Ancient North Eurasian

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. 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). The Ancient North Eurasians are deeply related to Paleolithic and Mesolithic European hunter-gatherers, but also derive a significant amount of their ancestry from a deep East Eurasian source, which they received in Siberia.

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.

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. 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.

Definition
The ANE lineage, also known as Paleolithic Siberians, 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.

The Ancient North Eurasians represent a Paleolithic Siberian cluster, more closely related to European hunter-gatherers than to East and Southeast Asian populations. It is suggested that the ANE ancestry found among modern human populations was largely contributed from a population linked to Afontova Gora (AG2/3), rather than Malta (MA1) or Yana.

Ancient North Eurasian associated Y-chromosome haplogroups are P-M45, and its subclades R and Q. Haplogroup P is inferred to have originated around 46,000 years ago, before or during the break up of Ancient West and East Eurasian lineages.

Formation
The formation of the Ancient North Eurasian/Siberian (ANE/ANS) gene pool likely occurred very early by the Upper Paleolithic dispersal of an 'Ancient West Eurasian' population via the 'northern route' through Central Asia into Siberia. There, they received geneflow from a population associated with the 'southern route' ('Ancient East Eurasians'). The West Eurasian source was distantly related to the Upper Paleolithic remains in Europe, such as the Goyet specimen, as well as the Kostenki-14 and Sungir individuals, and ultimately expanded from a population hub in the Iranian Plateau. The 'East Eurasian' source can be associated with ancestry found in the 40,000 year old Tianyuan man of Northern China, suggesting contact between these ancestral lineages in Northeastern Siberia.

Overall, Ancient North Eurasians are best described to have originated from an Ancient West Eurasian population, with approximately 22% geneflow from an East Eurasian source. Grebenyuk et al. argues that 'Ancient North Eurasians' were "Early Upper Paleolithic tribes of hunters" and linked to similar groups associated with Southern Siberian sites. These communities of Southern Siberian and Central Asian hunters belonged to one of the earliest migration waves of 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".

Studies on Ancient North Eurasian remains
The ANE/ANS-associated samples from the Yana Rhinoceros Horn Site (31,600 BP) in Northeastern Siberia and the closely related remains from the Mal'ta–Buret' culture (MA1), and the individuals associated with the Afontova Gora site (AG2/3), can be modeled to derive their ancestry from an Ancient West Eurasian lineage, with variable amounts of admixture from an East Eurasian lineage.

Lipson and Reich (2017) modeled the Mal'ta sample to be derived from a West Eurasian source (82%), with additional admixture from a lineage related to East Asians (18%), while also noting the possibility for a reversed geneflow from Mal'ta into East Asians, which however had less support with the available data. Posth et al. 2018 modeled the Mal'ta sample with around 84% West Eurasian and 16% East Eurasian ancestry. Yang et al. 2020 modeled the Yana specimens as a merger of a sister lineage of the 'European hunter-gatherer' Kostenki-14, contributing around 68% ancestry, and of a lineage contemporary to the 'Basal-East Asian' Tianyuan man, contributing around 32% ancestry, while finding no evidence for a reversed geneflow from ANE/ANS into Tianyuan or modern East Asians. Mao et al. 2021 models both Yana and Afontova Gora remains with around 73% West Eurasian and 27% East Eurasian ancestry. Sikora et al. 2019 analyzed the genetic remains of the Yana Rhinoceros Horn Site and found them to be closely related to the Ancient North Eurasians. The collectively named both populations as Ancient North Siberian. They modeled the ANE/ANS to derived around 78% West Eurasian ancestry and 22% East Eurasian ancestry. Sikora et al. also 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%). 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. By using a newly developed version of ADMIXTOOLS, they estimate around 76% West Eurasian ancestry and 24% East Eurasian ancestry for both the Yana and Mal'ta remains. Zhang et al. 2023 summarized that the Ancient North Siberians (Yana remains) are best described to derive 71% ancestry from a West Eurasian lineage and 29% ancestry from an East Eurasian lineage. The Yana remains are closely related to the Mal'ta and Afontova Gora remains, but not identical with them.

A different but geographically close specimen, known as the Salkhit individual (c. 34,000 BP) from Northern Mongolia was found to display a complex relation to the Yana individuals. While the Yana individuals derived between 25–33% of their ancestry from a Tianyuan-like source, the Salkhit individual derived around 25% ancestry from the Yana lineage and 75% from the Tianyuan lineage, suggesting bi-directional geneflow between Ancient West and East Eurasian populations in Northeastern Siberia.

Distribution
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.

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. 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.

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 and CHG ancestry. 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. Estimates for ANE ancestry among first wave Native Americans show higher percentages, such as 41% (36-45%) for those belonging to the Andean region in South America. 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. 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.

Genomic studies also indicate that the ANE component was brought to Western Europe by people related to the Yamnaya culture, long after the Paleolithic. It is reported in modern-day Europeans (10%–20%). 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. 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."

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.

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.

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.

Native American contribution
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. Vallini et al. 2024 notes that the "position of Native Americans suggests a primarily East Asian ancestry, with a smaller contribution from palaeolithic West Eurasian populations".

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.

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. 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. 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.

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. 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%. 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.

Jomon people
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.

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.

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. 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).

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.

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. 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 Asian Baikal populations).

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). 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.

North Han Chinese populations
Several studies reveal minor West Eurasian-derived admixture among Shaanxi Han Chinese, especially those living in Guanzhong and Shaanbei (2–5%). Ancient North Eurasian admixture is more dominant among Shaanxi Han Chinese compared to other Han subgroups.

West Asian populations
A model has been presented by Vallini et al. 2024, suggesting that Ancient Iranians (Iranian hunter-gatherers) formed from a deep Ancient West Eurasian lineage (WEC2, around 72%), and from minor geneflow from Basal Eurasian (around 18%) and Ancient East Eurasian (around 10%) sources. 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.

European populations
Lazaridis et al. (2014) detected ANE ancestry among modern European populations in proportions up to 20%. In ancient European populations, the ANE genetic component is visible in tests of the Yamnaya people but not of Western or Central Europeans predating the Corded Ware culture: 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). 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.

Villalba-Mouco et al. 2023 confirmed the strong affinity between the Eastern European Hunter-Gatherers (EHG) to the Ancient North Eurasians, and also found a low affinity to the Tianyuan man, explained by them having received significant amounts of ANE ancestry.

Scandinavian Hunter-Gatherer (SHG) is represented by several individuals buried at Motala, Sweden ca. 6000 BC. They were descended from Western Hunter-Gatherers who initially settled Scandinavia from the south, and received later admixture from EHG who entered Scandinavia from the north through the coast of Norway.

Western Steppe Herders (WSH) is the name given to a distinct ancestral component that represents descent closely related to the Yamnaya culture of the Pontic–Caspian steppe. This ancestry is often referred to as Yamnaya ancestry or Steppe ancestry, and was formed from EHG and CHG (Caucasus hunter-gatherer) in about equal proportions.

Phenotype prediction
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, while cautioning that this analysis was based on an extremely low coverage of DNA that might not give an accurate prediction of pigmentation. 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.

Anthropologic research
Han Kangxin (1994) Described the mummies found in the cementery of Gumugou as posessing "clear western racial characteristics" aproximating those of a so called "proto-european type". Whereas he asserted that among the seven mummies in the suburbs of Loulan cementery six posessed "clear european characteristics" aproximating the Saka population of the southern pamirs within the 6th century B.C.E; while one of them posessed "mongoloid" traits. Kozintsev (2020, 2022) argues that the historical Southern Siberian Okunevo population, and other Paleo-Siberians, which derive high amounts of their ancestry from Ancient North Eurasians, as possessing a distinct craniometric phenotype, which he dubbed "Americanoid", which represents the variation of the first humans in Siberia and should not be associated solely with ancient Caucasoids. The Ancient North Eurasians themselves originated among Ancient West Eurasians, and represent a "Boreal" variation of early humans. Craniometric data on ANE-rich remains (such as from Botai), show them to cluster most closely with remains from the Pit-Comb Ware culture in Eastern Europe, and to take up a more "western" position. Zhang et al. (2021) proposed that the 'Western' like features of the earlier Tarim mummies could be attributed to their Ancient North Eurasian ancestry. 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, but displayed an affinity for two specimens from the Harappan site of the Indus Valley Civilisation.

Evolution of blond hair
Blond hair is associated with a single nucleotide polymorphism, the mutated allele rs12821256 of the KITLG gene. 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). 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. Mathieson, et al. (2018) thus argued that this allele originated in the Ancient North Eurasian population, before spreading to western Eurasia.

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. 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. The gene was also found among the Tarim mummies.

Comparative mythology
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.

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.

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.