The domestication of horses reading answers

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The domestication of horses reading answers
The domestication of horses reading answers with explanation. The domestication of horses reading answers with location. The domestication of horses reading answers ielts. The domestication of horses
reading answers passage 1.
Your name and responses will be shared with TED Ed.To track your work across TED-Ed over time, Register or Login instead. Only students who are 13 years of age or older can create a TED-Ed account. You should spend about 20 minutes on Questions 1-14 which are based on this passage.There is no doubt that dogs are the oldest of all species
tamed by humans and their domestication was based on a mutually beneficial relationship with man. The conventional view is that the domestication of wolves began between 10,000 and 20,000 years ago. However, a recent ground-breaking paper by a group of international geneticists has pushed this date back by a factor of 10. Led by Dr. Robert
Wayne, at the University of California, Los Angeles, the team showed that all dog breeds had only one ancestor, the wolf. They did this by analysing the genetic history through the DINA of 162 wolves from around the world and 140 domestic dogs representing 67 breeds. The research also confirms, for the first time, that dogs are descended only from
wolves and do not share DNA with coyotes or jackals. The fact that our companionship with dogs now appears to go back at least 100,000 years means that this partnership may have played an important part in the development of human hunting techniques that developed 70,000 to 90,000 years ago. It also may even have affected the brain
development in both species.The Australian veterinarian David Paxton suggests that in that period of first contact, people did not so much domesticate wolves as wolves domesticated people. Wolves may have started living at the edge of human settlements as scavengers, eating scraps of food and waste. Some learned to live with human beings in a
mutually helpful way and gradually evolved into dogs.
At the very least, they would have protected human settlements, and given warnings by barking at anything approaching. The wolves that evolved into dogs have been enormously successful in evolutionary terms. They are found everywhere in the inhabited world, hundreds of millions of them. The descendants of the wolves that remained wolves are
now sparsely distributed, often in endangered populations.In return for companionship and food, the early ancestor of the dog assisted humans in tracking, hunting, guarding and a variety of other activities. Eventually humans began to selectively breed these animals for specific traits.
Physical characteristics changed and individual breeds began to take shape. As humans wandered across Asia and Europe, they took their dogs along, using them for additional tasks and further breeding them for selected qualities that would better enable them to perform specific duties.According to Dr. Colin Groves, of the Department of
Archaeology and Anthropology at Australian National University, early humans came to rely on dogs’ keen ability to hear, smell and see - allowing certain areas of the human brain to shrink in size relative to oilier areas. ‘Dogs acted as human's alarm systems, trackers and hunting aids, garbage disposal facilities, hot-water bottles and children's
guardians and playmates.
Humans provided dogs with food and security. This symbiotic relationship was stable for over 100,000 years and intensified into mutual domestication,’ said Dr. Groves. In his opinion, humans domesticated dogs and dogs domesticated humans.Dr. Groves repealed an assertion made as early as 1914 that humans have some of the same physical
characteristics as domesticated animals, the most notable being decreased brain size. The horse experienced a 16 percent reduction in brain size after domestication while pigs’ brains shrank by as much as 34 percent. The estimated brain-size reduction in domesticated dogs varies from 30 percent to 10 percent. Only in the last decade have
archaeologists uncovered enough fossil evidence to establish that brain capacity in humans declined in Europe and Africa by at least 10 percent beginning about 10,000 years ago. Dr. Groves believes this reduction may have taken place as the relationship between humans and dogs intensified. The close interaction between the two species allowed
for the diminishing of certain human brain functions like smell and hearing. Domestication is the process by which humans take wild species and acclimatize them to breeding and surviving in captivity. In many cases, domesticated animals serve some purpose for humans (food source, labor, companionship). The process of domestication results in
physiological and genetic changes in the organisms over generations. Domestication differs from taming in that tamed animals are born in the wild while domesticated animals are bred in captivity. The history of horses in human culture can be traced back as far as 30,000 BC when horses were depicted in Paleolithic cave paintings. The horses in the
paintings resembled wild animals and it is thought that true domestication of horses did not occur for tens of thousands of years to come. It is thought that the horses depicted in the Paleolithic cave paintings were hunted for their meat by humans. There are several theories as to when and where domestication of the horse occurred. Some theories
estimate that domestication occurred at about 2000 BC while other theories place domestication as early as 4500 BC. Evidence from mitochondrial DNA studies suggests that the domestication of horses occurred in multiple locations and at various times.
It is generally thought that Central Asia is among the sites that domestication occurred, with sites in Ukraine and Kazakhstan providing archeological evidence. Throughout history, horses have been used for riding and for pulling carriages, chariots, plows, and carts. They played a significant role in warfare by carrying soldiers into battle. Because the
first domesticated horses are thought to have been quite small, it is more likely that they were used to pull carts than for riding. The domestication of horses reading answers The domestication of horses reading answers with explanation. The domestication of horses reading answers passage 1. The domestication of horses reading answers with
location. The domestication of horsesreading answers ielts. 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'sHorse (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 particularhorse outside of its "normal range" (horses are very widespread) are not useful in helping resolve the question. The earliest possible hints for domestication would be the presence of what appears 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 evidencehas been found at Krasnyi Yar in Kazakhstan, in portions of the site dating to as early as 3600 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 andKozhai 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. The saddle was invented around 800 BC, and the stirrup (a matter of some debate among historians) was probably
invented around 200-300 AD. Krasnyi Yar includes over 50 residential pithouses, adjacent to which have been found dozens of postmolds. The postmolds—archaeological remnants of where posts have been set in the past—are arranged in circles, and these are interpreted as evidence of horse corrals. 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. 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) horsescompared 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 have had a specialplace in ancient history-according to Herodotus, they were held as sacred animals in the Achaemenid court of Xerxes the Great (ruled 485-465 BC). 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. 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 supports that history. Horse races in the 17th and 18th centuries ran 3,200-6,400 meters (2-4miles), 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.

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The genetic study looked atthe 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. In 2013, researchers led by Ludovic Orlando and
Eske Willerslev of theCentre 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 researchersfound 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. The researchers then compared the Thistle Creek specimen DNA to that of an Upper Paleolithic horse, a modern donkey, five modern domestic horse breeds, and one modern
Przewalski's horse.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-heldbelief that Przewalski's is the last remaining wild horse species.
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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
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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. The origins and spread of
domestichorses from the Western Eurasian steppes Pablo Librado1, Naveed Khan1,121 nAff120, Antoine Fages1, Mariya A. Kusliy1,2, Tomasz Suchan orcid.org/0000-0002-0811-87541,3, Laure Tonasso-Calvière1, Stéphanie Schiavinato1, Duha Alioglu1, Aurore Fromentier1, Aude Perdereau4, Jean-Marc Aury orcid.org/0000-0003-1718-30105,
Charleen Gaunitz1, Lorelei Chauvey1, Andaine Seguin-Orlando1, Clio Der Sarkissian1, John Southon6, Beth Shapiro7,8, Alexey A.Tishkin9, Alexey A. Kovalev orcid.org/0000-0003-2637-313110, Saleh Alquraishi11, Ahmed H. Alfarhan11, Khaled A. S. Al-Rasheid orcid.org/0000-0002-3404-339711, Timo Seregély12, Lutz Klassen13, Rune Iversen
orcid.org/0000-0001-7618-625X14, Olivier Bignon-Lau15, Pierre Bodu15, Monique Olive15, Jean-Christophe Castel16, Myriam Boudadi-Maligne17, Nadir Alvarez18,19, Mietje Germonpré orcid.org/0000-0001-8865-093720, Magdalena Moskal-del Hoyo3, Jarosław Wilczyński orcid.org/0000-0002-9786-069321, Sylwia Pospuła21, Anna Lasota-Kuś22,
Krzysztof Tunia22, Marek Nowak23, Eve Rannamäe24, Urmas Saarma25, Gennady Boeskorov26, Lembi Lōugas orcid.org/0000-0003-2011-214127, René Kyselý28, Lubomír Peške29, Adrian Bălășescu30, Valentin Dumitrașcu30, Roxana Dobrescu30, Daniel Gerber31,32, Viktória Kiss33, Anna Szécsényi-Nagy31, Balázs G. Mende31, Zsolt Gallina34,
Krisztina Somogyi35, Gabriella Kulcsár33, Erika Gál orcid.org/0000-0002-4226-321833, Robin Bendrey36, Morten E. Allentoft37,38,Ghenadie Sirbu39, Valentin Dergachev40, Henry Shephard41, Noémie Tomadini42, Sandrine Grouard42, Aleksei Kasparov43, Alexander E. Basilyan44, Mikhail A. Anisimov45, Pavel A. Nikolskiy44, Elena Y. Pavlova45,
Vladimir Pitulko orcid.org/0000-0001-5672-275643, Gottfried Brem46, Barbara Wallner orcid.org/0000-0003-4159-069546,Christoph Schwall orcid.org/0000-0002-6310-405647, Marcel Keller48,49, Keiko Kitagawa orcid.org/0000-0002-6998-845450,51,52, Alexander N. Bessudnov53, Alexander Bessudnov orcid.org/0000-0002-3785-634243, William
Taylor orcid.org/0000-0002-0836-781454, Jérome Magail orcid.org/0000-0002-5768-069855, Jamiyan-Ombo Gantulga56, Jamsranjav Bayarsaikhan57,58, Diimaajav Erdenebaatar59, Kubatbeek Tabaldiev60, Enkhbayar Mijiddorj59, Bazartseren Boldgiv orcid.org/0000-0003-0015-814261, Turbat Tsagaan orcid.org/0000-0001-6606-851656, Mélanie
Pruvost orcid.org/0000-0001-7824-215517, Sandra Olsen62, Cheryl A. Makarewicz orcid.org/0000-0002-1649-336X63,64,Silvia Valenzuela Lamas orcid.org/0000-0001-9886-037265, Silvia Albizuri Canadell orcid.org/0000-0001-6194-047566, Ariadna Nieto Espinet67, Ma Pilar Iborra orcid.org/0000-0002-4315-725768, Jaime Lira Garrido69,70, Esther
Rodríguez González71, Sebastián Celestino71, Carmen Olària72, Juan Luis Arsuaga70,73, Nadiia Kotova74, AlexanderPryor75, Pam Crabtree76, Rinat Zhumatayev77, Abdesh Toleubaev77, Nina L.
Morgunova78, Tatiana Kuznetsova79,80, David Lordkipanize81,82, Matilde Marzullo83, Ornella Prato83, Giovanna Bagnasco Gianni83, Umberto Tecchiati83, Benoit Clavel42, Sébastien Lepetz42, Hossein Davoudi orcid.org/0000-0002-5236-144484, Marjan Mashkour orcid.org/0000-0003-3630-945942,84, Natalia Ya. Berezina85, Philipp W.
Stockhammer86,87, Johannes Krause orcid.org/0000-0001-9144-392049,86, Wolfgang Haak orcid.org/0000-0003-2475-200749,86,88, Arturo Morales-Muñiz89, Norbert Benecke90, Michael Hofreiter91, Arne Ludwig orcid.org/0000-0001-7249-995392,93, Alexander S.Graphodatsky2, Joris Peters orcid.org/0000-0003-0894-262894,95, Kirill Yu.
Kiryushin9, Tumur-Ochir Iderkhangai59, Nikolay A. Bokovenko43, Sergey K. Vasiliev96, Nikolai N. Seregin9, Konstantin V. Chugunov97, Natalya A. Plasteeva98, Gennady F. Baryshnikov99, Ekaterina Petrova100, Mikhail Sablin orcid.org/0000-0002-2773-745499, Elina Ananyevskaya100, Andrey Logvin101, Irina Shevnina101, Victor Logvin102, Saule
Kalieva orcid.org/0000-0003-1901-1001102, Valeriy Loman orcid.org/0000-0001-6951-0509103,Igor Kukushkin103, Ilya Merz104, Victor Merz104, Sergazy Sakenov105, Victor Varfolomeyev103, Emma Usmanova103, Viktor Zaibert106, Benjamin Arbuckle orcid.org/0000-0002-5445-5516107, Andrey B. Belinskiy108, Alexej Kalmykov108, Sabine
Reinhold orcid.org/0000-0002-8107-630090, Svend Hansen90, Aleksandr I. Yudin109, Alekandr A. Vybornov110, Andrey Epimakhov111,112, Natalia S. Berezina113, Natalia Roslyakova orcid.org/0000-0002-1888-2713110, Pavel A.
Kosintsev98,114, Pavel F. Kuznetsov110, David Anthony115,116, Guus J. Kroonen orcid.org/0000-0002-3708-0476117,118, Kristian Kristiansen119,120, Patrick Wincker orcid.org/0000-0001-7562-34545, Alan Outram orcid.org/0000-0003-3360-089X75 & …Ludovic Orlando orcid.org/0000-0003-3936-18501 Nature 598, 634–640 (2021)Cite this article
110k Accesses 68Citations 2781 Altmetric Metrics Evolutionary geneticsPopulation genetics Domestication of horses fundamentally transformed long-range mobility and warfare1. However, modern domesticated breeds do not descend from the earliest domestic horse lineage associated with archaeological evidence of bridling, milking and
corralling2,3,4 at Botai,Central Asia around 3500 bc3. Other longstanding candidate regions for horse domestication, such as Iberia5 and Anatolia6, have also recently been challenged. Thus, the genetic, geographic and temporal origins of modern domestic horses have remained unknown. Here we pinpoint the Western Eurasian steppes, especially
the lower Volga-Donregion, as the homeland of modern domestic horses. Furthermore, we map the population changes accompanying domestication from 273 ancient horse genomes. This reveals that modern domestic horses ultimately replaced almost all other local populations as they expanded rapidly across Eurasia from about 2000 bc,
synchronously with equestrianmaterial culture, including Sintashta spoke-wheeled chariots.
We find that equestrianism involved strong selection for critical locomotor and behavioural adaptations at the GSDMC and ZFPM1 genes.
Our results reject the commonly held association7 between horseback riding and the massive expansion of Yamnaya steppe pastoralists into Europearound 3000 bc8,9 driving the spread of Indo-European languages10. This contrasts with the scenario in Asia where Indo-Iranian languages, chariots and horses spread together, following the early
second millennium bc Sintashta culture11,12. We gathered horse remains encompassing all suspected domestication centres, including Iberia, Anatolia and the steppes of Western Eurasia and Central Asia (Fig 1a).
The sampling targeted previously under-represented time periods, with 201 radiocarbon dates spanning 44426 to 202 bc, and five beyond 50250 to 47950 bc (Supplementary Table1).Fig.
1: Ancient horse remains and their genomic affinities.a, Temporal and geographic sampling. The red star indicates the location of the two TURG horses (late Yamnaya context) showing genetic continuity with DOM2. The dashed line indicates the inferred homeland of DOM2 horses in the lower Volga-Don region. Colours refer to regions and/or time
periods delineating genetically close horses. The radius of each cylinder is proportional to thenumber of samples analysed (for <10 specimens; radius constant above this), and the height refers to the time range covered. b, Neighbour-joining phylogenomic tree (100 bootstrap pseudo-replicates). Samples are coloured according to a and the main
phylogenetic clusters are numbered from 1 to 4. c, Fold difference between neighbour-joining-based and raw pairwise genetic distances. d, Pairwise distance matrix of Struct-f4 genetic affinities between samples. Increasing genetic affinities are indicated by a yellow-to-red gradient. e, Struct-f4 ancestry component profiles. f, Ancestry profiles of
selected key horse groups and samples. PRZE, Przewalski; UP-SFR, Upper Palaeolithic Southern France.The DNA quality enabled shotgun sequencing of 264 ancient genomes at 0.10× to 25.76× average coverage (239genomes above 1× coverage), including 16 genomes for which further sequencing added to previously reported data. Enzymatic13
and computational removal of post mortem DNA damage produced high-quality data with derived mutations decreasing with sample age, as expected if mutations accumulate through time (Extended Data Fig. 1). We added ten published modern genomes, and nine ancient genomes characterized with consistent technology or covering relevant time
periods and locations, to obtain the most extensive high-quality genome time series for horses.Neighbour-joining phylogenomic inference revealed four geographically defined monophyletic groups (Fig 1b). Theseclosely mirrored clusters identified using an extension of the Struct-f4 method5 (Fig 1d–f, Extended Data Fig. 2, Supplementary Methods),
except for the Neolithic Anatolia group (NEO-ANA), where the tree-to-data goodness of fit suggested phylogenetic misplacement (Fig 1c, Supplementary Methods).The most basal cluster included Equus lenensis(ELEN), a lineage identified in northeastern Siberia from the Late Pleistocene to the late fourth millennium bc5,14,15. A second group
covered Europe, including Late Pleistocene Romania, Belgium, France and Britain, and the region from Spain to Scandinavia and Hungary, Czechia and Poland during the sixth-to-third millennium bc. The third clustercomprised the earliest known domestic horses from Botai and Przewalski’s horses, as previously reported3, and extended to the Altai
and Southern Urals during the fifth-to-third millennium bc. Finally, modern domestic horses clustered within a group that became geographically widespread and prominent following about 2200 bc and during the second millennium bc (DOM2). This cluster appears genetically close to horses that lived in the Western Eurasia steppes (WE) but not
further west than the Romanian lower Danube, south of theCarpathians, before and during the third millennium bc. Significant correlation between genetic and geographic distances, and inference of limited long-distance connectivity with estimated effective migration surface16 (EEMS), confirmed the strong geographic differentiation of horse
populations before about 3000 bc (Fig 2a, Extended Data Fig.3a).Fig. 2: Horse geographic and genetic affinities.a–c, EEMS-predicted migration barriers16 and average ancestry components found in each archaeological site from before 3000 bc (a), during the third millennium bc (b) and after around 2000 bc (c). The size of the pie charts is
proportional to the number of samples analysed in a given location (<10, constant above). Pie chart colours refer to K = 6 ancestry components, averaged per location. Regions inferred as geographic barriers are shown in shades of brown, and regions affected by migrations are shown in shades of blue. The base map wasobtained from
rworldmap46.Horse ancestry profiles in Neolithic Anatolia and Eneolithic Central Asia, including at Botai, maximized a genetic component (coloured green in Fig. 1e, f) that was also substantial in Central and Eastern Europe during the Late Pleistocene (RONPC06_Rom_m34801) and the fourth or third millennium bc (Figs. 1e, 3a,Extended Data Fig.
4). It was, however, absent or moderately present in the Romanian lower Danube (ENEO-ROM), the Dnieper steppes (Ukr11_Ukr_m4185) and the western lower Volga-Don (C-PONT) populations during the sixth to third millennia bc. This indicates possible expansions of Anatolian horses into both Central and Eastern Europe andCentral Asia regions,
but not into the Western Eurasia steppes. The absence of typical NEO-ANA ancestry rules out expansion from Anatolia into Central Asia across the Caucasus mountains but supports connectivity south of the Caspian Sea prior to about 3500 bc.Fig.
3: Population genetic affinities, evolutionary history and geographic origins.a,Multi-dimensional scaling plot of f4-based genetic affinities. The age of the samples is indicated along the vertical axis. CA, Central Asia. b, Horse evolutionary history inferred by OrientAGraph19 with three migration edges and nine lineages representing key genomic
ancestries (coloured as in Fig 1a). The model explains 99.99% of the total variance.The triangular pairwise matrix provides model residuals. The external branch leading to donkey was set to zero to improve visualization. c, LOCATOR20 predictions of the geographic region where the ancestors of DOM2, tarpan and modern Przewalski’s horses lived.
The tarpan and modern Przewalski’s horses do not descend from the same ancestralpopulation as modern domestic horses. The map was drawn using the maps R package47.The C-PONT group not only possessed moderate NEO-ANA ancestry, but also was the first region where the typical DOM2 ancestry component (coloured orange in Fig. 1e, f)
became dominant during the sixth millennium bc. Multi-dimensional scaling furtheridentified three horses from the western lower Volga-Don region as genetically closest to DOM2, associated with Steppe Maykop (Aygurskii), Yamnaya (Repin) and Poltavka (Sosnovka) contexts, dated to about 3500 to 2600 bc (Figs. 2a, b, 3a). Additionally, genetic
continuity with DOM2 was rejected for all horses predating about 2200 bc, especiallythose from the NEO-ANA group (Supplementary Table 2), except for two late Yamnaya specimens from approximately 2900 to 2600 bc (Turganik (TURG)), located further east than the western lower Volga-Don region (Figs. 2a, b, 3a). These may therefore have
provided some of the direct ancestors of DOM2 horses.Modelling of the DOM2 population with qpADM17, rotating18 all combinations of 2, 3 or 4 population donors, eliminated the possibility of a contribution from the NEO-ANA population, but indicated possible formation within the WE population, including agenetic contribution of approximately
95% from C-PONT and TURG horses (Supplementary Table 3). This was consistent with OrientAGraph19 modelling from nine lineages representing key ancestry combinations, which confirmed the absence of NEO-ANA genetic ancestry in DOM2 and confirmed DOM2 as a sister population to the C-PONT horses(Fig. 3b).Identifying discrete
populations and modelling admixture as single unidirectional pulses, however, was highly challenging given the extent of spatial genetic connectivity. Indeed, the typical DOM2 ancestry component was maximized in the C-PONT group, but declined sharply eastwards (TURG and Central Asia) in the third millennium bc asthe proportion of NEO-ANA
ancestry increased (Fig. 2a). This suggests a cline of genetic connectivity east of the Western Eurasia steppes and Central Asia, ruling out DOM2 ancestors further east than the western lower Volga-Don and Turganik. A similar genetic cline characterized the region located west of C-PONT, where the typical DOM2 ancestry component declined
steadily in the Dnieper steppes, Poland, Turkish Thrace and Hungary in the fifth to third millennia bc. This eliminates the possibility of DOM2 ancestors further west than C-PONT and the Dnieper steppes. Furthermore,patterns of spatial autocorrelations in the genetic data20 indicated Western Eurasia steppes as the most likely geographic location of
DOM2 ancestors (Fig.
3c). Combined, our results demonstrate that DOM2 ancestors lived in the Western Eurasia steppes, especially the lower Volga-Don, but not in Anatolia, during the late fourth and early thirdmillennia bc.Analyses of ancient human genomes have revealed a massive expansion from the Western Eurasia steppes into Central and Eastern Europe during the
third millennium bc, associated with the Yamnaya culture8,9,11,12,21. This expansion contributed at least two thirds of steppe-related ancestry to populations of the Corded Ware complex(CWC) around 2900 to 2300 bc8. The role of horses in this expansion remained unclear, as oxen could have pulled Yamnaya heavy, solid-wheeled wagons7,22. The
genetic profile of horses from CWC contexts, however, almost completely lacked the ancestry maximized in DOM2 and Yamnaya horses (TURG and Repin) (Figs. 1e, f, 2a, b) and showed nodirect connection with the WE group, including both C-PONT and TURG, in OrientAGraph modelling (Fig. 3b, Extended Data Fig. 5).The typical DOM2 ancestry was
also limited in pre-CWC horses from Denmark, Poland and Czechia, associated with the Funnel Beaker and early Pitted Ware cultures (FB/PWC, FB/POL and ENEO-CZE, respectively). DOM2 ancestry reached a maximum 12.5% in one Hungarian horse dated to the mid-third millennium bc and associated with theSomogyvár-Vinkovci Culture
(CAR05_Hun_m2458). qpAdm17 modelling indicated that its DOM2 ancestry was acquired following gene flow from southern Thrace (Kan22_Tur_m2386), but not from the Dnieper steppes (Ukr11_Ukr_m4185) (Supplementary Table 3). Combined with the lack of increased horse dispersal during the early third millennium bc (Fig. 2b, Extended Data
Fig. 3b), these results suggest that DOM2 horses did not accompany the steppe pastoralist expansion north of the Carpathians.By around 2200–2000 bc, the typical DOM2 ancestry profile appeared outside the Western Eurasia steppes in