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]}

Neanderthal fossils have to date not been found in Africa, but rather close to Africa, found in the area of modern Israel. At some sites in the Holy Land region, 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.^[18]

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

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 and Italy in the south and from England and Portugal in the west to

A 2007 study suggested some Neanderthals may have had red hair and pale skin color.^[23][24]

Distinguishing physical traits

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. Nothing is certain (from unearthed bones) about the shape of soft parts such as eyes, ears, and lips of Neanderthals.^[25]

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.

• Cranial
  • Suprainiac fossa, a groove above the
    inion
  • Occipital bun, a protuberance of the occipital bone which looks like a hair knot ^[26]
  • Projecting mid-face
  • Low, flat, elongated skull
  • A flat basicranium^[27][28][29]
  • Supraorbital torus
    , a prominent, trabecular (spongy) brow ridge
  • 1,200–1,900 cm³ (73–116 cu in) skull capacity
  • Lack of a protruding chin (mental protuberance; although later specimens possess a slight protuberance)
  • Crest on the
    mastoid process
    behind the ear opening
  • No groove on
    canine teeth
  • A retromolar space posterior to the third molar
  • Bony projections on the sides of the nasal opening, projecting nose
  • Distinctive shape of the bony
    labyrinth
    in the ear
  • Larger mental foramen in mandible for facial blood supply
• Sub-cranial
  • Considerably more robust, stronger build
  • Long
    collar bones, wider shoulders
  • Barrel-shaped
    rib cage
  • Short, bowed shoulder blades
  • Larger round finger tips
  • Large
    kneecaps
  • Thick, bowed shaft of the
    thigh bones, bowed femur
  • Short shinbones and calf bones, shorter torus proportionally longer legs
  • Long, gracile pelvic
    superior pubic ramus
    )

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 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.^[30]

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.^[31]

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.^[32] 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.^[33] 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.^[34]

Although much has been made of the Neanderthals'

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.^[37]

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^[38] 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.
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.

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.

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

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^[40] 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.^[41] A 2007 study pushes the point of divergence back to around 800,000 years ago.^[42]

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.^[43][44]

On 16 November 2006,

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.[50]^[51] 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

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.[52] Competition from H. sapiens probably contributed to Neanderthal extinction.^[53] Jared Diamond has suggested a scenario of violent conflict and displacement.^[54]
2. Neanderthals were a contemporary subspecies which incidentally bred with
   Homo sapiens
   and disappeared through absorption.
3. Neanderthals never split from Homo sapiens 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.

Extinction

According to the

• 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.[55]
• 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.^[56]
• No incontrovertible fossils that demonstrate intermediate characteristics between modern humans and Neanderthals have been found.[55]
• 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.^[57][58]
• 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.^[59]
• A recent statistical simulation found either no or insignificant admixing between modern humans and Neanderthals.
  mtDNA analysis showed no evidence for Neanderthal contributions to the gene pool of modern humans.^[61]
  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 a while, 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

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.^[18]

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.^[18]

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.^[18]

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.^[18] 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."^[18]

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, prevented sexual attraction between these two human types; or that Neanderthals were different enough that there was little to no sexual contact at all, or if so, that there were no fertile offspring with any live offspring being sterile, like

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 Pestera 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.^[62] 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.^[63]

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.^[64] 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.^[64] 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.^[65]