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Neanderthals became extinct around 40,000 years ago. Hypotheses on the causes of the extinction include violence, transmission of diseases from modern humans which Neanderthals had no immunity to, competitive replacement, extinction by interbreeding with early modern human populations, natural catastrophes, climate change and inbreeding depression. It is likely that multiple factors caused the demise of an already low population.

Possible coexistence before extinction

In research published in Nature in 2014, an analysis of radiocarbon dates from forty Neanderthal sites from Spain to Russia found that the Neanderthals disappeared in Europe between 41,000 and 39,000 years ago with 95% probability. The study also found with the same probability that modern humans and Neanderthals overlapped in Europe for between 2,600 and 5,400 years. Modern humans reached Europe between 45,000 and 43,000 years ago. Improved radiocarbon dating published in 2015 indicates that Neanderthals disappeared around 40,000 years ago, which overturns older carbon dating which indicated that Neanderthals may have lived as recently as 24,000 years ago, including in refugia on the south coast of the Iberian peninsula such as Gorham's Cave. Zilhão et al. (2017) argue for pushing this date forward by some 3,000 years, to 37,000 years ago. Inter-stratification of Neanderthal and modern human remains has been suggested, but is disputed. Stone tools that have been proposed to be linked to Neanderthals have been found at Byzovya (ru:Бызовая) in the polar Urals, and dated to 31,000 to 34,000 years ago, but is also disputed. At Mandrin Cave the French palaeolontologist Ludovic Slimak [fr] and colleagues developed a new method of analysing soot from fires. They were able to distinguish between fires made by Neanderthals and modern humans based on the differing food residues in the soot as a result of their different diets. The researchers found that the last layer of soot from Neanderthal fires was a year or less before the first made by modern humans, and in Slimak's view this shows that the two species met and supports the hypothesis that the Neanderthals disappeared due to competitive replacement.

Possible causes of extinction

Violence

Kwang Hyun Ko discusses the possibility that Neanderthal extinction was either precipitated or hastened by violent conflict with Homo sapiens. Violence in early hunter-gatherer societies usually occurred as a result of resource competition following natural disasters. It is therefore plausible to suggest that violence, including primitive warfare, would have transpired between the two human species. The hypothesis that early humans violently replaced Neanderthals was first proposed by French paleontologist Marcellin Boule (the first person to publish an analysis of a Neanderthal) in 1912.

Parasites and pathogens

Infectious diseases carried by Homo sapiens may have passed to Neanderthals, who would have had poor protection to infections they had not previously been exposed to, leading to devastating consequences for Neanderthal populations. Homo sapiens were less vulnerable to Neanderthal diseases, partly because they had evolved to cope with the far higher disease load of the tropics and so were more able to cope with novel pathogens, and partly because the higher numbers of Homo sapiens meant that even devastating outbreaks would still have left enough survivors for a viable population. If viruses could easily jump between these two similar species, possibly because they lived near together, Homo sapiens might have infected Neanderthals and prevented the epidemic from burning out as Neanderthal numbers declined. The same process may also explain Homo sapiens' resilience to Neanderthal diseases and parasites. Novel human diseases likely moved from Africa into Eurasia. This purported "African advantage" remained until the agricultural revolution 10,000 years ago in Eurasia, after which domesticated animals surpassed other primates as the most prevalent source of new human infections, replacing the "African advantage" with a "Eurasian advantage". The catastrophic impact of Eurasian viruses on Native American populations in the historical past offers a sense of how modern humans may have affected hominin predecessor groups in Eurasia 40,000 years ago. Human and Neanderthal genomes and disease or parasite adaptations may give insight on this.

Infectious illness interactions may express the prolonged period of stagnation before the modification, as per disease ecology. Mathematical models have been used to make forecasts for future investigations, giving information about inter-species interactions during the shift between the Middle and Upper Paleolithic eras. This can be useful given the sparse material record from this time and the potential of DNA sequencing and dating technology. Such modeling, together with modern technology and prehistoric archaeological methodologies, may provide a fresh understanding of this time in human origins.

Competitive replacement

Species specific disadvantages

Slight competitive advantage on the part of modern humans may have accounted for Neanderthals' decline on a timescale of thousands of years.

Generally small and widely dispersed fossil sites suggest that Neanderthals lived in less numerous and socially more isolated groups than contemporary Homo sapiens. Tools such as Mousterian flint stone flakes and Levallois points are remarkably sophisticated from the outset, yet they have a slow rate of variability and general technological inertia is noticeable during the entire fossil period. Artifacts are of utilitarian nature, and symbolic behavioral traits are undocumented before the arrival of modern humans in Europe around 40,000 to 35,000 years ago.

The noticeable morphological differences in skull shape between the two human species also have cognitive implications. These include the Neanderthals' smaller parietal lobes and cerebellum, areas implicated in tool use, visuospatial integration, numeracy, creativity, and higher-order conceptualization. The differences, while slight, would have possibly been enough to affect natural selection and may underlie and explain the differences in social behaviors, technological innovation, and artistic output.

Jared Diamond, a supporter of competitive replacement, points out in his book The Third Chimpanzee that the replacement of Neanderthals by modern humans is comparable to patterns of behavior that occur whenever people with advanced technology clash with people with less developed technology.

Division of labour

In 2006, it was posited that Neanderthal Division of labour between the sexes was less developed than Middle paleolithic Homo sapiens. Both male and female Neanderthals participated in the single occupation of hunting big game, such as bison, deer, gazelles, and wild horses. This hypothesis proposes that the Neanderthal's relative lack of labour division resulted in less efficient extraction of resources from the environment as compared to Homo sapiens.

Anatomical differences and running ability

Researchers such as Karen L. Steudel of the University of Wisconsin have highlighted the relationship of Neanderthal anatomy (shorter and stockier than that of modern humans) and the ability to run and the requirement of energy (30% more).

Nevertheless, in the recent study, researchers Martin Hora and Vladimir Sladek of Charles University in Prague show that Neanderthal lower limb configuration, particularly the combination of robust knees, long heels, and short lower limbs, increased the effective mechanical advantage of the Neanderthal knee and ankle extensors, thus reducing the force needed and the energy spent for locomotion significantly. The walking cost of the Neanderthal male is now estimated to be 8–12% higher than that of anatomically modern males, whereas the walking cost of the Neanderthal female is considered to be virtually equal to that of anatomically modern females.

Other researchers, like Yoel Rak, from Tel-Aviv University in Israel, have noted that the fossil records show that Neanderthal pelvises in comparison to modern human pelvises would have made it much harder for Neanderthals to absorb shocks and to bounce off from one step to the next, giving modern humans another advantage over Neanderthals in running and walking ability. However, Rak also notes that all archaic humans had wide pelvises, indicating that this is the ancestral morphology and that modern humans underwent a shift towards narrower pelvises in the late Pleistocene.

Modern humans and alliance with dogs

Pat Shipman argues that the domestication of the dog gave modern humans an advantage when hunting. Evidence shows the oldest remains of domesticated dogs were found in Belgium (31,700 BP) and in Siberia (33,000 BP). A survey of early sites of modern humans and Neanderthals with faunal remains across Spain, Portugal and France provided an overview of what modern humans and Neanderthals ate. Rabbit became more frequent, while large mammals – mainly eaten by the Neanderthals – became increasingly rare. In 2013, DNA testing on the "Altai dog", a paleolithic dog's remains from the Razboinichya Cave (Altai Mountains), has linked this 33,000-year-old dog with the present lineage of Canis familiaris.

Interbreeding

At the time of the last Neanderthals, approximately 45 to 40 thousand years ago, genetic analysis suggests that there was a gene flow from Neanderthals to modern humans of around 10%, but almost no flow from modern humans to Neanderthals. This may be an artifact due to the small number of late Neanderthal genomes, or because hybrids were not viable in Neanderthal groups, or because fertile Neanderthals were being absorbed into modern human groups but not vice versa. If the effect was real over an extended period, it would have increased the size of the modern human gene pool and reduced that of the already sparse Neanderthals, contributing to reduce their numbers below a viable population and thus to their extinction.

Inbreeding

According to a study by Rios et al, kinship patterns among recovered Neanderthal remains suggests that there was inbreeding, such as pairings between half-siblings and/or uncle/aunt and niece/nephew. Researchers hypothesize that Neanderthals may have become isolated into small groups during harsh climatic conditions, which contributed to inbreeding behaviours. Due to the lack of genetic diversity, Neanderthal populations would have become more vulnerable to climatic changes, diseases, and other stressors, which may have contributed to their extinction. A similar model to the inbreeding hypothesis can be seen among endangered lowland gorillas. Their populations are so small that it has caused inbreeding, making them even more vulnerable to extinction.

Climate change

Neanderthals went through a demographic crisis in Western Europe that seems to coincide with climate change that resulted in a period of extreme cold in Western Europe. "The fact that Neanderthals in Western Europe were nearly extinct, but then recovered long before they came into contact with modern humans came as a complete surprise to us," said Love Dalén, associate professor at the Swedish Museum of Natural History in Stockholm. If so, this would indicate that Neanderthals may have been very sensitive to climate change.

The data reveal that sudden climatic change, although crucial locally, had a limited effect on the worldwide Neanderthal population. Interbreeding and assimilation, which were hypothesized as causes in the death of European Neanderthal populations, are successful only for low levels of food competition. Future research will examine models of interbreeding, and hybridization may be evaluated using genomic records from the last ice age (Fu et al., 2016).

Natural catastrophe

A number of researchers have argued that the Campanian Ignimbrite Eruption, a volcanic eruption near Naples, Italy, about 39,280 ± 110 years ago (older estimate ~37,000 years), erupting about 200 km (48 cu mi) of magma (500 km (120 cu mi) bulk volume) contributed to the extinction of Neanderthals. The argument has been developed by Golovanova et al. The hypothesis posits that although Neanderthals had encountered several Interglacials during 250,000 years in Europe, inability to adapt their hunting methods caused their extinction facing H. sapiens competition when Europe changed into a sparsely vegetated steppe and semi-desert during the last Ice Age. Studies of sediment layers at Mezmaiskaya Cave suggest a severe reduction of plant pollen. The damage to plant life would have led to a corresponding decline in plant-eating mammals hunted by the Neanderthals.

See also

• List of hominina (hominid) fossils
• Quaternary extinction event – Extinctions of large mammals in the Late PleistocenePages displaying short descriptions of redirect targets

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Further reading

• Slimak, Ludovic; et al. (2024). "Long genetic and social isolation in Neanderthals before their extinction". Cell Genomics. 4 (9). doi:10.1016/j.xgen.2024.100593.

• Neanderthals
• Upper Paleolithic
• Archaeology in Europe
• Pleistocene extinctions