Neanderthal

The Neanderthal (, also with, and ) or Neandertal was a species of the Homo (Homo neanderthalensis or Homo sapiens neanderthalensis) genus that inhabited Europe and parts of western Asia. The first proto-Neanderthal traits appeared in Europe as early as 350,000 years ago. 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.

Neanderthals had many adaptations to a cold climate, such as large braincase, short but robust builds, and large noses &mdash; 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 with 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.

Etymology and classification
The term Neanderthal Man was coined in 1863 by Anglo-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 1901, 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 nearby in Düsseldorf 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, evidence from mitochondrial DNA studies have been interpreted as evidence that Neanderthals were not a subspecies of H. sapiens. Recent genetic simulations suggested that 5% of human DNA can only be accounted for by assuming a substantial contribution of Neanderthaler to the European gene pool of up to 25%. 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".

Discovery
Neanderthal skulls were first discovered in Engis, Belgium (1829) and in Forbes' Quarry, Gibraltar (1848), both 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 Schaaffhausen. The discovery was jointly announced in 1857.

The original Neandertal 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.

Notable fossils

 * La Ferrassie 1
 * Shanidar 1
 * La Chappelle-aux-Saints 1
 * Le Moustier
 * Neanderthal 1
 * Kebara 2

Anatomy


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.

Compared to modern humans, Neanderthals were similar in height but with more robust bodies, 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.

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; however, many other researchers disagree.

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 possible. The presence of this bone implies that speech was anatomically possible. The bone that was found is virtually identical to that of modern humans.

The morphology of the outer and middle ear of Neanderthal ancestors, Homo heidelbergensis, found in Spain, suggests they had an auditory sensitivity similar to modern humans and very different from chimpanzees. They were probably able to differentiate between many different sounds.

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 australopithecines and modern chimpanzees. The canal carries the hypoglossal nerve, which controls the muscles of the tongue. This indicates that neanderthalensis had vocal capabilities similar to modern humans. 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.

Another anatomical difference between Neanderthals and humans that is relevant regarding speech is their lack of a mental protuberance (the point at the tip of the chin). While some Neandertal individuals do possess a mental protuberance, their chins never show the inverted T-shape of modern humans. In contrast, some Neanderthal individuals show inferior lateral mental tubercles (little bumps at the side of the chin). The mentalis muscle, one of the muscles that move the lower lip, is attached to the tip of the chin.

Tools
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.

Neanderthals are thought to have used tools of the Mousterian class, which 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. A result of this is that 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, whether they had implements that were used as projectile weapons is controversial. They had spears, in the sense of a long wooden shaft with a spearhead firmly attached to it, but they are thought by some to have been thrusting spears. Still, a Levallois point embedded in a vertebra shows an angle of impact suggesting that it entered by a "parabolic trajectory" suggesting that it was the tip of a projectile. Moreover, 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 Homo 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 Maori - modern Homo sapiens, who also rarely threw objects, but used spears and clubs instead. Nonetheless, the fact that it is much safer to strike prey or foes from a distance where they cannot strike back would put anyone depending on close quarter weapons at a tactical disadvantage.

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, has been questioned. 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 that contains holes that may have been deliberately bored into it. This bone was found in western Slovenia in 1995, near a Mousterian fireplace, 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. See: Divje Babe.

Habitat and range
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 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 that they occupied, since Middle-Palaeolithic looking artifacts have been found even further north, up to 60° on the Russian plain. Recent evidence has extended the Neanderthal habitat range by about 1250 miles east into Southern Siberia.

Ritual defleshing or cannibalism
Intentional burial and the inclusion of grave goods is the most typical representation of ritual behavior 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. However, results of technological tests reveal varied causes.

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.
 * 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.
 * According to some studies, fragments of bones from Krapina show 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.
 * According to others, the marks on the bones found at Krapina are indicative of cannibalism, although whether this was for nutritional or ritual purposes cannot be determined with certainty.
 * Analysis of bones from Abri Moula in France does seem to suggest cannibalism was practiced here. Cut-marks are concentrated in places expected 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.

The evidence indicating cannibalism would not distinguish Neanderthals from modern Homo sapiens. Ancient and existing Homo sapiens, including the Korowai, are known to have practiced cannibalism and/or mortuary defleshing.

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

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, 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.

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 8th and 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.

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).

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.

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 femora, tibiae and fibulae are indicative of a systemic infection or carcinoma (malignant tumour/cancer).

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. The last known population lived around a cave system on the remote south facing coast of Gibraltar, from 30,000 to 24,000 years ago.

Neanderthal findings in Abrigo do Lagar Velho, Portugal of 24,500 BP, featuring admixtures with early modern humans, have been published. However, the paleontological analysis of modern human emergence in Europe has been shifting from considerations of the Neanderthals to assessments of the biology and chronology of the earliest modern humans in western Eurasia. This focus, involving morphologically modern humans before 28,000 BP shows accumulating evidence that they present a variable mosaic of derived modern human, archaic human, and Neanderthal features.

Genome
While previous investigations concentrated on mitochondrial DNA (mtDNA), that due to strictly matrilineal inheritance and subsequent vulnerability to genetic drift is of limited value to disprove interbreeding, more recent investigations have access to growing strings of deciphered nuclear DNA (nDNA).

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.

DNA researcher Svante Pääbo 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 Vindija cave in Croatia in 1980 shows that Homo neanderthalensis and Homo sapiens share about 99.5% of their DNA. 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. From DNA records, scientists hope to falsify or confirm the theory that there was interbreeding between the species.

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.

On November 16, 2006, Science Daily published scientific test results demonstrating that Neanderthals and ancient humans probably did not interbreed. Scientists with the U.S. Department of Energy’s Lawrence Berkeley National Laboratory (Berkeley Lab) and the Joint Genome Institute (JGI) sequenced genomic nuclear DNA (nDNA) from a fossilized Neanderthal femur. Their results more precisely indicate a common ancestor about 706,000 years ago, and a complete separation of the ancestors of the species about 376,000 years ago. Their results show that the genomes of modern humans and Neanderthals are at least 99.5% identical, but despite this genetic similarity, and despite the two species having cohabitated the same geographic region for thousands of years, there is no evidence of any significant crossbreeding between the two. Edward Rubin, director of both JGI and Berkeley Lab’s Genomics Division: “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.”

A new investigation suggests that at least 5% of the genetic material of modern Europeans and West Africans has an archaic origin, due to interbreeding with Neanderthal and a hitherto unknown archaic African population. Plagnol and Wall arrived at this result by first calculating a "null model" of genetic characteristics that would fulfill the requirement of descendence from Homo sapiens sapiens in a straight line. Next they compared this model to the current distribution and characteristics of existing genetic polymorphisms, and concluded that this "null model" deviated considerably from what would be expected. Genetic simulations indicated this 5% of DNA not accounted for by the null model corresponds to a substantial contribution to the European gene pool of up to 25%. Future investigation - including a full scale Neanderthal genome project - is expected to cast more light on the subject of genetic polymorphisms to supply more details. Contrary to the investigation of mtDNA, the study of polymorph mutations has the potential to answer the question whether - and to what extent - Homo neanderthalensis and Homo sapiens interbred.

In November 2006, another 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. Trinkaus states, "Extinction through absorption is a common phenomenon." and "From my perspective, the replacement vs. continuity debate that raged through the 1990s is now dead".

The most accurate molecular estimates currently available suggest that H. sapiens and H. neanderthalensis direct lineages (excluding genetic elements from interbreeding/absorption) diverged around 800,000 years ago.

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 realised at the time.
 * 1856: Johann Karl Fuhlrott first recognised the fossil called “Neanderthal man”, discovered in Neanderthal 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: Thermoluminescence results 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: ESR dates showed that the Tabun Neanderthal was contemporaneous with modern humans from Skhul and Qafzeh.
 * 1997 Matthias Krings et al. are the first to amplify Neanderthal mitochondrial DNA (mtDNA) using a specimen from Feldhofer grotto in the Neander valley. Their work is published in the journal Cell.
 * 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 Neanderthal genome.

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, Isaac Asimov's The Ugly Little Boy and Jean M. Auel's Earth's Children series, or the more serious treatment by Swedish palaeontologist Björn Kurtén, in several works including Dance of the Tiger, and British psychologist Stan Gooch in his hybrid-origin theory of humans. A trilogy of science fiction novels dealing with neaderthals (called The Neanderthal Parallax) written by Robert J. Sawyer, explores a scenario where neanderthals are seen as a separate species from humans, survive in a parallel universe version of earth, and what happens when they, having developed a sophisticated technological culture of their own, open a portal to this version of the earth. Those three novels are titled Hominids, Humans, and Hybrids, respectively, and all form essentially one story.