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10-20-2021, 10:09 PM
https://media.springernature.com/full/springer-static/image/art%3A10.1038%2Fs41586-021-04018-9/MediaObjects/41586_2021_4018_Fig3_HTML.png?as=webp
https://www.nature.com/articles/s41586-021-04018-9#MOESM1
Abstract: Domestication of horses fundamentally transformed long-range mobility and warfare. However, modern domesticated breeds do not descend from the earliest domestic horse lineage associated with archaeological evidence of bridling, milking and corralling at Botai, Central Asia around 3500 bc. Other longstanding candidate regions for horse domestication, such as Iberia and Anatolia, 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 association between horseback riding and the massive expansion of Yamnaya steppe pastoralists into Europe around 3000 bc driving the spread of Indo-European languages. This contrasts with the scenario in Asia where Indo-Iranian languages, chariots and horses spread together, following the early second millennium bc Sintashta culture.
Expansion of steppe-related pastoralism
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 no direct 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 the Somogyvár-Vinkovci Culture (CAR05_Hun_m2458). qpAdm 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 22002000 bc, the typical DOM2 ancestry profile appeared outside the Western Eurasia steppes in Bohemia (Holubice), the lower Danube (Gordinesti II) and central Anatolia (Acemhöyük), spreading across Eurasia shortly afterwards, eventually replacing all pre-existing lineages (Fig 2c, Extended Data Fig. 3c). Eurasia became characterized by high genetic connectivity, supporting massive horse dispersal by the late third millennium and early second millennium bc. This process involved stallions and mares, indicated by autosomal and X-chromosomal variation (Extended Data Fig. 3d), and was sustained by explosive demographics apparent in both mitochondrial and Y-chromosomal variation (Extended Data Fig. 3e, f). Altogether, our genomic data uncover a high turnover of the horse population in which past breeders produced large stocks of DOM2 horses to supply increasing demands for horse-based mobility from around 2200 bc.
Of note, the DOM2 genetic profile was ubiquitous among horses buried in Sintashta kurgans together with the earliest spoke-wheeled chariots around 20001800 bc (Extended Data Fig. 6). A typical DOM2 profile was also found in Central Anatolia (AC9016_Tur_m1900), concurrent with two-wheeled vehicle iconography from about 1900 bc25,26. However, the rise of such profiles in Holubice, Gordinesti II and Acemhöyük before the earliest evidence for chariots supports horseback riding fuelling the initial dispersal of DOM2 horses outside their core region, in line with Mesopotamian iconography during the late third and early second millennia bc27. Therefore, a combination of chariots and equestrianism is likely to have spread the DOM2 diaspora in a range of social contexts from urban states to dispersed decentralized societies.
https://www.nature.com/articles/s41586-021-04018-9#MOESM1
Abstract: Domestication of horses fundamentally transformed long-range mobility and warfare. However, modern domesticated breeds do not descend from the earliest domestic horse lineage associated with archaeological evidence of bridling, milking and corralling at Botai, Central Asia around 3500 bc. Other longstanding candidate regions for horse domestication, such as Iberia and Anatolia, 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 association between horseback riding and the massive expansion of Yamnaya steppe pastoralists into Europe around 3000 bc driving the spread of Indo-European languages. This contrasts with the scenario in Asia where Indo-Iranian languages, chariots and horses spread together, following the early second millennium bc Sintashta culture.
Expansion of steppe-related pastoralism
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 no direct 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 the Somogyvár-Vinkovci Culture (CAR05_Hun_m2458). qpAdm 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 22002000 bc, the typical DOM2 ancestry profile appeared outside the Western Eurasia steppes in Bohemia (Holubice), the lower Danube (Gordinesti II) and central Anatolia (Acemhöyük), spreading across Eurasia shortly afterwards, eventually replacing all pre-existing lineages (Fig 2c, Extended Data Fig. 3c). Eurasia became characterized by high genetic connectivity, supporting massive horse dispersal by the late third millennium and early second millennium bc. This process involved stallions and mares, indicated by autosomal and X-chromosomal variation (Extended Data Fig. 3d), and was sustained by explosive demographics apparent in both mitochondrial and Y-chromosomal variation (Extended Data Fig. 3e, f). Altogether, our genomic data uncover a high turnover of the horse population in which past breeders produced large stocks of DOM2 horses to supply increasing demands for horse-based mobility from around 2200 bc.
Of note, the DOM2 genetic profile was ubiquitous among horses buried in Sintashta kurgans together with the earliest spoke-wheeled chariots around 20001800 bc (Extended Data Fig. 6). A typical DOM2 profile was also found in Central Anatolia (AC9016_Tur_m1900), concurrent with two-wheeled vehicle iconography from about 1900 bc25,26. However, the rise of such profiles in Holubice, Gordinesti II and Acemhöyük before the earliest evidence for chariots supports horseback riding fuelling the initial dispersal of DOM2 horses outside their core region, in line with Mesopotamian iconography during the late third and early second millennia bc27. Therefore, a combination of chariots and equestrianism is likely to have spread the DOM2 diaspora in a range of social contexts from urban states to dispersed decentralized societies.