Overview

Brief Summary

Taxonomy

Similar to Homo sapiens skeletally but the pelvis was more primitive in shape, broad and with an extended and thinner superior pubic ramus.The cranium was long and relatively low, with a double-arched supraorbital torus and a large nose, accompanied by mid-facial projection. The anterior mandible lacked the full development of a chin.

Morphology
Distinct in trunk shape and limb proportions, some of which may reflect cold adaptation.

Diagnostic description
Cranium displays mid-facial projection anteriorly, and a suprainiac fossa posteriorly.

Evolution
Fossil and genetic data suggest that Neanderthals shared a common ancestor with Homo sapiens in the Middle Pleistocene (perhaps the species Homo heidelbergensis).Primitive/ancestral Neanderthals are known from about 400,000 years ago, while the best-known examples date from between 35-130,000 years ago.

Genetics
Whole mitochondrial (mtDNA) genomes have been sequenced, indicating that Neanderthals represented a distinct lineage from all recent humans, with an inferred Middle Pleistocene divergence date. The newly published composite Neanderthal genome indicates a level of about 2% introgression of Neanderthal genes in modern humans outside Africa, perhaps from geneflow in the middle east about 60,000 years ago.
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Introduction

Homo neanderthalensis otherwise known as Neanderthals were probably our closest relatives. Neanderthals were humans but they were a different species from us, Homo sapiens. Homo neanderthalensis evolved in Europe and Asia while we were evolving in Africa.Fossils are known from the southerly regions of western Eurasia in both glacial and interglacial periods, while mitochondrial DNA data suggest they expanded as far as Siberia at times.
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Evolutionary Tree Information

Both fossil and genetic evidence indicate that Neanderthals and modern humans (Homo sapiens) evolved from a common ancestor between 500,000 and 200,000 years ago. Neandertals and modern humans belong to the same genus (Homo) and inhabited the same geographic areas in Asia for 30,000–50,000 years; genetic evidence indicate while they may have interbred with non-African modern humans, they are separate branches of the human family tree (separate species).

In fact, Neandertals and modern humans may have had little direct interaction for tens of thousands of years until during one very cold period, modern humans spread across Europe. Their presence may have prevented Neanderthals from expanding back into areas they once favored and served as a catalyst for the Neanderthal’s impending extinction. Over just a few thousand years after modern humans moved into Europe, Neanderthal numbers dwindled to the point of extinction. All traces of Neanderthals disappeared by about 28,000 years ago. The most recently dated Neanderthal fossils come from western Europe, which was likely where the last population of this early human species existed.

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Nickname: Neanderthal

Where Lived: Europe and southwestern to central Asia

When Lived: About 200,000 - 28,000 years ago

Neanderthals (Neander-thal, the ‘th’ pronounced as ‘t’) are our closest extinct human relative. Some defining features of their skulls include the large middle part of the face, angled cheek bones, and a huge nose for humidifying and warming cold, dry air. Their bodies were shorter and stockier than ours, another adaptation to living in cold environments. But their brains were just as large as ours and often larger - proportional to their brawnier bodies.

Neanderthals made and used a diverse set of sophisticated tools, controlled fire, lived in shelters, made and wore clothing, were skilled hunters of large animals and also ate plant foods, and occasionally made symbolic or ornamental objects. There is evidence that Neanderthals deliberately buried their dead and occasionally even marked their graves with offerings, such as flowers. No other primates, and no earlier human species, had ever practiced this sophisticated and symbolic behavior.

DNA has been recovered from more than a dozen Neanderthal fossils, all from Europe; the Neanderthal Genome Project is one of the exciting new areas of human origins research.

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Comprehensive Description

Biology

The Neanderthal species is inferred to have been largely similar to Homo sapiens, but with some distinct features as listed below.

Size
Neanderthal adult height averaged about 160cm, and average body mass is estimated at about 60-85kg

Growth
Most data suggest that the Neanderthals had an extended period of growth similar to, or only slightly accelerated compared with, Homo sapiens.

Life Expectancy
Skeletal data suggest that it was rare for a Neanderthal to reach 50 years in age.

Physiology
This is assumed to have been largely similar to that of Homo sapiens, but with the possibility of a somewhat higher metabolic rate.
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Physical Description

Size

Height:

Males: average 5 ft 5 in (164 cm); Females: average 5 ft 1 in (155 cm)

Weight:

Males: average 143 lbs (65 kg); Females: average 119 lbs (54 kg)

Height & Weight Supplemental Information:

While the tall, lean bodies of earlier human species like Homo erectus were adapted to tropical temperatures, the short, stocky bodies of Neanderthals were adapted to winter climates. With thick, short lower arm and leg bones making up a very strong and compact body, Neanderthals were more heavily built than modern humans and were more efficient at conserving body heat, allowing them to withstand living in cold climates during ice ages.

Neanderthal males were slightly taller and heavier than Neandertal females; however, sexual dimorphism was not nearly as extreme as what is seen in earlier human species.

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Type Information

Key Fossils

La Chapelle-aux-Saints

Nickname: The Old Man of La Chapelle

Site: La Chapelle-aux-Saints, France

Date of discovery: 1908

Discovered by: Amadee and Jean Bouyssonie and L. Bardon

Age: About 60,000 years old

Species: Homo neanderthalensis

The old man of La Chapelle

Discovered in 1908, the skeleton of "the old man of La Chapelle" was the first relatively complete skeleton of a Neanderthal individual that scientists had ever found. Buried in the limestone bedrock of a small cave near La Chapelle-aux-Saints, France, this skeleton includes the skull, jaw, most of the vertebrae, several ribs, most of the long bones of his arms and legs, plus some of the smaller bones of his hands and feet. The well-preserved skull shows the low, receding forehead, protruding midface, and heavy browridges typical of Homo neanderthalensis.

Scientists estimate he was quite old by the time he died, as bone had re-grown along the gums where he had lost several teeth, perhaps decades before. He lacked so many teeth in fact that it’s possible he needed his food ground down before he was able to eat it. Other Neanderthals in his social group may have supported him in his final years.

The original reconstruction of the ‘Old Man of La Chapelle’ by scientist Pierre Marcellin Boule led to the reason why popular culture stereotyped Neanderthals as dim-witted brutes for so many years. In 1911, Boule reconstructed this skeleton with a severely curved spine indicative of a stooped, slouching stance with bent knees, forward flexed hips, and the head jutted forward. He thought the low vaulted cranium and the large brow ridge, somewhat reminiscent of that seen in large apes such as gorillas, indicated a generally primitive early human and a lack of intelligence. However, additional discoveries of Neanderthal skeletons coupled with a re-examination of the Old Man’s skeleton in the 1950’s showed that many of the features thought to be unique in Neanderthals fall within the range of modern human variation, and that the Old Man suffered from “gross deforming osteoarthritis”. Thus, the slouching posture of the original reconstruction may have been based on an unfortunate individual with a deforming disability.

But this isn't quite the whole story. A more recent evaluation of the entire skeleton by scientist Erik Trinkaus has shown that, while the Old Man of La Chapelle did suffer from a degenerative joint disease, the deformation caused by this should not have affected Boule's original reconstruction of the individual’s posture. It appears that Boule's own preconceptions about early humans, and his rejection of the hypothesis that Neanderthals were the ancestors of modern humans, led him to reconstruct a stooped, brutish creature, effectively placing Neanderthals on a side branch of the human evolutionary tree. (Boule even gave his reconstruction an opposable big toe like the great apes, but there was no bone deformity that should or could have lead to this interpretation.)

3-D collection link: http://humanorigins.si.edu/evidence/3d-collection/f-la-chapelle-aux-saints

La Ferrassie

Site: La Ferrassie Cave, France

Date of discovery: 1909

Discovered by: Louis Capitan and Denis Peyrony

Age: Between 70,000 and 50,000 years old

Species: Homo neanderthalensis

Most complete Neanderthal skull

The excavations at the La Ferrassie rock shelter in the Dordogne Valley, France in the early 20th century produced the remains of an adult male and an adult female, providing scientists with the first evidence of sexual dimorphism in Neanderthals. In addition, the remains of the child and infant individuals help scientists understand the growth rates of Neanderthal children. A total of eight Neanderthal individuals -- including adults, children, infants, and two fetuses -- were found intentionally buried at La Ferrassie.

One of the most important individuals found at La Ferrassie is La Ferrassie 1, the skeleton of an adult male. His skull, the largest and most complete Neanderthal skull ever found (in 1909), has many of the typical Neanderthal traits such as the low, sloping forehead and large nasal opening. His teeth, which are all preserved, are heavily worn, indicating he was older at the time of his death. His front incisors show a slanted wear that does not occur from chewing; one hypothesis to explain this odd wear on his teeth is that he habitually held something in place between his front teeth, such as a hide, that he then scraped with a tool . Although this hypothesis has been debated, the use of the teeth as tools may represent a remarkable Neanderthal behavioral adaptation.

La Ferrassie 1 is considered by many scientists to exhibit the ‘classic’ example of Neanderthal anatomy. His leg and feet bones proved without a doubt that Neanderthals walked upright and with a gait very similar to modern humans. This debunked the earlier reconstruction of the La Chapelle-aux-Saints Neanderthal skeleton by French paleontologist Pierre Marcellin Boulethat portrayed this species as stooped, brutish creatures.

3-D collection link: http://humanorigins.si.edu/evidence/3d-collection/f-la-ferrassie-1-cranium

http://humanorigins.si.edu/evidence/3d-collection/f-la-ferrassie-1-mandible

Shanidar 1

Nickname: Nandy

Site: Shanidar, Iraq

Discovered by: Ralph Solecki

Age: Between 45,000 and 35,000 years old

Species: Homo neanderthalensis

Blow to the head

Through examining his skeletal remains, scientists found evidence that at a young age, Shanidar 1 experienced a crushing blow to his head. The blow damaged the left eye (possibly blinding him) and the brain area controling the right side of the body, leading to a withered right arm and possible paralysis that also crippled his right leg. One of Shanidar 1’s middle foot bones (metatarsal) on his right foot shows a healed fracture, which probably only enhanced his noticeable limp. All of Shanidar 1’s injuries show signs of healing, so none of them resulted in his death. In fact, scientists estimate he lived until 35–45 years of age. He would have been considered old to another Neandertal, and he would probably not have been able to survive without the care of his social group.

3-D collection link: http://humanorigins.si.edu/evidence/3d-collection/f-shanidar-1

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Ecology

General Ecology

Distribution ecology

Distribution
Neanderthal fossils are known from the southerly regions of western Eurasia in both glacial and interglacial periods, while mtDNA data suggest they expanded as far as Siberia at times.

Habitat
They ranged from temperate woodland to steppe-tundra environments.

Population biology
Population numbers are believed to have been low, particularly in glacial stages, mtDNA studies suggest an effective population size in later Neanderthals of less than 3000 females.

Trophic Strategy
From skeletal isotope data, the Neanderthals were top-level carnivores, preying on large mammals.
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Life History and Behavior

Behavior

Behaviour

This early human species was a large-brained biped, tool-making and tool-using, with a hunter-gatherer lifestyle. Behaviour included apparent burial of the dead.

Associations
The Neanderthals were evidently highly carnivorous, but they undoubtedly ate other foods, although only limited evidence of plant foods survives in the archaeological record.In Mediterranean regions the Neanderthals also exploited marine resources such as shellfish and seals, but their use of aquatic foods was certainly more limited than that of Homo sapiens.

Diseases
The Neanderthals evidently suffered from degenerative diseases such as osteoarthritis, and their skeletons also bear traces of lesions indicating systemic infection or carcinoma.
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How They Survived

Compared to early humans living in tropical Africa, with more abundant edible plant foods available year-round, the number of plant foods Neanderthals could eat would have dropped significantly during the winter of colder climates, forcing Neanderthals to exploit other food options like meat more heavily. There is evidence that Neanderthals were specialized seasonal hunters, eating animals were available at the time (i.e. reindeer in the winter and red deer in the summer). Scientists have clear evidence of Neanderthal hunting from uncovering sharp wooden spears and large numbers of big game animal remains were hunted and butchered by Neanderthals. There is also evidence from Gibraltar that when they lived in coastal areas, they exploited marine resources such as mollusks, seals, dolphins and fish. Isotopic chemical analyses of Neanderthal bones also tell scientists the average Neanderthal’s diet consisted of a lot of meat. Scientists have also found plaque on the remains of molar teeth containing starch grains—concrete evidence that Neandertals ate plants.

The Mousterian stone tool industry of Neanderthals is characterized by sophisticated flake tools that were detached from a prepared stone core. This innovative technique allowed flakes of predetermined shape to be removed and fashioned into tools from a single suitable stone. This technology differs from earlier ‘core tool’ traditions, such as the Acheulean tradition of Homo erectus. Acheulean tools worked from a suitable stone that was chipped down to tool form by the removal of flakes off the surface.

Neanderthals used tools for activities like hunting and sewing. Left-right arm asymmetry indicates that they hunted with thrusting (rather than throwing) spears that allowed them to kill large animals from a safe distance. Neanderthal bones have a high frequency of fractures, which (along with their distribution) are similar to injuries among professional rodeo riders who regularly interact with large, dangerous animals. Scientists have also recovered scrapers and awls (larger stone or bone versions of the sewing needle that modern humans use today) associated with animal bones at Neanderthal sites. A Neanderthal would probably have used a scraper to first clean the animal hide, and then used an awl to poke holes in it, and finally use strips of animal tissue to lace together a loose-fitting garment. Neanderthals were the first early humans to wear clothing, but it is only with modern humans that scientists find evidence of the manufacture and use of bone sewing needles to sew together tighter fitting clothing.

Neanderthals also controlled fire, lived in shelters, and occasionally made symbolic or ornamental objects. There is evidence that Neanderthals deliberately buried their dead and occasionally even marked their graves with offerings, such as flowers. No other primates, and no earlier human species, had ever practiced this sophisticated and symbolic behavior. This may be one of the reasons that the Neanderthal fossil record is so rich compared to some earlier human species; being buried greatly increases the chance of becoming a fossil!

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Evolution and Systematics

Fossil History

History of Discovery

Year of Discovery: 1829

Neandertal 1 was the first specimen to be recognized as an early human fossil. When it was discovered in 1856 in Germany, scientists had never seen a specimen like it: the oval shaped skull with a low, receding forehead and distinct browridges, the thick, strong bones. In 1864, it became the first fossil hominin species to be named—geologist William King suggested the name Homo neanderthalensis (Johanson and Edgar, 2006), after these fossils found in the Feldhofer Cave of the Neander Valley in Germany (tal—a modern form of thal—means “valley” in German). Several years after Neanderthal 1 was discovered, scientists realized that prior fossil discoveries—in 1829 at Engis, Belgium, and in 1848 at Forbes Quarry, Gibraltar—were also Neanderthals. Even though they weren’t recognized at the time, these two earlier discoveries were actually the first early human fossils ever found.

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Molecular Biology and Genetics

Molecular Biology

Barcode data: Homo sapiens neanderthalensis

The following is a representative barcode sequence, the centroid of all available sequences for this species.


No available public DNA sequences.

Download FASTA File
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Statistics of barcoding coverage: Homo sapiens neanderthalensis

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 6
Specimens with Barcodes: 6
Species With Barcodes: 1
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Wikipedia

Neanderthal

For other uses, see Neanderthal (disambiguation).

The Neanderthals (or Neandertals, from German: Neandertaler) (/niˈændərˌθɔːlz/, /niˈændərˌtɔːlz/, /niˈændərˌtɑːlz/, /nˈɑːndərˌtɑːlz/ or /niˈændərθəlz/)[3] are an extinct species of human in the genus Homo. They are closely related to modern humans,[4][5] differing in DNA by just 0.12%.[6] Remains left by Neanderthals include bone and stone tools, which are found in Eurasia, from Western Europe to Central and Northern Asia. The species is named after Neandertal ("Neander Valley"), the location in Germany where it was first discovered. Neanderthals are generally classified by biologists as the species Homo neanderthalensis, but a minority consider them to be a subspecies of Homo sapiens (Homo sapiens neanderthalensis).[7]

The first humans with proto-Neanderthal traits are believed to have existed in Eurasia as early as 350,000 - 600,000 years ago[8] with the first "true Neanderthals" appearing between 200,000 and 250,000 years ago.[9] The exact date of their extinction had been disputed. However, in 2014, Thomas Higham of the University of Oxford performed the most comprehensive dating of Neanderthal bones and tools ever carried out, which demonstrated that Neanderthals died out in Europe between 41,000 and 39,000 years ago - this coincides with the start of a very cold period in Europe and is 5,000 years after Homo sapiens reached the continent. This was based on improved radiocarbon dating of materials from 40 sites in Western Europe.[10]

Several cultural assemblages have been linked to the Neanderthals in Europe. The earliest, the Mousterian stone tool culture, dates to about 300,000 years ago.[11] Late Mousterian artifacts were found in Gorham's Cave on the south-facing coast of Gibraltar.[12][13]

With an average cranial capacity of 1600 cm3,[14] Neanderthal's cranial capacity is notably larger than the 1400 cm3 average for modern humans, indicating that their brain size was larger. However, owing to larger body size, Neanderthals are less encephalized.[15] Males stood 164–168 cm (65–66 in) and females about 152–156 cm (60–61 in) tall.[16]

Genetic evidence published in 2014 suggests that Neanderthals contributed to the DNA of anatomically modern humans, probably through interbreeding between 50,000 to 60,000[17] years ago with a population of anatomically modern humans. According to the study, by the time that population began dispersing across Eurasia, Neanderthal genes constituted as much as 1–4% of its genome (roughly equivalent to having one Neanderthal great-great-great-grandparent).[18][19][20] Ötzi the iceman, Europe's oldest preserved mummy, was found to possess an even higher percentage of Neanderthal ancestry.[21] Recent findings suggest there may be even more Neanderthal genes in non-African humans than previously expected: approximately 20% of the Neanderthal gene pool was present in a broad sampling of non-African individuals, though each individual's genome was on average only 2% Neanderthal.[22]

In December 2013, researchers reported evidence that Neanderthals practiced burial behavior and intentionally buried their dead.[23] In addition, scientists reported, for the first time, the entire genome of a Neanderthal. The genome was extracted from the toe bone of a 130,000-year-old Neanderthal found in a Siberian cave.[24][25]

Name[edit]

The species is named after the site of its first discovery, about 12 km (7.5 mi) east of Düsseldorf, Germany, in the Feldhofer Cave in the river Düssel's Neander valley named after Joachim Neander, a 17th-century German pastor and hymnist. Neander's own name was in turn a Greek translation of the German Neumann (lit. "New man"). Thal is the older spelling of Tal (both with the same pronunciation), the German word for 'valley' (cognate with English dale).[26][27][28]

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".[29] The binomial name Homo neanderthalensis – extending the name "Neanderthal man" from the individual type specimen to the entire species – was first proposed by the Anglo-Irish geologist William King in 1864 and this had priority over the proposal put forward in 1866 by Ernst Haeckel, Homo stupidus.[27] The practice of referring to "the Neanderthals" and "a Neanderthal" emerged in the popular literature of the 1920s.[30]

The German pronunciation of Neanderthaler and Neandertaler is [neˈandɐˌtʰaːlɐ] in the International Phonetic Alphabet. In British English, "Neanderthal" is pronounced with the /t/ as in German but different vowels (IPA: /niːˈændərtɑːl/).[31][32][33] In layman's American English, "Neanderthal" is pronounced with a /θ/ (the voiceless th as in thin) and /ɔ/ instead of the longer British /aː/ (IPA: /niːˈændərθɔːl/),[34] although scientists typically use the /t/ as in German.[35][36]

Classification[edit]

Crania: 1. Gorilla 2. Australopithecus 3. Homo erectus 4. Neanderthal (La Chapelle aux Saints) 5. Steinheim Skull 6. Modern Human

For some time, scientists have debated whether Neanderthals should be classified as Homo neanderthalensis or Homo sapiens neanderthalensis, the latter placing Neanderthals as a subspecies of H. sapiens.[37][38] Some morphological studies support the view that H. neanderthalensis is a separate species and not a subspecies.[39][40] Others, for example University of Cambridge Professor Paul Mellars, say "no evidence has been found of cultural interaction"[41] and evidence from mitochondrial DNA studies has been interpreted as evidence Neanderthals were not a subspecies of H. sapiens.[42]

Origin[edit]

Comparison of the DNA of Neanderthals and Homo sapiens suggests that they diverged from a common ancestor between 350,000 and 400,000 years ago. This ancestor was probably Homo heidelbergensis. Heidelbergensis originated between 800,000 and 1,300,000 years ago, and continued until about 200,000 years ago. It ranged over Eastern and South Africa, Europe and Western Asia. Between 350,000 and 400,000 years ago the African branch is thought to have started evolving towards modern humans and the Eurasian branch towards Neanderthals. Scientists do not agree when Neanderthals can first be recognised in the fossil record, with dates ranging between 200,000 and 300,000 years BP.[43][44][45][46]

Discovery[edit]

Neander Valley site
The site of Kleine Feldhofer Grotte where the type specimen was unearthed by miners in the 19th century
Location of Neander Valley, Germany, with the modern federal state of North Rhine-Westphalia highlighted

Neanderthal skulls were first discovered in Engis Caves (fr), in what is now Belgium (1829) by Philippe-Charles Schmerling and in Forbes' Quarry, Gibraltar, dubbed Gibraltar 1 (1848), both prior to the type specimen discovery in a limestone quarry of the Neander Valley in Erkrath near Düsseldorf in August 1856, three years before Charles Darwin's On the Origin of Species was published.[47]

The type specimen, dubbed Neanderthal 1, consisted of a skull cap, two femora, three bones from the right arm, two from the left arm, part of the left ilium, fragments of a scapula, and ribs. The workers who recovered this material originally thought it to be the remains of a bear. They gave the material to amateur naturalist Johann Carl Fuhlrott, who turned the fossils over to anatomist Hermann Schaaffhausen.

To date, the bones of over 400 Neanderthals have been found.[48]

Timeline[edit]

Neanderthal fossils
Skull, found in 1886 in Spy, Belgium
Frontal bone of a neanderthal child from the cave of La Garigüela
Skull from La Chapelle aux Saints
Semi-frontal view of a neanderthal skull from Gibraltar
  • 1829: Neanderthal skulls were discovered in Engis, in present-day Belgium.
  • 1848: Neanderthal skull Gibraltar 1 found in Forbes' Quarry, Gibraltar. Called "an ancient human" at the time.
  • 1856: Johann Karl Fuhlrott first recognized the fossil called "Neanderthal man", discovered in Neanderthal, a valley near Mettmann in what is now North Rhine-Westphalia, Germany.
  • 1880: The mandible of a Neanderthal child was found in a secure context and associated with cultural debris, including hearths, Mousterian tools, and bones of extinct animals.
  • 1886: Two nearly perfect skeletons of a man and woman were found at Spy, Belgium at the depth of 16 ft with numerous Mousterian-type implements.
  • 1899: Hundreds of Neanderthal bones were described in stratigraphic position in association with cultural remains and extinct animal bones.
  • 1899: Sand excavation workers found bone fragments on a hill in Krapina, Croatia called Hušnjakovo brdo. Local Franciscan friar Dominik Antolković requested Dragutin Gorjanović-Kramberger to study the remains of bones and teeth that were found there.
  • 1905: During the excavation in Krapina more than 5 000 items were found, of which 874 residue of human origin, including bones of prehistoric man and animals, artifacts.
  • 1908: A nearly complete Neanderthal skeleton was discovered in association with Mousterian tools and bones of extinct animals.[clarification needed]
  • 1925: Francis Turville-Petre finds the 'Galilee Man' or 'Galilee Skull' in the Zuttiyeh Cave in Wadi Amud in The British Mandate of Palestine (now Israel).
  • 1926 Skull fragments of Gibraltar 2, a four-year-old Neanderthal girl, discovered by Dorothy Garrod.
  • 1953–1957: Ralph Solecki uncovered nine Neanderthal skeletons in Shanidar Cave in the Kurdistan region of northern Iraq.
  • 1975: Erik Trinkaus' study of Neanderthal feet confirmed they walked like modern humans.
  • 1987: Thermoluminescence results from Israeli fossils date Neanderthals at Kebara to 60,000 BP and humans at Qafzeh to 90,000 BP. These dates were confirmed by electron spin resonance (ESR) dates for Qafzeh (90,000 BP) and Es Skhul (80,000 BP).
  • 1991: ESR dates showed the Tabun Neanderthal was contemporaneous with modern humans from Skhul and Qafzeh.
  • 1993: 127,000-year-old DNA is found on the child of Sclayn, found in Scladina (fr), Belgium.
  • 1997: Matthias Krings et al. are the first to amplify Neanderthal mitochondrial DNA (mtDNA) using a specimen from Feldhofer grotto in the Neander valley.[49]
  • 1998: A team led by pre-history archeologist João Zilhão discovered an early Upper Paleolithic human burial in Portugal, at Abrigo do Lagar Velho, which provided evidence of early modern humans from the west of the Iberian Peninsula. The remains, a largely complete skeleton of an approximately 4-year-old child, buried with pierced shell and red ochre, is dated to ca. 24,500 years BP.[50] The cranium, mandible, dentition, and postcrania present a mosaic of European early modern human and Neanderthal features.[50]
  • 2000: Igor Ovchinnikov, Kirsten Liden, William Goodman et al. retrieved DNA from a Late Neanderthal infant from Mezmaiskaya Cave in the Caucasus.[51]
  • 2005: The Max Planck Institute for Evolutionary Anthropology launched a project to reconstruct the Neanderthal genome, working with Connecticut-based 454 Life Sciences.[citation needed] In 2009, the Max Planck Institute announced the "first draft" of a complete Neanderthal genome is completed.[52]
  • 2010: Comparison of Neanderthal genome with modern humans from Africa and Eurasia shows that 1–4% of modern non-African human genome might come from the Neanderthals.[19][20]
  • 2010: Discovery of Neanderthal tools far away from the influence of H. sapiens indicate that the species might have been able to create and evolve tools on its own, and therefore be more intelligent than previously thought. Furthermore, it was proposed that the Neanderthals might be more closely related to Homo sapiens than previously thought and that may in fact be a sub species of it.[53] Evidence has more recently emerged that these artifacts are probably of H. sapiens sapiens origin.[54]
  • 2012: Charcoal found next to six paintings of seals in Nerja caves, Malaga, Spain, has been dated to between 42,300 and 43,500 years old. The paintings themselves will be dated in 2013, and if their pigment matches the date of the charcoal, they would be the oldest known cave paintings. José Luis Sanchidrián at the University of Cordoba, Spain believes the paintings are more likely to have been painted by Neanderthals than early modern humans.[55]
  • 2013: A jawbone found in Italy had features intermediate between Neanderthals and Homo sapiens suggesting it could be a hybrid. The mitochondrial DNA is Neanderthal.[56]
  • 2013: An international team of researchers reported evidence that Neanderthals practiced burial behavior and intentionally buried their dead.[23]
  • 2014: Researchers at the University of Colorado Museum in Boulder report that Neanderthals were not less intelligent than modern humans and "that single-factor explanations for the disappearance of the Neandertals are not warranted any more."[57]
  • 2014: Prof Thomas Higham of the University of Oxford performed the most comprehensive dating of Neanderthal bones and tools ever carried out, which demonstrated that Neanderthals died out in Europe between 41,000 and 39,000 years ago - this coincides with the start of a very cold period in Europe and is 5000 years after Homo sapiens reached the continent.[58]

Habitat and range[edit]

Further information: List of Neanderthal sites
Sites where typical Neanderthal fossils have been found

Early Neanderthals lived in the Last glacial period for a span of about 100,000 years. Because of the damaging effects the glacial period had on the Neanderthal sites, not much is known about the early species. Countries where their remains are known include most of Europe south of the line of glaciation, roughly along the 50th parallel north. This includes most of Western Europe, the south coast of Great Britain,[59] Central Europe, the Carpathians, and the Balkans,[60] some sites in Ukraine and in western Russia, Central and Northern Asia up to the Altai Mountains, and Western Asia from the Levant up to the Indus River. It is estimated that the total Neanderthal population across this habitat range numbered at around 70,000 at its peak.[61]

Neanderthal fossils have not been found to date in Africa, but there have been finds close to North Africa, both on Gibraltar and in the Levant. At some Levantine sites, Neanderthal remains date from after the same sites were vacated by modern humans. Mammal fossils of the same time period show cold-adapted animals were present alongside these Neanderthals in this region of the Eastern Mediterranean. This implies Neanderthals were better adapted biologically to cold weather than modern humans and at times displaced them in parts of the Middle East when the climate got cold enough.[62]

Homo sapiens sapiens appears to have been the only human type in the Nile River Valley during these periods, and Neanderthals are not known to have ever lived south-west of present-day Israel. When climate change caused warmer temperatures, the Neanderthal range likewise retreated to the north, along with the cold-adapted species of mammals. Apparently these weather-induced population shifts took place before modern people secured competitive advantages over the Neanderthal, as these shifts in range took place well over ten thousand years before modern people totally replaced the Neanderthal, despite the recent evidence of some successful interbreeding.[62]

Separate developments in the human line, in other regions such as Southern Africa, somewhat resembled the Eurasian Neanderthals, but these people were not Neanderthals. One such example is Rhodesian Man (Homo rhodesiensis), who existed long before any classic Eurasian Neanderthals, but had a more modern set of teeth. Some H. rhodesiensis populations appeared to be on the road to modern H. sapiens sapiens. At any rate, the populations in Eurasia underwent more and more "Neanderthalization" as time went on. There is some argument that H. rhodesiensis in general was ancestral to both modern humans and Neanderthals, and that at some point the two populations went their separate ways, but this supposes that H. rhodesiensis goes back to around 600,000 years ago.

To date, no intimate connection has been found between these similar archaic people and the Eurasian Neanderthals, at least during the same time as H. rhodesiensis seems to have lived about 600,000 years ago, long before the time of classic Neanderthals. This said, some researchers think that H. rhodesiensis may have lived much later than this period, depending on the method used to date the fossils, leaving this issue open to debate. Some H. rhodesiensis features, like the large brow ridge, may have been caused by convergent evolution.

It appears incorrect, based on present research and known fossil finds, to refer to any fossils outside Eurasia as true Neanderthals. They had a known range that possibly extended as far east as the Altai Mountains, but not farther to the east or south, and apparently not into Africa. At any rate, in North-East Africa the land immediately south of the Neanderthal range was possessed by modern humans Homo sapiens idaltu or Homo sapiens, since at least 160,000 years before the present. 160,000-year-old hominid fossils at Jebel Irhoud in Morocco were previously thought to be Neanderthal, but it is now clear that they are early modern humans.[63]

Classic Neanderthal fossils have been found over a large area, from northern Germany to Israel and Mediterranean countries like Spain[64] and Italy[65] in the south and from England and Portugal 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 farther north, up to 60° N, on the Russian plain.[66] Recent evidence has extended the Neanderthal range by about 1,250 miles (2,010 km) east into southern Siberia's Altai Mountains.[67][68]

Anatomy[edit]

Main article: Neanderthal anatomy
Reconstruction of the head of the Shanidar 1 fossil, a Neanderthal male who lived c. 70,000 years ago (John Gurche 2010).

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, particularly in certain isolated geographic regions. These include shorter limb proportions, a wider, barrel-shaped rib cage, a reduced chin and, perhaps most notably, a large nose, which was much larger in both length and width, and started somewhat higher on the face, than in modern humans.[9] Evidence suggests they were much stronger than modern humans, with particularly strong arms and hands,[69][70] while they were comparable in height; based on 45 long bones from at most 14 males and 7 females, Neanderthal males averaged 164–168 cm (65–66 in) and females 152–156 cm (60–61 in) tall.[16] 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.[71] A 2007 genetic study suggested some Neanderthals may have had red hair and blond hair, along with a light skin tone.[72]

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.[73] Neanderthals are known for their large cranial capacity, which at 1600 cm3 is larger on average than that of modern humans. One study has found that Neanderthal brains were more asymmetric than other hominid brains.[74] 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.[75]

Behavior[edit]

Main article: Neanderthal behavior

Neanderthals made advanced tools,[76] probably had a language (the nature of which is debated and likely unknowable) and lived in complex social groups. The Molodova archaeological site in eastern Ukraine suggests some Neanderthals built dwellings using animal bones. A building was made of mammoth skulls, jaws, tusks and leg bones, and had 25 hearths inside.[77]

Neanderthals may have been capable of building dugout boats since the Middle Paleolithic.[78][79] Mousterian stone tools discovered on the southern Ionian Islands suggests that Neanderthals were sailing the Mediterranean Sea as early as 110,000 years BP.[80][81] Quartz hand-axes, three-sided picks, and stone cleavers from Crete have also been recovered that date back about 170,000 years BP.[82]

It was once thought that Neanderthals lacked the sophistication for hunting, perhaps scavenging meat from carcasses,[9] but increasing evidence suggests they were apex predators,[83][84] capable of bringing down a wide range of prey from red deer, reindeer, ibex and wild boar, to larger animals such as aurochs and even, on occasion, mammoth, straight-tusked elephant and rhinoceros.[9][85] However, while largely carnivorous,[86][87] new studies indicate Neanderthals also had cooked vegetables in their diet.[84][88] In 2010, an isotope analysis of Neanderthal teeth found traces of cooked vegetable matter, and more recently a 2014 study of Neanderthal coprolites (fossilized feces) found substantial amounts of plant matter, contradicting the earlier belief they were exclusively (or almost exclusively) carnivorous.[86][89]

Violence[edit]

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. This wound was likely fatal, given the lack of medical care, causing the victim to bleed out, or through cranial concussion.[90]

Genome[edit]

Further information: Neanderthal genome project

Early investigations concentrated on mitochondrial DNA (mtDNA), which, owing to strictly matrilineal inheritance and subsequent vulnerability to genetic drift, is of limited value in evaluating the possibility of interbreeding of Neanderthals with Cro-Magnon people.

In 1997, geneticists were able to extract a short sequence of DNA from Neanderthal bones.[91] The extraction of mtDNA from a second specimen was reported in 2000, and showed no sign of modern human descent from Neanderthals.[51]

Scientist at the Max Planck Institute for Evolutionary Anthropology extracting the DNA.

In July 2006, the Max Planck Institute for Evolutionary Anthropology and 454 Life Sciences announced that they would sequence the Neanderthal genome over the next two years. This genome was expected to be roughly the size of the human genome, three-billion base pairs, and share most of its genes. It was hoped the comparison would expand understanding of Neanderthals, as well as the evolution of humans and human brains.[92]

Svante Pääbo has tested more than 70 Neanderthal specimens. Preliminary DNA sequencing from a bone fragment of a femur found at Vindija Cave, Croatia, in 1980 showed Neanderthals and modern humans 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[93] appearing in the journal Nature has calculated the species diverged about 516,000 years ago, whereas fossil records show a time of about 400,000 years ago.[94] A 2007 study pushes the point of divergence back to around 800,000 years ago.[95]

Edward Rubin of the Lawrence Berkeley National Laboratory states recent genome testing of Neanderthals suggests human and Neanderthal DNA are some 99.5% to nearly 99.9% identical.[96][97]

On 16 November 2006, Lawrence Berkeley National Laboratory issued a press release suggesting Neanderthals and ancient humans probably did not interbreed.[98] Edward M. Rubin, director of the U.S. Department of Energy's Lawrence Berkeley National Laboratory and the Joint Genome Institute (JGI), sequenced a fraction (0.00002) of genomic nuclear DNA (nDNA) from a 38,000-year-old Vindia Neanderthal femur. They calculated the common ancestor to be about 353,000 years ago, and a complete separation of the ancestors of the species about 188,000 years ago.[99]

Their results show the genomes of modern humans and Neanderthals are at least 99.5% identical, but despite this genetic similarity, and despite the two species having coexisted in the same geographic region for thousands of years, Rubin and his team did not find any evidence of any significant interbreeding between the two. Rubin said, "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."[99]

In 2008 Richard E. Green et al. from Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany, published the full sequence of Neanderthal mitochondrial DNA (mtDNA) and suggested "Neanderthals had a long-term effective population size smaller than that of modern humans."[100] Writing in Nature about Green et al.'s findings, James Morgan asserted the mtDNA sequence contained clues that Neanderthals lived in "small and isolated populations, and probably did not interbreed with their human neighbours."[101][102]

In the same publication, it was disclosed by Svante Pääbo that in the previous work at the Max Planck Institute that "Contamination was indeed an issue," and they eventually realized that 11% of their sample was modern human DNA.[103][104] Since then, more of the preparation work has been done in clean areas and 4-base pair 'tags' have been added to the DNA as soon as it is extracted so the Neanderthal DNA can be identified.

With 3 billion nucleotides sequenced, analysis of about ⅓ showed no sign of admixture between modern humans and Neanderthals, according to Pääbo. This concurred with the work of Noonan from two years earlier. The variant of microcephalin common outside Africa, which was suggested to be of Neanderthal origin and responsible for rapid brain growth in humans, was not found in Neanderthals. Nor was the MAPT variant, a very old variant found primarily in Europeans.[103]

However, an analysis of a first draft of the Neanderthal genome by the same team released in May 2010 indicates interbreeding may have occurred.[19][20] "Those of us who live outside Africa carry a little Neanderthal DNA in us," said Pääbo, who led the study. "The proportion of Neanderthal-inherited genetic material is about 1 to 4 percent. It is a small but very real proportion of ancestry in non-Africans today," says Dr. David Reich of Harvard Medical School, who worked on the study. This research compared the genome of the Neanderthals to five modern humans from China, France, sub-Saharan Africa, and Papua New Guinea. The finding is that about 1 to 4 percent of the genes of the non-Africans came from Neanderthals, compared to the baseline defined by the two Africans.[19]

This indicates a gene flow from Neanderthals to modern humans, i.e., interbreeding between the two populations. Since the three non-African genomes show a similar proportion of Neanderthal sequences, the interbreeding must have occurred early in the migration of modern humans out of Africa, perhaps in the Middle East. No evidence for gene flow in the direction from modern humans to Neanderthals was found. Gene flow from modern humans to Neanderthals would not be expected if contact occurred between a small colonizing population of modern humans and a much larger resident population of Neanderthals. A very limited amount of interbreeding could explain the findings, if it occurred early enough in the colonization process.[19]

While interbreeding is viewed as the most parsimonious interpretation of the genetic discoveries, the authors point out they cannot conclusively rule out an alternative scenario, in which the source population of non-African modern humans was already more closely related to Neanderthals than other Africans were, because of ancient genetic divisions within Africa.[19] Other studies carried out since the sequencing of the Neanderthal genome have cast doubt on the level of admixture between Neanderthals and modern humans, or even as to whether the species interbred at all. One study has asserted that the presence of Neanderthal or other archaic human genetic markers can be attributed to shared ancestral traits between the species originating from a 500,000-year-old common ancestor.[105][106][107]

Among the genes shown to differ between present-day humans and Neanderthals were RPTN, SPAG17, CAN15, TTF1 and PCD16.[19]

Extinction hypotheses[edit]

As 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, and that Homo sapiens arrived in Europe between 45,000 and 43,000 years ago, it is now apparent that the two different human populations shared Europe for as long as 5,000 years.[58] The exact nature of biological and cultural interaction between Neanderthals and other human groups between has been contested.[108]

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 humans) and were replaced by modern humans moving into their habitat between 45,000 and 40,000 years ago.[109] Jared Diamond has suggested a scenario of violent conflict and displacement.[110]
  2. Neanderthals were a contemporary subspecies that bred with modern humans and disappeared through absorption (interbreeding theory).
    mtDNA-based simulation of modern human expansion in Europe starting 1600 generations ago. Neanderthal range in light grey.[111]

As Paul Jordan notes: "A natural sympathy for the underdog and the disadvantaged lends a sad poignancy to the fate of the Neanderthal folk, however it came about." Jordan, though, does say that there was perhaps interbreeding to some extent, but that populations that remained totally Neanderthal were probably out-competed and marginalized to extinction by the Aurignacians.[62]

Climate change[edit]

About 55,000 years ago, the weather began to fluctuate wildly from extreme cold conditions to mild cold and back in a matter of a few decades. Neanderthal bodies were well suited for survival in a cold climate—their barrel chests and stocky limbs stored body heat better than the Cro-Magnons. However, the rapid fluctuations of weather caused ecological changes to which the Neanderthals could not adapt; familiar plants and animals would be replaced by completely different ones within a lifetime. Neanderthals' ambush techniques would have failed as grasslands replaced trees. Neanderthals died out in Europe between 41,000 and 39,000 years ago which coincides with the start of a very cold period.[58][112] Raw material sourcing and the examination of faunal remains by Adler et al. (2006) in the southern Caucasus region suggest that modern humans may have had a survival advantage during this period, being able to use social networks to acquire resources from a greater area. They found that in both the Late Middle Palaeolithic and Early Upper Palaeolithic more than 95% of stone artifacts were drawn from local material, suggesting Neanderthals were restricted to more local resources. Furthermore, excavations at Ortvale Klde Rockshelter discovered that there was a clear break between the Late Middle Paleolithic and the Early Upper Paleolithic lithic assemblages, which were attributed to Neanderthals and modern humans respectively. This would suggest that modern humans came in and replaced Neanderthals, rather than a slow shift or integration occurring in this region. [113]

Studies on Neanderthal body structures have shown that they needed more energy to survive than any other species of hominid. Their energy needs were up to 100 to 350 kcal (420 to 1,460 kJ) more per day comparing to projected anatomically modern human males weighing 68.5 kg (151 lb) and females 59.2 kg (131 lb).[114] When food became scarce, this difference may have played a major role in the Neanderthals' extinction.[112]

Skeleton and reconstruction of the La Ferrassie 1 Neanderthal man from the National Museum of Nature and Science.

Coexistence with Homo sapiens[edit]

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.[115] 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.[58] The exact nature of biological and cultural interaction between Neanderthals and other human groups has been contested.[108]

Neanderthals became classified as extinct roughly 30,000 years ago coinciding with the arrival of what is known today, as modern humans. According to The Sensuous Curmudgeon “Whatever dates one uses, it seems that Neanderthal was the first to arrive in Europe and the Middle East.”.[116] The modern humans co-existed with them in Europe starting around 35 kya, perhaps even earlier. Before the arrival of modern humans, Neanderthals inhabited those continents for many years before to the human arrival. The humans may have introduced a disease which contributed to the extinction hypothesis that complements other recent explanations for the extinction of Neanderthals. When Neanderthal ancestors left Africa roughly 100,000 years earlier 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 could have been devastating to the Neanderthal population because they would not have any immunity to the African pathogens. “Recent European and American history provides evidence for similar events”,[116] where the introduction of viral, or bacterial pathogens to new populations lead to mass mortality and local population extinction. Similar to when Christopher Columbus brought and introduced foreign diseases to the New World when he and his crew arrived to a native population with no immune system to foreign diseases.

Interbreeding hypotheses [edit]

Chris Stringer's hypothesis of the family tree of genus Homo, published 2012 in Nature: Homo floresiensis originated in an unknown location from unknown ancestors and reached remote parts of Indonesia. Homo erectus spread from Africa to western Asia, then east Asia and Indonesia; its presence in Europe is uncertain, but it gave rise to Homo antecessor, found in Spain. Homo heidelbergensis originated from Homo erectus in an unknown location and dispersed across Africa, southern Asia and southern Europe (other scientists interpret fossils, here named heidelbergensis, as late erectus). Homo sapiens spread from Africa to western Asia and then to Europe and southern Asia, eventually reaching Australia and the Americas. In addition to Neanderthals and Denisovans, a third gene flow of archaic Africa origin is indicated at the right.[117]

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, since it would imply that at least part of the genome of Europeans would descend from Neanderthals.

Hans Peder Steensby, while strongly emphasising that all modern humans are of mixed origins, proposed the interbreeding hypothesis in 1907, in the article Race studies in Denmark.[118] He held that this would best fit current observations, and attacked the widespread idea that Neanderthals were ape-like or inferior.

The most vocal proponent of the hybridization hypothesis is Erik Trinkaus of Washington University.[119] Trinkaus claims various fossils as products of hybridized populations, including the child of Lagar Velho, a skeleton found at Lagar Velho in Portugal.[120][121] In a 2006 publication co-authored by Trinkaus, the fossils found in 1952 in the cave of Peștera Muierii, Romania, are likewise claimed as descendants of previously hybridized populations.[122]

Genetic research has asserted that some admixture took place.[123] The genomes of all non-Africans include portions that are of Neanderthal origin,[124][125] due to interbreeding between Neanderthals and the ancestors of Eurasians in Northern Africa or the Middle East prior to their spread. Rather than absorption of the Neanderthal population, this gene flow appears to have been of limited duration and limited extent. An estimated 1 to 4 percent of the DNA in Europeans and Asians (French, Chinese and Papua probands) is non-modern, and shared with ancient Neanderthal DNA rather than with Sub-Saharan Africans (Yoruba people and San probands).[19] Nonetheless, more recent genetic studies seem to suggest that modern humans may have mated with "at least two groups" of ancient humans: Neanderthals and Denisovans.[126] Some researchers suggest admixture of 3.4%-7.9% in modern humans of non-African ancestry, rejecting the hypothesis of ancestral population structure.[127] Detractors have argued and continue to argue that the signal of Neanderthal interbreed may be due to ancient African substructure, meaning that the similarity is only a remnant of a common ancestor of both Neanderthals and modern humans and not the result of interbreeding.[128][129] John D. Hawks has argued that the genetic similarity to Neanderthals may indeed be the result of both structure and interbreeding, as opposed to just one or the other.[130]

While modern humans share some nuclear DNA with the extinct Neanderthals, the two species do not share any mitochondrial DNA,[131] which in primates is always maternally transmitted. This observation has prompted the hypothesis that whereas female humans interbreeding with male Neanderthals were able to generate fertile offspring, the progeny of female Neanderthals who mated with male humans were either rare, absent or sterile.[132] However, some researchers have argued that there is evidence of possible interbreeding between female Neanderthals and male modern humans.[133][134]

Specimens[edit]

Notable Specimens[edit]

  • Neanderthal 1: The first Neanderthal specimen found during an archaeological dig in August 1856. It was discovered in a limestone quarry at the Feldhofer grotto in Neanderthal, Germany. The find consisted of a skull cap, two femora, three right arm bones, two left arm bones, ilium, and fragments of a scapula and ribs.
  • 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.)
  • 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.
  • Le Moustier: A fossilized skull, discovered in 1909, at the archaeological 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.
Type Specimen, Neanderthal 1
  • Shanidar 1: Found in the Zagros Mountains in (Iraqi Kurdistan); a total of nine skeletons found believed to have lived in the Middle Paleolithic. One of the nine remains was missing part of its right arm, which is theorized to have been broken off or amputated. The find is also significant because it shows that stone tools were present among this tribe's culture. One of the skeletons was buried with flowers, signifying that some type of burial ceremony may have occurred.
  • Amud 1: Fossilized remains of an adult Neanderthal, dated to roughly 45,000 years ago, and one of several found in a cave at Nahal Amud, Israel, at least some of which may have been deliberately buried. A particularly notable feature of this find is its cranial capacity, which, at 1,740 cm3, is among the largest known for any hominid, living or extinct.[9][135]

Chronology[edit]

This section describes bones with Neanderthal traits in chronological order.

Mixed with H. heidelbergensis traits[edit]

Typical H. neanderthalensis traits[edit]

Homo sapiens with some neanderthal-like archaic traits[edit]

Popular culture[edit]

Neanderthals have been portrayed in popular culture including appearances in literature, visual media and comedy, often in an unflattering and inaccurate light.

Early artistic reconstructions mostly presented Neanderthals as beastly creatures, emphasizing hairiness and rough, dark complexion.[146] More recent reconstructions acknowledge that because of the lineage evolution in European latitude there is reason to believe that Neanderthals were fair-skinned and probably with no more facial hair than modern man. Archaeological evidence exists indicating that they probably communicated by speech and used tools. Artist renderings and reconstructions of Neanderthals have become much more intelligent-looking and closely resembling modern humans.[147][148]

See also[edit]

Lists:

References[edit]

  1. ^ Dictionary of Anthropology - Charles Winick - Google Books. Books.google.ca (1956-12-18). Retrieved on 2014-05-24.
  2. ^ Bibliography of Fossil Vertebrates 1954-1958 - C.L. Camp, H.J. Allison, and R.H. Nichols - Google Books. Books.google.ca. Retrieved on 2014-05-24.
  3. ^ Neanderthal. (2012). Dictionary.com. Retrieved September 7, 2012, from link.
  4. ^ Colin P.T. Baillie; University of California, Berkeley. "Neandertals: Unique from Humans, or Uniquely Human?". berkeley.edu. 
  5. ^ Smithsonian Institution National Museum of American History. "Ancient DNA and Neanderthals". si.edu. 
  6. ^ Eran Meshorer, Liran Carmel, et al. (2014). "Reconstructing the DNA Methylation Maps of the Neandertal and the Denisovan". Science. doi:10.1126/science.1250368. 
  7. ^ Hublin, J. J. (2009). "The origin of Neandertals". Proceedings of the National Academy of Sciences 106 (38): 16022–7. Bibcode:2009PNAS..10616022H. doi:10.1073/pnas.0904119106. JSTOR 40485013. PMC 2752594. PMID 19805257. 
  8. ^ Bischoff, James L.; Shamp, Donald D.; Aramburu, Arantza; Arsuaga, Juan Luis; Carbonell, Eudald; Bermudez de Castro, J.M. (2003). "The Sima de los Huesos Hominids Date to Beyond U/Th Equilibrium (>350kyr) and Perhaps to 400–500kyr: New Radiometric Dates". Journal of Archaeological Science 30 (3): 275–80. doi:10.1006/jasc.2002.0834. 
  9. ^ a b c d e Papagianni, Dmitra; Morse, Michael (2013). The Neandethals Rediscovered. Thames & Hudson. ISBN 978-0-500-05177-1. 
  10. ^ [1], BBC
  11. ^ Skinner, A., B. Blackwell, R. Long, M.R. Seronie-Vivien, A.-M. Tillier and J. Blickstein; New ESR dates for a new bone-bearing layer at Pradayrol, Lot, France; Paleoanthropology Society March 28, 2007
  12. ^ Finlayson, C; Pacheco, Fg; Rodríguez-Vidal, J; Fa, Da; Gutierrez, López, Jm; Santiago, Pérez, A; Finlayson, G; Allue, E; Baena, Preysler, J; Cáceres, I; Carrión, Js; Fernández, Jalvo, Y; Gleed-Owen, Cp; Jimenez, Espejo, Fj; López, P; López, Sáez, Ja; Riquelme, Cantal, Ja; Sánchez, Marco, A; Guzman, Fg; Brown, K; Fuentes, N; Valarino, Ca; Villalpando, A; Stringer, Cb; Martinez, Ruiz, F; Sakamoto, T (October 2006). "Late survival of Neanderthals at the southernmost extreme of Europe". Nature 443 (7113): 850–3. Bibcode:2006Natur.443..850F. doi:10.1038/nature05195. ISSN 0028-0836. PMID 16971951. 
  13. ^ Outside Europe, Mousterian tools were made by both Neanderthals and early modern Homo sapiens. (Donald Johanson & Blake Edgar (2006) From Lucy to Language, Simon & Schuster, p. 272)
  14. ^ "Neanderthal man". infoplease. 
  15. ^ Homo neanderthalensis - H. neanderthalensis is a widely known but poorly understood hominid ancestor. Archaeologyinfo.com. Retrieved on 2014-05-24.
  16. ^ a b Helmuth H (1998). "Body height, body mass and surface area of the Neanderthals". Zeitschrift Für Morphologie Und Anthropologie 82 (1): 1–12. PMID 9850627. 
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  145. ^ Hayes, Jacqui (2 November 2006). "Humans and Neanderthals interbred". Cosmos. Retrieved 17 May 2009. 
  146. ^ Neanderthal image by Kupka, based on Boule, 1909, in Humanity's Journeys Dr. Kathryn Denning, 2005, retrieved 2012-03-17
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Neanderthal behavior

Neanderthal cranial anatomy.

Neanderthal behavior is subject to much study and speculation. Neanderthals were thought to be almost exclusively carnivorous[1] and apex predators.[2] However, new studies indicate that they had cooked vegetables in their diet.[3][4] They made advanced tools, had language (the nature of which is debated) and lived in complex social groups.


Neanderthal brains were somewhat larger than humans but were shaped a bit differently[5] since they evolved separately for several hundred thousand years. The size and distribution of archaeological sites shows that Neanderthals spent their lives mostly in small groups of 5-10 individuals (compared to 20-30 individuals for Cro-Magnon man). Elders were rare as few Neanderthals lived past 35.

Skeletal evidence shows that injured individuals were often nursed back to health by others. Neanderthals rarely made contact with outsiders or traveled outside their small home territories. Although many Neanderthal sites have rare pieces of high-quality stone from more than 100 kilometers away, there is not enough to indicate trade or even regular contact with other communities.

It has been suggested by one pair of researchers that these stones may instead be "gifts" brought by adolescents wishing to join a new community (some form of "marrying out" was essential due to the small size of Neanderthal territories). In their view, this lack of trade could indicate that Neanderthals may have lacked some cognitive abilities for dealing with strangers, such as "cheater detection" and the ability to judge the value of one commodity in terms of another. Neanderthals had a smaller cognitive part of the brain and this would have limited them, including their ability to form larger groups.[6] The quality of tools found at archaeological sites is further said to suggest that Neanderthals were good at "expert" cognition, a form of observational learning and practice acquired through apprenticeship that relies heavily on long-term procedural memory.[7]

Neanderthal toolmaking supposedly changed little over hundreds of thousands of years. The lack of innovation was said to imply they may have had a reduced capacity for thinking by analogy and less working memory. The researchers further speculated that Neanderthal behavior would probably seem neophobic, dogmatic and xenophobic to modern humans.[7]

Language[edit]

There is some circumstantial evidence for thinking that Neanderthals had language with words and some kind of syntax; some of their tool-making and hunting tactics would have been difficult to learn and execute without it.[8] 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.[9]

The idea that Neanderthals lacked complex language was once widespread,[10] despite concerns about the accuracy of reconstructions of the Neanderthal vocal tract, until 1983, when a Neanderthal hyoid bone was found at the Kebara Cave in Israel. The hyoid is a small bone that connects the musculature of the tongue and the larynx, and by bracing these structures against each other, allows a wider range of tongue and laryngeal movements than would otherwise have been possible. The presence of this bone implies that structured speech was anatomically possible and that the repertory of sounds that might be produced was wide enough to contain well-defined sets of phonemes, and not simply inarticulate guttural grunts. The bone found is virtually identical to that of modern humans.[11]

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.[12]

Neurological evidence for potential speech in neanderthalensis may exist in the form of the hypoglossal canal. The canal of neanderthalensis is the same size or larger than in modern humans, which is significantly larger than the canal of australopithecines and modern chimpanzees. The canal carries the hypoglossal nerve, which controls the muscles of the tongue. This suggests to some theorists that neanderthalensis had vocal capabilities similar to modern humans.[13]

A research team from the University of California, Berkeley, led by David DeGusta, however, argues 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 that have an equal or larger hypoglossal canal.[14]

Another anatomical difference between Neanderthals and modern humans is the former's lack of a mental protuberance (the point at the tip of the chin). This may be relevant to speech, as the mentalis muscle contributes to moving the lower lip and is used to articulate a bilabial click. While some Neanderthal individuals do possess a mental protuberance, their chins never show the inverted T-shape of modern humans.[15] In contrast, some Neanderthal individuals show inferior lateral mental tubercles (small bumps at the side of the chin).

Steven Mithen (2006) speculates that the Neanderthals may have had an elaborate proto-linguistic system of communication that was more musical than modern human language, and that pre-dated the separation of language and music into two separate modes of cognition. He called this hypothetical lingual system 'hmmmmm' because it would be Holistic, manipulative, multi-modal, musical and mimetic.[16]

Tools[edit]

Neanderthal and Middle Paleolithic archaeological sites show a smaller and different toolkit than those found in Upper Paleolithic sites, which were perhaps occupied by modern humans that superseded them. Fossil evidence indicating who may have made the tools found in Early Upper Paleolithic sites is still missing. A survey of 332 archeological sites occupied over a period of 200,000 years under varying climatic conditions using lithic tool data from 190 layers at 103 sites showed that the Neanderthal toolkit changed little, showing technological inertia, a slower rate of variability compared to modern humans whose toolkits show more economic reactivity, variety in response to changing conditions. In addition to the obvious[to whom?] hypothesis that Neanderthals were not very creative despite having larger brains than modern humans, an alternate demographic hypothesis is that there were never very many Neanderthals, perhaps less than 10,000, making the probability of innovation low.[17]

An artist's impression of Neanderthal life.

Neanderthals 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 stone-flakes, task-specific hand axes, and spears. Many of these tools were very sharp. There is also good evidence that they used a lot of wood, but wooden objects are unlikely to have survived to the present.[18]

Neanderthals were capable of building dugout boats since the Middle Paleolithic.[19][20] Mousterian stone tools discovered on the southern Ionian Greek islands suggests that Neanderthals were navigating the Mediterranean Sea as early as 110,000 years BP.[21][22] Quartz hand-axes, three-sided picks, and stone cleavers from Crete have also been recovered that date back about 170,000 years BP.[23]

There is some evidence for interpersonal violence among Neanderthals. A 36,000-year-old Neanderthal skull found near St. Césaire has a healed fracture in its cranial vault that was most likely caused by the impact of a sharp implement. The location of the wound suggests interpersonal violence rather than an accident. Because the wound healed, it is known that the individual survived the attack.[24]

Also, while they had weapons, whether they had projectile weapons is controversial. They had spears, made of long wooden shafts with spearheads firmly attached, but some think these were thrusting spears and not projectiles.[25] Still, a Levallois point embedded in an animal vertebra shows an angle of impact suggesting that it entered by a "parabolic trajectory" suggesting that it was the tip of a projectile.[26]

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 Neanderthals’ ancestors, Homo erectus or Homo heidelbergensis. Generally, projectile weapons are more commonly associated with H. sapiens. The lack of projectile weaponry may be an indication of different sustenance methods, rather than inferior technology or abilities. The situation is identical to that of native New Zealand Māori—modern Homo sapiens, who also rarely threw objects, but used spears and clubs instead.[27] Neanderthals rarely produced bone tools from their prey. Neanderthals apparently did not have needles, but at best, bone awls to drill eyelets for lacing skins and furs together.[28] Some tools may have been due to trade or copying from Homo sapiens who coexisted with Neanderthals near the end of the latter's existence.[29]

Neanderthals also performed many sophisticated tasks normally associated only with modern humans. For example, 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.[30]

Particularly intriguing is a hollowed-out bear femur with holes that may have been deliberately bored into it, known as the Divje Babe flute. This bone was found in western Slovenia in 1995, near a Mousterian fireplace, but its significance is still disputed. Some paleoanthropologists hypothesize it was a musical instrument, others believe it was not the work of Neanderthals, or that the chomping action of another bear made the holes.

Pendants and other jewelry showing traces of ochre dye and of deliberate grooving have also been found[31] with later finds, particularly in France, but whether they were created by Neanderthals or traded to them by Cro-Magnons is a matter of controversy.

Burial practices[edit]

Although much has been made of the Neanderthals' burial of their dead, their burials were less elaborate than those of anatomically modern humans. The interpretation of the Shanidar IV burials as including flowers, and therefore being a form of ritual burial,[32] has been questioned.[33] On the other hand, five of the six flower pollens found with Shanidar IV are known to have had 'traditional' medical uses, even among relatively recent 'modern' populations. In some cases Neanderthal burials include grave goods, such as bison and aurochs bones, tools, and the pigment ochre although the evidence for this is disputed.[34]

Diet[edit]

Early studies indicated that Neanderthals were highly carnivorous and obtained most of the protein in their diet from animal sources.[35] Recently, however, traces of fossilized plants have been extracted from Neanderthal teeth found in Belgium, Iraq, and East Africa indicating they also ate plants such as grains and legumes in addition to meat.[36]

Cannibalism or ritual defleshing?[edit]

Neanderthal Burial of Kebara Cave (Carmel Range, Israel). Thermoluminescence dates place Neanderthal levels at Kebara at ca. 60,000 BP. Skeleton of an adult man nicknamed Moshe (25–35 years old, height 1.70 m) found in 1983

Neanderthals hunted large animals, such as the mammoth. Stone-tipped wooden spears were used for hunting and stone knives and poleaxes were used for butchering the animals or as weapons. However, they are believed to have practiced cannibalism or ritual defleshing. This hypothesis was formulated after researchers found marks on Neanderthal bones similar to the bones of a dead deer butchered 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.[37] However, the results of technological tests have revealed 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 brain) 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 similar to those 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.[38]
  • 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.[39]

Evidence indicating cannibalism would not distinguish Neanderthals from modern Homo sapiens. Ancient and existing Homo sapiens are known to have practiced cannibalism and/or mortuary defleshing (e.g., the sky burial of Tibet).

Grooves in bones are hypothesized to be cuts by Neanderthal tools, not animal teeth. The chances of them being random, as some writers attributing them to animals have proposed, is debated.

Body paint[edit]

A 2009 report in the Proceedings of the National Academy of Sciences (PNAS) on two archaeological sites in the Murcia province of southern Spain records the discovery of shells showing pigment residues, and concludes that these were used by the Neanderthals as make-up containers. Sticks of the black pigment manganese have previously been discovered in Africa. These may have been used as body paint by Neanderthals.[40] According to one study, Neanderthal use of pigments including red ochre was restricted to interglacial periods, disappearing during glacial periods.[41]

Cave art[edit]

Recent new dating techniques of cave art has suggested the possibility Neanderthals were cave painters. Many cave paintings are much older than previously thought, and possibly pre-date the arrival of H. sapiens.[42] Symbolism or simple art work has been found in 2012, inside the Gorham's Cave in Gibraltar; it may have been a sign indicating to others that the cave was already inhabited.[43]

Dwellings[edit]

While some Neanderthals used caves for shelter, the Molodova archaeological site in eastern Ukraine suggests others built dwellings using animal bones. A building was made of mammoth skulls, jaws, tusks and leg bones, and had 25 hearths inside.[44]

References[edit]

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References[edit]

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Neanderthal 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, particularly in certain isolated geographic regions. Evidence suggests they were much stronger than modern humans[2] while they were comparable in height; based on 45 long bones from at most 14 males and 7 females, Neanderthal males averaged 164–168 cm (65–66 in) and females 152–156 cm (60–61 in) tall.[3] 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.[4] A 2007 genetic study suggested some Neanderthals may have had red hair.[5][6]

Distinguishing physical traits[edit]

Ashley Montagu claimed humans have more neotenized skulls than Neanderthals.[7]
Neanderthal cranial anatomy
Neanderthal footprint in the Natural History Museum in Prague
A reconstruction of a Neanderthal male at Zagros Paleolithic Museum (2008)

The magnitude of autapomorphic traits in specimens differ in time. In the latest specimens, autapomorphy is unclear. The following is a list of physical traits that distinguish Neanderthals from modern humans. However, not all of them distinguish specific Neanderthal populations from various geographic areas, evolutionary periods, or other extinct humans. Also, many of these traits occasionally manifest in modern humans, particularly among certain ethnic groups traced to Neanderthal habitat ranges.[citation needed] Nothing is certain (from unearthed bones) about the shape of soft parts such as eyes, ears, and lips of Neanderthals.[8]

While the structure of the head and face were not very far removed from those of modern humans, there were still quite noticeable differences. Notably the modern human head is shorter, with a less pronounced facial front. The Neanderthal chin and forehead sloped backwards and the nose region protruded forward more than in modern humans. The common shapes of the nose are not known but in general it was likely more robust, and possibly slightly larger, than in modern humans. The brain space of the skull, and so most likely the brain itself, were larger than in modern humans.

When comparing traits to worldwide average present day human traits in Neanderthal specimens, the following traits are distinguished. The magnitude on particular trait changes with 300,000 years timeline. The large number of classic Neanderthal traits is significant because extreme examples of Homo sapiens may sometimes show one or more of these traits, but not most or all of them.

Cold-adapted theory[edit]

Some people thought that the large Neanderthal noses were an adaptation to the cold,[14] but primate and arctic animal studies have shown sinus size reduction in areas of extreme cold rather than enlargement in accordance with Allen's rule.[15] Todd C. Rae summarizes explanations about Neanderthal anatomy as trying to find explanations for the "paradox" that their traits are not cold-adapted.[15] Therefore Rae concludes that the design of the large and extensive Neanderthal nose was evolved for the hotter climate of the Middle East and went unchanged when the Neanderthals entered Europe.[15]

Pathology[edit]

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

Fractures[edit]

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 1), spine (Kebara 2) and skull (Shanidar I, Krapina, Sala 1). These fractures are often healed and show little or no sign of infection, suggesting that injured individuals were cared for during times of incapacitation. It has been remarked that Neanderthals showed a frequency of such injuries comparable to that of modern rodeo professionals, showing frequent contact with large, combative mammals. 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.[16]

Trauma[edit]

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

Degenerative disease[edit]

Arthritis was 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).

Developmental Stress[edit]

Neanderthal child

Two non-specific indicators of stress during development are found in teeth, which record stresses, such as periods of food scarcity or illness, that disrupt normal dental growth. One indicator is enamel hypoplasia, which appears as pits, grooves, or lines in the hard enamel covering of teeth. The other indicator, fluctuating asymmetry, manifests as random departures from symmetry in paired biological structures (such as right and left teeth). Teeth do not grow in size after they form nor do they produce new enamel, so enamel hypoplasia and fluctuating asymmetry provide a permanent record of developmental stresses occurring in infancy and childhood. A study of 669 Neanderthal crowns showed that 75% of individuals suffered some degree of hypoplasia. Two studies,[17][18] compared Neanderthals with the Tigara, coastal whale-hunting people from Point Hope Alaska, finding comparable levels of linear enamel hypoplasia (a specific form of hypoplasia) and higher levels of fluctuating asymmetry in Neanderthals.[non-primary source needed] Estimated stress episode duration from Neanderthal linear enamel hyoplasias suggest that Neandertals experienced stresses lasting from two weeks to up to three months.[citation needed]

Infection[edit]

Evidence of infections on Neanderthal skeletons is usually visible in the form of lesions on the bone, which are created by systemic 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).

Childhood[edit]

Neanderthal children may have grown faster than modern human children. Modern humans have the slowest body growth of any mammal during childhood (the period between infancy and puberty) with lack of growth during this period being made up later in an adolescent growth spurt.[19][20][21] The possibility that Neanderthal childhood growth was different was first raised in 1928 by the excavators of the Mousterian rock-shelter of a Neanderthal juvenile.[22] Arthur Keith in 1931 wrote, "Apparently Neanderthal children assumed the appearances of maturity at an earlier age than modern children."[23] The rate of body maturation can be inferred by comparing the maturity of a juvenile's fossil remains and the estimated age of death.

The age at which juveniles die can be indirectly inferred from their tooth morphology, development and emergence. This has been argued to both support[24] and question[25][26] the existence of a maturation difference between Neanderthals and modern humans. Since 2007, tooth age can be directly calculated using the noninvasive imaging of growth patterns in tooth enamel by means of x-ray synchrotron microtomography.[27]

This research supports the occurrence of much more rapid physical development in Neanderthals than in modern human children.[28] The x-ray synchrotron microtomography study of early H. sapiens sapiens argues that this difference existed between the two species as far back as 160,000 years before present.[29]

Footnotes[edit]

  1. ^ Bibliography of Fossil Vertebrates 1954-1958 - C.L. Camp, H.J. Allison, and R.H. Nichols - Google Books. Books.google.ca. Retrieved on 2014-05-24.
  2. ^ "Neanderthal". BBC. Retrieved 18 May 2009. 
  3. ^ Helmuth H (1998). "Body height, body mass and surface area of the Neanderthals". Zeitschrift Für Morphologie Und Anthropologie 82 (1): 1–12. PMID 9850627. 
  4. ^ Froehle, Andrew W; Chruchill, Steven E (2009). "Energetic Competition Between Neandertals and Anatomically Modern Humans" (PDF). PaleoAnthropology: 96–116. Retrieved 2011-10-31. 
  5. ^ Laleuza-Fox, Carles; Holger Römpler et al. (2007-10-25). "A Melanocortin 1 Receptor Allele Suggests Varying Pigmentation Among Neanderthals". Science 318 (5855): 1453–5. doi:10.1126/science.1147417. PMID 17962522. 
  6. ^ Rincon, Paul (25 October 2007). "Neanderthals 'were flame-haired'". BBC News. Retrieved 25 October 2007. 
  7. ^ a b Montagu, A. (1989). Growing Young. Bergin & Garvey: CT.[page needed]
  8. ^ Sawyer GJ, Maley B (March 2005). "Neanderthal reconstructed". Anatomical Record. Part B, New Anatomist 283 (1): 23–31. doi:10.1002/ar.b.20057. PMID 15761833. Lay summaryLiveScience (March 10, 2005). 
  9. ^ Gunz P, Harvati K (March 2007). "The Neanderthal 'chignon': variation, integration, and homology". Journal of Human Evolution 52 (3): 262–74. doi:10.1016/j.jhevol.2006.08.010. PMID 17097133. 
  10. ^ http://www.pajamacore.org/writings/origins.php[dead link]
  11. ^ Bower, Bruce (11 April 1992). "Neanderthals to investigators: can we talk? – vocal abilities in pre-historic humans". Science News. Retrieved 29 Apr 2014. 
  12. ^ Arensburg B, Schepartz LA, Tillier AM, Vandermeersch B, Rak Y (October 1990). "A reappraisal of the anatomical basis for speech in Middle Palaeolithic hominids". American Journal of Physical Anthropology 83 (2): 137–46. doi:10.1002/ajpa.1330830202. PMID 2248373. 
  13. ^ a b De Groote I (October 2011). "The Neanderthal lower arm". Journal of Human Evolution 61 (4): 396–410. doi:10.1016/j.jhevol.2011.05.007. PMID 21762953. 
  14. ^ Finlayson, C (2004). Neanderthals and modern humans: an ecological and evolutionary perspective. Cambridge University Press. pp. 84. ISBN 0-521-82087-1. 
  15. ^ a b c Rae TC, Koppe T, Stringer CB (February 2011). "The Neanderthal face is not cold adapted". Journal of Human Evolution 60 (2): 234–9. doi:10.1016/j.jhevol.2010.10.003. PMID 21183202. 
  16. ^ T.D. Berger and E. Trinkaus (1995). "Patterns of trauma among Neadertals". Journal of Archaeological Science 22 (6): 841–852. doi:10.1016/0305-4403(95)90013-6. Retrieved 2007-06-28. 
  17. ^ Guatelli-Steinberg D, Larsen CS, Hutchinson DL (2004). "Prevalence and the duration of linear enamel hypoplasia: a comparative study of Neandertals and Inuit foragers". Journal of Human Evolution 47 (1-2): 65–84. doi:10.1016/j.jhevol.2004.05.004. PMID 15288524. 
  18. ^ Barrett CK, Guatelli-Steinberg D, Sciulli PW (October 2012). "Revisiting dental fluctuating asymmetry in neandertals and modern humans". American Journal of Physical Anthropology 149 (2): 193–204. doi:10.1002/ajpa.22107. PMID 22791408. 
  19. ^ Walker R, Hill K, Burger O, Hurtado AM (April 2006). "Life in the slow lane revisited: ontogenetic separation between chimpanzees and humans". American Journal of Physical Anthropology 129 (4): 577–83. doi:10.1002/ajpa.20306. PMID 16345067. 
  20. ^ Bogin, Barry (1997). m682 "Evolutionary hypotheses for human childhood". American Journal of Physical Anthropology 104 (S25): 63–89. doi:10.1002/(SICI)1096-8644(1997)25+<63::AID-AJPA3>3.0.CO;2-8. 
  21. ^ Bogin, Barry (1999). Patterns of human growth. Cambridge: Cambridge University Press. ISBN 0-521-56438-7. OCLC 39692257. [page needed]
  22. ^ Garrod, D. A. E., Buxton, L. H. D., Elliot-Smith, G., Bate, D.M. A. (1928). "Excavation of a Mousterian rock-shelter at Devil's Tower, Gibraltar". Journal of the Royal Anthropological Institute 58: 33–113. JSTOR 4619528. 
  23. ^ Keith, Arthur (1931). New discoveries relating to the antiquity of man. London: W. W. Norton & Company. p. 346. OCLC 3665578. 
  24. ^ Ramirez Rozzi FV, Bermudez De Castro JM (April 2004). "Surprisingly rapid growth in Neanderthals". Nature 428 (6986): 936–9. doi:10.1038/nature02428. PMID 15118725. 
  25. ^ Macchiarelli R, Bondioli L, Debénath A, et al. (December 2006). "How Neanderthal molar teeth grew". Nature 444 (7120): 748–51. doi:10.1038/nature05314. PMID 17122777. 
  26. ^ Guatelli-Steinberg D, Reid DJ, Bishop TA, Larsen CS (October 2005). "Anterior tooth growth periods in Neandertals were comparable to those of modern humans". Proceedings of the National Academy of Sciences of the United States of America 102 (40): 14197–202. doi:10.1073/pnas.0503108102. PMC 1242286. PMID 16183746. 
  27. ^ Tafforeau P, Smith TM (February 2008). "Nondestructive imaging of hominoid dental microstructure using phase contrast X-ray synchrotron microtomography". Journal of Human Evolution 54 (2): 272–8. doi:10.1016/j.jhevol.2007.09.018. PMID 18045654. 
  28. ^ Smith TM, Toussaint M, Reid DJ, Olejniczak AJ, Hublin JJ (December 2007). "Rapid dental development in a Middle Paleolithic Belgian Neanderthal". Proceedings of the National Academy of Sciences of the United States of America 104 (51): 20220–5. doi:10.1073/pnas.0707051104. PMC 2154412. PMID 18077342. 
  29. ^ Smith TM, Tafforeau P, Reid DJ, et al. (April 2007). "Earliest evidence of modern human life history in North African early Homo sapiens". Proceedings of the National Academy of Sciences of the United States of America 104 (15): 6128–33. doi:10.1073/pnas.0700747104. PMC 1828706. PMID 17372199. 
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Neanderthal

The Neanderthal (pronounced /niːˈændərtɑːl/, /niːˈændərθɔːl/), or /neɪˈændərtɑːl/),[1] also spelled Neandertal,[2] is an extinct member of the Homo genus that is known from Pleistocene specimens found in Europe and parts of western and central Asia. Neanderthals are either classified as a subspecies (or race) of humans (Homo sapiens neanderthalensis) or as a separate species (Homo neanderthalensis).[3] The first proto-Neanderthal traits appeared in Europe as early as 600,000–350,000 years ago.[4] Current (as of 2010) genetic evidence suggests interbreeding took place with Homo sapiens during the period of some 80,000 to 50,000 years ago, and it is now believed that humans of Eurasian descent are anywhere netween 1% and 4% Neanderthal.

Proto-Neanderthal traits are occasionally grouped to another phenetic 'species', Homo heidelbergensis, or a migrant form, Homo rhodesiensis. By 130,000 years ago, complete Neanderthal characteristics had appeared. These characteristics then disappeared in Asia by 50,000 years ago and in Europe by 30,000 years ago.[5]

The youngest Neanderthal finds include Hyaena Den (UK), considered older than 30,000 years ago, while the Vindija (Croatia) Neanderthals have been re-dated to between 32,000 and 33,000 years ago. No definite specimens younger than 30,000 years ago have been found; however, evidence of fire by Neanderthals at Gibraltar indicate that they may have survived there until 24,000 years ago. Cro-Magnon or early modern human skeletal remains with 'Neanderthal traits' were found in Lagar Velho (Portugal), dated to 24,500 years ago and controversially interpreted as indications of extensively admixed populations.[6]

Neanderthal stone tools provide further evidence for their presence where skeletal remains have not been found. The last traces of Mousterian culture, a type of stone tools associated with Neanderthals, were found in Gorham's Cave on the remote south-facing coast of Gibraltar.[7] Other tool cultures sometimes associated with Neanderthal include Châtelperronian, Aurignacian, and Gravettian, with the latter extending to 22,000 years ago, the last indication of Neanderthal presence.

Neanderthal cranial capacity is thought to have been as large as that of Homo sapiens, perhaps larger, indicating that their brain size may have been comparable as well. In 2008, a group of scientists made a study using three-dimensional computer-assisted reconstructions of Neanderthal infants based on fossils found in Russia and Syria, showing that they had brains as large as ours at birth and larger than ours as adults.[8] On average, the height of Neanderthals was comparable to contemporaneous Homo sapiens. Neanderthal males stood about 165–168 cm (65–66 in), and were heavily built with robust bone structure. They were much stronger, having particularly strong arms and hands.[9] Females stood about 152–156 cm (60–61 in).[10] They were almost exclusively carnivorous[11] and apex predators.[12]

Contents

Etymology

The Neanderthal is named after the Neandertal valley, originally spelled Neanderthal, which is located about 12 km (7.5 mi) east of Düsseldorf. The spelling of the German word Thal ("valley"), was changed to Tal in 1901, and the spelling of the valley was also changed accordingly to Neandertal. The former spelling is however often retained in English for the hominid. The spelling with th is in addition always used in scientific names throughout the world. In German, the modern spelling with t is however otherwise used in referring both to the hominid and the valley.

The valley itself was named after the theologian Joachim Neander, who lived nearby in Düsseldorf in the late 17th century. "Neander" is a classicized form of the common German surname Neumann. In turn, the hominid was named after the valley, where the first Neanderthal remains were found. The term Neanderthal Man was coined in 1863 by the Anglo-Irish geologist William King.

The German pronunciation (regardless of spelling) is with the sound /t/. American English speakers commonly pronounce it as /θ/ (th as in thin), but American scientists usually use /t/. British English speakers usually pronounce it as /t/ followed by a long a as in tar,[1][13] matching the German pronounciation.

Classification

First reconstruction of Neanderthal man

For some time, scientists have debated whether Neanderthals should be classified as Homo neanderthalensis or as Homo sapiens neanderthalensis, the latter placing Neanderthals as a subspecies of Homo sapiens[14]. Some morphological studies support that Homo neanderthalensis is a separate species and not a subspecies.[15] Others, for example University of Cambridge Professor Paul Mellars, say "no evidence has been found of cultural interaction"[16] and evidence from mitochondrial DNA studies have been interpreted as evidence Neanderthals were not a subspecies of H. sapiens.[17]

The current state of sequence analysis of the Neanderthal genome suggest that there was no recent genetic exchange between Neanderthals and humans; previous results which showed some similarities have now been conclusively shown to be the result of contamination or improper phylogenetic assumptions. Consequently, Homo neanderthalensis at present appears to be the correct nomenclature.[18][19]

Neanderthals evolved from African apes along a path similar to that of humans. Sometime between 5 and 10 million years ago a common ancestral species between chimps and humans lived in Africa. The ancestor evolved along a path that might include Ardipithecus kadabba, Ardipithicus ramidus, Australopithecus anamensis, Australopithecus afarensis, Homo habilis, Homo ergaster (or Homo erectus). The last common ancestor between anatomically modern Homo sapiens and Neanderthals appears to be an African variant of Homo heidelbergensis known as Homo rhodesiensis, named after an archaic Homo sapiens, Broken hill 1 (Kabwe 1) discovered in the territory of Rhodesia in 1921.

Homo rhodesiensis arose in Africa an estimated 0.7 to 1 million years ago. The earliest estimates for Homo rhodesiensis reaching Europe are approximately 800 thousand years ago when a type of human referred to as Homo antecessor or Homo cepranensis already inhabited the region. These two human types may be forerunners to European Homo heidelbergensis, however stone tools dating from 1.2 to 1.56 million years ago of an unknown creator have been discovered in Southwestern Europe. The evidence at the Sima de los Huesos (in the Atapuerca cave system on the Iberian Peninsula) suggest that Homo heidelbergensis was already in Europe by 600,000 years ago.

Molecular phylogenetic analysis suggests that Homo rhodesiensis and Homo heidelbergensis continued to intermix until 350,000 years ago, after which they were separate species and sometime within the last 200,000 years Homo heidelbergensis evolved into Homo neanderthalensis, the classic Neanderthal man. If it is proven by further scientific research that Neanderthals provided no significant genetic input into modern populations of Homo sapiens, then it must be assumed that Neanderthal in fact is more distantly related to today's human than is Homo heidelbergensis.

Discovery

The site of Kleine Feldhofer Grotte where the first Neanderthal was unearthed by miners in the nineteenth century
Location of Neander Valley, Germany. (The highlighted area is the modern federal state of North Rhine-Westphalia.)

Neanderthal skulls were first discovered in Engis, Belgium (1829) by Philippe-Charles Schmerling and in Forbes' Quarry, Gibraltar (1848), both prior to the "original" discovery in a limestone quarry of the Neander Valley in Erkrath near Düsseldorf in August, 1856, three years before Charles Darwin's On the Origin of Species was published.[20]

The type specimen, dubbed Neanderthal 1, consisted of a skull cap, two femora, three bones from the right arm, two from the left arm, part of the left ilium, fragments of a scapula, and ribs. The workers who recovered this material originally thought it to be the remains of a bear. They gave the material to amateur naturalist Johann 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 Europeans who had played an important role in modern human origins. The bones of over 400 Neanderthals have been found since.[citation needed]

Timeline

Skull, found in 1886 in Spy, Belgium
Frontal bone of a neanderthal child from the cave of La Garigüela
Skull from La Chapelle aux Saints
  • 1829: Neanderthal skulls were discovered in Engis, Belgium.
  • 1848: Neanderthal skull found in Forbes' Quarry, Gibraltar. Called "an ancient human" at the time.
  • 1856: Johann Karl Fuhlrott first recognised the fossil called “Neanderthal man”, discovered in Neanderthal, a valley near Mettmann in what is now North Rhine-Westphalia, Germany.
  • 1880: The mandible of a Neanderthal child was found in a secure context and associated with cultural debris, including hearths, Mousterian tools, and bones of extinct animals.
  • 1886: two nearly perfect skeletons of a man and woman were found at Spy, Belgium at the depth of 16 ft. with numerous Mousterian-type implements.
  • 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 Israeli fossils date Neanderthals at Kebara to 60,000 BP and humans at Qafzeh to 90,000 BP. These dates were confirmed by Electron Spin Resonance (ESR) dates for Qafzeh (90,000 BP) and Es Skhul (80,000 BP).
  • 1991: ESR dates showed that the Tabun Neanderthal was contemporaneous with modern humans from Skhul and Qafzeh.
  • 1997: Matthias Krings et al. are the first to amplify Neanderthal mitochondrial DNA (mtDNA) using a specimen from Feldhofer grotto in the Neander valley.[21]
  • 2000: Igor Ovchinnikov, Kirsten Liden, William Goodman et al. retrieved DNA from a Late Neanderthal (29,000 BP) infant from Mezmaikaya Cave in the Caucasus.[22]
  • 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.
  • 2009: The Max Planck Institute for Evolutionary Anthropology announced that the "first draft" of a complete Neanderthal genome is completed.[23]


Habitat and range

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. Countries where their remains are known include most of Europe south of the line of glaciation, roughly along the 50th parallel north, including most of Western Europe, including the south coast of Great Britain,[24] Central Europe and the Balkans,[25] some sites in the Ukraine and in western Russia and outside of Europe in the Zagros Mountains and in the Levant.

Neanderthal fossils have to date not been found in Africa, but rather close to Africa, both at Gibraltar and in the Levant. At some Levantine sites, Neanderthal remains in fact date after the same sites were vacated by Homo sapiens. Mammal fossils of the same time period show that cold-adapted animals were present alongside these Neanderthals in this region of the Eastern Mediterranean. This implies Neanderthals were better adapted biologically to cold weather than H. sap. and at times displaced H. sap. in parts of the Middle East when the climate got cold enough. Homo sapiens appears to have been the only human type in the Nile River Valley during these periods, and Neanderthals are not known to have ever lived southwest of modern Israel. When further climate change caused warmer temperatures, the Neanderthal range likewise retreated to the north along with the cold-adapted species of mammals. Apparently these weather-induced population shifts took place before "modern" people secured competitive advantages over the Neanderthal, as these shifts in range took place well over ten thousand years before "moderns" totally replaced Neanderthals.[26]

There were separate developments in the human line, in other regions such as Southern Africa, that somewhat resembled the European and Western/Central Asian Neanderthals, but these people were not actually Neanderthals. One such example is Rhodesian Man (Homo rhodesiensis) who existed long before any classic European Neanderthals, but had a more modern set of teeth, and arguably some H. Rhodesiensis populations were on the road to modern Homo sapiens sapiens.

To date, no intimate connection has been found between these similar people and the Western/Central Eurasian Neanderthals, at least during the same time as classic Eurasian Neanderthals, and H. rhodesiensis seems to have evolved separately and earlier than classic Neanderthals in a case of convergent evolution.

It appears incorrect, based on present research and known fossil finds, to refer to any fossil outside of Europe or Western and Central Asia as a true Neanderthal. True Neanderthals had a known range that possibly extended as far east as the Altai Mountains, but not farther to the east or south, and apparently not into Africa. At any rate, in Africa the land immediately south of the Neanderthal range was possessed by "modern" H. sap., since at least 160,000 years before the present.

Classic Neanderthal fossils have been found over a large area, from northern Germany to Israel and Mediterranean countries like Spain[27] and Italy[28] in the south and from England and Portugal 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° N, on the Russian plain.[29] Recent evidence has extended the Neanderthal range by about 1,250 miles (2,010 km) east into southern Siberia's Altay Mountains.[30][31]

Anatomy

Reconstruction of a Neanderthal child from Gibraltar (Anthropological Institute, University of Zürich)
Neanderthal cranial anatomy.

Neanderthals were generally only 12–14 cm (5–6 in) shorter than modern humans, contrary to a common view of them as "very short" or "just over 5 feet". Based on 45 long bones from (at most) 14 males and 7 females, Neanderthal males averaged 164–168 cm (65–66 in) and females 152–156 cm (60–61 in) tall. Compared to Europeans some 20,000 years ago, it is nearly identical, perhaps slightly taller. Considering the body build of Neanderthals, new body weight estimates show they are only slightly above the cm/weight or the body mass index of modern Americans or Canadians.[32]

Neanderthals had more robust build and distinctive morphological features, especially of the cranium, which gradually accumulated more derived aspects, particularly in certain relatively isolated geographic regions. Evidence suggests they were much stronger than modern humans;[33] their relatively robust stature is thought to be an adaptation to the cold climate of Europe during the Pleistocene epoch.

A 2007 study suggested some Neanderthals may have had red hair and pale skin color.[34][35]

Distinguishing physical traits

Anatomical comparison of the skulls of anatomically modern humans and homo neanderthalensis
Comparison of crania, sapiens (left) and neanderthalensis (right).
Neanderthal footprint in the Natural History Museum in Prague

The magnitude of autapomorphic traits in specimens differ in time. In the latest specimens, autapomorphy is fuzzy. 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 traced to Neanderthal habitat ranges.[citation needed] Nothing is certain (from unearthed bones) about the shape of soft parts such as eyes, ears, and lips of Neanderthals.[36]

When comparing traits to worldwide average present day human traits in Neanderthal specimens, the following traits are distinguished. The magnitude on particular trait changes with 300,000 years timeline. The large number of classic Neanderthal traits is significant because extreme examples of Homo sapiens may sometimes show one or more of these traits, but not most or all of them.

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 1), spine (Kebara 2) and skull (Shanidar I, Krapina, Sala 1). These fractures are often healed and show little or no sign of infection, suggesting that injured individuals were cared for during times of incapacitation. It has been remarked that Neanderthals showed a frequency of such injuries comparable to that of modern rodeo professionals, showing frequent contact with large, combative mammals. 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.[41]

Trauma

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

Degenerative disease

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

Hypoplastic disease

Dental enamel hypoplasia is an indicator of stress during the development of teeth and records in the striations and grooves in the enamel periods of food scarcity, trauma or disease. A study of 669 Neanderthal dental crowns showed that 75% of individuals suffered some degree of hypoplasia and 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).

Childhood

Neanderthal child

Neanderthal children may have grown faster than modern human children. Modern humans have the slowest body growth of any mammal during childhood (the period between infancy and puberty) with lack of growth during this period being made up later in an adolescent growth spurt.[42][43][44] The possibility that Neanderthal childhood growth was different was first raised in 1928 by the excavators of the Mousterian rock-shelter of a Neanderthal juvenile.[45] Arthur Keith in 1931 wrote, "Apparently Neanderthal children assumed the appearances of maturity at an earlier age than modern children."[46] The earliness of body maturation can be inferred from the maturity of a juvenile's fossile remains and estimated age of death.

The age at which juveniles die can be indirectly inferred from their tooth morphology, development and emergence. This has been argued to both support[47] and question[48] the existence of a maturation difference between Neanderthals and modern humans. Since 2007 tooth age can be directly calculated using the noninvasive imaging of growth patterns in tooth enamel by means of x-ray synchrotron microtomography.[49]

This research supports the existence of a much quicker physical development in Neanderthals than in modern human children.[50] The x-ray synchrotron microtomography study of early H. sapiens sapiens argues that this difference existed between the two species as far back as 160,000 years before present.[51]

Behaviour

Language

The idea that Neanderthals lacked complex language was widespread,[52] 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.[53]

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.[54]

Neurological evidence for potential speech in neanderthalensis exists in the form of the hypoglossal canal. The canal of neanderthalensis is the same size or larger than in modern humans, which are significantly larger than the canal of australopithecines and modern chimpanzees. The canal carries the hypoglossal nerve, which controls the muscles of the tongue. This indicates that neanderthalensis had vocal capabilities similar to modern humans.[55] 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.[56]

Another anatomical difference between Neanderthals and modern humans is the former's lack of a mental protuberance (the point at the tip of the chin). This may be relevant to speech as the mentalis muscle contributes to moving the lower lip and is used to voice a bilabial click. While some Neanderthal individuals do possess a mental protuberance, their chins never show the inverted T-shape of modern humans.[57] 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.[58]

Steven Mithen (2006) proposes that the Neanderthals had an elaborate proto-linguistic system of communication which was more musical than modern human language, and which predated the separation of language and music into two separate modes of cognition.[59]

Tools

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

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 stone-flakes, task-specific hand axes, and spears. Many of these tools were very sharp. There is also good evidence that they used a lot of wood, objects which are unlikely to have been preserved until today.[60]

There is some evidence for interpersonal violence among Neandertals. A 36,000-year-old Neadertal skull found near St. Césaire has a healed fracture in its cranial vault that was most likely caused by the impact of a sharp implement. The location of the wound suggests interpersonal violence rather than an accident. Because the wound healed, we know that the individual survived the attack.[61]

Also, while they had weapons, whether they had implements which were used as projectile weapons is controversial. They had spears, made of long wooden shafts with spearheads firmly attached, but they are thought by some to have been thrusting spears.[62] Still, a Levallois point embedded in a vertebra shows an angle of impact suggesting that it entered by a "parabolic trajectory" suggesting that it was the tip of a projectile.[63] 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 Māori — modern Homo sapiens, who also rarely threw objects, but used spears and clubs instead.[64]

Although much has been made of the Neanderthals' burial of their dead, their burials were less elaborate than those of anatomically modern humans. The interpretation of the Shanidar IV burials as including flowers, and therefore being a form of ritual burial,[65] has been questioned.[66] On the other hand, five of the six flower pollens found with Shanidar IV are known to have had 'traditional' medical uses, even among relatively recent 'modern' populations. In some cases Neanderthal burials include grave goods, such as bison and aurochs bones, tools, and the pigment ochre.

Neanderthals 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.[24]

Particularly intriguing is a hollowed-out bear femur with holes which may have been deliberately bored into it, known as the Divje Babe flute. 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 musical instrument, while others believe it was created by accident through the chomping action of another bear; or possibly was in fact not the work of Neanderthals.

Pendants and other jewelry showing traces of ochre dye and of deliberate grooving have also been found[67] with later finds, particularly in France but whether or not they were created by Neanderthals or traded to them by Cro-Magnons is a matter of controversy.

Cannibalism or ritual defleshing?

Neanderthal Burial of Kebara Cave (Carmel Range, Israel). Thermoluminescence dates place Neanderthal levels at Kebara at ca. 60,000 BP. Skeleton of an adult man nicknamed Moshe (25–35 years old, height 1,70 m) found in 1983

Neanderthals hunted large animals, such as the mammoth. Stone-tipped wooden spears were used for hunting and stone knives and poleaxes were used for butchering the animals or as weapons. However, they are believed to have practiced cannibalism, or ritual defleshing. This hypothesis has been represented after researchers found marks on Neanderthal bones similar to the bones of a dead deer butchered 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.[68] 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.[69]
  • 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.[70]

The evidence indicating cannibalism would not distinguish Neanderthals from modern Homo sapiens. Ancient and existing Homo sapiens are known to have practiced cannibalism (e.g. the Korowai) and/or mortuary defleshing (e.g. the sky burial of Tibet).

Grooves in bones are hypothesized to be cuts by Neanderthal tools, not animal teeth. The chances of them being random, as some writers attributing them to animals have proposed, is debated.

Body paint

A 2009 report in the Proceedings of the National Academy of Sciences (PNAS) on two archaeological sites in the Murcia province of southern Spain, records the discovery of shells showing pigment residues and concludes that these were used by the Neanderthals as make-up containers. Sticks of the black pigment manganese have previously been discovered in Africa. These may have been used as body paint by Neanderthals.[71]

Genome

Previous investigations concentrated on mitochondrial DNA (mtDNA), which, owing to strictly matrilineal inheritance and subsequent vulnerability to genetic drift, is of limited value to disprove interbreeding of Neanderthals with Cro-Magnon people.

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. This genome is very likely to be roughly the size of the human genome, three-billion base pairs, and probably shares most of its genes. It is thought a comparison will expand understanding of Neanderthals as well as the evolution of humans and human brains.[72]

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 found at Vindija cave, 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[73] appearing in the journal Nature has calculated the species diverged about 516,000 years ago, whereas fossil records show a time of about 400,000 years ago. Scientists hope the DNA records will answer the question of whether there was interbreeding among the species.[74] A 2007 study pushes the point of divergence back to around 800,000 years ago.[75]

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% to nearly 99.9% identical.[76][77]

On 16 November 2006, Lawrence Berkeley National Laboratory issued a press release suggesting that Neanderthals and ancient humans probably did not interbreed.[78] Edward M. Rubin, director of the U.S. Department of Energy’s Lawrence Berkeley National Laboratory and the Joint Genome Institute (JGI), sequenced a fraction (0.00002) of genomic nuclear DNA (nDNA) from a 38,000-year-old Vindia Neanderthal femur bone. They calculated the common ancestor to be about 353,000 years ago, and a complete separation of the ancestors of the species about 188,000 years ago. Their results show the genomes of modern humans and Neanderthals are at least 99.5% identical, but despite this genetic similarity, and despite the two species having coexisted in the same geographic region for thousands of years, Rubin and his team did not find any evidence of any significant crossbreeding between the two. Rubin said, "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."[79]

In 2008 Richard E. Green et al. from Max Planck Institute for Evolutionary Anthropology published the full sequence of Neanderthal mitochondrial DNA (mtDNA) and suggested that "Neandertals had a long-term effective population size smaller than that of modern humans."[80] Writing in Nature about Green et al.'s findings, James Morgan asserted that the mtDNA sequence contained clues that Neanderthals lived in "small and isolated populations, and probably did not interbreed with their human neighbours."[81][82]

In the same publication, it was disclosed that the previous work at Max Plank Institute that according to Svante Pääbo "Contamination was indeed an issue," and eventually realized that 11% of their sample was modern human DNA.[18][19] Since then, more of the preparation work is done in clean areas and 4-base pair 'tags' are added to the DNA as soon as it is extracted so the Neanderthal DNA can be identified.

With 3 billion nucleotides sequenced, analysis of about 1/3rd shows that there is no sign of admixture between modern humans and Neanderthals, according to Pääbo. This concurs with the work of Noonan from two years earlier. The variant of Microcephalin common outside Africa, which was attributed to rapid brain growth in humans and suggested to be of Neanderthal origin, was not found in Neanderthals. Nor was the MAPT variant, a very old variant found primarily in Europeans.[18]

However, later research by the same Pääbo says there was in fact interbreeding. "Those of us who live outside Africa carry a little Neanderthal DNA in us," said Svante Paabo of the Max Planck Institute in Munich, Germany, who led the study. "The proportion of Neanderthal-inherited genetic material is about 1 to 4 percent. It is a small but very real proportion of ancestry in non-Africans today," says Dr. David Reich of Harvard Medical School in Boston, who worked on the study.

2010 research by Svante Paabo of the Max Planck Institute for Evolutionary Biology in Leipzig, Germany, compared the genome of the Neanderthals to five modern humans from China, France, Africa and Papua New Guinea. The conclusive result is that 1 percent to 4 percent of the genes of the Asians and Europeans came from Neanderthals, while Africans have no uniquely Neanderthal genes .

In May 2010 the first draft of the neandertal genome was published,[83] and among the genes shown to differ between present-day humans and Neanderthals were RPTN, SPAG17, CAN15, TTF1 and PCD16. Comparisons of genomes also revealed Neanderthals to be more closely related to non-Africans than to Africans. It would indicate a gene flow from Neanderthals to modern humans, i.e., interbreeding between the two populations.

Extinction

Possible hypotheses for the fate of Neanderthals include the following:

  1. Neanderthals were a separate species from modern humans, did not interbreed, and became extinct (due to climate change or interaction with humans) and were replaced by early modern humans traveling from Africa.[84] Competition from H. sapiens probably contributed to Neanderthal extinction.[85] Jared Diamond has suggested a scenario of violent conflict and displacement.[86]
  2. Neanderthals were a contemporary subspecies which incidentally bred with Homo sapiens and disappeared through absorption.
  3. Neanderthals never split from H. sap. and are the ancestors of some anatomically modern humans (see Multiregional origin of modern humans).

Since the 1990s there has been a consensus on the recent African origin of modern humans, based on evidence from mitochondrial DNA. This consensus precludes possibility three. However this consensus is likely to be challenged by evidence provided by Neanderthal Genome Project's draft sequencing which shows an admixture of up to four percent.[citation needed]

mtDNA-based simulation of modern human expansion in Europe starting 1600 generations ago. Neanderthal range in light grey.[87]

According to the Out of Africa theory, 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, such as modern-day Croatia, Iberia and the Crimean peninsula,[clarification needed] where they held on for thousands of years. The last traces of Mousterian culture (without human specimens) have been found in Gorham's Cave on the remote south-facing coast of Gibraltar, dated 30,000 to 24,500 years ago. Proponents of this model believe that modern humans and Neanderthals were separate species that were not inter-fertile. They cite the following evidence:

  • The Neanderthals and modern humans were contemporaneous species. The two species maintained distinct morphologies over hundreds of thousands of years. On a number of occasions the habitats of modern humans and the Neanderthals overlapped. However, despite this overlap, the respective morphologies remained distinct based on the available fossil record.[88]
  • For example, remains associated with modern human anatomy have been found at Qafzeh in Israel dating to 90,000 years ago. These remains predate Neanderthal remains such as those at Kebara Cave, also in Israel, by about 30,000 years. Since Neanderthals appear after modern humans, it is unlikely that these modern humans evolved from the Neanderthals.[89]
  • No incontrovertible fossils that demonstrate intermediate characteristics between modern humans and Neanderthals have been found.[88]
  • Studies using non-recombinant DNA point to a recent African origin of Europeans. Mitochondrial DNA studies of a Neanderthal specimen revealed modern humans and Neanderthals last shared a common ancestor circa 600 000 years ago.[90][91]
  • Currently all European mtDNA lineages trace back to African lineages. Haplogroup N (mtDNA), the ancestral haplogroup for all Europeans, is thought to have emerged in East Africa 60–80,000 years ago.
  • A study conducted in 2008 of 28,000-year-old Cro-Magnon remains found that the mtDNA haplogroup of the specimen was a common haplogroup in contemporary Europeans. The haplogroup differed substantially from known Neanderthal mtDNA sequences.[92]
  • A recent statistical simulation found either no or insignificant admixing between modern humans and Neanderthals.[93] Another mtDNA analysis showed no evidence for Neanderthal contributions to the gene pool of modern humans.[94] The authors of the study concede this does not exclude Neanderthal contributions of other genes. They nevertheless argue other genetic and morphological data also suggest little or no Neanderthal contribution.
  • The most recent patrilineal ancestor of all living humans (traced via Y-chromosome inheritance), Y-chromosomal Adam, is estimated to have lived in Africa 60,000 years ago.

Neanderthals, according to Jordan (2001), appear to have had psychological traits that worked well in their early history but finally placed them at a long-term disadvantage with regards to modern humans. Jordan is of the opinion that the Neanderthal mind was sufficiently different from that of Homo sapiens to have been "alien" in the sense of thinking differently from that of modern humans, despite the obvious fact that Neanderthals were highly intelligent, with a brain as large or larger than our own. This theory is supported by what Neanderthals possessed, and just as importantly, by what they lacked, in cultural attributes and manufactured artifacts. Essentially, the Neanderthals lost out because their behaviors and tools eventually became second-rate.

There once was a time when both human types shared essentially the same Mousterian tool kit and neither human type had a definite competitive advantage, as evidenced by the shifting Homo sapiens/Neanderthal borderland in the Middle East. But finally Homo sapiens started to attain behavioral or cultural adaptations that allowed "moderns" an advantage. There are early glimmers of this from Zaire where in the area of Katanda bone harpoon points have been found of fine workmanship, dating to perhaps 80,000 years ago. These featured backwards-pointing barbs and lateral grooves so they could be easily installed on a wooden shaft, used to harpoon local fish. These appear to have been made by modern humans and make for a more sophisticated spear than any that Neanderthals are known to have made. Jordan admits some of these innovations were "flash in the pan" local affairs that faded away for awhile, but there does not seem much question that Homo sapiens in Africa was taking steps toward better tools and a more complex social life, while the Neanderthal ways and technology remained the same. It is noted that fishing was never much of a Neanderthal accomplishment (they did eat fish on occasion) but is more a behavior of modern human types. There is an example of a barbed point made from bone, evidently made by a Neanderthal, but such finds are very rare. Per Jordan the Neanderthals made wooden and stone artifacts, but bone and ivory ones were not common, implying that the Neanderthal mind tended to be rather resistant to learning new methods or materials.

Neanderthal people mastered complex tasks such as the making of fire, shelters with post holes, and stone tools. In their later career Neanderthals appear to have sometimes buried their dead and their placement of Cave Bear bones in order shows some sort of reverence or perhaps religion toward this animal. Yet there were many Cro-Magnon tools and behaviors that the Neanderthals seem to have never developed: organized fishing, using fish hooks and fish nets; headgear or hats, shoes, sewn clothing, needle-and-thread, and long-distance trade. It is still debated whether Neanderthals had significant art or music.[26]

Other researchers think that the Neanderthals had little sexual division of labor, with Neanderthal women alongside the men hunting big game. Such a lifestyle was not as energy efficient as that of modern humans, whose hunter-gatherer lifestyle secured supplemental food of a much greater variety, including plant materials such as tubers or wild grains, fish, edible fungi, and small edible animals secured by women, young boys/girls and elderly men, while males in the prime of life could hunt big mammals. Since the Neanderthals were mostly carnivorous and targeting big mammals, a shortage of large mammals meant possible bouts of starvation or malnutrition, which affected Cro-Magnon people less. The Neanderthals appear to have stored food against lean times much less than Cro-Magnon people did. Neanderthals got food in a haphazard, catch-as-catch-can manner. In addition, the Cro-Magnon sites show a lot of animal remains of small creatures best hunted with traps and snares, such as squirrels and rabbits, whereas Neanderthal sites show few such fossils. In short, inferior methods shut Neanderthals out of many food sources that Cro-Magnons exploited.

Cro-Magnons could carry more people on the land than Neanderthals could, and one may infer that Cro-Magnons would have familial and tribal organization that Neanderthals could not match, if they had the latter at all.

Neanderthals appear to have never used boats or rafts, as evidenced by the lack of Neanderthal fossils from North Africa, yet in stark contrast Homo erectus, their more primitive ancestor, appears to have used rafts or some other sort of boat on occasion. Homo erectus, or some other hominid, used such craft to reach the island of Flores as evidenced by the discovery of Homo floresiensis in 2003. Flores and some other places Homo erectus reached have always been surrounded by very deep water, proving the use of watercraft of some sort.[26]

Since the Neanderthals evidently never used watercraft, but prior and/or arguably more primitive editions of humanity did, there is argument that Neanderthals represent a highly specialized side branch of the human tree, relying more on physiological adaptation than psychological adaptation in daily life than "moderns". Specialization has been seen before in other hominims, such as Paranthropus boisei which evidently was adapted to eat rough vegetation.

Additionally, Neanderthals evidently had little long-term planning when securing food. French caves show almost no salmon bones during Neanderthal occupancy but large numbers during Cro-Magnon occupancy. In contrast, Cro-Magnons planned for salmon runs months ahead of time, getting enough people together at just the right time and place to catch a lot of fish. Neanderthals appear to have had little to no social organization beyond the immediate family unit. Why Neanderthal psychology was different from the modern humans that they coexisted with for millennia is not known.[26]

Due to the paucity of symbolism that Neanderthal artifacts show, Neanderthal language probably did not deal much with a verbal future tense, again restricting Neanderthal exploitation of resources. Cro-Magnon people had a much better standard of living than the hardscrabble existence available to Neanderthals. With better language skills and bigger social groups, a better psychological repertoire, and better planning, Cro-Magnon people, living alongside the Neanderthals on the same land, outclassed them in terms of life span, population, available spare time (as shown by Cro-Magnon art), physical health and lower rate of injury, infant mortality, comfort, quality of life, and food procurement. The advantages held by Cro-Magnon people let them by this time to thrive in worse climatic conditions than their Neanderthal counterparts. As weather worsened about 30,000 years ago, Jordan notes it would have taken only one or two thousand years of inferior Neanderthal skills to cause them to go extinct, in light of better Cro-Magnon performance in all these areas.[26]

About 55,000 years ago, the weather began to fluctuate wildly from extreme cold conditions to mild cold and back in a matter of a few decades. Neanderthal bodies were well suited for survival in cold climate- their barrel chests and stocky limbs stored body heat better than the Cro-Magnons. However the rapid fluctuations of weather caused ecological changes that the Neanderthals could not adapt to. The weather changes were so rapid that within a lifetime the plants and animals that one had grown up would be replaced by completely different plants and animals. Neanderthal's ambush techniques would have failed as grasslands replaced trees. A large number of Neanderthals would have died during these fluctuations which maximized about 30,000 years ago.[95]

Studies on Neanderthal body structures have shown than they needed more energy to survive than the Cro-Magnon man. Their energy needs were up to 350 calories more per day compared to the Cro-Magnon man. When food became scarce this calorie for survival difference played a major role in Neanderthal extinction.[95]

Jordan states the Chatelperronian tool tradition suggests Neanderthals were making some attempts at advancement, as Chatelperronian tools are only associated with Neanderthal remains. It appears this tradition was connected to social contact with Cro-Magnons of some sort. There were some items of personal decoration found at these sites, but these are inferior to contemporary Cro-Magnon items of personal decoration and arguably were made more by imitation than by a spirit of original creativity. At the same time, Neanderthal stone tools were sometimes finished well enough to show some aesthetic sense.[26] As Jordan notes: "A natural sympathy for the underdog and the disadvantaged lends a sad poignancy to the fate of the Neanderthal folk, however it came about."[26]

Interbreeding hypotheses

One theory is that that Neanderthals and "moderns" could interbreed and produce fertile offspring, but that psychological/behavioural differences, as well as differences in language and appearance, made for little sexual attraction between these two human types; or if so, that any live offspring were sterile, like mules. If there were any fertile hybrid offspring, it is possible that the greater numbers of Homo sapiens simply drowned out the Neanderthal input or Neanderthal traits were later "weeded out" of the hybrid population by natural selection, by processes such as disease. It is noted that most of Neanderthal DNA and "modern" DNA would be identical. Therefore, DNA from Neanderthals may be present in some modern populations, but there would be no way of demonstrating the origin of such genes.

The validity of such an extensive period of cornered Neanderthal groups is recently questioned.[clarification needed] 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 Peştera cu Oase (south-west Romania), dated to 34–36 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.[96] As for now, the expansion of the first anatomically modern humans into Europe cannot be located by diagnostic and well-dated AMH fossils "west of the Iron Gates of the Danube" before 32 thousand years ago.[97]

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.[98] A new proposal resolves the issue by taking the Gravettians rather than the Aurignacians as the anatomically modern humans which contributed to the Eurasian genetic pool after 30 thousand years ago.[98] 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.[99]

Modern-human findings in Abrigo do Lagar Velho, Portugal of 24,500 years ago, allegedly featuring Neanderthal admixtures, have been published.[100] However the interpretation of the Portuguese specimen is disputed.[88]

Jordan in his work Neanderthal points out that without some interbreeding, certain features on some "modern" skulls of Eastern European Cro-Magnon heritage are hard to explain. In another study, researchers have recently found in Peştera Muierilor, Romania, remains of European humans from 30 thousand years ago 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.[101]

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.[97][102][103] Studies of fossils from the upper levels of the Sima de las Palomas, Murcia, Spain, dated to 40,000 years ago, establish the late persistence of Neanderthals in Iberia. This reinforces the conclusion that the Neanderthals were not merely swept away by advancing modern humans. In addition, the Palomas Neanderthals variably exhibit a series of modern-human features rare or absent in earlier Neanderthals. Either they were evolving on their own towards the modern-human pattern, or more likely, they had contact with early modern humans around the Pyrenees. If the latter, it implies that the persistence of the Middle Paleolithic in Iberia was a matter of choice, and not cultural retardation.[104]

Specimens

The Ferrassie skull
  • 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. One of the nine remains was missing part of its right arm; theorized to have been broken off or amputated. The find is also significant because it shows that stone tools were present among this tribe's culture. One was 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 archaeological 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.
Type Specimen, Neanderthal 1.
  • Neanderthal 1: Initial Neanderthal specimen found during an archaeological 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.

Chronology

Bones with Neanderthal traits in chronological order.[clarification needed]

Mixed with H. heidelbergensis traits

  • > 350 ka: Sima de los Huesos c. 500:350 ka ago[105][106]
  • 350–200 ka: Pontnewydd 225 ka ago.
  • 200–135 ka: Atapuerca,[107] Vértesszöllos, Ehringsdorf, Casal de'Pazzi, Biache, La Chaise, Montmaurin, Prince, Lazaret, Fontéchevade

Typical H. neanderthalensis traits

Popular culture

Statues at the Homo neanderthalensis finding site in Krapina
Neanderthal statue in Veringenstadt

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. Although they are frequently characterized in this manner, research showing Neanderthals were as intelligent as contemporaneous Homo sapiens, with early stone tool technologies of comparable efficiency, is debunking long-held beliefs.[113]

Counterbalancing this are sympathetic literary portrayals of Neanderthals, as in the novel The Inheritors by William Golding, Isaac Asimov's The Ugly Little Boy, and Jean M. Auel's Earth's Children series, 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 together form essentially one story.

In the Thursday Next series of novels by Jasper Fforde, a small population of Neanderthals were re-created in modern Britain by advanced cloning techniques in the later years of the twentieth century. These fictional Neanderthals have equivalent intelligence to normal humans, but have a radically different culture in which aggression and competition are virtually unthinkable.

See also

Lists:

Footnotes

  1. ^ a b The pronunciation /niːˈændərθɔːl/ is very common in the United States and is often listed first in US dictionaries, for example American Heritage Dictionary and Random House Dictionaries. The UK pronunciation is /niːˈændərtɑːl/, as shown in Cambridge Advanced Learner's Dictionary), and Oxford Advanced Learner's Dictionary.
  2. ^ Neandertal is a widespread alternative spelling in English (see also US dictionaries in previous footnote) becoming so common that it is sometimes now listed first in dictionaries, for example MSN Encarta. Archived 2009-11-01.
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References

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The Unknown

We don’t know everything about our early ancestors. But scientists are constantly in the field and the laboratory, excavating new areas and conducing analyses with groundbreaking technology, continually filling in some of the gaps about our understanding of human evolution. Below are some of the still unanswered questions about H. neanderthalensis that may be better answered with future discoveries:

1. Will more studies of Neanderthal DNA help us identify what is unique about the modern human genome compared with our closest extinct relatives, the Neanderthals?

2. Is there a close correlation between climate change and the extinction of the Neanderthals, or was competition with modern humans the most important factor?

3. What was the relative contribution of animal and plant sources to the average Neanderthal's diet?

4. Were Neanderthals routinely symbolic (e.g. making ornamental or decorative objects, burying the dead), or did this just occur in specific populations? If the latter is the case, why did those populations exhibit these behaviors?

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