Neanderthal

Neanderthal Temporal range: Middle–Late Pleistocene 0.43/0.25–0.04 Ma PreЄ Є O S D C P T J K Pg N ↓
Late Neanderthal skull (La Chapelle-aux-Saints 1)
An approximate reconstruction of a Neanderthal skeleton (The central ribcage, including the sternum, and parts of the pelvis are from modern humans.)
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Primates
Suborder:	Haplorhini
Infraorder:	Simiiformes
Family:	Hominidae
Subfamily:	Homininae
Tribe:	Hominini
Genus:	Homo
Species:	H. neanderthalensis
Binomial name
Homo neanderthalensis King, 1864
Known Neanderthal range in Europe (blue), Southwest Asia (orange), Uzbekistan (green), and the Altai mountains (violet).
Synonyms
Homo stupidus (Haeckel 1866)[1] Homo mousteriensis (Klaatsch 1909)[2] Palaeoanthropus neanderthalensis[3]

Neanderthals (UK: /niˈændərˌtɑːl/, also US: /neɪ-, -ˈɑːn-, -ˌtɔːl, -ˌθɔːl/;^[4] Homo neanderthalensis or Homo sapiens neanderthalensis^[5]) are an extinct species or subspecies of archaic humans in the genus Homo, who lived in Eurasia until 40,000 years ago.^[6][7][8][9]

Currently earliest fossils of Neanderthals in Europe are dated between 430,000 to 450,000 years ago, and thereafter Neanderthals expanded into Southwest and Central Asia. They are known from numerous fossils, as well as stone tool assemblages. Almost all assemblages younger than 160,000 years are of the so-called Mousterian techno-complex, which is characterised by tools made out of stone flakes.^[10] The type specimen is Neanderthal 1, found in Neander Valley in the German Rhineland, in 1856.

Compared to modern humans, Neanderthals were stockier, with shorter legs and bigger bodies. In conformance with Bergmann's rule, this likely was an adaptation to preserve heat in cold climates. Male and female Neanderthals had cranial capacities averaging 1,600 cm³ (98 cu in) and 1,300 cm³ (79 cu in), respectively,^[11] within the range of the values for anatomically modern humans.^[12] Males stood 164 to 168 cm (65 to 66 in) and females 152 to 156 cm (60 to 61 in) tall.^[13]

Since 2010, there has been growing evidence for admixture between Neanderthals and anatomically modern humans. This admixture is reflected in the genomes of modern European and Asian populations, but not in the genomes of most sub-Saharan Africans. This suggests that interbreeding between Neanderthals and anatomically modern humans took place after the recent "out of Africa" migration, likely between 60,000 and 40,000 years ago.^[14]

Name and classification

Neanderthals are named after one of the first sites where their fossils were discovered in the mid-19th century in the Neander Valley, just east of Düsseldorf, at the time in the Rhine Province of the Kingdom of Prussia (now in Northrhine-Westphalia, Germany). The valley itself was named for Joachim Neander, Neander being the graecicized form of the surname Neumann ("new man").^[15] The German spelling of Thal "Valley" was current in the 19th century (contemporary German Tal).^{[lower-alpha 1][1]}

Neanderthal 1 was known as the "Neanderthal cranium" or "Neanderthal skull" in anthropological literature, and the individual reconstructed on the basis of the skull was occasionally called "the Neanderthal man".^[16] The binomial name Homo neanderthalensis—extending the name "Neanderthal man" from the individual type specimen to the entire group—was first proposed by the Anglo-Irish geologist William King in 1864, although that same year King changed his mind and thought that the Neanderthal fossil was distinct enough from humans to warrant a separate species.^[17] King's name had priority over the proposal put forward in 1866 by Ernst Haeckel, Homo stupidus.^[1] Popular English usage of "Neanderthal" as shorthand for "Neanderthal man", as in "the Neanderthals" or "a Neanderthal", emerged in the popular literature of the 1920s.^[18]

Since the historical spelling -th- in German represents the phoneme /t/ or /tʰ/, not the fricative /θ/, standard British pronunciation of "Neanderthal" is with /t/ (IPA: /niːˈændərtɑːl/).^{[19][20][21][22]} Because of the usual sound of the digraph ⟨th⟩ in English, "Neanderthal" is also pronounced with the voiceless fricative /θ/, at least in "layman's American English" (as /niːˈændərθɔːl/).^[23]

The spelling Neandertal is occasionally seen in English, even in scientific publications. Since "Neanderthal", or "Neandertal", is a common name, there is no authoritative prescription on its spelling, unlike the spelling of the binominal name H. neanderthalensis, which is predicated by King 1864.^[24] The common name in German is always invariably Neandertaler (lit. "of the valley of Neander"), not Neandertal, but the spelling of the name of the Neander Valley itself (Neandertal vs. Neanderthal) has been affected by the species name, the names of the Neanderthal Museum and of Neanderthal station persisting with pre-1900 orthography.^[25]

Anatomical comparison of skulls of Homo sapiens (left) and Homo neanderthalensis (right)
(in Cleveland Museum of Natural History)
Features compared are the braincase shape, forehead, browridge, nasal bone, projection, cheek bone angulation, chin and occipital contour.

Ever since the discovery of the Neanderthal fossils, expert opinion has been divided as to whether Neanderthals should be considered a separate species (Homo neanderthalensis) or a subspecies (Homo sapiens neanderthalensis) relative to modern humans.^[26][27][28] Pääbo (2014) described such "taxonomic wars" as unresolveable in principle, "since there is no definition of species perfectly describing the case."^[5] The question depends on the definition of Homo sapiens as a chronospecies, which has also been in flux throughout the 20th century. Authorities preferring classification of Neanderthals as subspecies have introduced the subspecies name Homo sapiens sapiens for the anatomically modern Cro-Magnon population which lived in Europe at the same time as Neanderthals, while authorities preferring classification as separate species use Homo sapiens as equivalent to "anatomically modern humans".

During the early 20th century, a prevailing view of Neanderthals as "simian", influenced by Arthur Keith and Marcellin Boule,^[29] tended to exaggerate the anatomical differences between Neanderthals and Cro Magnon. Beginning in the 1930s, revised reconstructions of Neanderthals increasingly emphasized the similarity rather than differences from modern humans. From the 1940s throughout the 1970s, it was increasingly common to use the subspecies classification of Homo sapiens neanderthalensis vs. Homo sapiens sapiens.^[30] The hypothesis of "multiregional origin" of modern man was formulated in the 1980s on such grounds, arguing for the presence of an unbroken succession of fossil sites in both Europe and Asia.^[31] Hybridization between Neanderthals and Cro Magnon had been suggested on skeletal and craniological grounds since the early 20th century, and found increasing support in the later 20th century,^[32] until Neanderthal admixture was found to be present in modern populations genetics in the 2010s.

Evolution

Stage 1: Early pre-Neanderthal, possibly Homo erectus, (Tautavel Man, 450 ka)

Stage 2: Archaic Neanderthal (Miguelón, 430 ka)

Stage 3: Intermediate Neanderthal (Saccopastore I, 130 ka)

Stage 4: Classic European Neanderthal (La Chapelle-aux-Saints 1, 50 ka)

The evolution of Neanderthals according to the accretion model.

Both Neanderthals and anatomically modern humans were initially thought to have evolved from Homo erectus between 300,000 and 200,000 years ago. H. erectus had emerged around 1.8 million years ago, and had long been present, in various subspecies throughout Eurasia.

The divergence time between the Neanderthal and archaic Homo sapiens lineages is estimated at between 800,000 and 400,000 years ago. The more recent time depth has been suggested by Endicott et al. (2010)^[33] and Rieux et al. (2014).^[34]

The time of divergence between archaic Homo sapiens and ancestors of Neanderthals and Denisovans caused by a population bottleneck of the latter was dated at 744,000 years ago, combined with repeated early admixture events and Denisovans diverging from Neanderthals 300 generations after their split from Homo sapiens, was calculated by Rogers et al. (2017).^[35][36]

Homo heidelbergensis, dated 600,000 to 300,000 years ago, has long been thought to be a likely candidate for the last common ancestor of the Neanderthal and modern human lineages. However, genetic evidence from the Sima de los Huesos fossils published in 2016 seems to suggest that H. heidelbergensis in its entirety should be included in the Neanderthal lineage, as "pre-Neanderthal" or "early Neanderthal", while the divergence time between the Neanderthal and modern lineages has been pushed back to before the emergence of H. heidelbergensis, to about 600,000 to 800,000 years ago, the approximate age of Homo antecessor.^[37]

The taxonomic distinctions between H. heidelbergensis and Neanderthals is mostly due to a fossil gap in Europe between 300,000 and 243,000 years ago (MIS 8). "Neanderthals", by conventions, are fossils which date to after this gap.^[38][39] The quality of the fossil record greatly increases from 130,000 years ago onwards.^[40] Specimens younger than this date make up the bulk of known Neanderthal skeletons and were the first whose anatomy was comprehensively studied.^[41][42] In morphological studies, the term "classic Neanderthal" may be used in a narrower sense for Neanderthals younger than 71,000 years old (MIS 4 and 3).^[38]

Habitat and range

Approximate Neanderthal range; pre-Neanderthal and early Neanderthal range shown in purple, late Neanderthal range in blue.

Sites where "classic Neanderthal" fossils (70–40 ka) have been found. Ice sheets of the last glacial maximum are indicated (partly ice-free during the Eemian interglacial)

Early Neanderthals, living before the Eemian interglacial (130 ka), are poorly known and come mostly from European sites. From 130 ka onwards, the quality of the fossil record increases dramatically. From then on, Neanderthal remains are found in Western, Central, Eastern, and Mediterranean Europe,^[43] as well as Southwest, Central, and Northern Asia up to the Altai Mountains in Siberia. No Neanderthal has ever been found outside Central to Western Eurasia, namely neither to the south of 30°N (Shuqba, Levant), nor east of 85°E (Denisova, Siberia).

The limit of their northern range appears to have been south of 53°N (Bontnewydd, Wales),^[44] although it is difficult to assess because glacial advances destroy most human remains, the Bontnewydd tooth being exceptional. Middle Palaeolithic artifacts have been found up to 60°N on the Russian plains.^[45]

Total Neanderthal effective population size has been estimated at close to 15,000 individuals (corresponding to a total population of roughly 150,000 individuals), living in small, isolated, inbred groups.^[46]

Anatomy

Neanderthal anatomy differed from modern humans in that they had a more robust build and distinctive morphological features, especially on the cranium, which gradually accumulated more derived aspects as it was described by Marcellin Boule,^[47] particularly in certain isolated geographic regions. These include shorter limb proportions, a wider, barrel-shaped rib cage, a reduced chin, sloping forehead, and a large nose, being at the modern human higher end in both width and length,^{[lower-alpha 2]} and started somewhat higher on the face than in modern humans and a characteristic occipital bun.^[39] Evidence suggests they were much stronger than modern humans, with particularly strong arms and hands, while they were comparable in height; based on 45 long bones from at most 14 males and 7 females, Neanderthal males averaged 164 to 168 cm (65 to 66 in) and females 152 to 156 cm (60 to 61 in) tall.^[13] Samples of 26 specimens in 2010 found an average weight of 77.6 kg (171 lb) for males and 66.4 kg (146 lb) for females.^[48] A 2007 genetic study suggested some Neanderthals may have had red hair and blond hair, along with a light skin tone.^[49]

Gregory Cochran and Henry Harpending, in the book The 10,000 Year Explosion, investigated whether it is accurate to depict Neanderthals as having hair patterns similar to anatomically modern humans. They concluded that:

In 2017, researchers using 3D reconstructions of nasal cavities and Computational Fluid Dynamics techniques have found that Neanderthals and modern humans both adapted their noses (independently and in a convergent way) to help breathe in cold and dry conditions.^[51]  The large nose seen in Neanderthals, as well as Homo heidelbergensis, affected the shape of the skull and the muscle attachments, and gave them a weaker bite force than in modern humans.^[52]

In The Spread of Modern Humans in Europe (2002) John F. Hoffecker writes:

A 2013 study of Neanderthal skulls suggests that their eyesight may have been better than that of modern humans, owing to larger eye sockets and larger areas of the brain devoted to vision.^[54]

Neanderthals are known for their large cranial capacity, which at 1,600 cm³ (98 cu in) is larger on average than that of modern humans. One study has found that drainage of the dural venous sinuses (low pressure blood vessels that run between the meninges and skull leading down through the skull) in the occipital lobe region of Neanderthal brains appears more asymmetric than other hominid brains.^[55] In 2008, a group of scientists produced a study using three-dimensional computer-assisted reconstructions of Neanderthal infants based on fossils found in Russia and Syria. It indicated that Neanderthal and modern human brains were the same size at birth, but that by adulthood, the Neanderthal brain was larger than the modern human brain.^[56] They had almost the same degree of encephalisation (i.e. brain-to-body-size ratio) as modern humans.^[57][58]

The Neanderthal skeleton suggests they consumed 100 to 350 kcal (420 to 1,460 kJ) more per day than male modern humans of 68.5 kg (151 lb) and females of 59.2 kg (131 lb).^[48]

A group of researchers from the University of Tübingen and University of Basel deduced with evidence that Neandertals had precision grasping hand movements rather than power grasping ones as previously thought, based on studying the bones and muscles structure of their hands.^[59]

A 2018 study which examined dental remains from the sites of Fontana Fanuccio, located 50km southeast of Rome, and Visogliano, located 18km northwest of Trieste (450 km apart in central and northeastern Italy), concluded that Neanderthal dental features had evolved by around 450,000 years ago during the Middle Pleistocene epoch.^[60]

Behaviour

Levallois point – Beuzeville, France

Neanderthals made stone tools, used fire,^[61] and were hunters. The consensus on their behaviour ends there. It had long been debated whether Neanderthals were hunters or scavengers,^[39] but the discovery of the pre-Neanderthal Schöningen wooden spears in Germany helped settle the debate in favour of hunting. Most available evidence suggests they were apex predators,^[62] and fed on red deer, reindeer, ibex, wild boar, aurochs and on occasion mammoth, straight-tusked elephant and rhinoceros.^{[39][63][64][65]} They appear to have occasionally used vegetables as fall-back food,^[66] revealed in the 2000s and 2010s by isotope analysis of their teeth and study of their coprolites (fossilised faeces).^[64] Dental analysis of specimens from Spy, Belgium and El Sidrón, Spain^[67] in 2017 suggested that these Neanderthals had a wide-ranging diet, and that those "from El Sidrón showed no evidence of meat eating" at all and seemed to have lived on "a mixture of forest moss, pine nuts and a mushroom known as split gill".^[68]

The size and distribution of Neanderthal sites, along with genetic evidence, suggests Neanderthals lived in much smaller and more sparsely distributed groups than anatomically-modern Homo sapiens.^[69][70] The bones of twelve Neanderthals were discovered at El Sidrón cave in northwestern Spain. They are thought to have been a group killed and butchered about 50,000 years ago. Analysis of the mtDNA showed that the three adult males belonged to the same maternal lineage, while the three adult females belonged to different ones. This suggests a social structure where males remained in the same social group and females "married out".^[71]

The bones of the El Sidrón group show signs of defleshing, suggesting that they were victims of cannibalism.^[71] The St. Césaire 1 skeleton discovered in 1979 at La Roche à Pierrot, France, showed a healed fracture on top of the skull apparently caused by a deep blade wound, suggesting interpersonal violence.^[72]

Claims that Neanderthals deliberately buried their dead, and if they did, whether such burials had any symbolic meaning,^{[40]:158–60} are heavily contested.^[73][74][75] The debate on deliberate Neanderthal burials has been active since the 1908 discovery of the well-preserved Chapelle-aux-Saints 1 skeleton in a small hole in a cave in southwestern France. In this controversy's most recent installment, a team of French researchers reinvestigated the Chapelle-aux-Saints cave and in January 2014 reasserted the century-old claim that the 1908 Neanderthal specimen had been deliberately buried,^[76] and this has in turn been heavily criticised.^[77] A 2018 open access paper discussed, in light of recent developments in the fields of paleogenetics and paleoanthropology, whether or not Neanderthals were rational. The authors' argument focuses on the genetic evidence that supports interbreeding with Homo sapiens, language acquisition (including the FOXP2 gene), archaeological signs of cultural development and potential for cumulative cultural evolution.^[78]

Interbreeding with archaic and modern humans

Chris Stringer's hypothesis of the family tree of genus Homo, published 2012 in Nature – the horizontal axis represents geographic location, and the vertical axis represents time in millions of years ago.^{[lower-alpha 3]}

An alternative to extinction is that Neanderthals were absorbed into the Cro-Magnon population by interbreeding. This would be counter to strict versions of the recent African origin theory, since it would imply that at least part of the genome of Europeans would descend from Neanderthals.

Extinction

According to a 2014 study by Thomas Higham and colleagues of organic samples from European sites, Neanderthals died out in Europe between 41,000 and 39,000 years ago.^{[lower-alpha 4]} New dating in Iberia, where Neanderthal dates as late as 28,000 years had been reported, suggests evidence of Neanderthal survival in the peninsula after 42,000 years ago is almost non-existent.^[8]

Anatomically modern humans arrived in Mediterranean Europe between 45,000 and 43,000 years ago, so the two different human populations shared Europe for several thousand years.^[6][138] The exact nature of biological and cultural interaction between Neanderthals and other human groups is contested.^[139]

Possible scenarios for the extinction of the Neanderthals are:

1. Neanderthals were a separate species from modern humans, and became extinct (because of climate change or interaction with modern humans) and were replaced by modern humans moving into their habitat between 45,000 and 40,000 years ago.^[140] Jared Diamond has suggested a scenario of violent conflict and displacement.^[141]
2. Neanderthals were a contemporary subspecies that bred with modern humans and disappeared through absorption (interbreeding theory).
3. Volcanic catastrophe: see Campanian Ignimbrite Eruption

mtDNA-based simulation of modern human expansion in Europe starting 1,600 generations ago. Neanderthal range in light grey^[142]

Coexistence with modern humans

Skeleton and restoration model of the La Ferrassie 1 Neanderthal man (National Museum of Nature and Science, Tokyo, 2013 photograph)

In November 2011 tests conducted at the Oxford Radiocarbon Accelerator Unit in England on what were previously thought to be Neanderthal baby teeth, which had been unearthed in 1964 from the Grotta del Cavallo in Italy, were identified as the oldest modern human remains discovered anywhere in Europe, dating from between 43,000 and 45,000 years ago.^[146] Given that the 2014 study by Thomas Higham of Neanderthal bones and tools indicates that Neanderthals died out in Europe between 41,000 and 39,000 years ago, the two different human populations shared Europe for as long as 5,000 years.^[143] Nonetheless, the exact nature of biological and cultural interaction between Neanderthals and other human groups has been contested.^[139]

Modern humans co-existed with them in Europe starting around 45,000 years ago and perhaps even earlier. Neanderthals inhabited that continent long before the arrival of modern humans. These modern humans may have introduced a disease that contributed to the extinction of Neanderthals, and that may be added to other recent explanations for their extinction. When Neanderthal ancestors left Africa potentially as early as over 800,000 years ago they adapted to the pathogens in their European environment, unlike modern humans who adapted to African pathogens. This transcontinental movement is known as the Out of Africa model. If contact between humans and Neanderthals occurred in Europe and Asia the first contact may have been devastating to the Neanderthal population, because they would have had little if any immunity to the African pathogens. More recent historical events in Eurasia and the Americas show a similar pattern, where the unintentional introduction of viral or bacterial pathogens to unprepared populations has led to mass mortality and local population extinction.^[147] The most well-known example of this is the arrival of Christopher Columbus to the New World, which brought and introduced foreign diseases when he and his crew arrived to a native population who had no immunity.

Anthropologist Pat Shipman, of Pennsylvania State University, suggested that domestication of the dog could have played a role in Neanderthals' extinction.^[148]

History of research

Engis 2, child (1829)

Gibraltar 1, female (1848)

Neanderthal 1, male (upper skull 1856, left-cheek 2000)

Spy 2 skull, sex unclear^[149] (1886)

Krapina 3, female (1899)

Neanderthal fossils were first discovered in 1829 in the Engis caves (the partial skull dubbed Engis 2), in what is now Belgium by Philippe-Charles Schmerling and the Gibraltar 1 skull in 1848 in the Forbes' Quarry, Gibraltar. These finds were not, at the time, recognized as representing an archaic form of humans.

The first discovery which was recognized as representing an archaic form of humans was made in August 1856, three years before Charles Darwin's On the Origin of Species was published. This was the discovery of the type specimen, Neanderthal 1, in a limestone quarry (Feldhofer Cave), located in Neandertal Valley in the German Rhineland, about 12 km (7 mi) east of Düsseldorf). The find consisted of a skull cap, two femora, three bones of the right arm, two of the left arm, parts of the left ilium, fragments of a scapula, and ribs. The workers who recovered the objects originally thought them to be the remains of a cave bear. However, they eventually gave the material to amateur naturalist Johann Carl Fuhlrott, who turned the fossils over to anatomist Hermann Schaaffhausen.^[150]

To date, the bones of over 400 Neanderthals have been found.^[151]

• 1829: A damaged skull of a Neanderthal child, Engis 2, is discovered in Engis, Netherlands (now Belgium).
• 1848: A female Neanderthal skull, Gibraltar 1, is found in Forbes' Quarry, Gibraltar, but its importance is not recognised.
• 1856: Limestone miners discover the Neanderthal-type specimen, Neanderthal 1, in Neandertal, western Prussia (now Germany).
• 1864: William King is the first to recognise Neanderthal 1 as belonging to a separate species, for which he gives the scientific name Homo neanderthalensis. He then changed his mind on placing it in the genus Homo, arguing that the upper skull was different enough to warrant a separate genus since, to him, it had likely been "incapable of moral and theistic conceptions."^[17]
• 1880: The mandible of a Neanderthal child is discovered in a secure context in Šipka cave, in the Austro-Hungarian Empire (now the Czech Republic), associated with cultural debris, including hearths, Mousterian tools, and bones of extinct animals.
• 1886: Two well-preserved Neanderthal skeletons are found at Spy, Belgium, making the hypothesis that Neanderthal 1 was only a diseased modern human difficult to sustain.^[152]
• 1899: Sand excavation workers find hundreds of fragmentary Neanderthal remains representing at least 12 and likely as much as 70 individuals on a hill in Krapina, in the Austro-Hungarian Empire (now Croatia).
• 1908: A very well preserved Neanderthal, La Chapelle-aux-Saints 1, is found in its eponymous site in France,^[153] said by the excavators to be a burial, a claim still heatedly contested.^[77][73][74] For historical reasons it remains the most famous Neanderthal skeleton.^[40]:15
• 1912: Marcellin Boule publishes his now discredited influential study of Neanderthal skeletal morphology based on La Chapelle-aux-Saints 1.
• 1953–1957: Ten Neanderthal skeletons are excavated in Shanidar Cave, Iraqi Kurdistan, by Ralph Solecki and colleagues.
• 1975: Erik Trinkaus's study of Neanderthal feet strongly argues that Neanderthals walked like modern humans.
• 1981: The site of Bontnewydd, Wales yielded an early Neanderthal tooth, the most north-western Neanderthal remain ever.
• 1987: Israeli Neanderthal Kebara 2 is dated (by TL and ESR) to 60,000 BP, thus later than the Israeli anatomically modern humans dated to 90,000 and 80,000 BP at Qafzeh and Skhul.
• 1997: Matthias Krings et al. are the first to amplify Neanderthal mitochondrial DNA (mtDNA) using a specimen from Feldhofer grotto in the Neander valley.^[15]
• 2005: The Max Planck Institute for Evolutionary Anthropology and associated institutions launch the Neanderthal genome project to sequence the Neanderthal nuclear genome.^[154]
• 2010: Discovery of Neanderthal admixture in the genome of modern populations.^{[114][116][117]}
• 2014: A comprehensive dating of Neanderthal bones and tools from hundreds of sites in Europe dates the disappearance of Neanderthals to 41,000 and 39,000 years ago.^[155]
• 2018: Report on the complete genomic sequence of Denisova 11, a first generation of Neanderthal-Denisovan hybrid.

Specimens

Notable European Neanderthals

La Ferrassie 1, skull cast

Le Moustier 1 in 1909

Le Moustier 1 in 2011

Shanidar 1, skull cast

Kebara 2

Teshik-Tash 1

Remains of more than 300 European Neanderthals have been found. For the most important, see List of human evolution fossils.

• Neanderthal 1: The first human bones recognised as showing a non-modern anatomy. Discovered in 1856 in a limestone quarry at the Feldhofer grotto in Neanderthal, Germany, they consist of a skull cap, the two femora, the three right arm bones, two left arm bones, the ilium, and fragments of a scapula and ribs.
• La Chapelle-aux-Saints 1: Called the Old Man, a fossilised 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 long before death, leading some to suggest he was cared for by others.
• La Ferrassie 1: A fossilised skull discovered in La Ferrassie, France, by R. Capitan in 1909. It is estimated to be 70,000 years old. Its characteristics include a large occipital bun, low-vaulted cranium and heavily worn teeth.
• Le Moustier 1: One of the rare nearly complete Neanderthal skeletons to be discovered, it was excavated by a German team in 1908, at Peyzac-le-Moustier, France. Sold to a Berlin museum, the post cranial skeleton was bombed and mostly destroyed in 1945, and parts of the mid face were lost sometime after then. The skull, estimated to be less than 45,000 years old, includes a large nasal cavity and a less developed brow ridge and occipital bun than seen in other Neanderthals. The Mousterian tool techno-complex is named after its discovery site.

In popular culture

Neanderthals have been portrayed in popular culture including appearances in literature, visual media and comedy. Early 20th century artistic interpretations often presented Neanderthals as beastly creatures, emphasising hairiness and rough, dark complexion.^[165]

See also

Notes

References

Journals

• Boë, Louis-Jean; Heim, Jean-Louis; Honda, Kiyoshi; Maeda, Shinji (2002). "The potential Neandertal vowel space was as large as that of modern humans". Journal of Phonetics. 30 (3): 465–84. doi:10.1006/jpho.2002.0170.
• Lieberman, Philip (October 2007). "Current views on Neanderthal speech capabilities: A reply to Boe et al. (2002)". Journal of Phonetics. 35 (4): 552–63. doi:10.1016/j.wocn.2005.07.002.
• Serre, David; Langaney, André; Chech, Mario; Teschler-Nicola, Maria; Paunovic, Maja; Mennecier, Philippe; Hofreiter, Michael; Possnert, Göran; Pääbo, Svante (2004). "No Evidence of Neandertal mtDNA Contribution to Early Modern Humans". PLoS Biology. 2 (3): e57. doi:10.1371/journal.pbio.0020057. PMC 368159. PMID 15024415.
• Wild, Eva M.; Teschler-Nicola, Maria; Kutschera, Walter; Steier, Peter; Trinkaus, Erik; Wanek, Wolfgang (2005). "Direct dating of Early Upper Palaeolithic human remains from Mladeč". Nature. 435 (7040): 332–35. Bibcode:2005Natur.435..332W. doi:10.1038/nature03585. PMID 15902255.
• Zilhão, João; Davis, Simon J. M.; Duarte, Cidália; Soares, António M. M.; Steier, Peter; Wild, Eva (2010). Hawks, John, ed. "Pego do Diabo (Loures, Portugal): Dating the Emergence of Anatomical Modernity in Westernmost Eurasia". PLOS One. 5 (1): e8880. Bibcode:2010PLoSO...5.8880Z. doi:10.1371/journal.pone.0008880. PMC 2811729. PMID 20111705. Lay summary – ScienceDaily (January 27, 2010).

Bibliography

• Derev'anko, Anatoliy P.; Powers, William Roger; Shimkin, Demitri Boris (1998). The Paleolithic of Siberia: new discoveries and interpretations. Novosibirsk: Institute of Anthropology and Ethnography. ISBN 978-0-252-02052-0. OCLC 36461622.
• Lunine, Jonathan I. (2013). Earth: Evolution of a Habitable World. Cambridge University Press. 327. ISBN 978-0-521-85001-8.

Further reading

• Reich, David (2018). Who We Are And How We Got Here – Ancient DNA and the New Science of the Human Past. Pantheon Books. ISBN 978-1101870327. ^[1]
• Sankararaman, Sriram; Mallick, Swapan; Dannemann, Michael; Prüfer, Kay; Kelso, Janet; Patterson, Nick; Reich, David (2014). "The genomic landscape of Neanderthal ancestry in present-day humans". Nature. 507 (7492): 354–57. Bibcode:2014Natur.507..354S. doi:10.1038/nature12961. PMC 4072735. PMID 24476815.
• Vattathil, S.; Akey, J.M. (2015). "Small amounts of archaic admixture provide big insights into human history". Cell. 163 (2): 281–84. doi:10.1016/j.cell.2015.09.042. PMID 26451479.

External links

Wikimedia Commons has media related to Homo neanderthalensis.

Wikibooks has a book on the topic of: Introduction to Paleoanthropology

Wikispecies has information related to Homo neanderthalensis

Wikisource has the text of the 1905 New International Encyclopedia article Neanderthal Man.

• Kreger, C. David (June 30, 2000). "Homo neanderthalensis". ArchaeologyInfo.com. Retrieved May 23, 2009.
• O'Neil, Dennis (May 12, 2009). "Evolution of Modern Humans: Neandertals". Retrieved May 23, 2009.
• New Portuguese skull may be an early relative of Neandertals – article by Ann Gibbons at Science, March 13, 2017.
• In Neanderthal DNA, Signs of a Mysterious Human Migration – article by Carl Zimmer, NY Times, July 4, 2017
• "Homo neanderthalensis". The Smithsonian Institution.
• "Neanderthal DNA". International Society of Genetic Genealogy. Archived from the original on June 17, 2006. : Includes Neanderthal mtDNA sequences
• Panoramio – The Neandertal foot prints' (photo of ≈25K years old fossilised footprints discovered in 1970 on volcanic layers near Demirkopru Dam Reservoir, Manisa, Turkey)
• Did better mothering defeat the Neanderthals?
• My Great-great-great Grandfather's a Neanderthal
• Ancient tryst fortified human immune system
• Neanderthal-human hybridisation hypothesis
• Neanderthal hybridisation and Haldane's rule
• Neanderthal Studies Professional Online Service (NESPOS) fossil overview
• Human Timeline (Interactive) – Smithsonian, National Museum of Natural History (August 2016).