_ Homo mousteriensis_ _Palaeoanthropus neanderthalensis_
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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 went 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 species, _ 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. Male 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 in) in 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.
The Neanderthal genome project published papers in 2010 and 2014 stating that Neanderthals contributed to the DNA of modern humans , including most humans outside sub-Saharan Africa , as well as a few populations in sub-Saharan Africa, through interbreeding , likely between 50,000 and 60,000 years ago. Recent studies also show that some Neanderthals mated with ancestors of modern humans long before the "out of Africa migration" of present-day non-Africans, as early as 100,000 years ago. In 2016, research indicated that there were three distinct episodes of interbreeding between modern humans and Neanderthals: the first encounter involved the ancestors of non-African modern humans, probably soon after leaving Africa; the second, after the ancestral Melanesian group had branched off (and subsequently had a unique episode of interbreeding with Denisovans ); and the third, involving the ancestors of East Asians only.
Geneticists first sequenced the entire genome of a Neanderthal in 2013 by extracting it from the phalanx bone of a 50,000-year-old Siberian Neanderthal.
* 1 Name * 2 Classification * 3 Origins
* 4 Discovery
* 4.1 Timeline of research
* 5 Habitat and range * 6 Anatomy
* 7 Behavior
* 7.1 Claims of art and adornment
* 8 Genome
* 8.1 Background * 8.2 Interbreeding with modern humans * 8.3 Epigenetics
* 9 Extinction
* 9.1 Climate change * 9.2 Coexistence with modern humans
* 10 Interbreeding hypotheses
* 11 Specimens
* 11.4 Chronology
* 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 of 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 that the 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 . 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 _ Homo sapiens neanderthalensis_, placing Neanderthals as a subspecies of _H. sapiens_. DNA researcher Svante Pääbo referred to the ongoing "taxonomic wars" over whether Neanderthals were a separate species or subspecies as the type of debate that cannot be resolved, "since there is no definition of species perfectly describing the case."
During the early 20th century the prevailing view was heavily influenced by Arthur Keith and Marcellin Boule , who wrote the first scientific description of a nearly complete Neanderthal skeleton. 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.
Both Neanderthals and living humans are thought to have evolved from _ Homo erectus _. In the earliest known migration wave into Eurasia dated to 1.81 million years ago (Ma ), _ Homo erectus_ left Africa most probably via the Levant and reached Georgia (fossils of Dmanisi ). Hominins had reached China by 1.7 Ma and Iberia (Spain) by 1.4 Ma. The discoverers of fragmented bones in Spain (Iberia) dated to 1.2 million years, assigned to a new species _ Homo antecessor _, argue these are the remains of the ancestors of Neanderthals and of the older species _ Homo heidelbergensis _, an interpretation rejected by most anthropologists.
A large number of molecular clock genetic studies place the divergence time of the Neanderthal and modern human lineages between 800,000 and 400,000 years ago. For this reason, most scholars believe Neanderthals descend, via _ Homo heidelbergensis _, from another _ Homo erectus _ migration out of Africa that would have occurred in this time frame. Parts of the _ Homo erectus_ population that stayed in Africa would have evolved, perhaps through the intermediate _ Homo rhodesiensis _, into early anatomically modern humans by 200,000 years ago or earlier.
Neanderthal traits are present in _ Homo heidelbergensis _ specimens beginning between 600,000 and 350,000 years ago. 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 Neanderthals.
Neanderthal fossils were first discovered in 1829 in the Engis caves (the partial skull dubbed Engis 2 ), in what is now Belgium by Philippe-Charles Schmerling and the Gibraltar 1 skull in 1848 in the Forbes\' Quarry , Gibraltar , both prior to the type specimen discovery in a limestone quarry in the Neandertal in Germany in August 1856, three years before Charles Darwin 's _On the Origin of Species _ was published.
The type specimen , dubbed Neanderthal 1 , consisted of a skull cap, two femora , three bones of the right arm, two of the left arm, parts of the left ilium , fragments of a scapula , and ribs. The workers who recovered the objects originally thought them to be the remains of a cave bear. However, they eventually gave the material to amateur naturalist Johann Carl Fuhlrott , who turned the fossils over to anatomist Hermann Schaaffhausen .
To date, the bones of over 400 Neanderthals have been found.
TIMELINE OF RESEARCH
* 1829: A damaged skull of a Neanderthal child, Engis 2 , is discovered in Engis , Netherlands (now Belgium). * 1848: A female Neanderthal skull, Gibraltar 1 , is found in Forbes\' Quarry , Gibraltar, but its importance is not recognised. * 1856: Limestone miners discover the Neanderthal type specimen, Neanderthal 1 , in Neandertal , western Germany. * 1864: William King is the first to recognise Neanderthal 1 as belonging to a separate species, for which he gives the scientific name Homo neanderthalensis_. He then changed his mind on placing it in the _Homo_ genus, arguing that the upper skull was different enough to warrant a separate genus since, to him, it had likely been "incapable of moral and theistic conceptions." * 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 * 1912: Marcellin Boule publishes his now discredited influential study of Neanderthal skeletal morphology based on La Chapelle-aux-Saints 1 . * 1953–1957: Ten Neanderthal skeletons are excavated in Shanidar Cave, Iraqi Kurdistan , by Ralph Solecki and colleagues. * 1975: Erik Trinkaus 's study of Neanderthal feet strongly argues that Neanderthals walked like modern humans. * 1981: The site of Bontnewydd , Wales yielded an early Neanderthal tooth, the most north-western Neanderthal remain ever. * 1987: Israeli Neanderthal Kebara 2 is dated (by TL and ESR ) to 60,000 BP, thus later than the Israeli anatomically modern humans dated to 90,000 and 80,000 BP at Qafzeh and Skhul . * 1994: The site of Sidrón Cave , Spain, is discovered where the remains of 12 males and females were to be found. Mitochondrial DNA studies would show that the adult males were genetically related, but the adult females were not, suggesting female exogamy . * 1997: Matthias Krings _et al._ are the first to amplify Neanderthal mitochondrial DNA (mt 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 , 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 to sequence the Neanderthal nuclear genome. * 2010: 1–4% of the DNA of living non-African humans are found by the Max Planck Institute to likely come from Neanderthals, a result confirmed in 2012, and refined to 1.5–2.1% in 2014. * 2013: In the midst of a hundred-year old debate over the existence of Neanderthal deliberate burials, a team of French researchers reasserted a heavily contested 1908 claim that the Chapelle-aux-Saints 1 skeleton was deliberately buried. * 2014: Some researchers express worry that almost all theories for the Neanderthals' extinction assume that there is something modern humans 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 that European Neanderthals died out between 41,000 and 39,000 years ago, which coincides with the start of a very cold period in Europe and is no more than 5,000 years after anatomically-modern _Homo sapiens_ reached the continent.
HABITAT AND RANGE
Early Neanderthals, living before the Eemian interglacial (130 ka), are poorly known and come mostly from European sites. From 130 ka onwards, the quality of the fossil record increases dramatically. From then on, Neanderthal remains are found in Western, Central, Eastern, and Mediterranean Europe, as well as Southwest , Central, and Northern Asia up to the Altai Mountains in Siberia. No Neanderthal has ever been found outside Western Eurasia, namely neither to the south of Jerusalem (Shuqba ), nor further east than Kazakhstan (Denisova , Russia), nor to the north of Wales (Bontnewydd ), although it is difficult to assess the limits of their northern range because glacial advances destroy most human remains, the Bontnewydd tooth being exceptional. Middle Palaeolithic artifacts have been found up to 60° N on the Russian plains.
There likely never were more than 70,000 Neanderthals at any given time.
Main article: Neanderthal anatomy
Neanderthal anatomy differed from modern humans in that they had a more robust build and distinctive morphological features, especially on the cranium , which gradually accumulated more derived aspects as it was described by Marcellin Boule , 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, Neanderthal males averaged 164 to 168 cm (65 to 66 in) and females 152 to 156 cm (60 to 61 in) tall. Samples of 26 specimens in 2010 found an average weight of 77.6 kg (171 lb) for males and 66.4 kg (146 lb) for females. A 2007 genetic study suggested some 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 Neanderthals do."
In _The Spread of Modern Humans in Europe_ (2002) John F Hoffecker, writes " Neanderthal sites show no evidence of tools for making tailored clothing. There are only hide scrapers, which might have been used to make blankets or ponchos. This is in contrast to Upper Paleolithic (modern human) sites, which have an abundance of eyed bone needles and bone awls. Moreover, microwear analysis of Neanderthal hide scrapers shows that they were used only for the initial phases of hide preparation, and not for the more advanced phases of clothing production.
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 indicated that Neanderthal and modern human brains were the same size at birth, but that by adulthood, the Neanderthal brain was larger than the modern human brain. They had almost the same degree of encephalization (i.e. brain to body size ratio) as modern humans.
The Neanderthal skeleton suggests they consumed 100 to 350 kcal (420 to 1,460 kJ) more per day than male modern humans of 68.5 kg (151 lb) and females of 59.2 kg (131 lb).
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 studies suggest 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 Neanderthal coprolites (fossilized feces) found substantial amounts of plant matter. Dental analysis of specimens from Spy , Belgium and El Sidrón , Spain in 2017 argued these Neanderthals had a wide-ranging diet, and that those "from El Sidrón showed no evidence of meat eating" at all and seemed to have lived on "a mixture of forest moss, pine nuts and a mushroom known as split gill".
The size and distribution of Neanderthal sites, along with genetic evidence, suggests Neanderthals lived in much smaller and more sparsely distributed groups than anatomically-modern _ Homo sapiens_. The bones of twelve Neanderthals were discovered at El Sidrón cave in the Atapuerca Mountains of Spain. They are believed to have been a group killed and butchered about 50,000 years ago. Analysis of the mt 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.
Claims that 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 Neanderthals, Paul Mellars wrote that “the single most impressive and hitherto widely cited pillar of evidence for the presence of complex ‘symbolic’ behavior among the late Neanderthal populations in Europe has now effectively collapsed”. 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 Neanderthal art, adornment, and structures. These are often taken by the media as showing Neanderthals were capable of symbolic thought, or were “mental equals” to anatomically modern humans. As evidence of symbolism, none of them are widely accepted, although the same is true for Middle Palaeolithic anatomically modern humans. Among many others:
* Flower pollen on the body of pre- Neanderthal 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 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.
Further information: Neanderthal genome project
Early investigations concentrated on mitochondrial DNA (mtDNA), which, owing to strictly matrilineal inheritance and subsequent vulnerability to genetic drift , is of limited value in evaluating the possibility of interbreeding of Neanderthals with Cro-Magnon people.
In 1997, geneticists were able to extract a short sequence of DNA from 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 and 454 Life Sciences announced that they would sequence the Neanderthal genome over the next two years. This genome was expected to be roughly the size of the human genome , three-billion base pairs, and share most of its genes . It was hoped the comparison would expand understanding of Neanderthals, as well as the evolution of humans and human brains.
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.
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 Energy's Lawrence Berkeley National Laboratory and the Joint Genome Institute (JGI), sequenced a fraction (0.00002) of genomic nuclear DNA (nDNA) from a 38,000-year-old Vindia Neanderthal femur. They calculated the common ancestor to be about 353,000 years ago, and a complete separation of the ancestors of the groups about 188,000 years ago.
Their results show the genomes of modern humans and Neanderthals are at least 99.5% identical, but despite this genetic similarity, and despite the two groups having coexisted in the same geographic region for thousands of years, Rubin and his team did not find any evidence of any significant interbreeding between the two. Rubin said, "While unable to definitively conclude that interbreeding between the two species of humans did not occur, analysis of the nuclear DNA from the Neanderthal suggests the low likelihood of it having occurred at any appreciable level."
In 2008 Richard E. Green et al. from Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany, published the full sequence of Neanderthal mitochondrial DNA (mtDNA) and suggested " Neanderthals had a long-term effective population size smaller than that of modern humans."
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 Neanderthal DNA can be identified. Geneticist at the Max Planck Institute for Evolutionary Anthropology extracting ancient DNA
With 3 billion nucleotides sequenced, analysis of about ⅓ showed no sign of admixture between modern humans and Neanderthals, according to Pääbo. This concurred with the work of Noonan from two years earlier. The variant of microcephalin common outside Africa, which was suggested to be of Neanderthal origin and responsible for rapid brain growth in humans, was not found in Neanderthals. Nor was the MAPT variant, a very old variant found primarily in Europeans.
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 Neanderthal DNA in us," said Pääbo, who led the study. "The proportion of Neanderthal-inherited genetic material is about 1 to 4 percent. It is a small but very real proportion of ancestry in non-Africans today," says Dr. David Reich of Harvard Medical School , who worked on the study. This research compared the genome of the Neanderthals to five modern humans from China, France, sub-Saharan Africa, and Papua New Guinea. The finding is that about 1 to 4 percent of the genes of the humans outside sub-Saharan Africa came from Neanderthals, compared to the baseline defined by the two sub-Saharan Africans.
This indicates a gene flow from Neanderthals to modern humans, i.e., interbreeding between the two populations. Since the three non-African genomes show a similar proportion of Neanderthal sequences, the interbreeding must have occurred early in the migration of modern humans out of Africa , perhaps in the Middle East. No evidence for gene flow in the direction from modern humans to Neanderthals was found. Gene flow from modern humans to Neanderthals would not be expected if contact occurred between a small colonizing population of modern humans and a much larger resident population of Neanderthals. A very limited amount of interbreeding could explain the findings, if it occurred early enough in the colonization process.
It is suggested that 20 percent of Neanderthal DNA survived in modern humans, notably expressed in the skin, hair, and diseases of the modern people. 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 Neanderthal DNA comes from an unknown group of archaic humans, tantalizing hints of unknown groups from Asia and Africa that left genes in Denisovans and modern humans, respectively. 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 related to Neanderthals than other Africans were, because of ancient genetic divisions within Africa. Other studies carried out since the sequencing of the Neanderthal genome have cast doubt on the level of admixture between Neanderthals and modern humans, or even as to whether the groups interbred at all. One study has asserted that the presence of Neanderthal or other archaic human genetic markers can be attributed to shared ancestral traits between the lineages originating from a 500,000-year-old common ancestor.
More recent research suggests that Neanderthal–_ Homo sapiens sapiens_ interbreeding appears to have occurred asymmetrically among the ancestors of modern-day humans, and that this is a possible rationale for differing frequencies of Neanderthal-specific DNA in the genomes of modern humans. In 2015, researchers Benjamin Vernot and Joshua Akey at the University of Washington conclude in a paper in the American Journal of Human Genetics that the relatively greater quantity of Neanderthal-specific DNA in the genomes of individuals of East Asian descent (than those of European descent) cannot be explained by differences in selection. They further suggest that "two additional demographic models, involving either a second pulse of Neandertal gene flow into the ancestors of East Asians or a dilution of Neandertal lineages in Europeans by admixture with an unknown ancestral population" are parsimonious with their data. Similar conclusions were reached in a paper published in the same publication by researchers Bernard Kim and Kirk Lohmueller at UCLA: "Using simulations of a broad range of models of selection and demography, we have shown that this hypothesis cannot account for the higher proportion of Neandertal ancestry in East Asians than in Europeans. Instead, more complex demographic scenarios, most likely involving multiple pulses of Neandertal admixture, are required to explain the data."
In a subsequent interview, Dr. Lohmueller did note that these findings go against the commonly-held perception that Neanderthals were mostly 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 Neanderthals occurred. It's possible, for example, that all of the interbreeding with Neanderthals occurred in the Middle East, before the ancestors of modern non-Africans spread out across Eurasia. In the model from the paper, the ancestors of all non-Africans interbred with Neanderthals, and then split up into multiple groups that would later become Europeans, East Asians. Shortly after they split up, the ancestors of East Asians interbred with Neanderthals just a little bit more."
Studies published in March 2016 suggest that modern humans bred with hominins, including Neanderthals, on multiple occasions. Another study in April 2016 found differences between modern human and Neanderthal Y chromosomes that, they postulated, could cause female _ Homo sapiens sapiens_ to miscarry male babies that had Neanderthal fathers. This could explain why no modern man had to date been found with a Neanderthal Y chromosome. Melanesians and Australoid populations show evidence of only one interbreeding event, possibly about 100,000 years ago, occurring in the Middle East, Europeans show a second event, which may also be of Middle Eastern origin, occurring possibly 50,000 years ago, while East Asians show an additional third interbreeding event possibly 30,000 years ago occurring in Siberia. Evidence that Neanderthal genomic material is often found amongst genes of the immune system suggests that some of the interbreeding may have secured resistance to diseases that Neanderthal populations had bred resistance to.
In 2016 researchers reported that they had found Human DNA in the genome of a female Neanderthal from the Altai mountains region near the border between Mongolia and Russia. They calculated that the mating must have taken place about 100,000 years ago.
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 throughout the Neanderthal genome and compare them to modern humans. One of the major findings focused on the limb morphology of Neanderthals. Gokhman et al. found that changes in the activity levels of the HOX cluster of genes were behind many of the morphological differences between Neanderthals and modern humans, including shorter limbs, curved bones and more.
Main article: Neanderthal extinction
According to a 2014 study by Thomas Higham and colleagues of organic samples from European sites, Neanderthals died out in Europe between 41,000 and 39,000 years ago. New dating in Iberia, where Neanderthal dates as late as 28,000 years had been reported, suggests evidence of Neanderthal survival in the peninsula after 42,000 years ago is almost non-existent.
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 are:
* Neanderthals were a separate species from modern humans, and became extinct (because of climate change or interaction with modern humans) and were replaced by modern humans moving into their habitat between 45,000 and 40,000 years ago. 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 the Cro-Magnons. Neanderthals died out in Europe between 41,000 and 39,000 years ago, apparently coinciding with the start of a very cold period. Raw material sourcing and the examination of faunal remains by Adler et al. (2006) in the southern Caucasus suggest that modern humans may have had a survival advantage during this period, being able to use social networks to acquire resources from a greater area. They found that in both the Late Middle Palaeolithic and Early Upper Palaeolithic more than 95% of stone artifacts were drawn from local material, suggesting Neanderthals restricted themselves to more local sources.
COEXISTENCE WITH MODERN HUMANS
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 Higham of Neanderthal bones and tools indicates that Neanderthals died out in Europe between 41,000 and 39,000 years ago, the two different human populations shared Europe for as long as 5,000 years. Nonetheless, the exact nature of biological and cultural interaction between 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 arrival of Christopher Columbus to the New World, which brought and introduced foreign diseases when he and his crew arrived to a native population who had no immunity.
Anthropologist Pat Shipman, of Pennsylvania State University, suggested that domestication of the dog could have played a role in Neanderthals' extinction.
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.
An alternative to extinction is that Neanderthals were absorbed into the Cro-Magnon population by interbreeding . This would be counter to strict versions of the Recent African Origin , since it would imply that at least part of the genome of Europeans would descend from Neanderthals.
Hans Peder Steensby, while strongly emphasising that all modern humans are of mixed origins, proposed the interbreeding hypothesis in 1907, in the article _Race studies in Denmark_. He held that this would best fit current observations, and attacked the widespread idea that Neanderthals were ape-like or inferior.
The most vocal proponent of the hybridization hypothesis is Erik Trinkaus of Washington University . 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 cave of 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 ancient Neanderthal 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 : Neanderthals and 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 DNA of 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 female Neanderthals and male modern humans.
NOTABLE EUROPEAN NEANDERTHALS
* Neanderthal 1 : The first Neanderthal specimen found during mining in August 1856. It was discovered in a limestone quarry at the Feldhofer grotto in Neanderthal, Germany. The find consisted of a skull cap, two femora, three right arm bones, two left arm bones, ilium, and fragments of a scapula and ribs. * La Chapelle-aux-Saints 1 : Called the Old Man, a fossilized skull discovered in La Chapelle-aux-Saints , France, by A. and J. Bouyssonie, and L. Bardon in 1908. Characteristics include a low vaulted cranium and large browridge typical of Neanderthals. Estimated to be about 60,000 years old, the specimen was severely arthritic and had lost all his teeth long before death, leading some to suggest he was cared for by others. * La Ferrassie 1 : A fossilized skull discovered in La Ferrassie, France, by R. Capitan in 1909. It is estimated to be 70,000 years old. Its characteristics include a large occipital bun, low-vaulted cranium and heavily worn teeth. * Le Moustier 1: One of the rare nearly complete Neanderthal skeletons to be discovered, it was excavated by a German team in 1908, at Peyzac-le-Moustier, France. Sold to a Berlin museum, the post cranial skeleton was bombed and mostly destroyed in 1945, and parts of the mid face were lost sometime after then. The skull, estimated to be less than 45,000 years old, includes a large nasal cavity and a less developed brow ridge and occipital bun than seen in other Neanderthals. The Mousterian tool techno-complex is named after its discovery site.
NOTABLE SOUTHWEST ASIAN NEANDERTHALS
* 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 in the Shanidar microfauna and whose burrowing activity can be observed today". * 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 Neanderthal specimens.
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 order.
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, Fontéchevade
_H. Neanderthalensis_ Fossils
* 130–50 ka: Krapina , Saccopastore skulls , Malarnaud, Altamura , Gánovce, Denisova , Okladnikov , Pech de l'Azé, Tabun 120–7002100000000000000♠100±5 ka, Shanidar 1 to 9 80–60 ka, La Ferrassie 1 70 ka, Kebara 60 ka, Régourdou, Mt. Circeo, Combe Grenal , Erd 50 ka, La Chapelle-aux Saints 1 60 ka, Amud I 7001530000000000000♠53±8 ka, Teshik-Tash . * 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 , Velika Pećina.
_ Homo Sapiens Sapiens_ With Archaic Traits Reminiscent Of Neanderthals
Main article: Neanderthals in popular culture
Neanderthals have been portrayed in popular culture including appearances in literature, visual media and comedy. Early 20th century artistic interpretations often presented Neanderthals as beastly creatures, emphasizing hairiness and rough, dark complexion.
* _Dawn of Humanity_ (2015 PBS film) * Denisova hominin * Early human migrations * _ Homo heidelbergensis _ * Human evolution * Human timeline * Life timeline * List of Neanderthal sites * Neanderthal behaviour * Neanderthals in popular culture * Neanderthals of Gibraltar * _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 _Neandertal_ is used to refer not to the place but to the prehistoric humans, as is the case of the Neanderthal Museum , the _h_ is kept for the same reason as in English that it reflects the scientific name. * ^ The valley is named after Joachim Neander , whose Greek-style last name had been changed by his grandfather from "Neumann" ("new man"). * ^ 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 with both Neanderthal nose width and nose length. * ^ Higham et al did not study samples from sites outside Europe and they stated that further work was required to rule out later survival at Gorhams Cave, Gibraltar. * ^ _ Homo floresiensis _ originated in an unknown location from unknown ancestors and reached remote parts of Indonesia . _Homo erectus _ spread from Africa to western Asia, then east Asia and Indonesia; its presence in Europe is uncertain, but it gave rise to _ Homo antecessor _, found in Spain . _ Homo heidelbergensis _ originated from _ Homo erectus_ in an unknown location and dispersed across Africa, southern Asia and southern Europe (other scientists interpret fossils, here named _heidelbergensis_, as late _erectus_). _ Homo sapiens sapiens _ spread from Africa to western Asia and then to Europe and southern Asia, eventually reaching Australia and the Americas. In addition to Neanderthals and Denisovans , a third gene flow of archaic Africa origin is indicated at the right.
* ^ 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 of Fossil Vertebrates 1954-1958_. New York: The Geological Society of America, Inc. p. 556. * ^ "Neanderthal". _Wiley-Blackwell Encyclopedia of 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 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 patterning of 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 Human Evolution_. 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 Neanderthals die out?". _The Guardian_. Retrieved April 6, 2017. "It was once thought we appeared in Europe around 35,000 years ago and that we coexisted with Neanderthals for thousands of years after that. They may have hung on in pockets – including caves in Gibraltar – until 28,000 years ago " 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 News_. * ^ _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). berkeley.edu. * ^ _ Smithsonian Institution _ _National Museum of American History _. "Ancient 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 :2009PNAS..10616022H. 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 :10.1073/pnas.0308085100 . * ^ "Scientists Identify Neanderthal Genes in Modern Human DNA". Sci-News.com. January 30, 2014. Retrieved October 29, 2015. * ^ " Homo neanderthalensis_ King, 1864". _Wiley-Blackwell Encyclopedia of 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 1, 2016. * ^ 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 _ Homo sapiens 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 Neanderthal 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 human DNA in a Neanderthal individual". _ScienceDaily_. February 17, 2016. Retrieved March 6, 2016. * ^ The Combined Landscape of Denisovan and Neanderthal Ancestry in Present-Day Humans, _Current Biology, Sankararaman et al., 26, 1241–1247_, 2016. * ^ "Studies find Neanderthal genes in modern humans". * ^ Zimmer, Carl (December 18, 2013). "Toe Fossil Provides Complete Neanderthal Genome". _ 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 Hall, 2000) * ^ _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 :10.1086/401978 . * ^ Foley, Tim. TalkOrigins Archive . " Neanderthal or Neandertal?". 2005. * ^ 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 Define 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 :1999PNAS...96.7117T. JSTOR 48019 . PMC 33580 _. PMID 10377375 . doi :10.1073/pnas.96.13.7117 . * ^ Duarte, Cidália; Mauricio, João; Pettitt, Paul B.; Souto, Pedro; Trinkaus, Erik; Van Der Plicht, Hans; Zilhao, João (1999). "The early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern human emergence in Iberia" . Proceedings of the National Academy of Sciences_. 96 (13): 7604–9. Bibcode :1999PNAS...96.7604D. JSTOR 48106 . PMC 22133 _. PMID 10377462 . doi :10.1073/pnas.96.13.7604 . * ^ 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 26, 2016. * ^ "La Chapelle-Aux-Saints - The old man of La Chapelle - The original reconstruction of the ‘Old Man of La Chapelle’ by scientist Pierre Marcellin Boule led to the reason why popular culture stereotyped Neanderthals as dim-witted brutes for so many years.". Smithsonian Institution. Retrieved July 26, 2016. * ^ "Our Neandertal Brethren: Why They Were Not a Separate Species". SCIENTIFIC AMERICAN. August 1, 2010. Retrieved July 26, 2016. * ^ Wolpoff, MH; Hawks, J ; Caspari, R (2000). "Multiregional, not multiple origins" (pdf). _ American Journal of Physical Anthropology _. 112 (1): 129–36. PMID 10766948 . doi :10.1002/(SICI)1096-8644(200005)112:13.0.CO;2-K . * ^ 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)". _Journal of Human Evolution_. 65 (1): 1–9. doi :10.1016/j.jhevol.2013.01.012 . * ^ 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). " Neandertal 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 (1999). " DNA sequence of the mitochondrial hypervariable region II from the Neandertal type specimen". _Proceedings of the National Academy of Sciences USA_. 96 (10): 5581–5585. doi :10.1073/pnas.96.10.5581 . (subscription required) * ^ P. Beerli; S.V. Edwards (2002). "When did Neanderthals and modern humans diverge?". _Evolutionary Anthropology_. 11 (S1): 60–63. doi :10.1002/evan.10058 . (subscription required) * ^ I.V. Ovchinnikov; A. Götherström; G.P. Romanova; V.M. Kharitonov; K. Lidén; W. Goodwin (2002). "Molecular analysis of Neandertal DNA from the northern Caucasus". _Nature_. 404: 490–493. 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 Neandertal Mitochondrial Genome Sequence Determined by High-Throughput Sequencing" . _Cell_. 134 (3): 416–426. PMC 2602844 _. PMID 18692465 . doi :10.1016/j.cell.2008.06.021 . (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. (2009). "Targeted Retrieval and Analysis of Five Neandertal mtDNA 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 timing of Neanderthal and modern human origins". _Journal of Human Evolution_. 59 (1): 87–95. doi :10.1016/j.jhevol.2010.04.005 . (subscription required) * ^ 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 required) * ^ 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 :10.1006/jasc.2002.0834 . * ^ 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 . * ^ " Homo neanderthalensis". 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 - Homo neanderthalensis - Details - Key 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 (link ) * ^ _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 :10.1086/203725 . * ^ "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 El Sidrón Neanderthal site". _Annals of Anatomy - Anatomischer Anzeiger_. 194: 133–137. doi :10.1016/j.aanat.2011.01.014 . * ^ _A_ _B_ Krings, M; Stone, A; Schmitz, RW; Krainitzki, H; Stoneking, M; Pääbo, S (1997). " Neandertal DNA sequences and the origin of modern humans". _Cell_. 90 (1): 19–30. 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 :10.1073/pnas.192464099 . * ^ Duarte, C; Maurício, J; Pettitt, PB; Souto, P; Trinkaus, E; Van Der Plicht, H; Zilhão, J; et al. (1999). "The early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern human emergence in the Iberian Peninsula". Proceedings of the National Academy of Sciences of the United States of America_. PNAS . 96 (13): 7604–7609. Bibcode :1999PNAS...96.7604D. PMC 22133 _. PMID 10377462 . doi :10.1073/pnas.96.13.7604 . Retrieved June 21, 2009. * ^ 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. Bibcode :2010Sci...328..710G. 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 Neanderthal and Non-African Genetic Similarity" . Molecular Biology and Evolution_. 29 (10): 2987–2995. PMC 3457770 _. PMID 22513287 . doi :10.1093/molbev/mss117 . * ^ 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 Neanderthal movement" (Press release). Durham University . February 11, 2008. Retrieved May 18, 2009. * ^ Wade, Nicholas (October 2, 2007). " Fossil DNA Expands Neanderthal Range". _The New York Times _. Retrieved May 18, 2009. * ^ Ravilious, Kate (October 1, 2007). "Neandertals Ranged Much Farther East Than Thought". 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". _Journal of Human Evolution_. 47 (1–2): 3–17. PMID 15288521 . doi :10.1016/j.jhevol.2004.05.002 . * ^ 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" . _ Academia.edu _ (in 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). " Neanderthals \'were 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 \' Homo sapiens\'". * ^ " 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 humans." * ^ Abramiuk, Marc. _The Foundations of Cognitive Archaeology_, p. 199 (MIT Press 2012): "the encephalization quotient was slightly smaller". * ^ Froehle, Andrew W.; Churchill, Steven E. (2009). "Energetic Competition Between Neandertals and Anatomically Modern Humans" (PDF). _PaleoAnthropology_: 96–116. * ^ Moskvitch, Katia (September 24, 2010). " Neanderthals were able to \'develop their own tools\'". _ BBC News _. BBC. Retrieved October 1, 2010. * ^ 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 of Human Evolution_. 49 (1): 71–87. PMID 15869783 . doi :10.1016/j.jhevol.2005.03.003 . * ^ _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): 7663–6. Bibcode :2000pnas...97.7663r. 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 foods in Neanderthal diets ( Shanidar III, Iraq; Spy I and II, Belgium)" . Proceedings of the National Academy of Sciences_. 108 (2): 486–491. Bibcode :2011PNAS..108..486H. 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, 2017. * ^ Nicola Davis (March 8, 2017). " Neanderthal dental tartar reveals plant-based diet – and drugs". _The Guardian_. Retrieved March 9, 2017. * ^ 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 :10.1006/jhev.1999.0301 . * ^ 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 :10.1073/pnas.1014588107 * ^ 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 Re-evaluation of 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 (4). * ^ " 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 :10.1371/journal.pone.0045927 . * ^ 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) . "Early Neanderthal Constructions deep in Bruniquel Cave in Southwestern France". _Nature_. 534 (7605): 111–114. Bibcode :2016Natur.534..111J. ISSN 0028-0836 . PMID 27251286 . doi :10.1038/nature18291 . * ^ 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 bases)." * ^ Green RE, Krause J, Ptak SE, et al. (November 2006). "Analysis of one million base pairs of Neanderthal DNA". _Nature_. 444 (7117): 330–6. Bibcode :2006Natur.444..330G. PMID 17108958 . doi :10.1038/nature05336 . * ^ Wade, Nicholas (November 15, 2006). "New Machine Sheds Light on DNA of Neanderthals". _The 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 :10.1126/science.316.5827.967a . * ^ " 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 Neanderthal genes". MSNBC . Retrieved May 18, 2009. * ^ " Neanderthal Genome Sequencing Yields Surprising Results And Opens A New Door To Future Studies" (Press release). Lawrence Berkeley National Laboratory . November 16, 2006. Retrieved May 31, 2009. * ^ _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 :10.1126/science.323.5916.866 . * ^ 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 Neanderthal genome lives on in modern humans". Retrieved October 7, 2016. * ^ "Surprise! 20 Percent of Neanderthal Genome Lives On in Modern Humans, Scientists Find". National Geographic. Retrieved October 7, 2016. * ^ "Ancient DNA and Neanderthals - The Smithsonian Institution\'s Human Origins Program". * ^ "BBC NEWS - Science/Nature - Neanderthals \'were flame-haired\'". * ^ 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). " Neanderthals \'unlikely to have interbred with human ancestors\'". _The Guardian_. Retrieved May 24, 2014. * ^ _A_ _B_ Lowery, Robert K.; Uribe, Gabriel; Jimenez, Eric B.; Weiss, Mark A.; Herrera, Kristian J.; Regueiro, Maria; Herrera, Rene J. (2013). " Neanderthal and Denisova genetic affinities with contemporary humans: Introgression versus common ancestral polymorphisms". _Gene_. 530 (1): 83–94. ISSN 0378-1119 . PMID 23872234 . doi :10.1016/j.gene.2013.06.005 . * ^ Neanderthal breeding idea doubted. BBC. Retrieved March 2, 2016. * ^ _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 :10.1016/j.ajhg.2014.12.029 . * ^ 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 _. 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 of 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, 2016. * ^ _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. (subscription required) * ^ _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 : 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 :10.1371/journal.pbio.0020421 . * ^ 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 _. 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, 2016. * ^ 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 Neanderthal within._, 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): 17196–201. Bibcode :2006PNAS..10317196S. 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 Neandertal Origin is Present Among All Non-African Populations". _Molecular Biology and Evolution_. 28 (7): 1957–62. PMID 21266489 . doi :10.1093/molbev/msr024 . * ^ Viegas, Jennifer (July 18, 2011). "All Non-Africans Part Neanderthal, 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). " DNA Turning Human Story 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 human- Neanderthal interbreeding theory". The Guardian _. Retrieved February 19, 2015. * ^ Hawks, John (2013). "Significance of Neandertal and Denisovan Genomes in Human Evolution". _Annual Review of Anthropology_. 42: 433–49. doi :10.1146/annurev-anthro-092412-155548 . * ^ 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 :10.1371/journal.pone.0059781 . * ^ The Neanderthal Dead, exploring mortuary variability in middle paleolithic eurasia. 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 :10.1038/363252a0 . * ^ 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 Neanderthals from 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 (link ) * ^ 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, 2012.
* Boë, Louis-Jean; Heim, Jean-Louis; Honda, Kiyoshi; Maeda, Shinji (2002). "The potential Neandertal vowel space was as large as that of modern humans". _Journal of Phonetics_. 30 (3): 465–84. doi :10.1006/jpho.2002.0170 . * Lieberman, Philip (October 2007). "Current views on Neanderthal speech capabilities: A reply to Boe et al. (2002)". _Journal of Phonetics_. 35 (4): 552–63. doi :10.1016/j.wocn.2005.07.002 . * Serre, David; Langaney, André; Chech, Mario; Teschler-Nicola, Maria; Paunovic, Maja; Mennecier, Philippe; Hofreiter, Michael; Possnert, Göran; Pääbo, Svante (2004). "No Evidence of Neandertal mt 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): 332–5. Bibcode :2005Natur.435..332W. PMID 15902255 . doi :10.1038/nature03585 . * 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 27, 2010).
* 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 .
* Sankararaman, Sriram; Mallick,