The origins and spread of domestic horses from the Western Eurasian steppes.

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.

Morphology-based diagnostics of "protodogs." A commentary to Galeta et al., 2021, Anatomical Record, 304, 42-62, doi: 10.1002/ar.24500.

In a recent article in this journal, Galeta et al., (2020) discussed eight Pleistocene "protodogs" and seven Pleistocene wolves. Those "protodogs" had been diagnosed in earlier publications, based on skull morphology. We re-examined the Galeta et al. paper to offer comments on their observed outcomes, and the conclusion of presumed domestication. Of seven metrics that the authors used, five differed statistically between their two groups. However, from more elaborate studies, some of those same metrics had been rejected previously as not valid species-distinguishing traits. In this respect, we do accept cranium size and wider palate as species-distinguishing metrics. The physical size of their specimens was much larger than other archaeological specimens that have been accepted as dogs. Additionally, their sample size was small, compared to the number of available specimens, as shown from previous publications by the same group. Thus, we considered statistical differences that were found between groups in their study, and assessed whether the outcomes could have resulted from natural morphological variation. We examined a group of 73 dire wolves ((Aenocyon [Canis] dirus; Perri et al., 2021), using the same methods as used by Galeta et al., (2020). We could segregate two distinct morphological groups in our study, one having outcomes that were identical to the "protodogs" in Galeta et al. (2020). For the specimens of extinct dire wolves to segregate in the same way as the subjects from Galeta et al. indicates that natural variation probably was the driver of their observed outcomes, domestication being an unlikely assumption.