Approximate location of the Ancient North Eurasians
c. 24,000~16,000 BP.[3][4][5]
In
archaeogenetics, the term Ancient North Eurasian (ANE) is the name given to an ancestral component that represents the lineage of the people of the
Mal'ta–Buret' culture (
c. 24,000 BP) and populations closely related to them, such as the
Upper Paleolithic individuals from
Afontova Gora in Siberia.[6][7] Genetic studies also revealed that the ANE are closely related to the remains of the preceding
Yana Culture (
c. 32,000 BP), which were dubbed as 'Ancient North Siberians' (ANS), and which either are directly ancestral to the ANE, or both being closely related sister lineages, sharing a common origin from the merger of an 'Early West Eurasian' hunter-gatherer lineage (represented by
Kostenki-14,
c. 38,000 BP) and an '
Early East Eurasian' population (represented by the
Tianyuan man,
c. 40,000 BP).[8] The Ancient North Eurasians derive around 2/3 from an Early West Eurasian lineage and around 1/3 of their ancestry from an Early East Eurasian lineage.[a][b][9][10][11][12][13][14][15]
Around 20,000 to 25,000 years ago, a branch of Ancient North Eurasian people mixed with
Ancient East Asians, which led to the emergence of
Ancestral Native American,
Ancient Beringian and
Ancient Paleo-Siberian populations. It is unknown exactly where this population admixture took place, and two opposing theories have put forth different migratory scenarios that united the Ancient North Eurasians with ancient East Asian populations.[16]
ANE ancestry has spread throughout Eurasia and the Americas in various migrations since the
Upper Paleolithic, and more than half of the world's population today derives between 5 and 42% of their genomes from the Ancient North Eurasians.[17] Significant ANE ancestry can be found in
Native Americans, as well as in regions of northern
Europe,
South Asia,
Central Asia, and
Siberia. It has been suggested that their mythology may have featured narratives shared by both Indo-European and some Native American cultures, such as the existence of a metaphysical
world tree and a fable in which a dog guards the path to the afterlife.[18]
Genetic studies
Definition
The ANE lineage is defined by association with the "
Mal'ta boy" (MA-1), the remains of an individual who lived during the
Last Glacial Maximum, 24,000 years ago in central
Siberia, discovered in the 1920s. Together with the
Yana Rhinoceros Horn Site samples, and Afontova Gora individuals, they are collectively referred to as 'Ancient North Siberians', although 'Ancient North Eurasian' is also used as collective name for both MA-1 and Yana remains.[19][20][21]
The Ancient North Eurasians represent a distinct cluster of genetic diversity within the larger Eurasian gene pool, forming an early Siberian variation of modern humans, taking up an 'intermediate position' between West and East Eurasians.[24][25] It is suggested that the ANE ancestry found among modern human populations was largely contributed from a population linked to Afontova Gora (AG-3), rather than Malta (MA-1) or Yana.[26]
Formation
A qpGraph by Allentoft et al. 2024, showing the formation of Ancient North Siberians/Eurasians (ANS/ANE) and their subsequent contribution to Western Eurasian lineages.[14]
Formation of Ancient North Eurasians in the context of West and East Eurasian Core populations[21]
The formation of the Ancient North Eurasian/Siberian gene pool likely occurred very early by the combination of an '
Early West Eurasian' Upper Paleolithic European (UP) lineage, and an '
Early East Eurasian' Initial Upper Paleolithic (IUP) lineage, basal to contemporary East and Southeast Asian populations.[25][15] The formation of ANE ancestry may be linked to the 'northern route' from Western Eurasia, while the peopling of Eastern Asia happened via the 'southern route'.[27] The ANE/ANS-associated samples from the
Yana Rhinoceros Horn Site (31,600 BP) in Northeastern Siberia, the
Mal'ta–Buret' culture, and the individuals associated with the
Afontova Gora site, can be modeled to derive around 2/3 of their ancestry from Early West Eurasians (total estimated range between 50% and 78%), with varying but significant contributions from Early East Eurasians at around 1/3 of their ancestry (total estimated range between 22% and 50%). Their West Eurasian-related component can be associated with ancestry found among Paleolithic European hunter-gatherers, such as
Kostenki-14,
Sunghir, or the
Peștera Muierii woman,[28] while their East Eurasian-related component can be associated with ancestry found among a population related to the Paleolithic
Tianyuan man, who is basal to contemporary East/Southeast Asians.[29][30][31][32][33][34][c][d][35][13][14]
A different but geographically close specimen, known as the
Salkhit individual (c. 34,000 BP) from Northern Mongolia, was found to have approximately 83% Tianyuan-like ancestry and 17% Kostenki14-like ancestry, being the to date oldest found representative of contact between West and East Eurasians. The relationship between the Salkhit individual and Ancient North Eurasians is described as complex, with evidence for bidirectional geneflow between them.[36][25]
Lipson and Reich (2017) modeled the Malta sample to derive ancestry from a West Eurasian source, with additional admixture from a lineage related to East Asians (represented by
Amis), while also noting the possibility for a reversed geneflow from Malta into East Asians, which however had less support with the available data.[35] Yang et al. 2020a,b, corroborated that both the Malta and Yana specimens formed from the merger of a sister lineage of the 'European hunter-gatherer'
Kostenki-14, contributing around 60-68% ancestry, and from a lineage contemporary to the 'Basal-East Asian'
Tianyuan man, contributing between 32-40%, while finding no evidence for a reversed geneflow from ANE into Tianyuan or modern East Asians.[37][38] Other studies could reproduce significant geneflow between these two sources and increased affinity for the ANE to the Tianyuan man (or other Upper Paleolithic East/Southeast Asian specimens), and also confirmed higher affinity for
Eastern European Hunter-Gatherers (EHG) to the Tianyuan man, explained by them having received significant amounts of ANE ancestry.[29] The 'Basal East Asian' (Tianyuan-like) ancestry among EHGs (Sidelkino) has been estimated to be around 12,9–31%.[39][40]
Grebenyuk et al. summarized that the Ancient North Eurasians descended from the 'Ancient North Siberian' Yana population, which were "Early Upper Paleolithic tribes of hunters" and linked to similar groups associated with contemporaneous Southern Siberian sites. These communities of Southern Siberian and Central Asian hunters belonged to one of the earliest migration waves of the anatomically modern humans into Siberia. The authors summarized that "the initial peopling of Northeastern Asia by the anatomically modern humans could have happened both from West to East and from South to North".[41] Sikora et al. notes that the Ancient North Eurasians (Malta and Afontova Gora individuals) are unlikely to be direct descendants of the 'Ancient North Siberian' Yana population; rather, the study argues, both are sister lineages sharing a common ancestor. According to Sikora et al., the Malta sample may additionally also have received some 'early
Caucasus hunter-gatherer' geneflow (c. 11%).[20] This scenario is questioned by Maier et al. 2023, who state that this conclusion is contradicted by other published articles, and that the direction of gene flow as well as observed affinity between ANE and CHG populations cannot be demonstrated by analysis of admixture graphs, but need further investigation.[42]
Vallini et al. 2024 notes that the Ancient North Eurasian lineage (represented by the Mal'ta and Yana specimens) is taking up an intermediate position between 'Ancient West Eurasians' (represented by a
Kostenki-14-like lineage) and '
Ancient East Eurasians' (represented by a
Tianyuan-like lineage), and is the result of a Paleolithic admixture event between West and East Eurasians deriving around 50% ancestry from each lineage respectively.[21]
By c. 32kya, populations carrying ANE-related ancestry were probably widely distributed across northeast Eurasia. They may have expanded as far as Alaska and the Yukon, but were forced to abandon high latitude regions following the onset of harsher climatic conditions that came with the Last Glacial Maximum.[44]
Populations genetically similar to MA-1 and Afontova Gora were an important genetic contributor to
Native Americans,
Europeans,
Ancient Central Asians,
South Asians, and some East Asian groups, in order of significance.[45]
Lazaridis et al. (2016:10) note "a cline of ANE ancestry across the east-west extent of Eurasia". A 2016 study found that the global maximum of ANE ancestry occurs in modern-day
Kets,
Mansi,
Native Americans, and
Selkups.[6][45]
The ancient Bronze-age-steppe
Yamnaya and
Afanasevo cultures were found to have a significant ANE-like component at c. 25–50% via their EHG ancestry.[47][48] According to Moreno-Mayar et al. 2018 between 14% and 38% of Native American ancestry may originate from gene flow from the Mal'ta–Buret' (ANE) population. This difference is caused by the penetration of posterior "Neo-Siberian" migrations into the Americas, with the lowest percentages of ANE ancestry found in Inuit and Alaskan Natives, as these groups are the result of migrations into the Americas roughly 5,000 years ago.[49] Estimates for ANE ancestry among first wave Native Americans show higher percentages,[50] such as 42% for those belonging to the
Andean region in South America.[50] The other gene flow in Native Americans (the remainder of their ancestry) was of an East Asian-related origin, specifically diverged from other East Asians c. 30,000 years ago.[31] Gene sequencing of another south-central Siberian
people (Afontova Gora-2) dating to approximately 17,000 years ago, revealed similar autosomal genetic signatures to that of Mal'ta boy-1, suggesting that the region was continuously occupied by humans throughout the Last Glacial Maximum.[31]
Genomic studies also indicate that the ANE component was brought to Western Europe by people related to the
Yamnaya culture, long after the Paleolithic.[47][6] It is reported in modern-day Europeans (10%–20%).[47][6] Earlier ANE ancestry is found in European hunter-gatherer populations through Paleolithic interactions with
Eastern European Hunter-Gatherers, which resulted in populations such as Scandinavian Hunter-Gatherers. [51] Western Hunter-Gatherers of the
Villabruna cluster also carried the
Y-haplogroup R1b, derived from the Ancient North Eurasian
haplogroup R*, indicating "an early link between Europe and the western edge of the Steppe Belt of Eurasia."[52]
A deer tooth pendant impregnated with the genetic material of an ANE woman was found in the
Denisova Cave, and dated to circa 24,700 years before present. She is closely related to Mal'ta and Afontova Gora specimens, found further east.[46]
An early Neolithic Central Asian specimen (Tutkaul1) from
Tajikistan was found to be primarily derived from Ancient North Eurasians with some additional Neolithic Iranian-related inputs. The sample is closely related to Afontova Gora 3 (AG3) and Mal’ta 1, as well as to the West Siberian hunter-gatherers (Tyumen and Sosnoviy). While the sample also displays affinity for
Eastern hunter-gatherers (EHGs), AG3 was found to be closer to EHGs than Tutkaul1, who instead may be a good proxy for ANE-related ancestry among ancient populations from the Iran and the Turan region.[53]
The Ancient Tianyuan Man and modern East/Southeast Asian populations were found to lack Upper Paleolithic Western Eurasian or ANE-related admixture, suggesting "resistance of those groups to the incoming UP population movements", or alternatively a subsequent reexpansion from a genetically East Asian-like population reservoir.[25]
Groups partially derived from the Ancient North Eurasians
According to Jennifer Raff, the Ancient North Eurasian population mixed with a daughter population of ancient East Asians, who they encountered around 25,000 years ago, which lead to the emergence of
Native American ancestral populations. However, the exact location where the admixture took place is unknown, and the migratory movements that united the two populations are a matter of debate.[16]
One theory supposes that Ancient North Eurasians migrated south to
East Asia, or
Southern Siberia, where they would have encountered and mixed with ancient East Asians. Genetic evidence from
Lake Baikal in
Mongolia supports this area as the location where the admixture took place.[55]
However, a third theory, the "Beringian standstill hypothesis", suggests that East Asians instead migrated north to Northeastern Siberia, where they mixed with ANE, and later diverged in Beringia, where distinct Native American lineages formed. This theory is supported by
maternal and
nuclear DNA evidence.[56] According to Grebenyuk, after 20,000 BP, a branch of Ancient East Asians migrated to Northeastern Siberia, and mixed with descendants of the ANE, leading to the emergence of
Ancient Paleo-Siberian and
Native American populations in Extreme Northeastern Asia.[57][41] However, the Beringian standstill hypothesis is not supported by
paternal DNA evidence, which may reflect different population histories for paternal and maternal lineages in Native Americans, which is not uncommon and has been observed in other populations.[58]
The descendants of admixture between ANE and ancient East Asians include
Ancient Beringian/
Ancestral Native American, which are specific archaeogenetic lineages, based on the genome of an infant found at the Upward Sun River site (dubbed USR1), dated to 11,500 years ago.[59] The AB and the Ancestral Native American (ANA) lineage formed about 25,000 years ago, and subsequently diverged from each other, with the AB staying in the Beringian region, while the Ancestral Native Americans populated the Americas. The ANE genetic contribution to late-Paeolithic Ancestral Native Americans (USR1 specimen, dated to 11,500 BP in
Alaska, and
Clovis specimen, dated to 12,600 BP in Montana) is estimated at 36.8%.[60] There are also the
Ancient Paleo-Siberians, populations represented by the Late Upper Paeolithic Lake Baikal Ust'Kyakhta-3 (UKY) 14,050-13,770 BP. They carried 30% ANE ancestry and 70% East Asian ancestry.[61]
Jōmon people, the pre-
Neolithic population of Japan, mainly derived their ancestry from East Asian lineages, but also received geneflow from the ANE-related "Ancient North Siberians" (represented by samples from the
Yana Rhinoceros Horn Site) prior to the migration from the Asian mainland to the Japanese archipelago. Jōmon ancestry is still found among the inhabitants of present-day Japan: most markedly among the
Ainu people, who are considered the direct descendants of the Jōmon people, and to a small, but significant degree among the majority of the Japanese population.[62][63]
Siberian and Asian Holocene populations
Altai hunter-gatherer is the name given to Middle Holocene Siberian hunter-gatherers within the
Altai-Sayan region in Southern Siberia. They originated from the admixture of Paleo-Siberian and Ancient North Eurasian groups and show increased affinity towards Native Americans. Bronze Age groups from North and Inner Asia with significant ANE ancestry (e.g. Lake Baikal hunter-gatherers,
Okunevo pastoralists) can be successfully modeled with Altai hunter-gatherers as a proximal ANE-derived ancestry source.[64] The Okunevos and
Botai can be considered as direct descendents of the Ancient North Eurasians (ANE), specifically of the
Malta-Buret people.[65]
West Siberian Hunter-Gatherer (WSHG) is a specific archaeogenetic lineage that was first reported by Narasimhan et al. (2019). It can be modeled as 20% EHG, 73% ANE and 6%
Ancient Northeast Asian. Although only represented by three sampled hunter-gatherer individuals from
Tyumen Oblast in the Russian Forest Zone east of the
Urals dated ca. 5,000 BCE, high-levels of WSHG-like ancestry can be detected in various populations of Central Asia until the Bronze Age. The population of the
Botai culture, while probably not directly descended from WSHG, displays a high affinity with the WSHG lineage.[66] The European-Siberian cline defined by
Eastern hunter-gatherer-like ancestry streched from Central Europe to Siberia and was already established 10,000 years ago, including the West Siberian hunter-gatherers, all deriving their ancestry primarily from Paleolithic Siberians (ANE).[67]
Lake Baikal Holocene - Among the
Ancient Northeast Asians (ANA) of the Neolithic to Early Bronze Age period, Baikal Eneolithic (Baikal_EN) and Baikal Early Bronze Age (Baikal_EBA) derived 6.4% to 20.1% ancestry from ANE, while the rest of their ancestry was derived from ANA. Fofonovo_EN near by Lake Baikal were mixture of 12-17% ANE ancestry and 83-87% ANA ancestry.[69]
Tarim mummies
A 2021 genetic study on the
Tarim mummies found that they were primarily descended from a population represented by the Afontova Gora 3 specimen (AG3), genetically displaying "high affinity" with it.[68] The genetic profile of the Afontova Gora 3 individual represented about 72% of the ancestry of the Tarim mummies, while the remaining 28% of their ancestry was derived from a population represented by the
Baikal EBA (Early Bronze Age
Northeast AsianBaikal populations).[68]
The Tarim mummies are thus one of the rare
Holocene populations who derive most of their ancestry from the
Ancient North Eurasians (ANE, specifically the
Mal'ta and Afontova Gora populations), despite their distance in time (around 14,000 years).[68] Having survived in a type of "genetic bottleneck" in the Tarim basin where they preserved and perpetuated their ANE ancestry, the Tarim mummies, more than any other ancient populations, can be considered as "the best representatives" of the Ancient North Eurasians.[68]
West Asian populations
Mesolithic Iranian hunter-gatherers and Neolithic Iranian farmers as well as
Caucasus hunter-gatherers (CHG) are inferred to have derived significant amounts of their ancestry from Ancient North Eurasians. Allentoft et al. 2024 modeled the Neolithic Iranians to derive appopximately 52% ancestry from Ancient North Eurasians, with the remainder ancestry (48%) being derived from
Basal Eurasians. The CHG displayed additional geneflow from a Paleolithic Caucasus/Anatolian source (c. 20%) and additional ANE-like admixture (c. 10%).[70] An alternative model without the need of significant amounts of ANE ancestry has been presented by Vallini et al. 2024, suggesting that Ancient Iranians (Iranian hunter-gatherers) formed from a deep Ancient West Eurasian lineage (WEC2, at least 50%), and from varying degrees of
Ancient East Eurasian and
Basal Eurasian components. The Ancient West Eurasian component associated with Iranian hunter-gatherers is inferred to have diverged from the West Eurasian Core lineage (represented by
Kostenki-14; WEC), with the WEC2 component staying in the region of the
Iranian Plateau, while the proper WEC component expanded into Europe.[71]
The
Tarim Mummies have a strong genetic proximity with Ancient North Eurasians (represented by the MA-1 human specimen of the
Mal'ta-Buret' culture (
c. 24,000 BP),
Eastern Hunter Gatherers (EHGs) are the population with the highest genetic affinity in Europe and the main source for ANE-like ancestry among modern Europeans.
Modeling of modern European populatios, plotting them according to their proportions of ancestry from each of three inferred ancestral populations; Early European Farmers, Ancient Northern Euroasians and West European Hunter Gathers, as inferred from the model-based analysis.[72]
Lazaridis et al. (2014) detected ANE ancestry among modern European populations in proportions up to 20%.[73] In ancient European populations, the ANE genetic component is visible in tests of the
Yamnaya people[47] but not of Western or Central Europeans predating the
Corded Ware culture:[74] ANE ancestry was introduced in the European gene pool with the
Eastern Hunter-Gatherer (EHG) lineage which derived significant ancestry from the ANE, c. 70%, with the remaining ancestry from a group more closely related to, but distinct from,
Western Hunter-Gatherers (WHGs).[45][75][53] It is represented by multiple individuals, such as from
Yuzhny Oleny in
Karelia, one of Y-haplogroup R1a-M417, dated c. 8.4
kya, the other of Y-haplogroup J, dated c. 7.2 kya; and one individual from
Samara, of Y-haplogroup R1b-P297, dated c. 7.6 kya, as well as individuals from Sidelkino and Popovo. After the end of the Last Glacial Maximum, the
Western Hunter-Gatherers (WHG) and EHG lineages merged in Eastern Europe, accounting for early presence of ANE-derived ancestry in Mesolithic Europe. Evidence suggests that as Ancient North Eurasians migrated westward from Eastern Siberia, they absorbed Western Hunter-Gatherers and other West Eurasian populations as well.[76][53]
Genomic studies by Raghavan et al. (2014) and Fu et al. (2016) suggested that Mal'ta boy may have had brown eyes, and relatively dark hair and dark skin,[80][81] while cautioning that this analysis was based on an extremely low coverage of DNA that might not give an accurate prediction of pigmentation.[82] Mathieson, et al. (2018) could not determine if Mal'ta 1 boy had the derived allele associated with blond hair in ANE descendants, as they could obtain no coverage for this SNP.[83]
Anthropologic research
Kozintsev (2020) argues that the historical Southern Siberian
Okunevo population, which derives most of their ancestry from Ancient North Eurasians and their closest relatives, as possessing a distinct craniometric phenotype, which he dubbed "
Americanoid", which represents the variation of the first humans in Siberia. He further argues that "As the geography and chronology of the ANE component show, it is misleading to describe it as Western Eurasian and associate it solely with ancient Caucasoids. To all appearances, it emerged before the Caucasoid-Mongoloid split."[48]
Zhang et al. (2021) proposed that the 'Western' like features of the earlier
Tarim mummies could be attributed to their Ancient North Eurasian ancestry.[84] Previous craniometric analyses on the early Tarim mummies found that they formed their own cluster, and clustered with neither European-related Steppe pastoralists of the
Andronovo and
Afanasievo cultures, nor with inhabitants of the Western Asian
BMAC culture, nor with
East Asian populations further east.[85]
Evolution of blond hair
Blond hair is associated with a
single nucleotide polymorphism, the mutated allele rs12821256 of the KITLG gene.[86][87][88][89][90] The earliest known individual with this allele is a female south-central Siberian ANE individual from the
Afontova Gora 3 site, which is dated to
c. 17,000 before present (the earlier ANE
Mal'ta boy lacks the sequence coverage to make this determination).[91] The allele then appears later in ANE-derived
Eastern Hunter-Gatherer (EHG) populations at
Samara, Motala and Ukraine, circa 10,000 BP, and then in populations with
Steppe ancestry.[83] Mathieson, et al. (2018) thus argued that this allele originated in the Ancient North Eurasian population, before spreading to western Eurasia.[83]
Geneticist
David Reich said that the
KITLG gene for blond hair probably entered continental Europe in a population migration wave from the Eurasian steppe, by a population carrying substantial Ancient North Eurasian ancestry.[92] Hanel and Carlberg (2020) likewise report that populations derived Ancient North Eurasian ancestry, specifically the
Eastern Hunter-Gatherers and the
Yamnayas, were responsible for transmitting this gene to Europeans.[86] The gene was also found among the
Tarim mummies.[93]
Comparative mythology
Mal'ta–Buret' culture centrally perforated ivory plaques with abstract circles, and three snakes[1] According to archeologist Don Hitchcock "the snake is rare in northern hemisphere Paleolithic art, presumably because the cold conditions precluded a wide distribution of snakes. In addition, it can be seen that the snakes have very broad heads, as though they belong to the
Cobra group - yet Cobras are now known only in
southern asian localities." It has yet to be clarrified how the creators of these ivory plaques know snakes, or if there are other possible interpretations for these.[94]
Since the term 'Ancient North Eurasian' refers to a genetic bridge of connected mating networks, scholars of
comparative mythology have argued that they probably shared myths and beliefs that could be reconstructed via the comparison of stories attested within cultures that were not in contact for millennia and stretched from the
Pontic–Caspian steppe to the
American continent.[18]
The
mytheme of the dog guarding the
Otherworld possibly stems from an older Ancient North Eurasian belief, as suggested by similar motifs found in
Indo-European,
Native American and
Siberian mythology. In
Siouan,
Algonquian,
Iroquoian, and in Central and South American beliefs, a fierce guard dog was located in the
Milky Way, perceived as the path of souls in the afterlife, and getting past it was a test. The Siberian
Chukchi and
Tungus believed in a guardian-of-the-afterlife dog and a spirit dog that would absorb the dead man's soul and act as a guide in the afterlife. In Indo-European myths, the figure of the dog is embodied by
Cerberus,
Sarvarā, and
Garmr. In Zoroastrianism, two four-eyed dogs guard the bridge to the afterlife called
Chinvat Bridge.
Anthony and Brown note that it might be one of the oldest mythemes recoverable through
comparative mythology.[18]
A second canid-related series of beliefs, myths and rituals connected dogs with healing rather than death. For instance, Ancient Near Eastern and
Turkic-
Kipchaq myths are prone to associate dogs with healing and generally categorised dogs as impure. A similar myth-pattern is assumed for the Eneolithic site of
Botai in Kazakhstan, dated to 3500 BC, which might represent the dog as absorber of illness and guardian of the household against disease and evil. In Mesopotamia, the goddess
Nintinugga, associated with healing, was accompanied or symbolized by dogs. Similar absorbent-puppy healing and sacrifice rituals were practiced in Greece and Italy, among the
Hittites, again possibly influenced by Near Eastern traditions.[18]
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