Neanderthal 45347 224963010 2008-07-11T05:39:54Z Rokus01 3149596 Restored to version Vargenau at 05:31, 5 July 2008 following the rules of [[WP:NPOV]]. {{otheruses}} {{Taxobox | name = Neanderthal | fossil_range = [[Pleistocene]] | image = Homo sapiens neanderthalensis.jpg | image_caption = ''H. neanderthalensis'' La Chapelle-aux-Saints. |status=Fossil |extinct= | regnum = [[Animal]]ia | phylum = [[Chordate|Chordata]] | classis = [[Mammal]]ia | ordo = [[Primate]]s | familia = [[Hominidae]] | genus = ''[[Homo (genus)|Homo]]'' | species = '''''H. neanderthalensis''''' | binomial = †''Homo neanderthalensis'' | binomial_authority = [[William King (geologist)|King]], 1864 | synonyms = ''Palaeoanthropus neanderthalensis''<br>''H. s. neanderthalensis'' |<!-- range_map = Range of Homo neanderthalensis.png | range_map_width = 220px | range_map_caption = Neanderthal range --> }} The '''Neanderthal''' ({{IPAEng|niːˈændərθɔːl}}, also with {{IPA|/neɪ-/}}, and {{IPA|/-tɑːl/}}), or '''Neandertal''', was a species of the ''[[Homo (genus)|Homo]]'' genus (''Homo neanderthalensis'' or ''Homo sapiens neanderthalensis'')<ref>{{cite journal |author=Tattersall I, Schwartz JH |title=Hominids and hybrids: the place of Neanderthals in human evolution |journal=Proc. Natl. Acad. Sci. U.S.A. |volume=96 |issue=13 |pages=7117–9 |year=1999 |pmid=10377375 | doi = 10.1073/pnas.96.13.7117 <!--Retrieved from CrossRef by DOI bot-->| quote =on the view that these distinctive hominids merit species recognition in their own right as '''Homo neanderthalensis''' (e.g., refs. 4 and 5), at least as many still regard them as no more than a strange variant of our own species, '''Homo sapiens''' (6, 7)}} Available [http://www.pnas.org/cgi/content/full/96/13/7117 on-line]</ref> which inhabited [[Europe]] and parts of western and [[central Asia]]. The first proto-Neanderthal traits appeared in Europe as early as 350-500 thousand years ago.<ref>{{cite journal | author = J. L. Bischoff et al. | year = 2003 | title = The Sima de los Huesos Hominids Date to Beyond U/Th Equilibrium (>350 kyr) and Perhaps to 400–500 kyr: New Radiometric Dates | journal = J. Archaeol. Sci. | issue = 30 | pages = 275 | volume = 30 | doi = 10.1006/jasc.2002.0834 <!--Retrieved from CrossRef by DOI bot-->}}</ref> By 130,000 years ago, complete Neanderthal characteristics had appeared and by 50,000 years ago, Neanderthals disappeared from Asia, although they did not reach extinction in Europe until 30,000 years ago.<ref>[http://dsc.discovery.com/news/2008/06/23/neanderthal-tools.html Last Neanderthals Were Smart], Discovery Channel</ref> No Neanderthal skeletons of younger dating have been found, though it has been suggested that Neanderthals survived longer in Southern Iberia.<ref name=SouthIberia>[https://pmc.ucsc.edu/~apaytan/projects/papers_PDF/2007_Articles/Jimenez-EspejoEtAl_QSR_2007.pdf] Climate forcing and Neanderthal extinction in Southern Iberia: insights from a multiproxy marine record - Francisco J. Jiménez-Espejo et al.</ref><ref>The early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern human emergence in Iberia - Cidália Duarte, João Maurício, Paul B. Pettitt, Pedro Souto, Erik Trinkaus, Hans van der Plicht, and João Zilhão, PNAS Vol. 96, Issue 13, 7604-7609, June 22, 1999 [http://www.pnas.org/cgi/content/full/96/13/7604]</ref> Neanderthal may have coexisted with modern humans up to 15,000 years after ''[[Homo sapiens]]'' had migrated into Europe.<ref>{{cite web | url = http://news.bbc.co.uk/2/hi/science/nature/5343266.stm | title = Neanderthals' 'last rock refuge' | author = Rincon, Paul | date = [[2006-09-13]] | accessdate = 2006-09-19 | work = [http://news.bbc.co.uk/ BBC News]}}</ref><ref>{{cite web | author = Mcilroy, Anne | title = Neanderthals may have lived longer than thought | work = [http://www.theglobeandmail.com/ Globe and Mail] | url = http://www.theglobeandmail.com/servlet/story/RTGAM.20060913.wneanAM0913/BNStory/Science/home | date = [[2006-09-13]] | accessdate = 2006-09-19}}</ref><ref>{{cite journal | author = Richard G. Klein | year = 2003 | month = March | title = PALEOANTHROPOLOGY: Whither the Neanderthals? | journal = Science | volume = 299 | issue = 5612 | pages = 1525–1527 | doi = 10.1126/science.1082025 | pmid = 12624250 }}</ref> It is believed that the population of Neanderthals was never much more than 10,000 individuals.<ref>"Stone Age feminism? Females joining hunt may explain Neanderthals' end." Nickerson, Colin. ''Boston Globe'', November 10, 2007[http://www.boston.com/news/science/articles/2007/11/10/stone_age_feminism/?page=2]</ref> Neanderthals had many adaptations to a cold [[climate]]: short, robust builds, and rather large [[nose]]s &mdash; traits [[natural selection|selected]] by nature in cold climates.{{Fact|date=January 2008}} Their [[cranial capacity]] was larger than modern humans, indicating that their brains may have been larger. On average, the height of Neanderthals was comparable to contemporaneous ''homo sapiens''. Neanderthal males stood about 165–68 cm tall (about 5'5") and were heavily built with robust bone structure. Females stood about 152–56 cm tall (about 5'1").<ref>{{cite journal | last = Helmuth | first = H. | title = Body height, body mass and surface area of the Neanderthals | journal = Zeitschrift für Morphologie und Anthropologie | volume = 82 | issue = 1 | pages = 1–12 | date = [[1998]] | url = http://www.ncbi.nlm.nih.gov/pubmed/9850627 | accessdate = [[2008-05-26]] }}</ref> 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 [[rock (geology)|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== [[Image:Neandertal 1856.jpg|thumb|right|Type Specimen, Neanderthal 1.]] The [[Neandertal]] is a small valley of the river [[Düssel]] in the [[Germany|German]] Federal State of [[North Rhine-Westphalia]], located about {{convert|12|km|mi|abbr=on}} west of [[Düsseldorf]], the capital city of North Rhine-Westphalia. ''Neanderthal'' is now spelled two ways: the old spelling of the [[German language|German]] word ''Thal'', meaning "[[valley]] or dale", was changed to ''Tal'' in 1901, but the former spelling is often retained in [[English language|English]] and always in [[scientific name]]s, while the modern spelling is used in German while referring to the valley itself. The [[Neandertal|Neander Valley]] was named after theologian [[Joachim Neander]], who lived nearby in [[Düsseldorf]] in the late seventeenth century. In turn, Neanderthals were named after "Neander Valley", where the first Neanderthal remains were found. The term ''Neanderthal Man'' was coined in 1863 by [[Anglo-Irish]] [[geologist]] [[William King (geologist)|William King]]. The original German pronunciation (regardless of spelling) is with the sound /t/. (See [[German phonology]].) In English the term is commonly anglicised to /θ/ (''th'' as in ''thin''), though scientists frequently use /t/. "Neander" is a classicized form of the common German surname Neumann. For some time, professionals debated whether Neanderthals should be classified as ''Homo neanderthalensis'' or as ''Homo sapiens neanderthalensis'', the latter placing Neanderthals as a [[subspecies]] of ''[[Homo 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%.<ref name=Plagnol>Plagnol V, Wall JD: Possible ancestral structure in human populations. PLoS Genet 2006, 2:e105.[http://genetics.plosjournals.org/perlserv/?request=get-document&doi=10.1371/journal.pgen.0020105.eor]</ref> Additionally, morphological studies using latest techniques support that ''Homo neanderthalensis'' is a separate species and not a subspecies. <ref name=Harvati>Harvati & all: Neanderthal taxonomy reconsidered: implications of 3D primate models of intra- and interspecific differencesProc Natl Acad Sci U S A. 2004 Feb 3;101(5):1147-52)</ref> Some scientists, for example [[University of Michigan]] Professor [[Milford Wolpoff]], claim that [[fossil]] evidence suggests that the two species [[Hybrid (biology)|interbred]]. This would support the argument that the two were the same biological [[species]]. Others, for example [[University of Cambridge]] Professor Paul Mellars, say "no evidence has been found of cultural interaction"<ref>{{cite web | url = http://www.abc.net.au/news/newsitems/200509/s1450949.htm | title = Modern humans, Neanderthals shared earth for 1,000 years | date = [[2005-09-01]] | accessdate = 2006-09-19 | work = [http://abc.net.au/news/default.htm ABC News (Australia)]}}</ref> and evidence from [[mitochondrial DNA]] studies have been interpreted as evidence that [[Evolution of Homo sapiens#H. neanderthalensis|Neanderthals]] were not a subspecies of ''H. sapiens''.<ref>{{cite book | author = Hodges, S. Blair | title = Human Evolution: A start for population genomics | publisher = Nature Publishing Group (Nature. 2000 Dec 7;408(6813):652-3) | date = [[2000-12-07]]}}</ref> ==Discovery== [[Image:Neanderthal position.png|thumb|left|Location of [[Neander Valley]], Germany. (The highlighted area is the modern federal state of [[North Rhine-Westphalia]].)]] [[Craniometry|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 [[Neanderthal, Germany|Neander Valley]] in [[Erkrath]] near [[Düsseldorf]] in August, 1856, three years before [[Charles Darwin]]'s ''[[Origin of Species|On the Origin of Species]]'' was published.<ref>[http://anthropology.si.edu/humanorigins/ha/neand.htm Human Ancestors Hall: Homo neanderthalensis]</ref> The [[type specimen]], dubbed [[Neanderthal 1]], consisted of a skull cap, two [[femur|femora]], three bones from the right arm, two from the left arm, part of the left [[pelvis|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]]. The discovery was jointly announced in 1857. The original Neanderthal discovery is now considered the beginning of [[paleoanthropology]]. These and other discoveries led to the idea that these remains were from ancient [[Europe]]ans who had played an important role in modern [[human evolution|human origins]]. The bones of over 400 Neanderthals have been found since. ===Notable fossils=== * [[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. * [[Shanidar 1]]: Found in the Zagros Mountains in northern Iraq; a total of nine skeletons found believed to have lived in the Middle Paleolithic Period. One of the nine remains had an amputated arm. This is significant due to the fact that it shows that stone tools were present in that era. Also, another Neanderthal had been buried with flowers, showing that some type of burial ceremony may have occurred. * [[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 would have required that someone process his food for him, one of the earliest examples of Neanderthal altruism (similar to Shanidar I.) * [[Le Moustier]]: A fossilized skull, discovered in 1909, at the archeological 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. * [[Neanderthal 1]]: Initial Neanderthal specimen found during an archaeology dig in August 1856. Discovered in a limestone quarry at the Feldhofer grotto in Neanderthal, Germany. The find consisted of a skull cap, two femora, the three right arm bones, two of the left arm bones, ilium, and fragments of a scapula and ribs. * [[Kebara 2]] ==Anatomy== Compared to modern humans, Neanderthals were similar in height but with more robust bodies, and had distinct [[Comparative anatomy|morphological]] features, especially of the [[skull|cranium]], which gradually accumulated more derived aspects, particularly in certain relatively isolated geographic regions. Evidence suggests that they were much stronger than modern humans;<ref>[http://www.bbc.co.uk/nature/wildfacts/factfiles/459.shtml Science & Nature - Wildfacts - Neanderthal]</ref> their relatively robust stature is thought to be an adaptation to the cold climate of Europe during the [[Pleistocene]] epoch. A 2007 study confirmed that some Neanderthals had red hair and [[human skin color|pale skin color]]; however, the mutation in the [[melanocortin 1 receptor|MC1R]] gene arose independently of the mutation which causes a similar pigmentation pattern in modern humans.<ref>{{cite journal |last=Laleuza-Fox |first=Carles |coauthors=Holger Römpler ''et al'' |date=[[2007-10-25]] |title=A Melanocortin 1 Receptor Allele Suggests Varying Pigmentation Among Neanderthals |journal=[[Science (magazine)|Science]] |doi=10.1126/science.1147417 |accessdate= 2007-10-26 | volume = 318 | pages = 1453 |pmid=17962522}}</ref><ref>{{cite web | url = http://news.bbc.co.uk/2/hi/science/nature/7062415.stm | title = Neanderthals 'were flame-haired' | publisher = [[BBC.co.uk]] | author = Paul Rincon | date = [[2007-10-25]] | accessdate = 2007-10-25}}</ref> ===Distinguishing physical traits=== [[Image:Neanderthall cranial anatomy.JPG|thumb|right|350px|Neanderthal cranial anatomy.]] [[Image:Neandertal vs Sapiens.jpg|thumb|right|Comparison of [[skull|crania]], ''sapiens'' (left) and ''neanderthalensis'' (right).]] The following is a list of physical traits which 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 shape of soft parts such as eyes, ears, and lips of Neanderthals.<ref>{{cite web | url = http://www.livescience.com/history/050310_neanderthal_reconstruction.html | title = Scientists Build 'Frankenstein' Neanderthal Skeleton}}</ref> *'''Cranial''' **Suprainiac fossa, a groove above the [[inion]] **[[Occipital bun]], a protuberance of the [[occipital bone]] which looks like a hair knot <ref name=Gunz>Gunz and Harvati: The Neanderthal "chignon": variation, integration, and homology J Hum Evol. 2007 Mar;52(3):262-74 </ref> **Projecting mid-face **Low, flat, elongated skull **A flat basicranium[http://www.pajamacore.org/writings/origins.php][http://findarticles.com/p/articles/mi_m1200/is_n15_v141/ai_12140517]<ref>{{cite journal | author= B. Arensburg et al. |year = 1990 | title = A reappraisal of the anatomical basis for speech in Middle Palaeolithic hominids | journal = American Journal of Physical Anthropology | volume = '''83''' (2) | pages = 137–146 | doi = 10.1002/ajpa.1330830202}}</ref> **[[Supraorbital torus]], a prominent, trabecular (spongy) browridge **1200-1750 cm³ skull capacity (10% greater than modern human average) **Lack of a protruding chin (mental protuberance; although later specimens possess a slight protuberance) **Crest on the [[mastoid process]] behind the ear opening **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 (inner ear)|labyrinth]] in the ear **Larger [[mental foramen]] in mandible for facial blood supply **Broad, projecting [[nose]] *'''Sub-cranial''' **Considerably more robust **Large round [[finger]] tips **Barrel-shaped [[human rib cage|rib cage]] **Large kneecaps **Long collar bones **Short, bowed [[scapula|shoulder blades]] **Thick, bowed shaft of the thigh bones **Short shinbones and calf bones **Long, gracile pelvic pubis (superior pubic ramus) **Bowed Femur ==Language== {{seealso|Origin of language}} <!-- Image with unknown copyright status removed: [[Image:Neandertals48.jpg|thumb|Reconstruction of a Neandertal man made in 1948.]] --> The idea that Neanderthals lacked complex [[language]] was widespread,{{Fact|date=September 2007}} 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 which 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 which was found is virtually identical to that of modern humans.<ref>{{cite journal | author = B. Arensburg, A. M. Tillier, B. Vandermeersch, H. Duday, L. A. Schepartz & Y. Rak | year = 1989 | month = April | title = A Middle Palaeolithic human hyoid bone | journal = Nature | issue = 338 | pages = 758–760 | doi = 10.1038/338758a0 | volume = 338}}</ref> 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.<ref>{{cite journal | author = 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 | year = 2004 | month = July | title = Auditory capacities in Middle Pleistocene humans from the Sierra de Atapuerca in Spain | journal = [[Proceedings of the National Academy of Sciences|PNAS]] | volume = 101 | issue = 27 | pages = 9976–9981 | doi = 10.1073/pnas.0403595101 | pmid = 15213327}}</ref> 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 [[australopithecine]]s 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.<ref>{{cite journal | author = Richard F. Kay, Matt Cartmill, and Michelle Balow | year = 1998 | month = April | title = The hypoglossal canal and the origin of human vocal behavior | journal = [[Proceedings of the National Academy of Sciences|PNAS]] | volume = 95 | issue = 9 | pages = 5417–5419 | doi = 10.1073/pnas.95.9.5417 | pmid = 9560291}}</ref> 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.<ref> {{cite journal | author = David DeGusta, W. Henry Gilbert and Scott P. Turner | year = 1999 | month = Feb | title = Hypoglossal canal size and hominid speech | journal = [[Proceedings of the National Academy of Sciences|PNAS]] | volume = 96 | issue = 4 | pages = 1800–1804 | doi = 10.1073/pnas.96.4.1800 | pmid = 9990105}}</ref> Another anatomical difference between Neanderthals and Modern humans is their lack of a [[mental protuberance]] (the point at the tip of the chin). This may be relevant to speech as the mentalis muscle, one of the muscles which move the lower lip, is attached to the tip of the chin. While some Neanderthal individuals do possess a mental protuberance, their chins never show the inverted T-shape of modern humans.<ref> {{cite journal | author = Jeffrey Schwartz, Ian Tattersall| year = 2000 | month = May | title = The human chin revisited: What is it, and who has it?| journal = [[Journal of Human Evolution]] | volume = 38 | pages = 367–409 | doi = 10.1006/jhev.1999.0339}}</ref> In contrast, some Neanderthal individuals show inferior lateral mental tubercles (little bumps at the side 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.<ref>[http://www.nytimes.com/2007/10/19/science/19speech-web.html?ref=world Neanderthals Had Important Speech Gene, DNA Evidence Shows]</ref> [[Steven Mithen]] (2006) proposes that the Neanderthals had an elaborate [[proto-linguistic]] system of communication which was more [[origin of music|musical]] than modern human language, and which predated the separation of [[language]] and [[music]] into two separate modes of cognition.<ref>Steven Mithen, ''The Singing Neanderthals: the Origins of Music, Language, Mind and Body'' (2006).</ref> ==Tools== [[Image:Neanderthal 2D.jpg|thumb|Neanderthal Hunter, (American Mus. Nat. Hist.) {{3d alt|Neanderthal 3D.jpg}}.]] [[Image:Neandertal.jpg|thumb|right|Neanderthal Tool Maker.]] Neanderthal and [[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 which 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. Neanderthal ([[Mousterian]]) tools most often consisted of sophisticated [[lithic flake|stone-flakes]], task-specific [[hand axe]]s, and [[spear]]s. 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.<ref>Henig, Martin. "Odd man out: Neanderthals and modern humans." ''British Archeology'', #51, Feb 2000. [http://www.britarch.ac.uk/BA/ba51/ba51feat.html]</ref> Also, while they had [[weapon]]s, whether they had implements which were used as [[projectile]] weapons is controversial. They had [[spear]]s, made of long wooden shafts with spearheads firmly attached, but they are thought by some to have been thrusting spears.<ref>Churchill S.E. (2002) Of assegais and bayonets: Reconstructing prehistoric spear use. ''Evolutionary Anthropology'', 11, 185-186</ref> Still, a [[Levallois technique|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.<ref>Boëda E., Geneste J.M., Griggo C., Mercier N., Muhesen S., Reyss J.L., Taha A. & Valladas H. (1999) A Levallois point embedded in the vertebra of a wild ass (Equus africanus): Hafting, projectiles and Mousterian hunting. ''Antiquity'', 73, 394-402</ref> 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.<ref>Schwimmer, E.G. "Warfare of the Maori." ''Te Ao Hou: The New World'', #36, Sept 1961, pp. 51-53. [http://teaohou.natlib.govt.nz/teaohou/issue/Mao36TeA/c29.html]</ref> 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 [[flower]]s, and therefore being a form of [[ritual]] burial,<ref>{{cite journal | author= R. S. Solecki | year= 1975 | title = Shanidar IV, a Neanderthal flower burial in northern [[Iraq]] | journal = [[Science (journal)|]] | volume = 190 | issue = 28 | pages = 880}}</ref> has been questioned.<ref>{{cite journal | author= J. D. Sommer |year = 1999 | title = The Shanidar IV 'Flower Burial': A Reevaluation of Neanderthal Burial Ritual | journal = Cambridge Archæological Journal | volume = 9 | pages = 127–129}}</ref> 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 [[wisent|bison]] and [[aurochs]] bones, tools, and the [[pigment]] [[ochre]]. Neanderthals also performed many sophisticated tasks which are normally associated only with humans. For example, it is known that they controlled [[fire]], constructed complex shelters, and skinned animals. A trap excavated at [[La Cotte de St Brelade]] in [[Jersey]] gives testament to their intelligence and success as hunters <ref>A History of Britain, Richard Dargie (2007), p. 9</ref>. Particularly intriguing is a hollowed-out bear [[femur]] with holes which 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 hypothesized that it was a flute, while others believe it was created by accident through the chomping action of another bear. ''See: [[Divje Babe]]''. Pendants and other jewelry showing traces of ochre dye and of deliberate grooving have also been found[http://www.asa3.org/archive/ASA/200202/0560.html] with later finds, particularly in France but whether or not they were created by Neaderthals or traded to them by Cro-Magnons is a matter of controversy. ==Habitat and range== [[Image:Carte Neandertaliens.jpg|thumb|300px|Sites where typical Neanderthal fossils have been found.]] Early Neanderthals lived in the Last Glacial age for a span of about 100,000 years. Because of the damaging effects which the glacial period had on the Neanderthal sites, not much is known about the early species. Places where their remains are known include [[Portugal]], [[France]]<ref>[http://www.timesonline.co.uk/tol/life_and_style/court_and_social/the_hitch/article561030.ece Neanderthals and modern man shared a cave]</ref> and [[Spain]], as well as [[Great Britain|Britain]] <ref>A History of Britain, Richard Dargie (2007), p. 9</ref>, [[Germany]], [[Czech Republic]], [[Slovakia]], [[Croatia]], [[Greece]]<ref>[http://www.dur.ac.uk/news/newsitem/?itemno=6163 Ancient tooth provides evidence of Neanderthal movement]</ref> and [[Israel]]. 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 they occupied, since Middle-Palaeolithic looking artifacts have been found even further north, up to 60° on the Russian plain.<ref>{{cite journal | author = Pavlov, P., W. Roebroeks, and J. I. Svendsen | year = 2004 | title = The Pleistocene colonization of northeastern Europe: A report on recent research | journal = Journal of Human Evolution | volume = 47 | issue = 1-2 | pages = 3–17 | doi = 10.1016/j.jhevol.2004.05.002 }}</ref> Recent evidence has extended the Neanderthal range by about {{convert|1250|mi|km}} east into southern [[Siberia]]'s [[Altay Mountains]].<ref>[http://www.nytimes.com/2007/10/02/science/02nean.html?ref=science Fossil DNA Expands Neanderthal Range]</ref><ref>[http://news.nationalgeographic.com/news/2007/10/070930-neanderthals.html Neandertals Ranged Much Farther East Than Thought]</ref> ==Cannibalism or Ritual defleshing?== Neanderthals hunted large animals, such as the mammoth. Stone-tipped wooden spears were used for hunting and stone knives were used for butchering the animals. They are also believed to have gathered wild plants like their ancestors, ''[[Homo erectus]]''. However, they are believed to have practiced ritual defleshing. This hypothesis has been represented after researchers found marks on Neanderthal bones similar to the bones of a dead deer eaten by Neanderthals. Intentional burial and the inclusion of grave goods are the most typical representations 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 ritual defleshing. 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.<ref name="livescience">{{cite web |title=Neanderthals Were Cannibals, Study Confirms |url=http://www.livescience.com/humanbiology/061204_neanderthal_lifestyle.html |author=Andrea Thompson |date=2006-12-04 |work=Health SciTech |publisher=LiveScience}}</ref> 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 which 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 defleshing, although whether this was for nutritional or ritual purposes cannot be determined with certainty.<ref>{{cite journal | author = Pathou-Mathis M | year = 2000 | title = Neanderthal subsistence behaviours in Europe | journal = International Journal of Osteoarchaeology | volume = 10 | pages = 379–395 | doi = 10.1002/1099-1212(200009/10)10:5<379::AID-OA558>3.0.CO;2-4}}</ref> * 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.<ref>{{cite journal | author = Defleur A, White T, Valensi P, Slimak L, Cregut-Bonnoure E | year = 1999 | title = Neanderthal cannibalism at Moula-Guercy, Ardèche, France | journal = Science | volume = 286 | pages = 128–131 | doi = 10.1126/science.286.5437.128 <!--Retrieved from CrossRef by DOI bot-->}}</ref> 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 1]] ‘Old Man’), the femur ([[La Ferrassie]] 1), fibulae (La Ferrassie 2 and [[Tabun, Israel|Tabun]] 1), spine ([[Kebara Cave|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.<ref>{{cite journal|author = T.D. Berger and E. Trinkaus|title = Patterns of trauma among Neadertals|journal = Journal of Archaeological Science|volume = 22|date= 1995|pages = 841–852|url = http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6WH8-4FRCV9P-F&_user=994540&_coverDate=11%2F30%2F1995&_fmt=full&_orig=search&_cdi=6844&view=c&_acct=C000050024&_version=1&_urlVersion=0&_userid=994540&md5=a958d2c59d5c6d934e9844f46f275e0e&ref=full|accessdate = 2007-06-28 | doi = 10.1016/0305-4403(95)90013-6 <!--Retrieved from CrossRef by DOI bot-->}}</ref> ===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|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|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 that 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). [[Image:Neanderthal child.jpg|thumb|right|Reconstruction of a NeanderthaI child from [[Gibraltar]] (Anthropological Institute, University of Zürich).]] ==The fate of the Neanderthals== Possible hypotheses for the fate of Neanderthals include the following: # Neanderthals evolved to a separate species which became extinct (see [[Neanderthal extinction hypotheses]]) and were replaced by early modern humans traveling from Africa.<ref>[http://english.pravda.ru/science/mysteries/24-10-2007/99419-genocide-0 First genocide of human beings occurred 30,000 years ago]</ref> # Neanderthals was a contemporary subspecies which incidentally bred with ''Homo sapiens'' and disappeared through absorption (see [[Neanderthal interaction with Cro-Magnons]]) # Neanderthals never split from ''Homo sapiens'' and most of their populations transformed into anatomically modern humans between 50-30 kya (see [[Multiregional origin of modern humans]]). According to the oldest view (#1), modern humans (''[[Homo sapiens]]'') began replacing Neanderthals around 45,000 years ago, as the [[Cro-Magnon]] people appeared in [[Europe]], pushing populations of Neanderthals into regional pockets, where they held on for thousands of years, such as modern-day [[Croatia]], [[Iberian peninsula|Iberia]], and the [[Crimean peninsula]].{{Clarifyme|date=January 2008}} The last traces of Neanderthal ''culture'' have been found around a [[Gorham's Cave|cave system]] on the remote south-facing coast of [[Gibraltar]], from 30,000 to 24,000 years ago. The validity of such an extensive period of cornered Neanderthal groups is recently questioned. There is no longer certainty regarding the identity of the humans who produced the [[Aurignacian]] culture, even though the presumed westward spread of anatomically modern humans (AMHs) across Europe is still based on the controversial first dates of the Aurignacian. Currently, the oldest European anatomically modern Homo sapiens is represented by a robust modern human mandible discovered at Pestera cu Oase (south-west Romania), dated to 34–36 kya (thousand years ago). Human skeletal remains from the German site of Vogelherd, so far regarded the best association between anatomically modern Homo sapiens and [[Aurignacian]] culture, were revealed to represent intrusive Neolithic burials into the Aurignacian levels and subsequently all the key Vogelherd fossils are now dated to 3.9–5.0 thousand years ago instead.<ref>Conard, N.J. et al. (2004) Unexpectedly recent dates for human remains from Vogelherd. Nature 430, 198–201 [http://www.nature.com/nature/journal/v430/n6996/full/nature02690.html]; [http://www.urgeschichte.uni-tuebingen.de/fileadmin/downloads/Conard/vogelherdNature04.pdf]</ref> As for now, the expansion of the first anatomically modern humans into Europe can't be located by diagnostic and well-dated anatomically modern human fossils "west of the Iron Gates of the Danube" before 32 kya.<ref>Trinkaus, E. (2005) Early modern humans. Annu. Rev. Anthropol. 34, 207–230 [http://www.pnas.org/cgi/content/full/103/46/17196?maxtoshow=&HITS=10&hits=10&RESULTFORMAT=&fulltext=trinkaus&searchid=1&FIRSTINDEX=0&sortspec=date&resourcetype=HWCIT]</ref> Moreover, researchers have recently found in Pestera Muierii, Romania, remains of European humans from 30 kya who possessed mostly diagnostic "modern" anatomical features, but ''also'' had distinct Neanderthal features not present in ancestral modern humans in Africa, including a large bulge at the back of the skull, a more prominent projection around the elbow joint, and a narrow socket at the shoulder joint. Analysis of one skeleton's shoulder showed that these humans, like Neanderthal, did not have the full capability for throwing spears.<ref name=Cosmos200611>[http://www.cosmosmagazine.com/node/814] Humans and Neanderthals interbred, 2 November 2006 by Jacqui Hayes, Cosmos Online</ref> Consequently, the exact nature of biological and cultural interactions between Neanderthals and other human groups between 50 and 30 thousand years ago is currently hotly contested. A new proposal resolves the issue by taking the Gravettians rather than the Aurignacians as the anatomically modern humans which contributed to the post-30 kya Eurasian genetic pool.<ref>Rapid ecological turnover and its impact on Neanderthal and other human populations - Clive Finlayson and Jose´ S. Carrión, Trends in Ecology & Evolution, Volume 22, Issue 4 , April 2007, Pages 213-222 [http://www.jscarrion.com/pdf/finlayson_carrion.pdf] </ref> Correspondingly, the human skull fragment found at the Elbe River bank at Hahnöfersand near Hamburg was once radiocarbon dated to 36,000 years ago and seen as possible evidence for the intermixing of Neanderthals and anatomically modern humans. It is now dated to the more recent [[Mesolithic]].<ref name=Terberger>From the First Humans to the Mesolithic Hunters in the Northern German Lowlands, Current Results and Trends - THOMAS TERBERGER. From: Across the western Baltic, edited by: Keld Møller Hansen & Kristoffer Buck Pedersen, 2006, ISBN 87-983097-5-7, Sydsjællands Museums Publikationer Vol. 1 [http://www.uni-greifswald.de/~histor/~ufg/mitarbeiter/terberger/Terberger1_LoRes.pdf]</ref> Modern human findings in Abrigo do [[Lagar Velho]], [[Portugal]] of 24,500 years ago, allegedly featuring Neanderthal admixtures, have been published.<ref>The early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern human emergence in Iberia - Cidália Duarte, João Maurício, Paul B. Pettitt, Pedro Souto, Erik Trinkaus, Hans van der Plicht, and João Zilhão, PNAS Vol. 96, Issue 13, 7604-7609, June 22, 1999 [http://www.pnas.org/cgi/content/full/96/13/7604]</ref> 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 years ago shows accumulating evidence that they present a variable mosaic of derived modern human, archaic human, and Neanderthal features.<ref>An early modern human from the Petera cu Oase, Romania - Erik Trinkaus et al, PNAS | September 30, 2003 | vol. 100 | no. 20 | 11231-11236 [http://www.pnas.org/cgi/content/full/100/20/11231]</ref><ref>Early modern humans from the Petera Muierii, Baia de Fier, Romania - Andrei Soficaru*, Adrian Dobo, and Erik Trinkaus, PNAS | November 14, 2006 | vol. 103 | no. 46 | 17196-17201 [http://www.pnas.org/cgi/content/abstract/103/46/17196]</ref><ref>{{cite journal | last = Trinkaus | first = Erik | title = European early modern humans and the fate of the Neandertals | journal = PNAS | volume = 104 | issue = 18 | pages = 7367–7372 | date = [[2007-04-23]] | url = http://www.pnas.org/cgi/content/full/104/18/7367 | doi = 10.1073/pnas.0702214104 | accessdate = 2008-05-26 | pmid = 17452632 }}Reprint available at: [http://www.pnas.org/cgi/reprint/0702214104v1]</ref> On the other hand, a [[mtDNA]] analysis has shown no evidence for Neanderthal contributions to the gene pool of modern humans.<ref>[http://www.eva.mpg.de/english/ presskits/mtDNANeandertalarticle.pdf Krings et al., ''Neandertal DNA sequences and the origin of modern humans'' Cell. 1997 Jul 11;90(1):19-30.]; [http://sciencenow.sciencemag.org/cgi/content/full/2004/316/2 No Neandertals in the Gene Pool], Science (2004).</ref> The authors of the study concede that this does not exclude Neanderthal contributions of other genes. They nevertheless argue that other genetic and morphological data also suggest little or no Neanderthal contribution. ==Genome== {{see|Neanderthal Genome Project}} While previous investigations concentrated on mitochondrial DNA (mtDNA) &mdash; which, due to strictly matrilineal inheritance and subsequent vulnerability to [[genetic drift]], is of limited value to disprove interbreeding &mdash; 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. The [[Neanderthal genome]] very likely is roughly the size of the [[human genome]], three-billion base pairs, and probably shares most of its [[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.<ref>{{cite news | title = Neanderthal genome project launches | url = http://www.msnbc.msn.com/id/13955661/ | accessdate = 2006-08-22 | author = Moulson, Geir | publisher = Associated Press | work = [http://www.msnbc.msn.com/ MSNBC.com]}}</ref> DNA researcher [[Svante Pääbo]] has tested more than 70 Neanderthal specimens and found only one which 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 (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.<ref>[http://www.nytimes.com/2006/11/15/science/16neanderthalcnd.html New Machine Sheds Light on DNA of Neanderthals - New York Times<!-- Bot generated title -->]</ref> A 2007 study pushes the point of divergence back to around 800,000 years ago.<ref>{{cite journal | journal = Science | date = [[2007-05-18]] | volume = 316 | issue = 5827 | pages = 967 | title = ANCIENT DNA: No Sex Please, We're Neandertals | author = Pennisi, Elizabeth | accessdate = 2007-05-18 | doi = 10.1126/science.316.5827.967a | pmid = 17510332}}</ref> [[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.<ref>[http://www.cnn.com/2006/TECH/science/11/15/neanderthal.ap/index.html Neanderthal bone gives DNA clues]</ref><ref>[http://www.msnbc.msn.com/id/15732243/ Scientists decode Neanderthal genes]</ref> 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.” On the other hand, a 2006 investigation suggested 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.<ref name=Plagnol>Plagnol V, Wall JD: Possible ancestral structure in human populations. PLoS Genet 2006, 2:e105.[http://genetics.plosjournals.org/perlserv/?request=get-document&doi=10.1371/journal.pgen.0020105.eor]</ref> Plagnol and Wall arrived at this result by first calculating a "null model" of genetic characteristics which 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| 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.<ref>Review: Archaic admixture in the human genome - Jeffrey D Wall and Michael F Hammer, Elsevier Ltd., 2006 {{PDFlink|[http://www.aseanbiotechnology.info/Abstract/21020590.pdf]|90.0&nbsp;[[Kibibyte|KiB]]<!-- application/pdf, 92212 bytes -->}}</ref> A main proponent of the interbreeding hypothesis is [[Erik Trinkaus]] of [[Washington University]]. In a 2006 study published in ''[[Proceedings of the National Academy of Sciences]]'', Trinkaus and his co-authors report a possibility that Neanderthals and humans did interbreed. 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."<ref>[http://www.cosmosmagazine.com/node/814 Humans and Neanderthals interbred]</ref> and "From my perspective, the replacement vs. continuity debate that raged through the 1990s is now dead".<ref>[http://news.yahoo.com/s/hsn/20061030/hl_hsn/modernhumansneanderthalsmayhaveinterbred Modern Humans, Neanderthals May Have Interbred]</ref> There is a possibility that Neanderthals and Cro-Magnons interbred but left little genetic evidence of that. That is because there is an ongoing debate about whether the hunter-gatherers of the middle stone age started farming when they came in contact with agriculture, or were completely replaced by the farmers moving in from the Middle East. If modern Europeans are mainly descendents of these farming people with little or no genetic input of the hunter gatherers of the middle stone age, then possible interbreeding between them and the Neanderthals would not have had a great effect on the modern gene-pool.<ref>[http://www.nature.com/hdy/journal/v97/n2/full/6800852a.html Where do Europeans come from?]</ref> The case for interbreeding recently revived by studies that claim signs of admixture ([[introgression]]), finding unusually deep genealogies in highly divergent [[clades]] (genetic branches). The genetic variation at the [[microcephalin]] gene, a critical regulator of brain size whose loss-of-function by damaging mutations may also cause primary [[microcephaly]], is claimed to be the most compelling evidence of admixture thus far. One type of the gene,dubbed haplogroup D having an exceptionally high worldwide frequency (~70%), was shown to have a remarkably young coalescence age to its most recent common ancestor ~37,000 years ago. The remaining types (non-D) coalesce to ~990,000 years ago, while the separation time between D and non-D was estimated at ~1,100,000 years ago. An evolutionary advance was assumed, even though positive selection was never as all-decisive as to wipe out the remaining 30% of non-D haplogroups (in which case no introgression could have been suggested) and as for now, a measurable genetic advance has not been attested.<ref>{{cite journal | author = Mekel-Bobrov, N., ''et al.'' | title = The ongoing adaptive evolution of ASPM and Microcephalin is not explained by increased intelligence | year = 2007 | journal = [[Hum. Mol. Genet.]] | pages = adv. access | doi = 10.1093/hmg/ddl487 | volume = 16 | pmid = 17220170}} </ref> Both the worldwide frequency distribution of the D allele, exceptionally high outside of Africa but low in sub-Saharan Africa (29%) that suggests involvement of an archaic Eurasian population, and current estimates of the divergence time between modern humans and Neanderthals based on mitochondrial DNA (mtDNA), are in favor of the Neanderthal lineage as the most likely archaic Homo population from which introgression into the modern human gene pool took place. <ref>{{cite journal | last = Evans | first = Patrick D. | coauthors = et al. | title = Evidence that the adaptive allele of the brain size gene microcephalin introgressed into Homo sapiens from an archaic Homo lineage | journal = PNAS | volume = 10 | issue = 48 | pages = 18178–18183 | date = [[2006-11-07]] | url = http://www.pnas.org/cgi/content/full/103/48/18178 | doi = 10.1073/pnas.0606966103 | accessdate = 2008-05-26 | pmid = 17090677 }}</ref><ref>{{cite journal | last = Evans | first = Patrick D. | coauthors = et al. | title = Microcephalin, a Gene Regulating Brain Size, Continues to Evolve Adaptively in Humans | journal = Science | volume = 309 | issue = 5741 | pages = 1717–1720 | date = [[2005-09-09]] | url = http://www.sciencemag.org/cgi/content/abstract/309/5741/1717 | doi = 10.1126/science.1113722 | accessdate = | pmid = 16151009 }}</ref> ==Key dates== *1829: Neanderthal skulls were discovered in [[Engis]], [[Belgium]]. *1848: Skull of an ancient human was found in [[Gorham's Cave|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, Germany|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 Cave|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: [[Electron Spin Resonance|ESR]] dates showed that the [[Tabun, Israel|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== {{Refimprove|section|date=April 2008}} {{main|Neanderthals in popular culture}} 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" is also used. Counterbalancing this are sympathetic literary portrayals of Neanderthals, as in the novel ''[[The Inheritors (William Golding)|The Inheritors]]'' by [[William Golding]], [[Isaac Asimov]]'s ''[[The Ugly Little Boy]]'', and [[Jean M. Auel]]'s ''[[Earth's Children]]'' series, though Auel repeatedly compares Neanderthals to modern humans unfavorably within the series, showing them to be less advanced in nearly every facet of their lives. Instead she gives them access to a 'race memory' and uses it to explain both their cultural richness and eventual stagnation. A more serious treatment is offered 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. ''[[The Neanderthal Parallax]]'', a trilogy of science fiction novels dealing with neanderthals, written by [[Robert J. Sawyer]], explores a scenario where neanderthals are seen as a distinct species from humans and survive in a parallel universe version of earth. The novels explore what happens when they, having developed a sophisticated technological culture of their own, open a portal to this version of the earth. The three novels are titled ''Hominids'', ''Humans'', and ''Hybrids'', respectively, and all form essentially one story. ==See also== * [[List of Neanderthal sites]] * [[Neanderthal Genome Project]] * [[Neanderthal extinction hypotheses]] * [[List of fossil sites]] ''(with link directory)'' * [[List of primate and hominin fossils]] ''(with images)'' * [[Physical anthropology]] * [[Caveman]] * [[Abrigo do Lagar Velho]] - More about "the Lapedo child" * [[Almas (cryptozoology)|Almas: wild man of Mongolia]] * ''[[Homo floresiensis]]'' * [[Pleistocene megafauna]] ==Footnotes== {{reflist|2}} Solecki, Ralph S. "Shanidar." Grolier Multimedia Encyclopedia. 2007. Grolier Online. 25 Nov. 2007 <http://gme.grolier.com/cgi-bin/article?assetid=0264140-0>. ==References== * Derev’anko, Anatoliy P. 1998 The Paleolithic of Siberia. New Discoveries and Interpretations. University of Illinois Press, Urbana. * C. David Kreger ([[2000-06-30]]) [http://www.archaeologyinfo.com/homoneaderthalensis.htm Homo Neanderthalensis] * Dennis O'Neil ([[2004-12-06]]) [http://anthro.palomar.edu/homo2/mod_homo_2.htm Evolution of Modern Humans Neandertals] retrieved 12/26/2004 * Fink, Bob (1997) [http://www.greenwych.ca/fl-compl.htm ''The Neanderthal Flute...''] (Greenwich, Canada) ISBN 0-912424-12-5 * Hickmann, Kilmer, Eichmann (ed.) (2003) [http://www.greenwych.ca/studies.htm ''Studies in Music Archaeology III''] International Study Group on Music Archaeology's 2000 symposium. ISBN 3-89646-640-2 *{{cite journal|author=Serre ''et al.''|year=2004|title=No evidence of Neandertal mtDNA contribution to early modern humans|journal=PLoS Biology|volume=2|issue=3|pages=313–7|pmid=15024415 | doi = 10.1371/journal.pbio.0020057 <!--Retrieved from CrossRef by DOI bot-->}} * {{cite journal|author=Eva M. Wild, Maria Teschler-Nicola, Walter Kutschera, Peter Steier, Erik Trinkaus & Wolfgang Wanek|year=2005|month=05|day=19|title=Direct dating of Early Upper Palaeolithic human remains from [[Mladeč]]|journal=[[Nature (journal)|Nature]]|volume=435|pages=332–5|doi=10.1038/nature03585}} [http://www.nature.com/nature/journal/v435/n7040/full/nature03585.html link for ''Nature'' subscribers] *Boë, Louis-Jean, Jean-Louis Heim, Kiyoshi Honda and Shinji Maeda. (2002) "The potential Neandertal vowel space was as large as that of modern humans." Journal of Phonetics, Volume 30, Issue 3, July 2002, Pages 465-484 *Lieberman, Philip. (2007). "Current views on Neanderthal speech capabilities: A reply to Boe et al. (2002)". Journal of Phonetics, Volume 35, Issue 4, Pages 552-563. * [http://jqjacobs.net/anthro/paleo/neanderthal.html Neanderthal DNA Sequencing] ==External links== {{commonscat|Homo neanderthalensis}} {{Wikibooks|Introduction to Paleoanthropology}} {{wikispecies|Homo neanderthalensis}} * [http://www.eva.mpg.de/ Max Planck Institute for Evolutionary Anthropology] * [http://www.mnh.si.edu/anthro/humanorigins/ha/neand.htm Smithsonian] * [http://www.archaeologyinfo.com/homoneaderthalensis.htm Archaelogy Info] * [http://www.mnsu.edu/emuseum/biology/humanevolution/neanderthalensis.html MNSU] * [http://www.cosmosmagazine.com/node/814 "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) * [http://news.bbc.co.uk/hi/english/sci/tech/newsid_323000/323657.stm 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) :* [http://news.bbc.co.uk/2/hi/science/nature/2884801.stm 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) * [http://www.geocities.com/veldwezelt_2002/index.html GeoCities.com] - 'The Neanderthal Sites at [[Veldwezelt-Hezerwater]], [[Belgium]]' * [http://www.greenwych.ca/fl-compl.htm Greenwych.ca] - 'Neanderthal Flute: Oldest Musical Instrument's 4 Notes Matches 4 of Do, Re, Mi Scale - Evidence of Natural Foundation to [[Diatonic scale|Diatonic Scale]] (oldest known musical instrument), [[Greenwich Publishing]] :* [http://www.greenwych.ca/chewchip.htm Greenwych.ca] - 'Chewed or Chipped? Who Made the Neanderthal Flute? Humans or Carnivores?' Bob Fink, Greenwich Publishing (March, 2003) * [http://sapphire.indstate.edu/~ramanank/ IndState.edu] - 'Neanderthals: A Cyber Perspective', Kharlena María Ramanan, [[Indiana State University]] (1997) * [http://www.krapina.com/neandertals/index_en.htm Krapina.com] - '[[Krapina]]: The World's Largest Neanderthal Finding Site' * [http://www.neanderthal.de/ Neanderthal.de] - 'Neanderthal Museum' * [http://www.isogg.org/neanderthaldna.htm Neanderthal DNA] - 'Neanderthal DNA' Includes Neanderthal mtDNA sequences * [http://www.newanimal.org/neandertals.htm The Cryptid Zoo] - 'Neanderthals and Neanderthaloids in Cryptozoology' (modern sightings promoted by the [[pseudoscience]] of [[cryptozoology]]) * [http://www.ifi.unizh.ch/staff/zolli/CAP/comparingNeand.htm UniZH.ch] - 'Comparing Neanderthals and modern humans: Neanderthals differ from anatomically modern Homo sapiens in a suite of cranial features' (cranio-facial reconstructions), Institut für Informatik der [[Universität Zürich]] * [http://community.webshots.com/photo/407833162/1409245320063963747UfDUTi WebShots.com] - '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]]) * [https://nespos-live01.pxpgroup.com/display/openspace/Home interactive database on the archaeology and anthropology of Neanderthals] * [http://www.newswise.com/p/articles/view/510666/ Did free trade cause the extinction of Neanderthals?] * [http://www.pnas.org/cgi/content/abstract/99/25/16122 Mitochondrial and nuclear DNA can show conflicting phylogenetic histories] * [http://eja.sagepub.com/cgi/content/abstract/4/3/385 Neanderthal manifactured pitch] * [http://anthro.amnh.org/anthropology/research/recon_wiki.htm ''Homo neanderthalensis'' reconstruction] - Electronic articles published by the Division of Anthropology, American Museum of Natural History. * [http://www.cbsnews.com/stories/2006/09/13/ap/tech/mainD8K43TS81.shtml CBS article on latest scientific speculation about Neanderthals in Gibraltar.] *[http://www.cnn.com/2006/TECH/science/11/15/neanderthal.ap/index.html Neanderthal bone gives DNA clues] *[http://www.msnbc.msn.com/id/15732243/ Scientists decode Neanderthal genes] *[http://www.livescience.com/history/050310_neanderthal_reconstruction.html Scientists Build 'Frankenstein' Neanderthal Skeleton] *[http://cosmiclog.msnbc.msn.com/archive/2006/06/06/182.aspx A NEANDERTHAL'S DNA TALE] * [http://www.boneandstone.com 'Bone and Stone'] A digitally enhanced single frame philatelic exhibit dedicated to the Neanderthal. *[http://www.nature.com/nature/journal/v444/n7120/abs/nature05314.html How Neanderthal molar teeth grew] *[http://worldmuseumofman.org/mousterianeuropeanartifacts1.htm Mousterian Tools of Neanderthals From Europe - World Museum of Man] * [http://www.aulis.com/twothirds2.htm The Way We Are] * [http://www.panoramio.com/photo/322489 Link to picture of the Neanderthal trace near Gediz River in Turkey] * [http://www.panoramio.com/photo/5579262 Link to Cross-Eyed stereoview of Neanderthal fossil cast in Chicago Field Museum] {{Human Evolution}} [[Category:Early hominids]] [[Category:European archaeology]] [[Category:Pleistocene extinctions]] [[Category:Pleistocene mammals]] [[Category:Recent single origin hypothesis]] [[Category:German loanwords]] [[Category:Paleoanthropology]] [[Category:Human evolution]] {{Link FA|hu}} {{Link FA|fr}} [[als:Neandertaler]] [[ar:نياندرتال]] [[roa-rup:Homo sapiens neanderthalensis]] [[ast:Homo sapiens neanderthalensis]] [[bg:Неандерталец]] [[ca:Home de Neandertal]] [[cs:Homo neanderthalensis]] [[cy:Dyn Neanderthal]] [[da:Neandertaler]] [[de:Neandertaler]] [[et:Neandertallane]] [[es:Homo neanderthalensis]] [[eo:Neandertala homo]] [[eu:Neandertaleko gizaki]] [[fa:نئاندرتال]] [[fr:Homme de Néandertal]] [[gl:Home de Neanderthal]] [[ko:네안데르탈인]] [[hr:Neandertalac]] [[is:Neanderdalsmenn]] [[it:Homo neanderthalensis]] [[he:אדם ניאנדרתלי]] [[la:Homo neanderthalensis]] [[lb:Neandertaler]] [[lt:Neandertalietis]] [[hu:Neandervölgyi ember]] [[mk:Неандерталец]] [[nl:Neanderthaler]] [[ja:ネアンデルタール人]] [[no:Neandertaler]] [[pl:Neandertalczyk]] [[pt:Neandertal]] [[ro:Omul de Neanderthal]] [[ru:Неандерталец]] [[scn:Homo neanderthalensis]] [[simple:Neanderthal]] [[sd:نينڊرٿل]] [[sk:Človek neandertálsky]] [[sl:Neandertalec]] [[sr:Неандерталац]] [[sh:Neandertalac]] [[fi:Neandertalinihminen]] [[sv:Neandertalmänniska]] [[ta:நியண்டர்தால் மனிதன்]] [[tr:Neandertal adamı]] [[uk:Неандертальці]] [[zh-yue:尼安德特人]] [[zh:尼安德特人]]