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Homo
Homo
mousteriensis[1] Palaeoanthropus neanderthalensis[2]

Neanderthals
Neanderthals
(UK: /niˈændərˌtɑːl/, also US: /neɪ-, -ˈɑːn-, -ˌtɔːl, -ˌθɔːl/),[3][4] more rarely known as Neandertals,[a] were archaic humans that became extinct about 40,000 years ago.[8][9][10][11][12][13] They seem to have appeared in Europe and later expanded into Southwest, Central and Northern Asia. There, they left hundreds of stone tool assemblages. Almost all of those younger than 160,000 years are of the so-called Mousterian
Mousterian
techno-complex, which is characterised by tools made out of stone flakes.[14] Neanderthals
Neanderthals
are considered either a distinct species, Homo neanderthalensis,[15][16][17] or more rarely[18] a subspecies of Homo sapiens (H. s. neanderthalensis).[19][20] A 2010 report found that non-Africans and Neanderthals
Neanderthals
share 99.7% of their DNA[21] and are hence much more closely related than to their closest non-human relative, the chimpanzee (98.8%). Another report in 2017 concluded that 1.8 to 2.6 percent of the genes of non-Africans come from Neanderthals.[22] Compared to modern humans, Neanderthals
Neanderthals
were stockier, with shorter legs and a bigger body. In conformance with Bergmann's rule, this likely was a Darwinian adaptation to preserve heat in cold climates. Male and female Neanderthals
Neanderthals
had cranial capacities averaging 1,600 cm3 (98 cu in) and 1,300 cm3 (79 cu in), respectively,[23][24] extending to 1,736 cm3 (105.9 cu in) in the male Amud 1.[25] This is within range of anatomically modern humans, whose cranial capacity range from 1,230 to 1,880 cm3 (75 to 115 cu in).[26] Males stood 164 to 168 cm (65 to 66 in) and females 152 to 156 cm (60 to 61 in) tall.[27] The Neanderthal genome project
Neanderthal genome project
revealed in 2010 that, through interbreeding, Neanderthals
Neanderthals
may have contributed to the DNA
DNA
of modern humans, likely between 50,000 and 60,000 years ago.[28][29][30][31] Today, this is apparent in the genome of most people living outside sub-Saharan Africa, as well as in some sub-Saharan Africans. Subsequent studies suggested there may have been three episodes of interbreeding. The first would have occurred soon after modern humans left Africa. The second would have occurred after the ancestral Melanesians had branched off—these people seem to have thereafter bred with Denisovans. The third would have involved Neanderthals
Neanderthals
and the ancestors of East Asians only.[32][33][34]

Contents

1 Name 2 Classification 3 Origins 4 Discovery

4.1 Timeline of research

5 Habitat and range 6 Anatomy 7 Behaviour

7.1 Question of art and adornment

8 Genome

8.1 Background 8.2 Interbreeding with modern humans 8.3 Epigenetics

9 Extinction

9.1 Climate change 9.2 Coexistence with modern humans

10 Interbreeding hypotheses

10.1 Pre-2010 interbreeding hypotheses 10.2 Interbreeding hypotheses since 2010

11 Specimens

11.1 Notable European Neanderthals 11.2 Notable Southwest Asian Neanderthals 11.3 Notable Central Asian Neanderthal 11.4 Chronology

11.4.1 Mixed with H. heidelbergensis
H. heidelbergensis
traits 11.4.2 H. neanderthalensis fossils 11.4.3 H. s. sapiens with traits reminiscent of Neanderthals

12 In popular culture 13 See also 14 Notes 15 References 16 Further reading 17 External links

Name[edit] Neanderthals
Neanderthals
are named after one of the first sites where their fossils were discovered in the 19th century in the Neandertal in Erkrath, Germany.[b] Thal is an older spelling of the German word Tal (with the same pronunciation), which means "valley" (cognate with English dale).[c][36][37] Neanderthal 1
Neanderthal 1
was known as the " Neanderthal
Neanderthal
cranium" or "Neanderthal skull" in anthropological literature, and the individual reconstructed on the basis of the skull was occasionally called "the Neanderthal man".[38] The binomial name Homo
Homo
neanderthalensis—extending the name " Neanderthal
Neanderthal
man" from the individual type specimen to the entire group—was first proposed by the Anglo-Irish geologist William King in 1864, although that same year King changed his mind and thought that the Neanderthal
Neanderthal
fossil was distinct enough from humans to warrant a separate genus.[6] Nevertheless, King's name had priority over the proposal put forward in 1866 by Ernst Haeckel, Homo
Homo
stupidus.[36] The practice of referring to "the Neanderthals" and "a Neanderthal" emerged in the popular literature of the 1920s.[39] The German pronunciation of Neandertaler is [neˈandɐˌtʰaːlɐ] in the International Phonetic Alphabet. In British English, "Neanderthal" is pronounced with the /t/ as in German, but different vowels (IPA: /niːˈændərtɑːl/).[40][41][42] 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/),[43] although scientists typically use the /t/ as in German.[44][45] Classification[edit]

Anatomical comparison of skulls of Homo
Homo
sapiens (left) and Homo neanderthalensis (right) (in Cleveland Museum of Natural History) Features compared are the braincase shape, forehead, browridge, nasal bone, projection, cheek bone angulation, chin and occipital contour

Scientists still dispute whether Neanderthals
Neanderthals
should be classified as a distinct species – Homo
Homo
neanderthalensis – or as Homo sapiens neanderthalensis, placing Neanderthals
Neanderthals
as a subspecies of H. sapiens.[46][47] DNA
DNA
researcher Svante Pääbo
Svante Pääbo
referred to the ongoing "taxonomic wars" over whether Neanderthals
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."[48] During the early 20th century, the prevailing view was heavily influenced by Arthur Keith
Arthur Keith
and Marcellin Boule, who wrote the first scientific description of a nearly complete Neanderthal
Neanderthal
skeleton.[49] Senior members of their respective national paleontological institutes and among the most eminent paleoanthropologists of their time,[50][51] both men argued that the primitive traits of this skeleton meant it could not be a direct ancestor of Homo
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.[52] During the 1930s, scholars Ernst Mayr, George Gaylord Simpson and Theodosius Dobzhansky
Theodosius Dobzhansky
reinterpreted the existing fossil record and came to different conclusions.[53] Neanderthal
Neanderthal
man was classified as Homo
Homo
sapiens neanderthalensis – an early subspecies contrasted with what was now called Homo
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
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.[54] Origins[edit] See also: Accretion model of Neanderthal
Neanderthal
origins

Early pre-Neanderthal (stage 1, 450 ka, male)

Pre-Neanderthal (stage 2, 350 ka, male)

Early Neanderthal (stage 3, 130 ka, female)

Classic Neanderthal (stage 4, male, 50 ka)

The evolution of Neanderthals
Neanderthals
according to the accretion model. Note the larger nose and rounder eyes of the Classic Neanderthal.

Both Neanderthals
Neanderthals
and living humans are thought to have evolved from Homo
Homo
erectus. In the earliest known migration wave into Eurasia dated to 1.81 million years ago (Ma), Homo
Homo
erectus left Africa most probably via the Levant
Levant
and reached Georgia (fossils of Dmanisi). Hominins had reached China by 1.7 Ma[55] and Iberia (Spain) by 1.4 Ma.[56] The discoverers of fragmented bones in Spain
Spain
(Iberia) dated to 1.2 million years, assigned to a new species Homo
Homo
antecessor, argue these are the remains of the ancestors of Neanderthals
Neanderthals
and of the older species Homo heidelbergensis, an interpretation rejected by most anthropologists.[57] A large number of molecular clock genetic studies place the divergence time of the Neanderthal
Neanderthal
and modern human lineages between 800,000 and 400,000 years ago.[58][35][59][60][61][62][63][64][65] For this reason, most scholars believe Neanderthals
Neanderthals
descend, via Homo heidelbergensis, from another Homo
Homo
erectus migration out of Africa that would have occurred in this time frame. Parts of the Homo
Homo
erectus population that stayed in Africa would have evolved, perhaps through the intermediate Homo
Homo
rhodesiensis, into early anatomically modern humans by 200,000 years ago or earlier. Neanderthal
Neanderthal
traits are present in Homo
Homo
heidelbergensis specimens beginning between 600,000 and 350,000 years ago.[66][67][68][69] As of 1998[update], there is a fossil gap in Europe between 300 and 243 ka (MIS 8); no hominin has ever been dated to this period.[70] Conventionally, therefore, European hominins younger than 243,000 years old are called Neanderthals.[70][71] The quality of the fossil record greatly increases from 130,000 years ago onwards.[72] Specimens younger than this date make up the bulk of known Neanderthal
Neanderthal
skeletons and were the first whose anatomy was comprehensively studied. They are known as typical or Classic Neanderthals.[73][74] In morphological studies, the latter term may also be used in a narrower sense for Neanderthals
Neanderthals
younger than 71,000 years old (MIS 4 and 3).[70] Discovery[edit] Neanderthal
Neanderthal
fossils were first discovered in 1829 in the Engis
Engis
caves (the partial skull dubbed Engis
Engis
2), in what is now Belgium by Philippe-Charles Schmerling
Philippe-Charles Schmerling
and the Gibraltar 1
Gibraltar 1
skull in 1848 in the Forbes' Quarry, Gibraltar, both prior to the type specimen discovery in a limestone quarry (Feldhofer Cave), located in the Düssel
Düssel
River's Neandertal in Erkrath, Germany
Germany
(about 12 km (7 mi) east of Düsseldorf), in August 1856, three years before Charles Darwin's On the Origin of Species was published.[75][76][77] The type specimen, dubbed Neanderthal
Neanderthal
1, consisted of a skull cap, two femora, three bones of the right arm, two of the left arm, parts of the left ilium, fragments of a scapula, and ribs. The workers who recovered the objects originally thought them to be the remains of a cave bear. However, they eventually gave the material to amateur naturalist Johann Carl Fuhlrott, who turned the fossils over to anatomist Hermann Schaaffhausen. To date, the bones of over 400 Neanderthals
Neanderthals
have been found.[78] Timeline of research[edit]

Engis
Engis
2, child (1829)

Gibraltar
Gibraltar
1, female (1848)

Neanderthal
Neanderthal
1, male (upper skull 1856, left-cheek 2000)

Spy 2 skull, sex unclear[79] (1886)

Krapina
Krapina
3, female (1899)

1829: A damaged skull of a Neanderthal
Neanderthal
child, Engis
Engis
2, is discovered in Engis, Netherlands (now Belgium). 1848: A female Neanderthal
Neanderthal
skull, Gibraltar
Gibraltar
1, is found in Forbes' Quarry, Gibraltar, but its importance is not recognised. 1856: Limestone
Limestone
miners discover the Neanderthal-type specimen, Neanderthal
Neanderthal
1, in Neandertal, western Prussia (now Germany). 1864: William King is the first to recognise Neanderthal 1
Neanderthal 1
as belonging to a separate species, for which he gives the scientific name Homo
Homo
neanderthalensis. He then changed his mind on placing it in the Homo
Homo
genus, arguing that the upper skull was different enough to warrant a separate genus since, to him, it had likely been "incapable of moral and theistic conceptions."[6] 1880: The mandible of a Neanderthal
Neanderthal
child is discovered in a secure context in Šipka
Šipka
cave, in the Austro-Hungarian Empire (now the Czech Republic), associated with cultural debris, including hearths, Mousterian
Mousterian
tools, and bones of extinct animals. 1886: Two well-preserved Neanderthal
Neanderthal
skeletons are found at Spy, Belgium, making the hypothesis that Neanderthal 1
Neanderthal 1
was only a diseased modern human difficult to sustain.[80] 1899: Sand excavation workers find hundreds of fragmentary Neanderthal remains representing at least 12 and likely as much as 70 individuals on a hill in Krapina, in the Austro-Hungarian Empire (now Croatia). 1908: A very well preserved Neanderthal, La Chapelle-aux-Saints
La Chapelle-aux-Saints
1, is found in its eponymous site in France,[81] said by the excavators to be a burial, a claim still heatedly contested.[82][83][84] For historical reasons it remains the most famous Neanderthal skeleton.[72]:15 1912: Marcellin Boule
Marcellin Boule
publishes his now discredited influential study of Neanderthal
Neanderthal
skeletal morphology based on La Chapelle-aux-Saints
La Chapelle-aux-Saints
1. 1953–1957: Ten Neanderthal
Neanderthal
skeletons are excavated in Shanidar
Shanidar
Cave, Iraqi Kurdistan, by Ralph Solecki and colleagues. 1975: Erik Trinkaus's study of Neanderthal
Neanderthal
feet strongly argues that Neanderthals
Neanderthals
walked like modern humans. 1981: The site of Bontnewydd, Wales
Wales
yielded an early Neanderthal tooth, the most north-western Neanderthal
Neanderthal
remain ever. 1987: Israeli Neanderthal
Neanderthal
Kebara 2
Kebara 2
is dated (by TL and ESR) to 60,000 BP, thus later than the Israeli anatomically modern humans dated to 90,000 and 80,000 BP at Qafzeh
Qafzeh
and Skhul. 1994: The site of Sidrón Cave, Spain, is discovered where the remains of 12 males and females were to be found. Mitochondrial DNA
DNA
studies would show that the adult males were genetically related, but the adult females were not, suggesting female exogamy.[85][86] 1997: Matthias Krings et al. are the first to amplify Neanderthal mitochondrial DNA
DNA
(mtDNA) using a specimen from Feldhofer grotto in the Neander valley.[35] 1997–2000: A small part of the skull of Neanderthal
Neanderthal
1, the Neanderthal
Neanderthal
type fossil discovered in 1856, is found in a dump of limestone mining debris near the original discovery site.[87] 1998: The body of a c. 24,000-year-old early Upper Paleolithic
Upper Paleolithic
child is discovered at Abrigo do Lagar Velho, Portugal, and is described as presenting a mosaic of anatomically modern human and more archaic features, reminiscent of Neanderthals.[47] 2005: The Max Planck Institute for Evolutionary Anthropology
Max Planck Institute for Evolutionary Anthropology
and associated institutions launch the Neanderthal genome project
Neanderthal genome project
to sequence the Neanderthal
Neanderthal
nuclear genome.[88] 2010: 1–4% of the DNA
DNA
of living non-African humans are found by the Max Planck Institute to likely come from Neanderthals,[29][89] a result confirmed in 2012,[90][91] and refined to 1.5–2.1% in 2014.[58] 2013: In the midst of a hundred-year-old debate over the existence of Neanderthal
Neanderthal
deliberate burials, a team of French researchers reasserted a heavily contested 1908 claim that the Chapelle-aux-Saints 1 skeleton was deliberately buried.[92] 2014: Some researchers express worry that almost all theories for the Neanderthals' extinction assume that there is something modern humans had that Neanderthals
Neanderthals
did not.[93] 2014: The most comprehensive dating ever of Neanderthal
Neanderthal
bones and tools from hundreds of sites in Europe is carried out. It suggested that European Neanderthals
Neanderthals
died out between 41,000 and 39,000 years ago, which coincides with the start of a very cold period in Europe and is no more than 5,000 years after anatomically-modern Homo
Homo
sapiens reached the continent.[13][94]

Habitat and range[edit] Further information: Southwest Asian Neanderthals
Southwest Asian Neanderthals
and List of Neanderthal
Neanderthal
sites

Sites where typical Neanderthal
Neanderthal
fossils have been found

Early Neanderthals, living before the Eemian
Eemian
interglacial (130 ka), are poorly known and come mostly from European sites. From 130 ka onwards, the quality of the fossil record increases dramatically. From then on, Neanderthal
Neanderthal
remains are found in Western, Central, Eastern, and Mediterranean Europe,[95] as well as Southwest, Central, and Northern Asia up to the Altai Mountains
Altai Mountains
in Siberia. No Neanderthal
Neanderthal
has ever been found outside Western Eurasia, namely neither to the south of Jerusalem (Shuqba), nor further east than Kazakhstan (Denisova, Russia), nor to the north of Wales
Wales
(Bontnewydd),[96][97] although it is difficult to assess the limits of their northern range because glacial advances destroy most human remains, the Bontnewydd tooth being exceptional. Middle Palaeolithic
Middle Palaeolithic
artifacts have been found up to 60° N on the Russian plains.[98] There likely never were more than 70,000 Neanderthals
Neanderthals
at any given time.[99] Anatomy[edit] Main article: Neanderthal
Neanderthal
anatomy

Neanderthal anatomy
Neanderthal anatomy
differed from modern humans in that they had a more robust build and distinctive morphological features, especially on the cranium, which gradually accumulated more derived aspects as it was described by Marcellin Boule,[100] particularly in certain isolated geographic regions. These include shorter limb proportions, a wider, barrel-shaped rib cage, a reduced chin, and a large nose, being at the modern human higher end in both width and length,[d] and started somewhat higher on the face than in modern humans.[71] Evidence suggests they were much stronger than modern humans, with particularly strong arms and hands,[101][102] while they were comparable in height; based on 45 long bones from at most 14 males and 7 females, Neanderthal
Neanderthal
males averaged 164 to 168 cm (65 to 66 in) and females 152 to 156 cm (60 to 61 in) tall.[27] 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.[103] A 2007 genetic study suggested some Neanderthals
Neanderthals
may have had red hair and blond hair, along with a light skin tone.[104] Gregory Cochran and Henry Harpending, in the book The 10,000 Year Explosion, investigated whether it is accurate to depict Neanderthals as having hair patterns similar to anatomically modern humans. They concluded that, "We don't yet know for sure, but it seems likely that, as part of their adaptation to cold, Neanderthals
Neanderthals
were furry. Chimpanzees have ridges on their finger bones that stem from the way that they clutch their mothers' fur as infants. Modern humans don't have these ridges, but Neanderthals
Neanderthals
do."[105] In 2017, researchers using 3D reconstructions of nasal cavities and Computational Fluid Dynamics
Computational Fluid Dynamics
techniques have found that Neanderthals and modern humans both adapted their noses (independently and in a convergent way) to help breathe in cold and dry conditions.[106]  The large nose seen in Neanderthals, as well as Homo
Homo
heidelbergensis, affected the shape of the skull and the muscle attachments, and gave them a weaker bite force than in modern humans.[107] In The Spread of Modern Humans in Europe (2002) John F. Hoffecker, writes " Neanderthal
Neanderthal
sites show no evidence of tools for making tailored clothing. There are only hide scrapers, which might have been used to make blankets or ponchos. This is in contrast to Upper Paleolithic (modern human) sites, which have an abundance of eyed bone needles and bone awls. Moreover, microwear analysis of Neanderthal hide scrapers shows that they were used only for the initial phases of hide preparation, and not for the more advanced phases of clothing production.[108] A 2013 study of Neanderthal
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.[109] Neanderthals
Neanderthals
are known for their large cranial capacity, which at 1,600 cm3 (98 cu in) is larger on average than that of modern humans. One study has found that drainage of the dural venous sinuses (low pressure blood vessels that run between the meninges and skull leading down through the skull) in the occipital lobe region of Neanderthal
Neanderthal
brains appears more asymmetric than other hominid brains.[110] In 2008, a group of scientists produced a study using three-dimensional computer-assisted reconstructions of Neanderthal infants based on fossils found in Russia and Syria. It indicated that Neanderthal
Neanderthal
and modern human brains were the same size at birth, but that by adulthood, the Neanderthal
Neanderthal
brain was larger than the modern human brain.[111] They had almost the same degree of encephalisation (i.e. brain-to-body-size ratio) as modern humans.[112][113] The Neanderthal
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).[103]

Behaviour[edit]

Levallois point – Beuzeville, France

Main article: Neanderthal
Neanderthal
behavior See also: Pleistocene human diet Neanderthals
Neanderthals
made stone tools,[114] used fire,[115] and were hunters. The consensus on their behaviour ends there. It had actually long been debated whether Neanderthals
Neanderthals
were hunters or scavengers,[71] but the discovery of the pre- Neanderthal
Neanderthal
Schöningen wooden spears in Germany helped settle the debate in favour of hunting. Most available evidence suggests they were apex predators,[116][117] and fed on red deer, reindeer, ibex, wild boar, aurochs and on occasion mammoth, straight-tusked elephant and rhinoceros.[71][118][119][120] They appear to have occasionally used vegetables as fall-back food,[117][121] revealed in the 2000s and 2010s by isotope analysis of their teeth and study of their coprolites (fossilised faeces).[119][122] Dental analysis of specimens from Spy, Belgium
Spy, Belgium
and El Sidrón, Spain[123] in 2017 suggested that these Neanderthals
Neanderthals
had a wide-ranging diet, and that those "from El Sidrón showed no evidence of meat eating" at all and seemed to have lived on "a mixture of forest moss, pine nuts and a mushroom known as split gill".[124] The size and distribution of Neanderthal
Neanderthal
sites, along with genetic evidence, suggests Neanderthals
Neanderthals
lived in much smaller and more sparsely distributed groups than anatomically-modern Homo sapiens.[125][126] The bones of twelve Neanderthals
Neanderthals
were discovered at El Sidrón cave in northwestern Spain. They are believed to have been a group killed and butchered about 50,000 years ago. Analysis of the mt DNA
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".[127] The bones of the El Sidrón group show signs of defleshing, suggesting that they were victims of cannibalism.[127] 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.[128] Claims that Neanderthals
Neanderthals
deliberately buried their dead, and if they did, whether such burials had any symbolic meaning,[72]:158–60 are heavily contested.[83][84][129] 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
Neanderthal
specimen had been deliberately buried,[92] and this has in turn been heavily criticised.[82]

Question of art and adornment[edit] Whether Neandethals created art and used adornments, which would indicate a capability for complex symbolic thought, remains unresolved. A 2010 paper on radiocarbon dates cast doubt on the association of Châtelperronian
Châtelperronian
beads with Neanderthals,[130] and Paul Mellars considered the evidence for symbolic behavior to have been refuted.[131] This conclusion, however, is controversial, and others such as Jean-Jacques Hublin
Jean-Jacques Hublin
and colleagues have re-dated material associated with the Châtelperronian
Châtelperronian
artefacts[132] and used proteomic evidence to restate the challenged association with Neanderthals.[133] A very large number of other claims of Neanderthal
Neanderthal
art, adornment, and structures have been made. These are often taken by the media as showing Neanderthals
Neanderthals
were capable of symbolic thought,[134][135] or were "mental equals" to anatomically modern humans.[136][137] As evidence of symbolism, none of them are widely accepted,[138] although the same is true for Middle Palaeolithic
Middle Palaeolithic
anatomically modern humans. Among many others:

Flower pollen on the body of pre- Neanderthal
Neanderthal
Shanidar
Shanidar
4, Iraq, had in 1975 been argued to be a flower burial.[139] Once popular, this theory is no longer accepted.[140][141] Pigmented shells from Murcia, Spain, were argued in 2009 to be Neanderthal
Neanderthal
make-up containers.[142] 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[143] took to mean Neanderthals
Neanderthals
wore bird feathers as personal adornments.[144] Deep scratches were found in 2012 on a cave floor underlying Neanderthal
Neanderthal
layer in Gorham's Cave, Gibraltar, which some have controversially[145] interpreted as art.[146] Two 176,000-year-old stalagmite ring structures, several metres wide, were reported in 2016 more than 300 metres from the entrance within Bruniquel Cave, France. The authors claim artificial lighting would have been required as this part of the cave is beyond the reach of daylight and that the structures had been made by early Neanderthals, the only humans in Europe at this time.[147] In 2015, a study argued that a number of 130,000-year-old eagle talons found in a cache near Krapina, Croatia along with Neanderthal
Neanderthal
bones, had been modified to be used as jewellery.[148][149]

All of these appeared only in single locations. Yet in 2018, using uranium-thorium dating methods,[150] red painted symbols comprising a scalariform (ladder shape), a negative hand stencil, and red lines and dots on the cave walls of three Spanish caves 700 km (430 mi) apart were dated to at least 64,000 years old.[151] If the dating is correct, they were painted before the time anatomically modern humans are thought to have arrived in Europe. Paleoanthropologist John D. Hawks argues these findings demonstrate Neanderthals
Neanderthals
were capable of symbolic behavior previously thought to be unique to modern humans.[152] Genome[edit] Further information: Neanderthal genome
Neanderthal genome
project Background[edit] Early investigations concentrated on mitochondrial DNA
DNA
(mtDNA), which, owing to strictly matrilineal inheritance and subsequent vulnerability to genetic drift, is of limited value in evaluating the possibility of interbreeding of Neanderthals
Neanderthals
with Cro-Magnon
Cro-Magnon
people. In 1997, geneticists were able to extract a short sequence of DNA
DNA
from Neanderthal
Neanderthal
bones.[153] The extraction of mt DNA
DNA
from a second specimen was reported in 2000, and showed no sign of modern human descent from Neanderthals.[61] In July 2006, the Max Planck Institute for Evolutionary Anthropology and 454 Life Sciences announced that they would sequence the Neanderthal genome
Neanderthal genome
over the next two years. This genome was expected to be roughly the size of the human genome, three-billion base pairs, and share most of its genes. It was hoped the comparison would expand understanding of Neanderthals, as well as the evolution of humans and human brains.[154] Svante Pääbo
Svante Pääbo
has tested more than 70 Neanderthal
Neanderthal
specimens. The Neanderthal genome
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.[155] From mt DNA
DNA
analysis estimates, the two shared a common ancestor about 500,000 years ago. An article[88] 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.[156] A 2007 study pushes the point of divergence back to around 800,000 years ago.[157] Edward Rubin of the Lawrence Berkeley National Laboratory
Lawrence Berkeley National Laboratory
states recent genome testing of Neanderthals
Neanderthals
suggests human and Neanderthal DNA
DNA
are some 99.5% to nearly 99.9% identical.[158][159] Geneticists first sequenced the entire genome of a Neanderthal
Neanderthal
in 2013 by extracting it from the phalanx bone of a 50,000-year-old Siberian Neanderthal.[160][161][58] Interbreeding with modern humans[edit] On November 16, 2006, Lawrence Berkeley National Laboratory
Lawrence Berkeley National Laboratory
issued a press release suggesting Neanderthals
Neanderthals
and ancient humans probably did not interbreed.[162] Edward M. Rubin, director of the U.S. Department of Energy's Lawrence Berkeley National Laboratory
Lawrence Berkeley National Laboratory
and the Joint Genome Institute (JGI), sequenced a fraction (0.00002) of genomic nuclear DNA (nDNA) from a 38,000-year-old Vindia Neanderthal
Neanderthal
femur. They calculated the common ancestor to be about 353,000 years ago, and a complete separation of the ancestors of the groups about 188,000 years ago.[163] Their results show the genomes of modern humans and Neanderthals
Neanderthals
are at least 99.5% identical, but despite this genetic similarity, and despite the two groups having coexisted in the same geographic region for thousands of years, Rubin and his team did not find any evidence of any significant interbreeding between the two. Rubin said, "While unable to definitively conclude that interbreeding between the two species of humans did not occur, analysis of the nuclear DNA
DNA
from the Neanderthal
Neanderthal
suggests the low likelihood of it having occurred at any appreciable level."[163] In 2008 Richard E. Green et al. from Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany, published the full sequence of Neanderthal
Neanderthal
mitochondrial DNA
DNA
(mtDNA) and suggested " Neanderthals
Neanderthals
had a long-term effective population size smaller than that of modern humans."[62] In the same publication, it was disclosed by Svante Pääbo
Svante Pääbo
that in the previous work at the Max Planck Institute, "Contamination was indeed an issue," and they eventually realised that 11% of their sample was modern human DNA.[164][165] Since then, more of the preparation work has been done in clean areas and 4-base pair 'tags' have been added to the DNA
DNA
as soon as it is extracted so the Neanderthal
Neanderthal
DNA
DNA
can be identified.

Geneticist at the Max Planck Institute for Evolutionary Anthropology extracting ancient DNA

With 3 billion nucleotides sequenced, analysis of about ⅓ showed no sign of admixture between modern humans and Neanderthals, according to Pääbo. This concurred with the work of Noonan from two years earlier. The variant of microcephalin common outside Africa, which was suggested to be of Neanderthal
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.[164] However, an analysis of a first draft of the Neanderthal genome
Neanderthal genome
by the same team released in May 2010 indicates interbreeding may have occurred.[29][89] "Those of us who live outside Africa carry a little Neanderthal
Neanderthal
DNA
DNA
in us," said Pääbo, who led the study. "The proportion of Neanderthal-inherited genetic material is about 1 to 4 percent [later refined to 1.5 to 2.1 percent].[58] It is a small but very real proportion of ancestry in non-Africans today," says Dr. David Reich of Harvard Medical School, who worked on the study. This research compared the genome of the Neanderthals
Neanderthals
to five modern humans from China, France, sub-Saharan Africa, and Papua New Guinea.[89] This indicates a gene flow from Neanderthals
Neanderthals
to modern humans, i.e., interbreeding between the two populations. Since the three non-African genomes show a similar proportion of Neanderthal
Neanderthal
sequences, the interbreeding must have occurred early in the migration of modern humans out of Africa, perhaps in the Middle East. No evidence for gene flow in the direction from modern humans to Neanderthals
Neanderthals
was found. Gene flow
Gene flow
from modern humans to Neanderthals
Neanderthals
would not be expected if contact occurred between a small colonising population of modern humans and a much larger resident population of Neanderthals. A very limited amount of interbreeding could explain the findings, if it occurred early enough in the colonisation process.[89] It is suggested that 20 percent of Neanderthal
Neanderthal
DNA
DNA
survived in modern humans, notably expressed in the skin, hair and diseases of modern people.[166][167][unreliable source?] Modern human genes involved in making keratin—the protein found in skin, hair, and nails—have specially high levels of Neanderthal
Neanderthal
DNA.[167] For example, around 66% of East Asians contain the Neanderthal
Neanderthal
skin gene, while 70% of Europeans possess the Neanderthal
Neanderthal
gene which affects skin colour. POU2F3 is found in around 66 percent of East Asians, while the Neanderthal
Neanderthal
version of BNC2, which affects skin color, among other traits, is found in 70 percent of Europeans. Neanderthal
Neanderthal
are the variants in genes that affect the risk of several diseases, including lupus, biliary cirrhosis, Crohn's disease, and type 2 diabetes. Eight percent of Neanderthal
Neanderthal
DNA
DNA
comes from an unknown group of archaic humans, tantalising hints of unknown groups from Asia and Africa that left genes in Denisovans
Denisovans
and modern humans, respectively.[167][166] The genetic variant of the MC1R gene linked to red hair in Neanderthals
Neanderthals
has not been found in modern humans; hence, red hair may be an example of convergent evolution.[168][169][104] While interbreeding is viewed as the most parsimonious interpretation of the genetic discoveries, the authors point out they cannot conclusively rule out an alternative scenario, in which the source population of non-African modern humans was already more closely related to Neanderthals
Neanderthals
than other Africans were, because of ancient genetic divisions within Africa.[89] Other studies carried out since the sequencing of the Neanderthal genome
Neanderthal genome
have cast doubt on the level of admixture between Neanderthals
Neanderthals
and modern humans, or even as to whether the groups interbred at all. One study has asserted that the presence of Neanderthal
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.[170][171][172][173] Among the genes shown to differ between present-day humans and Neanderthals
Neanderthals
were RPTN, SPAG17, CAN15, TTF1, FOXP2
FOXP2
and PCD16.[89] Specifically, a visualisation map of the reference modern-human containing the genome regions with high degree of similarity or with novelty according to a Neanderthal
Neanderthal
of 50k[58] has been built by Pratas et al.[174] More recent research suggests that Neanderthal– Homo
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
DNA
in the genomes of modern humans. In 2015, researchers Benjamin Vernot and Joshua Akey at the University of Washington conclude in a paper in the American Journal of Human
Human
Genetics
Genetics
that the relatively greater quantity of Neanderthal-specific DNA
DNA
in the genomes of individuals of East Asian descent (than those of European descent) cannot be explained by differences in selection.[175] They further suggest that "two additional demographic models, involving either a second pulse of Neandertal gene flow into the ancestors of East Asians or a dilution of Neandertal lineages in Europeans by admixture with an unknown ancestral population" are parsimonious with their data.[175] Similar conclusions were reached in a paper published in the same publication by researchers Bernard Kim and Kirk Lohmueller at UCLA: "Using simulations of a broad range of models of selection and demography, we have shown that this hypothesis [that the greater proportion of Neandertal ancestry in East Asians than in Europeans is due to the fact that purifying selection is less effective at removing weakly deleterious Neandertal alleles from East Asian populations] cannot account for the higher proportion of Neandertal ancestry in East Asians than in Europeans. Instead, more complex demographic scenarios, most likely involving multiple pulses of Neandertal admixture, are required to explain the data."[176] In a subsequent interview, Dr. Lohmueller did note that these findings go against the commonly-held perception that Neanderthals
Neanderthals
were mostly localised to modern-day Europe and western Asia: "It’s very hard to put these findings into spatial context. The key idea is that there would have to have been some additional interbreeding events involving East Asians, but not Europeans. These interbreeding events could have been directly between Neanderthals
Neanderthals
and East Asians, maybe in some other indirect way."[177][better source needed] Vernot also noted that "[H]umans have been constantly migrating throughout their history—this makes it hard to say exactly where interactions with Neanderthals
Neanderthals
occurred. It's possible, for example, that all of the interbreeding with Neanderthals
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
Neanderthals
just a little bit more."[177][better source needed] Studies published in March 2016 suggest that modern humans bred with hominins, including Neanderthals, on multiple occasions.[178] Another study in April 2016 found differences between modern human and Neanderthal
Neanderthal
Y chromosomes that, they postulated, could cause female Homo
Homo
sapiens sapiens to miscarry male babies that had Neanderthal fathers.[179] This could explain why no modern man had to date been found with a Neanderthal
Neanderthal
Y chromosome.[180] Melanesians and Australoid populations show evidence of only one interbreeding event, possibly about 100,000 years ago, occurring in the Middle East, Europeans show a second event, which may also be of Middle Eastern origin, occurring possibly 50,000 years ago, while East Asians show an additional third interbreeding event possibly 30,000 years ago occurring in Siberia. Evidence that Neanderthal
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
Neanderthal
populations had bred resistance to.[178] In 2016 researchers reported that they had found Human
Human
DNA
DNA
in the genome of a female Neanderthal
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.[181] Epigenetics[edit] In April 2014, a first glimpse into the epigenetics of the Neanderthal was obtained with the publication of the full DNA
DNA
methylation of the Neanderthal
Neanderthal
and the Denisovan.[178][182] The reconstructed DNA methylation map allowed researchers to assess gene activity levels throughout the Neanderthal genome
Neanderthal genome
and compare them to modern humans. One of the major findings focused on the limb morphology of Neanderthals. Gokhman et al. found that changes in the activity levels of the HOX cluster
HOX cluster
of genes were behind many of the morphological differences between Neanderthals
Neanderthals
and modern humans, including shorter limbs, curved bones and more.[182] Extinction[edit] Main article: Neanderthal
Neanderthal
extinction According to a 2014 study by Thomas Higham and colleagues of organic samples from European sites, Neanderthals
Neanderthals
died out in Europe between 41,000 and 39,000 years ago.[e] New dating in Iberia, where Neanderthal
Neanderthal
dates as late as 28,000 years had been reported, suggests evidence of Neanderthal
Neanderthal
survival in the peninsula after 42,000 years ago is almost non-existent.[11] Anatomically modern humans arrived in Mediterranean Europe between 45,000 and 43,000 years ago, so the two different human populations shared Europe for several thousand years.[8][183] The exact nature of biological and cultural interaction between Neanderthals
Neanderthals
and other human groups is contested.[184] Possible scenarios for the extinction of the Neanderthals
Neanderthals
are:

Neanderthals
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.[185] Jared Diamond
Jared Diamond
has suggested a scenario of violent conflict and displacement.[186] Neanderthals
Neanderthals
were a contemporary subspecies that bred with modern humans and disappeared through absorption (interbreeding theory). Volcanic catastrophe: see Campanian Ignimbrite Eruption

mtDNA-based simulation of modern human expansion in Europe starting 1,600 generations ago. Neanderthal
Neanderthal
range in light grey[187]

Climate change[edit] About 55,000 years ago, the climate began to fluctuate wildly from extreme cold conditions to mild cold and back in a matter of decades. Neanderthal
Neanderthal
bodies were well-suited for survival in a cold climate—their stocky chests and limbs stored body heat better than the Cro-Magnons. Neanderthals
Neanderthals
died out in Europe between 41,000 and 39,000 years ago, apparently coinciding with the start of a very cold period.[13][188] 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
Middle Palaeolithic
and Early Upper Palaeolithic more than 95% of stone artifacts were drawn from local material, suggesting Neanderthals
Neanderthals
restricted themselves to more local sources.[189] Coexistence with modern humans[edit]

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

In November 2011 tests conducted at the Oxford Radiocarbon
Radiocarbon
Accelerator Unit in England on what were previously thought to be Neanderthal
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.[190] Given that the 2014 study by Thomas Higham of Neanderthal bones and tools indicates that Neanderthals
Neanderthals
died out in Europe between 41,000 and 39,000 years ago, the two different human populations shared Europe for as long as 5,000 years.[13] Nonetheless, the exact nature of biological and cultural interaction between Neanderthals
Neanderthals
and other human groups has been contested.[184] Modern humans co-existed with them in Europe starting around 45,000 years ago and perhaps even earlier. Neanderthals
Neanderthals
inhabited that continent long before the arrival of modern humans. These modern humans may have introduced a disease that contributed to the extinction of Neanderthals, and that may be added to other recent explanations for their extinction. When Neanderthal
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
Neanderthals
occurred in Europe and Asia the first contact may have been devastating to the Neanderthal
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.[191] The most well-known example of this is the arrival of Christopher Columbus
Christopher Columbus
to the New World, which brought and introduced foreign diseases when he and his crew arrived to a native population who had no immunity. Anthropologist Pat Shipman, of Pennsylvania State University, suggested that domestication of the dog could have played a role in Neanderthals' extinction.[192] Interbreeding hypotheses[edit] Main article: Archaic human admixture with modern humans

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

An alternative to extinction is that Neanderthals
Neanderthals
were absorbed into the Cro-Magnon
Cro-Magnon
population by interbreeding. This would be counter to strict versions of the recent African origin theory, since it would imply that at least part of the genome of Europeans would descend from Neanderthals. Pre-2010 interbreeding hypotheses[edit] Until the early 1950s, most scholars believed Neanderthals
Neanderthals
were not in the ancestry of living humans.[194]:232–234[195] Nevertheless, Thomas H. Huxley in 1904 saw among Frisians the presence of what he believed to be Neanderthaloid skeletal and cranial characteristics as an evolutionary development from Neanderthal
Neanderthal
rather than as a result of interbreeding, saying that "the blond long-heads may exhibit one of the lines of evolution of the men of the Neanderthaloid type," yet he raised the possibility that the Frisians alternatively "may be the result of the admixture of the blond long-heads with Neanderthal
Neanderthal
men," thus separating "blond" from "Neanderthaloid."[196] Hans Peder Steensby proposed interbreeding in 1907 in the article Race studies in Denmark. He strongly emphasised that all living humans are of mixed origins.[197] He held that this would best fit observations, and challenged the widespread idea that Neanderthals
Neanderthals
were ape-like or inferior. Basing his argument primarily on cranial data, he noted that the Danes, like the Frisians and the Dutch, exhibit some Neanderthaloid characteristics, and felt it was reasonable to "assume something was inherited" and that Neanderthals
Neanderthals
"are among our ancestors." Carleton Stevens Coon in 1962 found it likely, based upon evidence from cranial data and material culture, that Neanderthal
Neanderthal
and Upper Paleolithic peoples either interbred or that the newcomers reworked Neanderthal
Neanderthal
implements "into their own kind of tools."[198] Christopher Thomas Cairney in 1989 went further, laying out a rationale for hybridisation and adding a broader discussion of physical characteristics as well as commentary on interbreeding and its importance to adaptive European phenotypes. Cairney specifically discussed the "intermixture of racial elements" and "hybridisation."[199] By the early 2000s, the majority of scholars supported the Out of Africa hypothesis,[200][201] according to which anatomically modern humans left Africa about 50,000 years ago and replaced Neanderthals with little or no interbreeding. Yet some scholars still argued for hybridisation with Neanderthals. The most vocal proponent of the hybridisation hypothesis was Erik Trinkaus of Washington University.[202] Trinkaus claimed various fossils as products of hybridised populations, including the child of Lagar Velho, a skeleton found at Lagar Velho in Portugal[203][204][205] and the Peștera Muierii in Romania.[206] Interbreeding hypotheses since 2010[edit] In 2010, geneticists announced that interbreeding had likely taken place,[89][207] a result confirmed in 2012.[90][91][208][page needed] The genomes of all non-Africans include portions that are of Neanderthal
Neanderthal
origin,[209][210] a share estimated in 2014 to 1.5–2.1%.[58] This DNA
DNA
is absent in Sub-Saharan Africans ( Yoruba people
Yoruba people
and San subjects).[89] Ötzi
Ötzi
the iceman, Europe's oldest preserved mummy, was found to possess an even higher percentage of Neanderthal
Neanderthal
ancestry.[211] The two percent of Neanderthal
Neanderthal
DNA
DNA
in Europeans and Asians is not the same in all Europeans and Asians: In all, approximately 20% of the Neanderthal genome appears to survive in the modern human gene pool.[212] 2012 genetic studies seem to suggest that modern humans may have mated with "at least two groups" of archaic humans: Neanderthals
Neanderthals
and Denisovans.[213] Some researchers suggest admixture of 3.4–7.9% in modern humans of non-African ancestry, rejecting the hypothesis of ancestral population structure.[214] Detractors have argued and continue to argue that the signal of Neanderthal
Neanderthal
interbreeding may be due to ancient African substructure, meaning that the similarity is only a remnant of a common ancestor of both Neanderthals
Neanderthals
and modern humans and not the result of interbreeding.[172][215] John D. Hawks has argued that the genetic similarity to Neanderthals
Neanderthals
may indeed be the result of both structure and interbreeding, as opposed to just one or the other.[216] While some modern human nuclear DNA
DNA
has been linked to the extinct Neanderthals, no mitochondrial DNA
DNA
of Neanderthal
Neanderthal
origin has been detected,[35] which in primates is always maternally transmitted. This observation has prompted the hypothesis that whereas female humans interbreeding with male Neanderthals
Neanderthals
were able to generate fertile offspring, the progeny of female Neanderthals
Neanderthals
who mated with male humans were either rare, absent or sterile.[217] Specimens[edit] Notable European Neanderthals[edit]

La Ferrassie
La Ferrassie
1, skull cast

Le Moustier
Le Moustier
1 in 1909

Le Moustier
Le Moustier
1 in 2011

Shanidar
Shanidar
1, skull cast

Kebara 2

Teshik-Tash
Teshik-Tash
1

Remains of more than 300 European Neanderthals
Neanderthals
have been found. For the most important, see List of human evolution fossils.

Neanderthal
Neanderthal
1: The first human bones recognised as showing a non-modern anatomy. Discovered in 1856 in a limestone quarry at the Feldhofer grotto in Neanderthal, Germany, they consist of a skull cap, the two femora, the three right arm bones, two left arm bones, the ilium, and fragments of a scapula and ribs. La Chapelle-aux-Saints
La Chapelle-aux-Saints
1: Called the Old Man, a fossilised skull discovered in La Chapelle-aux-Saints, France, by A. and J. Bouyssonie, and L. Bardon in 1908. Characteristics include a low vaulted cranium and large browridge typical of Neanderthals. Estimated to be about 60,000 years old, the specimen was severely arthritic and had lost all his teeth long before death, leading some to suggest he was cared for by others. La Ferrassie
La Ferrassie
1: A fossilised skull discovered in La Ferrassie, France, by R. Capitan in 1909. It is estimated to be 70,000 years old. Its characteristics include a large occipital bun, low-vaulted cranium and heavily worn teeth. Le Moustier
Le Moustier
1: One of the rare nearly complete Neanderthal
Neanderthal
skeletons to be discovered, it was excavated by a German team in 1908, at Peyzac-le-Moustier, France. Sold to a Berlin museum, the post cranial skeleton was bombed and mostly destroyed in 1945, and parts of the mid face were lost sometime after then. The skull, estimated to be less than 45,000 years old, includes a large nasal cavity and a less developed brow ridge and occipital bun than seen in other Neanderthals. The Mousterian
Mousterian
tool techno-complex is named after its discovery site.

Notable Southwest Asian Neanderthals[edit] Remains of more than 70 Southwest Asian Neanderthals
Southwest Asian Neanderthals
have been found. For a complete list see List of Southwest Asian Neanderthals.

Shanidar
Shanidar
1 to 10: Eight Neanderthals
Neanderthals
and two pre-Neanderthals ( Shanidar
Shanidar
2 and 4) were discovered in the Zagros Mountains
Zagros Mountains
in Iraqi Kurdistan. One of the skeletons, Shanidar
Shanidar
4, was once thought to have been buried with flowers, a theory no longer accepted. To Paul B. Pettitt the "deliberate placement of flowers has now been convincingly eliminated", since "[a] recent examination of the microfauna from the strata into which the grave was cut suggests that the pollen was deposited by the burrowing rodent Meriones tersicus, which is common in the Shanidar
Shanidar
microfauna and whose burrowing activity can be observed today".[218] Amud 1: A male adult Neanderthal, dated to roughly 55,000 BP, and one of several found in a cave at Nahal Amud, Israel. At 178 cm (70 in), it is the tallest known Neanderthal. It also has the largest cranial capacity of all extinct hominins: 1,736 cm3.[71][219] Kebara 2: A male adult post-cranial skeleton, dated to roughly 60,000 BP, that was discovered in 1983 in Kebara Cave, Israel. It has been studied extensively, for its hyoid, ribcage, and pelvis are much better preserved than in all other Neanderthal
Neanderthal
specimens.

Notable Central Asian Neanderthal[edit]

Teshik-Tash
Teshik-Tash
1: An 8–11 year old skeleton discovered in Uzbekistan
Uzbekistan
by Okladnikov in 1938. This is the only fairly complete skeleton discovered to the east of Iraq. Okladnikov claimed it was a deliberate burial, but this is debated.

Chronology[edit] This section describes bones with Neanderthal
Neanderthal
traits in chronological order. Mixed with H. heidelbergensis
H. heidelbergensis
traits[edit]

> 350 ka: Sima de los Huesos c. 500:350 ka ago[67][220] 350–200 ka: Pontnewydd 225 ka ago. 200–135 ka: Atapuerca,[221] Vértesszőlős, Ehringsdorf, Casal de'Pazzi, Biache, La Chaise, Montmaurin, Prince, Lazaret, Fontéchevade

H. neanderthalensis fossils[edit]

130–50 ka: Krapina, Saccopastore skulls, Malarnaud, Altamura, Gánovce, Denisova, Okladnikov, Pech de l'Azé, Tabun 120–7002100000000000000♠100±5 ka,[222] Shanidar
Shanidar
1 to 9 80–60 ka, La Ferrassie 1
La Ferrassie 1
70 ka, Kebara 60 ka, Régourdou, Mt. Circeo, Combe Grenal, Erd 50 ka, La Chapelle-aux Saints 1 60 ka, Amud I 7001530000000000000♠53±8 ka,[223][224] Teshik-Tash. In radiocarbon range, > 50 ka: Le Moustier, Feldhofer, La Quina, l'Hortus, Kulna, Šipka, Saint Césaire, Bacho Kiro, El Castillo, Bañolas, El Sidrón (48±3 cal ka),[225] Arcy-sur-Cure, Châtelperron, Figueira Brava, Mezmaiskaya (41±1 cal ka),[10] Zafarraya, Vindija, Velika Pećina.

H. s. sapiens with traits reminiscent of Neanderthals[edit]

< 35 Pestera cu Oase
Pestera cu Oase
35 ka, Mladeč
Mladeč
31 ka, Pestera Muierii 30 ka (n/s),[226] Lapedo Child 24.5 ka.

In popular culture[edit] Main article: Neanderthals
Neanderthals
in popular culture Neanderthals
Neanderthals
have been portrayed in popular culture including appearances in literature, visual media and comedy. Early 20th century artistic interpretations often presented Neanderthals
Neanderthals
as beastly creatures, emphasising hairiness and rough, dark complexion.[227] See also[edit]

Homo
Homo
heidelbergensis Denisova hominin Early human migrations Human
Human
evolution Human
Human
timeline Life timeline List of Neanderthal
Neanderthal
sites Neanderthals
Neanderthals
in Gibraltar Prehistoric Autopsy
Prehistoric Autopsy
(2012 BBC documentary) Dawn of Humanity (2015 PBS film)

Notes[edit]

^ The common species name Neanderthal
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
Neanderthal
comes from the binomial scientific name[5] established by King in 1864, Homo neanderthalensis.[6] The binomial name is indeed taken from German but because binomial names are normally unalterable, the binomial still reflects the pre-1901 German spelling and hence so does, for most authors, the common name. The Neandertal region in Germany
Germany
is in English written without an h. Note that in German the common species name is almost always Neandertaler (lit. "of the valley of Neander") not Neandertal, but in the few instances where the word Neandertal is used to refer not to the place but to the prehistoric humans, as is the case of the Neanderthal
Neanderthal
Museum, the h is kept for the same reason as in English that it reflects the scientific name.[7] ^ The valley is named after Joachim Neander, whose Greek-style last name had been changed by his grandfather from "Neumann" ("new man").[35] ^ Some words beginning with th in older varieties of German were the result of a spelling embellishment that had no connection to English th. (Teil, meaning 'part,' was sometimes spelled Theil in the 18th and 19th centuries.) Tal became standardized with the German spelling reform of 1901, thus the German name Neandertal for both the valley and species/subspecies. ^ There are modern humans with noses as wide as those of Neanderthals and modern humans with similar nose lengths, but none with both Neanderthal
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.[8] ^ Homo
Homo
floresiensis originated in an unknown location from unknown ancestors and reached remote parts of Indonesia. Homo
Homo
erectus spread from Africa to western Asia, then east Asia and Indonesia; its presence in Europe is uncertain, but it gave rise to Homo
Homo
antecessor, found in Spain. Homo
Homo
heidelbergensis originated from Homo
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
Homo
sapiens sapiens spread from Africa to western Asia and then to Europe and southern Asia, eventually reaching Australia and the Americas. In addition to Neanderthals
Neanderthals
and Denisovans, a third gene flow of archaic Africa origin is indicated at the right.[193]

References[edit]

^ Romeo, Luigi (1979). Ecce Homo!:A Lexicon of Man. Amsterdam: John Benjamins Publishing Company. p. 92. ISBN 9027220069.  ^ Camp, C. L; Allison, H. J.; Nichols, R. H. (1964). Bibliography of Fossil Vertebrates 1954–1958. New York: The Geological Society of America, Inc. p. 556. ISBN 9780813710921.  ^ " Neanderthal
Neanderthal
in ODE". Oxford Dictionaries.  ^ ""Neanderthal" in Random House Dictionary (US) & Collins Dictionary (UK)". Dictionary.com.  ^ "Neanderthal". Wiley-Blackwell Encyclopedia of Human
Human
Evolution. 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?". mettmann.de. Neanderthal
Neanderthal
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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): 252–55. Bibcode:1993Natur.363..252M. doi:10.1038/363252a0. ISSN 0028-0836. PMID 8387643.  ^ Rink, W. Jack; Schwarcz, H.P.; Lee, H.K.; Rees-Jones, J.; Rabinovich, R.; Hovers, E. (August 2002). "Electron spin resonance (ESR) and thermal ionization mass spectrometric (TIMS) 230Th/234U dating of teeth in Middle Paleolithic layers at Amud Cave, Israel". Geoarchaeology. 16 (6): 701–17. doi:10.1002/gea.1017.  ^ Valladas, Hélène; Merciera, N.; Frogeta, L.; Hoversb, E.; Joronc, J.L.; Kimbeld, W.H.; Rak, Y. (March 1999). "TL Dates for the Neanderthal
Neanderthal
Site of the Amud Cave, Israel". Journal of Archaeological Science. 26 (3): 259–68. 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 Neanderthals
Neanderthals
from El Sidrón Cave (Asturias, Northern Spain)". Archaeometry. 55 (1): 148–58. doi:10.1111/j.1475-4754.2012.00671.x. CS1 maint: Uses authors parameter (link) ^ Hayes, Jacqui (November 2, 2006). "Humans and Neanderthals interbred". Cosmos. Archived from the original on February 22, 2009. Retrieved May 17, 2009.  ^ Neanderthal
Neanderthal
image by Kupka, based on Boule, 1909, in Humanity's Journeys Dr. Kathryn Denning, 2005. Retrieved March 17, 2012.

Journals

Boë, Louis-Jean; Heim, Jean-Louis; Honda, Kiyoshi; Maeda, Shinji (2002). "The potential Neandertal vowel space was as large as that of modern humans". Journal of Phonetics. 30 (3): 465–84. doi:10.1006/jpho.2002.0170.  Lieberman, Philip (October 2007). "Current views on Neanderthal
Neanderthal
speech capabilities: A reply to Boe et al. (2002)". Journal of Phonetics. 35 (4): 552–63. doi:10.1016/j.wocn.2005.07.002.  Serre, David; Langaney, André; Chech, Mario; Teschler-Nicola, Maria; Paunovic, Maja; Mennecier, Philippe; Hofreiter, Michael; Possnert, Göran; Pääbo, Svante (2004). "No Evidence of Neandertal mtDNA Contribution to Early Modern Humans". PLoS Biology. 2 (3): e57. doi:10.1371/journal.pbio.0020057. PMC 368159 . PMID 15024415.  Wild, Eva M.; Teschler-Nicola, Maria; Kutschera, Walter; Steier, Peter; Trinkaus, Erik; Wanek, Wolfgang (2005). "Direct dating of Early Upper Palaeolithic human remains from Mladeč". Nature. 435 (7040): 332–35. Bibcode:2005Natur.435..332W. doi:10.1038/nature03585. PMID 15902255.  Zilhão, João; Davis, Simon J. M.; Duarte, Cidália; Soares, António M. M.; Steier, Peter; Wild, Eva (2010). Hawks, John, ed. "Pego do Diabo (Loures, Portugal): Dating the Emergence of Anatomical Modernity in Westernmost Eurasia". PLOS One. 5 (1): e8880. Bibcode:2010PLoSO...5.8880Z. doi:10.1371/journal.pone.0008880. PMC 2811729 . PMID 20111705. Lay summary – ScienceDaily (January 27, 2010). 

Bibliography

Derev'anko, Anatoliy P.; Powers, William Roger; Shimkin, Demitri Boris (1998). The Paleolithic of Siberia: new discoveries and interpretations. Novosibirsk: Institute of Anthropology and Ethnography. ISBN 978-0-252-02052-0. OCLC 36461622.  Lunine, Jonathan I. (2013). Earth: Evolution of a Habitable World. Cambridge University Press. 327. ISBN 978-0-521-85001-8. 

Further reading[edit]

Sankararaman, Sriram; Mallick, Swapan; Dannemann, Michael; Prüfer, Kay; Kelso, Janet; Patterson, Nick; Reich, David (2014). "The genomic landscape of Neanderthal
Neanderthal
ancestry in present-day humans". Nature. 507 (7492): 354–57. Bibcode:2014Natur.507..354S. doi:10.1038/nature12961. PMC 4072735 . PMID 24476815.  Vattathil, S.; Akey, J.M. (2015). "Small amounts of archaic admixture provide big insights into human history". Cell. 163 (2): 281–84. doi:10.1016/j.cell.2015.09.042. PMID 26451479. 

External links[edit]

Wikimedia Commons has media related to Homo
Homo
neanderthalensis.

Wikibooks has a book on the topic of: Introduction to Paleoanthropology

Wikispecies
Wikispecies
has information related to Homo
Homo
neanderthalensis

Wikisource
Wikisource
has the text of the 1905 New International Encyclopedia article Neanderthal
Neanderthal
Man.

Kreger, C. David (June 30, 2000). " Homo
Homo
neanderthalensis". ArchaeologyInfo.com. Retrieved May 23, 2009.  O'Neil, Dennis (May 12, 2009). "Evolution of Modern Humans: Neandertals". Retrieved May 23, 2009.  New Portuguese skull may be an early relative of Neandertals – article by Ann Gibbons at Science, March 13, 2017. In Neanderthal
Neanderthal
DNA, Signs of a Mysterious Human
Human
Migration – article by Carl Zimmer, NY Times, July 4, 2017 " Homo
Homo
neanderthalensis". The Smithsonian Institution.  " Neanderthal
Neanderthal
DNA". International Society of Genetic Genealogy. Archived from the original on June 17, 2006. : Includes Neanderthal
Neanderthal
mt DNA
DNA
sequences Panoramio – The Neandertal foot prints' (photo of ≈25K years old fossilised footprints discovered in 1970 on volcanic layers near Demirkopru Dam Reservoir, Manisa, Turkey) Did better mothering defeat the Neanderthals? My Great-great-great Grandfather's a Neanderthal Ancient tryst fortified human immune system Neanderthal-human hybridisation hypothesis Neanderthal
Neanderthal
hybridisation and Haldane's rule Neanderthal
Neanderthal
Studies Professional Online Service (NESPOS) fossil overview Human
Human
Timeline (Interactive) – Smithsonian, National Museum of Natural History (August 2016).

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