Of horses, camels and extinct lineages


After a rather long hiatus, and while we wait for the long anticipated final version of Narasimhan et al. (hopefully out very soon), here’s a quick post commenting on a few things that have been published lately.


Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series

Fages et al. 2019

A follow up to their previous paper, briefly commented on a previous post, that brings some new information about the history of domestic horses, though far from clarifying things it makes everything less clear opening new questions. Here is their graphical abstract:

And the key findings regarding early history of domestication:

  • Two now-extinct horse lineages lived in Iberia and Siberia some 5,000 years ago
  • Iberian and Siberian horses contributed limited ancestry to modern domesticates

These newly found extinct lineages of early domestic horses add to the Botai ones, which didn’t go extinct but apparently went feral and survive in the form of Przewalski horses, as found in an even earlier paper (Gaunitz et al. 2018).

So let’s start to look at this puzzle and try to put a few pieces together. We have a horse lineage in Siberia living up until 5000 years ago that didn’t contribute any significant ancestry to the main (and for a long time the only) domestic lineage. Then we have another lineage further west, in Kazakhstan, that was domesticated c. 3500 BC but that also didn’t contribute much to main domestic horses. And finally we have a lineage in Iberia that we domesticated somewhere in the late Chalcolithic, and while it seems to have been used for a while as an early domestic horse in Europe it also ultimately went extinct without contributing much to the main domestic lineage.

A closer look at the data from this latter Iberian lineage brings us some additional interesting information: There is a sample from 2600 BC (note that this predates the arrival of R1b/steppe people) belonging to this domestic lineage (native to Iberia). But then another Iberian samples from 1900 BC (note that his is 500 years after the arrival of R1b/steppe Bell Beaker folk) is still of the same kind. Furthermore, a sample from Hungary c. 2100 BC shows some 12% admixture from this Iberian lineage (the rest of its makeup being from main domestic horses and another unknown lineage).

What this suggests is something rather surprising: R1b/steppe Bel Beakers don’t seem to have carried horses with them to Western Europe. Otherwise those would have largely replaced the local Iberian ones. Instead, it seems that Bell Beaker horses were those who they found in Iberia and might have traded with them all the way to Hungary. Indeed, this comes to reinforce the scarce evidence for the use of domestic horses in EBA steppe related cultures. Not much evidence in the Corded Ware Culture (for example, domestic horses only arrived to the east Baltic in the Iron Age, in spite of the area being occupied by early CWC population).

So where did domestic horses come from? That’s the min question right now. Traditionally, the Pontic-Caspian steppe has been the main candidate for it. However, the previous paper from this same team, based on climatic simulations and found remains, seemed to discard that area as suitable for horses at the time of probable domestication. Now this new study adds to that hypothesis by providing no evidence of the people coming from that area to Europe in the early 3rd mill. bringing their own domestic horses. Unfortunately, this study had 2 or 3 samples that could represent the Pontic-Caspian steppe wild horses, but they all yielded a very low amount of endogenous DNA to be included in any autosomal analysis. So that question remains open.

The thing is that the first good evidence of clear and extensive use of domestic horses of the main lineage comes from Sintashta, c. 2000 BC. So where did those horses came from? It’s unlikely that they were local, given the proximity with the Botai Culture area (and in west Siberia), where we know that very divergent lineages existed. To me it seems that they could have arrived there from anywhere. And it might not be that relevat after all. The place where the main domestic lineage originated might be rather unimportant, given that for all we know this domestication seems to have occurred quite later than first thought (closer to 2500 BC than to the previously suggested 3500 BC), and it may have been the Sintashta people the first ones who found a real use for them, no matter where they got them from. Whatever the case, it seems that the history of domestic horses is also turning out to be quite different from what was previously thought. Hopefully the next paper before the year’s end will shed some light in all of this.


Whole-genome sequencing of 128 camels across Asia provides insights into origin and migration of domestic Bactrian camels

Lian Ming et al. 2019 (preprint)

This one is a modern DNA study dealing with the domestication fo the Bactrian camel. Though I’m not sold on their idea that Bactrian camels were domesticated 10.000 year ago, the rest of the hypothesis looks good (as far as modern DNA can be informative). An image and a few excerpts summarise it well.

The origin of domestic dromedaries was recently revealed by world-wide sequencing of modern and ancient mitochondrial DNA (mtDNA), which suggested that they were at first domesticated in the southeast Arabian Peninsula [11]. However, the origin of domestic Bactrian camels is still a mystery. One intuitive possibility was the extant wild Bactrian camels were the progenitor of the domestic form, which were then dispersed from the Mongolian Plateau to west gradually [7, 12].


Another possible place of origin was Iran [1], where early skeletal remains of domestic Bactrain camels (around 2,500-3,000 BC) were discovered [14].


The wild Bactrian camels made also little contribution to the ancestry of domestic ones. Among the domestic Bactrian camels, those from Iran exhibited the largest genetic distance from others, and were the first population to separate in the phylogeny. Although evident admixture was observed between domestic Bactrian camels and dromedaries living around the Caspian Sea, the large genetic distance and basal position of Iranian Bactrian camels could not be explained by introgression alone. Taken together, our study favored the Iranian origin of domestic Bactrian camels, which were then immigrated eastward to Mongolia where the native wild Bactrian camels inhabited.


This scenario could well resolve the mystery why the wild and domestic Bactrian camels from the Mongolian Plateau have so large genetic distance.


Despite the insights gleaned from our data, it was important to note that the direct wild progenitor of domestic Bactrian camels were not found in Iran now, which may no longer exist.


In future work, sequencing of ancient genomes from camel fossils will add to the picture of their early domestication.

Not much to add, really. They looked with an extensive set of modern camel DNA at the two different scenarios proposed for domestication and concluded that their data favoured the Iranian one, in spite of wild camels no longer existing in Iran, contrary to Mongolia (where they exist but are very divergent from domestic ones, just like the horses).

We’ll wait for ancient DNA to either confirm or deny this.


The population history of northeastern Siberia since the Pleistocene

Sikora et al. 2019

I already commented quite extensively on the very interesting and well written paper when the preprint was out last year. Now it’s been finally published and hopefully the genomes will be made available (if they aren’t already). I was glad to see that the only problem I found with the preprint (their hypothesis about ANS surviving somewhere in Beringia and mixing with East Asian populations to form the Native American one, which seemed to me incompatible with the genetic data presented, due to Native Americans sharing more alleles with Malta and AfontovaGora3 than with the Yana samples) has been addressed in the final versions:

For both Ancient Palaeo-Siberians and Native Americans, ANS-related ancestry is more closely related to Mal’ta than to the Yana individuals (Extended Data Fig. 3f), which rejects the hypothesis that the Yana lineage contributed directly to later Ancient Palaeo-Siberians or Native American groups.

So unfortunately for them, these Yana population seems to have died out during the LGM, something not too surprising when such climatic event catches you in the Arctic.

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29 thoughts on “Of horses, camels and extinct lineages

  1. @raj

    Yes, even further reason to think that the 10.000 y.a. domestication date is out of reality.

    Interesting article by Talageri, which is in line with the latest archaeological findings from Sanauli. I hope they keep digging the areas east of it, around Yamuna river t see if they find more about that culture.

    Anyway we’ll have time to discuss this fairly soon when the final version of Narasimhan et al. is out.

  2. There’s a new Paleolithic sample from Crimea:

    The origin of the Gravettians: genomic evidence from a 36,000-year-old Eastern European

    Bennett et al. (preprint)

    The sample in question is from about 36.000 y.a. and has (probably) Y DNA C and mtDNA N1b. Autosomally it seems closer to Kostenki12 and Sunghir3 samples from European Russia and then to Central European samples like Vestonice16 and Pavlov1.

    It’s not any more related to Villabruna than to Tianyuan, or to MA-1 than to Ust-Ishim. So these Paleo-European populations seem to have had very little, if any, role in the formation of later WHG and ANE. However, more western WHGs that have some small amount of Paleo-European admixture (more related to Goyet-Q116) do have a slightly higher share with this new sample too.

  3. There’s also an interesting paper with high coverage modern genomes that brings some insights into population structure:

    Insights into human genetic variation and population history from 929 diverse genomes

    Bergström et al. (preprint)

    The paper deals in quite detail about archaic admixture in AMH populations. The main finding I think is that Neanderthal introgression seems to be from a single Neanderthal population (with a very small number of admixture events needed, around 2-4) while Denisovan introgression is much more complex and from divergent Denisoan populations. Again it seems that preservation bias has lead us to think that Neanderthals were the main Eurasian population before AMH came in, while it may have been Denisovans who were the largest population (living in warmer areas of southern Eurasia). At least they reached the Altai where we have been able to find them.

    Unrelated to the above, there’s an interesting finding regarding the bias in traditionally used SNPs arrays when it comes to African populations:

    Some of the f4-statistics commonly used to study population history and admixture (13) even shift sign when using array SNPs compared to when using all discovered SNPs, thus incorrectly reversing the direction of the ancestry relationship one would infer from the same set of genomes (for example: f4(BantuKenya,San;Mandenka,Sardinian) is positive (Z=2.9) using all variants but negative (Z=-3.11) when using commonly employed array sites).

    Also they find again Eurasian admixture in Yoruba (at least when compared to Mbuti). Gene flow which seems to be bidirectional between Africa and Eurasia and which is most likely responsible for the so called “Basal Eurasian” admixture.

    There are many other things to read in the paper, which is quite technical and detailed.

  4. From archaeological news, it seems that excavations in Iraqi Kurdistan have resumed after a long hiatus because of the situation in the area.

    Archaeologists uncover ancient palace of the Mittani Empire in Kurdistan Region


    They also found there clay tablets with cuneiform writings, so that may bring new information too. Hopefully we can get ancient DNA from this area too in the near future.

  5. Great to see Bennett et. al. managed to sequence the UP sample from Crimea
    And whilst sympathetic to their proposal, it’s not completely convincing, as Early Gravettian sites are lacking in eastern Europe (apart from this one site in Crimea). In fact, Gravettian occupation occurs in the late phase, and arguably arrives from central Europe. Nor does the samples from B.K. seem to be particularly ”ancestral” to late Paleolithic groups

    The mystery of the so-called ”Villabruna” cluster is still out. I wonder if Dzudzuana genomes will help shed light, when theyre released

  6. @Rob

    Though looking at the current data the mystery is not too deep regarding the Villabruna cluster. With Dzudzuana being aroun 80% (? from memory) Villabruna-related, and like later Wet Asians being roughly a similar mix of Villabruna-related + Basal Eurasian (with some Iran-related admixture too), it’s not too hard to imagine that the Villabruna cluster had to be somewhere in or near Anatolia on the way to the Balkans.

    And since we’re about animals in this post, the paper you shared about the Bison seems to contain some interesting clues.

  7. Alberto
    Yes I realise what the data is pointing to; but the problem is the dearth of late paleolithic sites in the east Balkans. There are enough in the west Balkans , but they all point to Italy (quite literally) & Central Europe. So we’re missing a thread from Anatolia – Caucasus -> Balkans -> Europe for 20-14 kya . Pinarbasi AHG doesn’t seem to be the fit either
    Have you any ideas ?

  8. Archaeologically speaking I don’t have any specific idea. But with the extremely low population in Europe during the LGM it’s not too strange that even a very small and hardly visible migration (most likely predating the LGM) could be responsible for the post-LGM repopulation.

    Another factor could be the much lower levels of the Black Sea during the Paleolithic, which would make any movements around it impossible to detect.

    I guess that any post-LGM site from the Balkans or Italy must contain WHG remains. The question is where exactly where they during (or just before) the LGM.

  9. Hi Alberto, un saludo. There is a very interesting paper by Jaime Lira (Atapuerca project) on the mitochondrial DNA of the Lusitanian horses that shows the genetic continuity of this lineage in Iberia. The myths that have been built for years regarding the theory of Kurgan by Gimbutas will fall one after another. (Origin of R1b-L51 / P312 in the Yamnaya culture, expansion of metallurgy from Eastern Europe, linking R1b to the IE language) Regarding the issue of horses, domestication occurred in several places and in different historical moments, and of course in Iberia it has nothing to do with R1b or the Indo-Europeans.

    On the other hand, the case of Szigentzenmiklós is another one of the many arguments that we used to demonstrate the migration of Iberian BBs to Hungary (together with uniparental markers, autosomal DNA, and some unique cooper objects) where they were mixed with Yamnaya migrants and native neolithic people.

  10. Hi Gaska, bienvenido.

    Yes, it seems some mtDNA lineages from early Iberian domestic horses have survived until the present. Even some fraction of native admixture can be detected, I believe. A sample from the LBA/IA still had 10% native Iberian admixture, but it probably vanished soon after that down to modern levels.

    Many of the myths of the Kurgan Hypothesis by Gimbutas had already been dispelled. What can be new with this study is the much later and therefor better documented hypothesis by David Anthony regarding the importance of domestic horses in the steppe expansion (which he saw mostly as cultural, not so much demic) may be a myth too, which is more surprising if it gets confirmed.

    In any case, it doesn’t matter that much, since after all the steppe migrations happened with or without horses and had a big impact in north and western Europe. A different issue is the language they spoke, which is unknown and indeed could be ancestral to Basque and Iberian and not to Indo-European languages. That’s something we also have to wait for more data to really know, though.

    I personally don’t see much evidence of a migration out of Iberia to the rest of Europe. We’d need a good sampling of France and Italy before we could have any decent knowledge about such putative migration.

    Many questions are still open, so let’s not get ahead of the data and just swap one myth with another one. Here I do my best to keep objective information and not favour any special hypothesis. So I hope you’ll keep following this blog to have another point of view that frequently differs from other more adventurous ones.

  11. I think it’s good that you’re as impartial as you can be. Few people outside of Spain think that there was genetic exchange between Iberia and the rest of Europe with the expansion of BB culture especially after the first Olalde’s paper. I think there were Iberian migrations (in fact it has been shown with the appearance of Df27 in Sicily in BB sites with Ciempozuelos pottery) but I do not try to convince anyone and although it is true that it is necessary to wait for France and Italy, the truth is that we have enough evidence pointing in that direction. I don’t know if you have read the new book that has been published regarding the chalcolithic graves of Humanejos- In that site and in Szigentzenmiklós identical copper halberds of an archaic type with three-nailed rivets have been found-These halberds are unique throughout Europe (this is more of a coincidence do not you think?). If to this, you join the horse, that in spite of the few studied sites at least 11 mitochondrial haplogroups are identical in Hungary and in Iberia, and the autosomal composition of some Hungarian samples (there are some that are more Spanish than me), only we can conclude that the migrations took place

    Necesitamos gente independiente que tenga tiempo y pueda trabajar con las muestras de ADN antiguo para de esta manera formar nuestro propio criterio (sean cuales sean las conclusiones).

  12. @Gaska

    I can just say that both myself and the people who read and comment here try to keep a more astringent interpretation of the data and avoid creating theories that are not strongly supported by enough data.

    I can’t go much into details here (you’ll find information about those topics in older entries and their comments), but keep in mind that the halberd from an Humanejos burial is post-2500 BC and associated to a population that had just arrived to Iberia (for more on halberds check this paper).

    The DF27 from Sicily may well have arrived there from Iberia, but again it belonged to a population that was new in Iberia.

    An out of Iberia migration should refer to the population that lived in Iberia from the early Neolithic to ca. 2500 BCE, since that’s the population that we could call “Iberian” at that point in time. The people who arrived from 2500 BCE onwards were very different genetically speaking, and they were already all over northern and western Europe.

  13. It will be a pleasure to discuss all this with you. One of the important issues of Humanejos is that R1b-P312 and I2a share tombs and grave goods, there is no distinction between them and therefore the deposit is not only associated to a new population (according to your criteria) but also to the local chalcolithic population. The archaeologists who have studied the site doubt the theories of Iñigo Olalde (although he has written the chapter related to the genetic study of the analyzed skeletons) because they consider them very hasty (Spaniards may be more incredulous, more difficult to convince and need more evidence than the Anglo-Saxons need when we talk about our genetic origin)

    As you can understand I have doubts that R1b-P312 is a newcomer population to Iberia, because I believe that this haplogroup originated in the Franco-Cantabricanregion, that is, they were already at home. Both P312 and L51 are Western (understanding Germany as Western Europe)

    The BB culture did not speak IE (at least in Iberia where the genetic continuity between the chalcolithic and the Iron age has been sufficiently demonstrated), and its expansion towards the east, stopped the expansion of the steppe migrations (CWC). Surely you have read in many forums that the guardians of Kurganist orthodoxy are trying to hide P312 in the SGC, recovering the old and outdated Dutch model and have gone so far as to propose that the CWC gave birth to the BBC. At the moment we also know that this lineage is older in Iberia than in the Netherlands, then the criteria could be perfectly the opposite. The last trench of the Kurganists will also end up falling and we will start talking about the German and French Neolithic cultures as the solution to the origin of L51

    Regarding the dating of the tomb where the halberd was found is 2,388 BC, at the moment older than the tomb of Hungary where the other halberd was found. What was the direction of the migration? BB culture lasted 1000 years in Iberia (approx 2,750-1,750 BC), surely you can imagine the number of migrations and small population movements that occurred during a millennium (both directions), so I am convinced that the Iberian BBs migrated to other BB regions (Morocco , Balearics islands, Sicily, Liguria, French Mediterranean coast etc) and the Df27 – Z195 of Sicily are the genetic demonstration of what I have been defending for years.

    But you are right, we will comment on issues as they arise, and I hope you keep the debate going for a long time

    Un saludo

  14. @ Alberto

    I get the impression that the recent horse DNA study, as good as it was, did not settle the question. And also lets remember that the dominant domestic horse lineage might be a different, although related question, to expansion of steppe ancestry, PIEs & other linguistic groups, etc
    During the Eneolithic, horse bones dominated the faunal assemblage at Dereivka and lipid resideu analysis confirms they were eaten. There is of course also Botai; and generally, hroses roamed around northern Eurasia since the Paleolithic.
    So its probable that horses were domesticated in a few places, and how one particular lineages survives today might be a different ‘horse story to human expansions.
    So I think horse riders didnt come swarming in from the Volga steppe, but I think there is good circumstantial evidence for horse riding by the 3rd millenium. I think the rapidity of BB expansion does support that, albeit indirectly.

  15. Deliberately being simplistic and a bit “jumpy” on conclusions based on Alberto’s various inputs… BB had a steppe component acquired during the Bronze Age . At least a subset of them spoke photo Iberian. they acquired Iberian(Spanish) horses for greater mobility..

  16. @Rob

    Yes, I agree that the study didn’t settle the question (quite the contrary, it brought more questions to the table). I also agree that the expansion of the main domestic horse and steppe ancestry may not be related (and this is exactly one of the questions the study brought, since the main hypothesis before it was that it was intimately related).

    We do know now that horses were domesticated in several places, but only one specific type became important. The reason of this is probably not irrelevant (unlike the survival of just one male lineage after a bottleneck, which is something circumstantial). The most probable cause for the main domestic horse to have been the one that survived is because that’s the one that became useful enough. And that could be related to chariots.

    I guess we have circumstantial evidence for horse riding in the bronze age, but we don’t have evidence that whole populations were horse riders. Maybe a horse could be used by a small person as a means of fast transport for some specific tasks, but other than that I don’t think it was a relevant thing until the Scythians (I don’t think that the rapid expansion of Bell Beakers can be related to horses in any significant way, at least with the data we have up until now).

    On the other hand, chariots started to be used in conjunction with horses and other equids in the second half of the 3rd mill. And it’s this what seems to have been the first important use for real horses. The ones that were first domesticated and trained for this task seem to have been the ones that expanded and replaced all others.

    How this (if correct, since it’s still just something I’m speculating about based on the clues we’re getting) may have influenced human or linguistic expansions is a different question. The “coming of the Greeks” may (or may not) be related to it. The Celtic expansion could well be related to it, at least in some areas. I guess I’ll have to take a second look at Robert Drews latest book for more clues about it, though IIRC he referred more to horse riders than to charioteers.

  17. Alberto,
    Drews outline of the use of horse is well argued. A progression from use as meat then traction then trialing riding single then finally mounted equestiranism.

    Im not sure how it might link to later human groups. The genesis of Celtic cultures certainly appears to have had some eastern input, before they themselves expanded east & southeast. With Greeks, its often proposed on blogs but, I doubt we’ll see a Sintashta conquest (although I don;t know this for sure). Much more realistically, we can propose that individuals like the Z93 guy from MBA Thrace might have been involved in their trade. I think Drews idea of a conquest of Europe (from Sintasha for northern Europe & Trialeti-like groups for southern Parts) is a bit radical, however, the effects these contacts had within Europe, which enable certain individuals and groups to manifest power & forge alliances is probably worth looking at.

  18. Re: Horses An interesting fact I came across by accident is that in a late BB settlement pit in Vienna-Rennweg*, more than 80% of the 37.8 kg(!) determinable bone material was from horses, suggesting a horse breeding specialisation that is also considered for contemporary W. Hungary. The Vienna-Rennweg bone assemblage contrasts markedly with preceding Upper Austrian MN assemblages, e.g. from the Jenisovice Culture (post-Baden), where game still dominated over domesticates.

    https://www.academia.edu/11592570/Ostösterreich_im_Endneolithikum_Am_Ende_der_Welt (p.32)

    These finds suggest that horse breeding was introduced into Upper Austria/ W. Hungary by BB incoming from the West (i.e. down the Danube). Central European horse-breeding may as such well go back to Iberian rather than Steppe traditions. If so, Vienna’s famous Spanish Riding School would (unintentionally) have re-established an already ancient connection..

    * Vienna’s Rennweg is the old Roman Limes Road leading eastwards to Carnuntium (Hainburg) and ultimately Budapest. In the vicinity of the BB settlement pit lay the late Roman civilian suburb (vicus), on a terrace over the east bank of the Vienna river. As such, we are dealing here with a strategic location, not just some minor farming outpost.

  19. @ Frank
    “”These finds suggest that horse breeding was introduced into Upper Austria/ W. Hungary by BB incoming from the West (i.e. down the Danube). Central European horse-breeding may as such well go back to Iberian rather than Steppe traditions. ”

    I don’t think the evidence supports that contention (in its entirety), as per Alberto –

    “Furthermore, a sample from Hungary c. 2100 BC shows some 12% admixture from this Iberian lineage (the rest of its makeup being from main domestic horses and another unknown lineage”.

    The Iberian lineage only contributed 12% to the Hungarian horse

  20. Yes, the data available is still too scarce to draw any sort of definitive conclusions. I just think that if Bell Beakers (and/or CWC) brought domestic horses from the steppe it would be more evident by now, both archaeologically and with the few samples we have. The Hungarian horse from c. 2100 BC had something like 40% from the main domestic type, and the Iberian one from 1900 BC nothing at all. That’s quite a surprising twist in the story of horse domestication in itself, even if we don’t know further details yet. It will be interesting to see more horse DNA from BB sites around Europe.

    Interesting too that the old paper (Mashkour 2003) I mentioned in the previous post dedicated to horse domestication was not only right about Botai horses as mentioned there, but now also it seems about Bell Beaker horses being different from the steppe ones and more similar to wild horses from Iberia, southern France, Italy… Her final suggestion was to look for the origin of domestic horses in more southern locations rather than the steppe. Soon enough we should know if she was right about that too.

    @Rob, regarding Greeks I was certainly not suggesting any direct or even indirect connection with Sintashta. All the cultural influences are from the eastern Mediterranean and deeper into West Asia, including the weapons and chariots (the stelae from the shaft graves depict some chariots with four spokes, which according to David Anthony himself was an innovation from the Near east). I would guess that the chariots came together with the horses trained to pull them.

    In general, I’m very interested in a good sampling of Thrace during the second half of the 3rd. mill. and the possible expansion from there to nearby areas in Europe (Romania/W.Ukraine, Carpathian Basin, Greece and Adriatic/N.Italy).

  21. Continuing with animal aDNA:

    Ancient cattle genomics, origins, and rapid turnover in the Fertile Crescent

    Pereira Verdugo et al. (No free access)

    Genome-wide analysis of 67 ancient Near Eastern cattle, Bos taurus, remains reveals regional variation that has since been obscured by admixture in modern populations. Comparisons of genomes of early domestic cattle to their aurochs progenitors identify diverse origins with separate introgressions of wild stock. A later region-wide Bronze Age shift indicates rapid and widespread introgression of zebu, Bos indicus, from the Indus Valley. This process was likely stimulated at the onset of the current geological age, ~4.2 thousand years ago, by a widespread multicentury drought. In contrast to genome-wide admixture, mitochondrial DNA stasis supports that this introgression was male-driven, suggesting that selection of arid-adapted zebu bulls enhanced herd survival. This human-mediated migration of zebu-derived genetics has continued through millennia, altering tropical herding on each continent.

    The shift is visible in samples from the Levant, Iraq and Iran, where there are samples predating and postdating 4200 BC. Presumably in Georgia too, but there are only postdating samples. The ones from Central Asia have the shift too, but they postdate 4200 BC and we don’t know if they were different before that date or not.


  22. Alberto,

    Glad to see that you’ve already noted that cattle aDNA paper.

    There is something very interesting about the spread of the Zebu cattle in the Near East in the 2nd millennium BC – it is also the timeframe when the Mitannis and arguably the Hittites also show their presence or become prominent at any rate.

    More significantly Asian Elephants also appear in Syria in the Mitanni domain around the same time and go extinct around 800 BC.

    Co-incidence ?

    Also worth remembering the discovery of chariots dating to 2200-2000 BC in Sanauli. Might help putting things in context.

    As for Zebu in Central Asia and Eastern Iran, these regions are clearly very closely related to the Quetta pottery culture and the Nal Pottery culture of Early Harappan period that existed in Baluchistan. So Zebu should have been present in these 2 regions quite early (around 3000 BC or earlier)

  23. Hi Jaydeep,

    Yes, a very interesting paper this one, bringing some new data that we didn’t know about, and regarding such an important domesticate as cattle.

    It’s hard to know how this relates to the movement of humans, though. There could well be a connection with the Mitanni, who are supposed to be an elite and their DNA cannot be traced from modern populations. I hope that now that apparently excavations in the Mitanni area have resumed we can get aDNA that sheds some light into it.

    The Asian elephants in Syria may be a different case. Those are supposed to be native to Syria. Asian elephant ivory is found in Iberia since the beginning of the 3rd mill. It’s only during antiquity that the Indian elephant was widely imported to the Near East (though it could be the case that Syrian elephants could show admixture from Indian ones around the same time as cattle does, but we’d need ancient DNA of the elephants to know it).

    Back to cattle, I remember you mentioned before that the steppe cattle had Zebu admixture. Do you know the date of whichever samples show such admixture? If it’s before 2200-2000 BC. then that must have arrived via Central Asia (where it probably was present quite earlier, as you said).

  24. Alberto,

    Well, regarding the Syrian Elephant it is now the majority view of archaeologists working on near east Bronze Age that the Asian elephant in 2nd millennium BC Syria was imported, likely from the Indus civilization. Here are a couple of links :-

    1. https://www.tandfonline.com/doi/full/10.1080/00758914.2016.1198068


    Ofcourse these archaeologists are oblivious to the idea that these import of Asian Elephants from the Indus could have something to do with the Mitannis who were Indo-Iranian.

    About Zebu admixture in the steppe we do not have any specific timeframe but it has been speculated to derive from the Yamnaya period by a recent study


  25. @Jaydeep

    Thanks for the links. It’s a bit mysterious this thing about the Asian elephant. Those studies you mentioned are about the LBA/IA, and seem to ignore earlier presence of Ivory in the Near East.

    From Iberia there is ample and well studied evidence of elephant ivory since the late 4th mill. (this one of African provenance, which was the type found in the SW of Iberia) and early 3rd mill., which in the SE of Iberia is without a doubt from Asian elephant.



    However, there’s a problem to determine the route followed by this Asian elephant ivory (mostly imported as raw material and worked in the Iberian peninsula, though the know-how was probably learned from oriental sources too) because of the lack of studies in intermediate areas such as Sardinia or Italy or eastern Mediterranean. Not just lack of studies, but even lack of conclusive evidence of elephant ivory at all. One of the very few sites mentioned is Bir es-Sadafi, but there’s not much information available about it either:


    From the conclusions on the second link provided above:

    But where exactly did this Asian elephant ivory come from and how did it reach the West?

    The Asian elephant in the Early Holocene had a more widespread distribution than today, ranging from China throughout India and Mesopotamia and as far as Syria. However the available information in Levant and Mediterranean is insufficient to establish a direct connexion with the context from Valencina [Seville, Spain]. The problem is not only the exact source but also the routes and the mode of distribution of this material and their chronology. To date, almost no analysis on prehistoric ivory has been undertaken outside the Western Mediterranean to discriminate between Asian and African elephant ivory (there have only been attempts to differentiate between hippopotamus and elephant ivory in general). In addition, their occurrence in the far west of the Mediterranean in the 3rd millennium BC is, at present, a millennium before their occurrence in the central and eastern Mediterranean.

    In fact, in the Levant we do not hear about elephant hunts and elephant bones datable before ca. 2000 BC, and they are currently restricted to the ones from Çatal Hüyük and some fragments from the Early Bronze Age III in Ras Shamra and the most ivory comes from hippopotamus. This seems to coincide with the situation in Crete, [and] the Greek
    mainland. In Sicily, Sardinia and the Italian peninsula, the earliest known ivory objects are dated around the middle of the 2nd millennium BC.

    So, our analysis conducted on material from Valencina in the first half of the 3rd millennium BC only speaks about Asian elephant ivory. But, at present, we are unable to trace back a possible route throughout the Mediterranean. At the same time, with the available data, it is still not possible to distinguish between the different geographic origins among this type of prehistoric ivory. The question of the exact geographic origin of the ancient distribution of Asian elephants will have to be solved by future analysis, as could be systematically undertaken by carbon (13C/12C), nitrogen (15N/14N), oxygen (18O/16O) and strontium isotope (87Sr/86Sr) analysis on recent and archaeological ivory.

    Whatever the case, it’s clear that Asian elephant ivory could not have reached SE Iberia in the 1st half of the 3rd. mill. directly from India, avoiding everywhere in between.

    As for the Zebu admixture in steppe cattle, I didn’t find much evidence about it there. There’s this sentence there:

    Using whole-genome sequences of ancient human DNA, Jones et al. (2015) and Haak et al. (2015) suggested massive migration of Yamnaya steppe herders as a source of dispersion of Indo-European languages to both northern-central Europe and India. These herders might also have mediated gene flow between Indian zebu and Ukrainian steppe cattle.

    But no information about which Ukrainian steppe cattle they are referring to, since I couldn’t find any samples in their study. There is, however, zebu admixture in (modern) cattle from the Balkans and Italy. We’d need to know when that admixture arrived there too.

  26. Alberto,

    Thanks for your reply. It’s an interesting topic.

    About Elephants, yes I am aware of that early Asian elephant ivory on the SE Iberian coast, a most fascinating topic.

    It appears we Iberians and Indians have known each other since long

    On a more serious note, Yes it is clear that the Asian elephant ivory could not have come directly from South Asia. One way to explain it could be that the demand of ivory in Iberia was much greater than even in the Near East. I am pretty sure Asian ivory was in use in the Near East during the 3rd millennium BC, but on a smaller scale.

    But certainly the evidence of Asian elephant ivory in Iberia in the 3rd millennium BC is not a proof of the existence of the Syrian elephant during that period. The evidence shows the absence of elephant bones right through the Holocene in the Near East until it’s sudden appearance at the end of the 3rd millennium BC around Turkey/Syria. So clearly the Pleistocene Syrian Elephant had gone extinct by around 10 kya if not earlier and the late Bronze Age Elephant is not descended from it. The only place it could have come from was from South Asia.

    Here is an interesting article by Talageri on this topic that is very well written –


    About cows,

    The Balkan and Italian breed of cattle which show Indicine admixture are known as the Podolian breed descended directly from primitive steppe cattle. The steppe cattle which is very closely related and is also perhaps considered as Podolian is the Ukrainian Grey and perhaps other varieties of Grey steppe such as Hungarian Grey.

    Since these Podolian are considered to not have been admixed with other cattle ever since they were brought to the Balkans and Italy, this indicine admixture is considered to have rcvd by them in the steppe itself.

  27. @ Alberto

    ”However, there’s a problem to determine the route followed by this Asian elephant ivory (mostly imported as raw material and worked in the Iberian peninsula, though the know-how was probably learned from oriental sources too) ”

    Despite the continued critique by some archaeologists, aDNA can inform not only about large migrations, but also the the lack of such migration. In earlier decades, scholars had often proposed that the south Iberian Chalcolithic cultures were of East mediterranean or Aegean origin. Although this had already been doubted on the basis of chronology, we now know that they were a ‘native’ development from the local Middle Neolithic basis (well, its more complex than that, but we can say it was not mediated by a significantly extraneous import).

    So we can suppose that those citadels and exotica were trans-cultural diffusions mediated by key individuals, by ‘down the line” exchange, perhaps. With a large enough sample set, some foreigners will undoubtedly appear

  28. @Rob

    Yes, I agree. The early days of ancient DNA studies (and that’s only 4-5 years ago) probably scared off archaeologists with the over interpretation of that early data. But things have been maturing since then and now it’s clear that the Manichaean pots or people debate is a thing of the past.

    Ancient DNA has already uncovered as many presumed migrations that never happened as it has confirmed others that did, or found new ones no one predicted.

    It’s now obvious that only with a good amount of cooperation between different experts it will be possible to make good use of the new data and obtain the best results.

    I’m hoping that the long period between the Narasimhan et al. preprint and the final version will show this process of maturation given the ambitious nature of the study with its venturing into a new geographical territory, which showed that much help was needed from more experts in the field . Let’s see if this closer cooperation has happened in the process and we see a much improved paper when it finally comes out.

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