Compared to modern humans, Neanderthals were stockier, with shorter legs and a bigger body. In conformance with Bergmann's rule, this likely was an adaptation to preserve heat in cold climates. Male and female Neanderthals had cranial capacities averaging 1,600 cm3 (98 cu in) and 1,300 cm3 (79 cu in), respectively,
within the range of the values for anatomically modern humans.
Males stood 164 to 168 cm (65 to 66 in) and females 152 to 156 cm (60 to 61 in) tall.
Since 2010, evidence for substantial admixture of Neanderthals DNA in modern populations has accumulated.
Evidence of admixture was found in both European and Asian populations, but not in Africans,
suggesting that interbreeding between Neanderthals and anatomically modern humans took place after the recent "out of Africa" migration, likely between 60,000 and 40,000 years ago.
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 species.
King's name had priority over the proposal put forward in 1866 by Ernst Haeckel, Homo stupidus.
Popular English usage of "Neanderthal" as shorthand for "Neanderthal man", as in "the Neanderthals" or "a Neanderthal", emerged in the popular literature of the 1920s.
Since the historical spelling -th- in German represents the phoneme /t/ or /tʰ/, not the fricative /θ/, standard British pronunciation of "Neanderthal" is with /t/ (IPA: /niːˈændərtɑːl/).
Due to the usual significance of the digraph ⟨th⟩ in English, "Neanderthal" is also pronounced with the voiceless fricative /θ/, at least in "layman's American English" (as /niːˈændərθɔːl/).
The spelling Neandertal is occasionally seen in English, even in scientific publications.
Since "Neanderthal", or "Neandertal", is a common name, there is no authoritative prescription on its spelling, unlike the spelling of the binomional name H. neanderthalensis, which is predicated by King 1864.
The common name in German is always invariably Neandertaler (lit. "of the valley of Neander"), not Neandertal,
but the spelling of the name of the Neander Valley itself (Neandertal vs. Neanderthal) has been affected by the
species name, the name of the Neanderthal Museum as well as Neanderthal station being retained in pre-1900 orthography.
Ever since the discovery of the Neanderthal fossils, expert opinion has been divided as to whether Neanderthals should be considered a separate species (Homo neanderthalensis) or a subspecies (Homo sapiens neanderthalensis) relative to modern humans.Pääbo (2014) described such "taxonomic wars" as unresolveable in principle, "since there is no definition of species perfectly describing the case."
The question depends on the definition of Homo sapiens as a chronospecies, which has also been in flux throughout the 20th century. Authorities preferring classification of Neanderthals as subspecies have introduced the subspecies name Homo sapiens sapiens for the anatomically modern Cro Magnon population which lived in Europe at the same time as Neanderthals, while authorities preferring classification as separate species use Homo sapiens as equivalent to "anatomically modern humans".
During the early 20th century, a prevailing view of Neanderthals as "simian", influenced by Arthur Keith and Marcellin Boule, tended to exaggerate the anatomical differences between Neanderthals and Cro Magnon.
Beginning in the 1930s, revised reconstructions of Neanderthals increasingly emphasized the similarity rather than differences from modern humans. From the 1940s throughout the 1970s, it was increasingly common to use the subspecies classification of
Homo sapiens neanderthalensis vs. Homo sapiens sapiens.
The hypothesis of "multiregional origin" of modern man was formulated in the 1980s on such grounds, arguing for the presence of an unbroke succession of fossil sites in both Europe and Asia.
Hybridization between Neanderthals and Cro Magnon had been suggested on skeletal and craniological grounds since the early 20th century, and found increasing support in the later 20th century, until Neanderthal admixture was found to be present in modern populations genetics in the 2010s.
Mainstream opinion by the 2010s appears to favour derivation of both Neanderthals and anatomically modern humans from Homo heidelbergensis.
which derived from H. erectus by about 0.6 million years ago.Homo antecessor, an archaic human species postulated in 1997, and assumed to be the immediate predecessor of H. heidelbergensis, has also been suggested as the last common ancestor of Neanderthals and anatomically modern humans.
The taxonomic distinctions between H. heidelbergensis and Neanderthals is mostly due to a fossil gap in Europe between
300,000 and 243,000 years ago (MIS 8). "Neanderthals" by conventions are fossils which date to after this gap.
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. In morphological studies, the term "classic Neandertrhal" may be used in a narrower sense for Neanderthals younger than 71,000 years old (MIS 4 and 3).
Habitat and range
Sites where typical Neanderthal fossils have been found. Ice sheets of the Last Glacial Maximum are indicated (partly ice-free during the Eemian interglacial)
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 Central to Western Eurasia, namely neither to the south of 35° N (Shuqba, Levant), nor east of 85° E (Denisova, Siberia), nor north of 55° N (Bontnewydd, Wales),
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.
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,[b] 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 patterns similar to anatomically modern humans. They concluded that, "We don't yet know for sure, but it seems likely that, as part of their adaptation to cold, Neanderthals were furry. Chimpanzees have ridges on their finger bones that stem from the way that they clutch their mothers' fur as infants. Modern humans don't have these ridges, but Neanderthals do."
In 2017, researchers using 3D reconstructions of nasal cavities and Computational Fluid Dynamics techniques have found that Neanderthals and modern humans both adapted their noses (independently and in a convergent way) to help breathe in cold and dry conditions. The large nose seen in Neanderthals, as well as Homo heidelbergensis, affected the shape of the skull and the muscle attachments, and gave them a weaker bite force than in modern humans.
In The Spread of Modern Humans in Europe (2002) John F. Hoffecker, 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 drainage of the dural venous sinuses (low pressure blood vessels that run between the meninges and skull leading down through the skull) in the occipital lobe region of Neanderthal brains appears more asymmetric than other hominid brains. In 2008, a group of scientists produced a study using three-dimensional computer-assisted reconstructions of Neanderthal infants based on fossils found in Russia and Syria. It indicated that Neanderthal and modern human brains were the same size at birth, but that by adulthood, the Neanderthal brain was larger than the modern human brain. They had almost the same degree of encephalisation (i.e. brain-to-body-size ratio) as modern humans.
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 and used fire, and were hunters. The consensus on their behaviour ends there. It had actually long been debated whether Neanderthals were hunters or scavengers, but the discovery of the pre-Neanderthal Schöningen wooden spears in Germany helped settle the debate in favour of hunting. Most available evidence suggests they were apex predators, and fed on red deer, reindeer, ibex, wild boar, aurochs and on occasion mammoth, straight-tusked elephant and rhinoceros. They appear to have occasionally used vegetables as fall-back food, revealed in the 2000s and 2010s by isotope analysis of their teeth and study of their coprolites (fossilised faeces). Dental analysis of specimens from Spy, Belgium and El Sidrón, Spain in 2017 suggested that these Neanderthals had a wide-ranging diet, and that those "from El Sidrón showed no evidence of meat eating" at all and seemed to have lived on "a mixture of forest moss, pine nuts and a mushroom known as split gill".
The size and distribution of Neanderthal sites, along with genetic 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 northwestern Spain. They are believed to have been a group killed and butchered about 50,000 years ago. Analysis of the mtDNA showed that the three adult males belonged to the same maternal lineage, while the three adult females belonged to different ones. This suggests a social structure where males remained in the same social group and females "married out".
The bones of the El Sidrón group show signs of defleshing, suggesting that they were victims of cannibalism. The St. Césaire 1 skeleton discovered in 1979 at La Roche à Pierrot, France, showed a healed fracture on top of the skull apparently caused by a deep blade wound, suggesting interpersonal violence.
Claims that Neanderthals deliberately buried their dead, and if they did, whether such burials had any symbolic meaning,:158–60 are heavily contested. The debate on deliberate Neanderthal burials has been active since the 1908 discovery of the well-preserved Chapelle-aux-Saints 1 skeleton in a small hole in a cave in southwestern France. In this controversy's most recent installment, a team of French researchers reinvestigated the Chapelle-aux-Saints cave and in January 2014 reasserted the century-old claim that the 1908 Neanderthal specimen had been deliberately buried, and this has in turn been heavily criticised.
Question of art and adornment
Whether Neandethals created art and used adornments, which would indicate a capability for complex symbolic thought, remains unresolved. A 2010 paper on radiocarbon dates cast doubt on the association of Châtelperronian beads with Neanderthals, and Paul Mellars considered the evidence for symbolic behavior to have been refuted. 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.
A very large number of other claims of Neanderthal art, adornment, and structures have been made. 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.
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.
Two 176,000-year-old stalagmite ring structures, several metres wide, were reported in 2016 more than 300 metres from the entrance within Bruniquel Cave, France. The authors claim artificial lighting would have been required as this part of the cave is beyond the reach of daylight and that the structures had been made by early Neanderthals, the only humans in Europe at this time.
In 2015, a study argued that a number of 130,000-year-old eagle talons found in a cache near Krapina, Croatia along with Neanderthal bones, had been modified to be used as jewellery.
All of these appeared only in single locations. Yet in 2018, using uranium-thorium dating methods, red painted symbols comprising a scalariform (ladder shape), a negative hand stencil, and red lines and dots on the cave walls of three Spanish caves 700 km (430 mi) apart were dated to at least 64,000 years old. If the dating is correct, they were painted before the time anatomically modern humans are thought to have arrived in Europe. Paleoanthropologist John D. Hawks argues these findings demonstrate Neanderthals were capable of symbolic behavior previously thought to be unique to modern humans.
Until the early 1950s, most scholars believed Neanderthals were not in the ancestry of living humans.:232–234 Nevertheless, Thomas H. Huxley in 1904 saw among Frisians the presence of what he believed to be Neanderthaloid skeletal and cranial characteristics as an evolutionary development from Neanderthal rather than as a result of interbreeding, saying that "the blond long-heads may exhibit one of the lines of evolution of the men of the Neanderthaloid type," yet he raised the possibility that the Frisians alternatively "may be the result of the admixture of the blond long-heads with Neanderthal men," thus separating "blond" from "Neanderthaloid."
Hans Peder Steensby proposed interbreeding in 1907 in the article Race studies in Denmark. He strongly emphasised that all living humans are of mixed origins. He held that this would best fit observations, and challenged the widespread idea that Neanderthals were ape-like or inferior. Basing his argument primarily on cranial data, he noted that the Danes, like the Frisians and the Dutch, exhibit some Neanderthaloid characteristics, and felt it was reasonable to "assume something was inherited" and that Neanderthals "are among our ancestors."
Carleton Stevens Coon in 1962 found it likely, based upon evidence from cranial data and material culture, that Neanderthal and Upper Paleolithic peoples either interbred or that the newcomers reworked Neanderthal implements "into their own kind of tools." Christopher Thomas Cairney in 1989 went further, laying out a rationale for hybridisation and adding a broader discussion of physical characteristics as well as commentary on interbreeding and its importance to adaptive European phenotypes. Cairney specifically discussed the "intermixture of racial elements" and "hybridisation."
In 2010, geneticists announced that interbreeding had likely taken place, a result confirmed in 2012. The genomes of all non-Africans include portions that are of Neanderthal origin, a share estimated in 2014 to 1.5–2.1%. This DNA is absent in Sub-Saharan Africans (Yoruba people and San subjects).Ötzi the iceman, Europe's oldest preserved mummy, was found to possess an even higher percentage of Neanderthal ancestry. The two percent of Neanderthal DNA in Europeans and Asians is not the same in all Europeans and Asians: In all, approximately 20% of the Neanderthal genome appears to survive in the modern human gene pool.
2012 genetic studies seem to suggest that modern humans may have mated with "at least two groups" of archaic humans: 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.
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.[d] 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).
mtDNA-based simulation of modern human expansion in Europe starting 1,600 generations ago. Neanderthal range in light grey
About 55,000 years ago, the climate began to fluctuate wildly from extreme cold conditions to mild cold and back in a matter of decades. Neanderthal bodies were well-suited for survival in a cold climate—their stocky chests and limbs stored body heat better than 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.
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 long before the arrival of modern humans. These modern humans may have introduced a disease that contributed to the extinction of Neanderthals, and that may be added to other recent explanations for their extinction. When Neanderthal ancestors left Africa roughly 100,000 years earlier they adapted to the pathogens in their European environment, unlike modern humans who adapted to African pathogens. This transcontinental movement is known as the Out of Africa model. If contact between humans and Neanderthals occurred in Europe and Asia the first contact may have been devastating to the Neanderthal population, because they would have had little if any immunity to the African pathogens. More recent historical events in Eurasia and the Americas show a similar pattern, where the unintentional introduction of viral or bacterial pathogens to unprepared populations has led to mass mortality and local population extinction. 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.
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.
1856: Limestone miners discover the Neanderthal-type specimen, Neanderthal 1, in Neandertal, western Prussia (now Germany).
1864: William King is the first to recognise Neanderthal 1 as belonging to a separate species, for which he gives the scientific name Homo neanderthalensis. He then changed his mind on placing it in the genus Homo, 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 the Czech Republic), associated with cultural debris, including hearths, Mousterian tools, and bones of extinct animals.
1886: Two well-preserved Neanderthal skeletons are found at Spy, Belgium, making the hypothesis that Neanderthal 1 was only a diseased modern human difficult to sustain.
1899: Sand excavation workers find hundreds of fragmentary Neanderthal remains representing at least 12 and likely as much as 70 individuals on a hill in Krapina, in the Austro-Hungarian Empire (now Croatia).
Neanderthal 1: The first human bones recognised as showing a non-modern anatomy. Discovered in 1856 in a limestone quarry at the Feldhofer grotto in Neanderthal, Germany, they consist of a skull cap, the two femora, the three right arm bones, two left arm bones, the ilium, and fragments of a scapula and ribs.
La Chapelle-aux-Saints 1: Called the Old Man, a fossilised skull discovered in La Chapelle-aux-Saints, France, by A. and J. Bouyssonie, and L. Bardon in 1908. Characteristics include a low vaulted cranium and large browridge typical of Neanderthals. Estimated to be about 60,000 years old, the specimen was severely arthritic and had lost all his teeth long before death, leading some to suggest he was cared for by others.
La Ferrassie 1: A fossilised skull discovered in La Ferrassie, France, by R. Capitan in 1909. It is estimated to be 70,000 years old. Its characteristics include a large occipital bun, low-vaulted cranium and heavily worn teeth.
Le Moustier 1: One of the rare nearly complete 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.
Shanidar 1 to 10: Eight Neanderthals and two pre-Neanderthals (Shanidar 2 and 4) were discovered in the Zagros Mountains in Iraqi Kurdistan. One of the skeletons, Shanidar 4, was once thought to have been buried with flowers, a theory no longer accepted. To Paul B. Pettitt the "deliberate placement of flowers has now been convincingly eliminated", since "[a] recent examination of the microfauna from the strata into which the grave was cut suggests that the pollen was deposited by the burrowing rodent Meriones tersicus, which is common in the Shanidar microfauna and whose burrowing activity can be observed today".
Amud 1: A male adult Neanderthal, dated to roughly 55,000 BP, and one of several found in a cave at Nahal Amud, Israel. At 178 cm (70 in), it is the tallest known Neanderthal. It also has the largest cranial capacity of all extinct hominins: 1,736 cm3.
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.
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 mtDNA from a second specimen was reported in 2000, and showed no sign of modern human descent from Neanderthals.
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.
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 realised that 11% of their sample was modern human DNA. Since then, more of the preparation work has been done in clean areas and 4-base pair 'tags' have been added to the DNA as soon as it is extracted so the Neanderthal DNA can be identified.
With 3 billion nucleotides sequenced, analysis of about one third 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.
A analysis of a first draft of the Neanderthal genome by the same team released in May 2010 indicated 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 [later refined to 1.5 to 2.1 percent]. It is a small but very real proportion of ancestry in non-Africans today," says Dr. David Reich of Harvard Medical School, who worked on the study. This research compared the genome of the Neanderthals to five modern humans from China, France, sub-Saharan Africa, and Papua New Guinea.
This indicates a gene flow from Neanderthals to modern humans, i.e., interbreeding between the two populations. Since the three non-African genomes show a similar proportion of Neanderthal sequences, the interbreeding must have occurred early in the migration of modern humans out of Africa, perhaps in the Middle East. No evidence for gene flow in the direction from modern humans to Neanderthals was found. Gene flow from modern humans to Neanderthals would not be expected if contact occurred between a small colonising population of modern humans and a much larger resident population of Neanderthals. A very limited amount of interbreeding could explain the findings, if it occurred early enough in the colonisation process.
It is suggested that 20 percent of Neanderthal DNA survived in modern humans, notably expressed in the skin, hair and diseases of modern people. Modern human genes involved in making keratin—the protein found in skin, hair, and nails—have specially high levels of Neanderthal DNA. For example, around 66% of East Asians contain the Neanderthal skin gene, while 70% of Europeans possess the Neanderthal gene which affects skin colour. POU2F3 is found in around 66 percent of East Asians, while the Neanderthal version of BNC2, which affects skin color, among other traits, is found in 70 percent of Europeans. Neanderthal are the variants in genes that affect the risk of several diseases, including lupus, biliary cirrhosis, Crohn's disease, and type 2 diabetes. Eight percent of Neanderthal DNA comes from an unknown group of archaic humans, tantalising hints of unknown groups from Asia and Africa that left genes in Denisovans and modern humans, respectively. The genetic variant of the MC1R gene linked to red hair in Neanderthals has not been found in modern humans; hence, red hair may be an example of convergent evolution.
While interbreeding is viewed as the most parsimonious interpretation of the genetic discoveries, the authors point out they cannot conclusively rule out an alternative scenario, in which the source population of non-African modern humans was already more closely 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.
Among the genes shown to differ between present-day humans and Neanderthals were RPTN, SPAG17, CAN15, TTF1, FOXP2 and PCD16.
Specifically, a visualisation map of the reference modern-human containing the genome regions with high degree of similarity or with novelty according to a Neanderthal of 50k has been built by Pratas et al.
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 [that the greater proportion of Neandertal ancestry in East Asians than in Europeans is due to the fact that purifying selection is less effective at removing weakly deleterious Neandertal alleles from East Asian populations] cannot account for the higher proportion of Neandertal ancestry in East Asians than in Europeans. Instead, more complex demographic scenarios, most likely involving multiple pulses of Neandertal admixture, are required to explain the data."
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.
There have been at least three episodes of interbreeding. The first would have occurred soon after modern humans left Africa. The second would have occurred after the ancestral Melanesians had branched off—these people seem to have thereafter bred with Denisovans. The third would have involved Neanderthals and the ancestors of East Asians only.
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.
A 2014 study on epigenetics of the Neanderthal published 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.
In popular culture
Neanderthals have been portrayed in popular culture including appearances in literature, visual media and comedy. Early 20th century artistic interpretations often presented Neanderthals as beastly creatures, emphasising hairiness and rough, dark complexion.
The German noun is cognate with English dale.
The German /t/ phoneme was frequently spelled th throughout the 15th to 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.
^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.
^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.
Reich, David (2018). Who We Are And How We Got Here - Ancient DNA and the New Science of the Human Past. Pantheon Books. ISBN 978-1101870327.
^ abPääbo, Svante (2014). Neanderthal Man: In Search of Lost Genomes. New York: Basic Books. p. 237.
^ abcT. Higham, K. Douka, R. Wood, C.B. Ramsey, F. Brock, L. Basell, M. Camps, A. Arrizabalaga, J. Baena, C. Barroso-Ruíz, C. Bergman, C. Boitard, P. Boscato, M. Caparrós, N.J. Conard, C. Draily, A. Froment, B. Galván, P. Gambassini, A. Garcia-Moreno, S. Grimaldi, P. Haesaerts, B. Holt, M.-J. Iriarte-Chiapusso, A. Jelinek, J.F. Jordá Pardo, J.-M. Maíllo-Fernández, A. Marom, J. Maroto, M. Menéndez, L. Metz, E. Morin, A. Moroni, F. Negrino, E. Panagopoulou, M. Peresani, S. Pirson, M. de la Rasilla, J. Riel-Salvatore, A. Ronchitelli, D. Santamaria, P. Semal, L. Slimak, J. Soler, N. Soler, A. Villaluenga, R. Pinhasi, R. Jacobi (2014). "The timing and spatiotemporal patterning of Neanderthal disappearance". Nature. 512 (7514): 306–09. Bibcode:2014Natur.512..306H. doi:10.1038/nature13621. PMID25143113. We show that the Mousterian [the Neanderthal tool-making tradition] ended by 41,030–39,260 calibrated years BP (at 95.4% probability) across Europe. We also demonstrate that succeeding 'transitional' archaeological industries, one of which has been linked with Neanderthals (Châtelperronian), end at a similar time.
T. Higham (2011). "European Middle and Upper Palaeolithic radiocarbon dates are often older than they look: problems with previous dates and some remedies"(PDF). Antiquity. 85 (327): 235–49. doi:10.1017/s0003598x00067570. Few events of European prehistory are more important than the transition from ancient to modern humans around 40 000 years ago, a period that unfortunately lies near the limit of radiocarbon dating. This paper shows that as many as 70 per cent of the oldest radiocarbon dates in the literature may be too young, due to contamination by modern carbon.
^ abR. Pinhasi, T.F.G. Higham, L.V. Golovanova, V.B. Doronichev (2011). "Revised age of late Neanderthal occupation and the end of the Middle Paleolithic in the northern Caucasus". Proceedings of the National Academy of Sciences USA. 108 (21): 8611–16. Bibcode:2011PNAS..108.8611P. doi:10.1073/pnas.1018938108. The direct date of the fossil (39,700 ± 1,100 14C BP) is in good agreement with the probability distribution function, indicating at a high level of probability that Neanderthals did not survive at Mezmaiskaya Cave after 39 ka cal BP. [...] This challenges previous claims for late Neanderthal survival in the northern Caucasus. [...] Our results confirm the lack of reliably dated Neanderthal fossils younger than ≈40 ka cal BP in any other region of Western Eurasia, including the Caucasus.
^McKie, Robin (June 2, 2013). "Why did the Neanderthals die out?". The Guardian. Retrieved April 6, 2017. "It was once thought we appeared in Europe around 35,000 years ago and that we coexisted with Neanderthals for thousands of years after that. They may have hung on in pockets – including caves in Gibraltar – until 28,000 years ago [said Chris Stringer]" Previous research on Neanderthal sites which suggested that they were more recent than 40,000 years old appears to be wrong," said Stringer. "That is a key finding that will be discussed at the conference."[...] However, scientists have set out to get round these problems. At Oxford University, scientists led by Tom Higham have developed new methods to remove contamination and have been able to make much more precise radiocarbon dating for this period.
^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.
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extending to 1,736 cm3 (105.9 cu in) in the male Amud 1.
Amano, H.; Kikuchi, T.; Morita, Y.; Kondo, O.; Suzuki, Hiromasa; et al. (August 2015). "Virtual Reconstruction of the Neanderthal Amud 1 Cranium". American Journal of Physical Anthropology. 158 (2): 185–97. doi:10.1002/ajpa.22777.
^"Neanderthal". Wiley-Blackwell Encyclopedia of Human Evolution. Chichester, West Sussex: Wiley-Blackwell. 2013.
^"Neandertal oder Neanderthal? – Was ist denn nun richtig?". mettmann.de. Neanderthal museum. Retrieved February 1, 2017. Heute sollten Ortsbezeichnungen das „Neandertal“ ohne „h“ bezeichnen. Alle Namen, die sich auf den prähistorischen Menschen beziehen, führen das „h“. [Today one should write for place names "Neandertal" without an "h". All names related to the prehistoric humans keep the "h".]
^ abD. Dean; J.-J. Hublin; R. Holloway; R. Ziegler (1998). "On the phylogenetic position of the pre-Neandertal specimen from Reilingen, Germany". Journal of Human Evolution. 34 (5). pp. 485–508. doi:10.1006/jhev.1998.0214.
^ abcdePapagianni, Dmitra; Morse, Michael (2013). The Neanderthals Rediscovered. Thames & Hudson. ISBN 978-0-500-05177-1.
^ abcStringer, C.; Gamble, C. (1993). In Search of the Neanderthals. London: Thames and Hudson. ISBN 978-0500050705.
^B. Vandermeersch; M.D. Garralda (2011). S. Condemi; G.-C. Weniger, eds. "Continuity or Discontinuity in the Peopling of Europe: One Hundred and Fifty Years of Neanderthal Study". Vertebrate Paleobiology and Paleoanthropology. Springer Netherlands. pp. 113–25. doi:10.1007/978-94-007-0492-3_10.
^N.J. Conard; J. Richter, eds. (2011). "2". Neanderthal Lifeways, Subsistence and Technology. Vertebrate Paleobiology and Paleoanthropology. 19. Springer. pp. 7–14. doi:10.1007/978-94-007-0415-2_2. ISBN 978-94-007-0414-5.
^Silberman, Neil. The Oxford Companion to Archaeology, p. 455 (Oxford University Press 2012): "[I]t is with the Neanderthals that we see the full achievement, for the first time, of the degree of encephalization (brain to body size ratio) that characterizes modern humans."
^Abramiuk, Marc. The Foundations of Cognitive Archaeology, p. 199 (MIT Press 2012): "the encephalization quotient was slightly smaller".
^Bocherens, Hervé; Drucker, Dorothée G.; Billiou, Daniel; Patou-Mathis, Marylène; Vandermeersch, Bernard (2005). "Isotopic evidence for diet and subsistence pattern of the Saint-Césaire I Neanderthal: Review and use of a multi-source mixing model". Journal of Human Evolution. 49 (1): 71–87. doi:10.1016/j.jhevol.2005.03.003. PMID15869783.
^ abRendu 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.
^ abGargett, R.H. (1989). "Grave Shortcomings: The Evidence for Neandertal Burial". Current Anthropology. 30 (2): 157–90. doi:10.1086/203725.
^Gargett, R.H. (1999). "Middle Palaeolithic burial is not a dead issue: the view from Qafzeh, Saint-Césaire, Kebara, Amud, and Dederiyeh". Journal of Human Evolution. 37 (1): 27–90. doi:10.1006/jhev.1999.0301. PMID10375476.
^Rendu W, Beauval C, Crevecoeur I, Bayle P, Balzeau A, Bismuth T, Bourguignon L, Delfour G, Faivre JP, Lacrampe-Cuyaubère F, Tavormina C, Todisco D, Turq A, Maureille B (January 2014). "Evidence supporting an intentional Neandertal burial at La Chapelle-aux-Saints". Proceedings of the National Academy of Sciences. 111 (1): 81–86. Bibcode:2014PNAS..111...81R. doi:10.1073/pnas.1316780110.
^ abH. Dibble and V. Aldeias and P. Goldberg and D. Sandgathe and T.E. Steele (2015). "A critical look at evidence from La Chapelle-aux-Saints supporting an intentional burial". Journal of Archaeological Science. 53: 649–57. doi:10.1016/j.jas.2014.04.019.
^Higham T, Jacobi R, Julien M, David F, Basell L, Wood R, Davies W, Ramsey CB.C (2010). "Chronology of the Grotte du Renne (France) and implications for the context of ornaments and human remains within the Chatelperronian". Proc Natl Acad Sci USA. doi:10.1073/pnas.1007963107PMID20956292
^Mellars P. (2010). "Neanderthal symbolism and ornament manufacture: The bursting of a bubble?" Proc Natl Acad Sci USA 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.
^Pike, A. W.; Hoffmann, D. L.; Pettitt, P. B.; García-Diez, M.; Zilhão, J. (2017). "Dating Palaeolithic cave art: Why U–Th is the way to go". Quaternary International. 432: 41–49. doi:10.1016/j.quaint.2015.12.013.
^D. L. Hoffmann; C. D. Standish; M. García-Diez; P. B. Pettitt; J. A. Milton; J. Zilhão; J. J. Alcolea-González; P. Cantalejo-Duarte; H. Collado; R. de Balbín; M. Lorblanchet; J. Ramos-Muñoz; G.-Ch. Weniger; A. W. G. Pike (2018). "U-Th dating of carbonate crusts reveals Neandertal origin of Iberian cave art". Science. 359 (6378): 912–915. doi:10.1126/science.aap7778.
^Liu, Prugnolle et al. (2006). "Currently available genetic and archaeological evidence is supportive of a recent single origin of modern humans in East Africa. However, this is where the consensus on human settlement history ends, and considerable uncertainty clouds any more detailed aspect of human colonization history."
^Lohse, Konrad; Frantz, Laurent A. F. (2013). "Maximum likelihood evidence for Neandertal admixture in Eurasian populations from three genomes". Populations and Evolution. 1307: 8263. arXiv:1307.8263. Bibcode:2013arXiv1307.8263L.
^ abLowery, Robert K.; Uribe, Gabriel; Jimenez, Eric B.; Weiss, Mark A.; Herrera, Kristian J.; Regueiro, Maria; Herrera, Rene J. (2013). "Neanderthal and Denisova genetic affinities with contemporary humans: Introgression versus common ancestral polymorphisms". Gene. 530 (1): 83–94. doi:10.1016/j.gene.2013.06.005. ISSN0378-1119. PMID23872234.
^ abFinlayson, C., Carrión, J.S. (April 2007). "Rapid ecological turnover and its impact on Neanderthal and other human populations". Trends in Ecology & Evolution (Personal Edition). 22 (4): 213–22. doi:10.1016/j.tree.2007.02.001. PMID17300854.
^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.
^S. Genovés (1954). "The problem of the sex of certain fossil hominids, with special reference to the Neandertal skeletons from Spy". The Journal of the Royal Anthropological Institute of Great Britain and Ireland. 84 (1/2). pp. 131–44.
^"Neanderthal Man". Encyclopedia Britannica (15th ed.). 1982.
^Rink, W. Jack; Schwarcz, H.P.; Lee, H.K.; Rees-Jones, J.; Rabinovich, R.; Hovers, E. (August 2002). "Electron spin resonance (ESR) and thermal ionization mass spectrometric (TIMS) 230Th/234U dating of teeth in Middle Paleolithic layers at Amud Cave, Israel". Geoarchaeology. 16 (6): 701–17. doi:10.1002/gea.1017.
^Valladas, Hélène; Merciera, N.; Frogeta, L.; Hoversb, E.; Joronc, J.L.; Kimbeld, W.H.; Rak, Y. (March 1999). "TL Dates for the Neanderthal Site of the Amud Cave, Israel". Journal of Archaeological Science. 26 (3): 259–68. doi:10.1006/jasc.1998.0334.
^R.E. Wood, T.F.G. Higham, T. de Torres, N. Tisnérate-Laborde, H. Valladas, J.E. Ortiz, C. Lalueza-Fox, S. Sánchez-Moral, J.C. Cañaveras, A. Rosas, D. Santamaría, M. de la Rasilla (March 20, 2012). "A new date for the Neanderthals from El Sidrón Cave (Asturias, Northern Spain)". Archaeometry. 55 (1): 148–58. doi:10.1111/j.1475-4754.2012.00671.x.
^Lunine 2013, p. 251: "The Neanderthal genome is about the same size as the human genome, and is identical to that of present-day humans to a level of 99.7% (this is comparing the ordering of the lettering in the nucleotide bases)."
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