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horsewheellanguage
Update: The preprint is out. End update

… and also after that, when the sons of God came in unto the daughters of men, and they bare children to them, the same became mighty men which were of old, men of renown.

Genesis, 6:4

A Battle-Axe

A Battle-Axe

An emanation from the one most high…uh, I mean, David Reich, has given his talk at Oxford. Thanks to Jean Manco we have a pretty good report of what he said. The core element seems to be that a paper will soon be published using ancient DNA results to conclude that Indo-European languages came to Europe from the Yamna culture of the Pontic Steppe ~4,000 years ago. Roughly, the argument laid out by David Anthony in The Horse, the Wheel, and Language: How Bronze-Age Riders from the Eurasian Steppes Shaped the Modern World. Where Reich’s conclusions differ from those that Anthony presented in the book is that this eruption from the Eurasian heartland resulted in a genetic-demographic transformation of Europe ~4,000 years ago. Manco confirms that the genetic data from the ancient sites that Reich’s team has access too indicate that the two dominant Y chromosomal haplogroups in Europe, R1b, and R1a, arrived with the Yamna people. This is not surprising, as previous ancient DNA exhibited a surprising death of these two lineages from both hunter-gatherer and early to late Neolithic populations in Europe. And, recent whole genome sequencing of Y chromosomes indicates that both R1a and R1b lineages across Eurasia have undergone recent explosive demographic expansion on a Holocene timescale (closer to 5 than 10 thousand years). In terms of total genome ancestry it looks as if the transition to Yamna successor cultures in Central Europe (e.g., Corded Ware) was accompanied by substantial genetic turnover. In the initial Corded Ware burial grounds 60-80 percent of the ancestry seems to have derived from the Yamna. The modern Yamna-derived proportion seems to be closer to ~50 percent in a region like Germany.

The Yamna themselves are a compound population, a mix of ancient hunter-gatherer groups (analogous to the modern Karelians), and an intrusive population with Near Eastern affinities, likely from the Caucasus. I am not clear whether the Near Eastern group had “Ancestral North Eurasian” ancestry, but the hunter-gatherers almost certainly did. The 1 to 18 percent “Ancestral North Eurasian” ancestry across continental Western and Central Europe today dates to the arrival of these Indo-European speakers. The very low fractions in Southern Europe, and its near absence in Sardinia, may suggest that the Indo-Europeans were demographically more significant in Northern Europe, even though they were clearly culturally effectual along the northern rim of the Mediterranean, and into Anatolia. In line with the Mal’ta results Manco relays that Reich believes that the R lineages, which are the dominant ones across Indo-European speaking populations from the North Sea down to South Asia, came into Europe via the Yamna, but ultimately derive from an “Ancestral North Eurasian” group.

220px-Yamna-en.svgThere are some phenotypic tidbits in the talk apparently. The Yamna were tall in terms of their genomic potential. Additionally, the very high frequency of lactase persistence may date to their arrival in Europe (there is some lack of clarity here). I doubt the high lactase persistence frequency and genotypes which result in greater final height are together by coincidence. Large people need a larger nutrient pipe, and adult digestion of lactose sugar would enable that.

There are two aspects which are not widely address in this talk. First, what was the exact dynamic of how the Indo-Europeans replaced the original populations? The idea of “demic diffusion” by waves of “demographic advance” promoted by Colin Renfrew seem to gradual and continuous to be responsible for this. This is basically an argument predicated on individual fitness, summed over groups. In this case I suspect that a better analogy may be the future that Genghis Khan had in mind for Northern China before his adviser Yelü Chucai dissuaded him: the North European plain was cleared out of people and turned over to pastureland. Genghis Khan and his Mongols were convinced of the value of Chinese as tax paying peasants, who could support the Mongol elite with their surplus. I suspect in a pre-state society such considerations were less relevant, as the institutional frameworks which would allow for the smooth absorption of subordinate groups were less elaborated, or even non-existent.

Hittites

Hittites

In both the Late Neolithic and after the Bronze Age the Reich group alludes to a return of the primal populations which were marginalized by the farmers, and later the Indo-European agro-pastoralists. One way to look at this is that there were larger migrations which were overlain upon the palimpsest. But, I believe one might also consider a model whereby there is ascertainment bias in the sorts of burial sites being explored and sampled, and one might be witnessing a patchy occupation of the landscape by intrusive cultures. For example, the newcomers might monopolize the rich bottom-lands for thousands of years, but huge swaths of the hinterland might be occupied by marginalized and less developed people, who over time drift into the core and become culturally absorbed. Instead of imagining the expansion of these people as purely ones of a vast uniform wave front, it might be better to conceptualize them as penetrating into virgin territory along the optimal avenues of settlement, and producing a patchy archipelago of habitation.

Second, there is the issue that though Reich and company focus on Indo-Europeans and the Yamna culture, the genetics leaves may loose threads that are difficult to tie back up. At ASHG Mait Metspalu express to me some misgivings about the term “Ancestral North Eurasian.” How do we truly know the locus and distribution of this ancestral component across Eurasia ~10,000 years ago? The Kalash of Pakistan exhibit signals of admixture with this group as high as Northern Europeans, so it is not limited to West Eurasia proper. The highest fractions today seem to be found in the North Caucasus, among many groups which are not Indo-European. If R1a was brought by Indo-Europeans to Europe, it is harder to conclude that this was the case in South Asia. Though the frequency of these lineages is higher in the Indo-Aryan North, there are relatively high fractions of R1a even among some South Indian tribal populations. R1b is found in appreciable fractions in Sardinia and among the Basques (one argument for the old idea that R1b was the legacy of European hunter-gatherers!). Obviously some of this could be due to admixture between Indo-Europeans and non-Indo-Europeans. But I think a major issue here is that Indo-European groups were a synthetic population which arose in a world where there were many synthetic populations, with ancient and recent affinities to them. I doubt the “Ancestral North Eurasian” ancestral component was limited purely to Indo-Europeans. So it seems unlikely that the R1 lineages would be purely Indo-European, even if recent expansion of some of their sub-lineages is a function of the Indo-European cultural explosion.

Of course there are only so many ancient DNA samples one can retrieve from a finite number of sites. The age of new genomic discoveries will start to close over the next few years as the paleo-demography inferred will start to exhibit some predictable solidity. That means that a deep knowledge of the archaeology, and what history there is, is essential.

 
• Category: History, Science • Tags: Corded Ware, Indo-Europeans 
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  1. The European R1a-Z282 and the Indo-Iranian (+ Indian) R1a-Z93 have the same parent: R1a-Z645. My view is that this mutation arose in an Indo-European speaker who lived no later than the late Neolithic, probably on the steppe.

    If so, then the Indo-European expansion in South Asia was almost as massive as in Europe, but perhaps even more male biased?

    Those Y-chromosome sequences from the Yamnaya males will be very interesting, especially if some turn out to be R1a-Z645*.

    • Replies: @Razib Khan
    If so, then the Indo-European expansion in South Asia was almost as massive as in Europe, but perhaps even more male biased?


    perhaps. one issue is that the difference between non-IE and IE groups in europe/central asia is going to smaller than IE and "ASI/Dravidian", so not sure we have as much power to talk about sex bias in europe.... though yes, seems plausible.
  2. “The Kalash of Pakistan exhibit signals of admixture with this group as high as Northern Europeans, so it is not limited to West Eurasia proper.”

    Can you give me a reference or tutorial of how you are able to relate South Asian groups (ANI-ASI admixture) in terms of ANE and EEF.

    “Though the frequency of these lineages is higher in the Indo-Aryan North, there are relatively high fractions of R1a even among some South Indian tribal populations”

    There are few or no “unreached” peoples in India. R1A and R1B is present across the board, and reduction of Indian populations to invaders and natives is extremely hard(This may be the reason why Reich went to Andaman to search for untouched people). Several tribes and Scheduled caste have even later admixture dates than the original dates of admixture 3500-4000 years BP. This is probably more due to “the newcomers might monopolize the rich bottom-lands for thousands of years, but huge swaths of the hinterland might be occupied by marginalized and less developed people, who over time drift into the core and become culturally absorbed”. In Indian case, they pick up percentages of invader genetic signatures, at later and later dates.

    • Replies: @Razib Khan
    Can you give me a reference or tutorial of how you are able to relate South Asian groups (ANI-ASI admixture) in terms of ANE and EEF.


    they used D-statistics. it was on a poster at ASHG 2014. nothing about EEF. but the ANE signal was clear.

    In Indian case, they pick up percentages of invader genetic signatures, at later and later dates.

    i think there might need to be revision in how we understand the arrival of dravidian peoples, and their early relationship to indo-aryans (i don't think dravidian languages are ancient to south asia anymore btw).
  3. The “Karelians” you are talking about are from 5000 bc and have little to do with modern Karelians, who have only around 20% ANE and considerable EEF/ENF.

    • Replies: @Razib Khan
    yeah, good point. perhaps might be better to stick with neologism EHG.
  4. Apparently, Reich believes ANE is correlated with R1, except when it’s not. Sounds plausible.

    • Replies: @notanon
    Maybe that implies something e.g. that R1 were adjacent to and mixed with ANE in some places but not others?
  5. @Davidski
    The European R1a-Z282 and the Indo-Iranian (+ Indian) R1a-Z93 have the same parent: R1a-Z645. My view is that this mutation arose in an Indo-European speaker who lived no later than the late Neolithic, probably on the steppe.

    If so, then the Indo-European expansion in South Asia was almost as massive as in Europe, but perhaps even more male biased?

    Those Y-chromosome sequences from the Yamnaya males will be very interesting, especially if some turn out to be R1a-Z645*.

    If so, then the Indo-European expansion in South Asia was almost as massive as in Europe, but perhaps even more male biased?

    perhaps. one issue is that the difference between non-IE and IE groups in europe/central asia is going to smaller than IE and “ASI/Dravidian”, so not sure we have as much power to talk about sex bias in europe…. though yes, seems plausible.

  6. @Bultare
    The "Karelians" you are talking about are from 5000 bc and have little to do with modern Karelians, who have only around 20% ANE and considerable EEF/ENF.

    yeah, good point. perhaps might be better to stick with neologism EHG.

  7. @vijay
    "The Kalash of Pakistan exhibit signals of admixture with this group as high as Northern Europeans, so it is not limited to West Eurasia proper."

    Can you give me a reference or tutorial of how you are able to relate South Asian groups (ANI-ASI admixture) in terms of ANE and EEF.

    "Though the frequency of these lineages is higher in the Indo-Aryan North, there are relatively high fractions of R1a even among some South Indian tribal populations"

    There are few or no "unreached" peoples in India. R1A and R1B is present across the board, and reduction of Indian populations to invaders and natives is extremely hard(This may be the reason why Reich went to Andaman to search for untouched people). Several tribes and Scheduled caste have even later admixture dates than the original dates of admixture 3500-4000 years BP. This is probably more due to "the newcomers might monopolize the rich bottom-lands for thousands of years, but huge swaths of the hinterland might be occupied by marginalized and less developed people, who over time drift into the core and become culturally absorbed". In Indian case, they pick up percentages of invader genetic signatures, at later and later dates.

    Can you give me a reference or tutorial of how you are able to relate South Asian groups (ANI-ASI admixture) in terms of ANE and EEF.

    they used D-statistics. it was on a poster at ASHG 2014. nothing about EEF. but the ANE signal was clear.

    In Indian case, they pick up percentages of invader genetic signatures, at later and later dates.

    i think there might need to be revision in how we understand the arrival of dravidian peoples, and their early relationship to indo-aryans (i don’t think dravidian languages are ancient to south asia anymore btw).

    • Replies: @Hipster
    Razib,

    I am reading In Gods We Trust and Atran mentions Dravidians as separate from Aboriginals in India. I had always equated the two. Who are Aboriginal Indians supposed to be, Austonesian people who spoke an Austronesian language who were conquered by Dravidians?
  8. anonymous • Disclaimer says:

    If you care…

    Manco confirms that the genetic data from the ancient sites that Reich’s team has access too indicate that the two dominant Y chromosomal haplogroups in Europe, R1b, and R1a, arrived with the Yamna people. This is not surprising, as previous ancient DNA exhibited a surprising dearth of these two lineages from both hunter-gatherer and early to late Neolithic populations in Europe.
    ….
    There are two aspects which are not widely addressed in this talk.

  9. Good post with good caveats, e.g. about the possible bias of data from high status graves.

    I also share some skepticism that R1b in Western Europe has a source in Indo-Europeans linguistically, despite the fact that one can make a case for an Eastern Europe to Central Europe to Southern Europe pathway based on R1b phylogeny.

    On the other hand, it does seem pretty clear that the source of Europe’s R1a and R1b were not that far from each other geographically, and probably absorbed fitness enhancing technologies with a common source.

    Also, re Genesis 6:4, one of the groups associated with Giants in Biblical times were the Philistines (e.g. Goliath). And archaeology and Egyptian historical sources now strongly point to them being Bronze Age collapse era migrants (ca. 1200 BCE) who were ethnically Mycenean Greeks, i.e. Indo-Europeans, who started to arrive in the Greek mainland ca. 2000 BCE (800 years earlier) and in all of the Aegean including Crete by about 1500 BCE.

    • Replies: @hbm
    Arguably the passage was written by a Hellenized Jew, the Nephilim being a sort of reductive appropriation of Homer's "men of reknown."

    Certainly whoever authored Goliath had seen a hoplite.

  10. According to acient Chinese souce, Genghis Khan genocide every body except craftsman (blacksmith, carpenters ect) from the very beggining. Only later, mongols started sparing farmers as tax payers.

    He never killed craftsman even before Yelü Chucai advice.

  11. Razib, I don’t know if this is helpful, but I did my thesis on the kinds of contact, and affects of contact, between Khoisan hunter-gatherer and Bantu-speaking tribal pastoral/farming societies. In the 1970s, after some 800 years of contact, the two economies were still distinct. There had been gene flow between the two populations, resulting in a small mixed population that had largely adopted farming and kept livestock. Intermarriage between people in this metis group and the dominant Tswana tribes were virtually non-existent. I certainly recorded none.

    Marriage of women from the hunter-gatherer group into the metis group accounted for less than half a percent of all formal unions, or informal concubinage relationships, in either group, and were exclusively movement of females who became second wives in existing metis households.

    Defectors from this mixed population, usually children of women who had married in from the hunter-gatherers, returned to hunting and gathering, often with explicit declarations to the effect that such a life and identity was preferable to being looked down on. These defectors were, in my own sample, all male. This is with a four generation genealogical backdrop, but I am fairly confident that this pattern was not unusual.

    This means that Y chromosome lineages can probably flow – possibly via a metis community – from a politically and militarily dominant pastoral society into the hunter-gatherers more easily than they flow in the opposite direction.

    I am suggesting this here to introduce the caution that a high proportion of certain Y variants in an existing population is less likely to represent population replacement, than it is to indicate a similar dynamic as I have documented. A very high incidence of such variants might be due to a combination of drift (- in that the founders of the metis communities appear to have represented a small proportion of the total pastoral people who migrated into the area once exclusively inhabited by hunter-gatherers -) and also to relatively lower proportion of men who successfully reproduced over long periods of contact.

    To sum up, in a situation of frequent contact between economic systems, exchange of goods, labour, and personnel (or just genes) tends to be fairly ubiquitous but gene flow is neither random nor necessarily even in both directions.

    One other thought – about differential mortality and morbidity at the zones of contact between foragers and post-Neolithic peoples. Cattle and other livestock are the source of some significant zoonotic infectious diseases. Brucellosis and tuberculosis were widespread in the Kalahari due to contact with cattle, and it struck me that men, among the hunter-gatherers, appeared to frequently be infected with one or both. Fatalities after long debilitating illnesses were about 10% and highest among men who had frequently herded cattle for the Bantu or in the metis community. Melioidosis (a form of what used to be called pseudomonas) is carried by horses and donkeys and causes up to 95% fatalities. The various pox viruses, including measles, chicken pox, cow pox and small pox, all derive from contact with infected livestock. All of these would be in play during an invasion of a hunter-gatherer territory by post-neolithic livestock-owning cultures.

    Patterns of differential mortality due to variation in genetic immunity, could well have been linked to the events that left some Y alleles and their lineages with numerous descendants and others virtually extinct. Certainly it seems logical that the hunter-gatherers would have had a swift upward tick of mortality after first contacts, events which might well have led to cultural immunity protocols restricting contact and even leading to hostility.

  12. Matt says:

    Razib: There are some phenotypic tidbits in the talk apparently. The Yamna were tall in terms of their genomic potential.

    This seems plausible – apparently I think Corded Ware burials worked out about 2.5 inches taller by direct measurement in some study compared to Early Neolithic Farmers in Europe.

    Although I wonder if any of this comes from tendencies for burials to be more disproportionately of higher SES compared to earlier cultures who buried with less regard to social status? Or for burials in high status kurgans to be more archaeologically visible. More social stratification / chiefs / warleaders in Yamna? Height genetic / realised usually seems to go with higher SES. So the Yamna fossils might be tall in their genomic potential more than the Yamna population, as such.

    • Replies: @notanon
    On the other hand maybe the dudes with more brawn in their ancestry had a better chance of attaining higher SES in a metal age meritocracy and then passing that on?

    If there were "giants" similar in some ways to those of mythology - bigger but maybe a shade dumber - then maybe 20% giant ancestry would be just right (for the metal ages).
  13. @The Beakerist
    Apparently, Reich believes ANE is correlated with R1, except when it's not. Sounds plausible.

    Maybe that implies something e.g. that R1 were adjacent to and mixed with ANE in some places but not others?

    • Replies: @Bultare
    R(and its brother Q) correlate with ANE everywhere except probably Africa. Of course there's also groups with high ANE but low RQ, but there's no patterns in their haplogroups that can be associated with ANE.
  14. @Razib Khan
    Can you give me a reference or tutorial of how you are able to relate South Asian groups (ANI-ASI admixture) in terms of ANE and EEF.


    they used D-statistics. it was on a poster at ASHG 2014. nothing about EEF. but the ANE signal was clear.

    In Indian case, they pick up percentages of invader genetic signatures, at later and later dates.

    i think there might need to be revision in how we understand the arrival of dravidian peoples, and their early relationship to indo-aryans (i don't think dravidian languages are ancient to south asia anymore btw).

    Razib,

    I am reading In Gods We Trust and Atran mentions Dravidians as separate from Aboriginals in India. I had always equated the two. Who are Aboriginal Indians supposed to be, Austonesian people who spoke an Austronesian language who were conquered by Dravidians?

  15. @Matt
    Razib: There are some phenotypic tidbits in the talk apparently. The Yamna were tall in terms of their genomic potential.

    This seems plausible - apparently I think Corded Ware burials worked out about 2.5 inches taller by direct measurement in some study compared to Early Neolithic Farmers in Europe.

    Although I wonder if any of this comes from tendencies for burials to be more disproportionately of higher SES compared to earlier cultures who buried with less regard to social status? Or for burials in high status kurgans to be more archaeologically visible. More social stratification / chiefs / warleaders in Yamna? Height genetic / realised usually seems to go with higher SES. So the Yamna fossils might be tall in their genomic potential more than the Yamna population, as such.

    On the other hand maybe the dudes with more brawn in their ancestry had a better chance of attaining higher SES in a metal age meritocracy and then passing that on?

    If there were “giants” similar in some ways to those of mythology – bigger but maybe a shade dumber – then maybe 20% giant ancestry would be just right (for the metal ages).

  16. @notanon
    Maybe that implies something e.g. that R1 were adjacent to and mixed with ANE in some places but not others?

    R(and its brother Q) correlate with ANE everywhere except probably Africa. Of course there’s also groups with high ANE but low RQ, but there’s no patterns in their haplogroups that can be associated with ANE.

  17. Razib,

    As a frequent lurker, I just wanted to thank you for your outstanding summaries and helpful links. I’m reading Anthony’s book now (on your recommendation) and it is spectacular. Sort of off topic to this post, but totally cool anyway, here is Anthony on what we know by studying Sanskrit:

    Old Indic, the language of the Rig Veda, was recorded in inscription not long after 1500 BCE but in a puzzling place. Most Vedic specialists agree that the 1,028 hymns of the Rig Veda were compiled into what became the sacred form in the Punjab, in northwestern India and Pakistan, probably between about 1500 and 1300 BCE. But the deities, moral concepts, and Old Indic language of the Rig Veda first appeared in written documents not in India but in northern Syria.

    The Mitanni dynasty ruled over what is today northern Syria between 1500 and 1350 BCE. The Mitanni kings regularly spoke a non-Indo-European language, Hurrian, then the dominant local language in much of northern Syria and eastern Turkey. Like Hattic, Hurrian was a native language of the Anatolian uplands, related to the Caucasian languages. But all the Mitanni kings, first to last, took Old Indic throne names, even if they had Hurrian names before being crowned.

    [...]

    Why did Hurrian-speaking kings in Syria use Old Indic names, words, and religious terms in these ways? A good guess is that the Mitanni kingdom was founded by Old Indic speaking mercenaries, perhaps charioteers, who regularly recited the kinds of hymns and prayers that were collected at about the same time far to the east by the compilers of the Rig Veda. Hired by the Hurrian king about 1500 BCE, they usurped his throne and founded a dynasty, a very common pattern in Near Eastern and Iranian dynastic histories.

    - loc 1019 – 1045

  18. “Maybe that implies something e.g. that R1 were adjacent to and mixed with ANE in some places but not others?”

    I think this would be properly called “a weak, positive correlation”. Also, even a positive correlation is not causation.

    Cameroonians, Sardinians and South Asians should have more ANE if the two indicators are dependent. Welsh, Basques, Irishmen should have higher ANE given the non-recombining chromosome in question is in the nineteenth percentile.

    In the last twenty thousand years there is a “weak, positive correlation”. If a Cameroonians can have high prevalence of R-M343 and low/or no ANE, then the same argument is possible for Late Pleistocene individuals in the Southern Pontic-Capsian even though their Siberian ancestors were more positively correlated.

    I’m playing the devil’s advocate, but isn’t it strange that the individuals with the highest saturation of R1b tend to have low or no ANE? (Cameroonians, Basque, Irish)

    • Replies: @Bultare
    Sardinian can mean two things. There are highlander Sardinians who are similar to Neolithic farmers, and the lowlanders who are more like Italians. I would guess it's the lowlanders who have R1b since they have more ANE.
    If you think South Asians and Irish have unusually low ANE you're very confused.
    , @notanon

    I’m playing the devil’s advocate, but isn’t it strange that the individuals with the highest saturation of R1b tend to have low or no ANE? (Cameroonians, Basque, Irish)
     
    Personally I think almost all of it is strange but I'd mention the offspring of male only groups from a high ANE population who married locally might have only half the ANE of males and females from the same group (or possibly more if the local women had more).
    , @Paul Ó Duḃṫaiġ
    I'm Irish (born and living here -- bearing a "native surname" to boot) and on Eurogenes K8 I came in as 15.28% ANE, if anything most of what I've seen points at quite high levels of ANE in NorthWest Europe.

    -Paul
  19. @Helga

    Thanks for the comment. The insights from your comment are the most worthwhile thing I’ve learned this month.

    The distinction, I suspect is between “tribal” people who are outside the caste system and are not really Dalits either, who are called aboriginals somewhat misleadingly, since genetic evidence suggests that they some tribals are fairly recent migrants to India who have “reverted” to pre-farming means of food production (for example, some tribal populations are rich in Y-DNA T compared to non-tribal populations), and Dravidians who would be within the caste system and speak a Dravidian language. Dravidians (especially non-Brahmin Dravidians) have on average less Indo-Aryan genetic ancestry than Indo-Aryan language speakers in India, but all South Asians, regardless of linguistic affiliation have ancestry (some combination of ANI and ASI) that predated Indo-Aryan invasion ca. 2000 BCE, in the vast majority of South Asians, “aboriginal” South Asian ancestry would be a majority.

    There is probably a more respectable term for “tribals” than “tribals” (perhaps Razib knows), but “aboriginal” while a less derogatory word in PC circles, is less accurate because it presumes a prehistory that certainly has not been proven. Calling a tribal South Asian an “aboriginal” is roughly equivalent in historical accuracy to calling a Sardinian a “Caucasian” or calling a Native American an “Indian” or calling a Romani individual a “Gypsy”, all of which are based on ancestry assumptions that have since been proved to be mostly wrong.

    • Replies: @Hipster
    Reading Wikipedia the outline I got was that the first people in the Indian subcontinent were related to Andaman Islanders, and those people could be rights be called "aboriginal". These people today apparently live in "scheduled tribes". These people apparently brought MtDNA haplogroup M, which over half of Indians to this day have.

    Apparently before the arrival of Dravidians or Aryans, Australo-Asiatic speaking people also arrived in India, and these people too are called Aborigonees because they predate Dravidians.

    Then came Dravidians, then Indo-Aryans.

    That is just the wikipedia summary, does that seem to match reality in a rough way, for those who know?
  20. @The Beakerist
    @notanon

    "Maybe that implies something e.g. that R1 were adjacent to and mixed with ANE in some places but not others?"

    I think this would be properly called "a weak, positive correlation". Also, even a positive correlation is not causation.

    Cameroonians, Sardinians and South Asians should have more ANE if the two indicators are dependent. Welsh, Basques, Irishmen should have higher ANE given the non-recombining chromosome in question is in the nineteenth percentile.

    In the last twenty thousand years there is a "weak, positive correlation". If a Cameroonians can have high prevalence of R-M343 and low/or no ANE, then the same argument is possible for Late Pleistocene individuals in the Southern Pontic-Capsian even though their Siberian ancestors were more positively correlated.

    I'm playing the devil's advocate, but isn't it strange that the individuals with the highest saturation of R1b tend to have low or no ANE? (Cameroonians, Basque, Irish)

    Sardinian can mean two things. There are highlander Sardinians who are similar to Neolithic farmers, and the lowlanders who are more like Italians. I would guess it’s the lowlanders who have R1b since they have more ANE.
    If you think South Asians and Irish have unusually low ANE you’re very confused.

  21. @ohwilleke
    @Helga

    Thanks for the comment. The insights from your comment are the most worthwhile thing I've learned this month.

    @Hipster

    The distinction, I suspect is between "tribal" people who are outside the caste system and are not really Dalits either, who are called aboriginals somewhat misleadingly, since genetic evidence suggests that they some tribals are fairly recent migrants to India who have "reverted" to pre-farming means of food production (for example, some tribal populations are rich in Y-DNA T compared to non-tribal populations), and Dravidians who would be within the caste system and speak a Dravidian language. Dravidians (especially non-Brahmin Dravidians) have on average less Indo-Aryan genetic ancestry than Indo-Aryan language speakers in India, but all South Asians, regardless of linguistic affiliation have ancestry (some combination of ANI and ASI) that predated Indo-Aryan invasion ca. 2000 BCE, in the vast majority of South Asians, "aboriginal" South Asian ancestry would be a majority.

    There is probably a more respectable term for "tribals" than "tribals" (perhaps Razib knows), but "aboriginal" while a less derogatory word in PC circles, is less accurate because it presumes a prehistory that certainly has not been proven. Calling a tribal South Asian an "aboriginal" is roughly equivalent in historical accuracy to calling a Sardinian a "Caucasian" or calling a Native American an "Indian" or calling a Romani individual a "Gypsy", all of which are based on ancestry assumptions that have since been proved to be mostly wrong.

    Reading Wikipedia the outline I got was that the first people in the Indian subcontinent were related to Andaman Islanders, and those people could be rights be called “aboriginal”. These people today apparently live in “scheduled tribes”. These people apparently brought MtDNA haplogroup M, which over half of Indians to this day have.

    Apparently before the arrival of Dravidians or Aryans, Australo-Asiatic speaking people also arrived in India, and these people too are called Aborigonees because they predate Dravidians.

    Then came Dravidians, then Indo-Aryans.

    That is just the wikipedia summary, does that seem to match reality in a rough way, for those who know?

    • Replies: @Vijay
    There is some massive confusion in this trail of comments and responses.

    1. The scheduled tribes do not have adirect relationship to aborigines. Even the most ancient of tribals like Pulliyars, Irulas and Malayans, are predominantly Dravidan. The recent tribals who intruded into India from the North and east. None of the tribals have any Austro-Asiatic in them.

    2. Neither the scheduled castes or the BC (or the Sudras) have any significant amount of Austro-Asiatic in them.

    3. The (few) Andaman Islanders who are remnants of Austro-Asiatics are one of a kind. To be precise, Andaman Isalns are closer to the Indonesian Archipealgo in flora, fauna, and peoples than they are to India. The Indians relocation into Andamans is at the best 2200 years, whereas the group that you are talking about inhabited Andamans as early as middle Paleolithic.

    OhWilleke is mixing up things here confusing "Aboriginal" and "Dravidian"; Sravidans populated India much later than Onge. ASI and Onge are close; but they are not the same. Look at "http://www.harappadna.org/wp-content/uploads/2011/05/cline_onge11_sa11_sa12_pca.png". The Pulliyars, Irulars, Malayans will fill in the gap betwee the rest of the Indians and Onge.
  22. @The Beakerist
    @notanon

    "Maybe that implies something e.g. that R1 were adjacent to and mixed with ANE in some places but not others?"

    I think this would be properly called "a weak, positive correlation". Also, even a positive correlation is not causation.

    Cameroonians, Sardinians and South Asians should have more ANE if the two indicators are dependent. Welsh, Basques, Irishmen should have higher ANE given the non-recombining chromosome in question is in the nineteenth percentile.

    In the last twenty thousand years there is a "weak, positive correlation". If a Cameroonians can have high prevalence of R-M343 and low/or no ANE, then the same argument is possible for Late Pleistocene individuals in the Southern Pontic-Capsian even though their Siberian ancestors were more positively correlated.

    I'm playing the devil's advocate, but isn't it strange that the individuals with the highest saturation of R1b tend to have low or no ANE? (Cameroonians, Basque, Irish)

    I’m playing the devil’s advocate, but isn’t it strange that the individuals with the highest saturation of R1b tend to have low or no ANE? (Cameroonians, Basque, Irish)

    Personally I think almost all of it is strange but I’d mention the offspring of male only groups from a high ANE population who married locally might have only half the ANE of males and females from the same group (or possibly more if the local women had more).

  23. Instead of imagining the expansion of these people as purely ones of a vast uniform wave front, it might be better to conceptualize them as penetrating into virgin territory along the optimal avenues of settlement, and producing a patchy archipelago of habitation.

    I like this proposed model a lot. Are there good examples of this in existing historical/anthropological data?

  24. But I think a major issue here is that Indo-European groups were a synthetic population which arose in a world where there were many synthetic populations, with ancient and recent affinities to them. I doubt the “Ancestral North Eurasian” ancestral component was limited purely to Indo-Europeans. So it seems unlikely that the R1 lineages would be purely Indo-European, even if recent expansion of some of their sub-lineages is a function of the Indo-European cultural explosion.

    After studying the history of Plains Indian tribes, that’s how I assumed Indo Europeans came to be. When the Indians got the horse from the Spaniards, they rapidly developed nomadic cultures based on living in the American version of the steppe. But that isn’t where they originally came from. They were from forests to the east and in some cases west.

    So I thought the Indo Europeans probably emerged from the forests upon domestication of the horse, possibly after developing lactose tolerance (drinking horse milk would allow them to live in part off grass). And then when they multiplied and grew in strength, they came charging back in. The entire process, based on what happened in North America, could have happened very quickly. Proto-Indo European could have been born and begun to spread almost at the same time.

    I’m an amateur, but I wonder why I haven’t seen others make the same comparison. Maybe they have and I just missed it.

  25. Oh, one more thing. I just had a thought about using crop genetics to track Indo-European expansion. I have no idea whether anyone has ever looked into this, but there’s a particular kind of fodder native to Europe that follows herds of horses around. It’s called “timothy grass,” and is grown extensively in Kittitas County. Since you lived in Oregon for a while, you may have heard of it, and almost certainly saw bales of it in fields and on trucks.

    It came to the US with settlers, and wasn’t intentionally planted, but was rapidly made use of. It seems that this grass follows herd animals, and horses in particular. It’s also widespread in Ukraine, and may well have originated there.

    Maybe someone could use the genetics of this grass to document the spread of pastoralists across Europe, if indeed it traveled in that manner (as it came to the Americas). It’s an idea, and if it pans out it might shed some light on a thing or two, just as the study of teosinte (ancestor of corn) has furthered our understanding of meso-American agriculture.

  26. @The Beakerist
    @notanon

    "Maybe that implies something e.g. that R1 were adjacent to and mixed with ANE in some places but not others?"

    I think this would be properly called "a weak, positive correlation". Also, even a positive correlation is not causation.

    Cameroonians, Sardinians and South Asians should have more ANE if the two indicators are dependent. Welsh, Basques, Irishmen should have higher ANE given the non-recombining chromosome in question is in the nineteenth percentile.

    In the last twenty thousand years there is a "weak, positive correlation". If a Cameroonians can have high prevalence of R-M343 and low/or no ANE, then the same argument is possible for Late Pleistocene individuals in the Southern Pontic-Capsian even though their Siberian ancestors were more positively correlated.

    I'm playing the devil's advocate, but isn't it strange that the individuals with the highest saturation of R1b tend to have low or no ANE? (Cameroonians, Basque, Irish)

    I’m Irish (born and living here — bearing a “native surname” to boot) and on Eurogenes K8 I came in as 15.28% ANE, if anything most of what I’ve seen points at quite high levels of ANE in NorthWest Europe.

    -Paul

  27. @Hipster
    Reading Wikipedia the outline I got was that the first people in the Indian subcontinent were related to Andaman Islanders, and those people could be rights be called "aboriginal". These people today apparently live in "scheduled tribes". These people apparently brought MtDNA haplogroup M, which over half of Indians to this day have.

    Apparently before the arrival of Dravidians or Aryans, Australo-Asiatic speaking people also arrived in India, and these people too are called Aborigonees because they predate Dravidians.

    Then came Dravidians, then Indo-Aryans.

    That is just the wikipedia summary, does that seem to match reality in a rough way, for those who know?

    There is some massive confusion in this trail of comments and responses.

    1. The scheduled tribes do not have adirect relationship to aborigines. Even the most ancient of tribals like Pulliyars, Irulas and Malayans, are predominantly Dravidan. The recent tribals who intruded into India from the North and east. None of the tribals have any Austro-Asiatic in them.

    2. Neither the scheduled castes or the BC (or the Sudras) have any significant amount of Austro-Asiatic in them.

    3. The (few) Andaman Islanders who are remnants of Austro-Asiatics are one of a kind. To be precise, Andaman Isalns are closer to the Indonesian Archipealgo in flora, fauna, and peoples than they are to India. The Indians relocation into Andamans is at the best 2200 years, whereas the group that you are talking about inhabited Andamans as early as middle Paleolithic.

    OhWilleke is mixing up things here confusing “Aboriginal” and “Dravidian”; Sravidans populated India much later than Onge. ASI and Onge are close; but they are not the same. Look at “http://www.harappadna.org/wp-content/uploads/2011/05/cline_onge11_sa11_sa12_pca.png”. The Pulliyars, Irulars, Malayans will fill in the gap betwee the rest of the Indians and Onge.

    • Replies: @Razib Khan
    The (few) Andaman Islanders who are remnants of Austro-Asiatics are one of a kind.

    no. austro-asiatics are almost certainly agriculturalists who arrived in southeast asia recently, and some of them pushed into india. this is obviously in their Y chr and non-trivial east asian ancestry. the andaman ppl r part of the continuum of 'melanesians' who were absorbed by the austro-asiatics (in vietnam this can be seen in the physical remains 4000 bc as the farmers moved south). & the "ASI" were probably the westward part of this continuum, though they are diverged from the andaman islanders. all of us (S Asians) have part aboriginal, as in ancestry earlier than 10,000 years bp. on the order of half. but none of the aboriginals left a pure genetic and likely cultural signal...
  28. Thanks again Razib. You make interesting times more interesting by allowing fans of good non fiction a far closer look at some of the most interesting cutting edges of science. I am a long time lurker who loves to be updated by the your link laden threads. As they say in the main stream news biz, this is a story with legs. Bronze Age Buttkickers (I’m thinking of a violent horribly politically incorrect video game along the lines of Grand Theft Auto) where you leave your homeland killing men, raping women, and watch your score, total number of descendants living in the present grow and grow. :)

    If Razib deletes this thread I won’t take it too personally. For people who haven’t read “The Horse, the Wheel, and Language I recommend this introduction by David Anthony here https://www.youtube.com/watch?v=HliaR2Ep24s. I will let Anthony speak for himself as to why this population was able to spread so successfully. Yea, going back to being a lurker might be a real good idea.

  29. Razib,

    As one of your more math-challenged readers, I have a question for you:

    It seems that the proto-Indo-Europeans had elevated ANE compared to modern Europeans, but that they were otherwise a mixture of WHG and EEF (or something close to EEF). Given that’s the case, can we estimate what percentage of WHG ancestry in Europe is “indigenous” versus from PIE? Greg Cochran, at least, seems to have been intimating over the last year that he believes virtually no WHG populations survived in the agriculturalist parts of “Old Europe,” and that all WHG is due to later reintroduction. A bold assertion, but one which would align with what happened to hunter-gatherer groups elsewhere in the world.

    • Replies: @Razib Khan
    they are going in that direction too karl. right now the SNP not dense enough/sample not big enough to be able to discern the distinction between EHG non-ANI and WHG. but there is something there, and whole genome sequence could fix this. they talk about this in the supplements. the short of it is that it looks like europe was settled by a small WHG-like population after the ice age. very homogeneous, lots of haplogroup I in males and U5 in females. on the eastern fringe it mixed with ANI-like populations creating the cline. since the non-ANI in EHG is a lot like WHG it is hard to detect, but it will be possible since they separated for several thousand years... also, when you have thicker SNP densities you'll see different patterns of ancestry tracts...
  30. @Vijay
    There is some massive confusion in this trail of comments and responses.

    1. The scheduled tribes do not have adirect relationship to aborigines. Even the most ancient of tribals like Pulliyars, Irulas and Malayans, are predominantly Dravidan. The recent tribals who intruded into India from the North and east. None of the tribals have any Austro-Asiatic in them.

    2. Neither the scheduled castes or the BC (or the Sudras) have any significant amount of Austro-Asiatic in them.

    3. The (few) Andaman Islanders who are remnants of Austro-Asiatics are one of a kind. To be precise, Andaman Isalns are closer to the Indonesian Archipealgo in flora, fauna, and peoples than they are to India. The Indians relocation into Andamans is at the best 2200 years, whereas the group that you are talking about inhabited Andamans as early as middle Paleolithic.

    OhWilleke is mixing up things here confusing "Aboriginal" and "Dravidian"; Sravidans populated India much later than Onge. ASI and Onge are close; but they are not the same. Look at "http://www.harappadna.org/wp-content/uploads/2011/05/cline_onge11_sa11_sa12_pca.png". The Pulliyars, Irulars, Malayans will fill in the gap betwee the rest of the Indians and Onge.

    The (few) Andaman Islanders who are remnants of Austro-Asiatics are one of a kind.

    no. austro-asiatics are almost certainly agriculturalists who arrived in southeast asia recently, and some of them pushed into india. this is obviously in their Y chr and non-trivial east asian ancestry. the andaman ppl r part of the continuum of ‘melanesians’ who were absorbed by the austro-asiatics (in vietnam this can be seen in the physical remains 4000 bc as the farmers moved south). & the “ASI” were probably the westward part of this continuum, though they are diverged from the andaman islanders. all of us (S Asians) have part aboriginal, as in ancestry earlier than 10,000 years bp. on the order of half. but none of the aboriginals left a pure genetic and likely cultural signal…

    • Replies: @Vijay
    This is news to me. Are you saying that ASI arrived in India from the east. I assumed based on Y dna that it arrived from the Middle East. I just wish there will be as much focus on ASI as much as on ANI.
    , @Vijay
    From running Dienekes K12b calculator a while ago, I was thinking (http://dienekes.blogspot.com/2012/08/inter-relationships-of-dodecad-k12b-and.html) that ASI (somewhat similar to Dienekes South Asian) was equal parts Caucasus-Gedrosian and East Asian with just a small portion of Australo-Asiatic. However, it has been a while since I have run that calculator, and so I might be wrong.
  31. @Karl Zimmerman
    Razib,

    As one of your more math-challenged readers, I have a question for you:

    It seems that the proto-Indo-Europeans had elevated ANE compared to modern Europeans, but that they were otherwise a mixture of WHG and EEF (or something close to EEF). Given that's the case, can we estimate what percentage of WHG ancestry in Europe is "indigenous" versus from PIE? Greg Cochran, at least, seems to have been intimating over the last year that he believes virtually no WHG populations survived in the agriculturalist parts of "Old Europe," and that all WHG is due to later reintroduction. A bold assertion, but one which would align with what happened to hunter-gatherer groups elsewhere in the world.

    they are going in that direction too karl. right now the SNP not dense enough/sample not big enough to be able to discern the distinction between EHG non-ANI and WHG. but there is something there, and whole genome sequence could fix this. they talk about this in the supplements. the short of it is that it looks like europe was settled by a small WHG-like population after the ice age. very homogeneous, lots of haplogroup I in males and U5 in females. on the eastern fringe it mixed with ANI-like populations creating the cline. since the non-ANI in EHG is a lot like WHG it is hard to detect, but it will be possible since they separated for several thousand years… also, when you have thicker SNP densities you’ll see different patterns of ancestry tracts…

  32. I see no mention of Y haplogroup I1. Were they the earlier hunter gatherers (not Indo-Europeans) that the R1b pushed around in Western Europe? Hard to imagine those Vikings getting pushed around too much.

    • Replies: @Razib Khan
    haplogroup I is from the original WHG population, yes. nick patterson told me he believes it is pretty diagnostic of someone having paleolithic european ancestry in ancient DNA.
  33. @Razib Khan
    The (few) Andaman Islanders who are remnants of Austro-Asiatics are one of a kind.

    no. austro-asiatics are almost certainly agriculturalists who arrived in southeast asia recently, and some of them pushed into india. this is obviously in their Y chr and non-trivial east asian ancestry. the andaman ppl r part of the continuum of 'melanesians' who were absorbed by the austro-asiatics (in vietnam this can be seen in the physical remains 4000 bc as the farmers moved south). & the "ASI" were probably the westward part of this continuum, though they are diverged from the andaman islanders. all of us (S Asians) have part aboriginal, as in ancestry earlier than 10,000 years bp. on the order of half. but none of the aboriginals left a pure genetic and likely cultural signal...

    This is news to me. Are you saying that ASI arrived in India from the east. I assumed based on Y dna that it arrived from the Middle East. I just wish there will be as much focus on ASI as much as on ANI.

  34. Apologies if this comes off as a bit cranky, but more on the timothy grass:

    In RAPD, UP-PCR and combination dendrograms, the northern Europe geographical races (i.e. Norwegian and Finnish), the central Europe geographical races (i.e. German and Dutch), Canadian geographical races and Japanese geographical races were distinguished. The distribution of these races seemed no correlation with climatic or soil data. Perhaps the present-day distribution of the population of timothy is not the result of a process of selection for different gene combinations in different geographical areas, but as the result of a past migration [emphasis mine]. This hypothesis of timothy is similar with that of the weed Silene latifolia which spread over Europe (vellekoop et al. 1996) and Finnish tansy (keskitalo et al. 1998).

  35. @ohwilleke
    Good post with good caveats, e.g. about the possible bias of data from high status graves.

    I also share some skepticism that R1b in Western Europe has a source in Indo-Europeans linguistically, despite the fact that one can make a case for an Eastern Europe to Central Europe to Southern Europe pathway based on R1b phylogeny.

    On the other hand, it does seem pretty clear that the source of Europe's R1a and R1b were not that far from each other geographically, and probably absorbed fitness enhancing technologies with a common source.

    Also, re Genesis 6:4, one of the groups associated with Giants in Biblical times were the Philistines (e.g. Goliath). And archaeology and Egyptian historical sources now strongly point to them being Bronze Age collapse era migrants (ca. 1200 BCE) who were ethnically Mycenean Greeks, i.e. Indo-Europeans, who started to arrive in the Greek mainland ca. 2000 BCE (800 years earlier) and in all of the Aegean including Crete by about 1500 BCE.

    Arguably the passage was written by a Hellenized Jew, the Nephilim being a sort of reductive appropriation of Homer’s “men of reknown.”

    Certainly whoever authored Goliath had seen a hoplite.

  36. @Hannah Katz
    I see no mention of Y haplogroup I1. Were they the earlier hunter gatherers (not Indo-Europeans) that the R1b pushed around in Western Europe? Hard to imagine those Vikings getting pushed around too much.

    haplogroup I is from the original WHG population, yes. nick patterson told me he believes it is pretty diagnostic of someone having paleolithic european ancestry in ancient DNA.

    • Replies: @Bultare
    C-V20 might be a better candidate if our definition of the original WHG population extends further back in time than the LGM.
    Its rarity might not be the result of population replacement but perhaps something about it that makes it just a little bit less fit, perhaps an unfortunate but not crippling mutation that was followed by a bottleneck. Its nearest cousin C-M8 in Japan isn't doing all that great either.
  37. @Razib Khan
    The (few) Andaman Islanders who are remnants of Austro-Asiatics are one of a kind.

    no. austro-asiatics are almost certainly agriculturalists who arrived in southeast asia recently, and some of them pushed into india. this is obviously in their Y chr and non-trivial east asian ancestry. the andaman ppl r part of the continuum of 'melanesians' who were absorbed by the austro-asiatics (in vietnam this can be seen in the physical remains 4000 bc as the farmers moved south). & the "ASI" were probably the westward part of this continuum, though they are diverged from the andaman islanders. all of us (S Asians) have part aboriginal, as in ancestry earlier than 10,000 years bp. on the order of half. but none of the aboriginals left a pure genetic and likely cultural signal...

    From running Dienekes K12b calculator a while ago, I was thinking (http://dienekes.blogspot.com/2012/08/inter-relationships-of-dodecad-k12b-and.html) that ASI (somewhat similar to Dienekes South Asian) was equal parts Caucasus-Gedrosian and East Asian with just a small portion of Australo-Asiatic. However, it has been a while since I have run that calculator, and so I might be wrong.

  38. @Razib Khan
    haplogroup I is from the original WHG population, yes. nick patterson told me he believes it is pretty diagnostic of someone having paleolithic european ancestry in ancient DNA.

    C-V20 might be a better candidate if our definition of the original WHG population extends further back in time than the LGM.
    Its rarity might not be the result of population replacement but perhaps something about it that makes it just a little bit less fit, perhaps an unfortunate but not crippling mutation that was followed by a bottleneck. Its nearest cousin C-M8 in Japan isn’t doing all that great either.

  39. How would the increasing frequency of blood type O in western Europe be related to R1b? The two do corelate. My understanding is that the Yamna proto indo European people spread blood type b in eastern europe, Scandinavia (to a lesser extent) and into the sub continent and Iranian plateau.

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