PMC 4375557 . PMID 25683122 . doi
* ^ A B . The Daily Mail. Retrieved January 26, 2016.
* ^ A B Zimmer, Carl (March 17, 2016). "Humans Interbred With
Hominins on Multiple Occasions, Study Finds". The
New York Times
New York Times .
Retrieved March 17, 2016.
* ^ Mendez, Fernando L.; et al. (April 7, 2016). "The Divergence of
Neandertal and Modern
Human Y Chromosomes" (PDF). The American Journal
Human Genetics. 98 (4): 728–734. PMC 4833433 . PMID 27058445 .
doi :10.1016/j.ajhg.2016.02.023 . Retrieved April 12, 2016.
* ^ "
DNA points to
Neanderthal breeding barrier". BBC News, Science
& Environment. April 8, 2016. Retrieved April 12, 2016.
* ^ A B Sample, Ian (February 17, 2016). "Oldest known case of
Neanderthal-human sex revealed by
DNA test". the Guardian. Retrieved
May 10, 2016.
* ^ Zimmer, Carl (March 17, 2016). "Humans Interbred With Hominins
on Multiple Occasions, Study Finds". New York Times. Retrieved May 5,
* ^ A B Gokhman D, Lavi E, Prüfer K, Fraga MF, Riancho JA, Kelso
J, Pääbo S, Meshorer E, Carmel L (2014). "Reconstructing the DNA
methylation maps of the
Neandertal and the Denisovan.". Science. 344
(6183): 523–7. PMID 24786081 . doi :10.1126/science.1250368 .
* ^ A B Higham, Tom et al. (August 21, 2014). "The timing and
spatiotemporal patterning of
Neanderthal disappearance". Nature. 512
(512): 306–309. PMID 25143113 . doi :10.1038/nature13621 . CS1
maint: Uses authors parameter (link )(subscription required)
* ^ 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.
* ^ 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. PMID 17300854 . doi :10.1016/j.tree.2007.02.001 . CS1
maint: Uses authors parameter (link )
* ^ "First genocide of human beings occurred 30,000 years ago".
Pravda . October 24, 2007. Retrieved May 18, 2009.
* ^ Diamond, Jared M. (1992). The third chimpanzee: the evolution
and future of the human animal.
New York City
New York City :
HarperCollins . p. 52.
ISBN 0-06-098403-1 .
OCLC 60088352 .
* ^ Currat, Mathias; Excoffier, Laurent (2004). "Modern Humans Did
Not Admix with
Neanderthals during Their Range Expansion into Europe"
. PLoS Biology. 2 (12): e421. PMC 532389 . PMID 15562317 . doi
* ^ Wong, Kate (August 1, 2009). "The Mysterious Downfall of the
Neandertals". Scientific American.
* ^ 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 .
* ^ Wilford, John Noble (November 2, 2011). "
Fossil Teeth Put
Humans in Europe Earlier Than Thought".
New York Times
New York Times . Retrieved
August 27, 2014.
* ^ Wolff, H. "Result Filters." National Center for Biotechnology
Information. U.S. National Library of Medicine, July 2010. Web.
October 22, 2014.
* ^ McKie, Robin (March 1, 2015). "How hunting with wolves helped
humans outsmart the Neanderthals". The Guardian. Retrieved May 5,
* ^ Stringer, Chris (2012). "Evolution: What makes a modern human".
Nature. 485 (7396): 33–5.
Bibcode :2012Natur.485...33S. PMID
22552077 . doi :10.1038/485033a .
* ^ Steensby, H. P. (1907). "Racestudier i Danmark" (PDF).
Geographical Journal (in Danish). Royal Library, Denmark. Retrieved
July 6, 2017.
* ^ Dan Jones: The
New Scientist 193.2007, H.
2593 (March 3), 28–32. Modern Humans,
Neanderthals May Have
Interbred; Humans and
Neanderthals interbred Archived February 22,
2009, at the
Wayback Machine .
* ^ Foley, Jim (July 31, 2000). "The
Lagar Velho 1 Skeleton".
Fossil Hominids FAQ.
TalkOrigins Archive . Retrieved July 6, 2017.
* ^ 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.
* ^ "Not a lasting last for the Neandertals". john hawks weblog .
September 13, 2006. Retrieved July 6, 2017.
* ^ 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):
JSTOR 30052409 . PMC
1859909 . PMID 17085588 . doi :10.1073/pnas.0608443103 .
* ^ Wade, Nicholas (July 26, 2012). "Genetic Data and Fossil
Evidence Tell Differing Tales of
Human Origins". The New York Times.
Retrieved May 5, 2015.
* ^ 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
is Present Among All Non-African Populations". Molecular Biology and
Evolution. 28 (7): 1957–62. PMID 21266489 . doi
* ^ Viegas, Jennifer (July 18, 2011). "All Non-Africans Part
Genetics Confirm". DNews.
* ^ "
Neandertal ancestry "Iced"". john hawks weblog. August 15,
2012. Retrieved July 6, 2017.
* ^ "Resurrecting Surviving
Neandertal Lineages from Modern Human
Genomes". Science. January 29, 2014. Retrieved February 2, 2014.
* ^ Mitchell, Alanna (January 30, 2012). "
Into a Tell-All".
NYTimes . Retrieved January 31, 2012.
* ^ Lohse, Konrad; Frantz, Laurent A. F. (2013). "Maximum
likelihood evidence for
Neandertal admixture in Eurasian populations
from three genomes". Populations and Evolution. 1307: 8263. Bibcode
:2013arXiv1307.8263L. arXiv :1307.8263 .
* ^ Jha, Alok (August 14, 2012). "Study casts doubt on
Neanderthal interbreeding theory".
The Guardian . Retrieved
February 19, 2015.
* ^ Hawks, John (2013). "Significance of
Neandertal and Denisovan
Human Evolution". Annual Review of Anthropology. 42:
433–49. doi :10.1146/annurev-anthro-092412-155548 .
* ^ Mason, Paul H.; Short, Roger V. (2011). "Neanderthal-human
Hybrids". Hypothesis. 9: e1. doi :10.5779/hypothesis.v9i1.215 .
* ^ Viegas, Jennifer (March 27, 2013). "First Love Child of Human,
Neanderthal Found". Discovery.
* ^ 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. PMC 3609795 . PMID 23544098 . doi
* ^ The
Neanderthal Dead, exploring mortuary variability in middle
Paul B. Pettitt (2002)
* ^ "
Homo neanderthalensis–The Neanderthals". Australian Museum.
Retrieved June 26, 2014.
* ^ Bischoff, J; Shamp, Donald D.; Aramburu, Arantza; Arsuaga, Juan
Luis; Carbonell, Eudald; Bermudez De Castro, J.M. (2003). "The Sima de
los Huesos Hominids Date to Beyond U/Th Equilibrium (>350kyr) and
Perhaps to 400–500kyr: New Radiometric Dates". Journal of
Archaeological Science. 30 (3): 275–280. doi :10.1006/jasc.2002.0834
* ^ 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. PMID 9300343
. doi :10.1006/jhev.1997.0133 .
* ^ Kreger, C. David. "
Homo neanderthalensis". ArchaeologyInfo.com.
Retrieved May 16, 2009.
* ^ Mcdermott, F; Grün, R; Stringer, Cb; Hawkesworth, Cj (May
1993). "Mass-spectrometric U-series dates for Israeli
Neanderthal/early modern hominid sites". Nature. 363 (6426): 252–5.
Bibcode :1993Natur.363..252M. ISSN 0028-0836 . PMID 8387643 . doi
* ^ Rink, W. Jack; Schwarcz, H.P.; Lee, H.K.; Rees-Jones, J.;
Rabinovich, R.; Hovers, E. (August 2002). "Electron spin resonance
(ESR) and thermal ionization mass spectrometric (TIMS) 230Th/234U
dating of teeth in Middle Paleolithic layers at Amud Cave, Israel".
Geoarchaeology. 16 (6): 701–717. doi :10.1002/gea.1017 .
* ^ Valladas, Hélène; Merciera, N.; Frogeta, L.; Hoversb, E.;
Joronc, J.L.; Kimbeld, W.H.; Rak, Y. (March 1999). "TL Dates for the
Neanderthal Site of the Amud Cave, Israel". Journal of Archaeological
Science. 26 (3): 259–268. doi :10.1006/jasc.1998.0334 .
* ^ 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
El Sidrón Cave
(Asturias, Northern Spain)". Archaeometry. 55 (1): 148–158. doi
:10.1111/j.1475-4754.2012.00671.x . CS1 maint: Uses authors parameter
* ^ Hayes, Jacqui (November 2, 2006). "Humans and Neanderthals
interbred". Cosmos. Archived from the original on February 22, 2009.
Retrieved May 17, 2009.
Neanderthal image by Kupka, based on Boule, 1909, in
Humanity\'s Journeys Dr. Kathryn Denning, 2005. Retrieved March 17,
* 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
* 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
DNA Contribution to Early Modern Humans" . PLoS Biology. 2 (3): e57.
PMC 368159 . PMID 15024415 . doi :10.1371/journal.pbio.0020057 .
* 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):
Bibcode :2005Natur.435..332W. PMID 15902255 . doi
* 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. PMC 2811729 . PMID 20111705 . doi
:10.1371/journal.pone.0008880 . Lay summary – ScienceDaily (January
* Derev'anko, Anatoliy P.; Powers, William Roger; Shimkin, Demitri
Boris (1998). The Paleolithic of Siberia: new discoveries and
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 .
* Sankararaman, Sriram; Mallick, Swapan; Dannemann, Michael;
Prüfer, Kay; Kelso, Janet; Patterson, Nick; Reich, David (2014). "The
genomic landscape of
Neanderthal ancestry in present-day humans" .
Nature. 507 (7492): 354–357.
Bibcode :2014Natur.507..354S. PMC
4072735 . PMID 24476815 . doi :10.1038/nature12961 .
* Vattathil, S.; Akey, J.M. (2015). "Small amounts of archaic
admixture provide big insights into human history". Cell. 163 (2):
281–284. PMID 26451479 . doi :10.1016/j.cell.2015.09.042 .
Wikimedia Commons has media related to HOMO NEANDERTHALENSIS .
Wikibooks has a book on the topic of: INTRODUCTION TO