Neanderthal

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For other uses, see Neanderthal (disambiguation).
iNeanderthals
H. neanderthalensis La Ferrassie 1
H. neanderthalensis La Ferrassie 1
Conservation status
Prehistoric
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Family: Hominidae
Genus: Homo
Species: H. neanderthalensis
Binomial name
Homo neanderthalensis
King, 1864
Synonyms

Palaeoanthropus neanderthalensis
H. s. neanderthalensis

The Neanderthal (IPA pronunciation: [niː.ændə(ɚ)θɒl, neɪ.ændə(ɚ)θɒl, niː.ændə(ɚ)tɒl, neɪ.ændə(ɚ)tɒl,]), (Homo neanderthalensis) or Neandertal was a species of the Homo genus that inhabited Europe and parts of western Asia. The first proto-Neanderthal traits appear in Europe as early as 350,000 years ago.[1] By 130,000 years ago, full blown Neanderthal characteristics had appeared and by 50,000 years ago, Neanderthals disappeared from Asia, although they did not reach extinction in Europe until 33,000 to 24,000 years ago, perhaps 15,000 years after Homo sapiens had migrated into Europe.[2][3][4]

Neanderthals had many adaptations to a cold climate, such as large braincase, short but robust builds, and large noses — traits selected by nature in cold climates. Their brain sizes have been estimated to be larger than modern humans, although such estimates have not been adjusted for their more robust builds. On average, Neanderthal males stood about 1.65 m tall (just under 5' 5") and were heavily built, and muscular due to their physical activity and robust bone structure. Females were about 1.53 to 1.57 m tall (about 5'–5'2").

The characteristic style of stone tools in the Middle Paleolithic is called the Mousterian Culture, after a prominent archaeological site where the tools were first found. The Mousterian culture is typified by the wide use of the Levallois technique. Mousterian tools were often produced using soft hammer percussion, with hammers made of materials like bones, antlers, and wood, rather than hard hammer percussion, using stone hammers. Near the end of the time of the Neanderthals, they created the Châtelperronian tool style, considered more "advanced" than that of the Mousterian. They either invented the Châtelperronian themselves or "borrowed" elements from the incoming modern humans who are thought to have created the Aurignacian.

Contents

[edit] Etymology and classification

The term Neanderthal Man was coined in 1863 by Irish anatomist William King. Neanderthal is now spelled two ways: the spelling of the German word Thal, meaning "valley or dale", was changed to Tal in the early 20th century, but the former spelling is often retained in English and always in scientific names, while the modern spelling is used in German.

The Neanderthal or "Neander valley" was named after theologian Joachim Neander, who lived there in the late seventeenth century.

The original German pronunciation (regardless of spelling) is with the sound /t/. (See German phonology.) When used in English, the term is usually anglicised to /θ/ (as in thin), though speakers more familiar with German use /t/.

For many years, professionals vigorously debated about whether Neanderthals should be classified as Homo neanderthalensis or as Homo sapiens neanderthalensis, the latter placing Neanderthals as a subspecies of Homo sapiens. However, recent evidence from mitochondrial DNA studies have been interpreted as evidence that Neanderthals were not a subspecies of H. sapiens.[5] Some scientists, for example Milford Wolpoff, argue that fossil evidence suggests that the two species interbred, and hence were the same biological species. Others, for example Cambridge Professor Paul Mellars, say "no evidence has been found of cultural interaction".[6]

[edit] Discovery

Location of Neander Valley, Germany
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Location of Neander Valley, Germany
Type Specimen, Neanderthal 1
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Type Specimen, Neanderthal 1

Neanderthal skulls were discovered in Engis, Belgium, in 1829 and Forbes' Quarry, Gibraltar, in 1848 prior to the "original" discovery in a limestone quarry of the Neander Valley (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 Karl Fuhlrott, who turned the fossils over to anatomist Hermann Schaafhausen. The discovery was jointly announced in 1857.

That discovery is now considered the beginning of paleoanthropology. These and other discoveries led to the idea that these remains were from ancient Europeans who had played an important role in modern human origins. The bones of over 400 Neanderthals have been found since.

[edit] Notable fossils

[edit] Anatomy

Neanderthal cranial anatomy
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Neanderthal cranial anatomy
Comparison of crania, sapiens (left) and neanderthalensis (right)
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Comparison of crania, sapiens (left) and neanderthalensis (right)

The following is a list of physical traits that distinguish Neanderthals from modern humans; however, not all of them can be used to distinguish specific Neanderthal populations, from various geographic areas or periods of evolution, from other extinct humans. Also, many of these traits occasionally manifest in modern humans, particularly among certain ethnic groups. Nothing is known about the skin color, the hair, or the shape of soft parts such as eyes, ears, and lips of Neanderthals.[7]

Compared to modern humans, Neanderthals were shorter in size and had distinct morphological features, especially of the cranium, which gradually accumulated more derived aspects, particularly in certain relatively isolated geographic regions. Evidence suggests that they were much stronger than modern humans; their relatively robust stature is thought to be an adaptation to the cold climate of Europe during the Pleistocene epoch.

Neanderthal physical traits
Cranial Sub-cranial
Suprainiac fossa, a groove above the inion Considerably more robust
Occipital bun, a protuberance of the occipital bone that looks like a hair knot Large round finger tips
Projecting mid-face Barrel-shaped rib cage
Low, flat, elongated skull Large kneecaps
A flat basic cranium Long collar bones
Supraorbital torus, a prominent browridge Short, bowed shoulder blades
1200-1750 cm³ skull capacity (10% greater than modern human average) Thick, bowed shaft of the thigh bones
Lack of a protruding chin (mental protuberance; although later specimens possess a slight protuberance) Short shinbones and calf bones
Crest on the mastoid process behind the ear opening Long, gracile pelvic pubis (superior pubic ramus)
No groove on canine teeth
A retromolar space posterior to the third molar
Bony projections on the sides of the nasal opening
Distinctive shape of the bony labyrinth in the ear
Larger mental foramen in mandible for facial blood supply
A broad, projecting nose

Based on a 2001 study, some commentators speculated that Neanderthals had red hair, and that some red-headed and freckled humans today share some heritage with Neanderthals;[8] however, many other researchers disagree.[9]

[edit] Language

See also: Origin of language

The idea that Neanderthals lacked complex language was widespread, despite concerns about the accuracy of reconstructions of the Neanderthal vocal tract, until 1983, when a Neanderthal hyoid bone was found at the Kebara Cave in Israel. The hyoid is a small bone that connects the musculature of the tongue and the larynx, and by bracing these structures against each other, allows a wider range of tongue and laryngeal movements than would otherwise be the case. Therefore, it seems to imply the presence of anatomical conditions for speech to occur. The bone that was found is virtually identical to that of modern humans.[10]

Furthermore, the morphology of the outer and middle ear of Neanderthal ancestors, Homo heidelbergensis, found in Spain, suggests they had similar auditory sensitivity as modern humans and very different from chimpanzees. Therefore, they were not only able to produce a wide range of sounds, they were also able to differentiate between these sounds. [11]

Aside from the morphological evidence above, neurological evidence for potential speech in neanderthalensis exists in the form of the hypoglossal canal. The canal of neanderthalensis is the same size or larger than in modern humans, which are significantly larger than the canal of modern chimpanzees and australopithecines. The canal carries the hypoglossal nerve, which supplies the muscles of the tongue with motor coordination. Researchers indicate that this evidence suggests that neanderthalensis had vocal capabilities similar to, or possibly exceeding that of, modern humans. [12]

A research team from the University of California, Berkeley, led by David DeGusta, suggests that the size of the hypoglossal canal is not an indicator of speech. His team's research, which shows no correlation between canal size and speech potential, shows there are a number of extant non-human primates and fossilized australopithecines which have equal or larger hypoglossal canal. [13]

Many people believe that even without the hyoid bone evidence, it is obvious that tools as advanced as those of the Mousterian Era, attributed to Neanderthals, could not have been developed without cognitive skills encompassing some form of spoken language.

A great many phonetically ill-informed comments surround the reconstruction of the Neanderthal vocal tract and the quality of Neanderthal speech. The popular view that the Neanderthals had a high larynx and therefore could not have produced the range of vowels supposedly essential for human speech is based on a disputed reconstruction of the vocal tract from the available fossil evidence, and a debatable interpretation of the acoustic characteristics of the reconstructed vocal tract. A larynx position as low as that found for modern female humans may have been present in adult male Neanderthals. Furthermore, the vocal tract is a plastic thing, and larynx movement is possible in many mammals. Finally, the suggestion that the vowels /i, a, u/ are essential for human language (and that if Neanderthals lacked them, they could not have evolved a human-like language) ignores the absence of one of these vowels in very many human languages, and the occurrence of 'vertical vowel systems' which lack both /i/ and /u/.

More doubtful suggestions about Neanderthal speech suggest that it would have been nasalised either because i) the tongue was high in the throat (for which there is no universally accepted evidence), or ii) because the Neanderthals had large nasal cavities. Nasalisation depends on neither of these things, but on whether or not the soft palate is lowered during speech. Nasalisation is therefore controllable, and we have no idea whether Neanderthal speech was nasalised or not. Comments on the lower intelligibility of nasalised speech ignore the fact that many varieties of English habitually have nasalised vowels, particularly low vowels, with no apparent effect on intelligibility.

Finally, suggestions that a 'stout larynx' would result in a higher rate of vibration of the vocal folds and hence a higher percept of pitch are erroneous. If the existence of a 'stout larynx' suggests large vocal folds, these would vibrate relatively slowly, and therefore give a percept of a lower pitch. Any comment about 'pitch levels' ignores the fact that the rate of vocal fold vibration can be changed by altering the tension in the vocal folds and by changing subglottal pressure. In other words, whatever the biological characteristics of the vocal folds, Neanderthal speech would be likely to have shown variation in the rate of vocal fold vibration (perceived as pitch), just as human speech or other mammalian vocalisations, and a non-biologically determined average or default pitch level could have been adopted by actively changing the 'neutral' state of the vocal folds.

[edit] Tools

Neanderthal Hunter, (American Mus. Nat. Hist.)    
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Neanderthal Hunter, (American Mus. Nat. Hist.)

Neanderthal (Middle Paleolithic) archaeological sites show a smaller and different toolkit than those which have been found in Upper Paleolithic sites, which were perhaps occupied by modern humans that superseded them. Fossil evidence indicating who may have made the tools found in Early Upper Paleolithic sites is still missing.

There is little evidence that Neanderthals used antlers, shell, or other bone materials to make tools; their bone industry was relatively simple. However, there is good evidence that they routinely constructed a variety of stone implements. The Neanderthal (Mousterian) tool kits consisted of sophisticated stone-flakes, task-specific hand axes, and spears. Many of these tools were very sharp. There is also good evidence that they used a lot of wood, objects which are unlikely to have been preserved until today.

Also, while they had weapons, none have yet been found that were used as projectile weapons. They had spears, in the sense of a long wooden shaft with a spearhead firmly attached to it, but these were not spears specifically crafted for flight (perhaps better described as a javelin). However, a number of 400,000 year old wooden projectile spears were found at Schöningen in northern Germany. These are thought to have been made by the Neanderthal's ancestors, Homo erectus or H. heidelbergensis. Generally, projectile weapons are more commonly associated with H. sapiens. The lack of projectile weaponry is an indication of different sustenance methods, rather than inferior technology or abilities. The situation is identical to that of native New Zealand Maoris - modern H. sapiens, who also rarely threw objects, but used spears and clubs instead.[citation needed]

Although much has been made of the Neanderthal's burial of their dead, their burials were less elaborate than those of anatomically modern humans. The interpretation of the Shanidar IV burials as including flowers, and therefore being a form of ritual burial,[14] has been questioned.[15] On the other hand, five of the six flower pollens found with Shanidar IV are known to have had 'traditional' medical uses, even among relatively recent 'modern' populations. In some cases Neanderthal burials include grave goods, such as bison and aurochs bones, tools, and the pigment ochre.

Neanderthals performed a sophisticated set of tasks normally associated with humans alone. For example, they constructed complex shelters, controlled fire, and skinned animals. Particularly intriguing is a hollowed-out bear femur with four holes spaced like four holes in the diatonic scale, claimed by many to have been deliberately bored into it. This bone was found in western Slovenia in 1995, near a Mousterian fireplace used by Neanderthals, but its significance is still a matter of dispute : some paleoanthropologists have postulated that it might have been a flute while some others have expressed that it is natural bone modified by bears. However, the holes are all roughly the same size and bored into the bone in a straight horizontal line along its shaft which would not seem to support the latter assertion.

See also: prehistoric music and Divje Babe.

[edit] Habitat and range

Distribution of the Neanderthals
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Distribution of the Neanderthals

Classic Neanderthal fossils have been found over a large area. From northern Germany in the north, to Israel and Mediterranean countries like Spain and Italy in the south and from England 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 especially 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 that they occupied, since Middle-Palaeolithic looking artifacts have been found even further north, up to 60° on the Russian plain.[16]

[edit] Ritual defleshing or cannibalism

Intentional burial and the inclusion of grave goods is the most typical representation of ritual behaviour in the Neanderthals and denote a developing ideology. However, another much debated and controversial manifestation of this ritual treatment of the dead comes from the evidence of cut-marks on the bone which has historically been viewed as evidence of cannibalism. Neanderthal bones from various sites (Combe-Grenal and Abri Moula in France, Krapina in Croatia and Grotta Guattari in Italy) have all been cited as bearing cut marks made by stone tools.[1] However, re-evaluation of these marks using high-powered microscopes, comparisons to contemporary butchered animal remains and recent ethnographic cases of excarnation mortuary practises have shown that perhaps this was a case of ritual defleshing. Fragments of bones from Krapina bear marks that are similar to those seen on bones from secondary burials at a Michigan ossuary (14th century AD) and are indicative of removing the flesh of a partially decomposed body. At Grotta Guattari, the apparently purposefully widened base of the skull (for access to the brains) has been shown to be caused by carnivore action, with hyena tooth marks found on the skull and mandible. However, analysis of the bones from Abri Moula in France does seem to suggest cannibalism was practiced here. This is the case since cut-marks are concentrated in the places where one would expect them in the case of butchery, instead of defleshing. Additionally the treatment of the bones was similar to that of roe deer bones, assumed to be food remains, found in the same shelter.[17] The evidence indicating cannibalism would not necessarily distinguish Neanderthals from modern Homo sapiens. Existing Homo sapiens tribes are known to practice cannibalism and mortuary defleshing.

[edit] Pathology

Within the west Asian and European record there are five broad groups of pathology or injury noted in Neanderthal skeletons.

[edit] Fractures

Neanderthals seemed to suffer a high frequency of fractures, especially common on the ribs (Shanidar IV, La Chapelle-aux-Saints ‘Old Man’), the femur (La Ferrassie 1), fibulae (La Ferrassie 2 and Tabun 1), spine (Kebara 2) and skull (Shanidar I, Krapina, Sala 1). These fractures are often healed and show little or no sign of infection, suggesting that injured individuals were cared for during times of incapacitation. The pattern of fractures has been found to be similar to modern rodeo clowns which, along with the absence of throwing weapons, suggests that they may have hunted by leaping onto their prey and stabbing or even wrestling it to the ground.

[edit] Trauma

Particularly related to fractures are cases of trauma seen on many skeletons of Neanderthals. These usually take the form of stab wounds, as seen on Shanidar III, whose lung was probably punctured by a stab wound to the chest between the 8-9th ribs. This may have been an intentional attack or merely a hunting accident; either way the man survived for some weeks after his injury before being killed by a rock fall in the Shanidar cave. Other signs of trauma include blows to the head (Shanidar I and IV, Krapina), all of which seemed to have healed, although traces of the scalp wounds are visible on the surface of the skulls.

[edit] Degenerative Disease

Arthritis is particularly common in the older Neanderthal population, specifically targeting areas of articulation such as the ankle (Shanidar III), spine and hips (La Chapelle-aux-Saints ‘Old Man’), arms (La Quina 5, Krapina, Feldhofer) knees, fingers and toes. This is closely related to degenerative joint disease, which can range from normal, use-related degeneration to painful, debilitating restriction of movement and deformity and is seen in varying degree in the Shanidar skeletons (I-IV).

[edit] Hypoplastic Disease

Dental enamel hypoplasia is an indicator of stress during the development of teeth and records in the striations and grooves in the enamel periods of food scarcity, trauma or disease. A study of 669 Neanderthal dental crowns showed that 75% of individuals suffered some degree of hypoplasia and the nutritional deficiencies were the main cause of hypoplasia and eventual tooth loss. All particularly aged skeletons show evidence of hypoplasia and it is especially evident in the Old Man of La Chapelle-aux-Saints and La Ferrassie 1 teeth.

[edit] Infection

Evidence of infections on Neanderthal skeletons is usually visible in the form of lesions on the bone, which are created by systematic infection on areas closest to the bone. Shanidar I has evidence of the degenerative lesions as does La Ferrassie 1, whose lesions on both femurs, tibiae and fibulae are indicative of a systemic infection or carcinoma (malignant tumour/cancer).

Reconstruction of a Neanderthal woman
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Reconstruction of a Neanderthal woman

[edit] The fate of the Neanderthals

The Neanderthals began to be displaced around 45,000 years ago by modern humans (Homo sapiens), as the Cro-Magnon people appeared in Europe. Despite this, populations of Neanderthals held on for thousands of years in regional pockets such as modern-day Croatia and the Iberian and Crimean peninsulas.

[edit] Genome

In July 2006, the Max Planck Institute for Evolutionary Anthropology and 454 Life Sciences announced that they would be sequencing the Neanderthal genome over the next two years. At three billion base pairs, the Neanderthal genome is roughly the size of the human genome and likely shares many identical genes. It is thought that a comparison of the Neanderthal genome and Human genome will expand understanding of Neanderthals as well as the evolution of humans and human brains.[18]

DNA researcher Svante Paabo has tested more than 70 Neanderthal specimens and found only one that had enough DNA to sample. Preliminary DNA sequencing from a 38,000 year old bone fragment of a femur bone found at a Croatian Vindija Cave in 1980 shows that Neanderthalis and Homo sapiens share about 99.5% of their DNA. It is believed that the two species shared a common ancestor about 500,000 years ago. Nature has calculated the species diverged about 516,000 years ago, whereas fossil records show a time of about 400,000 years ago. From DNA records, scientists hope to falsify or confirm the theory that there was interbreeding between the species.[19]

Edward Rubin of the Lawrence Berkeley National Laboratory in Berkeley, California states that recent genome testing of Neanderthals suggests human and Neanderthal DNA are some 99.5 percent to nearly 99.9 percent identical.[20][21]

In November 2006, a paper was published in the U.S. journal Proceedings of the National Academy of Sciences, in which a team of European researchers report that Neanderthals and humans interbred. Co-author Erik Trinkaus from Washington University explains, "Closely related species of mammals freely interbreed, produce fertile viable offspring, and blend populations." The study claims to settle the extinction controversy; according to researchers, the human and neanderthal populations blended together through sexual reproduction. Erik Trinkaus states, "Extinction through absorption is a common phenomenon."[22] and "From my perspective, the replacement vs. continuity debate that raged through the 1990s is now dead".[23]

[edit] Key dates

  • 1829: Neanderthal skulls were discovered in Engis, Belgium.
  • 1848: Skull of an ancient human was found in Forbes' Quarry, Gibraltar. Its significance was not realized at the time.
  • 1856: Johann Karl Fuhlrott first recognized the fossil called “Neanderthal man”, discovered in Neandertal a valley near Mettmann in what is now North Rhine-Westphalia, Germany.
  • 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.
  • 1899: Hundreds of Neanderthal bones were described in stratigraphic position in association with cultural remains and extinct animal bones.
  • 1908: A nearly complete Neanderthal skeleton was discovered in association with Mousterian tools and bones of extinct animals.
  • 1953-1957: Ralph Solecki uncovered nine Neanderthal skeletons in Shanidar Cave in northern Iraq.
  • 1975: Erik Trinkaus’s study of Neanderthal feet confirmed that they walked like modern humans.
  • 1987: New thermoluminescence resulted from Palestine fossils date Neanderthals at Kebara to 60,000 BP and modern 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: New ESR dates showed that the Tabun Neanderthal was contemporaneous with modern humans from Skhul and Qafzeh.
  • 2000: Igor Ovchinnikov, Kirsten Liden, William Goodman et al. retrieved DNA from a Late Neanderthal (29,000 BP) infant from Mezmaikaya Cave in the Caucausus.
  • 2005: The Max Planck Institute for Evolutionary Anthropology launched a project to reconstruct the Neanderthal genome.
  • 2006: The Max Planck Institute for Evolutionary Anthropology announced that it planned to work with Connecticut-based 454 Life Sciences to reconstruct the Neaderthal genome.

[edit] Popular culture

Popular literature has tended to greatly exaggerate the ape-like gait and related characteristics of the Neanderthals. It has been determined that some of the earliest specimens found in fact suffered from severe arthritis. The Neanderthals were fully bipedal and had a slightly larger average brain capacity than a typical modern human, though it is thought the brain structure may have been organized differently.

In popular idiom the word neanderthal is sometimes used as an insult, to suggest that a person combines a deficiency of intelligence and an attachment to brute force, as well as perhaps implying the person is old fashioned or attached to outdated ideas, much in the same way as "dinosaur" or "Yahoo" is also used. Counterbalancing this are sympathetic literary portrayals of Neanderthals, as in the novel The Inheritors by William Golding and Jean M. Auel's Earth's Children series, or the more serious treatment by palaeontologist Björn Kurtén, in several works including Dance of the Tiger, and British psychologist Stan Gooch in his hybrid-origin theory of humans.

[edit] See also

[edit] External links

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  • Smithsonian
  • Archaelogy Info
  • MNSU
  • "Humans and Neanderthals interbred": Modern humans contain a little bit of Neanderthal, according to a new theory, because the two interbred and became one species. (Cosmos magazine, November 2006)
  • BBC.co.uk - 'Neanderthals "mated with modern humans": A hybrid skeleton showing features of both Neanderthal and early modern humans has been discovered, challenging the theory that our ancestors drove Neanderthals to extinction', BBC (April 21, 1999)
  • BBC.co.uk - 'Neanderthals "had hands like ours": The popular image of Neanderthals as clumsy, backward creatures has been dealt another blow', Helen Briggs, BBC (March 27, 2003)
  • Greenwych.ca - 'Chewed or Chipped? Who Made the Neanderthal Flute? Humans or Carnivores?' Bob Fink, Greenwich Publishing (March, 2003)

[edit] References

  1. ^ J. L. Bischoff et al. (2003). "Neanderthals". J. Archaeol. Sci. (30): 275.
  2. ^ Rincon, Paul (2006-09-13). Neanderthals' 'last rock refuge'. BBC News. Retrieved on 2006-09-19.
  3. ^ Mcilroy, Anne (2006-09-13). Neanderthals may have lived longer than thought. Globe and Mail. Retrieved on 2006-09-19.
  4. ^ Richard G. Klein (March 2003). "PALEOANTHROPOLOGY: Whither the Neanderthals?". Science 299 (5612): 1525-1527. DOI:10.1126/science.1082025.
  5. ^ Hodges, S. Blair (2000-12-07). Human evolution: A start for population genomics. Nature Publishing Group.
  6. ^ Modern humans, Neanderthals shared earth for 1,000 years. ABC News (Australia) (2005-09-01). Retrieved on 2006-09-19.
  7. ^ Scientists Build 'Frankenstein' Neanderthal Skeleton
  8. ^ Red-Heads and Neanderthals (May 2001). Retrieved on 2005-10-28.
  9. ^ Nicole's hair secrets (2002-02-10). Retrieved on 2005-11-02.
  10. ^ B. Arensburg, A. M. Tillier, B. Vandermeersch, H. Duday, L. A. Schepartz & Y. Rak (April 1989). "A Middle Palaeolithic human hyoid bone". Nature (338): 758-760. DOI:10.1038/338758a0.
  11. ^ Martinez, I., L. Rosa, J.-L. Arsuaga, P. Jarabo, R. Quam, C. Lorenzo, A. Gracia, J.-M. Carretero, J.M. Bermúdez de Castro, E. Carbonell (July 2004). "Auditory capacities in Middle Pleistocene humans from the Sierra de Atapuerca in Spain = PNAS" 101 (27): 9976-9981.
  12. ^ Richard F. Kay, Matt Cartmill, and Michelle Balow (April 1998). "The hypoglossal canal and the origin of human vocal behavior". PNAS 95 (9): 5417-5419.
  13. ^ David DeGusta, W. Henry Gilbert and Scott P. Turner (Feb 1999). "Hypoglossal canal size and hominid speech". PNAS 96 (4): 1800-1804.
  14. ^ R. S. Solecki (1975). "Shanidar IV, a Neanderthal flower burial in northern Iraq". [[Science (journal)|]] 190 (28): 880.
  15. ^ J. D. Sommer (1999). "The Shanidar IV 'Flower Burial': A Reevaluation of Neanderthal Burial Ritual". Cambridge Archæological Journal 9: 127–129.
  16. ^ Pavlov, P., W. Roebroeks, and J. I. Svendsen (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.
  17. ^ Defleur A, White T, Valensi P, Slimak L, Cregut-Bonnoure E, 1999. Neanderthal cannibalism at Moula-Guercy, Ardèche, France. Science 286: 128-131.
  18. ^ Moulson, Geir. "Neanderthal genome project launches", MSNBC.com, Associated Press. Retrieved on 2006-08-22.
  19. ^ http://www.nytimes.com/2006/11/15/science/16neanderthalcnd.html
  20. ^ Neanderthal bone gives DNA clues
  21. ^ Scientists decode Neanderthal genes
  22. ^ Humans and Neanderthals interbred
  23. ^ Modern Humans, Neanderthals May Have Interbred