The 1912 ‘discovery’ of Piltdown Man (a hoax), appearing much more similar to modern humans than Neanderthals, was used as evidence that multiple different and unrelated branches
of primitive humans existed, and supported Boule’s reconstruction of H. neanderthalensis as a far distant relative and an evolutionary dead-end.
Their easternmost presence is recorded at Denisova Cave, Siberia 85°E; the southeast Chinese Maba Man, a skull, shares several physical attributes with Neanderthals, although
these may be the result of convergent evolution rather than Neanderthals extending their range to the Pacific Ocean.
 It is unclear when the line of Neanderthals split from that of modern humans; studies have produced various intervals ranging from 315,000 to more than 800,000 years
 Neanderthal hunters depicted in the Gallo-Romeins Museum Tongeren The neck vertebrae of Neanderthals are longer and thicker than those of (most) modern humans, leading
to stability,[clarification needed] possibly due to different head shape and size.
 Body proportions are usually cited as being “hyperarctic” as adaptations to the cold, because they are similar to those of human populations which developed in cold
climates—the Neanderthal build is most similar to that of Inuit and Siberian Yupiks among modern humans—and shorter limbs result in higher retention of body heat.
Using the latter dates, the split had likely already occurred by the time hominins spread out across Europe, and unique Neanderthal features had begun evolving by 600–500
thousand years ago.
 A large part of the controversy stems from the vagueness of the term “species”, as it is generally used to distinguish two genetically isolated populations, but admixture
between modern humans and Neanderthals is known to have occurred.
 Map of Europe during the Würm glaciation 70–20 thousand years ago Population Like modern humans, Neanderthals probably descended from a very small population with
an effective population—the number of individuals who can bear or father children—of 3,000 to 12,000 approximately.
 Archaeological evidence suggests that there was a tenfold increase in the modern human population in Western Europe during the period of the Neanderthal/modern human
transition, and Neanderthals may have been at a demographic disadvantage due to a lower fertility rate, a higher infant mortality rate, or a combination of the two.
 Although many of the gene variants inherited from Neanderthals may have been detrimental and selected out, Neanderthal introgression appears to have affected the
modern human immune system, and is also implicated in several other biological functions and structures, but a large portion appears to be non-coding DNA.
The date of around 250,000 years ago cites “H. helmei” as being the last common ancestor (LCA), and the split is associated with the Levallois technique of making stone tools.
 In 2019, English anthropologist John Stewart and colleagues suggested Neanderthals instead were adapted for sprinting, because of evidence of Neanderthals preferring
more warmer wooded areas over the colder mammoth steppe, and DNA analysis indicating a higher proportion of fast-twitch muscle fibres in Neanderthals than in modern humans.
 However, it has also been estimated that the age pyramids for Neanderthals and contemporary modern humans were the same.
 Neanderthals featured a rather large jaw which was once cited as a response to a large bite force evidenced by heavy wearing of Neanderthal front teeth (the “anterior
dental loading” hypothesis), but similar wearing trends are seen in contemporary humans.
 The fossil record is much more complete from 130,000 years ago onwards, and specimens from this period make up the bulk of known Neanderthal skeletons.
In 1939, American anthropologist Carleton Coon reconstructed a Neanderthal in a modern business suit and hat to emphasise that they would be, more or less, indistinguishable
from modern humans had they survived into the present.
 The northernmost bound is generally accepted to have been 55°N, with unambiguous sites known between 50–53°N, although this is difficult to assess because glacial advances
destroy most human remains, and palaeoanthropologist Trine Kellberg Nielsen has argued that a lack of evidence of Southern Scandinavian occupation is (at least during the Eemian interglacial) due to the former explanation and a lack of research
in the area.
Likewise, Neanderthals and Denisovans share a more recent last common ancestor (LCA) than to modern humans, based on nuclear DNA (nDNA).
 Middle Palaeolithic artefacts have been found up to 60°N on the Russian plains, but these are more likely attributed to modern humans.
 Evolution It is largely thought that H. heidelbergensis was the last common ancestor of Neanderthals, Denisovans, and modern humans before populations became isolated
in Europe, Asia, and Africa, respectively.
 The taxonomic distinction between H. heidelbergensis and Neanderthals is mostly based on a fossil gap in Europe between 300 and 243 thousand years ago during marine
isotope stage 8.
 Following Charles Darwin’s On the Origin of Species, Fuhlrott and Schaaffhausen argued the bones represented an ancient modern human form; Schaaffhausen,
a social Darwinist, believed that humans linearly progressed from savage to civilised, and so concluded that Neanderthals were barbarous cave-dwellers.
 Demographics Range Neanderthal skull from Tabun Cave, Israel, at the Israel Museum Pre- and early Neanderthals, living before the Eemian interglacial (130,000
years ago), are poorly known and come mostly from Western European sites.
 However, similar anatomy could also have been caused by adapting to a similar environment rather than interbreeding.
 Neanderthals are known from numerous fossils, especially from after 130,000 years ago.
 DNA studies have yielded various results for the Neanderthal/human divergence time, such as 538–315, 553–321, 565–503, 654–475, 690–550, 765–550, 741–317, and 800–520
thousand years ago; and a dental analysis concluded before 800,000 years ago.
In 2012, British-American geneticist Graham Coop hypothesised that they instead found evidence of a different archaic human species interbreeding with modern humans, which
was disproven in 2013 by the sequencing of a high-quality Neanderthal genome preserved in a toe bone from Denisova Cave, Siberia.
 In 2021, Israeli anthropologist Israel Hershkovitz and colleagues suggested the 140 to 120 thousand years old Israeli Nesher Ramla remains, which feature a mix of Neanderthal
and more ancient H. erectus traits, represent one such source population which recolonised Europe following a glacial period.
 Further, a computer reconstruction of the Neanderthal nose and predicted soft tissue patterns shows some similarities to those of modern Arctic peoples, potentially
meaning the noses of both populations convergently evolved for breathing cold, dry air.
 Dental remains from the Italian Visogliano and Fontana Ranuccio sites indicate that Neanderthal dental features had evolved by around 450–430 thousand years ago
during the Middle Pleistocene.
Also, the sinuses reconstructed wide are not grossly large, being comparable in size to those of modern humans.
This involved either introgression coming from an unknown archaic human into Denisovans, or introgression from an earlier unidentified modern human
wave from Africa into Neanderthals.
Estimates of 600,000 years ago assume that “H. rhodesiensis” was the LCA, which split off into modern human lineage and a Neanderthal/H.
 However, 430,000-year old bones at Sima de los Huesos could represent early Neanderthals or a closely related group, and the 400,000-year old Aroeira
3 could represent a transitional phase.
 However this study was rejected by other researchers who concluded that eyeball size does not offer any evidence the cognitive abilities of Neanderthal or modern humans.
It is claimed by some that this feature would be normal for all Homo, even tropically-adapted Homo ergaster or erectus, with the condition of a narrower thorax in most modern
humans being a unique characteristic.
The Neanderthal skull is typically more elongated, but also wider, and less globular than that of most modern humans, and features much more of an occipital bun, or “chignon”,
a protrusion on the back of the skull, although it is within the range of variation for humans who have it.
Two-phase argues that a single major environmental event—such as the Saale glaciation—caused European H. heidelbergensis to increase rapidly in body size and robustness, as
well as undergoing a lengthening of the head (phase 1), which then led to other changes in skull anatomy (phase 2).
 Numerous dates for the Neanderthal/human split have been suggested.
 There are two main hypotheses regarding the evolution of Neanderthals following the Neanderthal/human split: two-phase and accretion.
For much of the early 20th century, European researchers depicted Neanderthals as primitive, unintelligent, and brutish.
The oldest potential Neanderthal bones date to 430,000 years ago, but the classification remains uncertain.
 Various studies, using mtDNA analysis, yield varying effective populations, such as about 1,000 to 5,000; 5,000 to 9,000 remaining constant; or 3,000
to 25,000 steadily increasing until 52,000 years ago before declining until extinction.
 A 2017 study claimed the presence of Homo at the 130,000 year old Californian Cerutti Mastodon site in North America, but this is largely considered implausible.
 Neanderthals and Denisovans are more closely related to each other than they are to modern humans, meaning the Neanderthal/Denisovan split occurred after their split
with modern humans.
However, there was approximately 1 error for every 200 letters (base pairs) based on the implausibly high mutation rate, probably due to the preservation of the sample.
 A number of examples of symbolic thought and Palaeolithic art have been inconclusively attributed to Neanderthals, namely possible ornaments made from bird claws
and feathers or shells, collections of unusual objects including crystals and fossils, engravings, music production (possibly indicated by the Divje Babe flute), and Spanish cave paintings contentiously dated to
before 65,000 years ago.
 Anatomy Build Comparisons of a modern eurasian male example (left) and a Neanderthal (right) skull reconstruction at the Cleveland Museum of Natural History Neanderthal
skull features Neanderthals had more robust and stockier builds than typical modern humans, wider and barrel-shaped rib cages; wider pelvises; and proportionally shorter forearms and forelegs.
However, if sinus size is not an important factor for breathing cold air, then the actual function would be unclear, so they may not be a good indicator of evolutionary pressures
to evolve such a nose.
This likely resulted from an interbreeding event subsequent to the Neanderthal/Denisovan split which introduced another mtDNA line.
However, Neanderthals maintained this very low population, proliferating weakly harmful genes due to the reduced effectivity of natural selection.
 Neanderthals were likely capable of speech, possibly articulate, although the complexity of their language is not known.
 800,000 years ago has H. antecessor as the LCA, but different variations of this model would push the date back to 1 million years ago.
 By the middle of the century, based on the exposure of Piltdown Man as a hoax as well as a reexamination of La Chapelle-aux-Saints 1 (who had osteoarthritis
which caused slouching in life) and new discoveries, the scientific community began to rework its understanding of Neanderthals.
Pre- and early Neanderthals, on the other hand, seem to have continuously occupied only France, Spain, and Italy, although some appear to have moved out of this “core-area”
to form temporary settlements eastward (although without leaving Europe).
 Before splitting, Neanderthal/Denisovans (or “Neandersovans”) migrating out of Africa into Europe apparently interbred with an unidentified “superarchaic” human species
who were already present there; these superarchaics were the descendants of a very early migration out of Africa around 1.9 mya.
 The total population of Neanderthals remained low, proliferating weakly harmful gene variants and precluding effective long-distance networks.
 Theories for their extinction include demographic factors such as small population size and inbreeding, competitive replacement, interbreeding and assimilation
with modern humans, climate change, disease, or a combination of these factors.
 The binomial name Homo neanderthalensis—extending the name “Neanderthal man” from the individual specimen to the entire species, and formally recognising it as distinct
from humans—was first proposed by Irish geologist William King in a paper read to the 33rd British Science Association in 1863.
Nonetheless, southwestern France has the highest density of sites for pre-, early, and classic Neanderthals.
 It is also possible that there was gene flow between Western Europe and Africa during the Middle Pleistocene, obscuring Neanderthal characteristics in such specimens,
namely from Ceprano, Italy, and Sićevo Gorge, Serbia.
He explained their body proportions and greater muscle mass as adaptations to sprinting as opposed to the endurance-oriented modern human physique, as persistence hunting
may only be effective in hot climates where the hunter can run prey to the point of heat exhaustion (hyperthermia).
Researchers often explain these features as adaptations to conserve heat in a cold climate, but they may also have been adaptations for sprinting in the warmer, forested landscape
that Neanderthals often inhabited.
 The type specimen, Neanderthal 1, was found in 1856 in the Neander Valley in present-day Germany.
 Reconstruction of an elderly Neanderthal man Brain The Neanderthal braincase averages 1,640 cc for males and 1,460 cc for females, which is significantly
larger than the averages for all groups of extant humans; for example, mo
[‘After being mined for limestone, the cave caved in and was lost by 1900. It was rediscovered in 1997 by archaeologists Ralf Schmitz and Jürgen Thissen.
2. ^ The German spelling Thal (“valley”) was current until 1901 but has been Tal since then.
(The German noun is cognate with English dale.) The German /t/ phoneme was frequently spelled th from the 15th to 19th centuries, but the spelling Tal became standardized in 1901 and the old spellings of the German names Neanderthal for the valley
and Neanderthaler for the species were both changed to the spellings without h.
3. ^ In Mettmann, “Neander Valley”, there is a local idiosyncrasy in use of the outdated spellings with th, such as with the Neanderthal Museum (but the name
is in English [German would require Neandertalermuseum]), the Neanderthal station (Bahnhof Neanderthal), and some other rare occasions meant for tourists. Beyond these, city convention is to use th when referring to the species.
4. ^ King made
a typo and said “theositic”.
5. ^ The bones were discovered by workers of Wilhelm Beckershoff and Friedrich Wilhelm Pieper. Initially, the workers threw the bones out as debris, but Beckershoff then told them to store the bones. Pieper asked Fuhlrott
to come up to the cave and investigate the bones, which Beckershoff and Pieper believed belonged to a cave bear.
6. ^ OAS1 and STAT2 both are associated with fighting viral inflections (interferons), and the listed toll-like receptors
(TLRs) allow cells to identify bacterial, fungal, or parasitic pathogens. African origin is also correlated with a stronger inflammatory response.
7. ^ Higher levels of Neanderthal-derived genes are associated with an occipital and parietal
bone shape reminiscent to that of Neanderthals, as well as modifications to the visual cortex and the intraparietal sulcus (associated with visual processing).
8. ^ 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). Modern humans 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. The chart is missing superarchaic (which diverged from erectus 1.9
mya) introgression into Neanderthal/Denisovan common ancestor.
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