Ancient East Eurasians

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The term Ancient East Eurasian, alternatively also known as East Eurasian or Eastern Eurasian, is used in population genomics to describe the genetic ancestry and phylogenetic relationship of diverse populations primarily living in the Asia-Pacific region, belonging to the "Eastern Eurasian clade" of human genetic diversity, [1] [2] [3] [4] [5] [6] and which can be associated with the Initial Upper Paleolithic (IUP) wave, following the Out of Africa migration at least 60,000 years ago. [4] [6]

Contents

Early dispersals

Repetitive expansions into Eurasia from a population Hub OoA. Representative samples dated between 45 and 40 ka across Eurasia can be ascribed to a population movement with uniform genetic features and material culture consistent with an IUP affiliation. Repetitive expansions into Eurasia from a population Hub OoA.jpg
Repetitive expansions into Eurasia from a population Hub OoA. Representative samples dated between 45 and 40 ka across Eurasia can be ascribed to a population movement with uniform genetic features and material culture consistent with an IUP affiliation.
Microblade and core & flake sites in Eastern Asia Microblade and core & flake sites in Eastern Asia.webp
Microblade and core & flake sites in Eastern Asia

Ancient East Eurasians, modern humans of the Initial Upper Paleolithic (IUP) wave, are suggested to have diverged from Ancient West Eurasians around 48,000 years ago from a population hub likely on the Persian plateau, and expanded across Eurasia through a star-like expansion pattern at least 45,000 years ago. [7] [6]

Ancient East Eurasians started to diversify among themselves as early as 45,000 years ago. [6] This diversification may possibly be related to the development of two major IUP-affiliated types of material culture: "microlithic blade" (or microblade) and "core & flake" (or CAF assemblages). Specific IUP populations represented by specimens found in Europe, Central Asia, and Siberia, such as the Ust'-Ishim man, Bacho Kiro, Oase 2, and Kara-Bom, that are associated with the spread of microblade sites, are inferred to have used inland routes northward into Eurasia. [8] [9] [10] In contrast, IUP populations represented by specimens found in South, Southeast, and East Asia, as well as Oceania, such as the Tianyuan man, that are associated with the spread of core & flake sites, are inferred to have used a Southern dispersal route along the southern coast of Asia. [11] [12] [13]

The IUP populations that expanded through Southern dispersal route are inferred to be the ancestors of all modern East Eurasian populations (dubbed as "East Eurasian Core"; EEC). EEC populations subsequently diversified rapidly from South and Southeast Asia at least 40,000 years ago, and became ancestral to modern populations in Eastern Eurasia, Oceania, and the Americas, notably East Asians, Southeast Asians, Indigenous Siberians, Aboriginal Australians, Papuans, Pacific Islanders, Indigenous Americans, and partly South and Central Asians. [7] [14] [15] [16] [17] [18] [19] [20] [21] South Asia may have acted as central hub for the peopling of Eastern Asia and Australasia. [22]

Inferred model for the phylogenetic substructure of Eastern Eurasian populations. Phylogenetic structure of Eastern Eurasians.png
Inferred model for the phylogenetic substructure of Eastern Eurasian populations.

East Eurasian lineages

Major East Eurasian ancestry lineages which contributed to modern human populations include the following: [14]

Estimated ancestry components among selected modern populations per Changmai et al. (2022). K6 human genetic ancestries.png
Estimated ancestry components among selected modern populations per Changmai et al. (2022).

The "Australasian," "Ancient Ancestral South Indian," and "East and Southeast Asian" lineages display a closer genetic relationship to each other than to any non-Asian lineages, as well as being closer to each other than to any of the early East Eurasian IUP lineages (Bacho Kiro etc.), and together represent the main branches of "Asian-related ancestry", which diverged from each other at least 40,000 years ago. [14] The Australasian lineage however received higher archaic admixture in the Oceania region, and may also harbor some small amounts of "xOoA" admixture from an earlier human dispersal, which did not contribute to any other human population. Alternatively, Australasians can be described as nearly equally admixture between a "Basal East Asian" source (represented by Tianyuan or Onge) and a deeper East Eurasian lineage not sampled yet. [7] [1] [14] : 11 

Traces of an unsampled deeply diverged East Eurasian lineage can be observed in the genome of ancient and modern inhabitants of the Tibetan Plateau. While modern Tibetans mostly derive their ancestry from a northern East Asian source (specifically Yellow River farmers), a minor, but significant contribution stems from a deeply diverged East Eurasian local "Ghost population" that was distinct from other deeply diverged lineages such as Ust'Ishim, Hoabinhian/Onge or Tianyuan, representing the local Paleolithic population of the Tibetan Plateau. [26] [27] The remains of a 7,100 year old specimen from Yunnan, known as Xingyi_EN, were found to represent a new deeply branching Basal Asian lineage, which is closely related to the inferred Ghost ancestry among Tibetans. [28]

Deeper IUP-associated East Eurasian lineages have been associated with the remains of the Oase and Bacho Kiro cave specimens in southeastern Europe, and represent an inland migration northwards associated with the dispersal of IUP material culture, deeply diverged from all other East Eurasian populations (EEC). These deep East Eurasian populations did not contribute significantly to later Eurasian populations, except variable amounts of geneflow to subsequent Upper Paleolithic European groups, such as the Goyet Caves specimen (GoyetQ116-1; c. 17–23%) associated with the Aurignacian culture, or the Sungir specimen (c. 0–14%) associated with the Gravettian culture (via Aurignacian geneflow). In contrast, the Upper Paleolithic European Kostenki-14 specimen, did not display evidence for IUP-affilated admixture. [7] [29] [21] [14] [30] [31]

A deep IUP-affilated lineage may have also contributed ancestry (up to 39%) to the Tianyuan man, explaining the observed affinity between Tianyuan and GoyetQ116-1, as well as GoyetQ116-1 and BachoKiro_IUP. This variegated East Eurasian substrate found among the GoyetQ116-1 specimen may hint to "yet undescribed complexities within the IUP population branch". [31] [7] This lineage may also be affilated with IUP sites in Siberia and Northwest China, but currently no archaeogenetic data for these sites is available. [21]

The exact relationship of the Ust'-Ishim man from Siberia to other IUP/East Eurasian lineages is not well resolved yet. The Ust'-Ishim man forms a near trifurication between West Eurasian (Kostenki-14) and IUP/East Eurasian lineages, but shares a short period of evolutionary drift with Eastern Eurasians. [7] [6] The Ust'Ishim lineage is inferred to have not contributed ancestry to modern human populations. [21]

References

  1. 1 2 Lipson, Mark; Reich, David (2017). "A working model of the deep relationships of diverse modern human genetic lineages outside of Africa". Molecular Biology and Evolution. 34 (4): 889–902. doi:10.1093/molbev/msw293. ISSN   0737-4038. PMC   5400393 . PMID   28074030. Archived from the original on 20 December 2022. Retrieved 11 May 2023.
  2. ↑ Skoglund, Pontus; Mathieson, Iain (31 August 2018). "Ancient Genomics of Modern Humans: The First Decade". Annual Review of Genomics and Human Genetics. 19 (1): 381–404. doi: 10.1146/annurev-genom-083117-021749 . ISSN   1527-8204. PMID   29709204. S2CID   19933330.
  3. ↑ Zhang, Ming; Fu, Qiaomei (1 June 2020). "Human evolutionary history in Eastern Eurasia using insights from ancient DNA". Current Opinion in Genetics & Development. Genetics of Human Origin. 62: 78–84. doi:10.1016/j.gde.2020.06.009. ISSN   0959-437X. PMID   32688244. S2CID   220671047.
  4. 1 2 Vallini, Leonardo; Pagani, Luca (2022). "The future of the Eurasian past: highlighting plotholes and pillars of human population movements in the Late Pleistocene". Journal of Anthropological Sciences. 100 (100): 231–241. doi:10.4436/JASS.10013. ISSN   1827-4765. PMID   36565457.
  5. ↑ 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.
  6. 1 2 3 4 5 Vallini, Leonardo; Zampieri, Carlo; Shoaee, Mohamed Javad; Bortolini, Eugenio; Marciani, Giulia; Aneli, Serena; Pievani, Telmo; Benazzi, Stefano; Barausse, Alberto; Mezzavilla, Massimo; Petraglia, Michael D.; Pagani, Luca (25 March 2024). "The Persian plateau served as hub for Homo sapiens after the main out of Africa dispersal". Nature Communications. 15 (1): 1882. Bibcode:2024NatCo..15.1882V. doi:10.1038/s41467-024-46161-7. ISSN   2041-1723. PMC   10963722 . PMID   38528002. ... and the split between EEC and WEC, with the former leaving the Hub18, 46 kya (allowing the time for them to reach Ust'Ishim and Bacho Kiro by ~45 kya).
  7. 1 2 3 4 5 6 Vallini et al. 2022 (4 July 2022). "Genetics and Material Culture Support Repeated Expansions into Paleolithic Eurasia from a Population Hub Out of Africa". Genome Biology and Evolution. 14 (4). doi:10.1093/gbe/evac045. hdl: 2318/1855565 . PMID   35445261. Archived from the original on 21 April 2023. Retrieved 16 April 2023.{{cite journal}}: CS1 maint: numeric names: authors list (link)
  8. ↑ Vallini, Leonardo; Zampieri, Carlo; Shoaee, Mohamed Javad; Bortolini, Eugenio; Marciani, Giulia; Aneli, Serena; Pievani, Telmo; Benazzi, Stefano; Barausse, Alberto; Mezzavilla, Massimo; Petraglia, Michael D.; Pagani, Luca (25 March 2024). "The Persian plateau served as hub for Homo sapiens after the main out of Africa dispersal". Nature Communications. 15 (1): 1882. Bibcode:2024NatCo..15.1882V. doi:10.1038/s41467-024-46161-7. ISSN   2041-1723. PMC   10963722 . PMID   38528002.
  9. ↑ Bennett, E. Andrew; Liu, Yichen; Fu, Qiaomei (3 December 2024). "Reconstructing the Human Population History of East Asia through Ancient Genomics". Elements in Ancient East Asia. doi: 10.1017/9781009246675 . ISBN   978-1-009-24667-5. Archived from the original on 25 January 2025. Retrieved 25 January 2025.
  10. ↑ Yang, Ziyi; Jin, Yingshuai; Wang, Shejiang; Zhang, Xiaoling (2023). "Discovery and research review of knapped lithics of the South Asian subcontinent". 人类学学报. 42 (3). doi:10.16359/j.1000-3193/AAS.2023.0019.
  11. ↑ Bennett, E. Andrew; Liu, Yichen; Fu, Qiaomei (3 December 2024). "Reconstructing the Human Population History of East Asia through Ancient Genomics". Elements in Ancient East Asia. doi: 10.1017/9781009246675 . ISBN   978-1-009-24667-5. ... ancient and modern genomic studies appear to favor a southern route into East Asia for the majority of genetic diversity present there today. ... If these East Asian IUP sites were to be linked to populations related to Ust' Ishim it would appear these people left no detectable genetic legacy in modern East Asia. It may also be found that the material at some of these IUP sites was created by populations derived from East Asian lineages linked to or branching from Tianyuan.
  12. ↑ Abood, Steven; Oota, Hiroki (14 February 2025). "Human dispersal into East Eurasia: ancient genome insights and the need for research on physiological adaptations". Journal of Physiological Anthropology. 44 (1): 5. doi: 10.1186/s40101-024-00382-3 . ISSN   1880-6805. PMC   11829451 . PMID   39953642. The results indicated that all modern East Asians, Northeast Asians, and Native Americans, including this Jomon individual, are descendants of populations that reached the eastern side of the Eurasian continent via the southern route.
  13. ↑ "Following Pleistocene road signs of human dispersals across Eurasia". ResearchGate. Archived from the original on 2 March 2023. Retrieved 22 February 2025.
  14. 1 2 3 4 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. Archived from the original on 16 April 2023. Retrieved 11 May 2023.
  15. ↑ Sato, Takehiro; Adachi, Noboru; Kimura, Ryosuke; Hosomichi, Kazuyoshi; Yoneda, Minoru; Oota, Hiroki; Tajima, Atsushi; Toyoda, Atsushi; Kanzawa-Kiriyama, Hideaki; Matsumae, Hiromi; Koganebuchi, Kae (1 September 2021). "Whole-Genome Sequencing of a 900-Year-Old Human Skeleton Supports Two Past Migration Events from the Russian Far East to Northern Japan". Genome Biology and Evolution. 13 (9): evab192. doi:10.1093/gbe/evab192. ISSN   1759-6653. PMC   8449830 . PMID   34410389. the southern migration wave seems to have diversified into the local populations in East Asia (defined in this paper as a region including China, Japan, Korea, Mongolia, Taiwan and Southeast Asia), and the northern wave, which probably runs through the Siberian and Eurasian steppe regions and mixed with the southern wave, probably in Siberia.
  16. ↑ Osada, Naoki; Kawai, Yosuke (2021). "Exploring models of human migration to the Japanese archipelago using genome-wide genetic data". Anthropological Science. 129 (1): 45–58. doi: 10.1537/ase.201215 . S2CID   234247309. Archived from the original on 23 February 2023. Retrieved 10 November 2023. Via the southern route, ancestors of current Asian populations reached Southeast Asia and a part of Oceania around 70000–50000 years ago, probably through a coastal dispersal route (Bae et al., 2017). The oldest samples providing the genetic evidence of the northern migration route come from a high-coverage genome sequence of individuals excavated from the Yana RHS site in northeastern Siberia (Figure 2), which is about 31600 years old (Sikora et al., 2019).
  17. ↑ 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. Population genomic studies on present-day humans7,8 have exclusively supported the southern route origin of East Asian populations.
  18. ↑ Aoki, Kenichi; Takahata, Naoyuki; Oota, Hiroki; Wakano, Joe Yuichiro; Feldman, Marcus W. (30 August 2023). "Infectious diseases may have arrested the southward advance of microblades in Upper Palaeolithic East Asia". Proceedings of the Royal Society B: Biological Sciences. 290 (2005). doi:10.1098/rspb.2023.1262. ISSN   0962-8452. PMC   10465978 . PMID   37644833. A single major migration of modern humans into the continents of Asia and Sahul was strongly supported by earlier studies using mitochondrial DNA, the non-recombining portion of Y chromosomes, and autosomal SNP data [42–45]. Ancestral Ancient South Indians with no West Eurasian relatedness, East Asians, Onge (Andamanese hunter–gatherers) and Papuans all derive in a short evolutionary time from the eastward dispersal of an out-of-Africa population [46,47]. The HUGO (Human Genome Organization) Pan-Asian SNP consortium [44] investigated haplotype diversity within present-day Asian populations and found a strong correlation with latitude, with diversity decreasing from south to north. The correlation continues to hold when only mainland Southeast Asian and East Asian populations are considered, and is perhaps attributable to a serial founder effect [50]. These observations are consistent with the view that soon after the single eastward migration of modern humans, East Asians diverged in southern East Asia and dispersed northward across the continent.
  19. ↑ Demeter, Fabrice; Shackelford, Laura L.; Bacon, Anne-Marie; Duringer, Philippe; Westaway, Kira; Sayavongkhamdy, Thongsa; Braga, José; Sichanthongtip, Phonephanh; Khamdalavong, Phimmasaeng; Ponche, Jean-Luc; Wang, Hong; Lundstrom, Craig; Patole-Edoumba, Elise; Karpoff, Anne-Marie (4 September 2012). "Anatomically modern human in Southeast Asia (Laos) by 46 ka". Proceedings of the National Academy of Sciences. 109 (36): 14375–14380. Bibcode:2012PNAS..10914375D. doi: 10.1073/pnas.1208104109 . ISSN   0027-8424. PMC   3437904 . PMID   22908291. Inferences from nuclear (51), Y chromosome (52), and mitochondrial genome (53) data support an early migration of modern humans out of Africa and into Southeast Asia using a southern route by at least 60 ka. Patterns of genetic variation in recent human populations (11, 54, 55) recognize Southeast Asia as an important source for the peopling of East Asia and Australasia via a rapid, early settlement.
  20. ↑ Hao, Li (2022). "Exploring the southern dispersal routes of early modern humans". Acta Anthropologica Sinica. 41 (4): 630–648. Archived from the original on 25 December 2024. Retrieved 4 November 2024. Based on the above findings, some scholars have proposed that the specific diffusion path of early modern humans in the south was quite complex, possibly including both coastal routes (some sites may be submerged under the sea) and inland routes. Field J et al.'s minimum cost path analysis based on GIS technology confirmed this: early modern humans formed multiple paths during their diffusion, and their inland paths were mostly based on rivers as migration corridors, such as the Indus River and the Narmada River in the Indian Peninsula.
  21. 1 2 3 4 5 Bennett, E. Andrew; Liu, Yichen; Fu, Qiaomei (3 December 2024). "Reconstructing the Human Population History of East Asia through Ancient Genomics". Elements in Ancient East Asia. doi: 10.1017/9781009246675 . ISBN   978-1-009-24667-5. Archived from the original on 25 January 2025. Retrieved 25 January 2025.
  22. ↑ Mukhopadhyay, Sourav; Gupta, Antara; Kumar, Pankaj; Sukumaran, Prabhin; Sabale, Panduranga D. (1 March 2025). "Understanding the Microlithic technology in the Lower Ganga Basin, Eastern India: A chronological and ecological perspective". Quaternary Environments and Humans. 3 (1): 100059. doi: 10.1016/j.qeh.2025.100059 . ISSN   2950-2365. To understand the demographic pattern, current ancient DNA (aDNA) studies highlight India as a central hub for the later colonization of Asia and Australia
  23. ↑ Yelmen, Burak; Mondal, Mayukh; Marnetto, Davide; Pathak, Ajai K; Montinaro, Francesco; Gallego Romero, Irene; Kivisild, Toomas; Metspalu, Mait; Pagani, Luca (5 April 2019). "Ancestry-Specific Analyses Reveal Differential Demographic Histories and Opposite Selective Pressures in Modern South Asian Populations". Molecular Biology and Evolution. 36 (8): 1628–1642. doi:10.1093/molbev/msz037. ISSN   0737-4038. PMC   6657728 . PMID   30952160.
  24. ↑ Kusuma, Pradiptajati; Cox, Murray P.; Barker, Graeme; Sudoyo, Herawati; Lansing, J. Stephen; Jacobs, Guy S. (1 November 2023). "Deep ancestry of Bornean hunter-gatherers supports long-term local ancestry dynamics". Cell Reports. 42 (11): 113346. doi: 10.1016/j.celrep.2023.113346 . ISSN   2211-1247. PMID   37917587.
  25. ↑ Changmai, Piya; Pinhasi, Ron; Pietrusewsky, Michael; Stark, Miriam T.; Ikehara-Quebral, Rona Michi; Reich, David; Flegontov, Pavel (29 December 2022). "Ancient DNA from Protohistoric Period Cambodia indicates that South Asians admixed with local populations as early as 1st–3rd centuries CE". Scientific Reports. 12 (1): 22507. Bibcode:2022NatSR..1222507C. doi:10.1038/s41598-022-26799-3. ISSN   2045-2322. PMC   9800559 . PMID   36581666.
  26. ↑ Liu, Chi-Chun; Witonsky, David; Gosling, Anna; Lee, Ju Hyeon; Ringbauer, Harald; Hagan, Richard; Patel, Nisha; Stahl, Raphaela; Novembre, John; Aldenderfer, Mark; Warinner, Christina; Di Rienzo, Anna; Jeong, Choongwon (8 March 2022). "Ancient genomes from the Himalayas illuminate the genetic history of Tibetans and their Tibeto-Burman speaking neighbors". Nature Communications. 13 (1): 1203. Bibcode:2022NatCo..13.1203L. doi:10.1038/s41467-022-28827-2. ISSN   2041-1723. PMC   8904508 . PMID   35260549. our results reject previously suggested sources of gene flow into the Tibetan lineage13,35,36, including deeply branching Eastern Eurasian lineages, such as the 45,000-year-old Ust'-Ishim individual from southern Siberia, the 40,000-year-old Tianyuan individual from northern China, and Hoabinhian/Onge-related lineages in southeast Asia (Supplementary Fig. 10), suggesting instead that it represents yet another unsampled lineage within early Eurasian genetic diversity. This deep Eurasian lineage is likely to represent the Paleolithic genetic substratum of the Plateau populations.
  27. ↑ Wang, Hongru; Yang, Melinda A.; Wangdue, Shargan; Lu, Hongliang; Chen, Honghai; Li, Linhui; Dong, Guanghui; Tsring, Tinley; Yuan, Haibing; He, Wei; Ding, Manyu; Wu, Xiaohong; Li, Shuai; Tashi, Norbu; Yang, Tsho (15 March 2023). "Human genetic history on the Tibetan Plateau in the past 5100 years". Science Advances. 9 (11): eadd5582. Bibcode:2023SciA....9D5582W. doi:10.1126/sciadv.add5582. ISSN   2375-2548. PMC   10022901 . PMID   36930720.
  28. ↑ Wang, Tianyi; Yang, Melinda A.; Zhu, Zhonghua; Ma, Minmin; Shi, Han; Speidel, Leo; Min, Rui; Yuan, Haibing; Jiang, Zhilong; Hu, Changcheng; Li, Xiaorui; Zhao, Dongyue; Bai, Fan; Cao, Peng; Liu, Feng (29 May 2025). "Prehistoric genomes from Yunnan reveal ancestry related to Tibetans and Austroasiatic speakers". Science. 388 (6750): eadq9792. doi:10.1126/science.adq9792.
  29. ↑ Vallini, Pagani, Leonardo, Luca (December 2022). "The future of the Eurasian past: highlighting plotholes and pillars of human population movements in the Late Pleistocene". Journal of Anthropological Sciences. 100 (100): 231–241. doi:10.4436/JASS.10013. PMID   36565457.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  30. ↑ Massilani, Diyendo; Skov, Laurits; Hajdinjak, Mateja; Gunchinsuren, Byambaa; Tseveendorj, Damdinsuren; Yi, Seonbok; Lee, Jungeun; Nagel, Sarah; Nickel, Birgit; Devièse, Thibaut; Higham, Tom; Meyer, Matthias; Kelso, Janet; Peter, Benjamin M.; Pääbo, Svante (30 October 2020). "Denisovan ancestry and population history of early East Asians". Science. 370 (6516): 579–583. doi:10.1126/science.abc1166. PMID   33122380.
  31. 1 2 Hajdinjak, Mateja; Mafessoni, Fabrizio; Skov, Laurits; Vernot, Benjamin; Hübner, Alexander; Fu, Qiaomei; Essel, Elena; Nagel, Sarah; Nickel, Birgit; Richter, Julia; Moldovan, Oana Teodora; Constantin, Silviu; Endarova, Elena; Zahariev, Nikolay; Spasov, Rosen (7 April 2021). "Initial Upper Palaeolithic humans in Europe had recent Neanderthal ancestry". Nature. 592 (7853): 253–257. Bibcode:2021Natur.592..253H. doi:10.1038/s41586-021-03335-3. ISSN   1476-4687. PMC   8026394 . PMID   33828320.
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