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Ancient Northeast Asian

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Not to be confused with Ancient Northern East Asian (ANEA).
The Ancient Northeast Asians (ANA,   yellow area) are defined as a cluster of Neolithic populations from the Altai Mountains to the Pacific coast. They were bordered by Western Eurasian populations to the west, which combined BMAC, Afanasievo and Ancient North Eurasian (ANE) ancestry.[1]

In archaeogenetics, the term Ancient Northeast Asian (ANA),[2][3] also known as Amur ancestry,[4] is the name given to an ancestral component that represents the lineage of the hunter-gatherer people of the 7th-4th millennia before present, in far eastern Siberia, Mongolia and the Baikal regions. They are inferred to have diverged from Ancient East Asians about 24kya ago,[5] and are represented by several ancient human specimens found in archaeological excavations east of the Altai Mountains. They are a sub-group of the Ancient Northern East Asians (ANEA).[6]

Neolithic populations

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class=notpageimage|
Ancient Northeast Asians, with contemporary cultures c. -3000 BCE
Position of Ancient Northeast Asians ( ) in a principal component analysis (PCA) of non-African modern human genomes (grey), and other ancient populations (colors).[7]

The Prehistoric populations of Eastern Siberia are poorly understood, mainly due to the lack of archaeological specimens. So far, the oldest populations for which genomic data have been obtained are the Upper Paleolithic Ancient North Eurasians (c. 24,000 BP) from Central Siberia, and Upper-Paleolithic populations related to the "Basal-East Asian" Tianyuan man (c. 40,000 BP), specifically the Salkhit (c. 34,000 BP) and AR33K (c. 33,000 BP) samples from Mongolia and the Amur region, or Manchuria. There is then a large gap until the Neolithic period, where the specific ANA gene pool has been identified. Ancestry basal to the ANA gene pool, but significantly closer to them than to the Upper-Paleolithic Tianyuan-related gene pool or other East Asian lineages (such as Southern East Asians), has been found among a sample in the Amur region (AR19K; c. 19 000 BP), suggesting that Ancient Northeast Asians diverged from other East Asian populations sometimes between 19kya to 26kya.[8][9][10]

The first individual to be identified with the specific ANA gene pool came from the Russian Far East, near the Pacific coast, at the Devil’s Gate Cave ("DevilsCave_N", c. 5700 BCE).[11] More Neolithic individuals with the ANA/Amur-like gene pool have been identified in eastern Mongolia (SOU001, "EastMongolia_preBA" 4686–4495 cal. BCE), in central Mongolia (ERM003, "CentralMongolia_preBA" 3781–3639 cal. BCE).[12]

The closely related hunter-gatherers from the Baikal region and adjacent regions of Siberia are associated with the Early Neolithic eastern Baikal Fofonovo culture ("Fofonovo_EN"), and the western Baikal Kitoi culture ("Baikal_EN", 5200–4200 BCE or Shamanka_EN),[13][14] as well as in conjunction with Ancient Paleo-Siberians (APS), the Early Bronze Age Baikal populations associated with the Glazkovo culture ("Baikal_EBA", circa 2500 BCE or Shamanka_EBA) and Cisbaikal_LNBA.[15][12] They cluster broadly with other Ancient Northeast Asian (ANA) populations, but are differentiated from them via drift associated with an earlier inland expansion route, and a minor Ancient North Eurasians (ANE) component at c. 11% (5-20%).[16][17][18][19] The ANE-like component is best explained via Ancient Paleo-Siberian-rich groups.[20] They also display genetic affinities with the Yumin hunter-gatherers from Northeast China, as well as the Neolithic and Bronze Age groups in Yakutia (Yakutia_LNBA) and Krasnoyarsk (kra001) in the Altai-Sayan region. These populations are sometimes described as "Neo-Siberians" and can be differentiated from proper ANA/Amur populations represented by the Neolithic Devils Cave specimen, but share a common recent origin via their Ancient Northern East Asian ancestor. Neo-Siberians are inferred to have expanded prior to the expansion of Neolithic Amur ancestry.[21][22][23]

The Devils_Cave_N sample was found to display genetic continuity with a c. 14kya old sample (AR14K) from the Amur region, suggesting that the specific ANA gene pool formed as early as 14,000 BP.[9] Neolithic ANA remains have been found as far as the Altai Mountains, 1,500 km further to the west than previously understood.[24]

Later populations

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Ulaanzuukh and Slab Grave cultures

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The people of the Ulaanzuukh (1450–1150 BCE) and Slab Grave (1100–300 BCE) cultures were closely associated with the Ancient Northeast Asians (Amur ancestry) and can be modeled as direct descendants of them. They largely replaced the previous Neolithic and Early Bronze Age Baikal hunter-gatherers, although geneflow between them has been proposed, particularly between a Neolithic Eastern Mongolian population (East_Mongolia_preBA) having primarily Amur_N-like ancestry and local Baikal hunter-gatherers (Baikal_EBA).[25]

Altai MLBA and Khövsgöl LBA

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Khövsgöl LBA in the Middle-Late Bronze Age (1400 to 1100 BC) in Mongolia. Khövsgöl LBA is essentially composed of Baikal EBA ancestry   (itself essentially Ancient Northeast Asian, ANA  , with a small admixture from Ancient North Eurasian), and a relatively small admixture from a Sintashta-like source  .

Several successor groups of the Neolithic and Early Bronze Age Baikal hunter-gatherers with varying degrees of Western Steppe Herders/Sintashta-like admixture started to appear in the Altai region during the Late Bronze Age. These groups formed from the Neolithic and Early Bronze Age Baikal populations from the Eastern Steppe and subsequent admixture from Western Steppe Herder migrant groups. This includes the Khövsgöl LBA[26] herders from northern Mongolia and the Altai MLBA hunter-gatherers from the Altai region.[27][12]

The Khövsgöl LBA herders are descended from Early Bronze Age Baikal hunter-gatherers (Baikal EBA or Shamanka EBA, c. 93-96%) with small amounts of admixture from Western Steppe Herders (Sintashta, c. 4-7%). Genetic analyses revealed that while dairy pastoralism seems to have been adopted by them from the Western Steppe Herders, they were primarily of local Northern East Asian origin, implying cultural transmission. Modern day Tuvans and Nganasans, followed by Nanais, Yukaghirs, Evens, Itelmens, Ulchis, Koryaks, Nivkhs, and Chukchis, are among the people sharing the highest genetic affinities with the Late Bronze Age herders of Khövsgöl, but are not identical with them.[28][29][30][19]

The Altai MLBA gene pool further West can be associated with Eastern Scythians (Saka), who can be modeled as deriving significant amounts of ancestry (c. 40-55%) from the Baikal/Shamanka EBA groups, with the remainder being derived from Sintashta-like admixture (c. 45-60%) associated with early Indo-Iranians.[31][32]

  • Deer stones are often associated with Khövsgöl LBA burials.[33] Probably c.1400-1000 BCE.[34]
    Deer stones are often associated with Khövsgöl LBA burials.[33] Probably c.1400-1000 BCE.[34]
  • Detail of deer stone, with weapons
    Detail of deer stone, with weapons

Tarim mummies

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A genomic study published in 2021 found that the Tarim mummies (c. 2000 BCE) had high levels of Ancient North Eurasian ancestry (c. 72%), with a smaller admixture from an East Asian-like population (particularly the Baikal_EBA, at c. 28%), but no detectable Western Steppe Herder-related ancestry.[35][36]

Sakas, Xiongnus, Huns, Avars

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The Slab Grave culture was essentially Ancient Northeast Asian (ANA,  ), while the neighbouring Sakas combined in almost equal parts Western Eurasian (Sintashta,  ) with Ancient Northeast Asian (Baikal EBA,  ) ancestry, with a smaller Iranian contribution (BMAC,  ).[37]
Genomic evidence from human remains shows that the Avars were essentially derived from Ancient Northeast Asians (ANA).[38]
Nivkh people are closely related to Ancient Northeast Asians.

The Baikal EBA populations, also contributed to a large extent to the formation of the hybrid Eurasian Scytho-Siberian cultures, such as the Arzhan and Pazyryk (Eastern Saka) as well as the Tasmola (Central Saka) cultures of Central Asia from around 1,000 BCE, contributing about half of their genetic profile (40-55%), highlighting the increase in genetic diversity during the late Bronze Age and the following Iron Age.[37]

The hybrid Saka cultures in turn played an important role in the formation of the Xiongnu Empire (3rd century BCE-1st century CE), which combined specific Saka ancestries (particularly Chandman/Uyuk-related ones), with Neolithic Amur-derived Ulaanzuukh and Slab Grave ancestries, to which Sarmatian and Han ancestry was further added at a later stage.[37] High status Xiongnu individuals tended to have less genetic diversity, and their ancestry was essentially derived from the Eastern Eurasian Ulaanzuukh/Slab Grave culture, while low status individuals tended to be more diverse and having higher Saka-like ancestry.[39] A likely chanyu, a male ruler of the Empire identified by his prestigious tomb, was shown to have had similar ancestry as a high status female in the "western frontiers", deriving about 39.3% Slab Grave genetic ancestry, 51.9% Han ancestry, with the rest (8.8%) being Saka (Chandman) ancestry.[39]

Elite Hun burial genetic ancestry (350 CE, Budapest)

A later different Eastern influx is evident in three outlier samples of the Saka Tasmola culture (Tasmola Birlik) and one of the Pazyryk culture (Pazyryk Berel), which displayed c. 70-83% additional Amur-derived ancestry, suggesting them to be recent migrants from further East. The same additional Eastern ancestry is found among the later groups of Huns (Hun Berel 300CE, Hun elite 350CE), and the Karakaba remains (830CE) and may be associated with the westwards expansion of Xiongnu tribes. A Hun individual from an elite burial of the mid-4th century CE in Budapest, Hungary, was reconstructed as 60% Ancient Northeast Asian/Amur (ANA) and 40% Saka.[40]

The 7-8th century Avars in Europe, particularly as regards the Avar elite, were also confirmed to have essentially Ancient Northeast Asian ancestry (c. 90%), with some additions from other sources.[41]

Göktürks

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The Turkic princess Ashina (551–582 CE), whose remains were sequenced, was found to be genetically closely associated with Ancient Northeast Asians (with 97.7% Northeast Asian ancestry, 2.3% West Eurasian ancestry dating back to around 3000 years ago, and no Chinese ("Yellow River") admixture), which according to Yang et al supports a Northeast Asian origin of the Ashina tribe and the Göktürk Khanate.[42] These findings refute "the western Eurasian origin and multiple origin hypotheses" in favor of an East Asian origin for the Göktürks.[43] However, the authors also observed that the population of the "Türkic Empire" as a whole, particularly Central Steppe and Medieval Türks, had a high but variable degree of West Eurasian admixture, suggesting genetic sub-structure within the empire:[44][45] for example, the ancestry of early medieval Turks was derived from Ancient Northeast Asians for about 62,2% of their genome, while the remaining 37,8% was derived from West Eurasians (BMAC and Afanasievo), with the admixture occurring around the year 500 CE.[46][47] The ruling clan of the Turkic peoples, the Ashina tribe, was found to display close genetic affinities with the earlier Slab Grave and Ulaanzuukh culture remains.[48]

ANA ancestry today

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Genetically, ANA/Amur ancestry peaks among modern Tungusic, Mongolic and Nivkh-speaking populations of Northeast Asia.[12] ANA ancestry (represented by the Tungusic-speaking Ulchi people) overall forms the main ancestry of the early and contemporary speakers of Turkic, Mongolic and Tungusic languages, which supports their spread from Northeast Asia westwards, discernable in the Lake Baikal region since at least 6kya. An earlier wave of Northern East Asian ancestry into Siberia is associated with "Neo-Siberians" (represented by Uralic-speaking Nganasans), which may be associated with the expansion of Yukaghir and Uralic languages, and the partial displacement of Paleo-Siberians, starting around 11kya.[49]

See also

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References

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  1. ^ Jeong et al. 2020, Figure S4A.
  2. ^ Wang, Ke; Yu, He; Radzevičiūtė, Rita; Kiryushin, Yuriy F.; Tishkin, Alexey A.; Frolov, Yaroslav V.; Stepanova, Nadezhda F.; Kiryushin, Kirill Yu.; Kungurov, Artur L.; Shnaider, Svetlana V.; Tur, Svetlana S.; Tiunov, Mikhail P.; Zubova, Alisa V.; Pevzner, Maria; Karimov, Timur (6 February 2023). "Middle Holocene Siberian genomes reveal highly connected gene pools throughout North Asia". Current Biology. 33 (3): 423–433.e5. Bibcode:2023CBio...33E.423W. doi:10.1016/j.cub.2022.11.062. ISSN 0960-9822. PMID 36638796. S2CID 255750546. We find the presence of ancient Northeast Asian (ANA) ancestry—initially described in Neolithic groups from the Russian Far East ... first identified in the Russian Far East in Neolithic hunter-gatherers from Devil's Gate Cave (DevilsCave_N)
  3. ^ Jeong et al. 2020: "In this study, we analyzed six pre-Bronze Age individuals from three sites dating to the fifth and fourth millennia BCE: one from eastern Mongolia (SOU001, "eastMongolia_preBA", 4686-4495 cal. BCE), one from central Mongolia (ERM003, "centralMongolia_preBA", 3781-3643 cal. BCE), and four from the eastern Baikal region ("Fofonovo_EN"). By comparing these genomes to previously published ancient and modern data across Eurasia (Fig. 2) (see Methods and Materials), we found that they are most closely related to contemporaneous hunter-gatherers from the western Baikal region ("Baikal_EN", 5200-4200 BCE) and the Russian Far East ("DevilsCave_N", ca. 5700 BCE), filling in the geographic gap in the distribution of this genetic profile (Fig. 3a). We refer to this profile as "Ancient Northeast Asian" (ANA)"
  4. ^ Yang, Melinda A. (6 January 2022). "A genetic history of migration, diversification, and admixture in Asia". Human Population Genetics and Genomics. 2 (1): 1–32. doi:10.47248/hpgg2202010001. ISSN 2770-5005. Amur ancestry—ancestry associated with populations in the Amur River region, Mongolia, and Siberia, with the oldest individual sampled to date represented by a 14,000-year-old individual from the Amur River region, i.e. Amur14K [61]. Populations associated with this ancestry likely contributed to the ancestors of Native Americans and populations associated with Paleosiberian ancestry.
  5. ^ Yang, Melinda A. (6 January 2022). "A genetic history of migration, diversification, and admixture in Asia". Human Population Genetics and Genomics. 2 (1): 1–32. doi:10.47248/hpgg2202010001. ISSN 2770-5005. 24,000 years ago may also indicate the split time separating Amur ancestry from other Asian ancestries.
  6. ^ Yang, Melinda A. (6 January 2022). "A genetic history of migration, diversification, and admixture in Asia". Human Population Genetics and Genomics. 2 (1): 1–32. doi:10.47248/hpgg2202010001. ISSN 2770-5005. Using f4-statistics, both DevilsCave_N and AR14K share a close genetic relationship to each other and group phylogenetically with other ancient northern East Asian individuals rather than ancient southern East Asian individuals [61,68].
  7. ^ Zhang, Fan (November 2021). "The genomic origins of the Bronze Age Tarim Basin mummies". Nature. 599 (7884): 256–261. Bibcode:2021Natur.599..256Z. doi:10.1038/s41586-021-04052-7. ISSN 1476-4687. PMC 8580821. PMID 34707286.
  8. ^ Wang, Ke (2023). "Middle Holocene Siberian genomes reveal highly connected gene pools throughout North Asia". Current Biology. 33 (3): 423–433.e5. Bibcode:2023CBio...33E.423W. doi:10.1016/j.cub.2022.11.062. PMID 36638796. S2CID 255750546. These individuals carry the typical ancient North Eurasian (ANE) ancestry, a widely distributed gene pool that was first identified in a 24,000-year-old individual (MA1) from the Mal'ta site near the Lake Baikal in south central Siberia. Thereafter, there is a large gap of 12,000 years, during which the genomic profile of human populations in the Altai region is unknown. (...) We find the presence of ancient Northeast Asian (ANA) ancestry —initially described in Neolithic groups from the Russian Far East— in another Neolithic Altai-Sayan individual associated with different cultural features, revealing the spread of ANA ancestry 1,500 km further to the west than previously observed.
  9. ^ a b Mao, Xiaowei; Zhang, Hucai; Qiao, Shiyu; Liu, Yichen; Chang, Fengqin; Xie, Ping; Zhang, Ming; Wang, Tianyi; Li, Mian; Cao, Peng; Yang, Ruowei; Liu, Feng; Dai, Qingyan; Feng, Xiaotian; Ping, Wanjing (10 June 2021). "The deep population history of northern East Asia from the Late Pleistocene to the Holocene". Cell. 184 (12): 3256–3266.e13. doi:10.1016/j.cell.2021.04.040. ISSN 0092-8674. PMID 34048699. S2CID 235226413.
  10. ^ Gakuhari, Takashi; Nakagome, Shigeki; Rasmussen, Simon; Allentoft, Morten E.; Sato, Takehiro; Korneliussen, Thorfinn; Chuinneagáin, Blánaid Ní; Matsumae, Hiromi; Koganebuchi, Kae; Schmidt, Ryan; Mizushima, Souichiro; Kondo, Osamu; Shigehara, Nobuo; Yoneda, Minoru; Kimura, Ryosuke (25 August 2020). "Ancient Jomon genome sequence analysis sheds light on migration patterns of early East Asian populations". Communications Biology. 3 (1): 437. doi:10.1038/s42003-020-01162-2. ISSN 2399-3642. PMC 7447786. PMID 32843717. S2CID 221284845.
  11. ^ Wang, Ke (2023). "Middle Holocene Siberian genomes reveal highly connected gene pools throughout North Asia". Current Biology. 33 (3): 423–433.e5. Bibcode:2023CBio...33E.423W. doi:10.1016/j.cub.2022.11.062. PMID 36638796. S2CID 255750546. Ancient Northeast Asian (ANA) ancestry. The latter gene pool was first identified in the Russian Far East in Neolithic hunter-gatherers from Devil's Gate Cave (DevilsCave_N), and, to date, the westernmost presence of ANA ancestry was reported in the Lake Baikal region and in Central Mongolia by 7,500 and 5,600 BP, respectively.
  12. ^ a b c d Jeong, Choongwon (2020). "A Dynamic 6,000-Year Genetic History of Eurasia's Eastern Steppe". Cell. 183 (4): 890–904. doi:10.1016/j.cell.2020.10.015. ISSN 0092-8674. PMC 7664836. PMID 33157037. S2CID 214725595. ...the western Baikal early Neolithic Kitoi culture (Baikal_EN) and the early Bronze Age Glazkovo culture (Baikal_EBA) (Damgaard et al., 2018a)...
  13. ^ Jeong et al. 2020, p. 891: "In this study, we analyzed six pre-Bronze Age hunter-gatherer individuals from three sites dating to the fifth and fourth millennia BCE: one from eastern Mongolia (SOU001, eastMongolia_preBA, 4686–4495 cal. BCE), one from central Mongolia (ERM003, centralMongolia_preBA, 3781–3639 cal. BCE), and four from the eastern Baikal region (Fofonovo_EN). By comparing these genomes to previously published ancient and modern data across Eurasia (Figure 2; Table S3C), we found that they are most closely related to contemporaneous huntergatherers from the western Baikal region (Baikal_EN, 5200–4200 BCE) and the Russian Far East (DevilsCave_N, ca. 5700 BCE), filling in the geographic gap in the distribution of this genetic profile (Figure 3A). We refer to this profile as Ancient Northeast Asian (ANA) to reflect its geographic distribution relative to another widespread mid-Holocene genetic profile known as Ancient North Eurasian (ANE), which is found among the Pleistocene hunter-gatherers of the Mal'ta (ca. 24500–24100 BP) and Afontova Gora (ca. 16900–16500 BP) sites in Siberia (Fu et al., 2016; Raghavan et al., 2015) and the horse-herders of Botai, Kazakhstan (ca. 3500–3300 BCE)"
  14. ^ Allen, Edward (2022). "Multidisciplinary lines of evidence reveal East/Northeast Asian origins of agriculturalist/pastoralist residents at a Han dynasty military outpost in ancient Xinjiang". Frontiers in Ecology and Evolution. 10. doi:10.3389/fevo.2022.932004. ISSN 2296-701X.
  15. ^ Gnecchi-Ruscone et al. 2021, Fig.2 for the date.
  16. ^ Zhang et al. 2021: "Using qpAdm, we modelled the Tarim Basin individuals as a mixture of two ancient autochthonous Asian genetic groups: the ANE, represented by an Upper Palaeolithic individual from the Afontova Gora site in the upper Yenisei River region of Siberia (AG3) (about 72%), and ancient Northeast Asians, represented by Baikal_EBA (about 28%)"
  17. ^ Wang, Ke (2023). "Middle Holocene Siberian genomes reveal highly connected gene pools throughout North Asia". Current Biology. 33 (3): 423–433.e5. Bibcode:2023CBio...33E.423W. doi:10.1016/j.cub.2022.11.062. PMID 36638796. S2CID 255750546.
  18. ^ Jeong et al. 2020.
  19. ^ a b Jeong, Choongwon; Wilkin, Shevan; Amgalantugs, Tsend; Bouwman, Abigail S.; Taylor, William Timothy Treal; Hagan, Richard W.; Bromage, Sabri; Tsolmon, Soninkhishig; Trachsel, Christian; Grossmann, Jonas; Littleton, Judith; Makarewicz, Cheryl A.; Krigbaum, John; Burri, Marta; Scott, Ashley (27 November 2018). "Bronze Age population dynamics and the rise of dairy pastoralism on the eastern Eurasian steppe". Proceedings of the National Academy of Sciences. 115 (48): E11248–E11255. Bibcode:2018PNAS..11511248J. doi:10.1073/pnas.1813608115. ISSN 0027-8424. PMC 6275519. PMID 30397125. Assuming that the early Neolithic populations of the Khövsgöl region resembled those of the nearby Baikal region, we conclude that the Khövsgöl main cluster obtained ∼11% of their ancestry from an ANE source during the Neolithic period and a much smaller contribution of WSH ancestry (4–7%) beginning in the early Bronze Age.
  20. ^ Wang, Ke; Yu, He; Radzevičiūtė, Rita; Kiryushin, Yuriy F.; Tishkin, Alexey A.; Frolov, Yaroslav V.; Stepanova, Nadezhda F.; Kiryushin, Kirill Yu.; Kungurov, Artur L.; Shnaider, Svetlana V.; Tur, Svetlana S.; Tiunov, Mikhail P.; Zubova, Alisa V.; Pevzner, Maria; Karimov, Timur (6 February 2023). "Middle Holocene Siberian genomes reveal highly connected gene pools throughout North Asia". Current Biology. 33 (3): 423–433.e5. Bibcode:2023CBio...33E.423W. doi:10.1016/j.cub.2022.11.062. ISSN 0960-9822. PMID 36638796.
  21. ^ Zeng, Tian Chen; et al. (2 October 2023). "Postglacial genomes from foragers across Northern Eurasia reveal prehistoric mobility associated with the spread of the Uralic and Yeniseian languages". BioRxiv. doi:10.1101/2023.10.01.560332. S2CID 263706090.
  22. ^ Childebayeva, Ainash; et al. (1 October 2023). "Bronze Age Northern Eurasian Genetics in the Context of Development of Metallurgy and Siberian Ancestry". BioRxiv. doi:10.1101/2023.10.01.560195. S2CID 263672903.
  23. ^ Sikora, Martin; Pitulko, Vladimir V.; Sousa, Vitor C.; Allentoft, Morten E.; Vinner, Lasse; Rasmussen, Simon; Margaryan, Ashot; de Barros Damgaard, Peter; de la Fuente, Constanza; Renaud, Gabriel; Yang, Melinda A.; Fu, Qiaomei; Dupanloup, Isabelle; Giampoudakis, Konstantinos; Nogués-Bravo, David (June 2019). "The population history of northeastern Siberia since the Pleistocene". Nature. 570 (7760): 182–188. Bibcode:2019Natur.570..182S. doi:10.1038/s41586-019-1279-z. hdl:1887/3198847. ISSN 1476-4687. PMID 31168093. S2CID 174809069. Most modern Siberian speakers of Neosiberian languages genetically fall on an East- West cline between Europeans and Early East Asians. Taking Even speakers as representatives, the Neosiberian turnover from the south, which largely replaced Ancient Paleosiberian ancestry, can be associated with the northward spread of Tungusic and probably also Turkic and Mongolic. However, the expansions of Tungusic as well as Turkic and Mongolic are too recent to be associable with the earliest waves of Neosiberian ancestry, dated later than ~11 kya, but discernible in the Baikal region from at least 6 kya onwards. Therefore, this phase of the Neosiberian population turnover must initially have transmitted other languages or language families into Siberia, including possibly Uralic and Yukaghir.
  24. ^ Wang, Ke (2023). "Middle Holocene Siberian genomes reveal highly connected gene pools throughout North Asia". Current Biology. 33 (3): 423–433.e5. Bibcode:2023CBio...33E.423W. doi:10.1016/j.cub.2022.11.062. PMID 36638796. S2CID 255750546. We find the presence of ancient Northeast Asian (ANA) ancestry —initially described in Neolithic groups from the Russian Far East— in another Neolithic Altai-Sayan individual associated with different cultural features, revealing the spread of ANA ancestry 1,500 km further to the west than previously observed.
  25. ^ Lee, Juhyeon (14 April 2023). "Genetic population structure of the Xiongnu Empire at imperial and local scales". Science Advances. 9 (15): eadf3904. Bibcode:2023SciA....9F3904L. doi:10.1126/sciadv.adf3904. ISSN 2375-2548. PMC 10104459. PMID 37058560. S2CID 258153066. Overall, individuals from the Ulaanzuukh and the Slab Grave cultures present a homogeneous genetic profile that has deep roots in the region and is referred to as Ancient Northeast Asian (ANA)
  26. ^ Gnecchi-Ruscone et al. 2021, Fig.2.
  27. ^ Gnecchi-Ruscone, Guido Alberto (26 March 2021). "Ancient genomic time transect from the Central Asian Steppe unravels the history of the Scythians". Science Advances. 7 (13): Fig.2. Bibcode:2021SciA....7.4414G. doi:10.1126/sciadv.abe4414. ISSN 2375-2548. PMC 7997506. PMID 33771866.
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  29. ^ Jeong, Choongwon; Wilkin, Shevan; Amgalantugs, Tsend; Bouwman, Abigail S.; Taylor, William Timothy Treal; Hagan, Richard W.; Bromage, Sabri; Tsolmon, Soninkhishig; Trachsel, Christian; Grossmann, Jonas; Littleton, Judith; Makarewicz, Cheryl A.; Krigbaum, John; Burri, Marta; Scott, Ashley (27 November 2018). "Bronze Age population dynamics and the rise of dairy pastoralism on the eastern Eurasian steppe". Proceedings of the National Academy of Sciences. 115 (48): E11248–E11255. Bibcode:2018PNAS..11511248J. doi:10.1073/pnas.1813608115. ISSN 0027-8424. PMC 6275519. PMID 30397125.
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  32. ^ Jeong, Choongwon (2020). "A Dynamic 6,000-Year Genetic History of Eurasia's Eastern Steppe". Cell. 183 (4): 890–904. doi:10.1016/j.cell.2020.10.015. ISSN 0092-8674. PMC 7664836. PMID 33157037.
  33. ^ Jeong, Choongwon; Wilkin, Shevan; Amgalantugs, Tsend; Warinner, Christina (2018). "Bronze Age population dynamics and the rise of dairy pastoralism on the eastern Eurasian steppe". Proceedings of the National Academy of Sciences. 115 (48). Proceedings of the National Academy of Sciences of the United States of America: E11248–E11255. Bibcode:2018PNAS..11511248J. doi:10.1073/pnas.1813608115. PMC 6275519. PMID 30397125. S2CID 53230942.
  34. ^ Rawson, Jessica; Huan, Limin; Taylor, William Timothy Treal (1 December 2021). "Seeking Horses: Allies, Clients and Exchanges in the Zhou Period (1045–221 BC)". Journal of World Prehistory. 34 (4): 489–530. doi:10.1007/s10963-021-09161-9. ISSN 1573-7802.
  35. ^ Zhang et al. 2021, "Using qpAdm, we modelled the Tarim Basin individuals as a mixture of two ancient autochthonous Asian genetic groups: the ANE, represented by an Upper Palaeolithic individual from the Afontova Gora site in the upper Yenisei River region of Siberia (AG3) (about 72%), and ancient Northeast Asians, represented by Baikal_EBA (about 28%) (Supplementary Data 1E and Fig. 3a). Tarim_EMBA2 from Beifang can also be modelled as a mixture of Tarim_EMBA1 (about 89%) and Baikal_EBA (about 11%).".
  36. ^ Nägele, Kathrin; Rivollat, Maite; Yu, He; Wang, Ke (2022). "Ancient genomic research - From broad strokes to nuanced reconstructions of the past". Journal of Anthropological Sciences. 100 (100): 193–230. doi:10.4436/jass.10017. PMID 36576953. Combining genomic and proteomic evidence, researchers revealed that these earliest residents in the Tarim Basin carried genetic ancestry inherited from local Upper Palaeolithic hunter-gatherers, carried no steppe-related ancestry, but consumed milk products, indicating communications of persistence practices independent from genetic exchange.
  37. ^ a b c Jeong, Choongwon (2020). "A Dynamic 6,000-Year Genetic History of Eurasia's Eastern Steppe". Cell. 183 (4): 890–904.e29. doi:10.1016/j.cell.2020.10.015. ISSN 0092-8674. PMC 7664836. PMID 33157037. S2CID 214725595.
  38. ^ Gnecchi-Ruscone, Guido Alberto (14 April 2022). "Ancient genomes reveal origin and rapid trans-Eurasian migration of 7th century Avar elites". Cell. 185 (8): 1402–1413.e21. doi:10.1016/j.cell.2022.03.007. ISSN 0092-8674. PMC 9042794. PMID 35366416. S2CID 247859905. All of these individuals, albeit variably mixed with other sources, have been shown to trace their eastern Eurasian ancestry component to a genetic profile referred to as the "ancient northeast Asians" (ANA) (...) All of the early-Avar-period individuals (DTI_early_elite), except for an infant and a burial with typical characteristics of the Transtisza group (Figure 2B), form a tight cluster with a high level of ANA ancestry.
  39. ^ a b Lee, Juhyeon; Miller, Bryan K.; Bayarsaikhan, Jamsranjav; Johannesson, Erik; Ventresca Miller, Alicia; Warinner, Christina; Jeong, Choongwon (14 April 2023). "Genetic population structure of the Xiongnu Empire at imperial and local scales". Science Advances. 9 (15): eadf3904. Bibcode:2023SciA....9F3904L. doi:10.1126/sciadv.adf3904. ISSN 2375-2548. PMC 10104459. PMID 37058560. "In this genome-wide archaeogenetic study, we find high genetic heterogeneity among late Xiongnu-era individuals at two cemeteries located along the far western frontier of the Xiongnu empire and describe patterns of genetic diversity related to social status. Overall, we find that genetic heterogeneity is highest among lower-status individuals. In particular, the satellite graves surrounding the elite square tombs at TAK show extreme levels of genetic heterogeneity, suggesting that these individuals, who were likely low-ranking retainers, were drawn from diverse parts of the empire. In contrast, the highest-status individuals at the two sites tended to have lower genetic diversity and a high proportion of ancestry deriving from EIA Slab Grave groups, suggesting that these groups may have disproportionately contributed to the ruling elite during the formation of the Xiongnu empire." (...) "a chanyu, or ruler of the empire. Like the elite women at the western frontier, he also had very high eastern Eurasian ancestry (deriving 39.3 and 51.9% from SlabGrave1 and Han_2000BP, respectively, and the rest from Chandman_IA; data file S2C)" (...) "Chandman_IA was representative of people in far western Mongolia associated with Sagly/Uyuk (ca. 500 to 200 BCE), Saka (ca. 900 to 200 BCE), and Pazyryk (ca. 500 to 200 BCE) groups in Siberia and Kazakhstan." (...) "This further suggests the existence of an aristocracy in the Xiongnu empire, that elite status and power was concentrated within specific subsets of the broader population."... Although not conclusive, this suggests that the ANA ancestry source of the Xiongnu-period individuals may not be exclusively traced back to the Slab Grave culture but may also include nearby groups with a similar ANA genetic profile, such as the Xianbei. ... Last, our findings also confirm that the highest-status individuals in this study were females, supporting previous observations that Xiongnu women played an especially prominent role in the expansion and integration of new territories along the empire's frontier.
  40. ^ Gnecchi-Ruscone, Guido Alberto (26 March 2021). "Ancient genomic time transect from the Central Asian Steppe unravels the history of the Scythians". Science Advances. 7 (13): Fig.3A. Bibcode:2021SciA....7.4414G. doi:10.1126/sciadv.abe4414. ISSN 2375-2548. PMC 7997506. PMID 33771866.
  41. ^ Gnecchi-Ruscone, Guido Alberto (14 April 2022). "Ancient genomes reveal origin and rapid trans-Eurasian migration of 7th century Avar elites". Cell. 185 (8): 1402–1413.e21. doi:10.1016/j.cell.2022.03.007. ISSN 0092-8674. PMC 9042794. PMID 35366416. All of these individuals, albeit variably mixed with other sources, have been shown to trace their eastern Eurasian ancestry component to a genetic profile referred to as the "ancient northeast Asians" (ANA) (...) All of the early-Avar-period individuals (DTI_early_elite), except for an infant and a burial with typical characteristics of the Transtisza group (Figure 2B), form a tight cluster with a high level of ANA ancestry.
  42. ^ Yang, Xiaomin (2023). "Ancient Genome of Empress Ashina reveals the Northeast Asian origin of Göktürk Khanate". Journal of Systematics and Evolution. 61 (6): 1056–1064. doi:10.1111/jse.12938. S2CID 255690237. In the principal component analysis (PCA) (Figs. 1B, S3), the Ashina individual clustered with modern Tungusic and Mongolic speakers, ancient populations from Northeast Asia and eastern Mongolia Plateau, and especially with the Northeast Asian hunter-gatherers previously referred to as "Ancient Northeast Asian" (ANA), that is, DevilsCave_N, Mongolia_N_North, Boisman_MN, AR_EN (Jeong et al., 2020; Ning et al., 2020; Wang et al., 2021), as well as post-Iron Age Eastern Steppe nomadic people including Xianbei, Rouran, Khitan, and part of the Mongol population. (...) In summary, we have unveiled the first genomic profile of theancient Türkic royal family. Our genomic analyses of EmpressAshina revealed Göktürk's Northeast Asian origin (97.7% Northeast Asian ancestry and 2.3% West Eurasian ancestry), refuting the western Eurasian origin and multiple originhypotheses. We found Ashina shared most genetic affinity with post-Iron Age Tungusic and Mongolic Steppe pastoralists, such as Rouran, Xianbei, Khitan, and Heshui_Mohe, and showedgenetic heterogeneity with other ancient Türkic people, suggesting the multiple sources of the Türkic Khanate populations. Furthermore, the limited contribution from ancient Göktürk found in modern Turkic-speaking populations once again validates a cultural diffusion model over a demic diffusion model for the spread of Turkic languages. (...) Table S2F: "The Ashina showed west Eurasian admixture dating to about 2000 years ago (~1330–1550 years, assuming 29 years/ generation. Mongolia_N_North + Sarmatian/Mongolia_N_North + Afanasievo), while earlyMed_Turk and CentralSteppe_Turk showed more recently mixing date."
  43. ^ Yang 2023, "In summary, we have unveiled the first genomic profile of the ancient Türkic royal family. Our genomic analyses of Empress Ashina revealed Göktürk's Northeast Asian origin (97.7% Northeast Asian ancestry and 2.3% West Eurasian ancestry), refuting the western Eurasian origin and multiple origin hypotheses.".
  44. ^ Yang 2023, pp. 3–4: "In contrast to Ashina, Central Steppe and early Medieval Türk exhibited a high but variable degree of West Eurasian ancestry, indicating there was a genetic substructure of the Türkic empire."
  45. ^ Yang 2023, p. 5: "We found Ashina shared most genetic affnity with post Iron Age Tungusic and Mongolic Steppe pastoralists, such as Rouran, Xianbei, Khitan, and Heshui_Mohe, and showed genetic heterogeneity with other ancient Türkic people, suggesting the multiple sources of the Türkic Khanate populations"
  46. ^ Jeong et al. 2020, p. 897: See figure 4, B for admixture proportions in earlyMed_Turk. "...it is clear that these individuals have genetic profiles that differ from the preceding Xiongnu period, suggesting new sources of gene flow into Mongolia at this time that displace them along PC3 (Figure 2)...The admixture dates estimated for the ancient Türkic and Uyghur individuals in this study correspond to ca. 500 CE: 8 ± 2 generations before the Türkic individuals and 12 ± 2 generations before the Uyghur individuals (represented by ZAA001 and Olon Dov individuals)."
  47. ^ Yang 2023, p. 4: "The early Medieval Türk (earlyMed_Turk) derived the major ancestry from ANA at a proportion of 62.2%, the remainder from BMAC (10.7%) and Western Steppe Afanasievo nomad (27.1%) (Figs. 1C, 1D; Table S2E)."
  48. ^ Yang, Xiao-Min; Meng, Hai-Liang; Zhang, Jian-Lin; Yu, Yao; Allen, Edward; Xia, Zi-Yang; Zhu, Kong-Yang; Du, Pan-Xin; Ren, Xiao-Ying; Xiong, Jian-Xue; Lu, Xiao-Yu; Ding, Yi; Han, Sheng; Liu, Wei-Peng; Jin, Li (9 January 2023). "Ancient genome of Empress Ashina reveals the Northeast Asian origin of Göktürk Khanate". Journal of Systematics and Evolution. 61 (6): 1056–1064. doi:10.1111/jse.12938. ISSN 1674-4918. In addition, Ashina showed close genetic affinity with population related to Bronze Age Slab Grave and Ulaanzukh culture in Mongolia.
  49. ^ Sikora, Martin; Pitulko, Vladimir V.; Sousa, Vitor C.; Allentoft, Morten E.; Vinner, Lasse; Rasmussen, Simon; Margaryan, Ashot; de Barros Damgaard, Peter; de la Fuente, Constanza; Renaud, Gabriel; Yang, Melinda A.; Fu, Qiaomei; Dupanloup, Isabelle; Giampoudakis, Konstantinos; Nogués-Bravo, David (2019). "The population history of northeastern Siberia since the Pleistocene". Nature. 570 (7760): 182–188. Bibcode:2019Natur.570..182S. doi:10.1038/s41586-019-1279-z. hdl:1887/3198847. ISSN 1476-4687. PMID 31168093. S2CID 174809069. Most modern Siberian speakers of Neosiberian languages genetically fall on an East- West cline between Europeans and Early East Asians. Taking Even speakers as representatives, the Neosiberian turnover from the south, which largely replaced Ancient Paleosiberian ancestry, can be associated with the northward spread of Tungusic and probably also Turkic and Mongolic. However, the expansions of Tungusic as well as Turkic and Mongolic are too recent to be associable with the earliest waves of Neosiberian ancestry, dated later than ~11 kya, but discernible in the Baikal region from at least 6 kya onwards. Therefore, this phase of the Neosiberian population turnover must initially have transmitted other languages or language families into Siberia, including possibly Uralic and Yukaghir.

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