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Title Bronze Age Population Dynamics and the Rise of Dairy 1 Title 2 Bronze Age population dynamics and the rise of dairy pastoralism on the eastern Eurasian steppe 3 4 Authors 5 Choongwon Jeong1,2*†, Shevan Wilkin3†, Tsend Amgalantugs4, Abigail Bouwman5, William 6 Taylor3, Richard Hagan1, Sabri Bromage6, Soninkhishig Tsolmon7, Christian Trachsel8, Jonas 7 Grossmann8, Judith Littleton9, Cheryl Makarewicz10, John Krigbaum11, Marta Burri1, Ashley 8 Scott1, Ganmaa Davaasambuu6, Joshua Wright12, Franziska Irmer3, Erdene Myagmar13, Nicole 9 Boivin3, Martine Robbeets2, Frank Rühli5, Johannes Krause1, Bruno Frohlich14,15, Jessica 10 Hendy3, Christina Warinner1,5,16* 11 12 Affiliations 13 1Department of Archaeogenetics, Max Planck Institute for the Science of Human History, Jena, 14 Germany 15 2The Eurasia3angle Project, Max Planck Institute for the Science of Human History, Jena, 16 Germany 17 3Department of Archaeology, Max Planck Institute for the Science of Human History, Jena, 18 Germany 19 4Institute of Archaeology, Mongolian Academy of Sciences, Ulanbaatar, Mongolia 20 5Institute of Evolutionary Medicine, University of Zürich, Zürich, Switzerland 21 6Department of Nutrition, Harvard T.H. Chan School of Public Health, Boston, Massachusetts, 22 USA 23 7Nutrition and Biotechnology Department, Mongolian University of Science and Technology, 24 Ulaanbaatar, Mongolia 25 8Functional Genomics Centre Zürich, University of Zürich/ETH Zürich, Zürich, Switzerland 26 9Department of Anthropology, University of Auckland, Auckland, New Zealand 27 10Institute of Prehistoric and Protohistoric Archaeology, Christian Albrechts University, Kiel, 28 Germany 29 11Department of Anthropology, University of Florida, Gainesville, Florida, USA 30 12Department of Archaeology, University of Aberdeen, Aberdeen, UK 31 13Department of Archaeology, National University of Mongolia, Ulaanbaatar, Mongolia 32 14Department of Anthropology, National Museum of Natural History, Smithsonian Institution, 33 Washington DC, USA 34 15Department of Anthropology, Dartmouth College, Hanover, New Hampshire, USA 35 16Department of Anthropology, University of Oklahoma, Norman, Oklahoma, USA 36 37 †These authors contributed equally to this work. 38 39 *Corresponding authors: 40 Christina Warinner, [email protected] 1 41 Kahlaische Strasse 10, 07745 Jena, Germany 42 43 Choongwon Jeong, [email protected] 44 Kahlaische Strasse 10, 07745 Jena, Germany 45 46 Abbreviated Title 47 Bronze Age Mongolian dairy pastoralism 48 49 Keywords 50 Paleogenomics; LC-MS/MS; Dental calculus; Beta-lactoglobulin; Alpha-S1-casein 51 52 Abstract 53 Recent paleogenomic studies have shown that migrations of Western steppe herders (WSH) 54 beginning in the Eneolithic (ca. 3300-2700 BCE) profoundly transformed the genes and cultures 55 of Europe and Central Asia. Compared to Europe, however, the eastern extent of this WSH 56 expansion is not well defined. Here we present genomic and proteomic data from 22 directly 57 dated Late Bronze Age khirigsuur burials putatively associated with early pastoralism in northern 58 Mongolia (ca. 1380-975 BCE). Genome-wide analysis reveals that they are largely descended 59 from a population represented by Early Bronze Age hunter-gatherers in the Baikal region, with 60 only a limited contribution (~7%) of WSH ancestry. At the same time, however, mass 61 spectrometry analysis of dental calculus provides direct protein evidence of bovine, sheep, and 62 goat milk consumption in 7 of 9 individuals. No individuals showed molecular evidence of 63 lactase persistence (LP), and only one individual exhibited evidence of >10% WSH ancestry, 64 despite the presence of WSH populations in the nearby Altai-Sayan region for more than a 65 millennium. Unlike the spread of Neolithic farming in Europe and the expansion of Bronze Age 66 pastoralism on the Western steppe, our results indicate that ruminant dairy pastoralism was 67 adopted on the Eastern steppe by local hunter-gatherers through a process of cultural 68 transmission and minimal genetic exchange with outside groups. 69 70 Significance statement: 71 Since the Bronze Age, pastoralism has been a dominant subsistence mode on the Western steppe, 72 but the origins of this tradition on the Eastern steppe are poorly understood. Here we investigate 73 a putative early pastoralist population in northern Mongolia and find that dairy production was 74 established on the Eastern steppe by 1300 BCE. Milk proteins preserved in dental calculus 75 indicate an early focus on Western domesticated ruminants rather than local species, but genetic 76 ancestry analysis indicates minimal admixture with Western steppe herders, suggesting that dairy 77 pastoralism was introduced through adoption by local hunter-gatherers rather than population 78 replacement. 79 2 80 \body 81 Introduction 82 83 Archaeogenetic studies provide evidence that the Eurasian Eneolithic-Bronze Age transition was 84 associated with major genetic turnovers by migrations of peoples from the Pontic-Caspian steppe 85 both in Europe and in Central Asia (1-5). The migration of these Western steppe herders (WSH), 86 with the Yamnaya horizon (ca. 3300-2700 BCE) as their earliest representative, contributed not 87 only to the European Corded Ware culture (ca. 2500-2200 BCE) but also to steppe cultures 88 located between the Caspian Sea and the Altai-Sayan mountain region, such as the Afanasievo 89 (ca. 3300-2500 BCE) and later Sintashta (2100-1800 BCE) and Andronovo (1800-1300 BCE) 90 cultures. Although burials typologically linked to the Afanasievo culture have been occasionally 91 reported in Mongolia (6), the genetic profile of Eastern steppe populations, as well as the timing 92 and nature of WSH population expansion and the rise of dairy pastoralism in Mongolia, remains 93 unclear. 94 95 The remarkable demographic success of WSH populations has been linked to mobile pastoralism 96 with dairying (7), a system that efficiently converts cellulose-rich wild grasses into protein- and 97 fat-rich dairy products. Dairy foods provide a rich source of nutrients and fresh water, and 98 function as an adaptive subsistence strategy in cold, dry steppe environments where most crop 99 cultivation is highly challenging. Dairy pastoralism became widely practiced in the Eastern 100 Eurasian steppe, the homeland from which subsequent historical nomadic dairying empires, such 101 as the Xiongnu (ca. 200 BCE to 100 CE) and the Mongols (ca. 1200-1400 CE) expanded; 102 however, it is not fully understood when, where, and how this subsistence strategy developed. At 103 Botai, in central Kazakhstan, evidence for Eneolithic dairying has been reported through the 104 presence of ruminant and equine dairy lipids in ceramic residues as early as 3500 BCE (8, 9). In 105 the Altai and Tarim basin, where WSH populations have left strong genetic footprints (1, 3, 10, 106 11), archaeological evidence supports the presence of dairy products by the early 2nd Millennium 107 BCE and later (8, 12, 13). In the Eastern steppe, however, no direct observations of dairy 108 consumption have been made for a comparable time period, despite the fact that skeletal remains 109 of domestic livestock (such as sheep, goats, cattle and horses) have been found at Mongolian 110 ritual sites and in midden contexts as early as the 14th century BCE (14-17). In the absence of 111 direct evidence for Bronze Age milk production or consumption on the Eastern steppe, it remains 112 unclear whether these animals are merely ritual in nature or signify a major shift in dietary 113 ecology towards dairy pastoralism, and whether their appearance is connected to possible WSH 114 migrations onto the Eastern steppe. 115 116 To understand the population history and context of dairy pastoralism in the eastern Eurasian 117 steppe, we applied genomic and proteomic analysis to individuals buried in Late Bronze Age 118 (LBA) khirigsuurs (burial mounds) associated with the Deer Stone-Khirigsuur Complex (DSKC) 119 in northern Mongolia (SI Appendix, Table S1, Figs. S1-S3). To date, DSKC sites contain the 3 120 clearest and most direct evidence for animal pastoralism in the Eastern steppe prior to ca. 1200 121 BCE (18). Focusing on six distinct burial clusters in Arbulag soum, Khövsgöl aimag, Mongolia 122 (Fig. 1), we produced genome-wide sequencing data targeting ~1.2M SNPs for 22 DSKC- 123 associated individuals directly dated to ca. 1380-975 calibrated BCE (SI Appendix, Table S2, 124 Fig. S4), as well as whole sequenced whole genomes for two individuals (>3x coverage). Nine of 125 the individuals in this group yielded sufficient dental calculus for proteomic analysis, and we 126 tested these deposits for the presence of milk proteins using liquid chromatography-tandem mass 127 spectrometry (LC-MS/MS). Overall, our results find that DSKC subsistence included dairying of 128 Western domesticated ruminants, but that there was minimal gene flow between analyzed DSKC 129 populations and WSH groups during the LBA. Thus, in contrast to patterns observed in Western 130 Europe, where, for example, the arrival of WSH is associated with population replacement and 131 continental-level genetic turnover (5), contact between WSH and Eastern steppe populations is 132 characterized by trans-cultural transmission of dairy pastoralism in the near absence of demic 133 diffusion. 134 135 Results 136 137 Ancient DNA Sequencing and Quality Assessment 138 We built and sequenced UDG-half (19), double-indexed Illumina libraries for genomic DNA 139 extracted from teeth or femora from DSKC-associated burials in Khövsgöl, Mongolia. Twenty
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