부산대학교 도서관

Keywords ancient DNA; domestication; animal breeding; horses; mules; extinct lineages; selection; diversity; management; equestrian civilizations Highlights * Two now-extinct horse lineages lived in Iberia and Siberia some 5,000 years ago * Iberian and Siberian horses contributed limited ancestry to modern domesticates * Oriental horses have had a strong genetic influence within the last millennium * Modern breeding practices were accompanied by a significant drop in genetic diversity Summary Horse domestication revolutionized warfare and accelerated travel, trade, and the geographic expansion of languages. Here, we present the largest DNA time series for a non-human organism to date, including genome-scale data from 149 ancient animals and 129 ancient genomes ([greater than or equal to]1-fold coverage), 87 of which are new. This extensive dataset allows us to assess the modern legacy of past equestrian civilizations. We find that two extinct horse lineages existed during early domestication, one at the far western (Iberia) and the other at the far eastern range (Siberia) of Eurasia. None of these contributed significantly to modern diversity. We show that the influence of Persian-related horse lineages increased following the Islamic conquests in Europe and Asia. Multiple alleles associated with elite-racing, including at the MSTN 'speed gene,' only rose in popularity within the last millennium. Finally, the development of modern breeding impacted genetic diversity more dramatically than the previous millennia of human management. Author Affiliation: (1) Laboratoire d'Anthropobiologie Moleculaire et d'Imagerie de Synthese, CNRS UMR 5288, Universite de Toulouse, Universite Paul Sabatier, 31000 Toulouse, France (2) Lundbeck Foundation GeoGenetics Center, University of Copenhagen, 1350K Copenhagen, Denmark (3) Department of Biotechnology, Abdul Wali Khan University, Mardan, Pakistan (4) Evolutionary Ecology Group, Department of Zoology, University of Cambridge, Cambridge CB2 3EJ, UK (5) Institute for Immunity, Transplantation and Infection, Stanford University, Stanford, CA 94305, USA (6) Zoology Department, College of Science, King Saud University, Riyadh 11451, Saudi Arabia (7) Seminari d'Estudis i Recerques Prehistoriques, HAR2017-87695-P, Universitat de Barcelona, Barcelona, Spain (8) Anthropology Department, Hartwick College 1, Oneonta, NY 13820, USA (9) Shejire DNA project, 050046 Almaty, Kazakhstan (10) McDonald Institute for Archaeological Research, Department of Archaeology, University of Cambridge, Cambridge CB2 3ER, UK (11) National Museum of Mongolia, Ulaanbaatar 210646, Mongolia (12) Deutsches Archaologisches Institut (DAI), 14195 Berlin, Germany (13) Laboratorio de Paleontologia y Paleobiologia, Instituto Andaluz del Patrimonio Historico, Sevilla, Spain (14) Departamento de Prehistoria y Arqueologia, Universidad Autonoma de Madrid, Madrid, Spain (15) Department of Paleolithic, National Museum of Iran, 1136918111, Tehran, Iran (16) Centre for Ecological and Evolutionary Synthesis (CEES), Department of Biosciences, University of Oslo, Postbox 1066, Blindern, 0316 Oslo, Norway (17) Ecology Group, Department of Biology, School of Arts and Sciences, National University of Mongolia, Ulaanbaatar 14201, Mongolia (18) Institute of Animal Breeding and Genetics, Department of Biomedical Sciences, Veterinary University of Vienna, 1210 Vienna, Austria (19) Palaeogenetics Group, Institute of Organismic and Molecular Evolution (iOME), Johannes Gutenberg-University Mainz, 55099 Mainz, Germany (20) Osteological material research laboratory, Klaipeda university, Klaipeda 92294, Lithuania (21) Department of Osteology, National Museum of Iran, 1136918111, Tehran, Iran (22) Department of Archaeology, Faculty of Humanities, Tarbiat Modares University, Tehran, Iran (23) Centro de Biologia Molecular Severo Ochoa (CSIC-UAM), E-28049, Madrid, Spain (24) Integrative prahistorische und naturwissenschaftliche Archaologie (IPNA), 4055 Basel, Switzerland (25) Uniarq, Centro de Arqueologia da Universidade de Lisboa, Faculdade de Letras da Universidade de Lisboa, 1600-214 Lisboa, Portugal (26) CE3C-Centre for Ecology, Evolution and Environmental Changes, Faculdade de CiA*ncias, Universidade de Lisboa, 1749-016 Lisboa, Portugal (27) Institute for Pre- and Protohistoric Archaeology and Archaeology of the Roman Provinces, Ludwig-Maximilians-University Munich, 80539 Munchen, Germany (28) ArchaeoBioCenter, Ludwig-Maximilians-University Munich, 80539 Munchen, Germany (29) Institute of Palaeoanatomy, Domestication Research and History of Veterinary Medicine, Ludwig-Maximilians-University Munich, 80539 Munchen, Germany (30) Explico Foundation, 6900 Floro, Norway (31) Department of Archaeology, Ulaanbaatar State University, Ulaanbaatar 51, Mongolia (32) Archaezoology section, Bioarchaeology Laboratory of the Central Laboratory, University of Tehran, Tehran CP1417634934, Iran (33) Departamento de Historia, Facultad de Filosofia y Letras, Universidad de Leon, Leon, Spain (34) Departamento de Sistemas Fisicos, Quimicos y Naturales, Universidad Pablo de Olavide, 41013 Sevilla, Spain (35) Operational Direction, Earth and History of Life, Royal Belgian Institute of Natural Sciences, 1000, Brussels, Belgium (36) Faculty of Agricultural and Environmental Sciences, The Agricultural University of Iceland, Keldnaholti - Arleyni 22, 112 Reykjavik, Iceland (37) University of Potsdam, Faculty of Mathematics and Natural Sciences, Institute for Biochemistry and Biology, 14476 Potsdam, Germany (38) Institute for the History of Material Culture, Russian Academy of Sciences, St. Petersburg 191186, Russia (39) Archaeology Institute of Samarkand, Uzbekistan (40) Institute of Plant and Animal Ecology, Urals Branch of the Russian Academy of Sciences, Ekaterinburg 620144, Russia (41) Department of Historical Studies, University of Gothenburg, Gothenburg, Sweden (42) Department of History, Kyrgyz-Turkish Manas University, Bishkek, Kyrgyzstan (43) Institut de Biologia Evolutiva, (CSIC-Universitat Pompeu Fabra), PRBB, Barcelona, Catalonia 08003, Spain (44) Samara State University of Social Science and Education, Samara, Russia (45) Department of Archaeology, Faculty of Humanities, University of Tehran, Iran (46) Conservation and Evolutionary Genetics Group, Estacion Biologica de Donana (EBD-CSIC), 41092 Sevilla, Spain (47) Laboratoire Archeorient, UMR 5133, Maison de l'Orient et de la Mediterranee, 69365 Lyon Cedex 7, France (48) Departamento de Prehistoria y Arqueologia, Universidad Autonoma de Madrid, Madrid, Spain (49) Laboratory for Archaeological Research, Faculty of History and Law, Kostanay State University, Kostanay, Kazakhstan (50) Archaeological Research Collection, Tallinn University, 10130 Tallinn, Estonia (51) Department of Evolutionary Genetics, Leibniz Institute for Zoo and Wildlife Research, 10315 Berlin, Germany (52) Faculty of Life Sciences, Albrecht Daniel Thaer-Institute, Humboldt University Berlin, 10115 Berlin, Germany (53) Museu Nacional de Historia Natural e da CiA*ncia, Universidade de Lisboa, Lisboa, Portugal (54) Centro Interuniversitario de Historia das CiA*ncias e da Tecnologia (CIUHCT), Faculdade de CiA*ncias, Universidade de Lisboa, Lisboa, Portugal (55) Instituto Universitario de Lisboa (ISCTE-IUL), CIES-IUL, Lisboa, Portugal (56) Catalan Institution of Research and Advanced Studies (ICREA), 08010 Barcelona, Spain (57) CNAG-CRG, Centre for Genomic Regulation (CRG), Barcelona Institute of Science and Technology (BIST), 08028 Barcelona, Spain (58) Institut Catala de Paleontologia Miquel Crusafont, Universitat Autonoma de Barcelona, Edifici ICTA-ICP, c/ Columnes s/n, 08193, Cerdanyola del Valles, Barcelona, Spain (59) S.Toraighyrov Pavlodar State University, Joint Research Center for Archeological Studies, 637000 Pavlodar, Kazakhstan (60) University of Oxford, Faculty of History, George Street, Oxford, OX1 2RL, UK (61) Centre National de la Recherche Scientifique, Museum National d'Histoire Naturelle, Archeozoologie, Archeobotanique, Societes, Pratiques et Environnements (UMR 7209), 75005 Paris, France (62) Laboratory of Archaeozoology, Department Biologia, Universidad Autonoma de Madrid, Madrid, Spain (63) Archaeology of Social Dynamics Group (ADS), Institucio Mila i Fontanals-Consejo Superior de Investigaciones Cientificas (IMF-CSIC), 08001 Barcelona, Spain (64) Grup d'Investigacio Prehistorica, HAR2016-78277-R, Universitat de Lleida, 25003 Lleida, Spain (65) Osteoarchaeology Practice and Research Center and Department of Anatomy, Faculty of Veterinary Medicine, Istanbul University-Cerrahpasa, 34320, Avcilar, Istanbul, Turkey (66) Ivane Javakhishvili Tbilisi State University, Tbilisi 0179, Georgia (67) Universite de Bordeaux, CNRS, UMR 5199-PACEA, 33615 Pessac Cedex, France (68) Institute for Anthropological Research, Gajeva 32, 10000 Zagreb, Croatia (69) Vinkovci Municipal Museum, 32 100 Vinkovci, Croatia (70) Iranian Center for Archaeological Research (ICAR), Iranian Cultural Heritage, Handicrafts, and Tourism Organization (ICHHTO), 1136918111, Tehran, Iran (71) School of History, Classics and Archaeology, The University of Edinburgh, Edinburgh, EH8 9AG, UK (72) Central Natural Science Collections (ZNS), Martin Luther University Halle-Wittenberg, Domplatz 4, 06108 Halle (Saale), Germany (73) INRAP Guadeloupe, Centre de recherches archeologiques, UMR 7209 CNRS/MNHN, 97113 Gourbeyre, Guadeloupe (74) Department of Ecology and Evolutionary Biology and Howard Hughes Medical Institute, University of California, Santa Cruz, Santa Cruz, CA 95060, USA (75) Saeedi Institute for Advanced Studies, University of Kashan, Kashan 87317-51167, Iran (76) Department Earth System Science, University of California, Irvine, Irvine, CA 92697, USA (77) Department of Archaeology, University of Nottingham, Nottingham, NG7 2RD, UK (78) Department of Archaeology, University of Exeter, Exeter, EX4 4QE, UK (79) Kermanshah Regional Water Authority, Kermanshah 67145-1466, Iran (80) Max Planck Institute for the Science of Human History, 07745 Jena, Germany (81) Department of Anatomy, Histology and Embryology, Faculty of Veterinary Medicine, University of Zagreb, 10 000 Zagreb, Croatia (82) Department of Anthropology and Archaeology, School of Arts and Sciences, National University of Mongolia, Ulaanbaatar 14201, Mongolia (83) Saryarka Archaeological Institute of Buketov Karaganda State University, Karaganda 100074, Kazakhstan (84) Faculty of Humanities (Archaeology), University of Southampton, Avenue Campus, Highfield, Southampton SO17 1BF, UK (85) Scientific Research Institute of Archaeology and Steppe Civilizations, Al Farabi Kazakh National University, 050040 Almaty, Kazakhstan (86) deCODE Genetics, 101 Reykjavik, Iceland (87) GABI UMR1313, INRA, AgroParisTech, Universite Paris-Saclay, Jouy-en-Josas, France * Corresponding author Article History: Received 19 October 2018; Revised 14 February 2019; Accepted 27 March 2019 (miscellaneous) Published: May 2, 2019 (footnote)88 These authors contributed equally (footnote)89 Lead Contact Byline: Antoine Fages (1,2,88), Kristian Hanghoj (1,2,88), Naveed Khan (2,3,88), Charleen Gaunitz (2), Andaine Seguin-Orlando (1,2), Michela Leonardi (2,4), Christian McCrory Constantz (2,5), Cristina Gamba (2), Khaled A.S. Al-Rasheid (6), Silvia Albizuri (7), Ahmed H. Alfarhan (6), Morten Allentoft (2), Saleh Alquraishi (6), David Anthony (8), Nurbol Baimukhanov (9), James H. Barrett (10), Jamsranjav Bayarsaikhan (11), Norbert Benecke (12), Eloisa Bernaldez-Sanchez (13), Luis Berrocal-Rangel (14), Fereidoun Biglari (15), Sanne Boessenkool (16), Bazartseren Boldgiv (17), Gottfried Brem (18), Dorcas Brown (8), Joachim Burger (19), Eric Crubezy (1), Linas Daugnora (20), Hossein Davoudi (21,22), Peter de Barros Damgaard (2), Maria de los Angeles de Chorro y de Villa-Ceballos (23), Sabine Deschler-Erb (24), Cleia Detry (25), Nadine Dill (24), Maria do Mar Oom (26), Anna Dohr (27,28,29), Sturla Ellingvag (30), Diimaajav Erdenebaatar (31), Homa Fathi (21,32), Sabine Felkel (18), Carlos Fernandez-Rodriguez (33), Esteban Garcia-Vinas (34), Mietje Germonpre (35), Jose D. Granado (24), Jon H. Hallsson (36), Helmut Hemmer (19), Michael Hofreiter (37), Aleksei Kasparov (38), Mutalib Khasanov (39), Roya Khazaeli (21,32), Pavel Kosintsev (40), Kristian Kristiansen (41), Tabaldiev Kubatbek (42), Lukas Kuderna (43), Pavel Kuznetsov (44), Haeedeh Laleh (32,45), Jennifer A. Leonard (46), Johanna Lhuillier (47), Corina Liesau von Lettow-Vorbeck (48), Andrey Logvin (49), Lembi Lougas (50), Arne Ludwig (51,52), Cristina Luis (53,54,55), Ana Margarida Arruda (25), Tomas Marques-Bonet (43,56,57,58), Raquel Matoso Silva (55), Victor Merz (59), Enkhbayar Mijiddorj (31), Bryan K. Miller (60), Oleg Monchalov (44), Fatemeh A. Mohaseb (21,32,61), Arturo Morales (62), Ariadna Nieto-Espinet (63,64), Heidi Nistelberger (16), Vedat Onar (65), Albina H. Palsdottir (16,36), Vladimir Pitulko (38), Konstantin Pitskhelauri (66), Melanie Pruvost (67), Petra Rajic Sikanjic (68), Anita Rapan Papesa (69), Natalia Roslyakova (44), Alireza Sardari (70), Eberhard Sauer (71), Renate Schafberg (72), Amelie Scheu (19), Jorg Schibler (24), Angela Schlumbaum (24), Nathalie Serrand (61,73), Aitor Serres-Armero (43), Beth Shapiro (74), Shiva Sheikhi Seno (21,32,61), Irina Shevnina (49), Sonia Shidrang (75), John Southon (76), Bastiaan Star (16), Naomi Sykes (77,78), Kamal Taheri (79), William Taylor (80), Wolf-Rudiger Teegen (27,28), Tajana Trbojevic Vukicevic (81), Simon Trixl (29), Dashzeveg Tumen (82), Sainbileg Undrakhbold (17), Emma Usmanova (83), Ali Vahdati (70), Silvia Valenzuela-Lamas (63), Catarina Viegas (25), Barbara Wallner (18), Jaco Weinstock (84), Victor Zaibert (85), Benoit Clavel (61), Sebastien Lepetz (61), Marjan Mashkour (21,32,61), Agnar Helgason (86), Kari Stefansson (86), Eric Barrey (87), Eske Willerslev (2), Alan K. Outram (78), Pablo Librado (1,2), Ludovic Orlando [ludovic.orlando@univ-tlse3.fr] (1,2,89,*)