The modern domesticated horse (Equus caballus) is today spread throughout the world and among the most diverse creatures on the planet. In North America, the horse was part of the megafaunal extinctions at the end of the Pleistocene. Two wild subspecies survived until recently, the Tarpan (Equus ferus ferus, died out ca 1919) and Przewalski's Horse (Equus ferus przewalskii, of which there are a few left).
Horse history, especially the timing of the domestication of the horse, is still being debated, partly because the evidence for domestication itself is debatable. Unlike other animals, criteria such as changes in body morphology (horses are extremely diverse) or the location of a particular horse outside of its "normal range" (horses are very widespread) are not useful in helping resolve the question.
Horse History and the Evidence for Horse Domestication
The earliest possible hints for domestication would be the presence of what appear to be a set of postmolds with lots of animal dung within the area defined by the posts, which scholars interpret as representing a horse pen. That evidence has been found at Krasni Yar in Kazakhstan, in portions of the site dating to as early as 5000 BC. The horses may have been kept for food and milk, rather than riding or load-bearing.
Accepted archaeological evidence of horseback riding includes bit wear on horse teeth--that has been found in the steppes east of the Ural mountains at Botai and Kozhai 1 in modern Kazakhstan, around 3500-3000 BC. The bit wear was only found on a few of the teeth in the archaeological assemblages, which might suggest that a few horses were ridden to hunt and collect wild horses for food and milk consumption. Finally, the earliest direct evidence of the use of horses as beasts of burden--in the form of drawings of horse-drawn chariots--is from Mesopotamia, about 2000 BC.
Horse History and Genetics
Genetic data, interestingly enough, has traced all extant domesticated horses to one founder stallion, or to closely related male horses with the same Y haplotype. At the same time, there is a high matrilineal diversity in both domestic and wild horses. At least 77 wild mares would be required to explain the diversity of the mitochondrial DNA (mtDNA) in current horse populations, which probably means quite a few more.
A 2012 study (Warmuth and colleagues) combining archaeology, mitochondrial DNA, and Y-chromosomal DNA supports the domestication of horse as occurring once, in the western part of the Eurasian steppe, and that because of the horse's wild natures, several repeated introgression events (restocking of horse populations by adding wild mares), must have occurred. As identified in earlier studies, that would explain the diversity of mtDNA.
Three Strands of Evidence for Domesticated Horses
In a paper published in Science in 2009, Alan K. Outram and colleagues looked at three strands of evidence supporting horse domestication at Botai culture sites: shin bones, milk consumption, and bitwear. These data support domestication of the horse between about 3500-3000 BC sites in what is today Kazakhstan.
Horses skeletons at Botai Culture sites have gracile metacarpals. The horses' metacarpals-the shins or cannon bones-are used as key indicators of domesticity. For whatever reason (and I won't speculate here), shins on domestic horses are thinner--more gracile--than those of wild horses. Outram et al. describe the shinbones from Botai as being closer in size and shape to those of Bronze age (fully domesticated) horses compared to wild horses.
Fatty lipids of horse milk were found inside of pots. Although today it seems a bit weird to westerners, horses were kept for both their meat and milk in the past--and still are in the Kazakh region as you can see from the photograph above. Evidence of horse milk was found at Botai in the form of fatty lipid residues on the insides of ceramic vessels; further, evidence for consumption of horse meat has been identified at Botai culture horse and rider burials.
Bit wear is in evidence on horse teeth. Researchers noted bitting wear on horses' teeth--a vertical strip of wear on the outside of horses' premolars, where the metal bit damages the enamel when it sits between the cheek and tooth. Recent studies (Bendrey) using scanning electron microscopy with energy dispersive X-ray microanalysis found microscopic-sized fragments of iron embedded on Iron Age horse teeth, resulting from metal bit use.
White Horses and History
White horses are associated with the Pegasus myth, the unicorn in the Babylonian myth of Gilgamesh, Arabian horses, Lipizzaner stallions, Shetland ponies, and Icelandic pony populations.
- Read more about White Horses and Genetics
The Thoroughbred Gene
A recent DNA study (Bower et al.) examined the DNA of Thoroughbred racing horses, and identified the specific allele which drives their speed and precocity. Thoroughbreds are a specific breed of horse, all of whom today are descended from the children of one of three foundation stallions: Byerley Turk (imported to England in the 1680s), Darley Arabian (1704) and Godolphin Arabian (1729). These stallions are all of Arab, Barb and Turk origin; their descendants are from one of only 74 British and imported mares. Horse breeding histories for Thoroughbreds have been recorded in the General Stud Book since 1791, and the genetic data certainly support that history.
Horse races in the 17th and 18th centuries ran 3,200-6,400 meters (2-4 miles), and horses were usually five or six years old. By the early 1800s, the Thoroughbred was bred for traits that enabled speed and stamina over distances from 1,600-2,800 meters at three years of age; since the 1860s, the horses have been bred for shorter races (1,000-1400 meters) and younger maturity, at 2 years.
The genetic study looked at the DNA from hundreds of horses and identified the gene as C type myostatin gene variant, and came to the conclusion that this gene originated from a single mare, bred to one of the three founder male horses about 300 years ago. See Bower et al for additional information.
Thistle Creek DNA and Deep Evolution
In 2013, researchers led by Ludovic Orlando and Eske Willerslev of the Centre for GeoGenetics, Natural History Museum of Denmark and University of Copenhagen (and reported in Orlando et al. 2013) reported on a metapodial horse fossil which had been found in permafrost within a Middle Pleistocene context in the Yukon territory of Canada and dated between 560,00-780,000 years ago. Amazingly, the researchers found that there were sufficiently intact molecules of collagen within the matrix of the bone to enable them to map the Thistle Creek horse's genome.
Orlando and Willerslev's team found that over the past 500,000 years, horse populations have been enormously sensitive to climate change, and that extremely low population sizes are associated with warming events. Further, using the Thistle Creek DNA as a baseline, they were able to determine that all modern existing equids (donkeys, horses and zebras) originated from a common ancestor some 4-4.5 million years ago. In addition, Przewalski's horse diverged from the breeds which became domestic some 38,000-72,000 years ago, confirming the long-held belief that Przewalski's is the last remaining wild horse species.
This article is part of the About.com Guide to the History of Animal Domestication.
Bendrey R. 2012. From wild horses to domestic horses: a European perspective. World Archaeology 44(1):135-157.
Bendrey R. 2011. Identification of metal residues associated with bit-use on prehistoric horse teeth by scanning electron microscopy with energy dispersive X-ray microanalysis. Journal of Archaeological Science 38(11):2989-2994.
Bower MA, McGivney BA, Campana MG, Gu J, Andersson LS, Barrett E, Davis CR, Mikko S, Stock F, Voronkova V et al. 2012. The genetic origin and history of speed in the Thoroughbred racehorse. Nature Communications 3(643):1-8.
Brown D, and Anthony D. 1998. Bit Wear, Horseback Riding and the Botai Site in Kazakstan. Journal of Archaeological Science 25(4):331-347.
Cassidy R. 2009. The horse, the Kyrgyz horse and the ‘Kyrgyz horse’. Anthropology Today 25(1):12-15.
Jansen T, Forster P, Levine MA, Oelke H, Hurles M, Renfrew C, Weber J, Olek, and Klaus. 2002. Mitochondrial DNA and the origins of the domestic horse. Proceedings of the National Academy of Sciences 99(16):10905–10910.
Levine MA. 1999. Botai and the origins of horse domestication. Journal of Anthropological Archaeology 18(1):29-78.
Ludwig A, Pruvost M, Reissmann M, Benecke N, Brockmann GA, Castaños P, Cieslak M, Lippold S, Llorente L, Malaspinas A-S et al. 2009. Coat Color Variation at the Beginning of Horse Domestication. Science 324:485.
Kavar T, and Dovc P. 2008. Domestication of the horse: Genetic relationships between domestic and wild horses. Livestock Science 116(1):1-14.
Orlando L, Ginolhac A, Zhang G, Froese D, Albrechtsen A, Stiller M, Schubert M, Cappellini E, Petersen B, Moltke I et al. 2013. Recalibrating Equus evolution using the genome sequence of an early Middle Pleistocene horse. Nature in press.
Outram AK, Stear NA, Bendrey R, Olsen S, Kasparov A, Zaibert V, Thorpe N, and Evershed RP. 2009. The Earliest Horse Harnessing and Milking. Science 323:1332-1335.
Outram AK, Stear NA, Kasparov A, Usmanova E, Varfolomeev V, and Evershed RP. 2011. Horses for the dead: funerary foodways in Bronze Age Kazakhstan. Antiquity 85(327):116-128.
Sommer RS, Benecke N, Lõugas L, Nelle O, and Schmölcke U. 2011. Holocene survival of the wild horse in Europe: a matter of open landscape? Journal of Quaternary Science 26(8):805-812.
Rosengren Pielberg G, Golovko A, Sundström E, Curik I, Lennartsson J, Seltenhammer MH, Drum T, Binns M, Fitzsimmons C, Lindgren G et al. 2008. A cis-acting regulatory mutation causes premature hair graying and susceptibility to melanoma in the horse. Nature Genetics 40:1004-1009.
Warmuth V, Eriksson A, Bower MA, Barker G, Barrett E, Hanks BK, Li S, Lomitashvili D, Ochir-Goryaeva M, Sizonov GV et al. 2012. Reconstructing the origin and spread of horse domestication in the Eurasian steppe. Proceedings of the National Academy of Sciences Early edition.