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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119 thoughts on “Of horses, camels and extinct lineages

  1. See also, as amendment to my previous post:


    Linguistic aspects are further elaborated upon by Zoller 2016. He concludes (emphasis is mine):

    “I have shown that at the time of Old Indo-Aryan there must have existed a linkage of lects, with Vedic just one of them. These lectal differentiations seem to suggest that the standard model of the three branches of Indo-Iranian is in need of a revision. Their existence also supports the idea of the earlier immigration of the ancestor(s) of the Outer Language which led to a strong encounter with Munda/Austro-Asiatic languages (but to a weak encounter in case of Vedic and Classical Sanskrit) which must have dominated the prehistoric linguistic area of northern India. This dominance must have extended far into prehistory because of the many parallels in the language isolate Burushaski. ”


  2. @ Alberto

    “Any new paper on that subject that deals with the Anatolian influence in Thrace and the demise your refer to? This is pretty much a work in progress (for what I know) so still many details are poorly known. Maybe some aDNA could help too…”

    Indeed; but as mentioned; they are clearer
    I’ve sent you the info (listed & outlined also on the ‘Bronze age’ thread previously).

  3. @Rob

    Got it, thanks.


    Yes, it would still be great to see that 3rd part of your CHG on the steppe series. After your HD crash I could never get to you by mail, but you still have access to continue with the post. If you have some new mail contact me when you’d like to write it or publish it.

  4. @Alberto
    I do agree with the spirit that you convey, but think language = culture (and vice-versa) inevitably.

    In this vein, Mallory’s attempt to define I-E and Swadesh lists are effective somewhat.

    The verboten topic in genetics is whether culture derives from genes. No use poking that bear for the moment.

    Personally, as I’ve shared with Rob before, I think it is more effective to think in terms of Complexes when discussing a subject as expansive as this one.

    Regarding: hata
    Don’t think it’s obvious that centum. s -> h more likely as noticed in other Iranic languages.

  5. @Frank

    In your theory what would be the actual PIE/Indo-Hittite culture? Hajji Firuz?

    “and I find it very hard to imagine another place in time and space where/when IIA and Proto-Uralic could have interacted.”

    The thing is we do not know the actual PU homeland to pin point the actual relation between PU and II. Currently there are many theories on the PU homeland that, in my opinion, it’s very easy to imagine many other scenarios.

    A few examples:

    1. BMAC- Seima Turbino interactions
    2. BMAC-Andronovo interactions (if CW is Uralic as some claim)
    3. Issedones and other Scythian groups

  6. @Alberto

    I do think that there is a PIE culture. We cannot fully construct it but we there are a few core ideas in all known IE cultures and religions. Eg. The stories of Kingship, the holy abode, the three brothers…etc. Some of them are unique to Indo-Europeans. Sure the stories of the thunder god change over time but at their core it’s a story of a god/king challenged by a serpent, and it is an okay representation of the ideology of those people.

  7. @FrankN “My understanding is there is some of that, especially in Dardic. Kashmiri hata “hundred”, e.g., is obviously not Satem, but Centum, as is Kashmiri hun “dog”. Similarly, Khowar kuy “where?”, kyobachen ” for what?” seem to have conserved non-satemised traces of PIE “kw””

    kyobachen, kuy of khowar is seen as preserving kw is debatable. If so there are more mainstream words:

    when(eng), kabhI(hindi), keMvhA(marathi), kabE(bengali) etc where the NIA forms cannot be directly derived from kadA(sanskrit) which is strangely closer to latin derived forms (quand, quando) and slavic (kada, kedy)

    Even the often used kahAN (where in Hindi) is an odd one out vs other NIA (kitthE, kothay, kuThE) and Sanskrit(kutra) English (whither, where). None of these is conclusive by itself but I suspect that a structured study of such phenomenon does not exist.

    I was told that the English pair whither/tither is not related to kuThe, tithe (Marathi), kothay, tathAy(bengali), kutra, tatra(Sanskrit).

    don’t fully buy this … seems like inadequate research. As far as other reflexes of *kw in NIA we see a backed vowel after k in some cases but not after t and so forth. Is this significant or just random traits?… I don’t know yet and won’t have bandwidth to explore for a long time

  8. @FrankN, if you imagine a PIE migration from Iran Chalc to Central Asia, then parts of South Asia are on the way and probably would not be circumvented.

  9. In regard to Franks model

    * I have been interested in long duree models, however I presently agree with Kristiina in that languages are more likely to have been burst phenomena,.

    * The concern is that the social-demographic phenomena pointed to in the model seem too disparate to have carried strong language-cohering forces. Even more problematically, it would mean that half of Asia was Indo-European speaking, until the Bronze Age when it all suddenly reversed

    * It is likely that CHG ”arrived in the steppe” via the Caucasus, because at c. 4500 BC it peaks in Progress Eneolithic, not the Samara bend. But in any case, as Alberto pointed out, it really doesn’t matter which exact direction they arrived from, as they were 6th millenium forager-hunters (even if Kelteminar had some goats, etc)

  10. https://www.cnn.com/2019/08/06/asia/turkmenistan-president-gateway-to-hell-intl/index.html

    I’m surprised by the cadence and meter of the language.


    It is strongly moratimed.
    resembling both dravidian languages with frequent gemination as well as classical Sanskrit because of frequent compounds.

    These traits are long lived and not easily transmitted. They transcend language families.

  11. Rob:
    “The concern is that the social-demographic phenomena pointed to in the model seem too disparate to have carried strong language-cohering forces. ”

    Well, I didn’t yet elaborate on social-demographic features. But some are obvious:

    – Spread of Copper metalurgy,
    – Long-distance trade in precious stones (Carneol, Lapis Lazuli) and metals (gold, tin),
    – Wool sheep as key agricultural innovation (plus, in general, (south-)eastward spread of Iranian domesticates,
    – Non-light sensitive barley, originating on the Iranian Plateau, better adapted to moderate/ continental/ mountain climates than Fertile Crescent/Mediterranean varieties, and less draught-sensitive than wheat, allowing for effective exploitation of the Caucasus uplands (Sioni, KA), the Steppe, plus low-precipitation parts of Central Europe such as Kujawy, Bohemia and the Saale area after the end of the (humid) Holocene Climate Optimum.

    “Even more problematically, it would mean that half of Asia was Indo-European speaking, until the Bronze Age when it all suddenly reversed.”

    Well, for once, a good part of Asia, including Tajikistan, is still IE speaking. The reversal occured rather during the late IA to the Medieval (in Anatolia as late as 1923) via the Turkic/Mongolic expansions. That reversal equally affected the Pontic-Caspian Steppe, pre-dominantly Turkic (Khazars, Tartars) until Catherine the Great’s times, so I don’t really get your point.

    Otherwise, the village of Jastorf (the eponymous culture is commonly regarded as West Germanic homeland) lies just on the border between predominantly Germanic and Slavic toponymy. Austronesian is well alive and kicking, except for its Taiwanese homeland, where it has been relegated to endangered minority language status. Algic had virtually disappeared from the presumed homeland around Kennewick already in pre-European times, to instead stretch all the way from the Rockies to Newfoundland, and so on. Such geographic shifts were apparently anything but uncommon…

  12. @tim
    I think Shakas were there by the time Alexander came in. Definitely in greater Punjab if Greek and Persian records are to be believed.

    Not sure what you were meaning by stating you are Indian but I gather you mean it’s of particular interest to you.

  13. @Marko
    The data is interesting.
    Their suppositions are idiotic.

    “Four individuals are very tall, with height in the range 184.41-187.55 cm.” Very tall? Plenty of people this tall in India.
    “it is tempting to hypothesize that the tall and robust individuals correspond to the Roopkund_B cluster of Mediterranean origin.”
    “It is tempting to hypothesize that the Roopkund_B individuals descend from Indo-Greek populations established after the time of Alexander the Great…”

    They should use their research money and travel to Crete and the Mediterranean to see all those exceptionally tall people. A cheaper option is to read historical records of the height of Alexander’s troops compared to the found locals in the Punjab.

    Also includes R1a, G2a, and T1. None of the haplogroups are surprising. Ignoring my above gripes, I’ve long suspected E1b (which is found in NW populations) is of Greek origin.

    This data will be useful to everyone other than the people on this paper. But still very good to have this data.

  14. @Atri∂r

    The R1a, G2 etc. are from 1800 C. E, the R1a being typically Slavic. . The Roopkund_A group on the other hand is dated to ca 800 C.E. . So for Medieval Uttarakhand we have:


    If the hypotheses regarding Proto-Bangani are correct, these samples are right from the area with a complex and supposedly very old presence of IE languages. See Franks comments as well.

    And yeah, Cretans probably aren’t particularly tall – weird conclusions. Seems very unlikely that they were Indo-Greeks, though. Maybe Ottoman merchants.

  15. @Marko

    Which haplogroup group do you find intriguing? Which do you find rare? All of these are present in India in fairly old communities.

    As for languages, it’s useless to try and equalize language and haplgroups, especially from 800 c.e. The only time I see that as useful, in respect to early Indo-European languages, is the Bronze Age. Before and after that, useless. This in part, is how I predicted J2 in Mycenaea and Hittite ahead of the samples. The power of prediction is what confirms the strength of a model. Unfortunately, modern-day academia does not work like this so instead, most are forced to plod and waddle towards a proper model.

    I saw FrankN’s comments. I disagree with his premises; mainly because I see many of his starting premises as incorrect.

  16. I find it interesting that the individuals from Group A (800 CE), and specifically those in cluster 1 within the group, are quite similar to the Swat Valley samples that go from 1200 BC to 1 CE. Now we see that population (with similar genetic structure and still no R1a) in a different location, 2000 years later than the earliest ones from Swat.

    One wonders if the R1a-rich groups were in very specific areas until after 800 CE and then started to become prevalent. The diversity of the haplogroup suggests a different story, so it might be just for random reasons that there’s still no R1a-rich samples from ancient India.

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