Temporal range: Middle–Late Pleistocene 0.25–0.040 Ma
|The Neanderthal skull of La Chapelle-aux-Saints 1|
|Mounted Neanderthal skeleton at American Museum of Natural History|
|Range of Homo neanderthalensis. Eastern and northern ranges may extend to include Okladnikov in Altai and Mamotnaia in Ural|
Neanderthals, or more rarely Neandertals,[a] (UK //, us also //-, -//-, -//, -//; 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. Due to accumulating genetic and fossil evidence suggesting Neanderthals evolved in Europe separately from modern humans in Africa for over 600,000 years, paleontologists generally classify them as a distinct species, Homo neanderthalensis. From the 1950s to the early 1980s, however, Neanderthals were widely considered a subspecies of Homo sapiens (H. s. neanderthalensis) and a minority of scholars still hold this view.
Several cultural assemblages have been linked to the Neanderthals in Europe. The earliest, the Mousterian stone tool culture, dates to about 160,000 years ago. Late Mousterian artifacts were found in Gorham's Cave on the south-facing coast of Gibraltar.
Compared to Homo sapiens, 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 1600 cm3, females 1300 cm3, extending to 1736 cm3 in Amud 1. This is notably larger than the 1250–1400 cm3 typical of modern humans. Males stood 164–168 cm (65–66 in) and females 152–156 cm (60–61 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 a few Neanderthals began mating with ancestors of modern humans long before the large "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.
Claims that Neanderthals deliberately buried their dead, and if they did, whether such burials had any symbolic meaning, 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.
In 2013, scientists sequenced the entire genome of a Neanderthal for the first time. The genome was extracted from the phalanx bone of a 50,000-year-old Neanderthal found in a Siberian cave. In 2016, elaborate constructions of rings of broken stalagmites made by early Neanderthals around 176,000 years ago were discovered 336 m (1,102 ft) inside Bruniquel Cave in southwestern France. This would have required a more advanced social structure than previously known for Neanderthals.
- 1 Name
- 2 Classification
- 3 Origin
- 4 Discovery
- 5 Habitat and range
- 6 Anatomy
- 7 Behavior
- 8 Genome
- 9 Extinction hypotheses
- 10 Interbreeding hypotheses
- 11 Specimens
- 12 Popular culture
- 13 See also
- 14 Notes
- 15 References
- 16 Further reading
- 17 External links
The species is named after one of the first sites where its 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.[b] Thal is an older spelling of the German word Tal (with the same pronunciation), which means "valley" (cognate with English dale).[c]
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 species – 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 [neˈandɐˌtʰaːlɐ] in the International Phonetic Alphabet. In British English, "Neanderthal" is pronounced with the /t/ as in German, but different vowels (IPA: /niːˈændərtɑːl/). In layman's American English, "Neanderthal" is pronounced with a /θ/ (the voiceless th as in thin) and /ɔ/ instead of the longer British /aː/ (IPA: /niːˈændərθɔːl/), although scientists typically use the /t/ as in German.
Scientists to this day debate over whether Neanderthals should be classified as a distinct species - Homo neanderthalensis - or as Homo sapiens neanderthalensis, the latter placing Neanderthals as a subspecies of H. sapiens. 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 this "primitive" Neanderthal could not be a direct ancestor of modern man. This idea was reflected in an erroneous and inaccurate reconstruction of the Neanderthal findings of La Chapelle-aux-Saints, mounted in a crooked pose with a deformed and heavily curved spine and legs buckled.
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 obviously unbroken succession of fossil sites of both subspecies in Europe was considered evidence that there was a slow and 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.
Current scientific ideas hold that both species evolved from a common African ancestor: Homo erectus. Fossil and tool finds support the idea that during the first migration wave around 2 million years ago, small groups of Homo erectus left Africa via the Levant and the Black Sea area into Georgia (fossils of Dmanisi) and possibly via north-west Africa towards southern Spain. The 1.2-million-year-old fragmented finds in Spain are referred to as Homo antecessor by their discoverers, and considered to be the ancestors of Neanderthal. This interpretation, however, is controversial.
Some 600,000 years ago a second propagation wave of African Homo erectus took place. Skulls from that period found in Spain for example, suggest a brain volume of between 1100 and 1450 cm3. The brain volume of fossils from the first propagation wave is however estimated only to be slightly above 1000 ccm. Homo erectus that arrived in Europe in the second wave of colonization developed into Neanderthals through the intermediate Homo heidelbergensis. Since about 200,000 years ago in Africa the so-called early anatomically modern humans descended from the local Homo erectus through the intermediate Homo rhodesiensis. Evidence from mitochondrial DNA studies has been interpreted as evidence that Neanderthals were not a subspecies of H. sapiens. Others, for example University of Cambridge Professor Paul Mellars, say "no evidence has been found of cultural interaction".
The period when the Neanderthal lineage separated from the lineage that led to modern man remains debated. In 2010 a period between 440,000 and 270,000 years B.P. was calculated by the molecular clock method. However, the reliability of molecular clocks is questionable as dating determined by stratigraphic methods varies considerably from those determined using the molecular clock. In 2012 recalculation of the mutation rates revealed evidence of a much earlier separation: between 800,000 and 400,000 years ago.
The first representatives of the genus Homo with proto-Neanderthal traits are believed to have existed in Eurasia as early as 350,000–600,000 years ago with the first "true Neanderthals" appearing between 200,000 and 250,000 years ago. The exact date of their extinction had been disputed but in 2014, a team led by Thomas Higham of the University of Oxford used an improved radiocarbon dating technique on material from 40 archaeological sites to show that Neanderthals died out in Europe between 41,000 and 39,000 years ago, with the last group disappearing from southern Spain 28,000 years ago. Radiocarbon dating performed on charcoal in Gorham's Cave in Gibraltar in 2006 suggests that Neanderthals lived there 24,000 to 28,000 years ago. This coincides with the start of a very cold period in Europe and is 5,000 to 18,000 years after Homo sapiens reached the continent.
Comparison of the DNA of Neanderthals and Homo sapiens suggests that they diverged from a common ancestor between 350,000 and 400,000 years ago. This ancestor was probably Homo heidelbergensis. Heidelbergensis originated between 800,000 and 1,300,000 years ago, and continued until about 200,000 years ago. It ranged over Eastern and South Africa, Europe and Western Asia. Between 350,000 and 400,000 years ago the African branch is thought to have started evolving towards modern humans and the Eurasian branch towards Neanderthals. Scientists do not agree when Neanderthals can first be recognised in the fossil record, with dates ranging between 200,000 and 300,000 years BP.
Neanderthal skulls were first discovered in the Engis Caves (the partial skull dubbed Engis 2) (fr), in what is now Belgium (1829) by Philippe-Charles Schmerling and in Forbes' Quarry, Gibraltar, dubbed Gibraltar 1 (1848), both prior to the type specimen discovery in a limestone quarry of the Neander Valley in Erkrath near Düsseldorf in August 1856, three years before Charles Darwin's On the Origin of Species was published.
The type specimen, dubbed Neanderthal 1, consisted of a skull cap, two femora, three bones from the right arm, two from the left arm, part of the left ilium, fragments of a scapula, and ribs. The workers who recovered this material originally thought it to be the remains of a bear. They 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: The Engis 2 Neanderthal skulls were discovered in Engis, in present-day Belgium.
- 1848: Neanderthal skull Gibraltar 1 found in Forbes' Quarry, Gibraltar. Called "an ancient human" at the time.
- 1856: Johann Karl Fuhlrott first recognized the fossil called "Neanderthal man", discovered in Neanderthal, a valley near Mettmann in what is now North Rhine-Westphalia, Germany.
- 1864: William King proposed the name Homo neanderthalensis at a meeting of the British Association for the Advancement of Science, but then changed his mind and argued that Neanderthals were different enough from humans to warrant a separate genus, under the assumption that they were likely "incapable of moral and theositic conceptions."
- 1880: The mandible of a Neanderthal child was found in a secure context and associated with cultural debris, including hearths, Mousterian tools, and bones of extinct animals.
- 1886: Two nearly perfect skeletons of a man and woman were found at Spy, Belgium at the depth of 16 ft with numerous Mousterian-type implements.
- 1899: Hundreds of Neanderthal bones were described in stratigraphic position in association with cultural remains and extinct animal bones.
- 1899: Sand excavation workers found bone fragments on a hill in Krapina, Croatia called Hušnjakovo brdo. Local Franciscan friar Dominik Antolković requested Dragutin Gorjanović-Kramberger to study the remains of bones and teeth that were found there.
- 1905: During the excavation in Krapina more than 5,000 items were found, of which 874 residue of human origin, including bones of prehistoric man and animals, artifacts.
- 1908: A nearly complete Neanderthal skeleton was discovered in La Chapelle-aux-Saints, France in association with Mousterian tools and bones of extinct animals.
- 1925: Francis Turville-Petre finds the 'Galilee Man' or 'Galilee Skull' in the Zuttiyeh Cave in Wadi Amud in The British Mandate of Palestine (now Israel).
- 1926: Skull fragments of Gibraltar 2, a four-year-old Neanderthal girl, discovered by Dorothy Garrod.
- 1953–1957: Ralph Solecki uncovered nine Neanderthal skeletons in Shanidar Cave in the Kurdistan region of northern Iraq.
- 1975: Erik Trinkaus' study of Neanderthal feet confirmed they walked like modern humans.
- 1981: The Bontnewydd Palaeolithic site, Wales yields the most north-western site in Eurasia.
- 1987: Thermoluminescence results from Israeli fossils dates Kebara 2 to 60,000 BP and humans at Qafzeh to 90,000 BP. These dates were confirmed by electron spin resonance (ESR) dates for Qafzeh (90,000 BP) and Es Skhul (80,000 BP).
- 1991: ESR dates showed the Tabun Neanderthal was contemporaneous with modern humans from Skhul and Qafzeh.
- 1993: 127,000-year-old DNA is found on the child of Sclayn, found in Scladina (fr), Belgium.
- 1994: Neanderthal remains inadvertently uncovered inside the Sidrón Cave in Piloña municipality, Asturias, northwestern Spain. Since then the remains of at least 12 individuals: three men, three adolescent boys, three women, and three infants have been found. In 2009 Neanderthal ancient mtDNA was partially sequenced in HVR region for three distinct Neanderthals there.
- 1997: Matthias Krings et al. are the first to amplify Neanderthal mitochondrial DNA (mtDNA) using a specimen from Feldhofer grotto in the Neander valley.
- 1997–2000: During new excavations in the Neandertal, additional bone fragments are found, some of which fit the fragments found in 1856, thus pinpointing the exact location of the original find. The exact location had previously been unknown, as the site of the find (the „Kleine Feldhofer Grotte“) was destroyed by limestone mining.
- 1998: A team led by pre-history archeologist João Zilhão discovered an early Upper Paleolithic human burial in Portugal, at Abrigo do Lagar Velho, which provided evidence of early modern humans from the west of the Iberian Peninsula. The remains, a largely complete skeleton of an approximately 4-year-old child, buried with pierced shell and red ochre, is dated to ca. 24,500 years BP. The cranium, mandible, dentition, and postcrania present a mosaic of European early modern human and Neanderthal features.
- 2000: Igor Ovchinnikov et al. retrieved mitochondrial DNA of a Neanderthal infant from Mezmaiskaya Cave in the Caucasus.
- 2005: The Max Planck Institute for Evolutionary Anthropology launched the Neanderthal genome project in co-operation with Connecticut-based 454 Life Sciences and the Institute of Quaternary Paleontology and Geology in Zagreb, Croatia. In 2009, the Max Planck Institute announced the "first draft" of a complete Neanderthal genome is completed.
- 2010: Comparison of Neanderthal genome with modern humans from Africa and Eurasia shows that 1–4% of modern non-African human genome might come from the Neanderthals.
- 2010: Discovery of Neanderthal tools far away from the influence of H. sapiens indicate that the species might have been able to create and evolve tools on its own, and therefore be more intelligent than previously thought. Furthermore, it was proposed that the Neanderthals might be more closely related to Homo sapiens than previously thought and that may in fact be a subspecies of it. Evidence has more recently emerged that these artifacts are probably of H. sapiens sapiens origin.
- 2012: Charcoal found next to six paintings of seals in Nerja caves, Malaga, Spain, has been dated to between 42,300 and 43,500 years old. The paintings themselves will be dated in 2013, and if their pigment matches the date of the charcoal, they would be the oldest known cave paintings. José Luis Sanchidrián at the University of Cordoba, Spain believes the paintings are more likely to have been painted by Neanderthals than early modern humans.
- 2013: A jawbone found in Italy had features intermediate between Neanderthals and Homo sapiens suggesting it could be a hybrid. The mitochondrial DNA is Neanderthal.
- 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: Researchers at the University of Colorado Museum in Boulder report that Neanderthals were not less intelligent than modern humans and "that single-factor explanations for the disappearance of the Neandertals are not warranted any more."
- 2014: Prof Thomas Higham of the University of Oxford performed the most comprehensive dating of Neanderthal bones and tools ever carried out, which demonstrated that Neanderthals died out in Europe between 41,000 and 39,000 years ago—this coincides with the start of a very cold period in Europe and is 5,000 years after Homo sapiens reached the continent.
Habitat and range
Early Neanderthals lived in the last glacial period for a span of about 100,000 years. Because of the damaging effects the glacial period had on the Neanderthal sites, not much is known about the early species. Countries where their remains are known include most of Europe south of the line of glaciation, roughly along the 50th parallel north. This includes most of Western Europe, Central Europe, the Carpathians, and the Balkans, some sites in Ukraine and in western Russia, Central and Northern Asia up to the Altai Mountains, and Western Asia from the Levant up to the Indus River. The Bontnewydd Palaeolithic site at Denbighshire, North Wales is the most north-western site of Neanderthal remains and one of the oldest remains in Britain (230,000 years ago). It is estimated that the total Neanderthal population across this habitat range numbered at around 70,000 at its peak.
Neanderthal fossils have not been found to date in Africa, but there have been finds close to North Africa, both on Gibraltar and in the Levant. At some Levantine sites, Neanderthal remains date from after the same sites were vacated by modern humans. Mammal fossils of the same time period show cold-adapted animals were present alongside these Neanderthals in this region of the Eastern Mediterranean. This implies Neanderthals were better adapted biologically to cold weather than modern humans and at times displaced them in parts of the Middle East when the climate got cold enough. However this has been disputed recently through studies of their cranial morphology and sinuses. Some scholars have also posited that Neanderthals could be an Asian species which expanded into Europe, making Neanderthals a tropical species rather than cold adaptive.
Homo sapiens appears to have been the only human type in the Nile River Valley during these periods, and Neanderthals are not known to have ever lived south-west of present-day Israel. When climate change caused warmer temperatures, the Neanderthal range likewise retreated to the north, along with the cold-adapted species of mammals. Apparently these weather-induced population shifts took place before modern people secured competitive advantages over the Neanderthal, as these shifts in range took place well over ten thousand years before modern people totally replaced the Neanderthal, despite the recent evidence of some successful interbreeding.
Separate developments in the human line, in other regions such as Southern Africa, somewhat resembled the Eurasian Neanderthals, but these people were not Neanderthals. One such example is Rhodesian Man (Homo rhodesiensis), who existed long before any classic Eurasian Neanderthals, but had a more modern set of teeth. Some H. rhodesiensis populations appeared to be on the road to modern H. sapiens sapiens. At any rate, the populations in Eurasia underwent increasing "Neanderthalization" over time. Some have suggested that H. rhodesiensis in general was ancestral to both modern humans and Neanderthals, and that at some point the two populations went their separate ways, but this argument presupposes that H. rhodesiensis existed around 600,000 years ago.
To date, no intimate connection has been found between these similar archaic people and the Eurasian Neanderthals, at least during the same time as H. rhodesiensis seems to have lived about 600,000 years ago, long before the time of classic Neanderthals. This said, some researchers think that H. rhodesiensis may have lived much later than this period, depending on the method used to date the fossils, so this question remains open to debate. Some H. rhodesiensis features, like the large brow ridge, may have been caused by convergent evolution.
It appears incorrect, based on present research and known fossil finds, to refer to any fossils outside Eurasia as true Neanderthals. They had a known range that possibly extended as far east as the Altai Mountains, but not farther to the east or south, and apparently not into Africa. At any rate, in North-East Africa the land immediately south of the Neanderthal range was possessed by modern humans Homo sapiens idaltu or Homo sapiens, since at least 160,000 years before the present. 160,000-year-old hominid fossils at Jebel Irhoud in Morocco were previously thought to be Neanderthal, but it is now clear that they are early modern humans.
Classic Neanderthal fossils have been found over a large area, from northern Germany to Israel and Mediterranean countries like Spain and Italy in the south and from England and Portugal in the west to Uzbekistan in the east. This area probably was not occupied all at the same time. The northern border of their range, in particular, would have contracted frequently with the onset of cold periods. On the other hand, the northern border of their range as represented by fossils may not be the real northern border of the area they occupied, since Middle Palaeolithic-looking artifacts have been found even farther north, up to 60° N, on the Russian plain. Recent evidence has extended the Neanderthal range by about 1,250 miles (2,010 km) east into southern Siberia's Altai Mountains.
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, which was much larger in both length and width, 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 164–168 cm (65–66 in) and females 152–156 cm (60–61 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 Homo Sapiens. 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 mother’s fur as infants. Modern humans don’t have these ridges, but Neanderthals do." (Cochran, G. & H. Harpending (2009).[full citation needed]
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 ."[full citation needed]
Neanderthals are known for their large cranial capacity, which at 1600 cm3 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.
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Neanderthals made advanced tools, apparently kindled fire on demand, may have had a language (the nature of which is debated and likely unknowable), and lived in complex social groups. The Molodova I archaeological site in eastern Ukraine suggests some Neanderthals built dwellings using animal bones. A building was made of mammoth skulls, jaws, tusks and leg bones, and had 25 hearths inside.
Circumstantial evidence suggests Neanderthals may have been building some form of watercraft since the Middle Paleolithic. Scientists have speculated that these watercraft may have been similar to dugout canoes, which are among the oldest known boats in the archaeological record. Mousterian stone tools discovered on the southern Ionian Islands suggests that Neanderthals were sailing the Mediterranean Sea as early as 110,000 years ago. Quartz hand-axes, three-sided picks, and stone cleavers from Crete have also been recovered that date back about 170,000 years ago.
It was once thought that Neanderthals lacked the sophistication for hunting, perhaps scavenging meat from carcasses, but increasing evidence suggests they were apex predators, capable of bringing down a wide range of prey from red deer, reindeer, ibex and wild boar, to larger animals such as aurochs and even, on occasion, mammoth, straight-tusked elephant and rhinoceros. However, while they were largely carnivorous, new studies indicate Neanderthals also had cooked vegetables in their diet. In 2010, an isotope analysis of Neanderthal teeth found traces of cooked vegetable matter, and more recently a 2014 study of Neanderthal coprolites (fossilized feces) found substantial amounts of plant matter, contradicting the earlier belief they were exclusively (or almost exclusively) carnivorous. A dental analysis of specimens from Spy, Belgium and El Sidrón, Spain by an international team, published in 2017 in Nature revealed, that these Neanderthals "had a wide-ranging diet", even claiming the "Neanderthals from El Sidrón showed no evidence of meat eating" at all while living 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 their Homo sapiens contemporaries. Some experts suggest that this disparity alone was a major contributing factor to their ultimate replacement by Homo sapiens, which may have outnumbered them by as much as 9 to 1 according to some estimates. Their lower population density may have also increased Neanderthal susceptibility to mutations caused by inbreeding.
The bones of twelve Neanderthals were discovered at El Sidrón cave in the Atapuerca Mountains of Spain. They are believed to be a social group which was massacred about 50,000 years ago, and analysis of the mtDNA shows 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, and as their bones also show signs that they suffered from food shortage, they may have been victims of "survival cannibalism" by another Neanderthal group. 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. Researchers have taken this as evidence of the presence of interpersonal violence among the Neanderthals.
In 2016, Jacques Jaubert and colleagues reported the discovery of 176,000 year old stalagmite constructions 336 meters deep in Bruniquel Cave near Bruniquel in south-western France. There were two annular (ring) structures, one 6.7 by 4.5 metres, and the other 2.2 by 2.1, with one to four layers of aligned broken stalagmites, and short pieces within the layers to support them. There were also four stack structures, two within the large ring and two outside it. Artificial lighting would have been required as the cave is far beyond the reach of daylight, and 57 of the stalagmite pieces are reddened and 66 blackened by fire. Burnt organic material was also found. The researchers concluded that the constructions must have been made by early Neanderthals, as they were the only human species in the area at the time. The elaborate structures show that the Neanderthals had a more sophisticated level of social organisation than previously thought possible.
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.
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 species shared a common ancestor about 500,000 years ago. An article appearing in the journal Nature has calculated the species 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 16 November 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 species 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 species 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.
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 unknown species, tantalizing hints of unknown groups from Asia and Africa that left genes in Denisovans and modern humans, respectively.  17% of the genes of Denisovians and Neanderthals are shared. According to a research led by an Oxford scientist, it is possible that the gene causing red hair in modern humans is inherited from the Neanderthals, which is up to 100,000 years old, although a different mutation on Neanderthals was found in the MRC1 gene than on modern humans.
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 species 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 species originating from a 500,000-year-old common ancestor. 
More recent research suggests that Neanderthal–Homo 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."
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 "[H]umans have been constantly migrating throughout their history - this makes it hard to say exactly where interactions with Neanderthals occurred. It's possible, for example, that all of the 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 Homo sapiens and Neanderthal Y chromosomes that, they postulated, could cause female Homo 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 ~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.
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] and Homo sapiens 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 has been 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
As Paul Jordan notes: "A natural sympathy for the underdog and the disadvantaged lends a sad poignancy to the fate of the Neanderthal folk, however it came about." Jordan, though, does say that there was perhaps interbreeding to some extent, but that populations that remained totally Neanderthal were probably out-competed and marginalized to extinction by the Aurignacians.
About 55,000 years ago, the weather began to fluctuate wildly from extreme cold conditions to mild cold and back in a matter of a few decades. Neanderthal bodies were well suited for survival in a cold climate—their barrel chests and stocky limbs stored body heat better than the Cro-Magnons. However, the rapid fluctuations of weather caused ecological changes to which the Neanderthals could not adapt; familiar plants and animals would be replaced by completely different ones within a lifetime. Neanderthals' ambush techniques would have failed as grasslands replaced trees. Neanderthals died out in Europe between 41,000 and 39,000 years ago which coincides 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 region 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 were restricted to more local resources. Furthermore, excavations at Ortvale Klde Rockshelter discovered that there was a clear break between the Late Middle Paleolithic and the Early Upper Paleolithic lithic assemblages, which were attributed to Neanderthals and modern humans respectively. This would suggest that modern humans came in and replaced Neanderthals, rather than a slow shift or integration occurring in this region. 
Studies on Neanderthal body structures have shown that they needed more energy to survive than any other species of hominid. Their energy needs were up to 100 to 350 kcal (420 to 1,460 kJ) more per day comparing to projected anatomically modern human males weighing 68.5 kg (151 lb) and females 59.2 kg (131 lb). When food became scarce, this difference may have played a major role in the Neanderthals' extinction.
Coexistence with Homo sapiens
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. 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. H. sapiens 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.
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.
- 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, with evidence of healing. For him to have lived on suggests that someone might have had to process his food for him. This is one possible example of Neanderthal altruism (see also Shanidar I).
- 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: A fossilized skull, discovered in 1909, at the archaeological site in Peyzac-le-Moustier, Dordogne, France. The Mousterian tool culture is named after Le Moustier. The skull, estimated to be less than 45,000 years old, includes a large nasal cavity and a somewhat less developed brow ridge and occipital bun as might be expected in a juvenile.
- Shanidar Cave: Found in the Zagros Mountains in Iraqi Kurdistan; a total of nine skeletons found believed to have lived in the Middle Paleolithic. One of the nine remains was missing part of its right arm, which is theorized to have been broken off or amputated. The find is also significant because it shows that stone tools were present among this tribe's culture. One of the skeletons was originally thought to have been buried with flowers, signifying that some type of burial ceremony may have occurred. This is no longer considered to be the case, and Paul B. Pettitt has stated that the "deliberate placement of flowers has now been convincingly eliminated", noting that "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: Fossilized remains of an adult Neanderthal, dated to roughly 45,000 years ago, and one of several found in a cave at Nahal Amud, Israel, at least some of which may have been deliberately buried. A particularly notable feature of this find is its cranial capacity, which, at 1,740 cm3, is among the largest known for any hominid, living or extinct.
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
Typical H. neanderthalensis traits
- 135–45 ka: Krapina, Saccopastore skulls, Malarnaud, Altamura, Gánovce, Denisova, Okladnikov, Pech de l'Azé, Tabun 120–±5 ka, 100 Qafzeh9 100, 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 ±8 ka, 53 Teshik-Tash.
- 45–35 ka: Le Moustier 45 ka, Feldhofer 42 ka, La Quina, l'Horus, Hortus, Kulna, Šipka, Saint Césaire, Bacho Kiro, El Castillo, Bañolas, Arcy-sur-Cure.
- < 35 ka: Châtelperron, Figueira Brava, Zafarraya 30 ka, Vogelherd 35 to 32 ka Vindija ±800 14C B.P. 32,400 (Vi-208 ±220, Vi-207 31,390±1,800 14C B.P.), 32,400 Velika Pećina,
Homo sapiens with some neanderthal-like archaic traits
Neanderthals have been portrayed in popular culture including appearances in literature, visual media and comedy, often in an unflattering and inaccurate light.
Early artistic reconstructions mostly presented Neanderthals as beastly creatures, emphasizing hairiness and rough, dark complexion.
- Abrigo do Lagar Velho—More about "the Lapedo child"
- Altamura Man
- Biological anthropology
- Bruniquel Cave
- Denisova hominin
- Early human migrations
- Engis 2
- Life timeline
- Max Planck Institute for Evolutionary Anthropology
- Neanderthal 1
- Neanderthal genome project
- Neanderthal Museum
- Neanderthals of Gibraltar
- Pleistocene megafauna
- Prehistoric Autopsy (2012 BBC documentary film)
- Roca dels Bous (archaeological site)
- Species problem
- Neanderthals in popular culture
- List of fossil sites (with link directory)
- List of human evolution fossils (with images)
- List of Neanderthal sites
- 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 (litt. "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").
- Tal after the German spelling reform of 1901, whence the German name Neandertal for both the valley and species.
- 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 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.
- Dictionary of Anthropology—Charles Winick - Google Books. Books.google.ca (1956-12-18). Retrieved on 2014-05-24.
- Bibliography of Fossil Vertebrates 1954-1958 - C.L. Camp, H.J. Allison, and R.H. Nichols - Google Books. Books.google.ca. Retrieved on 2014-05-24.
- "Neanderthal". Wiley-Blackwell Encyclopedia of Human Evolution. Chichester, West Sussex: Wiley-Blackwell. 2013.
- 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 1 February 2017.
- "Neanderthal in ODE". Oxford Dictionaries.
- ""Neanderthal" in Random House Dictionary (US) & Collins Dictionary (UK)". Dictionary.com.
- 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. doi:10.1073/pnas.0904119106. JSTOR 40485013. PMC . PMID 19805257.
- 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. doi:10.1073/pnas.0308085100. PMC . PMID 14745010.
- "Scientists Identify Neanderthal Genes in Modern Human DNA". Sci-News.com. January 30, 2014. Retrieved October 29, 2015.
- Shaw, Ian; Jameson, Robert, eds. (1999). A Dictionary of Archaeology. Blackwell. p. 408. ISBN 0-631-17423-0. Retrieved 1 August 2016.
- Finlayson, C; Pacheco, FG; Rodríguez-Vidal, J; Fa, DA; Gutierrez López, JM; Santiago Pérez, A; Finlayson, G; Allue, E; Baena Preysler, J; Cáceres, I; Carrión, JS; Fernández Jalvo, Y; Gleed-Owen, CP; Jimenez Espejo, FJ; López, P; López Sáez, JA; Riquelme Cantal, JA; Sánchez Marco, A; Guzman, FG; Brown, K; Fuentes, N; Valarino, CA; Villalpando, A; Stringer, CB; Martinez Ruiz, F; Sakamoto, T (October 2006). "Late survival of Neanderthals at the southernmost extreme of Europe". Nature. 443 (7113): 850–3. Bibcode:2006Natur.443..850F. doi:10.1038/nature05195. ISSN 0028-0836. PMID 16971951.
- Outside Europe, Mousterian tools were made by both Neanderthals and early modern Homo sapiens. (Donald Johanson & Blake Edgar (2006) From Lucy to Language, Simon & Schuster, p. 272
- 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. (Aug 2015). "Virtual Reconstruction of the Neanderthal Amud 1 Cranium". American Journal of Physical Anthropology. 158: 185–197. doi:10.1002/ajpa.22777.
- 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. doi:10.1371/journal.pone.0047765. PMC . PMID 23082212. Retrieved May 29, 2016.
- Rincon, Paul (2010-05-06). "Neanderthal genes 'survive in us'". BBC News. Retrieved 2010-05-07.
- 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 (23 October 2014). "Genome sequence of a 45,000-year-old modern human from western Siberia". Nature. 514 (7523): 445–449. doi:10.1038/nature13810. PMC . PMID 25341783.
- 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. 17 February 2016. Retrieved 2016-03-06.
- The Combined Landscape of Denisovan and Neanderthal Ancestry in Present-Day Humans, Current Biology, Sankararaman et al., 26, 1241–1247, 2016
- Stringer, C.; Gamble, C. (1993). In Search of the Neanderthals. London: Thames and Hudson. pp. 158–160. ISBN 978-0500050705.
- 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.
- 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.
- Gargett, R.H. (1989). "Grave Shortcomings: The Evidence for Neandertal Burial". Current Anthropology. 30 (2): 157–190. doi:10.1086/203725.
- 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 (Jan 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.
- 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.
- "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.
- Prüfer, Kay; et al. (2014). "The complete genome sequence of a Neanderthal from the Altai Mountains". Nature. 505 (7481): 43–49. doi:10.1038/nature12886. PMC . PMID 24352235.
- 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) [online 25 May 2016]. "Early Neanderthal Constructions deep in Bruniquel Cave in Southwestern France". Nature. 534 (7605): 111–114. doi:10.1038/nature18291. ISSN 0028-0836.
- 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)
- Howell, F. Clark (1957). "The evolutionary significance of variation and varieties of 'Neanderthal' man". The Quarterly Review of Biology. 32 (4): 330–47. doi:10.1086/401978. JSTOR 2816956. PMID 13506025.
- 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.
- "Neanderthal adjective—definition in British English Dictionary & Thesaurus—Cambridge Dictionary Online". Dictionary.cambridge.org. 2013-01-08. Retrieved 2013-01-22.
- "Oxford Learner's Dictionaries—Find pronunciation, clear meanings and definitions of words at OxfordLearnersDictionaries.com".[dead link]
- "Neanderthal | Define Neanderthal at Dictionary.com". Dictionary.reference.com. Retrieved 2013-01-22.
- Kurtén, Björn (10 October 1995). Dance of the Tiger: A Novel of the Ice Age. University of California Press. pp. xxi. ISBN 0-520-20277-5. Retrieved 9 May 2012.
- Pollet, Carl J. (September 21, 1991). "…And Etymology". Science News. 140 (12): 191. doi:10.2307/3975867. JSTOR 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. doi:10.1073/pnas.96.13.7117. JSTOR 48019. PMC . PMID 10377375.
- 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. doi:10.1073/pnas.96.13.7604. JSTOR 48106. PMC . PMID 10377462.
- "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. doi:10.1002/(SICI)1096-8644(200005)112:1<129::AID-AJPA11>3.0.CO;2-K. PMID 10766948.
- Hedges SB (December 2000). "Human evolution. A start for population genomics". Nature. 408 (6813): 652–3. doi:10.1038/35047193. PMID 11130051.
- "Modern humans, Neanderthals shared earth for 1,000 years". ABC News (Australia). 1 September 2005. Archived from the original on January 15, 2006. Retrieved 19 September 2006.
- 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.
- Papagianni, Dmitra; Morse, Michael (2013). The Neanderthals Rediscovered. Thames & Hudson. ISBN 978-0-500-05177-1.
- Higham, T.; et al. (2014). "The timing and spatiotemporal patterning of Neanderthal disappearance". Nature. 512 (7514): 306–309. doi:10.1038/nature13621. PMID 25143113.
- "Homo heidelbergensis: Evolutionary Tree". Smithsonian National Museum of Natural History. Retrieved 17 March 2013.
- Stringer, Chris. "The Ancient Human Occupation of Britain" (PDF). Natural History Museum, London. Archived from the original (PDF) on March 2, 2013. Retrieved 17 March 2013.
- 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.
- "Homo neanderthalensis". Smithsonian Institution. Archived from the original on May 21, 2009. Retrieved 18 May 2009.
- "New Evidence On The Role Of Climate In Neanderthal Extinction". Science Daily.
- "Neanderthal - Homo neanderthalensis - Details - Key Fossils - La Chapelle-aux-Saints". Encyclopedia of Life. Retrieved July 18, 2016.
- "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.
- 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. doi:10.1016/S0092-8674(00)80310-4. ISSN 0092-8674. PMID 9230299.
- 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. doi:10.1073/pnas.192464099. PMC . PMID 12232049.
- 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. doi:10.1073/pnas.96.13.7604. PMC . PMID 10377462. Retrieved 2009-06-21.
- 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. doi:10.1038/35006625. PMID 10761915.
- Green, RE; Krause, J; Ptak, SE; et al. (November 16, 2006). "Analysis of one million base pairs of Neanderthal DNA". Nature. 444: 330–6. doi:10.1038/nature05336. PMID 17108958. Retrieved July 18, 2016.
- Morgan, James (12 February 2009). "Neanderthals 'distinct from us'". BBC News. Retrieved 22 May 2009.
- 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. doi:10.1126/science.1188021. PMID 20448178.
- "Neanderthals More Intelligent Than Thought". September 22, 2010. Archived from the original on September 25, 2010. Retrieved September 27, 2010.
- Benazzi, S.; Douka, K.; Fornai, C.; Bauer, C. C.; Kullmer, O.; Svoboda, J. Í.; Pap, I.; Mallegni, F.; Bayle, P.; Coquerelle, M.; Condemi, S.; Ronchitelli, A.; Harvati, K.; Weber, G. W. (2011). "Early dispersal of modern humans in Europe and implications for Neanderthal behaviour". Nature. 479 (7374): 525–528. doi:10.1038/Nature10617. PMID 22048311.
- Fergal MacErlean (10 February 2012). "First Neanderthal cave paintings discovered in Spain". New Scientist. Retrieved 10 February 2012.
- Viegas, Jennifer (27 March 2013). "First Love Child of Human, Neanderthal Found". Discovery News. Retrieved 11 April 2013.
- "Neanderthals were not less intelligent than modern humans, scientists find". April 30, 2014. Retrieved April 30, 2014.
- "BBC News—New dates rewrite Neanderthal story". BBC News.
- "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. 11 February 2008. Retrieved 18 May 2009.
- O'Neill, Dennis. "Evolution of Modern Humans: Neanderthals", Palomar College, June 10, 2011, accessed August 21, 2011.
- Jordan, P. (2001) Neanderthal: Neanderthal Man and the Story of Human Origins. The History Press ISBN 978-0-7509-2676-8.
- Bilsborough, A, Rae, TC (2015) [REVISION OF] Hominoid cranial diversity and adaptation. In: Henke, W., Rothe, H. & Tattersall, I. (eds.) Handbook of Palaeoanthropology, New York: Springer
- "Fieldwork—Jebel Irhoud". Max Planck Institute, Department of Human Evolution.
- Arsuaga, J.L; Gracia, A; Martínez, I; Bermúdez de Castro, J.M; Rosas, A; Villaverde, V; Fumanal, M.P (1989). "The human remains from Cova Negra (Valencia, Spain) and their place in European Pleistocene human evolution". Journal of Human Evolution. 19: 55–92. doi:10.1016/0047-2484(89)90023-7.
- Mallegni, F., Piperno, M., and Segre, A (1987). "Human remains of Homo sapiens neanderthalensis from the Pleistocene deposit of Sants Croce Cave, Bisceglie (Apulia), Italy". American Journal of Physical Anthropology. 72 (4): 421–429. doi:10.1002/ajpa.1330720402. PMID 3111268.
- 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. doi:10.1016/j.jhevol.2004.05.002. PMID 15288521.
- Wade, Nicholas (2 October 2007). "Fossil DNA Expands Neanderthal Range". The New York Times. Retrieved 18 May 2009.
- Ravilious, Kate (1 October 2007). "Neandertals Ranged Much Farther East Than Thought". National Geographic Society. Retrieved 18 May 2009.
- L’Homme de Neanderthal par Paul Dardé : L’Homme Primitif https://www.academia.edu/11187487/L_Homme_de_Neanderthal_par_Paul_Dard%C3%A9_L_Homme_Primitif
- "Science & Nature—Wildfacts—Neanderthal". BBC. Archived from the original on January 15, 2009. Retrieved 2009-06-21.
- "Neanderthal". BBC. Archived from the original on January 15, 2009. Retrieved 18 May 2009.
- Froehle, Andrew W; Churchill, Steven E (2009). "Energetic Competition Between Neandertals and Anatomically Modern Humans" (PDF). PaleoAnthropology: 96–116. Retrieved 2011-10-31.
- Laleuza-Fox, Carles; Römpler, Holger; et al. (2007-10-25). "A Melanocortin 1 Receptor Allele Suggests Varying Pigmentation Among Neanderthals". Science. 318 (5855): 1453–5. Bibcode:2007Sci...318.1453L. doi:10.1126/science.1147417. PMID 17962522.; see also Rincon, Paul (25 October 2007). "Neanderthals 'were flame-haired'". BBC News. Retrieved 25 October 2007.
- "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. 2008-09-09. Retrieved 2009-09-19.
- 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".
- Moskvitch, Katia (2010-09-24). "Neanderthals were able to 'develop their own tools'". BBC News. BBC. Retrieved 2010-10-01.
- Heyes, Peter; Anastasakis, Konstantinos; de Jong, Wiebren (2016). "Selection and Use of Manganese Dioxide by Neanderthals". Scientific Reports. 6: 22159. doi:10.1038/srep22159. ISSN 2045-2322.
- "Neanderthals could speak like modern humans, study suggests". BBC. December 20, 2013. Retrieved July 19, 2016.
- "Language Abilities in Neanderthals". Annual Review of Linguistics. July 10, 2014. Retrieved July 19, 2016.
- "Neanderthals: Facts About Our Extinct Human Relatives - Social structure". LiveScience. April 13, 2016. Retrieved July 19, 2016.
- Gray, Richard (December 18, 2011). "Neanderthals built homes with mammoth bones". Telegraph.co.uk.
- Demay, Laëtitia; Péan, Stéphane; Patou-Mathis, Marylène (2012). "Mammoths used as food and building resources by Neanderthals: Zooarchaeological study applied to layer 4, Molodova I (Ukraine)". Quaternary International. 276–277: 212–226. doi:10.1016/j.quaint.2011.11.019. ISSN 1040-6182.
- Yirka, Bob (1 March 2012). "Evidence suggests Neanderthals took to boats before modern humans". phys.org. Retrieved 5 May 2016.
- Marshall, Michael (29 February 2012). "Neanderthals were ancient mariners". New Scientist. Retrieved 5 May 2016.
- Choi, Charles Q. (15 November 2012). "Ancient Mariners: Did Neanderthals Sail to Mediterranean?". LiveScience. Retrieved 5 May 2016.
- "Neanderthals May Have Sailed to Crete". DNews. 13 December 2012. Retrieved 5 May 2016.
- 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. PMID 15869783.
- 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.
- 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. doi:10.1073/pnas.120178997. JSTOR 122870. PMC . PMID 10852955.
- 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. doi:10.1371/journal.pone.0014769. PMC . PMID 21445243.
- 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. doi:10.1073/pnas.1016868108.
- Webb, Jonathan (25 June 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.
- 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. doi:10.1073/pnas.082111899. PMC . PMID 11972028.
- Brown, Cynthia Stokes. Big History. New York, NY: The New Press, 2008. Print.
- Moulson, Geir; Associated Press (20 July 2006). "Neanderthal genome project launches". MSNBC. Retrieved 22 August 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. doi:10.1038/nature05336. PMID 17108958.
- Wade, Nicholas (15 November 2006). "New Machine Sheds Light on DNA of Neanderthals". The New York Times. Retrieved 18 May 2009.
- Pennisi, E. (May 2007). "Ancient DNA. No sex please, we're Neandertals". Science. 316 (5827): 967. doi:10.1126/science.316.5827.967a. PMID 17510332.
- "Neanderthal bone gives DNA clues". CNN. Associated Press. 16 November 2006. Archived from the original on 18 November 2006. Retrieved 18 May 2009.
- Than, Ker; LiveScience (15 November 2006). "Scientists decode Neanderthal genes". MSNBC. Retrieved 18 May 2009.
- "Neanderthal Genome Sequencing Yields Surprising Results And Opens A New Door To Future Studies" (Press release). Lawrence Berkeley National Laboratory. 16 November 2006. Retrieved 31 May 2009.
- Hayes, Jacqui (15 November 2006). "DNA find deepens Neanderthal mystery". Cosmos. Archived from the original on March 26, 2009. Retrieved 18 May 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. doi:10.1016/j.cell.2008.06.021. ISSN 0092-8674. PMC . PMID 18692465.
- Elizabeth Pennisi (2009). "Tales of a Prehistoric Human Genome". Science. 323 (5916): 866–871. doi:10.1126/science.323.5916.866. PMID 19213888.
- 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. doi:10.1038/emboj.2009.222. PMC . PMID 19661919.
- "New studies reveal 20 Percent of Neanderthal genome lives on in modern humans". Retrieved 7 October 2016.
- "Surprise! 20 Percent of Neanderthal Genome Lives On in Modern Humans, Scientists Find". National Geographic. Retrieved 7 October 2016.
- "The ginger gene revealed".
- "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 (30 November 2007). "A Melanocortin 1 Receptor Allele Suggests Varying Pigmentation Among Neanderthals". Science. 318 (5855): 1453–1455. doi:10.1126/science.1147417. PMID 17962522 – via science.sciencemag.org.
- Neanderthals did not interbreed with humans, scientists find Archived April 19, 2014, at the Wayback Machine.. Telegraph. Retrieved on 2014-05-24.
- Press Association (4 February 2013). "Neanderthals 'unlikely to have interbred with human ancestors'". The Guardian. Retrieved 24 May 2014.
- 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. doi:10.1016/j.gene.2013.06.005. ISSN 0378-1119. PMID 23872234.
- Neanderthal breeding idea doubted. BBC. Retrieved on 2016-03-02.
- "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. doi:10.1016/j.ajhg.2014.12.029. PMC . PMID 25683122.
- . The Daily Mail. Retrieved 2016.01.26.
- Zimmer, Carl (17 March 2016). "Humans Interbred With Hominins on Multiple Occasions, Study Finds". The New York Times. Retrieved 17 March 2016.
- Mendez, Fernando L.; et al. (7 April 2016). "The Divergence of Neandertal and Modern Human Y Chromosomes" (PDF). The American Journal of Human Genetics. 98 (4): 728–734. doi:10.1016/j.ajhg.2016.02.023. Retrieved 12 April 2016.
- "DNA points to Neanderthal breeding barrier". BBC News, Science & Environment. 8 April 2016. Retrieved 12 April 2016.
- Sample, Ian (2016-02-17). "Oldest known case of Neanderthal-human sex revealed by DNA test". the Guardian. Retrieved 2016-05-10.
- Zimmer, Carl (March 17, 2016). "Humans Interbred With Hominins on Multiple Occasions, Study Finds". New York Times. Retrieved 5 May 2016.
- 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. doi:10.1126/science.1250368. PMID 24786081.
- Higham, Tom et al. (21 August 2014). "The timing and spatiotemporal patterning of Neanderthal disappearance". Nature. 512 (512): 306–309. doi:10.1038/nature13621. PMID 25143113. (subscription required)
- Stein, Richard A (1 October 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)
- 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. doi:10.1016/j.tree.2007.02.001. PMID 17300854.
- "First genocide of human beings occurred 30,000 years ago". Pravda. 24 October 2007. Retrieved 18 May 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. doi:10.1371/journal.pbio.0020421. PMC . PMID 15562317.
- Wong, Kate (1 August 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.
- Froehle, Andrew W.; Churchill, Steven E. (2009). "Energetic Competition Between Neandertals and Anatomically Modern Humans" (PDF). PaleoAnthropology: 96–116.
- 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. 22 Oct. 2014.
- McKie, Robin (1 March 2015). "How hunting with wolves helped humans outsmart the Neanderthals". The Guardian. Retrieved 5 May 2016.
- Stringer, Chris (2012). "Evolution: What makes a modern human". Nature. 485 (7396): 33–5. Bibcode:2012Natur.485...33S. doi:10.1038/485033a. PMID 22552077.
- http://img.kb.dk/tidsskriftdk/pdf/gto/gto_0019-PDF/gto_0019_67206.pdf[full citation needed]
- Dan Jones: The Neanderthal within., New Scientist 193.2007, H. 2593 (3 March), 28–32. Modern Humans, Neanderthals May Have Interbred[dead link]; Humans and Neanderthals interbred Archived February 22, 2009, at the Wayback Machine.
- ; [https://www.theguardian.com/science/story/0,,1871842,00.html[full citation needed]
- Not a lasting last for the Neandertals—John Hawks weblog, September 13, 2006[full citation needed]
- 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. doi:10.1073/pnas.0608443103. JSTOR 30052409. PMC . PMID 17085588.
- 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. doi:10.1093/molbev/msr024. PMID 21266489.
- Viegas, Jennifer (18 July 2011). "All Non-Africans Part Neanderthal, Genetics Confirm". DNews.
- Neandertal ancestry "Iced" - John Hawks weblog, August 15, 2012[full citation needed]
- "Resurrecting Surviving Neandertal Lineages from Modern Human Genomes". Science. 2014-01-29. Retrieved 2014-02-02.
- 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. arXiv: . Bibcode:2013arXiv1307.8263L.
- Jha, Alok (14 August 2012). "Study casts doubt on human-Neanderthal interbreeding theory". The Guardian. Retrieved 19 February 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 (27 March 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. doi:10.1371/journal.pone.0059781. PMC . PMID 23544098.
- The Neanderthal Dead, exploring mortuary variability in middle paleolithic eurasia. Paul B. Pettitt (2002)
- "Homo neanderthalensis–The Neanderthals". Australian Museum. Retrieved 26 June 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. doi:10.1006/jhev.1997.0133. PMID 9300343.
- Kreger, C. David. "Homo neanderthalensis". ArchaeologyInfo.com. Retrieved 16 May 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. doi:10.1038/363252a0. ISSN 0028-0836. PMID 8387643.
- Rink, W. Jack; Schwarcz, H.P.; Lee, H.K.; Rees-Jones, J.; Rabinovich, R.; Hovers, E. (August 2002). "Electron spin resonance (ESR) and thermal ionization mass spectrometric (TIMS) 230Th/234U dating of teeth in Middle Paleolithic layers at Amud Cave, Israel". Geoarchaeology. 16 (6): 701–717. 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.
- Rincon, Paul (13 September 2006). "Neanderthals' 'last rock refuge'". BBC News. Retrieved 18 May 2009.
- "Hohlenstein - Hohlenstein 4". NESPOS Space. Retrieved July 24, 2016.
- "Lonetal Sites, including Neanderthal and Middle Paleolithic sites in the Swabian Alb near the city of Ulm - View of Vogelherd about 35 000 years BP...The stratigraphy reveals deposits from [...] Mousterian...". Don's Maps. Retrieved July 24, 2016.
- Conard, Nj; Grootes, Pm; Smith, Fh (July 2004). "Unexpectedly recent dates for human remains from Vogelherd". Nature. 430 (6996): 198–201. Bibcode:2004Natur.430..198C. doi:10.1038/nature02690. ISSN 0028-0836. PMID 15241412.
- Higham T, Ramsey CB, Karavanić I, Smith FH, Trinkaus E (January 2006). "Revised direct radiocarbon dating of the Vindija G1 Upper Paleolithic Neandertals". Proceedings of the National Academy of Sciences. 103 (3): 553–7. Bibcode:2006PNAS..103..553H. doi:10.1073/pnas.0510005103. PMC . PMID 16407102.
- Hayes, Jacqui (2 November 2006). "Humans and Neanderthals interbred". Cosmos. Archived from the original on February 22, 2009. Retrieved 17 May 2009.
- Neanderthal image by Kupka, based on Boule, 1909, in Humanity's Journeys Dr. Kathryn Denning, 2005, retrieved 2012-03-17
- Boë, Louis-Jean; Heim, Jean-Louis; Honda, Kiyoshi; Maeda, Shinji (2002). "The potential Neandertal vowel space was as large as that of modern humans". Journal of Phonetics. 30 (3): 465–84. doi: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 mtDNA Contribution to Early Modern Humans". PLoS Biology. 2 (3): e57. doi:10.1371/journal.pbio.0020057. PMC . PMID 15024415.
- Wild, Eva M.; Teschler-Nicola, Maria; Kutschera, Walter; Steier, Peter; Trinkaus, Erik; Wanek, Wolfgang (2005). "Direct dating of Early Upper Palaeolithic human remains from Mladeč". Nature. 435 (7040): 332–5. Bibcode:2005Natur.435..332W. doi:10.1038/nature03585. PMID 15902255.
- Zilhão, João; Davis, Simon J. M.; Duarte, Cidália; Soares, António M. M.; Steier, Peter; Wild, Eva (2010). Hawks, John, ed. "Pego do Diabo (Loures, Portugal): Dating the Emergence of Anatomical Modernity in Westernmost Eurasia". PLoS ONE. 5 (1): e8880. doi:10.1371/journal.pone.0008880. PMC . PMID 20111705. 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, Swapan; Dannemann, Michael; Prüfer, Kay; Kelso, Janet; Patterson, Nick; Reich, David (2014). "The genomic landscape of Neanderthal ancestry in present-day humans". Nature. 507 (7492): 354–357. doi:10.1038/nature12961.
- Vattathil, S.; Akey, J.M. (2015). "Small amounts of archaic admixture provide big insights into human history". Cell. 163 (2): 281–284. doi:10.1016/j.cell.2015.09.042. PMID 26451479.
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- Kreger, C. David (30 June 2000). "Homo neanderthalensis". ArchaeologyInfo.com. Retrieved 23 May 2009.
- O'Neil, Dennis (12 May 2009). "Evolution of Modern Humans: Neandertals". Retrieved 23 May 2009.
- "Homo neanderthalensis". The Smithsonian Institution.
- "Neanderthal DNA". International Society of Genetic Genealogy. Archived from the original on June 17, 2006.: Includes Neanderthal mtDNA sequences
- Panoramio—'IMG_6922 The Neandertal foot prints' (photo of ~25K years old fossilized footprints discovered in 1970 on volcanic layers near Demirkopru Dam Reservoir, Manisa, Turkey)
- Did better mothering defeat the Neanderthals?
- My Great-great-great Grandfather's a Neanderthal
- Ancient tryst fortified human immune system
- Neanderthal-human hybridisation hypothesis
- Neanderthal hybridization and Haldane's rule
- Hominin interbreeding and the evolution of human variation – article by Kwang Hyun Ko, published online 16 July 2016
- Neanderthal Studies Professional Online Service (NESPOS) fossil overview
- Human Timeline (Interactive) – Smithsonian, National Museum of Natural History (August 2016).