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 corralling2–4 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-Don region, 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 equestrian material 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 Europe around 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.

BackgroundClimatic and environmental fluctuations as well as anthropogenic pressure have led to the extinction of much of Europe’s megafauna. The European bison or wisent (Bison bonasus), one of the last wild European large mammals, narrowly escaped extinction at the onset of the 20th century owing to hunting and habitat fragmentation. Little is known, however, about its origin, evolutionary history and population dynamics during the Pleistocene.ResultsThrough ancient DNA analysis we show that the emblematic European bison has experienced several waves of population expansion, contraction, and extinction during the last 50,000 years in Europe, culminating in a major reduction of genetic diversity during the Holocene. Fifty-seven complete and partial ancient mitogenomes from throughout Europe, the Caucasus, and Siberia reveal that three populations of wisent (Bison bonasus) and steppe bison (B. priscus) alternately occupied Western Europe, correlating with climate-induced environmental changes. The Late Pleistocene European steppe bison originated from northern Eurasia, whereas the modern wisent population emerged from a refuge in the southern Caucasus after the last glacial maximum. A population overlap during a transition period is reflected in ca. 36,000-year-old paintings in the French Chauvet cave. Bayesian analyses of these complete ancient mitogenomes yielded new dates of the various branching events during the evolution of Bison and its radiation with Bos, which lead us to propose that the genetic affiliation between the wisent and cattle mitogenomes result from incomplete lineage sorting rather than post-speciation gene flow.ConclusionThe paleogenetic analysis of bison remains from the last 50,000 years reveals the influence of climate changes on the dynamics of the various bison populations in Europe, only one of which survived into the Holocene, where it experienced severe reductions in its genetic diversity. The time depth and geographical scope of this study enables us to propose temperate Western Europe as a suitable biotope for the wisent compatible with its reintroduction.Electronic supplementary materialThe online version of this article (doi:10.1186/s12915-016-0317-7) contains supplementary material, which is available to authorized users.

Background: Climatic and environmental fluctuations as well as anthropogenic pressure have led to the extinction of much of Europe's megafauna. The European bison or wisent (Bison bonasus), one of the last wild European large mammals, narrowly escaped extinction at the onset of the 20th century owing to hunting and habitat fragmentation. Little is known, however, about its origin, evolutionary history and population dynamics during the Pleistocene.

Recent excavations at the cave site of Roc de Marsal (in the Dordogne region of SW France) have yielded several Mousterian assemblages rich in well-preserved faunal remains. The Layer 4 faunal assemblage, associated with a rich Quina Mousterian occupation, provides an opportunity to investigate Neandertal prey selection, transport decisions and reindeer carcass processing strategies. One of the most striking characteristics of the Roc de Marsal Layer 4 faunal assemblage is an apparent lack of spongy parts that cannot be explained by taphonomic processes or carnivore activities. Limb elements were selectively transported to the site, where they underwent considerable further processing, resulting in a notably high degree of fragmentation of almost all elements. Comparison of reindeer carcass processing strategies employed by Neandertals during the Quina Mousterian with those identified for the later periods of the Upper Palaeolithic gives rise to some unexpected interpretations of Neandertal consumption behaviours. We show that this kind of intensive fragmentation is associated with bone grease rendering in ethnographic and Upper Palaeolithic contexts, in most such cases with the aid of fire. However, evidence of fire is remarkably scarce in Roc de Marsal Layer 4. Layer 4 could reflect a systematic Neandertal practice of sucking and chewing on the spongy portions of bones in order to extract as much of the highly nutritious bone grease as possible. Additionally, Roc de Marsal Layer 4 shows examples of use of animal resources for nonalimentary purposes by Neandertals. The abundance of cutmarks on the anterior surfaces of reindeer metapodials seems to indicate the acquisition of tendons, and bones were frequently used as retouchers.