The Neanderthal (IPA: /niːˈændərθɑːl/, also with /neɪ-/, and /-tɑːl/) or Neandertal was a species of the Homo (Homo neanderthalensis or Homo sapiens neanderthalensis)[1] genus that inhabited Europe and parts of western
Asia. The first proto-Neanderthal traits appeared in Europe as early as 350,000 years ago.[2] 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.[3][4][5]
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 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
Type Specimen, Neanderthal 1
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.[6] 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%.[7] Some scientists, for example Milford
Wolpoff, argue that fossil evidence suggests that the two species interbred, and hence were the same biological species [citation needed]. Others, for example
Cambridge Professor Paul Mellars, say "no evidence has been found of cultural
interaction".[8]
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
Anatomy
Neanderthal cranial anatomy
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.[9]
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;[citation needed] 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, trabecular (spongy) 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;[10] however, many other researchers disagree.[11]
Language
- See also: Origin of language
|
|
The neutrality of this section is disputed.
Please see the discussion on the talk page. |
The idea that Neanderthals lacked complex language was widespread,[citation needed] 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.[12]
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. [13]
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. [14] 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. [15]
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.[16] 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.
A recent extraction of DNA from Neanderthal bones indicates that Neanderthals had the same version of the FOXP2 gene as modern humans. This gene is known to play a role in human language.[17]
Tools
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.
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. [18]
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 [19]. 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 [20]. 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. [21] 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,[22] has been
questioned.[23] 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
Sites where typical Neanderthal fossils have been found.
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.[24]Recent evidence has extended the
Neanderthal habitat range by about 1250 miles east into Southern Siberia.[25]
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.[26] 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.[27]
- 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.[28]
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.[29]
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).
Reconstruction of a Neanderthal child from
Gibraltar (Anthropological Institute, University of
Zürich)
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.[30] 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.[31] [32]
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.[33]
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.[34]
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.[35][36]
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.[7] 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.[37]
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."[38] and "From my perspective, the replacement vs. continuity debate that raged
through the 1990s is now dead".[39]
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.[40]
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 Finnish 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.
See also
References
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