Azərbaycanca
Deutsch
Русский
Türkçe
Українська
Neanderthals ( ;Homo neanderthalensis or H. sapiens neanderthalensis) are an extinct group of archaic humans (generally regarded as a distinct species, though some regard it as a subspecies of Homo sapiens) who lived in Eurasia until about 40,000 years ago. The type specimen, Neanderthal 1, was found in 1856 in the in present-day Germany.
| Neanderthal Temporal range: Middle to Late Pleistocene | |
|---|---|
 | |
| An approximate reconstruction of a Neanderthal skeleton. The central rib-cage (including the sternum) and parts of the pelvis are from modern humans. | |
| Scientific classification | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Mammalia |
| Order: | Primates |
| Suborder: | Haplorhini |
| Infraorder: | Simiiformes |
| Family: | Hominidae |
| Subfamily: | Homininae |
| Tribe: | Hominini |
| Genus: | Homo |
| Species: | †H. neanderthalensis |
| Binomial name | |
| †Homo neanderthalensis King, 1864 | |
 | |
| Known Neanderthal range in Europe (blue), (Southwest Asia) (orange), Uzbekistan (green), and the Altai Mountains (violet) | |
| Synonyms | |
| Homo
Palaeoanthropus Protanthropus
Acanthropus
| |
It is not clear when the line of Neanderthals split from that of modern humans; studies have produced various times ranging from 315,000 to more than 800,000 years ago. The date of divergence of Neanderthals from their ancestor (H. heidelbergensis) is also unclear. The oldest potential Neanderthal bones date to 430,000 years ago, but the classification remains uncertain. Neanderthals are known from numerous fossils, especially from after 130,000 years ago.
The reasons for Neanderthal extinction are disputed. Theories for their extinction include demographic factors such as small population size and inbreeding, competitive replacement, interbreeding and assimilation with modern humans, change of climate, disease, or a combination of these factors.
For much of the early 20th century, European researchers depicted Neanderthals as primitive, unintelligent and brutish. Although knowledge and perception of them has markedly changed since then in the scientific community, the image of the unevolved caveman archetype remains prevalent in popular culture. In truth, Neanderthal technology was quite sophisticated. It includes the Mousterian stone-tool industry as well as the abilities to create fire, build cave hearths (to cook food, keep warm, defend themselves from animals, placing it at the centre of their homes), make adhesive birch bark tar, craft at least simple clothes similar to blankets and ponchos, weave, go seafaring through the Mediterranean, make use of medicinal plants, treat severe injuries, store food, and use various cooking techniques such as roasting, boiling, and smoking. Neanderthals consumed a wide array of food, mainly hoofed mammals, but also (megafauna), plants, small mammals, birds, and aquatic and marine resources. Although they were probably apex predators, they still competed with cave lions, cave hyenas and other large predators. 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, 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. Some claims of religious beliefs have been made. Neanderthals were likely capable of speech, possibly articulate, although the complexity of their language is not known.
Compared with modern humans, Neanderthals had a more robust build and proportionally shorter limbs. 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. They had cold-specific adaptations, such as specialised body-fat storage and an enlarged nose to warm air (although the nose could have been caused by genetic drift). Average Neanderthal men stood around 165 cm (5 ft 5 in) and women 153 cm (5 ft 0 in) tall, similar to pre-industrial modern Europeans. The braincases of Neanderthal men and women averaged about 1,600 cm3 (98 cu in) and 1,300 cm3 (79 cu in), respectively, which is considerably larger than the modern human average (1,260 cm3 (77 cu in) and 1,130 cm3 (69 cu in), respectively). The Neanderthal skull was more elongated and the brain had smaller parietal lobes and cerebellum, but larger temporal, occipital and orbitofrontal regions.
The total population of Neanderthals remained low, proliferating weakly harmful gene variants and precluding effective long-distance networks. Despite this, there is evidence of regional cultures and regular communication between communities. They may have frequented caves and moved between them seasonally. Neanderthals lived in a high-stress environment with high trauma rates, and about 80% died before the age of 40.
The 2010 (Neanderthal genome project)'s draft report presented evidence for interbreeding between Neanderthals and modern humans. It possibly occurred 316,000 to 219,000 years ago, but more likely 100,000 years ago and again 65,000 years ago. Neanderthals also appear to have interbred with (Denisovans), a different group of archaic humans, in Siberia. Around 1–4% of genomes of Eurasians, Indigenous Australians, Melanesians, Native Americans and North Africans is of Neanderthal ancestry, while most inhabitants of sub-Saharan Africa have around 0.3% of Neanderthal genes, save possible traces from early sapiens-to-Neanderthal gene flow and/or more recent back-migration of Eurasians to Africa. In all, about 20% of distinctly Neanderthal gene variants survive in modern humans. 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.
Taxonomy
Etymology
Neanderthals are named after the Neander Valley in which the first identified specimen was found. The valley was spelled Neanderthal and the species was spelled Neanderthaler in German until the spelling reform of 1901. The spelling Neandertal for the species is occasionally seen in English, even in scientific publications, but the scientific name, H. neanderthalensis, is always spelled with th according to the principle of priority. The vernacular name of the species in German is always Neandertaler ("inhabitant of the Neander Valley"), whereas Neandertal always refers to the valley. The valley itself was named after the late 17th century German theologian and hymn writer Joachim Neander, who often visited the area. His name in turn means 'new man', being a learned Graecisation of the German surname Neumann.
Neanderthal can be pronounced using the /t/ (as in ) or the standard English pronunciation of th with the fricative /θ/ (as ).
Neanderthal 1, the type specimen, 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". 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. However, in 1864, he recommended that Neanderthals and modern humans be classified in different genera as he compared the Neanderthal braincase to that of a chimpanzee and argued that they were "incapable of moral and [theistic] conceptions".
Research history
The first Neanderthal remains—(Engis 2) (a skull)—were discovered in 1829 by Dutch/Belgian prehistorian (Philippe-Charles Schmerling) in the (Grottes d'Engis), Belgium. He concluded that these "poorly developed" human remains must have been buried at the same time and by the same causes as the co-existing remains of extinct animal species. In 1848, (Gibraltar 1) from (Forbes' Quarry) was presented to the Gibraltar Scientific Society by their Secretary Lieutenant Edmund Henry Réné Flint, but was thought to be a modern human skull. In 1856, local schoolteacher (Johann Carl Fuhlrott) recognised bones from (Kleine Feldhofer Grotte) in Neander Valley—Neanderthal 1 (the (holotype specimen))—as distinct from modern humans, and gave them to German anthropologist (Hermann Schaaffhausen) to study in 1857. It comprised the cranium, thigh bones, right arm, left humerus and ulna, left ilium (hip bone), part of the right shoulder blade, and pieces of the ribs. 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. Fuhlrott and Schaaffhausen met opposition namely from the prolific pathologist Rudolf Virchow who argued against defining new species based on only a single find. In 1872, Virchow erroneously interpreted Neanderthal characteristics as evidence of senility, disease and malformation instead of archaicness, which stalled Neanderthal research until the end of the century.
By the early 20th century, numerous other Neanderthal discoveries were made, establishing H. neanderthalensis as a legitimate species. The most influential specimen was (La Chapelle-aux-Saints 1) ("The Old Man") from (La Chapelle-aux-Saints), France. French palaeontologist (Marcellin Boule) authored several publications, among the first to establish palaeontology as a science, detailing the specimen, but reconstructed him as slouching, ape-like, and only remotely related to modern humans. 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). He fuelled the popular image of Neanderthals as barbarous, slouching, club-wielding primitives; this image was reproduced for several decades and popularised in science fiction works, such as the 1911 (The Quest for Fire) by (J.-H. Rosny aîné) and the 1927 The Grisly Folk by H. G. Wells in which they are depicted as monsters. In 1911, Scottish anthropologist Arthur Keith reconstructed La Chapelle-aux-Saints 1 as an immediate precursor to modern humans, sitting next to a fire, producing tools, wearing a necklace, and having a more humanlike posture, but this failed to garner much scientific rapport, and Keith later abandoned his thesis in 1915.
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. Ideas such as Neanderthal behaviour, intelligence and culture were being discussed, and a more humanlike image of them emerged. 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. William Golding's 1955 novel (The Inheritors) depicts Neanderthals as much more emotional and civilised. However, Boule's image continued to influence works until the 1960s. In modern-day, Neanderthal reconstructions are often very humanlike.
Hybridisation between Neanderthals and early modern humans had been suggested early on, such as by English anthropologist Thomas Huxley in 1890, Danish ethnographer (Hans Peder Steensby) in 1907, and Coon in 1962. In the early 2000s, supposed hybrid specimens were discovered: (Lagar Velho 1) and (Muierii 1). However, similar anatomy could also have been caused by adapting to a similar environment rather than interbreeding.(Neanderthal admixture) was found to be present in modern populations in 2010 with the mapping of the first Neanderthal genome sequence. This was based on three specimens in (Vindija Cave), Croatia, which contained almost 4% archaic DNA (allowing for near complete sequencing of the genome). 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. 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.
Classification
| |||||||||||||||||||||||||||||||||
| 2019 phylogeny based on comparison of ancient proteomes and genomes with those of modern species. |
Neanderthals are hominids in the genus Homo, humans, and generally classified as a distinct species, H. neanderthalensis, although sometimes as a subspecies of modern human as Homo sapiens neanderthalensis. This would necessitate the classification of modern humans as (H. sapiens sapiens).
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. However, the absence of Neanderthal-derived patrilineal Y-chromosome and matrilineal mitochondrial DNA (mtDNA) in modern humans, along with the underrepresentation of Neanderthal X chromosome DNA, could imply reduced fertility or frequent sterility of some hybrid crosses, representing a partial biological reproductive barrier between the groups, and therefore species distinction. In 2014 geneticist Svante Pääbo summarised the controversy, describing such "taxonomic wars" as unresolvable, "since there is no definition of species perfectly describing the case".
Neanderthals are thought to have been more closely related to Denisovans than to modern humans. Likewise, Neanderthals and Denisovans share a more recent last common ancestor (LCA) than to modern humans, based on nuclear DNA (nDNA). However, Neanderthals and modern humans share a more recent mitochondrial LCA (observable by studying mtDNA) and Y chromosome LCA. This likely resulted from an interbreeding event subsequent to the Neanderthal/Denisovan split. 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. The fact that the mtDNA of a ~430,000 years old early Neanderthal-line archaic human from (Sima de los Huesos) in Spain is more closely related to those of Denisovans that to other Neanderthals or modern humans has been cited as evidence in favour of the latter hypothesis.
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 243,000 years ago during marine isotope stage 8. "Neanderthals", by convention, are fossils which date to after this gap. DNA from archaic humans from the 430,000-year-old Sima de los Huesos site in Spain indicate that they are more closely related to Neanderthals than to Denisovans, indicating that the split between Neanderthals and Denisovans must predate this time. The 400,000-year-old (Aroeira 3) skull may also represent an early member of the Neanderthal line. It is possible that gene flow between Western Europe and Africa during the Middle Pleistocene, may have obscured Neanderthal characteristics in some Middle Pleistocene European hominin specimens, such those (from Ceprano), Italy, and (Sićevo Gorge), Serbia. 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. Dental remains from the Italian Visogliano and Fontana Ranuccio sites indicate that Neanderthal dental features had evolved by around 450–430,000 years ago during the Middle Pleistocene.
There are two main hypotheses regarding the evolution of Neanderthals following the Neanderthal/human split: two-phase and accretion. 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). However, Neanderthal anatomy may not have been driven entirely by adapting to cold weather. Accretion holds that Neanderthals slowly evolved over time from the ancestral H. heidelbergensis, divided into four stages: early-pre-Neanderthals ((MIS 12), (Elster glaciation)), pre-Neanderthals sensu lato (MIS (11)–9, Holstein interglacial), early Neanderthals (MIS 7–5, (Saale glaciation)–Eemian), and classic Neanderthals sensu stricto (MIS 4–3, Würm glaciation).
Numerous dates for the Neanderthal/human split have been suggested. 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. The date of about 400,000 years ago uses H. heidelbergensis as the LCA. Estimates of 600,000 years ago assume that "H. rhodesiensis" was the LCA, which split off into modern human lineage and a Neanderthal/H. heidelbergensis lineage. Eight hundred thousand years ago has (H. antecessor) as the LCA, but different variations of this model would push the date back to 1 million years ago. However, a 2020 analysis of H. antecessor enamel proteomes suggests that H. antecessor is related but not a direct ancestor. 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 800–520,000 years ago; and a dental analysis concluded before 800,000 years ago.
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. Assuming a mutation rate of 1 × 10−9 or 0.5 × 10−9 per base pair (bp) per year, the Neanderthal/Denisovan split occurred around either 236–190,000 or 473–381,000 years ago, respectively. Using 1.1 × 10−8 per generation with a new generation every 29 years, the time is 744,000 years ago. Using 5 × 10−10nucleotide sites per year, it is 616,000 years ago. 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 600–500,000 years ago. 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.
Demographics
Range
Pre- and early Neanderthals, living before the Eemian interglacial (130,000 years ago), are poorly known and come mostly from Western European sites. From 130,000 years ago onwards, the quality of the fossil record increases dramatically with classic Neanderthals, who are recorded from Western, Central, Eastern and Mediterranean Europe, as well as (Southwest), Central and Northern Asia up to the Altai Mountains in southern Siberia. 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). Nonetheless, southwestern France has the highest density of sites for pre-, early and classic Neanderthals. The Neanderthals were the first human species to permanently occupy Europe as the continent was only sporadically occupied by earlier humans.
The southernmost find was recorded at (Shuqba Cave), Levant; reports of Neanderthals from the North African Jebel Irhoud and (Haua Fteah) have been reidentified as H. sapiens. 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. 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. Middle Palaeolithic artefacts have been found up to 60°N on the Russian plains, but these are more likely attributed to modern humans. 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.
It is unknown how the rapidly fluctuating climate of the last glacial period (Dansgaard–Oeschger events) impacted Neanderthals, as warming periods would produce more favourable temperatures but encourage forest growth and deter megafauna, whereas frigid periods would produce the opposite. However, Neanderthals may have preferred a forested landscape. Stable environments with mild mean annual temperatures may have been the most suitable Neanderthal habitats. Populations may have peaked in cold but not extreme intervals, such as marine isotope stages 8 and 6 (respectively, 300,000 and 191,000 years ago during the Saale glaciation). It is possible their range expanded and contracted as the ice retreated and grew, respectively, to avoid permafrost areas, residing in certain refuge zones during glacial maxima. In 2021, Israeli anthropologist Israel Hershkovitz and colleagues suggested the 140- to 120,000-year-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.
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. However, Neanderthals maintained this very low population, proliferating weakly harmful genes due to the reduced effectivity of natural selection. 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. 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. Estimates giving a total population in the higher tens of thousands are contested. A consistently low population may be explained in the context of the "(Boserupian Trap)": a population's carrying capacity is limited by the amount of food it can obtain, which in turn is limited by its technology. Innovation increases with population, but if the population is too low, innovation will not occur very rapidly and the population will remain low. This is consistent with the apparent 150,000 year stagnation in Neanderthal lithic technology.
In a sample of 206 Neanderthals, based on the abundance of young and mature adults in comparison to other age demographics, about 80% of them above the age of 20 died before reaching 40. This high mortality rate was probably due to their high-stress environment. However, it has also been estimated that the (age pyramids) for Neanderthals and contemporary modern humans were the same. Infant mortality was estimated to have been very high for Neanderthals, about 43% in northern Eurasia.
