Figure - available from: Nature
This content is subject to copyright. Terms and conditions apply.
Relative error rates Missing mutations per site in a test genome (y-axis), relative to a modern Icelandic horse (P5782_Ice_Modern) used as high-quality reference. a) for the full data set and SNP_pval 0. b) for the downsampled data set and SNP_val 0.
Source publication
Horses revolutionized human history with fast mobility¹. However, the timeline between their domestication and their widespread integration as a means of transport remains contentious2–4. Here we assemble a collection of 475 ancient horse genomes to assess the period when these animals were first reshaped by human agency in Eurasia. We find that re...
Citations
... In the past, different parts of the world were connected with varying degrees of intensity over several millennia, such that in a loose way, globalisation is not a new phenomenon. Horse riding dates to c. 3000 BCE and while genetic evidence suggests that horses were first domesticated on the Pontic Caspian Steppes c. 2200 BCE (Librado et al. 2024), within 300 years, similar horses had spread as far as Spain as result of migration from the steppes. Long-distance movement of material culture similarly points to connections we associate with globalisation. ...
Long camel caravans of merchants trekking over sand dunes silhouetted by a setting sun form a common image trope used to promote tourism to China's far west. In these tourism brochures, posters, and web images for travel to Ningxia, Gansu, and Xinjiang, we see a romantic imagining of the silk roads. The Silk Road was not a road like those of the Romans paved with stones, let alone a modern tarmac highway. It was a network of connections that fanned out across Eurasia through deserts, grasslands, and mountains. Nor was it just an overland route. Sea routes were important from the early first millennium and later moved more goods and more people than the overland route. And while silk remained important throughout the centuries, it was only one among an array of goods that were produced and traded across Eurasia. People and goods, knowledge and technology, culture and religions, and crops, animals, and diseases, have moved in each direction from well before historical times. Archaeology, archaeological science, and geography allow us to reconstruct those flows for periods or places for which written documents do not exist or have not survived. The Silk Road is a modern term, coined by the German geographer and explorer Baron Ferdinand von Richthofen in 1877. It was not used by those who lived in the oasis towns of Central Asia or the nomadic pastoralists of the steppes. Nor was it used by the rulers, officials, and merchants of the literate settled empires in China, West Asia, or Europe. Richthofen used both the singular form (Seidenstrasse) and the plural (Seidenstrassen) for what he defined as the route linking Han Dynasty China to Rome at the end of the first millennium BCE. Nevertheless, in his imagination , the concept was to motivate building trans-Eurasian railways and highways that could join Europe and China. When Richthofen coined the term, the Silk Road or silk roads were part of the European imperial imagination of the late nineteenth century. In its most recent political appropriation, the Silk Road is again very much an ideological concept. It is a wrapper for the designs and aspirations of the contemporary Chinese government, which like the Han, Tang, Ming, and Qing dynasties before it, seeks to project political and economic power into its peripheries in Asia and beyond. Some scholars have recently criticised or even dismissed the idea of silk roads, because the literature deploying this term has privileged the sedentary and literate empires at either end of Eurasia, thereby ignoring the vitality of the contributions-if
... For instance, the early trajectory of Christianity in Central and East Asia, from Persia through Sogdian territories, and then into Xinjiang and Gansu, suggests the relevance of areas along the silk roads in the global history of Christianity during the millennium before the Jesuit missions. Patterns in the movement of animals, crops, and material cultures since prehistoric times (Hsu et al. 2016;Zhou et al. 2020;Librado et al. 2024) are equally significant to problematise the idea of silk roads. In other words, from our volume, a new image of the silk roads emerges, which encourages scholars working on different periods to recognise the complexity of Eurasian networks and the polycentric origins of changes in their structures, nodes, and flows. ...
... Evidence for horse husbandry, however, comes from laboratory analysis revealing a number of notable results. These include isotope ratios from lipid residues on ceramics argued to represent horse milk, post-hole features with elevated phosphorous levels interpreted as corrals, dental pathologies associated with bridle mouthpiece use, genomic indicators of selective pressures associated with coat color, and shortened inter-generational intervals suggestive of reproductive control (Librado et al., 2024;Outram & Bogaard, 2019;Outram, 2023). ...
... All modern domesticated horses arise from this secondary lineage founded in the lower Volga-Don region of the Pontic-Caspian steppe and are referred to by geneticists as DOM2 horses (Librado et al., 2021). The early to mid-3rd millennium BCE marks the earliest available evidence for the emergence of the DOM2 lineage (Librado et al., 2021:635;Librado et al., 2024). However, debate persists over the scope of horse use during this very early period. ...
... 2100-1800 BCE) attesting clearly to both horse domesticated status and their importance to steppe communities for traction, transportation, and prestigious display (Librado et al., 2021;Koryakova & Epimakhov, 2007:79-80;Hanks et al., 2007). Based on spatial and demographic modeling the dispersal of DOM2 horses began from ca. 2200 BCE and proceeded rapidly, eventually replacing most other horse lineages across Eurasia (Librado et al., 2024). The primary technology responsible for the overwhelming interest in and uptake of horses, as evidenced from Europe, North Africa, the Near East, and eastward to China, was the Sintashta spoke-wheeled chariot design (Anthony, 2007;Librado et al., 2021). ...
The timing, geographical pathways, and earliest uses for horses (Equus caballus) in East Asia have long been a scientific puzzle vigorously debated by scholars with different backgrounds across diverse disciplinary fields. Over the past two decades, a wide range of high-resolution evidence has been presented documenting the multi-regional dispersion of domesticated horses, but questions still abound, especially in the eastern regions of Eurasia. This study provides a rigorous critique of this body of evidence and advances hypotheses as to when and where domesticated horses first arrived in Mongolia and China. We present the earliest ancient DNA evidence for the DOM2 genetic lineage of modern domesticated horses in Mongolia and evaluate a comprehensive dataset of radiocarbon dates for East Asian domesticated horses using a Bayesian statistical approach that is powerful but seldom utilized by archaeologists. Our genetic and chronological evidence demonstrates that horses far to the east in Mongolia, interred in monumental prone burial contexts, were domesticated and contemporaneous with the earliest horses documented in western regions of Mongolia and Xinjiang. These results both complicate and clarify major questions related to the occurrence of domesticated horses within the greater region and help to explain the development of horse culture(s) and knowledge on the eastern steppe and at the Late Shang capital of Yinxu.
... Evidence for horse husbandry, however, comes from laboratory analysis revealing a number of notable results. These include isotope ratios from lipid residues on ceramics argued to represent horse milk, post-hole features with elevated phosphorous levels interpreted as corrals, dental pathologies associated with bridle mouthpiece use, genomic indicators of selective pressures associated with coat color, and shortened inter-generational intervals suggestive of reproductive control (Librado et al., 2024;Outram & Bogaard, 2019;Outram, 2023). ...
... All modern domesticated horses arise from this secondary lineage founded in the lower Volga-Don region of the Pontic-Caspian steppe and are referred to by geneticists as DOM2 horses (Librado et al., 2021). The early to mid-3rd millennium BCE marks the earliest available evidence for the emergence of the DOM2 lineage (Librado et al., 2021:635;Librado et al., 2024). However, debate persists over the scope of horse use during this very early period. ...
... 2100-1800 BCE) attesting clearly to both horse domesticated status and their importance to steppe communities for traction, transportation, and prestigious display (Librado et al., 2021;Koryakova & Epimakhov, 2007:79-80;Hanks et al., 2007). Based on spatial and demographic modeling the dispersal of DOM2 horses began from ca. 2200 BCE and proceeded rapidly, eventually replacing most other horse lineages across Eurasia (Librado et al., 2024). The primary technology responsible for the overwhelming interest in and uptake of horses, as evidenced from Europe, North Africa, the Near East, and eastward to China, was the Sintashta spoke-wheeled chariot design (Anthony, 2007;Librado et al., 2021). ...
The timing, geographical pathways, and earliest uses for horses (Equus caballus) in East Asia have long been a scientific puzzle vigorously debated by scholars with different backgrounds across diverse disciplinary fields. Over the past two decades, a wide range of high-resolution evidence has been presented documenting the multi-regional dispersion of domesticated horses, but questions still abound, especially in the eastern regions of Eurasia. This study provides a rigorous critique of this body of evidence and advances hypotheses as to when and where domesticated horses first arrived in Mongolia and China. We present the earliest ancient DNA evidence for the DOM2 genetic lineage of modern domesticated horses in Mongolia and evaluate a comprehensive dataset of radiocarbon dates for East Asian domesticated horses using a Bayesian statistical approach that is powerful but seldom utilized by archaeologists. Our genetic and chronological evidence demonstrates that horses far to the east in Mongolia, interred in monumental prone burial contexts, were domesticated and contemporaneous with the earliest horses documented in western regions of Mongolia and Xinjiang. These results both complicate and clarify major questions related to the occurrence of domesticated horses within the greater region and help to explain the development of horse culture(s) and knowledge on the eastern steppe and at the Late Shang capital of Yinxu.
... In the past, different parts of the world were connected with varying degrees of intensity over several millennia, such that in a loose way, globalisation is not a new phenomenon. Horse riding dates to c. 3000 BCE and while genetic evidence suggests that horses were first domesticated on the Pontic Caspian Steppes c. 2200 BCE (Librado et al. 2024), within 300 years, similar horses had spread as far as Spain as result of migration from the steppes. Long-distance movement of material culture similarly points to connections we associate with globalisation. ...
... For instance, the early trajectory of Christianity in Central and East Asia, from Persia through Sogdian territories, and then into Xinjiang and Gansu, suggests the relevance of areas along the silk roads in the global history of Christianity during the millennium before the Jesuit missions. Patterns in the movement of animals, crops, and material cultures since prehistoric times (Hsu et al. 2016;Zhou et al. 2020;Librado et al. 2024) are equally significant to problematise the idea of silk roads. In other words, from our volume, a new image of the silk roads emerges, which encourages scholars working on different periods to recognise the complexity of Eurasian networks and the polycentric origins of changes in their structures, nodes, and flows. ...
... К сожалению, пока остаётся неясным, является ли это результатом одомашнивания местной породы лошади или следами охоты на ещё дикий вид. Как известно, одомашнивание современной лошади (DOM 2) произошло сравнительно недавно, и новейшие данные относят этот момент примерно к XXII в. до н. э. (Librado et al, 2024). То есть на 300 лет позже финала афанасьевской культуры. ...
The article presents modern data on the stages of development of horse use by man in the Palaeometal epoch on the example of archaeological sites of the Minusinsk basin. The beginning was made in the Afanasievo era and can probably be considered as periodic domestication or early stages of domestication. In the Okunev culture, the first experiments in training and wider use of horses are already taking place. The Andronovo culture and the early stages of the Karasuk period demonstrate the use of the horse as a draft animal in light carts and chariots. Finally, the end of the Palaeometal epoch is characterized by horseback riding, which had a very significant impact on the entire economic life of steppe pastoralists.
... Compared with mitochondrial DNA, the nuclear genome data provide significantly more information. Nuclear genome analysis allows to assess in detail the demographic history of extinct species, make assumptions about the possibility of ancient introgression, causes of species adaptation and extinction, directions of animal domestication, and etc (Barlow et al., 2021;Sharko et al., 2021;Dehasque et al., 2024;Librado et al., 2024). ...
The Panthera genus originated in Africa and subsequently spread to Eurasia, North America and South America. Species within this genus occupy dominant positions in ecological and food chains, but due to climatic fluctuations, changing ecosystems and various other ecological factors, several Panthera species have become extinct. The cave lion (Panthera spelaea Goldfuss) is one of the iconic species of Pleistocene megafauna, and its genetic research began around twenty years ago. Despite the information available so far, a number of issues regarding taxonomy, microevolutionary processes in populations, and the causes of extinction at the boundary between the Pleistocene and Holocene for this top predator remain unclear. This review aims to systematize the available data on cave lion genetics and develop new directions for studying this animal, as well as considering the genetic prospects for lion survival in the Holocene period.
... In equids, most of the ancient DNA work has, however, focused on elucidating the domestication history of the horse (reviewed in Orlando et al., 2021), and the donkey (Todd et al., 2022), in which palaeogenomic archives from several hundreds of ancient animals have been characterized (Librado et al., 2021;Librado et al., 2024). While 26 ancient genomes and two almost complete mitochondrial genomes have been sequenced for E. ovodovi (Cai et al., 2022;Druzhkova et al., 2017;Vilstrup et al., 2013), the genetic characterization of E. hydruntinus has been limited to a handful of partial or pseudo-complete mitochondrial genomes (Bennett et al., 2017;Catalano et al., 2020;Orlando et al., 2006Orlando et al., , 2009, and three nuclear genomes (Özkan et al., 2024).Moreover, ancient genome data for their closest phylogenetic relatives, the Asian asses (E. ...
... mestic horses in Mongolia available so far remains in the Bronze Age, some ~3.2-3.1 Kya(Fages et al., 2019;Librado et al., 2024), in line with archaeological evidence for hippophagy(Taylor et al., 2018), horse milking(Ventresca Miller et al., 2022) and bridling(Taylor et al., 2017) around ~3.4-3.2 Kya, ~3.2 Kya and ~ 3.2-3.0 ...
The extremely rich palaeontological record of the horse family, also known as equids, has provided many examples of macroevolutionary change over the last ~55 Mya. This family is also one of the most documented at the palaeogenomic level, with hundreds of ancient genomes sequenced. While these data have advanced understanding of the domestication history of horses and donkeys, the palaeogenomic record of other equids remains limited. In this study, we have generated genome‐wide data for 25 ancient equid specimens spanning over 44 Ky and spread across Anatolia, the Caucasus, Central Asia and Mongolia. Our dataset includes the genomes from two extinct species, the European wild ass, Equus hydruntinus , and the sussemione Equus ovodovi . We document, for the first time, the presence of sussemiones in Mongolia and their survival around ~3.9 Kya, a finding that should be considered when discussing the timing of the first arrival of the domestic horse in the region. We also identify strong spatial differentiation within the historical ecological range of Asian wild asses, Equus hemionus , and incomplete reproductive isolation in several groups yet considered as different species. Finally, we find common selection signatures at ANTXR2 gene in European, Asian and African wild asses. This locus, which encodes a receptor for bacterial toxins, shows no selection signal in E. ovodovi , but a 5.4‐kb deletion within intron 7. Whether such genetic modifications played any role in the sussemione extinction remains unknown.
... The earliest evidence of horse domestication dates back to only 5500 years ago, from the Eneolithic site of Botai, in Northern Kazakhstan, where several independent lines of evidence point towards horse husbandry, although domestication at Botai remains a debated hypothesis (Outram et al., 2009;Taylor & Barr on-Ortiz, 2021;Outram, 2023;Librado et al., 2024). Importantly, horses managed at Botai do not represent the ancestors of present-day domesticated horses but instead those of Przewalski's horses (Gaunitz et al., 2018). ...
... Humans have thus attempted to take control over the reproduction and care of horses on multiple occasions, once more highlighting their interest in creating close interactions with horses. This is also supported by the swift spread of domesticated horses from the Western Eurasian steppes starting around 4000 years ago, suggesting that human communities across the entire Eurasian continent adopted the use of horses within only a few centuries (Librado et al., 2021(Librado et al., , 2024. ...
... Prior to extensive extinction events, there was great functional diversity in wild equids, herein within Equus ferus (Hulbert, 1993;Mihlbachler et al., 2011). Following local or total extinctions, the only allegedly wild caballine equid (sub)species that is still extant is the Przewalski's horsealthough its classification as truly wild or feral is under debate (Outram et al., 2009;Taylor & Barr on-Ortiz, 2021;Outram, 2023;Librado et al., 2024). Discovered in the late 19th century in the steppes of Central Asia, Przewalski's horses became extinct in the wild in the 1960s due to habitat loss and overhunting, but were saved from extinction thanks to massive conservation efforts and worldwide collaboration between zoos (Turghan, Jiang & Niu, 2022). ...
In recent decades, the integration of horses (Equus ferus) in European rewilding initiatives has gained widespread popularity due to their potential for regulating vegetation and restoring natural ecosystems. However, employing horses in conservation efforts presents important challenges, which we here explore and discuss. These challenges encompass the lack of consensus on key terms inherent to conservation and rewilding, the entrenched culture and strong emotions associated with horses, low genetic diversity and high susceptibility to hereditary diseases in animals under human selection, as well as insufficient consideration for the social behaviour of horses in wild-living populations. In addition, management of wild-living horses involves intricate welfare, ethics and legislative dimensions. Anthropocentric population-control initiatives may be detrimental to horse group structures since they tend to prioritise individual welfare over the health of populations and ecosystems. To overcome these challenges, we provide comprehensive recommendations. These involve a systematic acquisition of genetic information, a focus on genetic diversity rather than breed purity and minimal veterinary intervention in wild-living populations. Further, we advise allowing for natural top-down and bottom-up control-or, if impossible, simulating this by culling or non-lethal removal of horses-instead of using fertility control for population management. We advocate for intensified collaboration between conservation biologists and practitioners and enhanced communication with the general public. Decision-making should be informed by a thorough understanding of the genetic makeup, common health issues and dynamics, and social behaviour in wild-living horse populations. Such a holistic approach is essential to reconcile human emotions associated with horses with the implementation of conservation practices that are not only effective but also sustainable for the long-term viability of functional, biodiverse ecosystems, while rehabilitating the horse as a widespread wild-living species in Europe.
