Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series.


Journal

Cell
ISSN: 1097-4172
Titre abrégé: Cell
Pays: United States
ID NLM: 0413066

Informations de publication

Date de publication:
30 05 2019
Historique:
received: 19 10 2018
revised: 14 02 2019
accepted: 27 03 2019
pubmed: 6 5 2019
medline: 25 3 2020
entrez: 7 5 2019
Statut: ppublish

Résumé

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 (≥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.

Identifiants

pubmed: 31056281
pii: S0092-8674(19)30384-8
doi: 10.1016/j.cell.2019.03.049
pmc: PMC6547883
pii:
doi:

Substances chimiques

DNA, Ancient 0

Types de publication

Historical Article Journal Article Research Support, N.I.H., Extramural Research Support, Non-U.S. Gov't

Langues

eng

Sous-ensembles de citation

IM

Pagination

1419-1435.e31

Subventions

Organisme : European Research Council
ID : 681605
Pays : International
Organisme : NIMH NIH HHS
ID : U01 MH106874
Pays : United States
Organisme : Austrian Science Fund FWF
ID : W 1225
Pays : Austria
Organisme : Howard Hughes Medical Institute
Pays : United States

Commentaires et corrections

Type : CommentIn
Type : CommentIn

Informations de copyright

Copyright © 2019 The Author(s). Published by Elsevier Inc. All rights reserved.

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Auteurs

Antoine Fages (A)

Laboratoire d'Anthropobiologie Moléculaire et d'Imagerie de Synthèse, CNRS UMR 5288, Université de Toulouse, Université Paul Sabatier, 31000 Toulouse, France; Lundbeck Foundation GeoGenetics Center, University of Copenhagen, 1350K Copenhagen, Denmark.

Kristian Hanghøj (K)

Laboratoire d'Anthropobiologie Moléculaire et d'Imagerie de Synthèse, CNRS UMR 5288, Université de Toulouse, Université Paul Sabatier, 31000 Toulouse, France; Lundbeck Foundation GeoGenetics Center, University of Copenhagen, 1350K Copenhagen, Denmark.

Naveed Khan (N)

Lundbeck Foundation GeoGenetics Center, University of Copenhagen, 1350K Copenhagen, Denmark; Department of Biotechnology, Abdul Wali Khan University, Mardan, Pakistan.

Charleen Gaunitz (C)

Lundbeck Foundation GeoGenetics Center, University of Copenhagen, 1350K Copenhagen, Denmark.

Andaine Seguin-Orlando (A)

Laboratoire d'Anthropobiologie Moléculaire et d'Imagerie de Synthèse, CNRS UMR 5288, Université de Toulouse, Université Paul Sabatier, 31000 Toulouse, France; Lundbeck Foundation GeoGenetics Center, University of Copenhagen, 1350K Copenhagen, Denmark.

Michela Leonardi (M)

Lundbeck Foundation GeoGenetics Center, University of Copenhagen, 1350K Copenhagen, Denmark; Evolutionary Ecology Group, Department of Zoology, University of Cambridge, Cambridge CB2 3EJ, UK.

Christian McCrory Constantz (C)

Lundbeck Foundation GeoGenetics Center, University of Copenhagen, 1350K Copenhagen, Denmark; Institute for Immunity, Transplantation and Infection, Stanford University, Stanford, CA 94305, USA.

Cristina Gamba (C)

Lundbeck Foundation GeoGenetics Center, University of Copenhagen, 1350K Copenhagen, Denmark.

Khaled A S Al-Rasheid (KAS)

Zoology Department, College of Science, King Saud University, Riyadh 11451, Saudi Arabia.

Silvia Albizuri (S)

Seminari d'Estudis i Recerques Prehistoriques, HAR2017-87695-P, Universitat de Barcelona, Barcelona, Spain.

Ahmed H Alfarhan (AH)

Zoology Department, College of Science, King Saud University, Riyadh 11451, Saudi Arabia.

Morten Allentoft (M)

Lundbeck Foundation GeoGenetics Center, University of Copenhagen, 1350K Copenhagen, Denmark.

Saleh Alquraishi (S)

Zoology Department, College of Science, King Saud University, Riyadh 11451, Saudi Arabia.

David Anthony (D)

Anthropology Department, Hartwick College 1, Oneonta, NY 13820, USA.

Nurbol Baimukhanov (N)

Shejire DNA project, 050046 Almaty, Kazakhstan.

James H Barrett (JH)

McDonald Institute for Archaeological Research, Department of Archaeology, University of Cambridge, Cambridge CB2 3ER, UK.

Jamsranjav Bayarsaikhan (J)

National Museum of Mongolia, Ulaanbaatar 210646, Mongolia.

Norbert Benecke (N)

Deutsches Archäologisches Institut (DAI), 14195 Berlin, Germany.

Eloísa Bernáldez-Sánchez (E)

Laboratorio de Paleontologia y Paleobiologia, Instituto Andaluz del Patrimonio Historico, Sevilla, Spain.

Luis Berrocal-Rangel (L)

Departamento de Prehistoria y Arqueología, Universidad Autónoma de Madrid, Madrid, Spain.

Fereidoun Biglari (F)

Department of Paleolithic, National Museum of Iran, 1136918111, Tehran, Iran.

Sanne Boessenkool (S)

Centre for Ecological and Evolutionary Synthesis (CEES), Department of Biosciences, University of Oslo, Postbox 1066, Blindern, 0316 Oslo, Norway.

Bazartseren Boldgiv (B)

Ecology Group, Department of Biology, School of Arts and Sciences, National University of Mongolia, Ulaanbaatar 14201, Mongolia.

Gottfried Brem (G)

Institute of Animal Breeding and Genetics, Department of Biomedical Sciences, Veterinary University of Vienna, 1210 Vienna, Austria.

Dorcas Brown (D)

Anthropology Department, Hartwick College 1, Oneonta, NY 13820, USA.

Joachim Burger (J)

Palaeogenetics Group, Institute of Organismic and Molecular Evolution (iOME), Johannes Gutenberg-University Mainz, 55099 Mainz, Germany.

Eric Crubézy (E)

Laboratoire d'Anthropobiologie Moléculaire et d'Imagerie de Synthèse, CNRS UMR 5288, Université de Toulouse, Université Paul Sabatier, 31000 Toulouse, France.

Linas Daugnora (L)

Osteological material research laboratory, Klaipėda university, Klaipėda 92294, Lithuania.

Hossein Davoudi (H)

Department of Osteology, National Museum of Iran, 1136918111, Tehran, Iran; Department of Archaeology, Faculty of Humanities, Tarbiat Modares University, Tehran, Iran.

Peter de Barros Damgaard (P)

Lundbeck Foundation GeoGenetics Center, University of Copenhagen, 1350K Copenhagen, Denmark.

María de Los Ángeles de Chorro Y de Villa-Ceballos (M)

Centro de Biología Molecular Severo Ochoa (CSIC-UAM), E-28049, Madrid, Spain.

Sabine Deschler-Erb (S)

Integrative prähistorische und naturwissenschaftliche Archäologie (IPNA), 4055 Basel, Switzerland.

Cleia Detry (C)

Uniarq, Centro de Arqueologia da Universidade de Lisboa, Faculdade de Letras da Universidade de Lisboa, 1600-214 Lisboa, Portugal.

Nadine Dill (N)

Integrative prähistorische und naturwissenschaftliche Archäologie (IPNA), 4055 Basel, Switzerland.

Maria do Mar Oom (M)

CE3C-Centre for Ecology, Evolution and Environmental Changes, Faculdade de Ciências, Universidade de Lisboa, 1749-016 Lisboa, Portugal.

Anna Dohr (A)

Institute for Pre- and Protohistoric Archaeology and Archaeology of the Roman Provinces, Ludwig-Maximilians-University Munich, 80539 München, Germany; ArchaeoBioCenter, Ludwig-Maximilians-University Munich, 80539 München, Germany; Institute of Palaeoanatomy, Domestication Research and History of Veterinary Medicine, Ludwig-Maximilians-University Munich, 80539 München, Germany.

Sturla Ellingvåg (S)

Explico Foundation, 6900 Florø, Norway.

Diimaajav Erdenebaatar (D)

Department of Archaeology, Ulaanbaatar State University, Ulaanbaatar 51, Mongolia.

Homa Fathi (H)

Department of Osteology, National Museum of Iran, 1136918111, Tehran, Iran; Archaezoology section, Bioarchaeology Laboratory of the Central Laboratory, University of Tehran, Tehran CP1417634934, Iran.

Sabine Felkel (S)

Institute of Animal Breeding and Genetics, Department of Biomedical Sciences, Veterinary University of Vienna, 1210 Vienna, Austria.

Carlos Fernández-Rodríguez (C)

Departamento de Historia, Facultad de Filosofía y Letras, Universidad de León, León, Spain.

Esteban García-Viñas (E)

Departamento de Sistemas Físicos, Químicos y Naturales, Universidad Pablo de Olavide, 41013 Sevilla, Spain.

Mietje Germonpré (M)

Operational Direction, Earth and History of Life, Royal Belgian Institute of Natural Sciences, 1000, Brussels, Belgium.

José D Granado (JD)

Integrative prähistorische und naturwissenschaftliche Archäologie (IPNA), 4055 Basel, Switzerland.

Jón H Hallsson (JH)

Faculty of Agricultural and Environmental Sciences, The Agricultural University of Iceland, Keldnaholti - Árleyni 22, 112 Reykjavík, Iceland.

Helmut Hemmer (H)

Palaeogenetics Group, Institute of Organismic and Molecular Evolution (iOME), Johannes Gutenberg-University Mainz, 55099 Mainz, Germany.

Michael Hofreiter (M)

University of Potsdam, Faculty of Mathematics and Natural Sciences, Institute for Biochemistry and Biology, 14476 Potsdam, Germany.

Aleksei Kasparov (A)

Institute for the History of Material Culture, Russian Academy of Sciences, St. Petersburg 191186, Russia.

Mutalib Khasanov (M)

Archaeology Institute of Samarkand, Uzbekistan.

Roya Khazaeli (R)

Department of Osteology, National Museum of Iran, 1136918111, Tehran, Iran; Archaezoology section, Bioarchaeology Laboratory of the Central Laboratory, University of Tehran, Tehran CP1417634934, Iran.

Pavel Kosintsev (P)

Institute of Plant and Animal Ecology, Urals Branch of the Russian Academy of Sciences, Ekaterinburg 620144, Russia.

Kristian Kristiansen (K)

Department of Historical Studies, University of Gothenburg, Gothenburg, Sweden.

Tabaldiev Kubatbek (T)

Department of History, Kyrgyz-Turkish Manas University, Bishkek, Kyrgyzstan.

Lukas Kuderna (L)

Institut de Biologia Evolutiva, (CSIC-Universitat Pompeu Fabra), PRBB, Barcelona, Catalonia 08003, Spain.

Pavel Kuznetsov (P)

Samara State University of Social Science and Education, Samara, Russia.

Haeedeh Laleh (H)

Archaezoology section, Bioarchaeology Laboratory of the Central Laboratory, University of Tehran, Tehran CP1417634934, Iran; Department of Archaeology, Faculty of Humanities, University of Tehran, Iran.

Jennifer A Leonard (JA)

Conservation and Evolutionary Genetics Group, Estación Biológica de Doñana (EBD-CSIC), 41092 Sevilla, Spain.

Johanna Lhuillier (J)

Laboratoire Archéorient, UMR 5133, Maison de l'Orient et de la Méditerranée, 69365 Lyon Cedex 7, France.

Corina Liesau von Lettow-Vorbeck (C)

Departamento de Prehistoria y Arqueología, Universidad Autónoma de Madrid, Madrid, Spain.

Andrey Logvin (A)

Laboratory for Archaeological Research, Faculty of History and Law, Kostanay State University, Kostanay, Kazakhstan.

Lembi Lõugas (L)

Archaeological Research Collection, Tallinn University, 10130 Tallinn, Estonia.

Arne Ludwig (A)

Department of Evolutionary Genetics, Leibniz Institute for Zoo and Wildlife Research, 10315 Berlin, Germany; Faculty of Life Sciences, Albrecht Daniel Thaer-Institute, Humboldt University Berlin, 10115 Berlin, Germany.

Cristina Luis (C)

Museu Nacional de História Natural e da Ciência, Universidade de Lisboa, Lisboa, Portugal; Centro Interuniversitário de História das Ciências e da Tecnologia (CIUHCT), Faculdade de Ciências, Universidade de Lisboa, Lisboa, Portugal; Instituto Universitário de Lisboa (ISCTE-IUL), CIES-IUL, Lisboa, Portugal.

Ana Margarida Arruda (AM)

Uniarq, Centro de Arqueologia da Universidade de Lisboa, Faculdade de Letras da Universidade de Lisboa, 1600-214 Lisboa, Portugal.

Tomas Marques-Bonet (T)

Institut de Biologia Evolutiva, (CSIC-Universitat Pompeu Fabra), PRBB, Barcelona, Catalonia 08003, Spain; Catalan Institution of Research and Advanced Studies (ICREA), 08010 Barcelona, Spain; CNAG-CRG, Centre for Genomic Regulation (CRG), Barcelona Institute of Science and Technology (BIST), 08028 Barcelona, Spain; Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, c/ Columnes s/n, 08193, Cerdanyola del Vallès, Barcelona, Spain.

Raquel Matoso Silva (R)

Instituto Universitário de Lisboa (ISCTE-IUL), CIES-IUL, Lisboa, Portugal.

Victor Merz (V)

S.Toraighyrov Pavlodar State University, Joint Research Center for Archeological Studies, 637000 Pavlodar, Kazakhstan.

Enkhbayar Mijiddorj (E)

Department of Archaeology, Ulaanbaatar State University, Ulaanbaatar 51, Mongolia.

Bryan K Miller (BK)

University of Oxford, Faculty of History, George Street, Oxford, OX1 2RL, UK.

Oleg Monchalov (O)

Samara State University of Social Science and Education, Samara, Russia.

Fatemeh A Mohaseb (FA)

Department of Osteology, National Museum of Iran, 1136918111, Tehran, Iran; Archaezoology section, Bioarchaeology Laboratory of the Central Laboratory, University of Tehran, Tehran CP1417634934, Iran; Centre National de la Recherche Scientifique, Muséum National d'Histoire Naturelle, Archéozoologie, Archéobotanique, Sociétés, Pratiques et Environnements (UMR 7209), 75005 Paris, France.

Arturo Morales (A)

Laboratory of Archaeozoology, Department Biología, Universidad Autónoma de Madrid, Madrid, Spain.

Ariadna Nieto-Espinet (A)

Archaeology of Social Dynamics Group (ADS), Institució Milà i Fontanals-Consejo Superior de Investigaciones Científicas (IMF-CSIC), 08001 Barcelona, Spain; Grup d'Investigació Prehistòrica, HAR2016-78277-R, Universitat de Lleida, 25003 Lleida, Spain.

Heidi Nistelberger (H)

Centre for Ecological and Evolutionary Synthesis (CEES), Department of Biosciences, University of Oslo, Postbox 1066, Blindern, 0316 Oslo, Norway.

Vedat Onar (V)

Osteoarchaeology Practice and Research Center and Department of Anatomy, Faculty of Veterinary Medicine, Istanbul University-Cerrahpaşa, 34320, Avcılar, Istanbul, Turkey.

Albína H Pálsdóttir (AH)

Centre for Ecological and Evolutionary Synthesis (CEES), Department of Biosciences, University of Oslo, Postbox 1066, Blindern, 0316 Oslo, Norway; Faculty of Agricultural and Environmental Sciences, The Agricultural University of Iceland, Keldnaholti - Árleyni 22, 112 Reykjavík, Iceland.

Vladimir Pitulko (V)

Institute for the History of Material Culture, Russian Academy of Sciences, St. Petersburg 191186, Russia.

Konstantin Pitskhelauri (K)

Ivane Javakhishvili Tbilisi State University, Tbilisi 0179, Georgia.

Mélanie Pruvost (M)

Université de Bordeaux, CNRS, UMR 5199-PACEA, 33615 Pessac Cedex, France.

Petra Rajic Sikanjic (P)

Institute for Anthropological Research, Gajeva 32, 10000 Zagreb, Croatia.

Anita Rapan Papeša (A)

Vinkovci Municipal Museum, 32 100 Vinkovci, Croatia.

Natalia Roslyakova (N)

Samara State University of Social Science and Education, Samara, Russia.

Alireza Sardari (A)

Iranian Center for Archaeological Research (ICAR), Iranian Cultural Heritage, Handicrafts, and Tourism Organization (ICHHTO), 1136918111, Tehran, Iran.

Eberhard Sauer (E)

School of History, Classics and Archaeology, The University of Edinburgh, Edinburgh, EH8 9AG, UK.

Renate Schafberg (R)

Central Natural Science Collections (ZNS), Martin Luther University Halle-Wittenberg, Domplatz 4, 06108 Halle (Saale), Germany.

Amelie Scheu (A)

Palaeogenetics Group, Institute of Organismic and Molecular Evolution (iOME), Johannes Gutenberg-University Mainz, 55099 Mainz, Germany.

Jörg Schibler (J)

Integrative prähistorische und naturwissenschaftliche Archäologie (IPNA), 4055 Basel, Switzerland.

Angela Schlumbaum (A)

Integrative prähistorische und naturwissenschaftliche Archäologie (IPNA), 4055 Basel, Switzerland.

Nathalie Serrand (N)

Centre National de la Recherche Scientifique, Muséum National d'Histoire Naturelle, Archéozoologie, Archéobotanique, Sociétés, Pratiques et Environnements (UMR 7209), 75005 Paris, France; INRAP Guadeloupe, Centre de recherches archéologiques, UMR 7209 CNRS/MNHN, 97113 Gourbeyre, Guadeloupe.

Aitor Serres-Armero (A)

Institut de Biologia Evolutiva, (CSIC-Universitat Pompeu Fabra), PRBB, Barcelona, Catalonia 08003, Spain.

Beth Shapiro (B)

Department of Ecology and Evolutionary Biology and Howard Hughes Medical Institute, University of California, Santa Cruz, Santa Cruz, CA 95060, USA.

Shiva Sheikhi Seno (S)

Department of Osteology, National Museum of Iran, 1136918111, Tehran, Iran; Archaezoology section, Bioarchaeology Laboratory of the Central Laboratory, University of Tehran, Tehran CP1417634934, Iran; Centre National de la Recherche Scientifique, Muséum National d'Histoire Naturelle, Archéozoologie, Archéobotanique, Sociétés, Pratiques et Environnements (UMR 7209), 75005 Paris, France.

Irina Shevnina (I)

Laboratory for Archaeological Research, Faculty of History and Law, Kostanay State University, Kostanay, Kazakhstan.

Sonia Shidrang (S)

Saeedi Institute for Advanced Studies, University of Kashan, Kashan 87317-51167, Iran.

John Southon (J)

Department Earth System Science, University of California, Irvine, Irvine, CA 92697, USA.

Bastiaan Star (B)

Centre for Ecological and Evolutionary Synthesis (CEES), Department of Biosciences, University of Oslo, Postbox 1066, Blindern, 0316 Oslo, Norway.

Naomi Sykes (N)

Department of Archaeology, University of Nottingham, Nottingham, NG7 2RD, UK; Department of Archaeology, University of Exeter, Exeter, EX4 4QE, UK.

Kamal Taheri (K)

Kermanshah Regional Water Authority, Kermanshah 67145-1466, Iran.

William Taylor (W)

Max Planck Institute for the Science of Human History, 07745 Jena, Germany.

Wolf-Rüdiger Teegen (WR)

Institute for Pre- and Protohistoric Archaeology and Archaeology of the Roman Provinces, Ludwig-Maximilians-University Munich, 80539 München, Germany; ArchaeoBioCenter, Ludwig-Maximilians-University Munich, 80539 München, Germany.

Tajana Trbojević Vukičević (T)

Department of Anatomy, Histology and Embryology, Faculty of Veterinary Medicine, University of Zagreb, 10 000 Zagreb, Croatia.

Simon Trixl (S)

Institute of Palaeoanatomy, Domestication Research and History of Veterinary Medicine, Ludwig-Maximilians-University Munich, 80539 München, Germany.

Dashzeveg Tumen (D)

Department of Anthropology and Archaeology, School of Arts and Sciences, National University of Mongolia, Ulaanbaatar 14201, Mongolia.

Sainbileg Undrakhbold (S)

Ecology Group, Department of Biology, School of Arts and Sciences, National University of Mongolia, Ulaanbaatar 14201, Mongolia.

Emma Usmanova (E)

Saryarka Archaeological Institute of Buketov Karaganda State University, Karaganda 100074, Kazakhstan.

Ali Vahdati (A)

Iranian Center for Archaeological Research (ICAR), Iranian Cultural Heritage, Handicrafts, and Tourism Organization (ICHHTO), 1136918111, Tehran, Iran.

Silvia Valenzuela-Lamas (S)

Archaeology of Social Dynamics Group (ADS), Institució Milà i Fontanals-Consejo Superior de Investigaciones Científicas (IMF-CSIC), 08001 Barcelona, Spain.

Catarina Viegas (C)

Uniarq, Centro de Arqueologia da Universidade de Lisboa, Faculdade de Letras da Universidade de Lisboa, 1600-214 Lisboa, Portugal.

Barbara Wallner (B)

Institute of Animal Breeding and Genetics, Department of Biomedical Sciences, Veterinary University of Vienna, 1210 Vienna, Austria.

Jaco Weinstock (J)

Faculty of Humanities (Archaeology), University of Southampton, Avenue Campus, Highfield, Southampton SO17 1BF, UK.

Victor Zaibert (V)

Scientific Research Institute of Archaeology and Steppe Civilizations, Al Farabi Kazakh National University, 050040 Almaty, Kazakhstan.

Benoit Clavel (B)

Centre National de la Recherche Scientifique, Muséum National d'Histoire Naturelle, Archéozoologie, Archéobotanique, Sociétés, Pratiques et Environnements (UMR 7209), 75005 Paris, France.

Sébastien Lepetz (S)

Centre National de la Recherche Scientifique, Muséum National d'Histoire Naturelle, Archéozoologie, Archéobotanique, Sociétés, Pratiques et Environnements (UMR 7209), 75005 Paris, France.

Marjan Mashkour (M)

Department of Osteology, National Museum of Iran, 1136918111, Tehran, Iran; Archaezoology section, Bioarchaeology Laboratory of the Central Laboratory, University of Tehran, Tehran CP1417634934, Iran; Centre National de la Recherche Scientifique, Muséum National d'Histoire Naturelle, Archéozoologie, Archéobotanique, Sociétés, Pratiques et Environnements (UMR 7209), 75005 Paris, France.

Agnar Helgason (A)

deCODE Genetics, 101 Reykjavik, Iceland.

Kári Stefánsson (K)

deCODE Genetics, 101 Reykjavik, Iceland.

Eric Barrey (E)

GABI UMR1313, INRA, AgroParisTech, Université Paris-Saclay, Jouy-en-Josas, France.

Eske Willerslev (E)

Lundbeck Foundation GeoGenetics Center, University of Copenhagen, 1350K Copenhagen, Denmark.

Alan K Outram (AK)

Department of Archaeology, University of Exeter, Exeter, EX4 4QE, UK.

Pablo Librado (P)

Laboratoire d'Anthropobiologie Moléculaire et d'Imagerie de Synthèse, CNRS UMR 5288, Université de Toulouse, Université Paul Sabatier, 31000 Toulouse, France; Lundbeck Foundation GeoGenetics Center, University of Copenhagen, 1350K Copenhagen, Denmark.

Ludovic Orlando (L)

Laboratoire d'Anthropobiologie Moléculaire et d'Imagerie de Synthèse, CNRS UMR 5288, Université de Toulouse, Université Paul Sabatier, 31000 Toulouse, France; Lundbeck Foundation GeoGenetics Center, University of Copenhagen, 1350K Copenhagen, Denmark. Electronic address: ludovic.orlando@univ-tlse3.fr.

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