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The Genetic Prehistory of the Greater Caucasus

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The Genetic Prehistory of the Greater Caucasus bioRxiv preprint doi: https://doi.org/10.1101/322347; this version posted May 16, 2018. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 1 The genetic prehistory of the Greater Caucasus 2 3 4 Chuan-Chao Wang1,2,*, Sabine Reinhold3, Alexey Kalmykov4, Antje Wissgott1, Guido 5 Brandt1, Choongwon Jeong1, Olivia Cheronet5,6,7, Matthew Ferry8,9, Eadaoin 6 Harney8,9,10, Denise Keating5,7, Swapan Mallick8,9,11, Nadin Rohland8,11, Kristin 7 Stewardson8,9, Anatoly R. Kantorovich12, Vladimir E. Maslov13, Vladimira G. 8 Petrenko13, Vladimir R. Erlikh14, Biaslan Ch. Atabiev15, Rabadan G. Magomedov16, 9 Philipp L. Kohl17, Kurt W. Alt18,19,20, Sandra L. Pichler19, Claudia Gerling19, Harald 10 Meller21, Benik Vardanyan22,23, Larisa Yeganyan23, Alexey D. Rezepkin24, Dirk 11 Mariaschk3, Natalia Berezina25, Julia Gresky26, Katharina Fuchs27, Corina Knipper28, 12 Stephan Schiffels1, Elena Balanovska29, Oleg Balanovsky29,30, Iain Mathieson31, 13 Thomas Higham32, Yakov B. Berezin25, Alexandra Buzhilova25, Viktor Trifonov33, 14 Ron Pinhasi34, Andrej B. Belinskij4, David Reich8,9,11,35, Svend Hansen3,*, Johannes 15 Krause1,35,* & Wolfgang Haak1,36,* 16 17 18 1Department of Archaeogenetics, Max-Planck Institute for the Science of Human 19 History, Kahlaische Strasse 10, D-07745 Jena, Germany 20 2Department of Anthropology and Ethnology, Xiamen University 361005 Xiamen, 21 China 22 3German Archaeological Institute, Eurasia Department, Im Dol 2-6, D-14195 Berlin, 23 Germany 24 4’Nasledie’ Cultural Heritage Unit, 355006 Stavropol, Russia 25 5Earth Institute, University College Dublin, Dublin 4, Ireland 26 6Department of Anthropology, University of Vienna, 1090 Vienna, Austria 27 7School of Archaeology, University College Dublin, Dublin 4, Ireland 28 8Department of Genetics, Harvard Medical School, Boston 02115 MA, USA 29 9Howard Hughes Medical Institute, Harvard Medical School, Boston 02115 MA, USA 30 10Department of Organismic and Evolutionary Biology, Harvard University, 31 Cambridge, MA02138, USA 32 11Broad Institute of Harvard and MIT, Cambridge 02142 MA, USA 33 12Department of Archaeology, Faculty of History, Lomonosov Moscow State 34 University, Lomonosovsky pr. 27/4, 119192, Moscow, Russia 35 13Institute of Archaeology RAS, Ul. Dm. Ulyanova 19, 117036 Moscow, Russian 36 Federation 37 14State Museum of Oriental Art, 12a Nikitskiy Boulevard, 119019 Moscow, Russian 38 Federation 39 15Ltd. Institute for Caucasus Archaeology, Ul. Katkhanova 30, 361401 Nalchik, 40 Republic Kabardino-Balkaria, Russian Federation 41 16Institute of History, Archaeology and Ethnography DNC RAS, Ul. M. Jaragskogo 42 75, 367030 Makhachkala, Republic Dagestan, Russian Federation 43 17Department of Anthropology, Wellesley College, Pendleton East 331, 106 Central 44 Street, Wellesley, MA 02481, USA 45 18Danube Private University, A-3500 Krems-Stein, Austria 46 19IPAS – Institute of Prehistory and Archaeological Science, University of Basel, CH- 47 4055 Basel, Switzerland 48 20Department of Biomedical Engineering, University of Basel, CH-4123 Allschwil, 49 Switzerland 1 bioRxiv preprint doi: https://doi.org/10.1101/322347; this version posted May 16, 2018. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 50 21State Heritage Museum, Saxony-Anhalt, D-06114 Halle/Saale, Germany 51 22Martin-Luther-Universität Halle-Wittenberg, Germany 52 23Shirak Center for Armenological Studies of National Academy of Science RA, 53 Armenia 54 24Institute for the History of Material Culture, Russian Academy of Sciences, 55 Dvortsovaya nab.,18, 191186 Saint-Petersburg, Russia 56 25Research Institute and Museum of Anthropology of Lomonosov Moscow State 57 University, Mokhovaya 11, Moscow, Russia 58 26German Archaeological Institute, Department of Natural Sciences, Im Dol 2-6, D- 59 14195 Berlin, Germany 60 27CRC 1266 "Scales of Transformation", Institut für Ur- und Frühgeschichte, 61 Christian-Albrechts-Universität, Johanna-Mestorf-Straße 2-6, 24118 Kiel, Germany 62 28Curt Engelhorn Center for Archaeometry gGmbH, 68159 Mannheim, Germany 63 29Research Centre for Medical Genetics, Moscow 115478, Russia 64 30Vavilov Institute for General Genetics, Moscow 119991, Russia 65 31 Department of Genetics, Perelman School of Medicine, University of Pennsylvania, 66 Philadelphia PA 19104, USA 67 32Oxford Radiocarbon Accelerator Unit, RLAHA, University of Oxford, OX13QY, UK 68 33Institute for the History of Material Culture, Russian Academy of Sciences, 69 Dvortsovaya nab.,18, 191186 Saint-Petersburg, Russia 70 34Department of Evolutionary Anthropology, University of Vienna, 1010 Vienna, 71 Austria 72 35Max Planck-Harvard Research Center for the Archaeoscience of the Ancient 73 Mediterranean, Cambridge, MA 02138, USA 74 36School of Biological Sciences, The University of Adelaide, Adelaide 5005, Australia 75 76 77 78 *corresponding authors: [email protected], [email protected], 79 [email protected], [email protected] 80 2 bioRxiv preprint doi: https://doi.org/10.1101/322347; this version posted May 16, 2018. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 81 Abstract 82 Archaeogenetic studies have described the formation of Eurasian ‘steppe ancestry’ as 83 a mixture of Eastern and Caucasus hunter-gatherers. However, it remains unclear 84 when and where this ancestry arose and whether it was related to a horizon of cultural 85 innovations in the 4th millennium BCE that subsequently facilitated the advance of 86 pastoral societies likely linked to the dispersal of Indo-European languages. To 87 address this, we generated genome-wide SNP data from 45 prehistoric individuals 88 along a 3000-year temporal transect in the North Caucasus. We observe a genetic 89 separation between the groups of the Caucasus and those of the adjacent steppe. The 90 Caucasus groups are genetically similar to contemporaneous populations south of it, 91 suggesting that – unlike today – the Caucasus acted as a bridge rather than an 92 insurmountable barrier to human movement. The steppe groups from Yamnaya and 93 subsequent pastoralist cultures show evidence for previously undetected farmer- 94 related ancestry from different contact zones, while Steppe Maykop individuals 95 harbour additional Upper Palaeolithic Siberian and Native American related ancestry. 96 3 bioRxiv preprint doi: https://doi.org/10.1101/322347; this version posted May 16, 2018. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 97 The 1100-kilometre long Caucasus mountain ranges extend between the Black Sea 98 and the Caspian Sea and are bound by the rivers Kuban and Terek in the north and by 99 the Kura and Araxes rivers in the south. With Mount Elbrus in Russian Kabardino- 100 Balkaria rising to a height of 5642 metres and Mount Shkhara in Georgia to 5201 101 metres, the Caucasus mountain ranges form a natural barrier between the Eurasian 102 steppes and the Near East (Fig. 1). 103 104 The rich archaeological record suggests extensive periods of human occupation since 105 the Upper Palaeolithic1, 2, 3. The density of languages and cultures in the region is 106 mirrored by faunal and floral diversity, and the Caucasus has often been described as 107 a contact zone and natural refuge with copious ecological niches. However, it also 108 serves as a bio-geographic border between the steppe and regions to the south such as 109 Anatolia and Mesopotamia rather than a corridor for human4, 5 and animal movement6, 110 7, 8. The extent to which the Caucasus has played an important role for human 111 population movements between south and north over the course of human history is 112 thus a critical question, and one that until now has been unanswered by 113 archaeogenetic studies. 114 115 A Neolithic lifestyle based on food production began in the Caucasus after 6000 116 calBCE9. In the following millennia the Caucasus region began to play an 117 increasingly important role in the economies of the growing urban centres in northern 118 Mesopotamia10 as a region rich in natural resources such as ores, pastures and 119 timber11. In the 4th millennium BCE the archaeological record attests to the presence 120 of the Maykop and Kura-Araxes cultural complexes, with the latter being found on 121 both flanks of the Caucasus mountain range, thus clearly demonstrating the 122 connection between north and south11. The Maykop culture was an important player 123 in the innovative horizon of the 4th millennium BCE in Western Eurasia. It is well 124 known for its rich burial mounds, especially at the eponymous Maykop site in today’s 125 Adygea, which reflect the rise of a new system of social organization12. The 4th 126 millennium BCE witnesses a concomitant rise in commodities and technologies such 127 as the wheel and wagon including associated technology, copper alloys, new 128 weaponry, and new breeds of domestic sheep13, 14.
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