The origins of domestic horses

[Anfänger]

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

Abstract

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-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 BC8,9driving 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.

Link: https://www.nature.com/articles/s41586-021-04018-9

Apparently domestic horses were spread by Sintastha and Indo-Iranians. It wasn't Yamnaya as many have thought.

[olivand]

I would have thought that horses were more relevant to roaming pastoralists than to sedentary farmers such as the EEFs who spread west from Anatolia before the late Bronze-Iron Age yamnaya 'invaders', a theory that will be hard to crack until more evidence comes to light. An interesting development though - thanks for the link which I will read!

[johen]

I think this is a third big horse paper:

Yamnaya horses at Repin and Turganik carried more DOM2 genetic affinity than presumably wild horses from hunter-gatherer sites of the sixth millennium BC (NEO-NCAS, from approximately 5500–5200 BC), which may suggest early horse management and herding practices. Regardless, Yamnaya pastoralism did not spread horses far outside their native range, similar to the Botai horse domestication, which remained a localized practice within a sedentary settlement system2,36. The globalization stage started later, when DOM2 horses dispersed outside their core region, first reaching Anatolia, the lower Danube, Bohemia and Central Asia by approximately 2200 to 2000 BC, then Western Europe and Mongolia soon afterwards, ultimately replacing all local populations by around 1500 to 1000 BC. This process first involved horseback riding, as spoke-wheeled chariots represent later technological innovations, emerging around 2000 to 1800 BC in the Trans-Ural Sintashta culture7. The weaponry, warriors and fortified settlements associated with this culture may have arisen in response to increased aridity and competition for critical grazing lands, intensifying territoriality and hierarchy37. This may have provided the basis for the conquests over the subsequent centuries that resulted in an almost complete human and horse genetic turnover in Central Asian steppes11,21. The expansion to the Carpathian basin38, and possibly Anatolia and the Levant, involved a different scenario in which specialized horse trainers and chariot builders spread with the horse trade and riding. In both cases, horses with reduced back pathologies and enhanced docility would have facilitated Bronze Age elite long-distance trade demands and become a highly valued commodity and status symbol, resulting in rapid diaspora. We, however, acknowledge substantial spatiotemporal variability and evidential bias towards elite activities, so we do not discount additional, harder to evidence, factors in equine dispersal.

Our results also have important implications for mechanisms underpinning two major language dispersals. The expansion of the Indo-European language family from the Western Eurasia steppes has traditionally been associated with mounted pastoralism, with the CWC serving as a major stepping stone in Europe39,40,41. However, while there is overwhelming lexical evidence for horse domestication, horse-drawn chariots and derived mythologies in the Indo-Iranian branch of the Indo-European family, the linguistic indications of horse-keeping practices at the deeper Proto-Indo-European level are in fact ambiguous42 (Supplementary Discussion) . The limited presence of horses in CWC assemblages43 and the local genetic makeup of CWC specimens reject scenarios in which horses were the primary driving force behind the initial spread of Indo-European languages in Europe44. By contrast, DOM2 dispersal in Asia during the early-to-mid second millennium BC was concurrent with the spread of chariotry and Indo-Iranian languages, whose earliest speakers are linked to populations that directly preceded the Sintashta culture11,12,45. We thus conclude that the new package of chariotry and improved breed of horses, including chestnut coat colouration documented both linguistically (Supplementary Discussion) and genetically (Extended Data Fig. 8), transformed Eurasian Bronze Age societies globally within a few centuries after about 2000 BC.

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 was obtained from rworldmap46.

However, 1st research paper said that sintashta horse is related to arctic horse. So next time the horse/cattle bones in Ural east should be included, where they had contacted south east aral sea from mesolithic to eneolithic.

The 14 ancient genomes reported here have strong implications for the horse domestication process. First, it has recently been discovered that a now-extinct lineage of wild horses existed in the Arctic until at least ~5.2 ka and significantly contributed to the genetic makeup of present-day domesticates (14,15). The timing of the underlying admixture event(s) is, however, unknown. Using D statistics, we confirmed that this extinct lineage shared more derived polymorphisms with the Sintashta and especially Scythian horses than with present-day domesticates (Fig. 2

. The domestic horse lineage, thus, experienced a net loss of archaic introgressed tracts within the past ~2.3 ky.

[bicicleur 2]

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

Abstract

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-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 BC8,9driving 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.

Link: https://www.nature.com/articles/s41586-021-04018-9

Apparently domestic horses were spread by Sintastha and Indo-Iranians. It wasn't Yamnaya as many have thought.

The Yamna didn't spread any further than between the Carpathian basin and the Altaï Mts.
The R1b-L51 which spread in Europe since 4,5 ka were not Yamna descendants despite being similar in autosomal DNA.

[olivand]

Thanks for these findings which suggest cultural diffusion of horse domestication rather than demic diffusion of horse riding pastoralists, and their horses. Focusing on male ancestry only, I was under the impression that my L151 ancestor traced back to early CWC of around 3000BC in modern day North Eastern Ukraine/Southern Poland and quickly diversified in west central Europe over a period of several hundred years. Certainly the spread of R1b across western Europe was very rapid and pretty much wiped out earlier Y-haplogroups such as G2a and I2a prevalent before the late Bronze Age. However Bell Beaker folk and their culture were contemporaneous with CWC and expanded from the Atlantic coast eastwards spreading culture and genetics including horse domestication which these papers appear to show preceded the Yamnaya.

[Anfänger]

The Yamna didn't spread any further than between the Carpathian basin and the Altaï Mts.
The R1b-L51 which spread in Europe since 4,5 ka were not Yamna descendants despite being similar in autosomal DNA.

I agree, with the samples and data available today, it is unlikely that Yamnaya are the ancestors of R1b-L51 in Western Europe but there might R1b-L51 in unsampled Yamnaya burials, i think particularly those from the western Yamnaya horizon in Ukraine. R1b-L51 and R1b-Z2103 are brother clades so it could be that they share a common origin that is older than Yamnaya, maybe from Repin culture, who knows.

[bicicleur 2]

I agree, with the samples and data available today, it is unlikely that Yamnaya are the ancestors of R1b-L51 in Western Europe but there might R1b-L51 in unsampled Yamnaya burials, i think particularly those from the western Yamnaya horizon in Ukraine. R1b-L51 and R1b-Z2103 are brother clades so it could be that they share a common origin that is older than Yamnaya, maybe from Repin culture, who knows.

yes, there is no doubt that they had a common ancestor, but this common ancestor is older than Yamnaya
the common ancestor R1b-M269 probably lived in Khvalynsk, or even more likely in Repin culture

[Angela]

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

Abstract

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-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 BC8,9driving 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.

Link: https://www.nature.com/articles/s41586-021-04018-9

Apparently domestic horses were spread by Sintastha and Indo-Iranians. It wasn't Yamnaya as many have thought.

Well, that's not quite how I interpreted the abstract, although I'll have to read the whole paper.

The Sintashta culture didn't spread horses into Europe. They "came" from Europe and then spread into Asia with their horses and their language.

I steadfastly maintained for years that horse mounted Corded Ware men did not come into Europe brandishing their bronze swords and spreading their genes and their language a la "Conan the Barbarian", or as "blonde cowboys of the steppe", to quote Eurogenes. :) You have to go 1000 years into the future and a lot further east for anything resembling that.


David Anthony was wrong, as I always said. There were barely any horses found in Corded Ware sites, and their metallurgy was very primitive. Heck, they barely had any copper, much less bronze, and the copper of poor quality, so the whole trope never made any sense. Those wagons were pulled by oxen, not horses.

Anthony was overlaying the culture of much later Iron Age steppe people onto Yamnaya and Corded Ware, and because they wanted to believe it, the majority willingly adopted it, without ever checking the archeology to see what these cultures were really like.

"And another one bites the dust", as a famous musician sang. :)


That idea has to go into the book I keep updating of all of Eurogenes' incorrect predictions and conclusions.

[Vallicanus]

Well, that's not quite how I interpreted the abstract, although I'll have to read the whole paper.

The Sintashta culture didn't spread horses into Europe. They "came" from Europe and then spread into Asia with their horses and their language.

I steadfastly maintained for years that horse mounted Corded Ware men did not come into Europe brandishing their bronze swords and spreading their genes and their language a la "Conan the Barbarian", or as "blonde cowboys of the steppe", to quote Eurogenes. :) You have to go 1000 years into the future and a lot further east for anything resembling that.


David Anthony was wrong, as I always said. There were barely any horses found in Corded Ware sites, and their metallurgy was very primitive. Heck, they barely had any copper, much less bronze, and the copper of poor quality, so the whole trope never made any sense. Those wagons were pulled by oxen, not horses.

Anthony was overlaying the culture of much later Iron Age steppe people onto Yamnaya and Corded Ware, and because they wanted to believe it, the majority willingly adopted it, without ever checking the archeology to see what these cultures were really like.

"And another one bites the dust", as a famous musician sang. :)


That idea has to go into the book I keep updating of all of Eurogenes' incorrect predictions and conclusions.

These weren't actual predictions by Davidski but wishful thinking and blind prejudice on his part.

[johen]

yes, there is no doubt that they had a common ancestor, but this common ancestor is older than Yamnaya
the common ancestor R1b-M269 probably lived in Khvalynsk, or even more likely in Repin culture

How about a relationship between yamna and afanasievo?

author said "yamnaya pastoralism did not spread horses far outside their native range, similar to the botai horse domestication, which remained a localized practice within a sedentary settlement system2,36."


Yamna mounds:


Afanasievo burials are in pits surrounded by circular or rectangular

[Malaparte]

This coheres with Robert Drews' timetable for militarized horse technology. I think it likely that comparatively peaceful movements of R1b peoples into Europe had been ongoing for some two millennia before more advanced warrior elites swept in circa 1600 BC. The long-term movements were on a mass scale and transformed the population; the elite takeovers were sudden, dramatic, and had a limited (but not negligible) genetic legacy, as exemplified by Mycenaean Greece.

[Angela]

These weren't actual predictions by Davidski but wishful thinking and blind prejudice on his part.

Well, I give even the devil his due; some of his ideas can be found in David Anthony, although Davidski vastly exaggerated even Anthony's incorrect conclusions.

[ihype02]

Davidski's rumours in my experience were not inherently wrong but usually lack a very important detail, especially the dates. I don't know about all of his "inaccurate" predictions but there was the case of some Mycenaean samples having nearly 30% Steppe that we haven't seen yet. But it turns out in fact those were the sample from EBA/MBA and not appropriate for the overall image of Mycenaeans because back then people were extremely divergent genetically.

[Johane Derite]

"Modern domestic horses ultimately replaced almost all other local populations as they expanded rapidly across Eurasia from about 2000 BC"


1000 years after Yamnaya expansion, so not with Yamnaya...

Fascinating.

Macurdy believed the royal Trojan lineage of Laomedon came from horsebreeders.

In the Iliad, it mentions the Dardanian Anchises stealing the breed of horses of Laomedon, which had been given to Troy by Zeus i.e. the highest God.


Was this the breed from lower Don-Volga region that expanded 2000BC?

Seems to have been quite important to them and for good reason, makes sense that it was so.


Was the Trojan dynasty also from the lower Don-Volga region then?

[Angela]

Davidski's rumours in my experience were not inherently wrong but usually lack a very important detail, especially the dates. I don't know about all of his "inaccurate" predictions but there was the case of some Mycenaean samples having nearly 30% Steppe that we haven't seen yet. But it turns out in fact those were the sample from EBA/MBA and not appropriate for the overall image of Mycenaeans because back then people were extremely divergent genetically.

A week before the paper on the Mycenaeans came out he predicted they would be blonde and blue eyed and virtually indistinguishable from Corded Ware.

What later samples showed that to be true? It's a rhetorical question; the answer is none. The levels of steppe in modern Greeks had to wait for the Slavs to arrive. That's one hell of a lag time.

My bet has always been that the Greek speakers were R1b Z-2103. We shall see.

As for the rest of the hypothesis, about which David Anthony was also incorrect, the arrival of the people from the steppe bringing the Indo-European languages was not by horse riding, metallurgy bringing, sword yielding Yamnaya or even Corded Ware people. That's the salient point.

[Angela]

This coheres with Robert Drews' timetable for militarized horse technology. I think it likely that comparatively peaceful movements of R1b peoples into Europe had been ongoing for some two millennia before more advanced warrior elites swept in circa 1600 BC. The long-term movements were on a mass scale and transformed the population; the elite takeovers were sudden, dramatic, and had a limited (but not negligible) genetic legacy, as exemplified by Mycenaean Greece.

I doubt even they were more advanced.

The Etruscans were steppe admixed, but they adopted the language and the cultural achievements of the culture they found in Italy. No culture across the Alps was their equal in iron metallurgy. Razib Khan goes so far, in his substack document, as to say that the Etruscans, from whom the Latins adopted so much, were the last great flowering of the culture of "Old Europe".

[ihype02]

A week before the paper on the Mycenaeans came out he predicted they would be blonde and blue eyed and virtually indistinguishable from Corded Ware.

What later samples showed that to be true? It's a rhetorical question; the answer is none. The levels of steppe in modern Greeks had to wait for the Slavs to arrive. That's one hell of a lag time.

My bet has always been that the Greek speakers were R1b Z-2103. We shall see.

As for the rest of the hypothesis, about which David Anthony was also incorrect, the arrival of the people from the steppe bringing the Indo-European languages was not by horse riding, metallurgy bringing, sword yielding Yamnaya or even Corded Ware people. That's the salient point.

Old Greeks whenever it was the Bronze Age or the Classical Period left a large amount of artwork that depicted themselves as a strictly Mediterranean-looking people. You don't need science to figure that out. Why would anyone think they were essentially blonde has some real serious issues with bias and objectivity.

I agree about R1b.

[bicicleur 2]

"Modern domestic horses ultimately replaced almost all other local populations as they expanded rapidly across Eurasia from about 2000 BC"


1000 years after Yamnaya expansion, so not with Yamnaya...

Fascinating.

Macurdy believed the royal Trojan lineage of Laomedon came from horsebreeders.

In the Iliad, it mentions the Dardanian Anchises stealing the breed of horses of Laomedon, which had been given to Troy by Zeus i.e. the highest God.


Was this the breed from lower Don-Volga region that expanded 2000BC?

Seems to have been quite important to them and for good reason, makes sense that it was so.


Was the Trojan dynasty also from the lower Don-Volga region then?

what about the legendary centaurs?
didn't they invade from the north?

[Anfänger]

Well, that's not quite how I interpreted the abstract, although I'll have to read the whole paper.

The Sintashta culture didn't spread horses into Europe. They "came" from Europe and then spread into Asia with their horses and their language.

I think this is a misunderstanding. I didn't say that they came from Asia but the origins of modern horses are from "DOM2" horses from the Volga-Don region 2200-2000BC. That's exactly the region and time of the Sintashta culture.

I steadfastly maintained for years that horse mounted Corded Ware men did not come into Europe brandishing their bronze swords and spreading their genes and their language a la "Conan the Barbarian", or as "blonde cowboys of the steppe", to quote Eurogenes. :) You have to go 1000 years into the future and a lot further east for anything resembling that.


David Anthony was wrong, as I always said. There were barely any horses found in Corded Ware sites, and their metallurgy was very primitive. Heck, they barely had any copper, much less bronze, and the copper of poor quality, so the whole trope never made any sense. Those wagons were pulled by oxen, not horses.

Anthony was overlaying the culture of much later Iron Age steppe people onto Yamnaya and Corded Ware, and because they wanted to believe it, the majority willingly adopted it, without ever checking the archeology to see what these cultures were really like.

"And another one bites the dust", as a famous musician sang. :)


That idea has to go into the book I keep updating of all of Eurogenes' incorrect predictions and conclusions.

Yes, you are right. David Anthony was wrong with his idea of "horse riding hordes" conquering Europe, so was Davidski.

[Johane Derite]

I think this is a misunderstanding. I didn't say that they came from Asia but the origins of modern horses are from "DOM2" horses from the Volga-Don region 2200-2000BC. That's exactly the region and time of the Sintashta culture.

This is the region that DOM2 comes from. Sintashta is peripheral to this region at the most eastern.

This falls within the late "Catacomb culture" region

[Johane Derite]

what about the legendary centaurs?
didn't they invade from the north?

Peake, who she refers to, made that association explicitly.

Referencing him, she mentions how this western expansion of "southern Russian" steppemen at the ending of the third millenium bc / beginning of the second millenium BC (perfectly matches the horse breed expansion) was responsible for the "destruction" of the Tripolye culture (or pushing it west).

She then mentions that something of the Tripolye culture must have been picked up by them since after this pottery like that of Tripolye appears in Thessally, Troy, and Caicus...

I wonder if this end of Tripolye can be responsible for E-V13 introduction into IE groups, either by pushing some west into carpathians, or integrating some specialists.

[kingjohn]

So buttom line davidski was right or wrong in his predictions ?

[Anfänger]

This is the region that DOM2 comes from. Sintashta is peripheral to this region at the most eastern.

This falls within the late "Catacomb culture" region

For Asia the situation is clear according to this paper. Horses and chariots were spread at the same time from Sintashta culture:

Of note, the DOM2 genetic profile was ubiquitous among horses buried in Sintashta kurgans together with the earliest spoke-wheeled chariots around 2000–1800 BC7,9,23,24 (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.
...
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 societies28.

But MAYBE you are right for the time period 2200-2000BC when there wasn't any chariots and horseback riding was the only use for horses. There we need horse ancient DNA to prove an origin from late Catacomb culture. Still you have to account for the spread of chariots into Europe which is definitely from Sintastha(Indo-Iranians):

The globalization stage started later, when DOM2 horses dispersed outside their core region, first reaching Anatolia, the lower Danube, Bohemia and Central Asia by approximately 2200 to 2000 BC, then Western Europe and Mongolia soon afterwards, ultimately replacing all local populations by around 1500 to 1000 BC. This process first involved horseback riding, as spoke-wheeled chariots represent later technological innovations, emerging around 2000 to 1800 BC in the Trans-Ural Sintashta culture7. The weaponry, warriors and fortified settlements associated with this culture may have arisen in response to increased aridity and competition for critical grazing lands, intensifying territoriality and hierarchy37. This may have provided the basis for the conquests over the subsequent centuries that resulted in an almost complete human and horse genetic turnover in Central Asian steppes11,21. The expansion to the Carpathian basin38, and possibly Anatolia and the Levant, involved a different scenario in which specialized horse trainers and chariot builders spread with the horse trade and riding.
...
However, while there is overwhelming lexical evidence for horse domestication, horse-drawn chariots and derived mythologies in the Indo-Iranian branch of the Indo-European family, the linguistic indications of horse-keeping practices at the deeper Proto-Indo-European level are in fact ambiguous42

[Anfänger]

So buttom line davidski was right or wrong in his predictions ?

This time he was wrong :)

[Johane Derite]

For Asia the situation is clear according to this paper. Horses and chariots were spread at the same time from Sintashta culture:



But MAYBE you are right for the time period 2200-2000BC when there wasn't any chariots and horseback riding was the only use for horses. There we need horse ancient DNA to prove an origin from late Catacomb culture. Still you have to account for the spread of chariots into Europe which is definitely from Sintastha(Indo-Iranians):

Yes, for asia there seems to be no doubt that sintashta is the vector of horse spread.

Interestingly, Albanian only has isoglosses with Indo-Iranian branch in words related to Horse-breeding