{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,9]],"date-time":"2026-04-09T19:21:24Z","timestamp":1775762484339,"version":"3.50.1"},"reference-count":55,"publisher":"Proceedings of the National Academy of Sciences","issue":"7","content-domain":{"domain":["www.pnas.org"],"crossmark-restriction":true},"short-container-title":["Proc. Natl. Acad. Sci. U.S.A."],"published-print":{"date-parts":[[2012,2,14]]},"abstract":"<jats:p>Archaeological and genetic evidence concerning the time and mode of wild horse (<jats:italic>Equus ferus<\/jats:italic>) domestication is still debated. High levels of genetic diversity in horse mtDNA have been detected when analyzing the control region; recurrent mutations, however, tend to blur the structure of the phylogenetic tree. Here, we brought the horse mtDNA phylogeny to the highest level of molecular resolution by analyzing 83 mitochondrial genomes from modern horses across Asia, Europe, the Middle East, and the Americas. Our data reveal 18 major haplogroups (A\u2013R) with radiation times that are mostly confined to the Neolithic and later periods and place the root of the phylogeny corresponding to the Ancestral Mare Mitogenome at \u223c130\u2013160 thousand years ago. All haplogroups were detected in modern horses from Asia, but F was only found in<jats:italic>E. przewalskii<\/jats:italic>\u2014the only remaining wild horse. Therefore, a wide range of matrilineal lineages from the extinct<jats:italic>E. ferus<\/jats:italic>underwent domestication in the Eurasian steppes during the Eneolithic period and were transmitted to modern<jats:italic>E. caballus<\/jats:italic>breeds. Importantly, now that the major horse haplogroups have been defined, each with diagnostic mutational motifs (in both the coding and control regions), these haplotypes could be easily used to (<jats:italic>i<\/jats:italic>) classify well-preserved ancient remains, (<jats:italic>ii<\/jats:italic>) (re)assess the haplogroup variation of modern breeds, including Thoroughbreds, and (<jats:italic>iii<\/jats:italic>) evaluate the possible role of mtDNA backgrounds in racehorse performance.<\/jats:p>","DOI":"10.1073\/pnas.1111637109","type":"journal-article","created":{"date-parts":[[2012,1,31]],"date-time":"2012-01-31T06:24:54Z","timestamp":1327991094000},"page":"2449-2454","update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":209,"title":["Mitochondrial genomes from modern horses reveal the major haplogroups that underwent domestication"],"prefix":"10.1073","volume":"109","author":[{"given":"Alessandro","family":"Achilli","sequence":"first","affiliation":[{"name":"Dipartimento di Biologia Cellulare e Ambientale, Universit\u00e0 di Perugia, 06123 Perugia, Italy;"}]},{"given":"Anna","family":"Olivieri","sequence":"additional","affiliation":[{"name":"Dipartimento di Biologia e Biotecnologie \u201cL. Spallanzani\u201d, Universit\u00e0 di Pavia, 27100 Pavia, Italy;"}]},{"given":"Pedro","family":"Soares","sequence":"additional","affiliation":[{"name":"Instituto de Patologia e Imunologia Molecular da Universidade do Porto (IPATIMUP), 4200-465 Porto, Portugal;"}]},{"given":"Hovirag","family":"Lancioni","sequence":"additional","affiliation":[{"name":"Dipartimento di Biologia Cellulare e Ambientale, Universit\u00e0 di Perugia, 06123 Perugia, Italy;"}]},{"given":"Baharak Hooshiar","family":"Kashani","sequence":"additional","affiliation":[{"name":"Dipartimento di Biologia e Biotecnologie \u201cL. Spallanzani\u201d, Universit\u00e0 di Pavia, 27100 Pavia, Italy;"}]},{"given":"Ugo A.","family":"Perego","sequence":"additional","affiliation":[{"name":"Dipartimento di Biologia e Biotecnologie \u201cL. Spallanzani\u201d, Universit\u00e0 di Pavia, 27100 Pavia, Italy;"},{"name":"Sorenson Molecular Genealogy Foundation, Salt Lake City, UT 84115;"}]},{"given":"Solomon G.","family":"Nergadze","sequence":"additional","affiliation":[{"name":"Dipartimento di Biologia e Biotecnologie \u201cL. Spallanzani\u201d, Universit\u00e0 di Pavia, 27100 Pavia, Italy;"}]},{"given":"Valeria","family":"Carossa","sequence":"additional","affiliation":[{"name":"Dipartimento di Biologia e Biotecnologie \u201cL. Spallanzani\u201d, Universit\u00e0 di Pavia, 27100 Pavia, Italy;"}]},{"given":"Marco","family":"Santagostino","sequence":"additional","affiliation":[{"name":"Dipartimento di Biologia e Biotecnologie \u201cL. Spallanzani\u201d, Universit\u00e0 di Pavia, 27100 Pavia, Italy;"}]},{"given":"Stefano","family":"Capomaccio","sequence":"additional","affiliation":[{"name":"Centro di Studio del Cavallo Sportivo, Dipartimento di Patologia, Diagnostica e Clinica Veterinaria, Universit\u00e0 di Perugia, 06123 Perugia, Italy;"}]},{"given":"Michela","family":"Felicetti","sequence":"additional","affiliation":[{"name":"Centro di Studio del Cavallo Sportivo, Dipartimento di Patologia, Diagnostica e Clinica Veterinaria, Universit\u00e0 di Perugia, 06123 Perugia, Italy;"}]},{"given":"Walid","family":"Al-Achkar","sequence":"additional","affiliation":[{"name":"Department of Molecular Biology and Biotechnology, Atomic Energy Commission, 6091 Damascus, Syria;"}]},{"given":"M. Cecilia T.","family":"Penedo","sequence":"additional","affiliation":[{"name":"Veterinary Genetics Laboratory, University of California, Davis, CA 95616;"}]},{"given":"Andrea","family":"Verini-Supplizi","sequence":"additional","affiliation":[{"name":"Centro di Studio del Cavallo Sportivo, Dipartimento di Patologia, Diagnostica e Clinica Veterinaria, Universit\u00e0 di Perugia, 06123 Perugia, Italy;"}]},{"given":"Massoud","family":"Houshmand","sequence":"additional","affiliation":[{"name":"Department of Medical Genetics, National Institute for Genetic Engineering and Biotechnology (NIGEB), 14965\/161 Tehran, Iran;"}]},{"given":"Scott R.","family":"Woodward","sequence":"additional","affiliation":[{"name":"Sorenson Molecular Genealogy Foundation, Salt Lake City, UT 84115;"}]},{"given":"Ornella","family":"Semino","sequence":"additional","affiliation":[{"name":"Dipartimento di Biologia e Biotecnologie \u201cL. Spallanzani\u201d, Universit\u00e0 di Pavia, 27100 Pavia, Italy;"}]},{"given":"Maurizio","family":"Silvestrelli","sequence":"additional","affiliation":[{"name":"Centro di Studio del Cavallo Sportivo, Dipartimento di Patologia, Diagnostica e Clinica Veterinaria, Universit\u00e0 di Perugia, 06123 Perugia, Italy;"}]},{"given":"Elena","family":"Giulotto","sequence":"additional","affiliation":[{"name":"Dipartimento di Biologia e Biotecnologie \u201cL. Spallanzani\u201d, Universit\u00e0 di Pavia, 27100 Pavia, Italy;"}]},{"given":"Lu\u00edsa","family":"Pereira","sequence":"additional","affiliation":[{"name":"Instituto de Patologia e Imunologia Molecular da Universidade do Porto (IPATIMUP), 4200-465 Porto, Portugal;"},{"name":"Faculdade de Medicina, Universidade do Porto, 4200-319 Porto, Portugal; and"}]},{"given":"Hans-J\u00fcrgen","family":"Bandelt","sequence":"additional","affiliation":[{"name":"Department of Mathematics, University of Hamburg, 20146 Hamburg, Germany"}]},{"given":"Antonio","family":"Torroni","sequence":"additional","affiliation":[{"name":"Dipartimento di Biologia e Biotecnologie \u201cL. Spallanzani\u201d, Universit\u00e0 di Pavia, 27100 Pavia, Italy;"}]}],"member":"341","published-online":{"date-parts":[[2012,1,30]]},"reference":[{"key":"e_1_3_4_1_2","doi-asserted-by":"crossref","first-page":"700","DOI":"10.1038\/nature01019","article-title":"Evolution, consequences and future of plant and animal domestication","volume":"418","author":"Diamond J","year":"2002","unstructured":"J Diamond, Evolution, consequences and future of plant and animal domestication. Nature 418, 700\u2013707 (2002).","journal-title":"Nature"},{"key":"e_1_3_4_2_2","doi-asserted-by":"crossref","first-page":"6","DOI":"10.1111\/j.1365-2052.2010.02038.x","article-title":"Genetic diversity in farm animals\u2014a review","volume":"41","author":"Groeneveld LF","year":"2010","unstructured":"LF Groeneveld, et al., Genetic diversity in farm animals\u2014a review. 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Science 324, 485 (2009).","journal-title":"Science"},{"key":"e_1_3_4_6_2","first-page":"5","volume-title":"The Domestic Horse: The Origins, Development and Management of its Behaviour","author":"Levine MA","year":"2005","unstructured":"MA Levine The Domestic Horse: The Origins, Development and Management of its Behaviour, eds DS Mills, SM McDonnell (Cambridge University Press, Cambridge, UK), pp. 5\u201322 (2005)."},{"key":"e_1_3_4_7_2","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.livsci.2008.03.002","article-title":"Domestication of the horse: Genetic relationships between domestic and wild horses","volume":"116","author":"Kavar T","year":"2008","unstructured":"T Kavar, P Dov\u010d, Domestication of the horse: Genetic relationships between domestic and wild horses. 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