In archaeogenetics, the term Ancient North Eurasian (ANE) refers to an ancestral component that represents the lineage of the people of the Mal'ta–Buret' culture () 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 (), which were dubbed as Ancient North Siberians (ANS), and which either are directly ancestral to the ANE, or both being closely related sister lineages.

The ANE/ANS lineages both derive their ancestry from an admixture event between Ancient West Eurasians (about 65%, best represented by Upper Paleolithic Europeans such as Kostenki-14, c. 38,000 BP) and Ancient East Eurasians (about 35%, best represented by the Tianyuan man, c. 39,000 BP) during the Upper Paleolithic period. The Pleistocene ANE gene pool is likely associated with so-called "Western features", as visible in their descendants, the Tarim Mummies.: "The Tarim mummies' so-called Western physical features are probably due to their connection to the Pleistocene ANE gene pool"

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 (APS) 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.

Later, ANE populations migrated westward into Europe and admixed with European Western hunter-gatherer (WHG)-related groups to form the Eastern hunter-gatherer (EHG) group, which later admixed with Caucasus hunter-gatherers to form the Western Steppe Herder group, which became widely dispersed across Eurasia during the Bronze Age.

ANE ancestry has spread throughout Eurasia and the Americas in various migrations since the Upper Paleolithic. Significant ANE ancestry can be found in Native Americans, as well as in 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 dog which guards the path to the afterlife.

Genetic studies

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 a collective name for both MA-1 and Yana remains.

Ancient North Eurasian-associated Y-chromosome haplogroups are P-M45, and its subclades R and Q. Haplogroup P is inferred to have originated around 44,000 years ago in South or Southeast Asia and is observed with its highest frequency among indigenous Southeast Asian groups, such as the Andamanese or the Jahai people. P is downstream to haplogroup K2b found among the Upper Paleolithic Tianyuan man in Northern China, with its sister clade haplogroup MS-P397 being most common among Oceanian populations. Their maternal haplogroup belonged to subclades of haplogroup U commonly found among West Eurasian populations.

Formation

The formation of the Ancient North Eurasian/Siberian (ANE/ANS) gene pool occurred during the Upper Paleolithic period: an early West Eurasian population related to Upper Paleolithic Europeans, such as Kostenki-14 and Sungir, migrated via a northern route into Siberia where they encountered and admixed with an Ancient East Eurasian population which arrived earlier during the Initial Upper Paleolithic period via a southern route. The East Eurasian source is most closely related to the ancestry found among the 40,000 year old Tianyuan man of Northern China and other Upper Paleolithic Siberian and East/Southeast Asian groups. In total, the ANE/ANS derive around 50–71% ancestry from an early West Eurasian-affiliated source, and 29–50% from an East Eurasian source. On average, the ANE/ANS are usually modeled as roughly 2/3 early West Eurasian and 1/3 early East Eurasian.

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), consisted with them being the result of a Paleolithic admixture event. In terms of genome-wide structure the Ancient North Eurasian lineage appears roughly intermediate between modern western Eurasians and Native Americans, but distant from modern East Asians.

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".

Distribution and expansion patterns

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. 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.

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.

ANE-like ancestry was 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. 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.

The ancient Tianyuan Man, and by extension modern East/Southeast Asian populations, lacked 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. 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 around 50% of their ancestry from a Tianyuan-like source, the Salkhit individual derived around 25% ancestry from the Yana lineage, with the remainder 75% being from the Tianyuan lineage.

Groups partially derived from the Ancient North Eurasians

Native American contribution

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 ancestry component among Native Americans (the remainder of their ancestry) was of East Asian-like origin, specifically from a lineage that diverged from East Asians c. 30,000 years ago.

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 led to the emergence of ancestral Native American 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 APS 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 APS, 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. Despite extensive admixture with East Asian-related groups, APS ancestry persists in Siberian groups such as Yeniseian and Uralic-speaking groups.

Jōmon people