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Animals : an open access journal from MDPI2021; 11(7); 1859; doi: 10.3390/ani11071859

From the Eurasian Steppes to the Roman Circuses: A Review of Early Development of Horse Breeding and Management.

Abstract: The domestication of the horse began about 5500 years ago in the Eurasian steppes. In the following millennia horses spread across the ancient world, and their role in transportation and warfare affected every ancient culture. Ownership of horses became an indicator of wealth and social status. The importance of horses led to a growing interest in their breeding and management. Many phenotypic traits, such as height, behavior, and speed potential, have been proven to be a subject of selection; however, the details of ancient breeding practices remain mostly unknown. From the fourth millennium BP, through the Iron Age, many literature sources thoroughly describe horse training systems, as well as various aspects of husbandry, many of which are still in use today. The striking resemblance of ancient and modern equine practices leaves us wondering how much was accomplished through four thousand years of horse breeding.
Publication Date: 2021-06-22 PubMed ID: 34206575PubMed Central: PMC8300240DOI: 10.3390/ani11071859Google Scholar: Lookup
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  • Journal Article
  • Review

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.

The research article discusses the history of horse domestication, describing how it began about 5500 years ago in the Eurasian steppes, and how it spread across the ancient world, affecting every ancient culture through its significance in transportation and warfare. The article details the evolution of horse breeding and management through the years, highlighting the parallels between ancient practices and current ones.

Context and Significance

  • This study hinges on understanding the deep-rooted historical context of horse domestication and breeding, highlighting its beginning around 5500 years ago in the Eurasian steppes. This is significant as horses became instrumental in transportation and warfare, influencing all ancient cultures.
  • The ownership of horses was a marker of wealth and social status, further emphasizing the societal importance of horses during this time.

Horse Breeding and Management

  • The research elucidates the increasing interest in horse breeding and management, owing mainly to their importance in society. It also explores various phenotypic traits such as height, behavior, and speed potential that became subjects of selection.
  • However, the full details of ancient breeding practices remain largely unknown. This represents one of the challenges or gaps in understanding the full historical depth and evolution of horse breeding and management.

Training and Husbandry Practices

  • The article investigates literature sources from the fourth millennium BP through the Iron Age that offers detailed descriptions of both the training systems for horses and various aspects of husbandry.
  • Interestingly, many of these ancient practices are still in use in modern times, indicating a degree of continuity and effectiveness in these practices over the centuries.

Comparison of Ancient and Modern Practices

  • The research underscores a prominent resemblance between ancient horse breeding and management practices and those prevalent today. It brings up a question about the advancements in horse breeding over the past four thousand years, given these similarities.
  • This comparison deepens our understanding of the long-running tradition of horse breeding, management, and training over time and prompts further research on the development and changes in these practices over the centuries.

Cite This Article

APA
Klecel W, Martyniuk E. (2021). From the Eurasian Steppes to the Roman Circuses: A Review of Early Development of Horse Breeding and Management. Animals (Basel), 11(7), 1859. https://doi.org/10.3390/ani11071859

Publication

ISSN: 2076-2615
NlmUniqueID: 101635614
Country: Switzerland
Language: English
Volume: 11
Issue: 7
PII: 1859

Researcher Affiliations

Klecel, Weronika
  • Department of Animal Genetics and Conservation, Institute of Animal Sciences, Warsaw University of Life Sciences, 02-786 Warsaw, Poland.
Martyniuk, Elżbieta
  • Department of Animal Genetics and Conservation, Institute of Animal Sciences, Warsaw University of Life Sciences, 02-786 Warsaw, Poland.

Conflict of Interest Statement

The authors declare no conflict of interest.

References

This article includes 133 references
  1. Donaghy T. Horse Breeds and Breeding in the Greco-Persian World: 1st and 2nd Millennium BC. .
  2. Levine M.A. Botai and the origins of horse domestication. J. Anthropol. Archaeol. 1999;18:29–78.
    doi: 10.1006/jaar.1998.0332google scholar: lookup
  3. 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.
    doi: 10.1073/pnas.152330099pmc: PMC125071pubmed: 12130666google scholar: lookup
  4. Orlando L. The Evolutionary and Historical Foundation of the Modern Horse: Lessons from Ancient Genomics.. Annu Rev Genet 2020 Nov 23;54:563-581.
  5. Robinson IH. The human-horse relationship: how much do we know?. Equine Vet J Suppl 1999 Apr;(28):42-5.
  6. Hausberger M, Roche H, Henry S, Visser E.K. A review of the human-horse relationship. Appl. Anim. Behav. Sci. 2008;109:1–24.
  7. Thompson D.W. Historia Animalium. 1910. p. 633.
  8. Columella L.I.M. In: On Agriculture. Forster E.S., Heffner E.H., editors. Volume II. Loeb Class.; Harvard University Press; Cambridge, MA, USA: 1989. Books 5–9.
  9. Cato M.P., Varro M.T. In: De Re Rustica. Ash H.B., Page T.E., Hooper W.D., editors. Loeb Class.; Harvard University Press; Cambridge, MA, USA: 1967.
  10. Donaghy T. From Aristotle to Pelagonius: Greek and Latin writers on ancient horse breeding. Vet. Hist. 2016;18:320–344.
  11. Virgil . In: The Georgics. Thomas R.F., editor. Volume II. Cambridge University Press; Cambridge, UK: 2008. Book III–IV.
  12. Van Kolfschoten T. The palaeolithic locality Schöningen (Germany): A review of the mammalian record. Quat. Int. 2014;326–327:469–480.
  13. Fages A, Seguin-Orlando A, Germonpré M, Orlando L. Horse males became over-represented in archaeological assemblages during the Bronze Age. J. Archaeol. Sci. Rep. 2020;31:102364.
  14. Richardson C. The Horse Breakers. 1998.
  15. Chamberlin J.E. Horse. How The Horse Has Shaped Civilizations. 2006.
  16. Gaunitz C, Fages A, Hanghøj K, Albrechtsen A, Khan N, Schubert M, Seguin-Orlando A, Owens IJ, Felkel S, Bignon-Lau O, de Barros Damgaard P, Mittnik A, Mohaseb AF, Davoudi H, Alquraishi S, Alfarhan AH, Al-Rasheid KAS, Crubézy E, Benecke N, Olsen S, Brown D, Anthony D, Massy K, Pitulko V, Kasparov A, Brem G, Hofreiter M, Mukhtarova G, Baimukhanov N, Lõugas L, Onar V, Stockhammer PW, Krause J, Boldgiv B, Undrakhbold S, Erdenebaatar D, Lepetz S, Mashkour M, Ludwig A, Wallner B, Merz V, Merz I, Zaibert V, Willerslev E, Librado P, Outram AK, Orlando L. Ancient genomes revisit the ancestry of domestic and Przewalski's horses.. Science 2018 Apr 6;360(6384):111-114.
    doi: 10.1126/science.aao3297pubmed: 29472442google scholar: lookup
  17. Mileto S, Kaiser E, Rassamakin Y, Evershed R.P. New insights into the subsistence economy of the Eneolithic Dereivka culture of the Ukrainian North-Pontic region through lipid residues analysis of pottery vessels. J. Archaeol. Sci. Rep. 2017;13:67–74.
  18. Orlando L. Ancient Genomes Reveal Unexpected Horse Domestication and Management Dynamics.. Bioessays 2020 Jan;42(1):e1900164.
    doi: 10.1002/bies.201900164pubmed: 31808562google scholar: lookup
  19. Olsen S.L, Bradley B, Maki D, Outram A.K. Community organization among Copper Age sedentary horse pastoralists of Kazakhstan. 2006. pp. 89–111.
  20. Olsen S.L. The exploitation of horses at Botai, Kazakhstan. 2003. pp. 83–104.
  21. 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.
    doi: 10.1126/science.1168594pubmed: 19265018google scholar: lookup
  22. Anthony D.W, Brown D.R. Eneolithic horse exploitation in the Eurasian steppes: Diet, ritual and riding. Antiquity 2000;74:75–86.
    doi: 10.1017/S0003598X00066163google scholar: lookup
  23. Bökönyi S. The earliest waves of domestic horses in East Europe. J. Indo-Eur. Stud. 1978;6:17–76.
  24. Levine M.A. Dereivka and the problem of horse domestication. Antiquity 1990;64:727–740.
    doi: 10.1017/S0003598X00078832google scholar: lookup
  25. Bökönyi S. History of Domestic Mammals in Central and Eastern Europe. Akademiai Kiado; Budapest, Hungary: 1974.
  26. Lira J, Linderholm A, Olaria C, Brandström Durling M, Gilbert MT, Ellegren H, Willerslev E, Lidén K, Arsuaga JL, Götherström A. Ancient DNA reveals traces of Iberian Neolithic and Bronze Age lineages in modern Iberian horses.. Mol Ecol 2010 Jan;19(1):64-78.
  27. Arbuckle B.S. Animals and inequality in Chalcolithic central Anatolia. J. Anthropol. Archaeol. 2012;31:302–313.
    doi: 10.1016/j.jaa.2012.01.008google scholar: lookup
  28. Fages A, Hanghøj K, Khan N, Gaunitz C, Seguin-Orlando A, Leonardi M, McCrory Constantz C, Gamba C, Al-Rasheid KAS, Albizuri S, Alfarhan AH, Allentoft M, Alquraishi S, Anthony D, Baimukhanov N, Barrett JH, Bayarsaikhan J, Benecke N, Bernáldez-Sánchez E, Berrocal-Rangel L, Biglari F, Boessenkool S, Boldgiv B, Brem G, Brown D, Burger J, Crubézy E, Daugnora L, Davoudi H, de Barros Damgaard P, de Los Ángeles de Chorro Y de Villa-Ceballos M, Deschler-Erb S, Detry C, Dill N, do Mar Oom M, Dohr A, Ellingvåg S, Erdenebaatar D, Fathi H, Felkel S, Fernández-Rodríguez C, García-Viñas E, Germonpré M, Granado JD, Hallsson JH, Hemmer H, Hofreiter M, Kasparov A, Khasanov M, Khazaeli R, Kosintsev P, Kristiansen K, Kubatbek T, Kuderna L, Kuznetsov P, Laleh H, Leonard JA, Lhuillier J, Liesau von Lettow-Vorbeck C, Logvin A, Lõugas L, Ludwig A, Luis C, Arruda AM, Marques-Bonet T, Matoso Silva R, Merz V, Mijiddorj E, Miller BK, Monchalov O, Mohaseb FA, Morales A, Nieto-Espinet A, Nistelberger H, Onar V, Pálsdóttir AH, Pitulko V, Pitskhelauri K, Pruvost M, Rajic Sikanjic P, Rapan Papeša A, Roslyakova N, Sardari A, Sauer E, Schafberg R, Scheu A, Schibler J, Schlumbaum A, Serrand N, Serres-Armero A, Shapiro B, Sheikhi Seno S, Shevnina I, Shidrang S, Southon J, Star B, Sykes N, Taheri K, Taylor W, Teegen WR, Trbojević Vukičević T, Trixl S, Tumen D, Undrakhbold S, Usmanova E, Vahdati A, Valenzuela-Lamas S, Viegas C, Wallner B, Weinstock J, Zaibert V, Clavel B, Lepetz S, Mashkour M, Helgason A, Stefánsson K, Barrey E, Willerslev E, Outram AK, Librado P, Orlando L. Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series.. Cell 2019 May 30;177(6):1419-1435.e31.
    doi: 10.1016/j.cell.2019.03.049pmc: PMC6547883pubmed: 31056281google scholar: lookup
  29. 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.abb0030pmc: PMC7494339pubmed: 32938680google scholar: lookup
  30. Anthony D.W. The Horse, the Heel, and Language: How Bronze-Age Riders from the Eurasian Steppes Shaped the Modern World. 2007.
  31. Chechushkov I.V, Epimakhov A.V. Eurasian Steppe Chariots and Social Complexity during the Bronze Age. J. World Prehistory. 2018;31:435–483.
    doi: 10.1007/s10963-018-9124-0google scholar: lookup
  32. Bakker J.A, Kruk J, Lanting A.E, Milisauskas S. The earliest evidence of wheeled vehicles in Europe and the Near East. Antiquity 1999;73:778.
    doi: 10.1017/S0003598X00065522google scholar: lookup
  33. Kohl P.L. The Making of Bronze Age Eurasia. 2007.
  34. Koryakova L, Epimakhov A. The Urals and Western Siberia in the Bronze and Iron Ages. 2007.
  35. Anthony D.W, Telegin D.Y, Brown D.R. The Origin of Horseback Riding. Sci. Am. 1991;265:94–101.
  36. Bendrey R. New methods for the identification of evidence for bitting on horse remains from archaeological sites. J. Archaeol. Sci. 2007;34:1036–1050.
    doi: 10.1016/j.jas.2006.09.010google scholar: lookup
  37. Drews R. Early Riders: The Beginnings of Mounted Warfare in Asia and Europe. 2004.
  38. Weingarten J. Aegean Seals. An Introduction. Am. J. Archaeol. 2006;110:320–322.
  39. Littauer M.A, Crouwel J.H. The Earliest Evidence for Metal Bridle Parts. Oxford J. Archaeol. 2001;20:329–338.
    doi: 10.1111/1468-0092.00140google scholar: lookup
  40. Rossel S, Marshall F, Peters J, Pilgram T, Adams MD, O'Connor D. Domestication of the donkey: timing, processes, and indicators.. Proc Natl Acad Sci U S A 2008 Mar 11;105(10):3715-20.
    doi: 10.1073/pnas.0709692105pmc: PMC2268817pubmed: 18332433google scholar: lookup
  41. Kimura R, Tefera S. Taming requirements in the domestication process of donkeys: The methods and concepts of the training among the Afar pastoralists in Ethiopia. Pastoralism 2018;8.
    doi: 10.1186/s13570-018-0126-4google scholar: lookup
  42. Chondros T.G, Milidonis K.F, Rossi C, Zrnic N. The evolution of the double-horse chariots from the bronze age to the Hellenistic times. FME Trans. 2016;44:229–236.
    doi: 10.5937/fmet1603229Cgoogle scholar: lookup
  43. Drews R. The End of the Bronze Age. 1993.
  44. Littauer M.A, Crouwel J.H. The origin of the true chariot. Antiquity 1996;70:934–939.
    doi: 10.1017/S0003598X00084192google scholar: lookup
  45. de Barros Damgaard P, Martiniano R, Kamm J, Moreno-Mayar JV, Kroonen G, Peyrot M, Barjamovic G, Rasmussen S, Zacho C, Baimukhanov N, Zaibert V, Merz V, Biddanda A, Merz I, Loman V, Evdokimov V, Usmanova E, Hemphill B, Seguin-Orlando A, Yediay FE, Ullah I, Sjögren KG, Iversen KH, Choin J, de la Fuente C, Ilardo M, Schroeder H, Moiseyev V, Gromov A, Polyakov A, Omura S, Senyurt SY, Ahmad H, McKenzie C, Margaryan A, Hameed A, Samad A, Gul N, Khokhar MH, Goriunova OI, Bazaliiskii VI, Novembre J, Weber AW, Orlando L, Allentoft ME, Nielsen R, Kristiansen K, Sikora M, Outram AK, Durbin R, Willerslev E. The first horse herders and the impact of early Bronze Age steppe expansions into Asia.. Science 2018 Jun 29;360(6396).
    doi: 10.1126/science.aar7711pmc: PMC6748862pubmed: 29743352google scholar: lookup
  46. McMiken DF. Ancient origins of horsemanship.. Equine Vet J 1990 Mar;22(2):73-8.
  47. Anthony D.W. The Sintashta Genesis. The Roles of Climate Change, Warfare, and Long-Distance Trade. 2009. pp. 47–73.
  48. Littauer M.A, Crouwel J. Wheeled Vehicles and Ridden Animals in the Ancient Near East. 1987.
  49. Olsen S.L. Early Horse Domestication on the Eurasian Steppe. 2006. pp. 245–269.
  50. Moorey P.R.S. The emergence of the light, horse-drawn chariot in the Near-East c. 2000–1500 B.C.. World Archaeol. 1986;18:196–215.
  51. Latacz J. Troy and Homer: Towards a Solution of an Old Mystery. 2004.
  52. Oki H, Sasaki Y, Willham R.L. Genetic parameter estimates for racing time by restricted maximum likelihood in the Thoroughbred horse of Japan. J. Anim. Breed. Genet. 1995;112:146–150.
  53. Sobczyńska M. Environmental factors affecting the speed of Thoroughbred horses competing in Poland. Anim. Sci. Pap. Rep. 2011;29:303–312.
  54. Hitchens PL, Blizzard CL, Jones G, Day LM, Fell J. The association between jockey experience and race-day falls in flat racing in Australia.. Inj Prev 2012 Dec;18(6):385-91.
  55. Crouwel J.H. Chariots and Other Means of Land Transport in Bronze Age Greece. 1981. pp. 28–38.
  56. Anthony D.W, Brown D.R. The Secondary Products Revolution, Horse-Riding, and Mounted Warfare. J. World Prehistory. 2011;24:131–160.
    doi: 10.1007/s10963-011-9051-9google scholar: lookup
  57. Di Cosmo N. State formation and periodization in inner Asia. Int. Hist. Rev. 1999;20:287–309.
  58. Ewers J.C. In: The Horse In Blackfoot Indian Culture with Comparative Material from Other Western Tribes. Smithsonian Institution Bureau of American Ethnology, editor. Volume 159. United States Government Printing Office; Washington, DC, USA: 1955.
  59. Anderson J.K. Homeric, British and Cyrenaic Chariots. Am. J. Archaeol. 1965;69:349–352.
    doi: 10.2307/502184google scholar: lookup
  60. Littauer M.A. The Military Use of the Chariot in the Aegean in the Late Bronze Age. Am. J. Archaeol. 1972;76:145.
    doi: 10.2307/503858google scholar: lookup
  61. Crouwel J.H. Chariots and Other Wheeled Vehicles in Iron Age Greece. 1992.
  62. Littauer M.A, Crouwel J.H. Robert Drews and the role of chariotry in bronze age Greece. Oxf. J. Archaeol. 1996;15:297–305.
  63. Kelder J. Horseback Riding and Cavalry in Mycenaean Greece. Anc. West East. 2012;11:1–18.
    doi: 10.2143/AWE.11.0.2175875google scholar: lookup
  64. Gaebel R.E. Cavalry Operations in the Ancient Greek World. 2002.
  65. Greenhalgh P.A.L. Early Greek Warfare: Horsemen and Chariots in the Homeric and Archaic Ages. 1973.
  66. Stefaniuk-Szmukier M, Ropka-Molik K, Piórkowska K, Szmatoła T, Długosz B, Pisarczyk W, Bugno-Poniewierska M. Variation in TBX3 Gene Region in Dun Coat Color Polish Konik Horses. J. Equine Vet. Sci. 2017;49:60–62.
  67. May-Davis S, Brown WY, Shorter K, Vermeulen Z, Butler R, Koekkoek M. A Novel Non-Invasive Selection Criterion for the Preservation of Primitive Dutch Konik Horses.. Animals (Basel) 2018 Feb 1;8(2).
    doi: 10.3390/ani8020021pmc: PMC5836029pubmed: 29389896google scholar: lookup
  68. Stachurska A.M. Inheritance of primitive markings in horses. J. Anim. Breed. Genet. 1999;116:29–38.
  69. BENIRSCHKE K, MALOUF N, LOW RJ, HECK H. CHROMOSOME COMPLEMENT: DIFFERENCES BETWEEN EQUUS CABALLUS AND EQUUS PRZEWALSKII, POLIAKOFF.. Science 1965 Apr 16;148(3668):382-3.
    doi: 10.1126/science.148.3668.382pubmed: 14261533google scholar: lookup
  70. Wallner B, Brem G, Müller M, Achmann R. Fixed nucleotide differences on the Y chromosome indicate clear divergence between Equus przewalskii and Equus caballus.. Anim Genet 2003 Dec;34(6):453-6.
  71. Creel H.G. The Role of the Horse in Chinese History. Am. Hist. Rev. 1965;70:647.
    doi: 10.2307/1845936google scholar: lookup
  72. Azzaroli A. An Early History of Horsemanship. 1985.
  73. Delpeut L, Hettema H. Ancient Arabian horses? Revisiting ancient Egyptian equine imagery; Proceedings of the Current Research in Egyptology 2019: Proceedings of the Twentieth Annual Symposium, University of Alcalá; 17–21 June 2019; Gloucestershire, UK. 2021. p. 168.
  74. Wentworth J. Horses of The World. West. Horseman 1946;2:18–19, 50–51, 54–55.
  75. Czarnecki R. Effect of Polish and foreign purebred Arabian stallions on conformation traits of their progeny participating in shows in the last decade. J. Cent. Eur. Agric. 2014;15:115–123.
    doi: 10.5513/JCEA01/15.4.1517google scholar: lookup
  76. Kaproń M, Czerniak E, Łukaszewicz M, Danielewicz A. The breeding value of Wielkopolski horses belonging to particular stallion lineages in the successive volume of the stud book, as evidenced by their body conformation and performance traits. Acta Sci. Pol. Zootech. 2015;14:77–90.
  77. Francfort H.-P, Lepetz S. Les chevaux de Berel’ (Altaï)—Chevaux steppiques et chevaux achéménides: Haras et races. Histoire d’équidés 2010;4:57–104.
  78. Librado P, Gamba C, Gaunitz C, Der Sarkissian C, Pruvost M, Albrechtsen A, Fages A, Khan N, Schubert M, Jagannathan V, Serres-Armero A, Kuderna LFK, Povolotskaya IS, Seguin-Orlando A, Lepetz S, Neuditschko M, Thèves C, Alquraishi S, Alfarhan AH, Al-Rasheid K, Rieder S, Samashev Z, Francfort HP, Benecke N, Hofreiter M, Ludwig A, Keyser C, Marques-Bonet T, Ludes B, Crubézy E, Leeb T, Willerslev E, Orlando L. Ancient genomic changes associated with domestication of the horse.. Science 2017 Apr 28;356(6336):442-445.
    doi: 10.1126/science.aam5298pubmed: 28450643google scholar: lookup
  79. Signer-Hasler H, Flury C, Haase B, Burger D, Simianer H, Leeb T, Rieder S. A genome-wide association study reveals loci influencing height and other conformation traits in horses.. PLoS One 2012;7(5):e37282.
  80. Willekes C. Breeding success: The creation of the racehorse in antiquity. Mouseion 2019;16:453–469.
    doi: 10.3138/mous.16.3.006google scholar: lookup
  81. Johnstone C.J. A Biometric Study of Equids in the Roman World. 2004.
  82. Rieder S. Molecular tests for coat colours in horses.. J Anim Breed Genet 2009 Dec;126(6):415-24.
  83. Cieslak M, Reissmann M, Hofreiter M, Ludwig A. Colours of domestication.. Biol Rev Camb Philos Soc 2011 Nov;86(4):885-99.
  84. Sponenberg D.P, Bellone R. Equine Color Genetics. 2017.
  85. 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.
    doi: 10.1126/science.1172750pmc: PMC5102060pubmed: 19390039google scholar: lookup
  86. Pruvost M, Bellone R, Benecke N, Sandoval-Castellanos E, Cieslak M, Kuznetsova T, Morales-Muñiz A, O'Connor T, Reissmann M, Hofreiter M, Ludwig A. Genotypes of predomestic horses match phenotypes painted in Paleolithic works of cave art.. Proc Natl Acad Sci U S A 2011 Nov 15;108(46):18626-30.
    doi: 10.1073/pnas.1108982108pmc: PMC3219153pubmed: 22065780google scholar: lookup
  87. Kukekova AV, Trut LN, Chase K, Shepeleva DV, Vladimirova AV, Kharlamova AV, Oskina IN, Stepika A, Klebanov S, Erb HN, Acland GM. Measurement of segregating behaviors in experimental silver fox pedigrees.. Behav Genet 2008 Mar;38(2):185-94.
    doi: 10.1007/s10519-007-9180-1pmc: PMC2374754pubmed: 18030612google scholar: lookup
  88. Trut L.N. Early canid domestication: The farm-fox experiment: Foxes bred for tamability in a 40-year experiment exhibit remarkable transformations that suggest an interplay between behavioral genetics and development. Am. Sci. 1999;87:160–169.
    doi: 10.1511/1999.2.160google scholar: lookup
  89. van Dierendonck M, Goodwin D. Social contact in horses: Implications for human-horse interractions. 2005. pp. 65–81.
  90. Trut L.N, Oskina I.N, Kharlamova A.V. Experimental studies of early canid domestication. 2012. pp. 12–37.
  91. Prasolova L.A, Oskina I.N, Plyusnina I, Trut L.N. Maternal methyl supplements affect the phenotypic variation of the agouti gene in the offspring of rats with different behavioral types. Russ. J. Genet. 2009;45:587–592.
    doi: 10.1134/S1022795409050111google scholar: lookup
  92. Albert FW, Carlborg O, Plyusnina I, Besnier F, Hedwig D, Lautenschläger S, Lorenz D, McIntosh J, Neumann C, Richter H, Zeising C, Kozhemyakina R, Shchepina O, Kratzsch J, Trut L, Teupser D, Thiery J, Schöneberg T, Andersson L, Pääbo S. Genetic architecture of tameness in a rat model of animal domestication.. Genetics 2009 Jun;182(2):541-54.
    doi: 10.1534/genetics.109.102186pmc: PMC2691762pubmed: 19363126google scholar: lookup
  93. Jacobs LN, Staiger EA, Albright JD, Brooks SA. The MC1R and ASIP Coat Color Loci May Impact Behavior in the Horse.. J Hered 2016 May;107(3):214-9.
    doi: 10.1093/jhered/esw007pmc: PMC4885240pubmed: 26884605google scholar: lookup
  94. Rankins E.M, Wickens C.L. A systematic review of equine personality. Appl. Anim. Behav. Sci. 2020;231:105076.
  95. 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.1416991111pmc: PMC4284583pubmed: 25512547google scholar: lookup
  96. Librado P, Der Sarkissian C, Ermini L, Schubert M, Jónsson H, Albrechtsen A, Fumagalli M, Yang MA, Gamba C, Seguin-Orlando A, Mortensen CD, Petersen B, Hoover CA, Lorente-Galdos B, Nedoluzhko A, Boulygina E, Tsygankova S, Neuditschko M, Jagannathan V, Thèves C, Alfarhan AH, Alquraishi SA, Al-Rasheid KA, Sicheritz-Ponten T, Popov R, Grigoriev S, Alekseev AN, Rubin EM, McCue M, Rieder S, Leeb T, Tikhonov A, Crubézy E, Slatkin M, Marques-Bonet T, Nielsen R, Willerslev E, Kantanen J, Prokhortchouk E, Orlando L. Tracking the origins of Yakutian horses and the genetic basis for their fast adaptation to subarctic environments.. Proc Natl Acad Sci U S A 2015 Dec 15;112(50):E6889-97.
    doi: 10.1073/pnas.1513696112pmc: PMC4687531pubmed: 26598656google scholar: lookup
  97. Bell S.W, Willekes C. Horse Racing and Chariot Racing. 2014. pp. 478–490.
  98. Hyland A. The Horse in the Ancient World. 2003.
  99. Monard A.M, Duncan P. Consequences of natal dispersal in female horses. Anim. Behav. 1996;52:565–579.
    doi: 10.1006/anbe.1996.0198google scholar: lookup
  100. 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. Reconstructing the origin and spread of horse domestication in the Eurasian steppe.. Proc Natl Acad Sci U S A 2012 May 22;109(21):8202-6.
    doi: 10.1073/pnas.1111122109pmc: PMC3361400pubmed: 22566639google scholar: lookup
  101. Warmuth V, Manica A, Eriksson A, Barker G, Bower M. Autosomal genetic diversity in non-breed horses from eastern Eurasia provides insights into historical population movements.. Anim Genet 2013 Feb;44(1):53-61.
  102. Lippold S, Matzke NJ, Reissmann M, Hofreiter M. Whole mitochondrial genome sequencing of domestic horses reveals incorporation of extensive wild horse diversity during domestication.. BMC Evol Biol 2011 Nov 14;11:328.
    doi: 10.1186/1471-2148-11-328pmc: PMC3247663pubmed: 22082251google scholar: lookup
  103. 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.
    doi: 10.1073/pnas.1111637109pmc: PMC3289334pubmed: 22308342google scholar: lookup
  104. 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.
    doi: 10.1038/ng1326pubmed: 15034578google scholar: lookup
  105. 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.
    doi: 10.1093/jhered/esq047pubmed: 20497969google scholar: lookup
  106. 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.
    doi: 10.1038/ncomms1447pubmed: 21863017google scholar: lookup
  107. 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.aap9691pmc: PMC5906072pubmed: 29675468google scholar: lookup
  108. Levine MA. Investigating the origins of horse domestication.. Equine Vet J Suppl 1999 Apr;(28):6-14.
  109. McDowell K.J, Powell D.G, Baker C.B. Effect of book size and age of mare and stallion on foaling rates in thoroughbred horses. J. Equine Vet. Sci. 1992;12:364–367.
  110. Nagy P, Guillaume D, Daels P. Seasonality in mares.. Anim Reprod Sci 2000 Jul 2;60-61:245-62.
    doi: 10.1016/S0378-4320(00)00133-0pubmed: 10844199google scholar: lookup
  111. Mahboubi M, Mohammad Taghizadeh Kashani L, Mahboubi M. Squill (Drimia maritima L.) and its novel biological activity. Orient. Pharm. Exp. Med. 2019;19:227–234.
    doi: 10.1007/s13596-018-0341-ygoogle scholar: lookup
  112. Waran N.K, Clarke N, Farnworth M. The effects of weaning on the domestic horse (Equus caballus). Appl. Anim. Behav. Sci. 2008;110:42–57.
  113. Rodgers R. Cattle, Horses and Other Livestock: Textual Notes on Columella, Books 6 and 7. Riv. Cult. Class. Medioev. 2009;LI:1000–1041.
    doi: 10.1400/114909google scholar: lookup
  114. Raulwing P. The Kikkuli Text. Hittite Training Instructions for Chariot Horses in the Second Half of the 2nd Millennium BC and Their Interdisciplinary Context. .
  115. Amemiya T. Economy and Economics of Ancient Greece. 2007.
  116. Herz P. Finances and Costs of the Roman Army. 2007. p. 306.
  117. Heesch J. Van. Some Aspects of Wage Payments and Coinage in Ancient Rome, First to Third Centuries AD. 2007. pp. 77–96.
  118. Cioffi R.L. Travel in the Roman World. 2016.
  119. Golden M. Equestrian Competition in Ancient Greece: Difference, Dissent, Democracy. Phoenix 1997;51:327.
    doi: 10.2307/1192541google scholar: lookup
  120. Fischer KD. Ancient veterinary medicine. A survey of Greek and Latin sources and some recent scholarship.. Medizinhist J 1988;23(3-4):191-209.
    pubmed: 11622693
  121. Doyen-Higuet A.-M. The “Hippiatrica” and Byzantine Veterinary Medicine. Dumbart. Oaks Pap. 1984;38:111.
    doi: 10.2307/1291499google scholar: lookup
  122. Odder E., Hoppe K. In: Corpus Hippiatricorum Graecorum. I. Hippiatrica Berolinensia. Teubner B.G., editor. Teubneri; Leipzig, Germany: 1924. pp. 22–38.
  123. Odder K. Corpus Hippiatricorum Graecorum. II. Hippiatrica Parisina, Cantabrigiensia, Londinensia, Lugdunensia. Teubner; Leipzig, Germany: 1927. pp. 57–59.
  124. Nyland A. The Kikkuli Method of Horse Training. 2008.
  125. Nicholson N.J. Aristocracy and Athletics in Archaic and Classical Greece. 2005.
  126. Potter D.S. Entertainers in the Roman Empire. 1999. pp. 280–350.
  127. Hemingway S.A. The Horse and Jockey from Artemision. A Bronze Equestrian Monument of the Hellenistic Period. 2004.
  128. McBane S, McCarthy G. The Competition Horse: Breeding, Production and Management. 1991.
  129. Humphrey J.H. Roman Circuses: Arenas for Chariot Racing. 1986.
  130. Bell S.W. Horse Racing in Imperial Rome: Athletic Competition, Equine Performance, and Urban Spectacle. Int. J. Hist. Sport. 2020;37:1–50.
  131. Bell S. A Companion to Sport and Spectacle in Greek and Roman Antiquity. 2013. pp. 492–504.
  132. Lim R. Inventing Secular Space in the Late Antique City: Reading the Circus Maximus. 2012. pp. 61–81.
  133. Larson J. The Emperor, the Church and Chariot Races. The Imperial Struggles with Christianity and Entertainment in Late Antique Constantinople. 2012.

Citations

This article has been cited 10 times.
  1. Noble GK. Horse Husbandry-Nutrition, Management and Welfare. Animals (Basel) 2023 Jan 1;13(1).
    doi: 10.3390/ani13010169pubmed: 36611777google scholar: lookup
  2. Yordanov G, Mehandjyiski I, Palova N, Atsenova N, Neov B, Radoslavov G, Hristov P. Genetic Diversity and Structure of the Main Danubian Horse Paternal Genealogical Lineages Based on Microsatellite Genotyping. Vet Sci 2022 Jul 1;9(7).
    doi: 10.3390/vetsci9070333pubmed: 35878350google scholar: lookup
  3. Sabiniewicz A, Białek M, Tarnowska K, Świątek R, Dobrowolska M, Sorokowski P. A Preliminary Investigation of Interspecific Chemosensory Communication of Emotions: Can Humans (Homo sapiens) Recognise Fear- and Non-Fear Body Odour from Horses (Equus ferus caballus). Animals (Basel) 2021 Dec 8;11(12).
    doi: 10.3390/ani11123499pubmed: 34944275google scholar: lookup
  4. Liu X, Zhang Y, Liu W, Li Y, Pan J, Pu Y, Han J, Orlando L, Ma Y, Jiang L. A single-nucleotide mutation within the TBX3 enhancer increased body size in Chinese horses. Curr Biol 2022 Jan 24;32(2):480-487.e6.
    doi: 10.1016/j.cub.2021.11.052pubmed: 34906355google scholar: lookup
  5. Wang T, Yang X, Wang C, Wang J, Meng J, Yao X, Zeng Y, Ren W. Bridging Breeds: Transcriptomic Insights into Immune Traits of Yili, Thoroughbred, and Kazakh Horses. Life (Basel) 2025 Sep 23;15(10).
    doi: 10.3390/life15101496pubmed: 41157169google scholar: lookup
  6. Kassymbekova SN, Bimenova ZZ, Iskhan KZ, Sobiech P, Jastrzebski JP, Brym P, Babis W, Kalykova AS, Otebayev ZM, Kabylbekova DI, Baneh H, Romanov MN. Uncovering Genetic Diversity and Adaptive Candidate Genes in the Mugalzhar Horse Breed Using Whole-Genome Sequencing Data. Animals (Basel) 2025 Sep 11;15(18).
    doi: 10.3390/ani15182667pubmed: 41007912google scholar: lookup
  7. Ahmad I, Ijaz S, Usman MM, Safdar A, Khan IU, Zeeshan M, Bukhari SSUH. Evaluating Forelimb and Hindlimb Joint Conformation of Morna Racehorses (Equus caballus). Vet Sci 2025 Jan 5;12(1).
    doi: 10.3390/vetsci12010020pubmed: 39852895google scholar: lookup
  8. Radovic L, Remer V, Rigler D, Bozlak E, Allen L, Brem G, Reissman M, Brockmann GA, Ropka-Molik K, Stefaniuk-Szmukier M, Kalinkova L, Kalashnikov VV, Zaitev AM, Raudsepp T, Castaneda C, von Butler-Wemken I, Patterson Rosa L, Brooks SA, Novoa-Bravo M, Kostaras N, Abdurasulov A, Antczak DF, Miller DC, Lopes MS, da Câmara Machado A, Lindgren G, Juras R, Cothran G, Wallner B. The global spread of Oriental Horses in the past 1,500 years through the lens of the Y chromosome. Proc Natl Acad Sci U S A 2024 Dec 3;121(49):e2414408121.
    doi: 10.1073/pnas.2414408121pubmed: 39556761google scholar: lookup
  9. Arefnejad B, Zeinalabedini M, Talebi R, Mardi M, Ghaffari MR, Vahidi MF, Nekouei MK, Szmatoła T, Salekdeh GH. Unveiling the population genetic structure of Iranian horses breeds by whole-genome resequencing analysis. Mamm Genome 2024 Jun;35(2):201-227.
    doi: 10.1007/s00335-024-10035-6pubmed: 38520527google scholar: lookup
  10. Lebrasseur O, More KD, Orlando L. Equine herpesvirus 4 infected domestic horses associated with Sintashta spoke-wheeled chariots around 4,000 years ago. Virus Evol 2024;10(1):vead087.
    doi: 10.1093/ve/vead087pubmed: 38465241google scholar: lookup