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Proceedings of the National Academy of Sciences of the United States of America2012; 109(21); 8202-8206; doi: 10.1073/pnas.1111122109

Reconstructing the origin and spread of horse domestication in the Eurasian steppe.

Abstract: Despite decades of research across multiple disciplines, the early history of horse domestication remains poorly understood. On the basis of current evidence from archaeology, mitochondrial DNA, and Y-chromosomal sequencing, a number of different domestication scenarios have been proposed, ranging from the spread of domestic horses out of a restricted primary area of domestication to the domestication of numerous distinct wild horse populations. In this paper, we reconstruct both the population genetic structure of the extinct wild progenitor of domestic horses, Equus ferus, and the origin and spread of horse domestication in the Eurasian steppes by fitting a spatially explicit stepping-stone model to genotype data from >300 horses sampled across northern Eurasia. We find strong evidence for an expansion of E. ferus out of eastern Eurasia about 160 kya, likely reflecting the colonization of Eurasia by this species. Our best-fitting scenario further suggests that horse domestication originated in the western part of the Eurasian steppe and that domestic herds were repeatedly restocked with local wild horses as they spread out of this area. By showing that horse domestication was initiated in the western Eurasian steppe and that the spread of domestic herds across Eurasia involved extensive introgression from the wild, the scenario of horse domestication proposed here unites evidence from archaeology, mitochondrial DNA, and Y-chromosomal DNA.
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Publication Date: 2012-05-07 PubMed ID: 22566639PubMed Central: PMC3361400DOI: 10.1073/pnas.1111122109Google Scholar: Lookup
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Summary

This research summary has been generated with artificial intelligence and may contain errors and omissions. Refer to the original study to confirm details provided. Submit correction.

This research paper investigates the early history and development of horse domestication, proposing that horse domestication likely originated in the western part of the Eurasian steppe. The researchers used archaeological evidence, and mitochondrial and Y-chromosomal DNA from over 300 horses in northern Eurasia, to create a model of this early domestication and spread.

Research Process

  • The authors conducted a comprehensive research using archaeological findings, mitochondrial DNA, and Y-chromosomal sequencing. Their study aimed to provide a clearer understanding of how, where, and when horse domestication began in the Eurasian steppe.
  • They scrutinized various theories and scenarios that have been previously proposed about horse domestication. These ranged from hypotheses about domestic horses spreading from a single primary area to proposals about the domestication of distinct wild horse populations.
  • By compiling and analyzing genotypic data from more than 300 horses sampled across Northern Eurasia, they were able to construct a spatially explicit stepping-stone model, which allowed them to map the genetic structure across different horse populations and trace their routes of dispersal.

Findings

  • The researchers found substantial evidence pointing to a population expansion of Equus ferus, the wild precursor to domestic horses, from Eastern Eurasia around 160,000 years ago. This likely reflects the colonization of Eurasia by this species.
  • The best-fitting scenario according to their model suggests that horse domestication first started in the western part of the Eurasian steppe. As the domestic horse population spread out of this area, it appears that domestic herds were repeatedly replenished with local wild horeses.

Conclusion

  • The research concluded that horse domestication was likely to have been initiated in the western Eurasian steppe. They also found that the widespread dispersal of domestic horses across Eurasia involved extensive intermingling and breeding with local wild horse populations, otherwise known as introgression.
  • By substantiating the proposed scenario of the origins of horse domestication and its spread, the authors have managed to coherently unite evidence from multiple sources: archaeology, mitochondrial DNA, and Y-chromosomal DNA.

Cite This Article

APA
Warmuth V, Eriksson A, Bower MA, Barker G, Barrett E, Hanks BK, Li S, Lomitashvili D, Ochir-Goryaeva M, Sizonov GV, Soyonov V, Manica A. (2012). Reconstructing the origin and spread of horse domestication in the Eurasian steppe. Proc Natl Acad Sci U S A, 109(21), 8202-8206. https://doi.org/10.1073/pnas.1111122109

Publication

Proceedings of the National Academy of Sciences of the United States of America
ISSN: 1091-6490
NlmUniqueID: 7505876
Country: United States
Language: English
Volume: 109
Issue: 21
Pages: 8202-8206

Researcher Affiliations

Warmuth, Vera
  • Department of Zoology, University of Cambridge, Cambridge CB2 3EJ, United Kingdom. vmw23@cam.ac.uk
Eriksson, Anders
    Bower, Mim Ann
      Barker, Graeme
        Barrett, Elizabeth
          Hanks, Bryan Kent
            Li, Shuicheng
              Lomitashvili, David
                Ochir-Goryaeva, Maria
                  Sizonov, Grigory V
                    Soyonov, Vasiliy
                      Manica, Andrea

                        MeSH Terms

                        • Algorithms
                        • Animals
                        • Animals, Domestic / genetics
                        • Animals, Wild / genetics
                        • Archaeology
                        • Asia
                        • DNA, Mitochondrial / genetics
                        • Europe
                        • Genetics, Population
                        • Horses / genetics
                        • Microsatellite Repeats / genetics
                        • Models, Genetic
                        • Y Chromosome / genetics

                        Grant Funding

                        • BB/H005854/1 / Biotechnology and Biological Sciences Research Council

                        Conflict of Interest Statement

                        The authors declare no conflict of interest.

                        References

                        This article includes 29 references
                        1. Outram AK, Stear NA, Bendrey R, Olsen S, Kasparov A, Zaibert V, Thorpe N, Evershed RP. The earliest horse harnessing and milking.. Science 2009 Mar 6;323(5919):1332-5.
                          pubmed: 19265018doi: 10.1126/science.1168594google scholar: lookup
                        2. Olsen SL. In: Documenting Domestication: New Genetic and Archaeological Paradigms. Zeder MA, Bradley DG, Emshwiller E, Smith BD, editors. Los Angeles: Univ of California Press; 2006. pp. 245–269.
                        3. Clutton-Brock J. A Natural History of Domesticated Mammals. Cambridge, UK: Cambridge Univ Press; 1999.
                        4. Lindgren G, Backström N, Swinburne J, Hellborg L, Einarsson A, Sandberg K, Cothran G, Vilà C, Binns M, Ellegren H. Limited number of patrilines in horse domestication.. Nat Genet 2004 Apr;36(4):335-6.
                          pubmed: 15034578doi: 10.1038/ng1326google scholar: lookup
                        5. Vilà C, Leonard JA, Gotherstrom A, Marklund S, Sandberg K, Liden K, Wayne RK, Ellegren H. Widespread origins of domestic horse lineages.. Science 2001 Jan 19;291(5503):474-7.
                          pubmed: 11161199doi: 10.1126/science.291.5503.474google scholar: lookup
                        6. Jansen T, Forster P, Levine MA, Oelke H, Hurles M, Renfrew C, Weber J, Olek K. Mitochondrial DNA and the origins of the domestic horse.. Proc Natl Acad Sci U S A 2002 Aug 6;99(16):10905-10.
                          pmc: PMC125071pubmed: 12130666doi: 10.1073/pnas.152330099google scholar: lookup
                        7. Prothero DR, Schoch RM. Horns, Tusks, & Flippers: The Evolution of Hoofed Mammals. Baltimore: The John Hopkins Univ Press; 2002. pp. 213–227.
                        8. Brown D, Anthony D. Bit wear, horseback riding and the Botai site in Kazakhstan. J Archaeol Sci 1998;25:331–347.
                        9. Achilli A, Olivieri A, Soares P, Lancioni H, Hooshiar Kashani B, Perego UA, Nergadze SG, Carossa V, Santagostino M, Capomaccio S, Felicetti M, Al-Achkar W, Penedo MC, Verini-Supplizi A, Houshmand M, Woodward SR, Semino O, Silvestrelli M, Giulotto E, Pereira L, Bandelt HJ, Torroni A. Mitochondrial genomes from modern horses reveal the major haplogroups that underwent domestication.. Proc Natl Acad Sci U S A 2012 Feb 14;109(7):2449-54.
                          pmc: PMC3289334pubmed: 22308342doi: 10.1073/pnas.1111637109google scholar: lookup
                        10. Warmuth V, Eriksson A, Bower MA, Cañon J, Cothran G, Distl O, Glowatzki-Mullis ML, Hunt H, Luís C, do Mar Oom M, Yupanqui IT, Ząbek T, Manica A. European domestic horses originated in two holocene refugia.. PLoS One 2011 Mar 30;6(3):e18194.
                          pmc: PMC3068172pubmed: 21479181doi: 10.1371/journal.pone.0018194google scholar: lookup
                        11. Cieslak M, Pruvost M, Benecke N, Hofreiter M, Morales A, Reissmann M, Ludwig A. Origin and history of mitochondrial DNA lineages in domestic horses.. PLoS One 2010 Dec 20;5(12):e15311.
                          pmc: PMC3004868pubmed: 21187961doi: 10.1371/journal.pone.0015311google scholar: lookup
                        12. Luikart G, Gielly L, Excoffier L, Vigne JD, Bouvet J, Taberlet P. Multiple maternal origins and weak phylogeographic structure in domestic goats.. Proc Natl Acad Sci U S A 2001 May 8;98(10):5927-32.
                          pmc: PMC33315pubmed: 11344314doi: 10.1073/pnas.091591198google scholar: lookup
                        13. Boyd L, Houpt KA. In: Przewalski’s Horse, the History and Biology of an Endangered Species. Boyd L, Houpt KA, editors. Albany, NY: State Univ of New York Press; 1994. pp. 195–228.
                        14. Levine MA. Botai and the origins of horse domestication. J Anthropol Archaeol 1999;18:29–78.
                        15. Ling Y, Ma Y, Guan W, Cheng Y, Wang Y, Han J, Jin D, Mang L, Mahmut H. Identification of Y chromosome genetic variations in Chinese indigenous horse breeds.. J Hered 2010 Sep-Oct;101(5):639-43.
                          pubmed: 20497969doi: 10.1093/jhered/esq047google scholar: lookup
                        16. Lippold S, Knapp M, Kuznetsova T, Leonard JA, Benecke N, Ludwig A, Rasmussen M, Cooper A, Weinstock J, Willerslev E, Shapiro B, Hofreiter M. Discovery of lost diversity of paternal horse lineages using ancient DNA.. Nat Commun 2011 Aug 23;2:450.
                          pubmed: 21863017doi: 10.1038/ncomms1447google scholar: lookup
                        17. Ludwig A, Pruvost M, Reissmann M, Benecke N, Brockmann GA, Castaños P, Cieslak M, Lippold S, Llorente L, Malaspinas AS, Slatkin M, Hofreiter M. Coat color variation at the beginning of horse domestication.. Science 2009 Apr 24;324(5926):485.
                          pmc: PMC5102060pubmed: 19390039doi: 10.1126/science.1172750google scholar: lookup
                        18. Linz B, Balloux F, Moodley Y, Manica A, Liu H, Roumagnac P, Falush D, Stamer C, Prugnolle F, van der Merwe SW, Yamaoka Y, Graham DY, Perez-Trallero E, Wadstrom T, Suerbaum S, Achtman M. An African origin for the intimate association between humans and Helicobacter pylori.. Nature 2007 Feb 22;445(7130):915-918.
                          pmc: PMC1847463pubmed: 17287725doi: 10.1038/nature05562google scholar: lookup
                        19. Tanabe K, Mita T, Jombart T, Eriksson A, Horibe S, Palacpac N, Ranford-Cartwright L, Sawai H, Sakihama N, Ohmae H, Nakamura M, Ferreira MU, Escalante AA, Prugnolle F, Björkman A, Färnert A, Kaneko A, Horii T, Manica A, Kishino H, Balloux F. Plasmodium falciparum accompanied the human expansion out of Africa.. Curr Biol 2010 Jul 27;20(14):1283-9.
                          pubmed: 20656209doi: 10.1016/j.cub.2010.05.053google scholar: lookup
                        20. François O, Blum MG, Jakobsson M, Rosenberg NA. Demographic history of european populations of Arabidopsis thaliana.. PLoS Genet 2008 May 16;4(5):e1000075.
                          pmc: PMC2364639pubmed: 18483550doi: 10.1371/journal.pgen.1000075google scholar: lookup
                        21. Manica A, Amos W, Balloux F, Hanihara T. The effect of ancient population bottlenecks on human phenotypic variation.. Nature 2007 Jul 19;448(7151):346-8.
                          pmc: PMC1978547pubmed: 17637668doi: 10.1038/nature05951google scholar: lookup
                        22. Manica A, Prugnolle F, Balloux F. Geography is a better determinant of human genetic differentiation than ethnicity.. Hum Genet 2005 Dec;118(3-4):366-71.
                          pmc: PMC1805472pubmed: 16189711doi: 10.1007/s00439-005-0039-3google scholar: lookup
                        23. Ackland GJ, Signitzer M, Stratford K, Cohen MH. Cultural hitchhiking on the wave of advance of beneficial technologies.. Proc Natl Acad Sci U S A 2007 May 22;104(21):8714-9.
                          pmc: PMC1885568pubmed: 17517663doi: 10.1073/pnas.0702469104google scholar: lookup
                        24. Itan Y, Powell A, Beaumont MA, Burger J, Thomas MG. The origins of lactase persistence in Europe.. PLoS Comput Biol 2009 Aug;5(8):e1000491.
                          pmc: PMC2722739pubmed: 19714206doi: 10.1371/journal.pcbi.1000491google scholar: lookup
                        25. van Etten J, Hijmans RJ. A geospatial modelling approach integrating archaeobotany and genetics to trace the origin and dispersal of domesticated plants.. PLoS One 2010 Aug 11;5(8):e12060.
                          pmc: PMC2920323pubmed: 20711460doi: 10.1371/journal.pone.0012060google scholar: lookup
                        26. Allen M, Engström AS, Meyers S, Handt O, Saldeen T, von Haeseler A, Pääbo S, Gyllensten U. Mitochondrial DNA sequencing of shed hairs and saliva on robbery caps: sensitivity and matching probabilities.. J Forensic Sci 1998 May;43(3):453-64.
                          pubmed: 9608683
                        27. Sokolov VE, Orlov VN. The Przewalski Horse and Restoration to Its Natural Habitat in Mongolia. Rome: Food and Agriculture Organization of the United Nations, Agriculture and Consumer Protection Department; 1986.
                        28. Wegmann D, Leuenberger C, Neuenschwander S, Excoffier L. ABCtoolbox: a versatile toolkit for approximate Bayesian computations.. BMC Bioinformatics 2010 Mar 4;11:116.
                          pmc: PMC2848233pubmed: 20202215doi: 10.1186/1471-2105-11-116google scholar: lookup
                        29. Bowling AT, Eggleston-Stott ML, Byrns G, Clark RS, Dileanis S, Wictum E. Validation of microsatellite markers for routine horse parentage testing.. Anim Genet 1997 Aug;28(4):247-52.
                          pubmed: 9345720doi: 10.1111/j.1365-2052.1997.00123.xgoogle scholar: lookup

                        Citations

                        This article has been cited 66 times.
                        1. Kang Z, Shi J, Liu T, Zhang Y, Zhang Q, Liu Z, Wang J, Cheng S. Genome-wide single-nucleotide polymorphism data and mitochondrial hypervariable region 1 nucleotide sequence reveal the origin of the Akhal-Teke horse. Anim Biosci 2023 Oct;36(10):1499-1507.
                          doi: 10.5713/ab.23.0044pubmed: 37170508google scholar: lookup
                        2. Cardinali I, Giontella A, Tommasi A, Silvestrelli M, Lancioni H. Unlocking Horse Y Chromosome Diversity. Genes (Basel) 2022 Dec 2;13(12).
                          doi: 10.3390/genes13122272pubmed: 36553539google scholar: lookup
                        3. Han H, McGivney BA, Allen L, Bai D, Corduff LR, Davaakhuu G, Davaasambuu J, Dorjgotov D, Hall TJ, Hemmings AJ, Holtby AR, Jambal T, Jargalsaikhan B, Jargalsaikhan U, Kadri NK, MacHugh DE, Pausch H, Readhead C, Warburton D, Dugarjaviin M, Hill EW. Common protein-coding variants influence the racing phenotype in galloping racehorse breeds. Commun Biol 2022 Dec 13;5(1):1320.
                          doi: 10.1038/s42003-022-04206-xpubmed: 36513809google scholar: lookup
                        4. Jardat P, Calandreau L, Ferreira V, Gouyet C, Parias C, Reigner F, Lansade L. Pet-directed speech improves horses' attention toward humans. Sci Rep 2022 Mar 11;12(1):4297.
                          doi: 10.1038/s41598-022-08109-zpubmed: 35277552google scholar: lookup
                        5. Schleimer A, Richart L, Drygala F, Casabianca F, Maestrini O, Weigand H, Schwartz C, Mittelbronn M, Frantz AC. Introgressive hybridisation between domestic pigs (Sus scrofa domesticus) and endemic Corsican wild boars (S. s. meridionalis): effects of human-mediated interventions. Heredity (Edinb) 2022 Apr;128(4):279-290.
                          doi: 10.1038/s41437-022-00517-1pubmed: 35273382google scholar: lookup
                        6. Remer V, Bozlak E, Felkel S, Radovic L, Rigler D, Grilz-Seger G, Stefaniuk-Szmukier M, Bugno-Poniewierska M, Brooks S, Miller DC, Antczak DF, Sadeghi R, Cothran G, Juras R, Khanshour AM, Rieder S, Penedo MC, Waiditschka G, Kalinkova L, Kalashnikov VV, Zaitsev AM, Almarzook S, Reißmann M, Brockmann GA, Brem G, Wallner B. Y-Chromosomal Insights into Breeding History and Sire Line Genealogies of Arabian Horses. Genes (Basel) 2022 Jan 26;13(2).
                          doi: 10.3390/genes13020229pubmed: 35205275google scholar: lookup
                        7. Klecel W, Martyniuk E. From the Eurasian Steppes to the Roman Circuses: A Review of Early Development of Horse Breeding and Management. Animals (Basel) 2021 Jun 22;11(7).
                          doi: 10.3390/ani11071859pubmed: 34206575google scholar: lookup
                        8. Oleksyk TK, Wolfsberger WW, Weber AM, Shchubelka K, Oleksyk OT, Levchuk O, Patrus A, Lazar N, Castro-Marquez SO, Hasynets Y, Boldyzhar P, Neymet M, Urbanovych A, Stakhovska V, Malyar K, Chervyakova S, Podoroha O, Kovalchuk N, Rodriguez-Flores JL, Zhou W, Medley S, Battistuzzi F, Liu R, Hou Y, Chen S, Yang H, Yeager M, Dean M, Mills RE, Smolanka V. Genome diversity in Ukraine. Gigascience 2021 Jan 13;10(1).
                          doi: 10.1093/gigascience/giaa159pubmed: 33438729google scholar: lookup
                        9. Loog L. Sometimes hidden but always there: the assumptions underlying genetic inference of demographic histories. Philos Trans R Soc Lond B Biol Sci 2021 Jan 18;376(1816):20190719.
                          doi: 10.1098/rstb.2019.0719pubmed: 33250022google scholar: lookup
                        10. Cappellato A, Miletto Petrazzini ME, Bisazza A, Dadda M, Agrillo C. Susceptibility to Size Visual Illusions in a Non-Primate Mammal (Equus caballus). Animals (Basel) 2020 Sep 17;10(9).
                          doi: 10.3390/ani10091673pubmed: 32957449google scholar: lookup
                        11. Guimaraes S, Arbuckle BS, Peters J, Adcock SE, Buitenhuis H, Chazin H, Manaseryan N, Uerpmann HP, Grange T, Geigl EM. Ancient DNA shows domestic horses were introduced in the southern Caucasus and Anatolia during the Bronze Age. Sci Adv 2020 Sep;6(38).
                          doi: 10.1126/sciadv.abb0030pubmed: 32938680google scholar: lookup
                        12. Liu S, Yang Y, Pan Q, Sun Y, Ma H, Liu Y, Wang M, Zhao C, Wu C. Ancient Patrilineal Lines and Relatively High ECAY Diversity Preserved in Indigenous Horses Revealed With Novel Y-Chromosome Markers. Front Genet 2020;11:467.
                          doi: 10.3389/fgene.2020.00467pubmed: 32508879google scholar: lookup
                        13. Funk SM, Guedaoura S, Juras R, Raziq A, Landolsi F, Luís C, Martínez AM, Musa Mayaki A, Mujica F, Oom MDM, Ouragh L, Stranger YM, Vega-Pla JL, Cothran EG. Major inconsistencies of inferred population genetic structure estimated in a large set of domestic horse breeds using microsatellites. Ecol Evol 2020 May;10(10):4261-4279.
                          doi: 10.1002/ece3.6195pubmed: 32489595google scholar: lookup
                        14. Salamanca-Carreño A, Jordana J, Crosby-Granados RA, Bentez-Molano J, Parés-Casanova PM. Lineal Discrimination of Horses and Mules. A Sympatric Case from Arauca, Colombia. Animals (Basel) 2020 Apr 13;10(4).
                          doi: 10.3390/ani10040679pubmed: 32294995google scholar: lookup
                        15. Frantz LAF, Bradley DG, Larson G, Orlando L. Animal domestication in the era of ancient genomics. Nat Rev Genet 2020 Aug;21(8):449-460.
                          doi: 10.1038/s41576-020-0225-0pubmed: 32265525google scholar: lookup
                        16. Lönker NS, Fechner K, Wahed AAE. Horses as a Crucial Part of One Health. Vet Sci 2020 Feb 29;7(1).
                          doi: 10.3390/vetsci7010028pubmed: 32121327google scholar: lookup
                        17. Loog L, Thalmann O, Sinding MS, Schuenemann VJ, Perri A, Germonpré M, Bocherens H, Witt KE, Samaniego Castruita JA, Velasco MS, Lundstrøm IKC, Wales N, Sonet G, Frantz L, Schroeder H, Budd J, Jimenez EL, Fedorov S, Gasparyan B, Kandel AW, Lázničková-Galetová M, Napierala H, Uerpmann HP, Nikolskiy PA, Pavlova EY, Pitulko VV, Herzig KH, Malhi RS, Willerslev E, Hansen AJ, Dobney K, Gilbert MTP, Krause J, Larson G, Eriksson A, Manica A. Ancient DNA suggests modern wolves trace their origin to a Late Pleistocene expansion from Beringia. Mol Ecol 2020 May;29(9):1596-1610.
                          doi: 10.1111/mec.15329pubmed: 31840921google scholar: lookup
                        18. Manteca Vilanova X, De Briyne N, Beaver B, Turner PV. Horse Welfare During Equine Chorionic Gonadotropin (eCG) Production. Animals (Basel) 2019 Dec 1;9(12).
                          doi: 10.3390/ani9121053pubmed: 31805698google scholar: lookup
                        19. Ning T, Ling Y, Hu S, Ardalan A, Li J, Mitra B, Chaudhuri TK, Guan W, Zhao Q, Ma Y, Savolainen P, Zhang Y. Local origin or external input: modern horse origin in East Asia. BMC Evol Biol 2019 Nov 27;19(1):217.
                          doi: 10.1186/s12862-019-1532-ypubmed: 31775623google scholar: lookup
                        20. Raudsepp T, Finno CJ, Bellone RR, Petersen JL. Ten years of the horse reference genome: insights into equine biology, domestication and population dynamics in the post-genome era. Anim Genet 2019 Dec;50(6):569-597.
                          doi: 10.1111/age.12857pubmed: 31568563google scholar: lookup
                        21. Frantz LAF, Haile J, Lin AT, Scheu A, Geörg C, Benecke N, Alexander M, Linderholm A, Mullin VE, Daly KG, Battista VM, Price M, Gron KJ, Alexandri P, Arbogast RM, Arbuckle B, Bӑlӑşescu A, Barnett R, Bartosiewicz L, Baryshnikov G, Bonsall C, Borić D, Boroneanţ A, Bulatović J, Çakirlar C, Carretero JM, Chapman J, Church M, Crooijmans R, De Cupere B, Detry C, Dimitrijevic V, Dumitraşcu V, du Plessis L, Edwards CJ, Erek CM, Erim-Özdoğan A, Ervynck A, Fulgione D, Gligor M, Götherström A, Gourichon L, Groenen MAM, Helmer D, Hongo H, Horwitz LK, Irving-Pease EK, Lebrasseur O, Lesur J, Malone C, Manaseryan N, Marciniak A, Martlew H, Mashkour M, Matthews R, Matuzeviciute GM, Maziar S, Meijaard E, McGovern T, Megens HJ, Miller R, Mohaseb AF, Orschiedt J, Orton D, Papathanasiou A, Pearson MP, Pinhasi R, Radmanović D, Ricaut FX, Richards M, Sabin R, Sarti L, Schier W, Sheikhi S, Stephan E, Stewart JR, Stoddart S, Tagliacozzo A, Tasić N, Trantalidou K, Tresset A, Valdiosera C, van den Hurk Y, Van Poucke S, Vigne JD, Yanevich A, Zeeb-Lanz A, Triantafyllidis A, Gilbert MTP, Schibler J, Rowley-Conwy P, Zeder M, Peters J, Cucchi T, Bradley DG, Dobney K, Burger J, Evin A, Girdland-Flink L, Larson G. Ancient pigs reveal a near-complete genomic turnover following their introduction to Europe. Proc Natl Acad Sci U S A 2019 Aug 27;116(35):17231-17238.
                          doi: 10.1073/pnas.1901169116pubmed: 31405970google scholar: lookup
                        22. Pitt D, Sevane N, Nicolazzi EL, MacHugh DE, Park SDE, Colli L, Martinez R, Bruford MW, Orozco-terWengel P. Domestication of cattle: Two or three events?. Evol Appl 2019 Jan;12(1):123-136.
                          doi: 10.1111/eva.12674pubmed: 30622640google scholar: lookup
                        23. Cozzi MC, Strillacci MG, Valiati P, Rogliano E, Bagnato A, Longeri M. Genetic variability of Akhal-Teke horses bred in Italy. PeerJ 2018;6:e4889.
                          doi: 10.7717/peerj.4889pubmed: 30202639google scholar: lookup
                        24. Spence JP, Steinrücken M, Terhorst J, Song YS. Inference of population history using coalescent HMMs: review and outlook. Curr Opin Genet Dev 2018 Dec;53:70-76.
                          doi: 10.1016/j.gde.2018.07.002pubmed: 30056275google scholar: lookup
                        25. Yang L, Kong X, Yang S, Dong X, Yang J, Gou X, Zhang H. Haplotype diversity in mitochondrial DNA reveals the multiple origins of Tibetan horse. PLoS One 2018;13(7):e0201564.
                          doi: 10.1371/journal.pone.0201564pubmed: 30052677google scholar: lookup
                        26. Leonardi M, Boschin F, Giampoudakis K, Beyer RM, Krapp M, Bendrey R, Sommer R, Boscato P, Manica A, Nogues-Bravo D, Orlando L. Late Quaternary horses in Eurasia in the face of climate and vegetation change. Sci Adv 2018 Jul;4(7):eaar5589.
                          doi: 10.1126/sciadv.aar5589pubmed: 30050986google scholar: lookup
                        27. Honka J, Heino MT, Kvist L, Askeyev IV, Shaymuratova DN, Askeyev OV, Askeyev AO, Heikkinen ME, Searle JB, Aspi J. Over a Thousand Years of Evolutionary History of Domestic Geese from Russian Archaeological Sites, Analysed Using Ancient DNA. Genes (Basel) 2018 Jul 20;9(7).
                          doi: 10.3390/genes9070367pubmed: 30037043google scholar: lookup
                        28. Zhang C, Ni P, Ahmad HI, Gemingguli M, Baizilaitibei A, Gulibaheti D, Fang Y, Wang H, Asif AR, Xiao C, Chen J, Ma Y, Liu X, Du X, Zhao S. Detecting the Population Structure and Scanning for Signatures of Selection in Horses (Equus caballus) From Whole-Genome Sequencing Data. Evol Bioinform Online 2018;14:1176934318775106.
                          doi: 10.1177/1176934318775106pubmed: 29899660google scholar: lookup
                        29. Wutke S, Sandoval-Castellanos E, Benecke N, Döhle HJ, Friederich S, Gonzalez J, Hofreiter M, Lõugas L, Magnell O, Malaspinas AS, Morales-Muñiz A, Orlando L, Reissmann M, Trinks A, Ludwig A. Decline of genetic diversity in ancient domestic stallions in Europe. Sci Adv 2018 Apr;4(4):eaap9691.
                          doi: 10.1126/sciadv.aap9691pubmed: 29675468google scholar: lookup
                        30. Loog L, Mirazón Lahr M, Kovacevic M, Manica A, Eriksson A, Thomas MG. Estimating mobility using sparse data: Application to human genetic variation. Proc Natl Acad Sci U S A 2017 Nov 14;114(46):12213-12218.
                          doi: 10.1073/pnas.1703642114pubmed: 29087301google scholar: lookup
                        31. Anderson DG, Kvie KS, Davydov VN, Røed KH. Maintaining genetic integrity of coexisting wild and domestic populations: Genetic differentiation between wild and domestic Rangifer with long traditions of intentional interbreeding. Ecol Evol 2017 Sep;7(17):6790-6802.
                          doi: 10.1002/ece3.3230pubmed: 28904760google scholar: lookup
                        32. Burnik Šturm M, Ganbaatar O, Voigt CC, Kaczensky P. Sequential stable isotope analysis reveals differences in multi-year dietary history of three sympatric equid species in SW Mongolia. J Appl Ecol 2017 Aug;54(4):1110-1119.
                          doi: 10.1111/1365-2664.12825pubmed: 28717255google scholar: lookup
                        33. Elsner J, Hofreiter M, Schibler J, Schlumbaum A. Ancient mtDNA diversity reveals specific population development of wild horses in Switzerland after the Last Glacial Maximum. PLoS One 2017;12(5):e0177458.
                          doi: 10.1371/journal.pone.0177458pubmed: 28542345google scholar: lookup
                        34. Roux C, Fraïsse C, Romiguier J, Anciaux Y, Galtier N, Bierne N. Shedding Light on the Grey Zone of Speciation along a Continuum of Genomic Divergence. PLoS Biol 2016 Dec;14(12):e2000234.
                          doi: 10.1371/journal.pbio.2000234pubmed: 28027292google scholar: lookup
                        35. Wutke S, Benecke N, Sandoval-Castellanos E, Döhle HJ, Friederich S, Gonzalez J, Hallsson JH, Hofreiter M, Lõugas L, Magnell O, Morales-Muniz A, Orlando L, Pálsdóttir AH, Reissmann M, Ruttkay M, Trinks A, Ludwig A. Spotted phenotypes in horses lost attractiveness in the Middle Ages. Sci Rep 2016 Dec 7;6:38548.
                          doi: 10.1038/srep38548pubmed: 27924839google scholar: lookup
                        36. Librado P, Fages A, Gaunitz C, Leonardi M, Wagner S, Khan N, Hanghøj K, Alquraishi SA, Alfarhan AH, Al-Rasheid KA, Der Sarkissian C, Schubert M, Orlando L. The Evolutionary Origin and Genetic Makeup of Domestic Horses. Genetics 2016 Oct;204(2):423-434.
                          doi: 10.1534/genetics.116.194860pubmed: 27729493google scholar: lookup
                        37. Sánchez-Villagra MR, Geiger M, Schneider RA. The taming of the neural crest: a developmental perspective on the origins of morphological covariation in domesticated mammals. R Soc Open Sci 2016 Jun;3(6):160107.
                          doi: 10.1098/rsos.160107pubmed: 27429770google scholar: lookup
                        38. Reissmann M, Musa L, Zakizadeh S, Ludwig A. Distribution of coat-color-associated alleles in the domestic horse population and Przewalski's horse. J Appl Genet 2016 Nov;57(4):519-525.
                          doi: 10.1007/s13353-016-0352-7pubmed: 27194311google scholar: lookup
                        39. Almathen F, Charruau P, Mohandesan E, Mwacharo JM, Orozco-terWengel P, Pitt D, Abdussamad AM, Uerpmann M, Uerpmann HP, De Cupere B, Magee P, Alnaqeeb MA, Salim B, Raziq A, Dessie T, Abdelhadi OM, Banabazi MH, Al-Eknah M, Walzer C, Faye B, Hofreiter M, Peters J, Hanotte O, Burger PA. Ancient and modern DNA reveal dynamics of domestication and cross-continental dispersal of the dromedary. Proc Natl Acad Sci U S A 2016 Jun 14;113(24):6707-12.
                          doi: 10.1073/pnas.1519508113pubmed: 27162355google scholar: lookup
                        40. Cardinali I, Lancioni H, Giontella A, Capodiferro MR, Capomaccio S, Buttazzoni L, Biggio GP, Cherchi R, Albertini E, Olivieri A, Cappelli K, Achilli A, Silvestrelli M. An Overview of Ten Italian Horse Breeds through Mitochondrial DNA. PLoS One 2016;11(4):e0153004.
                          doi: 10.1371/journal.pone.0153004pubmed: 27054850google scholar: lookup
                        41. Frantz LA, Schraiber JG, Madsen O, Megens HJ, Cagan A, Bosse M, Paudel Y, Crooijmans RP, Larson G, Groenen MA. Evidence of long-term gene flow and selection during domestication from analyses of Eurasian wild and domestic pig genomes. Nat Genet 2015 Oct;47(10):1141-8.
                          doi: 10.1038/ng.3394pubmed: 26323058google scholar: lookup
                        42. Naundrup PJ, Svenning JC. A Geographic Assessment of the Global Scope for Rewilding with Wild-Living Horses (Equus ferus). PLoS One 2015;10(7):e0132359.
                          doi: 10.1371/journal.pone.0132359pubmed: 26177104google scholar: lookup
                        43. Lv FH, Peng WF, Yang J, Zhao YX, Li WR, Liu MJ, Ma YH, Zhao QJ, Yang GL, Wang F, Li JQ, Liu YG, Shen ZQ, Zhao SG, Hehua E, Gorkhali NA, Farhad Vahidi SM, Muladno M, Naqvi AN, Tabell J, Iso-Touru T, Bruford MW, Kantanen J, Han JL, Li MH. Mitogenomic Meta-Analysis Identifies Two Phases of Migration in the History of Eastern Eurasian Sheep. Mol Biol Evol 2015 Oct;32(10):2515-33.
                          doi: 10.1093/molbev/msv139pubmed: 26085518google scholar: lookup
                        44. Fort J. Demic and cultural diffusion propagated the Neolithic transition across different regions of Europe. J R Soc Interface 2015 May 6;12(106).
                          doi: 10.1098/rsif.2015.0166pubmed: 25977959google scholar: lookup
                        45. Balaresque P, Poulet N, Cussat-Blanc S, Gerard P, Quintana-Murci L, Heyer E, Jobling MA. Y-chromosome descent clusters and male differential reproductive success: young lineage expansions dominate Asian pastoral nomadic populations. Eur J Hum Genet 2015 Oct;23(10):1413-22.
                          doi: 10.1038/ejhg.2014.285pubmed: 25585703google scholar: lookup
                        46. Jun J, Cho YS, Hu H, Kim HM, Jho S, Gadhvi P, Park KM, Lim J, Paek WK, Han K, Manica A, Edwards JS, Bhak J. Whole genome sequence and analysis of the Marwari horse breed and its genetic origin. BMC Genomics 2014;15 Suppl 9(Suppl 9):S4.
                          doi: 10.1186/1471-2164-15-S9-S4pubmed: 25521865google scholar: lookup
                        47. Schubert M, Jónsson H, Chang D, Der Sarkissian C, Ermini L, Ginolhac A, Albrechtsen A, Dupanloup I, Foucal A, Petersen B, Fumagalli M, Raghavan M, Seguin-Orlando A, Korneliussen TS, Velazquez AM, Stenderup J, Hoover CA, Rubin CJ, Alfarhan AH, Alquraishi SA, Al-Rasheid KA, MacHugh DE, Kalbfleisch T, MacLeod JN, Rubin EM, Sicheritz-Ponten T, Andersson L, Hofreiter M, Marques-Bonet T, Gilbert MT, Nielsen R, Excoffier L, Willerslev E, Shapiro B, Orlando L. Prehistoric genomes reveal the genetic foundation and cost of horse domestication. Proc Natl Acad Sci U S A 2014 Dec 30;111(52):E5661-9.
                          doi: 10.1073/pnas.1416991111pubmed: 25512547google scholar: lookup
                        48. Schibler J, Elsner J, Schlumbaum A. Incorporation of aurochs into a cattle herd in Neolithic Europe: single event or breeding?. Sci Rep 2014 Jul 23;4:5798.
                          doi: 10.1038/srep05798pubmed: 25052335google scholar: lookup
                        49. Marshall FB, Dobney K, Denham T, Capriles JM. Evaluating the roles of directed breeding and gene flow in animal domestication. Proc Natl Acad Sci U S A 2014 Apr 29;111(17):6153-8.
                          doi: 10.1073/pnas.1312984110pubmed: 24753599google scholar: lookup
                        50. Gerbault P, Allaby RG, Boivin N, Rudzinski A, Grimaldi IM, Pires JC, Climer Vigueira C, Dobney K, Gremillion KJ, Barton L, Arroyo-Kalin M, Purugganan MD, Rubio de Casas R, Bollongino R, Burger J, Fuller DQ, Bradley DG, Balding DJ, Richerson PJ, Gilbert MT, Larson G, Thomas MG. Storytelling and story testing in domestication. Proc Natl Acad Sci U S A 2014 Apr 29;111(17):6159-64.
                          doi: 10.1073/pnas.1400425111pubmed: 24753572google scholar: lookup
                        51. Chuluunbat B, Charruau P, Silbermayr K, Khorloojav T, Burger PA. Genetic diversity and population structure of Mongolian domestic Bactrian camels (Camelus bactrianus). Anim Genet 2014 Aug;45(4):550-8.
                          doi: 10.1111/age.12158pubmed: 24749721google scholar: lookup
                        52. McCoy AM, Schaefer R, Petersen JL, Morrell PL, Slamka MA, Mickelson JR, Valberg SJ, McCue ME. Evidence of positive selection for a glycogen synthase (GYS1) mutation in domestic horse populations. J Hered 2014 Mar-Apr;105(2):163-72.
                          doi: 10.1093/jhered/est075pubmed: 24215078google scholar: lookup
                        53. Orlando L, Ginolhac A, Zhang G, Froese D, Albrechtsen A, Stiller M, Schubert M, Cappellini E, Petersen B, Moltke I, Johnson PL, Fumagalli M, Vilstrup JT, Raghavan M, Korneliussen T, Malaspinas AS, Vogt J, Szklarczyk D, Kelstrup CD, Vinther J, Dolocan A, Stenderup J, Velazquez AM, Cahill J, Rasmussen M, Wang X, Min J, Zazula GD, Seguin-Orlando A, Mortensen C, Magnussen K, Thompson JF, Weinstock J, Gregersen K, Røed KH, Eisenmann V, Rubin CJ, Miller DC, Antczak DF, Bertelsen MF, Brunak S, Al-Rasheid KA, Ryder O, Andersson L, Mundy J, Krogh A, Gilbert MT, Kjær K, Sicheritz-Ponten T, Jensen LJ, Olsen JV, Hofreiter M, Nielsen R, Shapiro B, Wang J, Willerslev E. Recalibrating Equus evolution using the genome sequence of an early Middle Pleistocene horse. Nature 2013 Jul 4;499(7456):74-8.
                          doi: 10.1038/nature12323pubmed: 23803765google scholar: lookup
                        54. Mwacharo JM, Nomura K, Hanada H, Han JL, Amano T, Hanotte O. Reconstructing the origin and dispersal patterns of village chickens across East Africa: insights from autosomal markers. Mol Ecol 2013 May;22(10):2683-97.
                          doi: 10.1111/mec.12294pubmed: 23611649google scholar: lookup
                        55. Wallner B, Vogl C, Shukla P, Burgstaller JP, Druml T, Brem G. Identification of genetic variation on the horse y chromosome and the tracing of male founder lineages in modern breeds. PLoS One 2013;8(4):e60015.
                          doi: 10.1371/journal.pone.0060015pubmed: 23573227google scholar: lookup
                        56. Vilstrup JT, Seguin-Orlando A, Stiller M, Ginolhac A, Raghavan M, Nielsen SC, Weinstock J, Froese D, Vasiliev SK, Ovodov ND, Clary J, Helgen KM, Fleischer RC, Cooper A, Shapiro B, Orlando L. Mitochondrial phylogenomics of modern and ancient equids. PLoS One 2013;8(2):e55950.
                          doi: 10.1371/journal.pone.0055950pubmed: 23437078google scholar: lookup
                        57. Petersen JL, Mickelson JR, Cothran EG, Andersson LS, Axelsson J, Bailey E, Bannasch D, Binns MM, Borges AS, Brama P, da Câmara Machado A, Distl O, Felicetti M, Fox-Clipsham L, Graves KT, Guérin G, Haase B, Hasegawa T, Hemmann K, Hill EW, Leeb T, Lindgren G, Lohi H, Lopes MS, McGivney BA, Mikko S, Orr N, Penedo MC, Piercy RJ, Raekallio M, Rieder S, Røed KH, Silvestrelli M, Swinburne J, Tozaki T, Vaudin M, M Wade C, McCue ME. Genetic diversity in the modern horse illustrated from genome-wide SNP data. PLoS One 2013;8(1):e54997.
                          doi: 10.1371/journal.pone.0054997pubmed: 23383025google scholar: lookup
                        58. Forster P, Hurles ME, Jansen T, Levine M, Renfrew C. Origins of the domestic horse. Proc Natl Acad Sci U S A 2012 Nov 13;109(46):E3148; author reply E3149.
                          doi: 10.1073/pnas.1210326109pubmed: 23064643google scholar: lookup
                        59. Eriksson A, Betti L, Friend AD, Lycett SJ, Singarayer JS, von Cramon-Ta뫞l N, Valdes PJ, Balloux F, Manica A. Late Pleistocene climate change and the global expansion of anatomically modern humans. Proc Natl Acad Sci U S A 2012 Oct 2;109(40):16089-94.
                          doi: 10.1073/pnas.1209494109pubmed: 22988099google scholar: lookup
                        60. Steensma MJ, Ducro BJ, Dibbits B, Doekes HP, van Schipstal JGC, Kalblfleisch T, Groenen MAM, Derks MFL. High-quality, haplotype-resolved reference genomes of the Dutch warmblood horse and Friesian horse using trio binning. BMC Genomics 2025 Sep 1;26(1):790.
                          doi: 10.1186/s12864-025-11985-0pubmed: 40890628google scholar: lookup
                        61. Di Lucrezia A, Scandurra A, Lotito D, Iervolino V, D'Aniello B, Mastellone V, Lombardi P, Pinelli C. The power of interspecific sociality: how humans provide social buffering for horses. Anim Cogn 2025 Mar 12;28(1):20.
                          doi: 10.1007/s10071-025-01942-5pubmed: 40072705google scholar: lookup
                        62. Li X, Wang Z, Zhu M, Wang B, Teng S, Yan J, Wang H, Yuan P, Cao S, Qu X, Wang Z, Zhan K, Choudhury MP, Yang X, Bao Q, He S, Liu L, Zhao P, Jiang J, Xiang H, Fang L, Tang Z, Liao Y, Yi G. Genomic Insights into Post-Domestication Expansion and Selection of Body Size in Ponies. Adv Sci (Weinh) 2025 Apr;12(16):e2413023.
                          doi: 10.1002/advs.202413023pubmed: 40009528google scholar: lookup
                        63. Arifianto D, Esfandiari A, Wibawan IWT, Amrozi A, Maharani M, Darsono D, Setiadi H, Setiyono A. Assessment of health impacts in retired antisera-producing horses: Blood biochemistry and serum amyloid A analysis. Vet World 2024 Sep;17(9):2136-2143.
                          doi: 10.14202/vetworld.2024.2136-2143pubmed: 39507779google scholar: lookup
                        64. Librado P, Tressières G, Chauvey L, Fages A, Khan N, Schiavinato S, Calvière-Tonasso L, Kusliy MA, Gaunitz C, Liu X, Wagner S, Der Sarkissian C, Seguin-Orlando A, Perdereau A, Aury JM, Southon J, Shapiro B, Bouchez O, Donnadieu C, Collin YRH, Gregersen KM, Jessen MD, Christensen K, Claudi-Hansen L, Pruvost M, Pucher E, Vulic H, Novak M, Rimpf A, Turk P, Reiter S, Brem G, Schwall C, Barrey É, Robert C, Degueurce C, Horwitz LK, Klassen L, Rasmussen U, Kveiborg J, Johannsen NN, Makowiecki D, Makarowicz P, Szeliga M, Ilchyshyn V, Rud V, Romaniszyn J, Mullin VE, Verdugo M, Bradley DG, Cardoso JL, Valente MJ, Telles Antunes M, Ameen C, Thomas R, Ludwig A, Marzullo M, Prato O, Bagnasco Gianni G, Tecchiati U, Granado J, Schlumbaum A, Deschler-Erb S, Mráz MS, Boulbes N, Gardeisen A, Mayer C, Döhle HJ, Vicze M, Kosintsev PA, Kyselý R, Peške L, O'Connor T, Ananyevskaya E, Shevnina I, Logvin A, Kovalev AA, Iderkhangai TO, Sablin MV, Dashkovskiy PK, Graphodatsky AS, Merts I, Merts V, Kasparov AK, Pitulko VV, Onar V, Öztan A, Arbuckle BS, McColl H, Renaud G, Khaskhanov R, Demidenko S, Kadieva A, Atabiev B, Sundqvist M, Lindgren G, López-Cachero FJ, Albizuri S, Trbojević Vukičević T, Rapan Papeša A, Burić M, Rajić Šikanjić P, Weinstock J, Asensio Vilaró D, Codina F, García Dalmau C, Morer de Llorens J, Pou J, de Prado G, Sanmartí J, Kallala N, Torres JR, Maraoui-Telmini B, Belarte Franco MC, Valenzuela-Lamas S, Zazzo A, Lepetz S, Duchesne S, Alexeev A, Bayarsaikhan J, Houle JL, Bayarkhuu N, Turbat T, Crubézy É, Shingiray I, Mashkour M, Berezina NY, Korobov DS, Belinskiy A, Kalmykov A, Demoule JP, Reinhold S, Hansen S, Wallner B, Roslyakova N, Kuznetsov PF, Tishkin AA, Wincker P, Kanne K, Outram A, Orlando L. Widespread horse-based mobility arose around 2200 BCE in Eurasia. Nature 2024 Jul;631(8022):819-825.
                          doi: 10.1038/s41586-024-07597-5pubmed: 38843826google scholar: lookup
                        65. Bazvand B, Rashidi A, Zandi MB, Moradi MH, Rostamzadeh J. Genome-wide analysis of population structure, effective population size and inbreeding in Iranian and exotic horses. PLoS One 2024;19(3):e0299109.
                          doi: 10.1371/journal.pone.0299109pubmed: 38442089google scholar: lookup
                        66. Orazymbetova Z, Ualiyeva D, Dossybayev K, Torekhanov A, Sydykov D, Mussayeva A, Baktybayev G. Genetic Diversity of Kazakhstani Equus caballus (Linnaeus, 1758) Horse Breeds Inferred from Microsatellite Markers. Vet Sci 2023 Sep 30;10(10).
                          doi: 10.3390/vetsci10100598pubmed: 37888550google scholar: lookup
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