Caucasus hunter-gatherer

Caucasus hunter-gatherer
PeriodUpper Paleolithic, Mesolithic
Dates13,000–6,000 BC

Caucasus hunter-gatherer (CHG), also called Satsurblia cluster,[1] is an anatomically modern human genetic lineage, first identified in a 2015 study,[2][3] based on the population genetics of several modern Western Eurasian (European, Caucasian and Near Eastern) populations.[4][5]

Genetic structure of ancient Europe. Caucasus hunter-gatherers are represented by the Satsurbila and Kotias specimens.
Genetic affinity of modern populations to the ancient Kotias specimen.
Admixture graph of deep Eurasian lineages (Allentoft et al. 2024)

The CHG lineage is suggested to have diverged from the ancestor of Western Hunter-Gatherers (WHGs) probably during the Last Glacial Maximum (sometimes between 45,000 to 26,000 years ago).[6] They further separated from the "Anatolian Hunter Gatherer" (AHG) lineage later, suggested to around 25,000 years ago during the late LGM period.[3][7] The Caucasus hunter-gatherers managed to survive in isolation since the late LGM period as a distinct population, and display high genetic affinities to Mesolithic and Neolithic populations on the Iranian plateau, such as Neolithic specimens found in Ganj Dareh. The CHG display higher genetic affinities to European and Anatolian groups than Iranian hunter-gatherers do, suggesting a possible cline and geneflow into the CHG and less into Mesolithic and Neolithic Iranian groups.[2][8] The Mesolithic/Neolithic Iranian lineage and the Caucasus hunter-gatherers derive significant amounts of their ancestry from Basal Eurasian, with the remainder ancestry being closer to Ancient North Eurasians (ANE). The CHG displayed a higher ANE component than the Neolithic Iranians do, suggesting they may have stood in continuous contact with Eastern Hunter-Gatherers to their North.[2][9]

At the beginning of the Neolithic, at c. 8000 BC, they were probably distributed across western Iran and the Caucasus,[10] and people similar to northern Caucasus and Iranian plateau hunter-gatherers arrived before 6000 BC in Pakistan and north-west India.[11] A roughly equal merger between the CHG and Eastern Hunter-Gatherers in the Pontic–Caspian steppe resulted in the formation of the Western Steppe Herders (WSHs). The WSHs formed the Yamnaya culture and subsequently expanded massively throughout Europe during the Late Neolithic and Early Bronze Age circa 3000-2000 BC.[12]

Research[edit]

One of the Caucasus hunters was unearthed at Satsurblia cave in Georgia.

Jones et al. (2015) analyzed genomes from males from western Georgia, in the Caucasus, from the Late Upper Palaeolithic (13,300 years old) and the Mesolithic (9,700 years old). These two males carried Y-DNA haplogroup: J* and J2a, later refined to J1-FT34521, and J2-Y12379*, and mitochondrial haplogroups of K3 and H13c, respectively.[13] Their genomes showed that a continued mixture of the Caucasians with Middle Eastern populations took place up to 25,000 years ago, when the coldest period in the last Ice Age started.[4]

CHG ancestry was also found in an Upper Palaeolithic specimen from Satsurblia cave (dated ca. 11000 BC), and in a Mesolithic one from Kotias Klde cave, in western Georgia (dated ca. 7700 BC). The Satsurblia individual is closest to modern populations from the South Caucasus.[2]

Margaryan et al. (2017) analysing South Caucasian ancient mitochondrial DNA found a rapid increase of the population at the end of the Last Glacial Maximum, about 18,000 years ago. The same study also found continuity in descent in the maternal line for 8,000 years.[14]

According to Narasimhan et al. (2019) Iranian farmer related people arrived before 6000 BCE in Pakistan and north-west India, before the advent of farming in northern India. They suggest the possibility that this "Iranian farmer–related ancestry [...] was [also] characteristic of northern Caucasus and Iranian plateau hunter-gatherers."[11]

Vallini et al. 2022 argues that the dispersal and split patterns of West Eurasian lineages was not earlier than c. 38,000 years ago, with older Upper-Paleolithic European specimens, such as Zlaty Kun and Bacho Kiro, being unrelated to Western Hunter-gatherers.[15]

Allentoft et al. 2024 modeled the CHG lineage to have formed as sister lienage of Mesolithic/Neolithic Iranians, deriving appopximately 52% ancestry from Ancient North Eurasians and 48% ancestry from Basal Eurasians, with additional geneflow for the CHG from a Paleolithic Caucasus/Anatolian source (c. 20%) and ANE-like ancestry (c. 10%).[16]

Proto-Indo Europeans[edit]

Main genetic ancestries of Western Steppe Herders (Yamnaya pastoralists): a confluence of Eastern Hunter-Gatherers (EHG) and Caucasus Hunter-Gatherers (CHG).[17]

The ancestry of the Yamnaya people can be mostly modelled as an admixture of Eastern Hunter-Gatherers (EHGs) and a Near Eastern component related to Caucasus hunter-gatherers, Iranian Chalcolithic people, or a genetically similar population.[18][2][19][4][note 1] Each of those two populations contributed about half the Yamnaya DNA.[20][4] According to co-author Andrea Manica of the University of Cambridge:

The question of where the Yamnaya come from has been something of a mystery up to now […] we can now answer that, as we've found that their genetic make-up is a mix of Eastern European hunter-gatherers and a population from this pocket of Caucasus hunter-gatherers who weathered much of the last Ice Age in apparent isolation.[4]

According to Jones et al. (2015), Caucasus hunter-gatherers (CHG) "genomes significantly contributed to the Yamnaya steppe herders who migrated into Europe ~3,000 BCE, supporting a formative Caucasus influence on this important Early Bronze Age culture. CHG left their imprint on modern populations from the Caucasus and also Central and South Asia possibly correlating with the arrival of Indo-Aryan languages."[21]

Lazaridis et al. (2016) proposes a different people, likely from Iran, as the source for the Middle Eastern ancestry of the Yamnaya people, finding that "a population related to the people of the Iran Chalcolithic contributed ~43% of the ancestry of early Bronze Age populations of the steppe".[22][note 2] That study asserts that these Iranian Chalcolithic people were a mixture of "the Neolithic people of western Iran, the Levant, and Caucasus Hunter Gatherers".[22]

Gallego-Llorente et al. (2016) conclude that Iranian populations are not a likelier source of the 'southern' component in the Yamnaya than Caucasus hunter-gatherers.[23]

Wang et al. (2018) analysed genetic data of the North Caucasus of fossils dated between the 4th and 1st millennia BC and found correlation with modern groups of the South Caucasus, concluding that "unlike today – the Caucasus acted as a bridge rather than an insurmountable barrier to human movement".[24]

While some argue that the Pre-Proto-Indo-European language may have originated among a CHG-rich population in Western Asia,[25] others, such as David W. Anthony, suggest that the Indo-European languages were initially spoken by EHGs living in Eastern Europe.[26]

Ancient Greece and Aegean[edit]

Beyond contributing to the population of mainland Europe through Bronze Age pastoralists of the Yamnaya, CHG also appears to have arrived on its own in the Aegean without Eastern European hunter–gatherer (EHG) ancestry and provided approximately 9–32% of ancestry to the Minoans. The origin of this CHG component might have been Central Anatolia.[27]

See also[edit]

Notes[edit]

References[edit]

  1. ^ Eisenmann, S.; Bánffy, E.; van Dommelen, P.; et al. (2018). "Reconciling material cultures in archaeology with genetic data: The nomenclature of clusters emerging from archaeogenomic analysis". Scientific Reports. 8 (1): 13003. Bibcode:2018NatSR...813003E. doi:10.1038/s41598-018-31123-z. PMC 6115390. PMID 30158639.
  2. ^ a b c d e Jones et al. 2015.
  3. ^ a b Fu et al. 2016.
  4. ^ a b c d e "Europe's fourth ancestral 'tribe' uncovered". BBC. 16 November 2015.
  5. ^ Dutchen, Stephanie (May 2, 2016). "History on Ice". Harvard Medical School. Retrieved 11 May 2016.
  6. ^ "'Fourth strand' of European ancestry originated with hunter-gatherers isolated by Ice Age". University of Cambridge. 16 November 2015. By reading the DNA, the researchers were able to show that the lineage of this fourth Caucasus hunter-gatherer strand diverged from the western hunter-gatherers just after the expansion of anatomically modern humans into Europe from Africa.
  7. ^ Marchi, Nina; Winkelbach, Laura; Schulz, Ilektra; Brami, Maxime; Hofmanová, Zuzana; Blöcher, Jens; Reyna-Blanco, Carlos S.; Diekmann, Yoan; Thiéry, Alexandre; Kapopoulou, Adamandia; Link, Vivian; Piuz, Valérie; Kreutzer, Susanne; Figarska, Sylwia M.; Ganiatsou, Elissavet (May 2022). "The genomic origins of the world's first farmers". Cell. 185 (11): 1842–1859.e18. doi:10.1016/j.cell.2022.04.008. ISSN 0092-8674. PMC 9166250. PMID 35561686.
  8. ^ Gallego-Llorente, M.; Connell, S.; Jones, E. R.; Merrett, D. C.; Jeon, Y.; Eriksson, A.; Siska, V.; Gamba, C.; Meiklejohn, C.; Beyer, R.; Jeon, S.; Cho, Y. S.; Hofreiter, M.; Bhak, J.; Manica, A. (2016-08-09). "The genetics of an early Neolithic pastoralist from the Zagros, Iran". Scientific Reports. 6 (1): 31326. Bibcode:2016NatSR...631326G. doi:10.1038/srep31326. ISSN 2045-2322. PMC 4977546. PMID 27502179.
  9. ^ Lazaridis et al. 2016.
  10. ^ Anthony 2009b, p. 29.
  11. ^ a b Narasimhan et al. 2019, p. 11.
  12. ^ Jeong et al. 2019.
  13. ^ "YFull | NextGen Sequence Interpretation".
  14. ^ Margaryan, Ashot; Derenko, Miroslava; Hovhannisyan, Hrant; Malyarchuk, Boris; Heller, Rasmus; Khachatryan, Zaruhi; Avetisyan, Pavel; Badalyan, Ruben; Bobokhyan, Arsen; Melikyan, Varduhi; Sargsyan, Gagik; Piliposyan, Ashot; Simonyan, Hakob; Mkrtchyan, Ruzan; Denisova, Galina; Yepiskoposyan, Levon; Willerslev, Eske; Allentoft, Morten E. (July 2017). "Eight Millennia of Matrilineal Genetic Continuity in the South Caucasus". Current Biology. 27 (13): 2023–2028.e7. doi:10.1016/j.cub.2017.05.087. PMID 28669760.
  15. ^ Vallini et al. 2022 (2022-07-04). "Genetics and Material Culture Support Repeated Expansions into Paleolithic Eurasia from a Population Hub Out of Africa". Retrieved 2023-04-16.{{cite web}}: CS1 maint: numeric names: authors list (link)
  16. ^ Allentoft, Morten E.; Sikora, Martin; Refoyo-Martínez, Alba; Irving-Pease, Evan K.; Fischer, Anders; Barrie, William; Ingason, Andrés; Stenderup, Jesper; Sjögren, Karl-Göran; Pearson, Alice; Sousa da Mota, Bárbara; Schulz Paulsson, Bettina; Halgren, Alma; Macleod, Ruairidh; Jørkov, Marie Louise Schjellerup (January 2024). "Population genomics of post-glacial western Eurasia". Nature. 625 (7994): 301–311. doi:10.1038/s41586-023-06865-0. ISSN 1476-4687. PMC 10781627. PMID 38200295.
  17. ^ Hanel, Andrea; Carlberg, Carsten (3 July 2020). "Skin colour and vitamin D: An update". Experimental Dermatology. 29 (9): 864–875. doi:10.1111/exd.14142. PMID 32621306.
  18. ^ Haak 2015, p. 3.
  19. ^ Lazaridis et al. 2016, p. 8: "The spread of Near Eastern ancestry into the Eurasian steppe was previously inferred without access to ancient samples, by hypothesizing a population related to present-day Armenians as a source."
  20. ^ Mathieson 2015.
  21. ^ Jones et al. 2015: "Caucasus hunter-gatherers (CHG) belong to a distinct ancient clade that split from western hunter-gatherers ~45 kya, shortly after the expansion of anatomically modern humans into Europe and from the ancestors of Neolithic farmers ~25 kya, around the Last Glacial Maximum."
  22. ^ a b Lazaridis et al. 2016, p. 5.
  23. ^ Gallego-Llorente, M.; Connell, S.; Jones, E. R.; Merrett, D. C.; Jeon, Y.; Eriksson, A.; et al. (2016). "The genetics of an early Neolithic pastoralist from the Zagros, Iran". Scientific Reports. 6: 31326. Bibcode:2016NatSR...631326G. doi:10.1038/srep31326. PMC 4977546. PMID 27502179.
  24. ^ Wang et al. 2018.
  25. ^ Lazaridis, Iosif; Alpaslan-Roodenberg, Songül; Acar, Ayşe; Açıkkol, Ayşen; Agelarakis, Anagnostis; Aghikyan, Levon; Akyüz, Uğur; Andreeva, Desislava; Andrijašević, Gojko; Antonović, Dragana; Armit, Ian; Atmaca, Alper; Avetisyan, Pavel; Aytek, Ahmet İhsan; Bacvarov, Krum (2022-08-26). "The genetic history of the Southern Arc: A bridge between West Asia and Europe". Science. 377 (6609): eabm4247. doi:10.1126/science.abm4247. ISSN 0036-8075. PMC 10064553. PMID 36007055.
  26. ^ Anthony, David (2019-01-01). "Archaeology, Genetics, and Language in the Steppes: A Comment on Bomhard". Journal of Indo-European Studies.
  27. ^ Lazaridis, Iosif; Mittnik, Alissa; Patterson, Nick; Mallick, Swapan; Rohland, Nadin; Pfrengle, Saskia; Furtwängler, Anja; Peltzer, Alexander; Posth, Cosimo; Vasilakis, Andonis; McGeorge, P. J. P.; Konsolaki-Yannopoulou, Eleni; Korres, George; Martlew, Holley; Michalodimitrakis, Manolis; Özsait, Mehmet; Özsait, Nesrin; Papathanasiou, Anastasia; Richards, Michael; Roodenberg, Songül Alpaslan; Tzedakis, Yannis; Arnott, Robert; Fernandes, Daniel M.; Hughey, Jeffery R.; Lotakis, Dimitra M.; Navas, Patrick A.; Maniatis, Yannis; Stamatoyannopoulos, John A.; Stewardson, Kristin; Stockhammer, Philipp; Pinhasi, Ron; Reich, David; Krause, Johannes; Stamatoyannopoulos, George (2017). "Genetic origins of the Minoans and Mycenaeans". Nature. 548 (7666): 214–218. Bibcode:2017Natur.548..214L. doi:10.1038/nature23310. PMC 5565772. PMID 28783727.

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