Tuesday, February 10, 2015

Massive migration from the steppe is a source for Indo-European languages in Europe (Haak et al. 2015 preprint)


I'll probably end up writing a whole series of posts on this paper. But for now, here's the abstract and a PCA.

We generated genome-wide data from 69 Europeans who lived between 8,000-3,000 years ago by enriching ancient DNA libraries for a target set of almost four hundred thousand polymorphisms. Enrichment of these positions decreases the sequencing required for genome-wide ancient DNA analysis by a median of around 250-fold, allowing us to study an order of magnitude more individuals than previous studies and to obtain new insights about the past. We show that the populations of western and far eastern Europe followed opposite trajectories between 8,000-5,000 years ago. At the beginning of the Neolithic period in Europe, ~8,000-7,000 years ago, closely related groups of early farmers appeared in Germany, Hungary, and Spain, different from indigenous hunter-gatherers, whereas Russia was inhabited by a distinctive population of hunter-gatherers with high affinity to a ~24,000 year old Siberian6. By ~6,000-5,000 years ago, a resurgence of hunter-gatherer ancestry had occurred throughout much of Europe, but in Russia, the Yamnaya steppe herders of this time were descended not only from the preceding eastern European hunter-gatherers, but from a population of Near Eastern ancestry. Western and Eastern Europe came into contact ~4,500 years ago, as the Late Neolithic Corded Ware people from Germany traced ~3/4 of their ancestry to the Yamnaya, documenting a massive migration into the heartland of Europe from its eastern periphery. This steppe ancestry persisted in all sampled central Europeans until at least ~3,000 years ago, and is ubiquitous in present-day Europeans. These results provide support for the theory of a steppe origin of at least some of the Indo-European languages of Europe.


Haak et al., Massive migration from the steppe is a source for Indo-European languages in Europe, bioRxiv, Posted February 10, 2015, doi: https://dx.doi.org/10.1101/013433

438 comments:

  1. " whereas Russia was inhabited by a distinctive population of hunter-gatherers with high affinity to a ~24,000 year old Siberian"


    Weren't some people arguing that Russian H&G must have been up to 80% WHG?

    The WHG story seems to lose on ground more and more.

    I stay at my estimation of 20 to max 30% WHG among Yamna.

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  2. Also

    "These results provide support for the theory of a steppe origin of at least some of the Indo-European languages of Europe."

    Seems to me like not all Indo European languages can be explained with a Steppe origin?

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  3. Kurti my friend, embrace your Eastern Euro heritage.

    The Karelian and Samara foragers are R1a and R1b. So a lot of people have Eastern Euro ancestors, it seems.

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  4. Looks like R1a was the northern brother of R1b here, and now it's the eastern brother, haha. So R1a took the shorter, northern route and acquired only a little Neolithic DNA, and i guess R1b took the southern route through more populous areas and acquired much more?

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  5. Amazing that Samara was R1B. Amazing.

    I love science. :)

    So how did Eastern Europe end up mostly R1a today? Did R1B invade Europe first while R1A followed a little later, eradicating their "cousins" in their wake? Or did R1B move south and around R1A territory?

    Interesting!

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  6. Jesus on Anthrogenica picked up on this:

    This pattern is also seen in ADMIXTURE analysis (Fig. 2b, SI6), which implies that the Yamnaya have ancestry from populations related to the Caucasus and South Asia that is largely absent in 38 Early or Middle Neolithic farmers but present in all 25 Late Neolithic or Bronze Age individuals.

    Sounds like the Caucasus/Gedrosia thing perhaps?

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  7. @David
    ''The Karelian and Samara foragers are R1a and R1b. So a lot of people have Eastern Euro ancestors, it seems.''
    From Which Date?

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  8. '' which implies that the Yamnaya have ancestry from populations related to the Caucasus and South Asia that is largely absent in 38 Early or Middle Neolithic farmers but present in all 25 Late Neolithic or Bronze Age individuals.''
    OMG Let me check.

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  9. Holy Bananas 5000 BC R1b in Spain!?!??!?!?!?!??!?!?!??!???!?!?!??!?!?!?!?!?!? Throw out everything!!!!!!!!!

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  10. Nirjhar,

    You can check the dates in the study. But the R1a Karelian hunter-gatherer is ~7,550 years old.

    It looks like R1 is native to Eastern Europe and maybe Western Siberia. Forget India or even Iran.

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  11. R1b in Europe is EHG, I'm very surprised. My maternal line is WHG and my paternal line is EHG.

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  12. "Holy Bananas 5000 BC R1b in Spain!?!??!?!?!?!??!?!?!??!???!?!?!??!?!?!?!?!?!? Throw out everything!!!!!!!!!"

    That R1b along with the R1b from Karelia(I was think N and I) is just labeled as R1b1. It's going to take hour and hours to understand this paper.

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  13. Congrats Davidski, for seeing the connection between MA-1 and modern R so long ago.

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  14. ''It looks like R1 is native to Eastern Europe and maybe Western Siberia. Forget India or even Iran.''
    NO Sir it all within my Conclusion in case you missed!

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  15. The Yamnaya is all Z2103, P312 is in Bell Beaker 2200 B.C., looks like R1b branched off into Europe before the Yamnaya existed......

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  16. "Western and Eastern Europe came into contact ~4,500 years
    ago, as the Late Neolithic Corded Ware people from Germany traced ~3/4 of their
    ancestry to the Yamnaya, documenting a massive migration into the heartland of
    Europe from its eastern periphery."

    The data in this paper do not support the assertion of "mass migration into Europe approximately 4,500 years ago" "from an eastern periphery".

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  17. Derived I2a1b clades, I2a2, T1a, and strange R clades in EEFs is big news. Don't forget about I1.

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  18. Just saw an H2 in Y DNA. Big surprise. No J though.

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  19. '' strange R clades in EEFs is big news.''
    yes but according to the Migration from West Asia likely ( The Earlier).

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  20. Davidski That allot of my ancestry came from the Pontic-Caspian Steppes is without doubt. But the paper indiciates that not every Indo European group originates from the Steppes or can be explained with a Steppe origin.

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  21. @Romulus That the Bell Beaker with P312 is predominantly Yamnaya would need to be explained. Whats the chance he got it from a rare R1b in western Europe, or one part of the group in which they couldn't move for M269. All that Yamnaya specific R1b would have had to die off in western Europe, with not much impact autosmally.

    Seems much more likely to me that there may have been R1b sub-groupings within Yamnaya, it's quite a big complex - perhaps the ones directly ancestral to Bell Beaker were west/northwest of the Black Sea, rather than north of the Caspian?

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  22. @PersonaMan

    Yup sounds like Caucasus_Gedrosia. More so Gedrosia.

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  23. @Krefter
    One I1 from the Neolithic in the Carpathian basin i think it was...seems to be elusive lol.

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  24. Romulus,

    Are you sure all of the Yamnaya are Z2103?

    Anyway, it's interesting that M412 is missing. Here's a very important quote from the study. Page 47:

    "Summarizing the results from the Yamnaya males, all seven belonged to haplogroup R1b1a. Six of these could be further assigned to haplogroup R1b1a2a, and five of these to haplogroup R1b1a2a2. The uniformity of R1b Y-chromosomes in this sample suggests a patrilineal organization of the Yamnaya, or at least of the people who were given expensive Kurgan burials. We cannot exclude the presence of other haplogroups in the general population, or in other individuals located elsewhere in the expansive Yamnaya horizon23. We also emphasize the absence of M412 (the dominant lineage within haplogroup R-M269 in Europe) in this sample, as well as the absence of the R1a haplogroup which was detected in the Corded Ware and Late Bronze Age Halberstadt individual from central Europe. A survey of other European steppe groups may reveal the more immediate patrilineal kin of the major founding lineages of modern European R1a and R1b chromosomes."

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  25. @Kurti, which makes it seem much more likely that P312 is a descendant of steppe R1b rather than Iberian Neolithic, especially when it peaks locally in NW Europe, cannot be explained unless their R1b came from the Steppe after ~3000bc.

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  26. Dienekes Will Be Pretty happy also!

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  27. Kurti,

    The paper most certainly doesn't indicate that not every Indo-European group originated from the steppe or can be explained with a steppe origin.

    What it says is that more data is needed to completely nail the steppe hypothesis.

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  28. @PersonaMan

    "@Kurti, which makes it seem much more likely that P312 is a descendant of steppe R1b rather than Iberian Neolithic, especially when it peaks locally in NW Europe, cannot be explained unless their R1b came from the Steppe after ~3000bc."

    Megalithicism appears almost simultaneously at the threshold of the Neolithic across the Mediterranean.

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  29. The more southern distribution of Gedrosia in Asia compared to say Caucasus also fits with Gedrosia being stronger in Europe in R1b-dominated populations, and with R1b being located to the south of R1a in antiquity. Perhaps also why it may have been picked up by early Neolithic peoples moving through part of the Near East to Europe (Iberian R1b1)?

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  30. Significant-
    ''the signal of admixture from the Yamnaya is not the strongest one for Armenians. Moreover, as shown in SI 7, the Yamnaya have a negative f3-
    statistic with (X, Y) = (Karelia_HG, Armenian). A negative statistic for both Armenians and Yamnaya
    with each other as a reference population may suggest that a third (unsampled) population admixed
    into both the Yamnaya and to Armenians. The question of directionality can only be furthered
    elucidated by the study of additional ancient samples from the Caucasus, Near East and the steppe.''
    SC-Asia Central Asia?

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  31. The list Jean posted makes sense now, it was based on levels of Yamnaya ancestry, not levels of purely ANE ancestry.

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  32. @Marnie

    Of what significance is this specifically? I was basically making the point that the authors mention an admixture in the Steppe of Caucasus-Gedrosia likeness at a certain date - that this doesn't seem to be present in the Neolithic Farmers of Europe to date (esp. Gedrosia) iirc, or at least not nearly as much as today. So the ancestors of western European R1b are likely to have been one of the groups affected by this, as Gedrosia-like components have a local peak in western Europe, which is highest in the British Isles - which also has extremely high levels of P312. So if local R1b (of which there seems to be a little at most) is the ancestor of western European R1b - how is this, and the substantial Yamnaya contribution explained?

    It almost puts a terminus ante quem on where western European R1b's ancestors must have been prior to the late Neolithic, in order to be affected by this. Unless these autosomal artefacts were carried into western Europe by members of R1b groups that mysteriously died out, or of other lineages which are not in evidence as of yet. Seems unlikely to me.

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  33. @Davidski,

    Is R1b1a2a2 not Z2103? all of them fall under it but 2 which are ancestral. If I am wrong please explain but this seems pretty straightforward.

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  34. @David
    ''The paper most certainly doesn't indicate that not every Indo-European group originated from the steppe or can be explained with a steppe origin.

    What it says is that more data is needed to completely nail the steppe hypothesis.''
    I Agree its the IE Folks of Europe that is mostly concerned here or should i say only? The thing however is that they Lack knowledge from Asian Side which biased and flattered their conclusions for sure....

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  35. They're not all z2103

    I0439 (Yamnaya) This individual could be assigned to haplogroup R1b1a (P297:18656508G→C), with upstream haplogroup R1 (M173:15026424A→C, M306:22750583C→A) also supported. It was ancestral for haplogroup R1b1a2a1 (L51:8502236G→A) and so could be designated R1b1a*(xR1b1a2a1).


    I0443 (Yamnaya) This individual could only be assigned to haplogroup R1b1a2a (L49.1:2842212T→A, L23:6753511G→A). It could also be assigned to the upstream haplogroups R1b1a2 (PF6399:2668456C→T, L150.1:10008791C→T, L1353:19179540G→A, PF6509:22190371A→G, M269:22739367T→C, CTS12478:28590278G→A). The individual was ancestral for haplogroup R1b1a2a1 (L51/M412:8502236G→A) and, unlike I0231, I0370 and I0438 also for R1b1a2a2 (Z2105:15747432C→A). Thus, it could be designated as R1b1a2a*(xR1b1a2a1, R1b1a2a2)

    I0443 is one step from L51 and could very well be under L51 with more digging.

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  36. That R1b in Spain is not ancestral to almost all of us. There are a few in modern Spain. Some EHG looks like it got into Europe.

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  37. If you look at the high def supplement of the admixtures, the Yamnaya are 50% WHG and 50% of a population most predominant in the Kalash. Most interestingly of all they are 0% EEF.

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  38. @Mike Thomas

    "But maybe there was no single source for European R1b"

    Yes, I don't think we should assume any "single" source. I'm just pointing out that there seems to have been something happening in the Mediterranean at the threshold of the Neolithic, where megalithic sites demonstrate very similar architectures and rock art.

    So, if R1b [and other lineages] were associated with this, there's no need to assume a directionality over time and space of associated haplogroups.

    For a good reference, see "Megalithic Rock Art of the Mediterranean and Atlantic Seaboard Europe" by George Nash, in "A Companion to Rock Art", Jo McDonald and Peter Veth, editors, 2012.

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  39. @Chad,

    So what if 2 of them are ancestral to either Z2103 or L51, 7 others are Z2103. Its clear from the data that there was an R1b population ancestral to both the Yamnaya and the 5000 B.C. R1b in Spain that spread out Prior to 5000 B.C.

    I think we are seeing evidence that L51 evolved in Western Europe Possibly as early as the Megalithic era.

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  40. @Romulus
    ''If you look at the high def supplement of the admixtures, the Yamnaya are 50% WHG and 50% of a population most predominant in the Kalash.''
    What is your judgement on that observation?

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  41. You can't get L51 without L49... how the hell do you think a very old clade of R1b can just jump to L51?? You're off your rocker. There are a few of these that survive in Spain. They did not turn into European R1b. m343, L278, L49 are on the Steppes.

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  42. @Nirjhar007

    in the K19 admixtures, the Yamnaya are 50% blue and 50% teal. WHGs like Loschbour or Motala are 100% blue. The population with the most teal is the Kalash. EEF is the orange admixture and the Yamnaya have none of it.

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  43. I0410 (Spain_EN) We determined that this individual belonged to haplogroup R1b1 (M415:9170545C→A), with upstream haplogroup R1b (M343:2887824C→A) also supported. However, the individual was ancestral for R1b1a1 (M478:23444054T→C), R1b1a2 (PF6399:2668456C→T, L265:8149348A→G, L150.1:10008791C→T and M269:22739367T→C), R1b1c2 (V35:6812012T→A), and R1b1c3 (V69:18099054C→T), and could thus be designated R1b1*(xR1b1a1, R1b1a2, R1b1c2, R1b1c3).

    He is a long ways from "European R1b". L51 can't come from him, but can from the Steppe L49.

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  44. The fact that R1b was found in Neolithic Spain as well Yamna. Speaks for the theory of Maciamo on Eupedia. That R1b came from North mesopotamia. Went to maykop and arrived in Yamna. This would explain the dual source of R1b.

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  45. Bell Beaker R1b 2,500YBP being specifically "Italo-Celtic" P312 is big news. It suggests an early spread of L11, because it had already split by that time.

    The diversity of P312 in west Europe today strongly suggests an early bronze age spread.

    The Y DNA from bell beaker in this study hasn't been tested before. This makes Bell beaker Y DNA 3/3 R1b.

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  46. The R1a1 in Karelia seems to have been put aside slightly, any interesting comments regarding that?

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  47. "The fact that R1b was found in Neolithic Spain as well Yamna. Speaks for the theory of Maciamo on Eupedia. That R1b came from North mesopotamia. Went to maykop and arrived in Yamna. This would explain the dual source of R1b."

    I agree it means R1bs story is more complicated. If the R1b1 individual from Spain has no ANE, that's big news. The R1b1 from Spain is about as old as the R1b1 from Russia, but the L23 from Yamna is good evidence L11 in Europe today comes from the steppe and isn't native.

    I'm very suprised about Z2103, which I always though was west Asian-specific is found way up north in Samara over 5,000 years ago. It could have come there with Yamna's "Armenian" ancestors or it could be from EHG.

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  48. no... EHG has no Basal Eurasian. R1b is not from the Middle East. You can't have a Mesolithic R1b with no Basal, if their ancestors were from there.

    What needs to happen is for David to test that R1b Neolithic for ANE, to see if some ANE or EHG was in Western Europe before farming, or a stray EHG that wandered into the path of farmers and transported to Iberia. Either way, that Iberian is not ancestral to almost all of us.

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  49. @Chad,

    I'm not an expert on R1b, however the 2200BC P312 in Bell Beaker and the R1b1 in 5000 B.C. spain seems to me to imply that P312 at least and likely L51 have a Western European origin.

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  50. @Chad

    "He is a long ways from "European R1b". L51 can't come from him, but can from the Steppe L49."

    Sure, if you're looking for your direct ancestor, then maybe this R1b in Iberia isn't very interesting.

    But some of us are not looking for a direct ancestor. We're trying to understand the formation of Europe.

    The fact that R1b (whether upstream from L51 or now) is interesting toward understanding mobility in the Early Neolithic.

    It speaks to the layer process that Mike Thomas has mentioned, and which I also support.

    . . . lost on you?

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  51. @krefter

    I honestly doubt that R1b in Spain had not already some ANE, simply because the source which brought the "Near Eastern" ancestry to Yamna was already "ANE rich" even if not as much as the "EHG" people.

    Also another thing from the paper.

    "The Armenian plateau hypothesis gains in plausibility by the fact that we have discovered
    evidence of admixture in the ancestry of Yamnaya steppe pastoralists, including gene flow
    from a population of Near Eastern ancestry for which Armenians today appear to be a
    reasonable surrogate "

    I doubt,like many people that the "Near Eastern" source was Armenian like. Seems like they were taken up simply from the fact that they speak Indo European. But the plausibility of an West Asian pastoralist origin of proto Indo European seems to have risen also.

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  52. Romulus, that Iberian R1b is very old! It is tons of snp's short of L51. The only one that is one step short is the Yamnaya sample. That Yamnaya could end up L51 or with more samples have an L51. There are plenty of his brother z2103. There should then be some L51 there.

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  53. David,
    Do you have the genomes?

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  54. In regards to the Karelian R1a, it appears to be from an early basal split and not closely related to the Kurgan, Balto-Slavic, Norse and Indo-Iranian R1a-M417.

    However, the presence of basal R1a and R1b in Eastern European foragers suggests that R1 is native to Europe, and probably expanded from there via different subclades, mainly during the Copper Age.

    By the way, those Yamnaya folks don't look ancestral to Western Europeans based on a quick look at their data. The seem to be proto-Anatolians and proto-Armenians IMO. But I'm sure that testing different parts of the Yamnaya horizon will find both R1b-M412 and also R1a-M417, which now dominate Europe and most Indo-European speaking groups.

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  55. I don't have the genomes. They won't be available until the paper is published in a journal.

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  56. @Wesolowski
    ''However, the presence of basal R1a and R1b in Eastern European foragers suggests that R1 is native to Europe, and probably expanded from there via different subclades, mainly during the Copper Age.''
    No R1a-420 if i'm not wrong? and how you can conclude R1 and R1b to be native? Without similar aDNA data from West Asian Areas?.
    ''By the way, those Yamnaya folks don't look ancestral to Western Europeans based on a quick look at their data. The seem to be proto-Anatolians and proto-Armenians''
    Yup.
    '' But I'm sure that testing different parts of the Yamnaya horizon will find both R1b-M412 and also R1a-M417, which now dominate Europe and most Indo-European speaking groups.''
    Lets see...
    BTW Where is Z-93 David?

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  57. One Yamnaya is one step from L51... Most Yamnaya are z2103, brother to L51!!! Beakers are 50% Yamnaya... it doesn't take Einstein to figure out this one!

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  58. @Davidski

    "However, the presence of basal R1a and R1b in Eastern European foragers suggests that R1 is native to Europe, and probably expanded from there via different subclades, mainly during the Copper Age."

    "By the way, those Yamnaya folks don't look ancestral to Western Europeans based on a quick look at their data. The seem to be proto-Anatolians and proto-Armenians IMO. But I'm sure that testing different parts of the Yamnaya horizon will find both R1b-M412 and also R1a-M417, which now dominate Europe and most Indo-European speaking groups."


    Important results that jibe with other papers (Myers 2010, in particular, and Balaresque Jan 2015) and the archaeological record.

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  59. Does anyone know what format Online Table 2 in the supplementary info is? I can't get it to open.

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  60. @Romulus IN the population paper here.

    Groups such as Lithuanians who have ~50% WHG are only ~25% blue, what brings us to the conclusion that the reast of the WHG came from the Yamna source. But this doesn't fit the ~50% WHG estimation.

    http://s1133.photobucket.com/user/jeanlohizun/media/Haaketal2015-Figure-3_zpsf94c99b9.jpg.html

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  61. The comments on the ADMIXTURE output are interesting:

    "An interesting pattern occurs at K=8, with all the late LN/BA groups from central Europe and the Yamnaya having some of the “light green” component that is lacking in earlier European farmers and hunter-gatherers; this component is found at high frequencies in South Asian populations...At K=9...Note, also, the persistent presence of the “light green” component that ties LN/EBA groups to South Asia between K=9 and K=15."

    I think it was premature of me to say that the early Indo-Europeans contributed no substantial ancestry to Central/South Asians. Based on these results, the early Indo-Europeans were quite genetically "Indo"! Then again, this could just be a spurious result, the product of extremely high ANE ancestry in South Asia creating an illusion of relationship between these ANE-rich groups, and not a reflection of truly direct ancestral connections between South Central Asians and the early Indo-Europeans.

    I think we'll find out, when David gets his hands on these samples.

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  62. I think one of the most unaddressed things in this paper is that the EEF admixture PEAKS in Europeans and is only found at small mixtures elsewhere in the world specifically the Near East. I think this is a great Candidate for the light skin Gene.

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  63. @Steinundzeit

    from the wording and paper.

    "A Near Eastern people"

    It seems like this "South Asian" component was definitely Gedrosia. What makes most sense. Since this component peaks in a region from the Caucasus all the way into South_Central Asia.

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  64. @Kurti

    The component isn't very Gedrosia-like until K=16. With K=16, they note:

    "A similar (darker green) component also distinguishes LN/EBA groups from earlier ones at K=16; this component appears to be highly represented in groups from South Asia, the Near East, and the Caucasus."

    This is a Gedrosian/Afghan/Pakistani component. The other quote involved K=8+9, which have a straightforward South Asian (peaks in South Indians) component.

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  65. @Chad
    ''One Yamnaya is one step from L51''
    Chad isn't R1b1a2a which is the Ancestor of L51 is found in very high frequency in Caucasus the variance of R1b-L23 (the main clade of M269) is very high in the Caucasus but it is highest in Pakistan!
    and Grugni et al. write: "the variance distribution of the rare R1b-M269* Y chromosomes, displaying decreasing values from Iran, Anatolia and the western Black Sea coastal region, is also suggestive of a westward diffusion from the Iranian plateau. Correct me if i'm wrong

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  66. Highest diversity of L23 is now Bulgaria. Who knows 5kya, yamnaya!?!? Iran?!? Please... Samara 5500bce with no basal Eurasian.

    David!!
    Look at that one MNE German!!! Might be 70% WHG on the K8!

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  67. @Chad
    Also Kurds and Iranians have high percentages of R1b-M343*, the ancestral haplogroup of R1b. Kurds from Kazakhstan, mainly deported there from Soviet Caucasian countries, have 13% of this haplogroup, a frequency with no match in other populations. However, this haplogroup is found also in Iranian populations: 4.3% in Persians of Yazd, 3.2% in Azeris,.

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  68. "EEF admixture PEAKS in Europeans and is only found at small mixtures elsewhere in the world specifically the Near East."

    So can we stop calling them farmers now? For Oetzi's sake?

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  69. Give it up Nirjhar.. We have R1b as EHG.. No near eastern ancestry..

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  70. @Chad
    '' Who knows 5kya, yamnaya!?!? Iran?!? Please... Samara 5500bce with no basal Eurasian. ''
    I don't understand:) what are you trying to say?.

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  71. Basal r1b is in Samara.... No near Eastern ancestry... Pure EHG... Got it??

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  72. What Chad is saying is that it is illogical and anachronistic to suggest that the R1b found in the Yamnaya remains was of non-local origin since we have an ancestral R1b in Eastern European Hunter-Gatherer remains from Samara Valley.

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  73. @ZGN

    "What Chad is saying is that it is illogical and anachronistic to suggest that the R1b found in the Yamnaya remains was of non-local origin since we have an ancestral R1b in Eastern European Hunter-Gatherer remains from Samara Valley."

    Huh?

    Wouldn't it then be reasonable to say that Yamnaya *could* be local, but *could* also be from anywhere between Eastern Europe and the Samara Valley? Maybe even Armenia?

    I think you guys should also dump the "basal" thing. How about "ancestral"?

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  74. @Chad

    YOU ARE WRONG.

    This is what you said: "Basal r1b is in Samara.... No near Eastern ancestry... Pure EHG... Got it??"

    Ebizur's comment from the other post:

    "All seven tested Yamnaya individuals belong to R1b1a-P297. One Yamnaya individual is confirmed R1b1a2-M269(xL51/M412, Z2105). The haplogroup of one Yamnaya individual could only be confirmed so far as R1b1a-P297(xL51/M412). The other five Yamnaya individuals are confirmed R1b1a2a2-CTS1078/Z2103."

    "R1b1a2a2-CTS1078/Z2103 is not the form of R1b that is common in (Western) Europe, but rather the form that is common in Armenia, northern Iran, the eastern Caucasus, etc. It is the typically "West Asian" sort of R1b. Thus, at least this sampled group of Yamnaya males is patrilineally more closely related to many Armenians, Assyrians, etc. than it is patrilineally related to Western Europeans."

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  75. Marnie, R1b is pure EHG in Samara. Think about it... It's like some jackass saying that they pick i2 as Neolithic even though we have pure WHG with it... You can't put the cart before the horse. R1b is not from west Asia..and it is connected to ANE....logic evades some

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  76. R1b is very old and there is no reason to suppose it was restricted to the Caspian steppe. So while obviously a local EHG source is plausible, there is nothing to rule out the Yamnaya R1b being intrusive from West or Central Asia. We won't know until we have aDNA from potential source regions - which sounds like it won't be long.

    ReplyDelete
  77. capra,

    Take a look at this PCA. Note how the West Asians are pulling towards an area of the plot that could be the Ural steppe.

    https://drive.google.com/file/d/0B9o3EYTdM8lQREh3dTBFS21FREE/view?usp=sharing

    This is where they got their ANE from, don't you think? So which paternal marker was involved in that migration from the steppe?

    Maybe R1, which we now know is native to Eastern Europe?

    ReplyDelete
  78. I don't think I can feel any more vindicated that I do now.

    https://distantconnections.wordpress.com/25-2/

    Not only did I think r1b was in yamnaya (as many others argued) but I said r1b dominated yamnaya and that r1a was lacking in yamnaya. Im quite sure thats true. I think the IE type of r1a was to the northwest.

    ReplyDelete
  79. Very old? This paper is matching snp dates pretty well. M343 is 8000bce in that method. We've got z2103 400 years younger than the expected but within range.

    ReplyDelete
  80. I organized all the Y DNA results in this paper. There are some edits that have to do with neatness I have correct but besides that it's all good.

    https://docs.google.com/document/d/1yZT6gFhWfWoOPAcmQe6edjmA-VM6F4BI4ezn0kOU91Q/edit?usp=sharing

    ReplyDelete
  81. Colin,

    I already told you why this was extremely unlikely.

    I'm betting on R1a all over Yamnaya to the west of Samara.

    ReplyDelete
  82. "Maybe R1, which we now know is native to Eastern Europe?"

    It was in Mesolithic east Europe, but it also could have been in parts of Asia.

    ReplyDelete
  83. Krefter,

    It's very difficult to argue now that R1 moved from the Near East to Europe, not only because we have European forages with R1, but also because ANE is not native to the Near East, and R is clearly an ANE paternal marker.

    Maybe R1 was in Westen Siberia and Central Asia too, but for the latter I'm rather betting on R2 and ancestral lineages.

    ReplyDelete
  84. It's pretty incredible all the R1b-Z1203(1 was not tested, 1 was negative) in Yamna. None of the Yamna males were related, meaning they all probably had pretty distant paternal lineages(we're not looking at a family).

    Such high R1b can only be compared to modern west Europeans, especially British-Irish and Iberians. High frequency of R1b was passed down from L23, to L150, to L11, to P312.

    ReplyDelete
  85. It's natural that Caucasian and Volga-Ural people will be drawn toward each other on the PCA - they are geographically adjacent. Regardless of which direction the genes were flowing (probably both).

    We know that R1 was found in Meoslithic Eastern Europe. We do not know where else it was at that time, because we don't have any aDNA (except to the west). The same is true of ANE. Going by present distributions, R1 was probably found over a large area already (V88 surely didn't come from the Caspian steppe after the Neolithic). Certainly this is true of ANE, which we know was already in the Americas.

    I'm not saying that the steppes weren't the main source of Indo-European R1b - that seems likely. But the evidence at hand is not solid yet.

    ReplyDelete
  86. The most interesting thing for me to see is R1a1 and R1b1 in Mesolithic Russia. Today R1a and R1b seem so different and separated, and we've finally traced to a time we they lived together.

    ReplyDelete
  87. In the PCA on Figure S5.2:

    Bell Beaker and Unetice are clustering with Northwest Europeans and Corded ware is just southeast of Balts. Also, Yamna is way east of Balts, and about as south as CWC.

    Just as we predicted!!

    ReplyDelete
  88. Sorry, for the many posts.

    Neolithic HGs from Sweden in that PCA are farther south than other HGs. Suggesting that had some EEF ancestry, like what their mtDNA suggests.

    ReplyDelete
  89. @Chad and others
    ''Give it up Nirjhar.. We have R1b as EHG.. No near eastern ancestry..''
    hahahahaha.
    Dude Urals already got intrusions from Zagros-Zarzian area on and before 6000 BC which Intensified with time there should be NO doubt that it brought R1b and also Some R1a!.
    About autosome yes it look EHG.

    ReplyDelete
  90. capra,

    Samples on a PCA aren't drawn together if there's gene flow in one direction only.

    In this case there had to be gene flow both ways. However, the R1b forager from the Samara Valley lacks Near Eastern admixture. So obviously R1b was on the Ural steppe before any Near Eastern farmers set foot there.

    Mike,

    Myself and others have already explained it. But I'll put up a post on it tomorrow, because those R1b-Z1203 Yamnaya nomads fit so well with the way the West Asians in my fateful triangle are behaving.

    ReplyDelete
  91. Krefter,

    No it suggests that the SHG had ANE, which we know they did. This is causing them to shift southeast. Same thing for the Yamnaya nomads.

    ReplyDelete
  92. It's a big jump south, not just east. mtDNA K1a1, H, T2b, and HV6-17 in PWC suggests some of them had EEF.

    ReplyDelete
  93. @David
    ''It's very difficult to argue now that R1 moved from the Near East to Europe, not only because we have European forages with R1, but also because ANE is not native to the Near East, and R is clearly an ANE paternal marker.''
    Pardon me as i'm not the expert here but Isn't the Ostrov Karella folk aka I0061 (Karelia_HG) is of R1a1? On The Other Hand Iran AND Turkey has older and basal R1a-420! with R1a1* Also?
    ''Maybe R1 was in Westen Siberia and Central Asia too, but for the latter I'm rather betting on R2 and ancestral lineages.''
    Sounds vague and wishful.

    ReplyDelete
  94. Nirjhar,

    Correction: Modern Turks and Iranians carry R1a-M420*. But of course they have steppe ancestry, so what's your point?

    The Karelian R1a is indeed R1a1, which means it's close to Indo-European R1a. Let's call it pre-proto-Indo-European. :)

    ReplyDelete
  95. Hi David,
    ''Correction: Modern Turks and Iranians carry R1a-M420*. But of course they have steppe ancestry, so what's your point?''
    My Point is Why Those basal clades are lacking? and another Where is R1a1a1b2? about component yes they have a bit.
    ''The Karelian R1a is indeed R1a1, which means it's close to Indo-European R1a. Let's call it pre-proto-Indo-European. :)''
    I have said earlier in your blog posts that E European area was already getting intrusions from West Asia from 80000+ YBP times.....

    ReplyDelete
  96. @Guys
    BTW A gross idea which emerges now is that Yamnaya was a R1b dominant area and it is interesting that Central Asian Sites like Andronovo, Mummies of Tarim Completely Lack the Y-DNA, any suggestions?.

    ReplyDelete
  97. Corded ware came out R1a1a1, and was mostly Yamna-like. Andronovo, the Tarim mummies, etc., were certainly mostly Yamna.

    ReplyDelete
  98. @Chad

    "Marnie, R1b is pure EHG in Samara. Think about it... It's like some jackass saying that they pick i2 as Neolithic even though we have pure WHG with it... You can't put the cart before the horse. R1b is not from west Asia..and it is connected to ANE....logic evades some"

    I am not saying that R1b is from West Asia.

    (Personally, I think that the R1b/R1a split happened either in the Balkans or Baltic, and the R1/R2 split happened in some place like Kazakhstan, but I don't state that, because I know there is not enough data to definitively make that statement.)

    You can't make a statement like R1b is from "Samara". It just happens that you have some samples from Samara that have R1b in them.

    And these particular Samara samples appear to be more related to Armenians than, say, Austrians. AND, Samara is almost directly north of Armenia, exactly where you would expect them to be if Samara were their Summer hunting ground and Armenia were their winter hunting ground.

    But these Samara samples are probably not the ancestors of Western European R1b, not even the ancestors of the Bulgarians.

    So again, you can't assume that descendants of MA1 only made it to Samara 6,000 years ago, end of story.

    You keep talking like you've got it all nailed down. I know you are on the message boards telling people who are not scientifically trained that you've got it all nailed down.

    But you don't.

    IN 20,000 YEARS, R DESCENDANTS WERE **NOT** CONFINED ONLY BETWEEN LAKE BAIKAL AND SAMARA JUST BECAUSE THEY DIDN'T HAVE F'ING CHARIOTS.

    END OF STORY.

    (until there are more samples)

    ReplyDelete
  99. Nirjhar,

    The early Indo-Europeans were highly patriarchal and patrilineal, and the Yamnaya horizon was made up of six different archeological parts. Think about that for a while.

    Nevertheless, and really concentrate on this, the data from this study point very clearly to the bifurcation in R1 lineages on the Eastern European steppe and their dominance in the early Indo-Europeans.

    Mmmmk?

    ReplyDelete
  100. @david

    Colin,

    I already told you why this was extremely unlikely.

    I'm betting on R1a all over Yamnaya to the west of Samara.


    Good luck with that theory. Not long ago you were so sure that yamnaya must have lacked r1b and could have only been r1a dominant. You thought the samara samples would all come back r1a. Instead they come back all r1b. How could you possibly think the non samara yamnaya will be r1a dominant? The evidence we got today contradicts the most obvious prediction of your theory, which is that the eastern yamnaya HAD to be r1a and not r1b.

    I don't think r1b could have tread through a bunch of r1a yamnaya territory to eventually dominate western europe. Also, we now have proof that r1a replaced r1b on much of the steppe, which you though was not possible.

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  101. Davidski,

    R1b1* is in Neolithic Spain and a likely R is in Neolithic Germany. So, I wouldn't call it an east European lineage just yet. It could have been in many other places, as far back as 7,000YBP.

    It is pretty obvious though that almost all R1b and R1a in Europe is Mesolithic East European derived., because Autosomally that was the new form of ancestry brought with modern R1b and R1a clades.

    As of far me and Chad are the only ones here who may very well have Yamna paternal lineages.

    ReplyDelete
  102. @davidski
    "However, the presence of basal R1a and R1b in Eastern European foragers suggests that R1 is native to Europe, and probably expanded from there via different subclades, mainly during the Copper Age."

    Was R1 some kind of Repunzel ? The more basal and greater the time depth (7000 BP) the harder it is to localize. For e.g. mal'ta and karelia are not next door neighbours.

    Also are you suggesting that the armenian like component in yamnaya only altered adna but left ydna untouched ?

    I recall you drew the same repunzel like scenario for R* in siberia which is even older.

    ReplyDelete
  103. I'm pretty sure we will find that r1a settled to the northwest of the steppe were it then gained IE and spread the IE related r1a to much of the IE world.

    ReplyDelete
  104. @krefter
    As of far me and Chad are the only ones here who may very well have Yamna paternal lineages.

    huh?

    ReplyDelete
  105. For nearlly 10 years now people have been debating R1bs connection with IEs and Yamna. Then every single Yamna male comes out R1b. It's unbelievable.

    ReplyDelete
  106. postneo,

    I don't know why European male foragers took Neolithic farmer wives and moved in with them? But it happened.

    Where's the Neolithic farmer Y-DNA you ask? The G2 and E? It's still there in little bits in the north of Europe, but better represented in the south.

    Krefter,

    The Iberian R1b might be from a wandering Eastern Euro hunter-gatherer. Maybe he was chasing a mammoth and ended up all the way in Iberia?

    Colin,

    Those mostly Z1203+ and M412- Yamnaya nomads aren't your ancestors. They probably have more relatives in Poland than Ireland.

    But don't worry, the Yamnaya horizon was huge...big enough for both of our tribes.

    ReplyDelete
  107. @Krefter
    ''Corded ware came out R1a1a1, and was mostly Yamna-like. Andronovo, the Tarim mummies, etc., were certainly mostly Yamna.''
    That is baseless R1a1a1 is found we know what that to do with R1b? Is There R1b in Andronovo?in Tarim? and Where is Z-93 in European aDNA?

    ReplyDelete
  108. @krefter

    Corded ware came out R1a1a1, and was mostly Yamna-like. Andronovo, the Tarim mummies, etc., were certainly mostly Yamna.

    Yamna related, necessarily yamna descended (probably not). Andronovo seems to lack r1b from samara which probably means it wasn't mostly yamna. I think there was some major spillage of r1a into the steppe with CW, CT, or both that eventually replaced much of the r1b.

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  109. Colin,

    Tying R1a to CT is really dumb.

    ReplyDelete
  110. The R1a1a1-M417* from Corded ware is very surprising. But anyways this makes Corded ware 3/3 R1a.

    ReplyDelete
  111. Davidski,

    You might be right, that would be my guess. Some of the EEFs look like they might have some ANE. Hopefully when the paper is published you can resolve that mystery.

    But don't forget the R from Neolithic Germany. It was positive for P1 and R.

    ReplyDelete
  112. @Nirjhar007

    Listen to Davidski. I would have never believed it.
    It's all starting to fit. Komi R1b, Rare Nepal/Bhutan R1b Rare Northern z2103 Pakistan/Punjab R1b, Mongol/Han Chinese R1b z2103 and ancient Karelian-R1a within same region it's just to way to much evidence.

    ReplyDelete
  113. @David
    ''The early Indo-Europeans were highly patriarchal and patrilineal, and the Yamnaya horizon was made up of six different archeological parts. Think about that for a while.''
    Practically NO R1a in Yamnaya aDNA thats just what happened for now! CWC yes but thats a 3000 BC Phenomenon after the Maykop Intrusion starting from ~4000 BC
    And West Asians were already entering E Europe from Neolithic that is UNDENIABLE.
    ''Nevertheless, and really concentrate on this, the data from this study point very clearly to the bifurcation in R1 lineages on the Eastern European steppe and their dominance in the early Indo-Europeans.''
    Not Without allowance from Asian aDNA leaving aside all other disputing data anyway this data is good ONLY for European perspective nothing else.
    The Data actually hints the PIE in West Asia you guys will get to that soon at least in your Subconscious levels:).

    ReplyDelete
  114. @david

    Those mostly Z1203+ and M412- Yamnaya nomads aren't your ancestors. They probably have more relatives in Poland than Ireland.

    Actually I had said a few times that the r1b in east europe was probably related what brought Indo European to the area...

    As for the samara valley people not being the direct ancestors of the irish that only pushes the closely related r1b of irish to more westerly yamnaya which leave less room for r1a.

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  115. this yamna males are not the ancestors of western europeans and it looks like this lineage moved from the caucasus or near east to the steppe. Again most western european r1b is not derived from this yamna males and r1b-2103 is rather typical for the near east and caucasus

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  116. @a
    this lineages there came more likely from west asia than directly from the steppe

    ReplyDelete
  117. Colin,

    How many square miles do you think it leaves for R1a? Enough for a few cows and sheep to graze in and a wagon to sleep in? Gee, I sure hope so.

    ReplyDelete
  118. @A
    ''It's all starting to fit. Komi R1b, Rare Nepal/Bhutan R1b Rare Northern z2103 Pakistan/Punjab R1b, Mongol/Han Chinese R1b z2103 and ancient Karelian-R1a within same region it's just to way to much evidence.''
    Not Enough trust me and interpretations can be multiple anyway Mongol-R1b is interesting can you link?.
    My Thought is that Central Asia-SC Asia Was R1a dominant from Archaic times Where West Asia was the Factory:) for both R1a and R1b,R2a etc.
    I'M pretty sure that R1a-Z93 will not be available in European aDNA which is Asian.

    ReplyDelete
  119. On Karelia

    "Thus, it can be designated as
    belonging to haplogroup R1a1*(xR1a1a) and it occupied a basal position to the vast majority of
    modern Eurasian R1a-related Y-chromosomes4
    , although more basal (R1a-M420*) Y-chromosomes
    have been detected in Iran and eastern Turkey4"

    I guess these are mordern samples but should this be discounted?

    R1b dominates yamnaya migration( with a small number of basal clades of R1a seen in england, scandinavia) and then east european R1a comes in at the tail end. they are probably linguistically and temporally separated.

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  120. @ this yamna males belonged to subclades which are very rare in western europe and r1b in western europe is clearly not derived from them so this males left almost no genetic traces in western europe. R1a was surely present in yamna and we even dont know if this yamna males were really indo-european or spoke another language, which was maybe orginally from the caucasus or west asia.

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  121. postneo,

    We don't know how and when those basal R1a clades got to Iran and Turkey. They're far removed from Indo-European R1a.

    The Karelian forager, who's obviously a European with less Near Eastern ancestry than any modern European in fact, has an R1a lineage that is much closer to the vast majority of European and Asian R1a.

    This puts you in a very difficult position. I suspect that at some level you know that.

    ReplyDelete
  122. Postneo, they say this in the paper:

    "The ultimate question of the Proto-Indo-European homeland is unresolved by our data.
    One important future direction for genetic research
    into Indo-European origins is to obtain ancient
    DNA data from India, Iran, northwestern China, and intervening regions to test hypotheses about the
    spread of Indo-European languages to the east. "

    They also discuss their data's implications regarding the Steppe, Anatolian, Balkan and Armenian Plateau theories of I-E origins, and say it decreases the plausibility of Anatolian and Balkan theories and may support both Steppe and Armenian theories (no mention of Iranian Plateau for the record). My guess is that the biggest deciding factor will be mesolithic or older Y-DNA from East Anatolia, S-C Asia and Iran.

    ReplyDelete

  123. @David
    ''The Karelian forager, who's obviously a European with less Near Eastern ancestry than any modern European in fact, has an R1a lineage that is much closer to the vast majority of European and Asian R1a.''
    R1a1* is also In Iran but The Karelian forager is also after the Jarmo Related intrusions from South Of Caspian if he was of 7500 BC then i would have no problems anyway the the Maykop Intrusion Likely brought the I-E folks to Europe which had roots in ANE Rich Central-SC Asia.

    ReplyDelete
  124. @Shai
    ''My guess is that the biggest deciding factor will be mesolithic or older Y-DNA from East Anatolia, S-C Asia and Iran.''
    Bingo! but still Asian origin way more heavy.

    ReplyDelete
  125. Davidski,

    I0443 could have been L51 with more digging so he he could be a direct ancestor of the Western Europeans.

    ReplyDelete
  126. "We don't know how and when those basal R1a clades got to Iran and Turkey. They're far removed from Indo-European R1a."

    Its closer to R1* and points to the split btw R1a and R1b near Iran. The steppe R1a and R1b are derived and ancestral to only Europe. Its still possible that the steppe R1a is ancestral to south asian but I doubt whether such an old lineage can be localized based on a sample.

    ReplyDelete
  127. Tesmos wrote,

    "I0443 could have been L51 with more digging so he he could be a direct ancestor of the Western Europeans."

    No. Every one of the seven Yamnaya individuals has been tested and found to be negative for L51:

    "I0439 (Yamnaya)
    This individual could be assigned to haplogroup R1b1a (P297:18656508G→C), with upstream
    haplogroup R1 (M173:15026424A→C, M306:22750583C→A) also supported. It was ancestral for
    haplogroup R1b1a2a1 (L51:8502236G→A) and so could be designated R1b1a*(xR1b1a2a1).

    I0443 (Yamnaya)
    This individual could only be assigned to haplogroup R1b1a2a (L49.1:2842212T→A,
    L23:6753511G→A). It could also be assigned to the upstream haplogroups R1b1a2
    (PF6399:2668456C→T, L150.1:10008791C→T, L1353:19179540G→A, PF6509:22190371A→G,
    M269:22739367T→C, CTS12478:28590278G→A). The individual was ancestral for haplogroup
    R1b1a2a1 (L51/M412:8502236G→A) and, unlike I0231, I0370 and I0438 also for R1b1a2a2
    (Z2105:15747432C→A). Thus, it could be designated as R1b1a2a*(xR1b1a2a1, R1b1a2a2)."

    ReplyDelete
  128. First of all, the monumentality of the effort and expertise to this paper requires nothing but commendation, respect and gratitude to its authors.

    ReplyDelete
  129. @MT
    ''First of all, the monumentality of the effort and expertise to this paper requires nothing but commendation, respect and gratitude to its authors. ''
    Yes.
    So what is your core review?.

    ReplyDelete
  130. WEll argued paper. However, Im not sure their estimated admixture rate necessarily reflect proportions of individual elements.


    Moreover, "A survey of other European steppe groups may reveal the more immediate patrilineal kin of the major founding lineages of modern European R1a and R1b chromosomes."

    So, there was a shift toward Yamnaya in central Europe, but this was in itself part of much larger processes (such as shift back to HG in the west, arrival of Caucasus, south Asian in the east). The evolution of the autosomal profile of Europeans we are detecting, therefore cannot be reduced to any single event. Certainly, at least from the immediate Y-DNA, the tested sample are nevertheless *not* the direct ancestors of modern Europeans.

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  131. L51 was probsbly already in those Carpathian Kurgans.

    ReplyDelete
  132. @ David

    "It's very difficult to argue now that R1 moved from the Near East to Europe, not only because we have European forages with R1, but also because ANE is not native to the Near East, and R is clearly an ANE paternal marker.

    Maybe R1 was in Westen Siberia and Central Asia too, but for the latter I'm rather betting on R2 and ancestral lineagese? "

    Maybe, maybe not . We'll see.

    "Nevertheless, and really concentrate on this, the data from this study point very clearly to the bifurcation in R1 lineages on the Eastern European steppe and their dominance in the early Indo-Europeans."

    Again, possibly. But why solely in eastenr European steppe and not also the central Asian one ? In fact, Id bet on a rather ever existing gradient and intermixture rather than sharp dividing lines.


    ReplyDelete
  133. @ Colin

    " think there was some major spillage of r1a into the steppe with CW, CT, or both that eventually replaced much of the r1b. "

    Perhaps the north, also.

    So then, and anti-Kurgan migration ?

    This study is great.

    ReplyDelete
  134. But Mike, why Central Asia, when we now have everything we need in Eastern Europe?

    - Karelian forager belonging to R1a M417-

    - Corded Ware herder belonging to R1a M417+ Z282-

    - Late Bronze Age German belonging to R1a M417+ Z282+

    See that, in a single study with just a handful of R1a samples they managed to catch the entire European story of R1a.

    No need for a migration from anywhere since that Karelian was alive. And he was clearly native to Eastern Europe anyway.

    ReplyDelete
  135. And there's no anti-Kurgan migration. Colin's just talking out of his ass.

    The Corded Ware samples don't look CT. They're obviously from the steppe, and have the same R1a as the Andronovo Kurgans and Tarim Basin mummies.

    ReplyDelete
  136. Yes, David. Im not necessarily saying there was a migration INTO Europe. But Im also stating that this genetic data plays havoc rather than proving the Kurgan hypothesis:

    We have Yamnaya being distinct branches of R1b, which share connections with West Asia / Caucasus rather than Europe.

    The direction of flow, if indeed only one, remains to be demonstrated, but it is suggestive that if there was new intrusion, it came from south of the Caspo-Pontic region, not vice-versa.

    We have R1a being spread in vast regions of Bronze Age Eurasia - from NE Europe to central Asia; as if arching around the western steppe.

    Between the Bronze Age, then, and the modern period, the steppe appears to have shifted rathern to a east-central European look c.f. the old western steppe type.

    Do you see my point ? The arrows don't add up to the movements suggested by the Yamnaya-steppe scenario. .

    ReplyDelete
  137. @David
    ''The Corded Ware samples don't look CT. They're obviously from the steppe, and have the same R1a as the Andronovo Kurgans and Tarim Basin mummies.''
    What R1a SNPs did Tarim,Andronovo have? You Know?

    ReplyDelete
  138. Krefter- thanks for compiling that useful Y - DNA summary

    ReplyDelete
  139. L51s brother and father are yamnaya. Bell beaker is 50% yamnaya. L51 Cotafeni looks pretty certain, as I said. Not rocket science!

    ReplyDelete
  140. Chad, Im not denying the gene flow scenario you are suggesting, in fact, I wasn;t directly making any statement on that. But I am sugesting that this complex scenario of gene flows are not the Kurgan hypothesis.

    ReplyDelete
  141. @ Chad Rohlfsen

    L51s brother and father are yamnaya. Bell beaker is 50% yamnaya. L51 Cotafeni looks pretty certain, as I said. Not rocket science!

    Do you have any proof that L51 orginated there? the yamna males were not ancestral to L51

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  142. Mike,

    Why are you ignoring the fact that the Eastern European foragers belonged to R1a and R1b, you know, both clades of R1?

    What makes them Near Eastern in your book? They don't look Near Eastern to me.

    The fact that the Yamnaya nomads were partly Near Eastern might be due to the foragers taking Neolithic farmer wives, like the I2 foragers did in Western and Central Europe. That never crossed your mind?

    Nirjhar,

    The Corded Ware, late Bronze Age German, Andronovo and Tarmin Basin R1a were all R1a-M417. You know, the main European and Indian clade.

    Oh, but the Karelian forager belonged to R1a M417-. Nice, huh?

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  143. This comment has been removed by the author.

    ReplyDelete
  144. Dave

    "Why are you ignoring the fact that the Eastern European foragers belonged to R1a and R1b, you know, both clades of R1?

    What makes them Near Eastern in your book? They don't look Near Eastern to me."

    Yes, Im aware that both R1a and R1b are found in Mesolithic E. Europe.

    Im not stating they're near eastern. There is, moreover a difference between the Near East - Levant, and the south Caucasian - Iranian highlands.

    "The fact that the Yamnaya nomads were Near Eastern might be due to the foragers taking Neolithic farmer wives, like the I2 foragers did in Western and Central Europe. That never crossed your mind?"

    No it didn't, because its a weak explanation, especially in the absence of explicit data to support such as definitive scenario.

    ReplyDelete
  145. @David
    ''The Corded Ware, late Bronze Age German, Andronovo and Tarmin Basin R1a were all R1a-M417. You know, the main European and Indian clade.

    Oh, but the Karelian forager belonged to R1a M417-. Nice, huh?''
    Yes very nice very Nice Since the CWC one predates the Andronovo,Tarim by ~1000 Years and Karelian one predates them by 4000 years! also what is important that CWC migrated towards deep into Europe not Asia!!!.
    WOW.

    ReplyDelete
  146. @Nirhjar007
    http://dienekes.blogspot.ca/2011/05/on-tocharian-origins.html


    A recent paper by Zhong et al. provides rich data on Uyghurs

    The region carries rare mtdna X2e also from Georgia X2e
    Georgia, Kyrgyz, Altai X2 X2 15310C
    https://www.familytreedna.com/public/x

    only 2 branches of R1b2103 moved south L277 and L584[Caucasus, Anatolia North Iran,Northrn India CTS 7822>9219 stayed within R1a 280 center core regions and spread to Albanians and Balkans region
    Somewhere along the way 277 and 584 are heavily mixed with 93's

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  149. Has anyone looked more closely where the R1bs presented in this study from Spain (EN) and Germany (MN, LN, EBA) sit with modern European samples, as well as the ancient ones from Samara ?

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  150. Maju Says'' the only R1b within the Euro-clade is from Spain.''
    Any remarks?

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  152. So ANE came to Europe via "farmer wives" then.

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  153. @Davidski

    Late Bronze Age person from Halberstadt is R1a-Z280

    and mesolithic Karelian is R1a-M459. We have such people in our R1a project :)

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  154. Wonderful!

    Gonna read the paper quite carefully. In any case, the tree models are gorgeous. The positions of Ust Ishim and Kostenki that we have reached on this blog so many months before have been confirmed! Congrats to all who put their heads together, and to you too David!

    Its also great that the're adding branches to ANE-WHG, making it more like the bush it was in real life. And also the new f4's. I expect such methodological progress to really pay off in the future.

    First comments.

    This throws YDNA into the air. R1b in yamnaya, but not Euro R1b? And found today in pops with almost no WHG too, many non-IE.
    And something extremely glaring. The R1b in Spain is very divergent, occurs in I0410 who is from a neolithic context, and I0410 does not differ from the rest of the neols at all in ADMIXTURE--they're really homogenous, suggesting the presence of this Y-marker in his pop at least >several hundred years there. Considering the fact that no basal R1b exists in Europe really, the closest archaeo R1b is so far away, I'm beginning to think that R1b in C+W+S Asia might be quite old? Such that ENF pops can have it with almost no ANE.

    But the autosomal picture confirms what we already know.

    Their autosomal affinities are estimated wrt modern components as West Asian and E.Euro, in approx equal parts, and their ENF ancestry is not from Europe.

    Why did they project their PCA samples? Pure ANE turns up in their PCA closer to E. Euros than French are to E Euros. That pca is useless. Facepalm.

    David, you need to redo some of their work lol.

    ANE levels turn out to be approx 27-28%, pulling off their f4 method and the tree, which is the low end of my estimated range and they're further east like I said, Hooray!, but we need to see how it turns out in David's analysis.

    In any case, I would caution agst paying too much attention to the YDNA results. The autosomal more or less shows that these ppl, or something very, very like them, influenced Europe to a great degree. RMB that these ppl are about as east and north in the Yamnaya horizon as you could go, so I highly doubt ppl from other places will have more of the Karelian side inst of the Armenian one.

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  155. About that sample from Corded Ware = it was also negative for L664, Z645, Z647. But CTS4385 and Z283 were not tested.

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  156. @Artur
    What about R1a-Z93 was the CWC Sample tested for that?

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  157. Impressive, that's what I call a seminal paper!
    I think it's all pretty much resolved now: The PIE homeland was on the PC steppe, and the forest steppe to the north of it. I used to have doubts about the derivation of western European R1b from the steppe and an association of it with IE, but this paper really has conclusive, unambiguous evidence.

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  158. Ironically, there really was R1b in Spain too, and at a very early date, at that. But it was upstream of P297, i.e. a very basal type of R1b.

    Whereas the Yamnaya sample includes even one L23. And a lot of M412's brother clade, which was ancestral to the Near Eastern L277.

    And no, the R1b didn't come from the "pseudo-Armenians", since already the Samara EHG had it.

    So on the paternal side this Yamnaya sample looks purely EHG. The West Asian admixture is restricted to the maternal side. And this suggests something obvious to me: The West Asian impact was rather the result of the capturing of West Asian women than the result of dominant West Asian males. Which obviously suggests that IE didn't come from the West Asians.

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  159. RK, the PCA's in figure 2 and figure S5.1 are not projections.

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  160. My congratulations to the Eupedia site; somehow they had guessed it right all the time.

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  161. Also I've read today that according to David Anthony the Fatyanovo culture was really predating the earliest Corded Ware. I used to learn it the other way round. A quick check on Wikipedia informs me that Fatyanovo pottery combined Globular Amphora and Corded Ware traits, and that it may be derived from the Middle Dnepr group. So it looks like the Middle Dnepr groups is the source for the central European Corded Ware, and for its R1a.

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  162. Skilur,
    One male was ancestral to L51. A straight L23. Yamnaya had L51s dad and brother. That is just a small part of Yamnaya. If we had 30-40 samples across the whole western steppe it would be there.

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  163. @Simon
    ''And no, the R1b didn't come from the "pseudo-Armenians", since already the Samara EHG had it.

    So on the paternal side this Yamnaya sample looks purely EHG. The West Asian admixture is restricted to the maternal side. And this suggests something obvious to me: The West Asian impact was rather the result of the capturing of West Asian women than the result of dominant West Asian males. Which obviously suggests that IE didn't come from the West Asians.''
    No it is well within the West Asian Projection you just lack the knowledge.

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  164. A little deductive reasoning with Bell Beaker being 50% Yamnaya, Skilur.

    Mike,
    I wasn't referring to your comment.

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  165. More female mediated, just as European Neolithic up North.

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  166. R1b and Italo-Celtic seem to stem from the Yamnaya in the Carpathian Basin. It really looks like Marija Gimbutas suggested: Bell Beaker (at least the eastern group) is an amalgam of Vucedol and Yamnaya. Vucedol was related with Baden, i.e. CO1-like. Yamnaya people settled east of the Tisza. Then they seem to have mixed. I used to be confused by the terminology of the Hungarian archeologists who have the Bronze Age starting already around 2750 BC, although at that date tin bronze wasn't yet in use. Vucedol did have arsenic copper. And occasionally that appeared also in eastern Bell Beaker sites. So there's no anachronism.

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  167. @ Chad Rohlfsen
    They were all negative for L51 and this yamna males did not migrated to western europe in great numbers because their subclade is quite rare there. It is most frequent among west asian populations, who are in many cases even not indo-europeans. Maybe the bell beaker orginated in the steppe but the existence of r1b-z2103 in yamna samples of samara is not proving or disproving that- Rich estimates about yamna ancestry look not realistic for me and eastern europeans like russians, poles and ukrainians should share much more dna with PIEs than any western european or even scandinavian population because they live quite close to the Indo-European homeland. But if we believe Rich than western europeans have almost the same amount of "steppe" dna like eastern europeans what makes no sense. The patriarchal indo-europeans were in most areas ,where they migrated, just an elite of males who took often local women and diluted their blood more and more as far they moved away from the eastern european homeland

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  168. @Nirjhar007

    Projection, you mean the PCA? I don't deny that Yamnaya had a lot of West Asian ancestry, but it was from the maternal side. The males all had R1b-P297 derived yDNA, and the latter was present in an EHG from Samara. So paternally they were EHG descended. This means they displaced the West Asian males. And patriarchalic as they were, did they take language lessons with their West Asian women? Rather not...

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  169. @Nirhjar007

    Sample can't be Z93 because it's negative for 2 SNPs that are upstream from Z93.

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  170. If R1b of West Europe is from Samara, some of those men did take language lessons from Basque women. We don't know the pseudo-Armenian Y-DNA distribution yet.

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  171. @Simon
    No not PCA.:)
    @Artur
    Ok. But please give your Suggestion on the case of R1b.:)

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  172. Let's make this clear: This Yamnaya sample is predominantly a side branch of M412, which later invaded West Asia, but there is also one L23 and one P297. What is L51? It's derived from L23. Even if it didn't exist back then, it may have arisen somewhere on the road. And since P297 was already present in Samara EHG it means that this branch of R1b was deeply rooted in the PC steppe, there is no reason to regard it as West Asian.

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  173. Yes indeed, perhaps there was a time when Basque women couldn't resist IE males, but their sons perhaps identified more with their mother's culture.

    There is no way that west European and Basque R1b could be descended from the early basal R1b in Iberian EEF, when there already was L23 in the steppe. Or do you really believe L23 arose twice, independently?

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  174. And not just L23, at that. But all markers inbetween the basal R1b1 in Spanish EEFs and L23. That is, P297, M269 and L23. That would be absurd.

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  175. Lets make it clear data has it R1b originates in West Asia and Yamnaya also show West Asian Mtdna, wives? Please there is too much Material data to topple that vague reasoning.
    Even the 6th Millennium BC r1b of Russia falls within the migrations from West Asia time frame which are detected from Archaeology etc.....

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  176. The Paper Concludes-
    ''The occurrence of a basal form of R1b1 in western Europe logically raises the possibility that presentday
    western Europeans (who belong predominantly to haplogroup R1b1a2-M269) may trace their
    origin to early Neolithic farmers of western Europe. However, we think this is not likely given the existence of R1b1a2-M269 not only in western Europe but also in the Near East; such a distribution
    implies migrations of M269 males from western Europe to the Near East which do not seem
    archaeologically plausible. We prefer the explanation that R-M269 originated in the eastern end of its
    distribution, given its first appearance in the Yamnaya males (below) and in the Near East''

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  177. If this study has taught us anything it's to abandon any idea of trying to use modern populations as references for prehistoric migrations. I myself got way too caught up in the R1a mania.

    IMO, it makes the steppe theory a lot more confusing, but still shows there was a massive demic impact from the steppe. IMO The other PIE theories are indeed alive, especially the Caucasian one. The actual PIE question, as in earliest Indo-European problem is a linguistic problem.

    Anyway, I still think we need more Yamna genomes and paternal markers, we only got 8? Why don't we have 100? I'm shocked at the lack of R1a and huge presence of R1b, but who knows.

    At least we can now say with authority the Steppe model has huge genetic diffusion power though, because before this it was looking weak from an archeological standpoint and years ago I had my doubts, not anymore.

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  178. Maybe R1b* originated in West Asia, or rather central Asia, it's quite irrelevant. European R1b is all in P297, which was so far only found in EHG. The mtDNA doesn't always parallel the yDNA, that's exactly the reason why mtDNA alone isn't enough.

    And you call it vague reasoning? We're dealing with actual ancient DNA, there's hardly anything more concrete.

    Yes, the EHG may have West Asian admixture too, but the ADMIXTURE analysis suggests that this is at most at trace levels.

    Also, as I said, it's impossible that the same chain of at least three mutations (P297, M269 and L23) occured independently in Western Europe and on the steppe.

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  179. I also found interesting that there's the rather eastern/ eastern-central European R1a-Z280 in the late Bronze Age near the Harz. To my knowledge the local culture was related with the contemporaneous Lusatian culture, so that makes sense. I suspect the sister branch (or rather brother branch) Z284 was present in the Nordic late Bronze Age.

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  180. another is-
    ''It is not possible to determine whether the appearance of R-Z2103 in the Yamnaya individual is due to
    (i) gene flow from the south to the steppe and related to the autosomal signal of “dilution” of Eastern
    European hunter-gatherers, or (ii) gene flow from the steppe to the south. Modern Armenians have a
    signal of admixture from the Yamnaya, as when we test...so the signal of admixture from the Yamnaya IS NOT the strongest one for Armenians. Moreover, as shown in SI 7, the Yamnaya have a negative f3-
    statistic with (X, Y) = (Karelia_HG, Armenian). A negative statistic for both Armenians and Yamnaya
    with each other as a reference population may suggest that a third (unsampled) population admixed
    into both the Yamnaya and to Armenians.''

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  181. This is also-
    ''Thus, it appears that before ~4,500 years ago, the frequency of R1a and R1b in Europe outside Russia
    was very low, and it rose in the Late Neolithic/Bronze Age period. The young, star-like phylogenies
    of these two haplogroups24 also suggest relatively recent expansions. The UBIQUITY of these
    haplogroups in Russia, Siberia, and Central Asia suggest that their rise in Europe was likely to have
    been due to A MIGRATION FROM THE EAST, although more work is needed to trace these migrations and
    also to correlate them with regions of the world that have not yet been studied with ancient DNA
    (such as southern Europe, the Caucasus, the Near East, Iran, and Central and South Asia).
    Nonetheless, the Y-chromosome results suggest the same EAST-TO-WEST migration as our analysis of
    autosomal DNA.''

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  182. This studies never have haplogroups anymore and to me that is irritating.

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  183. @david

    And there's no anti-Kurgan migration. Colin's just talking out of his ass.

    The Corded Ware samples don't look CT. They're obviously from the steppe, and have the same R1a as the Andronovo Kurgans and Tarim Basin mummies.


    So non of your theories on r1a have changed despite the fact that you were entirely wrong on your strongest prediction. That is, eastern yamnaya seem to be r1b dominant as I had predicted.

    Don't focus on the CT comment, for your sake you can just think of Corded Ware as being entirely responsible. Why is the CW r1a "obviously from the steppe"? And why do you think Tarim mummies and androvono have r1a from the yamnaya when they completely lack r1b that was likely dominant in eastern yamnaya? Nothing really points towards on origin of IE r1a on the western steppe.

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  184. @tone

    So how did Eastern Europe end up mostly R1a today? Did R1B invade Europe first while R1A followed a little later, eradicating their "cousins" in their wake? Or did R1B move south and around R1A territory?

    Id say both. R1a went north of steppe and R1b. Then R1b migrated out of western steppe which is when r1a filled the western steppe.

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  185. @ Colling Welling

    Daviski was not really wrong about anything and the Tanais burials who were 3000 years old show that R1a was present in the western steppe. I am not sure if this burials belonged to scythians or a pre-scythian population. Just because R1b-22103 dominated this samples we can not assume that this was true for the whole Yamna culture. Actually this is very unlikely because the yamna culture spreaded over a huge area. I think that yamna was also rich in r1a or r1a males killed almost all r1b males there because r1b is just not really frequent there anymore but ancient populations were not the same like modern populations.

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  186. K16 admixture results of Karelia and Samara Hunter-Gather samples:

    In Fig. S6.2 we show a box-and-whiskers plot of the cross-validation error for K=2 to K=20. This
    seems to plateau as K increases. The lowest median CV error is attained for K=16.
    p.56

    Karelia_HG
    Caucasus-Gedrosia 7,07%
    Amerindian 7,61%
    Beringian 3,26%
    WHG 78,26%
    Amerindian 3,80%

    Samara_HG
    Caucasus-Gedrosia 12,50%
    Amerindian 7,61%
    Beringian 0,00%
    WHG 78,26%
    Amerindian 1,63%

    http://abload.de/img/ehg5ezl5.png

    Interesting that even HG samples score beyond noise level West Asian values.

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  187. Interesting, so the Caucasus Gedrosia seen throughout Europe now (or generally after the Late Neolithic) is potentially directly linked with EHG?

    Makes sense.

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  188. Could be a case of EHG-related admixture in the "Gedrosia"-like component (meaning EHG in Caucasus and S-C Asia), just like the case of MA-1 which appears to have admixture from "unadmixed" Native Americans while other methods have shown the opposite is true. ADMIXTURE's problem is that a homogenous sample collection will form its own component that makes them appear unadmixed, whether those samples are mixed or not.

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  189. First to agree with all of those who have praised the paper. The amount of data and analysis it contains it's huge. I was quite sceptic about this paper, but I'm very happy to admit I was wrong. It was worth the wait.

    For me the best thing is how the base model has changed. It is a HUGE step in the right direction compared to the previous paper (that was too simplistic and quite biased).

    OLD MODEL:

    Mesolithic Europeans: WHG.
    Neolithic Europeans: WHG + ENF = EEF.
    Bronze Age Europeans: EEF + EHG (WHG*ANE)


    NEW MODEL:

    WHG
    WHG + ENF = EEF
    EEF + WHG = MNE (mid neolithic Europeans)
    Here a mysterious population of unknown origin enters into Europe. They are Armenian like, ANE rich (ENF+ANE), and enter to the steppe, mixing with EHG.
    EHG + 3rd-pop = Yamnaya
    Bronze age Euros: MNE + Yamnaya

    This new, much more realistic model, fits really good for most Europeans. Especially for Eastern and Northern Europeans (though some WHG has to be added after the Bronze Age, and some East Asian to Russians and Finns).

    For South Europeans it still has to be stretched a bit (and we don't know for NW euros, since we don't have samples from there). A piece or two from the puzzle might still be missing, but we're getting much, much closer than before. Let's hope that next month's conference about Greek civilizations brings us one more piece.


    The only thing I'd criticize from the paper is it being too Eurocentric (and I'm European myself). It largely ignores Asia, and the whole event of the intrusion of the Caucasus-like population is somehow treated as a "minor" event and not as the catalyst of the whole migrations of the period. Still too much "steppe" centric (I know that until we don't know the origin of that population is hard to give it a name, but this whole "Yamnaya" is like referring to the first wave of migrations from the Near East as "Balkan culture" (just because they first landed in Europe in the Balkans and there they mixed with native European populations).

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  190. Such a great paper.

    On the Yamnaya...they were stated as 50% EHG and 50% near eastern/Armenian like'obviously based on these samples.

    Setting aside the question for now of where these Yamnaya R1b clades are most frequent today(maybe Armenia,Caucasus today, but before?) and the obvious(and confusing!) Spanish R1b from about 5000BC, if Yamnaya can be modelled as 50/50 EHG/near eastern or armenian like, looking at the mtDNA in the table S2.2 in this publication, it lists all the mtDNA of Yamnaya samples so far collected. Of the 36 samples, about 60-70% seem to be from the near east, and the rest 'Hunter-Gatherer'. In this particular study, there were 6 near eastern haplogroups out of 9.

    Would this roughly 2/3rds near eastern mtDNA be solely enough to explain all of the near eastern in Yamnaya?

    All the males were R1b. If R1b is also near eastern, that means 2/3rds of mt is near eastern as well as all the r1b, which would make well over the 50/50 predicted mix for yamnaya dna. More like 80% near eastern, the remainder Hunter Gatherer.

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  191. Glad it's finally out and everyone is reading. I haven't read the paper fully yet (let alone the comments here), after being busy today; looks much more complex than what has come before!

    Initial impression is it strikes me that for immediate interpretation and "punch" this paper is much more reliant for its immediate analysis on the ADMIXTURE (as I thought would be the case given their expanded samples), and PCA and relatively few and simple admixture statistics to make their case.

    The phylogenic tree modelling has clearly got *much* more complex and less conclusive for this paper, given how much more adna is around. There are a lot of supplements with a lot of methods to reconfirm their analysis and a lot of questions.

    Notes about the non-projected PCA (I assume non-projected as a projected PCA exists for contrast) as it's relatively simple to interpret -

    - As I thought it is clearly the case that a "Fateful Triangle" model, although very close, doesn't fit *exactly* to approximate a West Eurasian PCA, as like I noticed in David's earlier PCA with Oetzi, many of the early-middle Neolithic samples are out beyond the edge of the line running through WHG to Sardinian to BedouinB (as you can see on Fig 2a).

    - As Krefer observes, the differences between Motala and the Skoglund MHG / NHG samples clearly do not resolve through ANE admixture (because they're shifted south from Motala with no east shift from Motala). So unless its a genotyping effect, something else is going on between them.

    - My interpretation of the position of MA-1 on that PCA is that much of MA-1's genetic uniqueness is simply unrelated to European variation. Thus it appears closer than it is (it would be much more different if we took PC3, PC4, PC5 into account (as with Kostenki14 and Ust-Ishim)).

    - One thing that would be interesting is that in Lazaridis they ran a model where they ran a simple PCA of the ancient samples (Loschbour, Stuttgart and MA-1) then projected West Eurasians onto that. A repitition of this would benefit from their expanded sample size. I may comment on bioXriv.

    - The samples are all great. In a perfect world (where we get everything we want) I wish they'd had some Spanish Beaker people as these might have less local admixture from Corded Ware compared to these Germans (although which one truly represents early Beaker people is probably a question for folk like Bell Beaker Blogger). The German Beakers mostly seem to fit clinal to Middle Neolithic German plus Corded Ware cline , though - they have a wide dispersal and clearly some have excess farmer and SHG ancestry.

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  192. RK: Gonna read the paper quite carefully. In any case, the tree models are gorgeous. The positions of Ust Ishim and Kostenki that we have reached on this blog so many months before have been confirmed! Congrats to all who put their heads together, and to you too David!

    p80, I'm glad they've attempted to include K14 in their models, which are the meat of their paper, still introduction of another layer / form of Basal in K14 is dubious to me.

    Still.... Why no inclusion of the D(Mbuti,Ust-Ishim;[ancient Eurasian],K14) stats? Also why no inclusion of the D(Chimpanzee,Mbuti;[ancient / modern Eurasian],Ust-Ishim) stats? Those are crucial for placing U-I on their tree.

    I will have to comment on the paper's bioXriv later today to ask these questions.

    The below comments are all on the phylogenic section (p78). (Comments all essentially about the paleolithic time scale modelling, so if you're just interested in the Neolithic and post Neolithic action will be of marginal interest).

    p81. The hypothesis of Basal Eurasian ancestry in Kostenki14 needs to be further tested, as the
    negative D(Mbuti, Han; Loschbour, Kostenki14) statistic could also reflect gene flow
    between Han and Loschbour a priori plausible, as these populations are much younger
    than Kostenki14 and may share intra-Eurasian genetic drift that Kostenki14 lacks because of
    its age. The possibility of later gene flow between Europeans and eastern non-Africans must
    be further tested with additional ancient samples from Upper Paleolithic Europe and Asia.


    This is welcome (I think it's a better option than another basal group), still it would be better phrased if they had acknowledged the note from Laziridis 2013 their comment about the extreme unlikelihood of symmetrical gene flow into ANE and "West Eurasian" from ENA, which is pretty much what the whole introduction of Basal Eurasian into their model is very dependent on.

    p84, Karelia HG fails (marginally) as a mix of WHG and ANE. Also given p7 Modeling of the ancient samples shows that while Karelia is genetically intermediate betweenLoschbour and MA1, the topology that considers Karelia as a mixture of these two elements
    is not the only one that can fit the data (SI8).


    which leads on to

    Still - "All three of these 2-way mixture models arrive at a similar inference of 38-40% ANE and 60-62% WHG ancestry in Karelia-HG, which is higher than the ~20% ANE ancestry inferred for
    Motala12 in a previous study4, consistent with the fact that EHG are the population sharing
    more alleles with “Ancient North Eurasians” (SI 7) than any other."


    39:61 ANE:WHG at least gets us a marginal EHG model, despite the fact this marginally fails.

    p85 - MA1 as admixed between a Karelia HG like population and a ghost population. Interesting models, however if one way gene flow was dubious for other populations, it's very dubious for Loschbour (flow from proto-EHG into proto-WHG, no reverse flow? dubious). Plus, how many of these models could cope of with K14? We'll have to see how well they cope when we get a MA1 age Upper Paleolithic European from Central Europe.

    (btw, anyone else notice Laziridis is the author of the pdf document? ; ) ).

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  193. Regarding Y-DNA:

    - Big surprise to see all R1b in Yamnaya. Just a couple of months ago I was asking to those who claimed that R1b was from the steppe if they expected to see it in the Samara samples, and all were finding excuses as to why it might not be there, but still be from the steppe))

    - Big surprise to see R1b in Spain in the Neolithic. No matter if he's not the ancestor of WE, it's till BIG. And he clusters with other Neolithic Iberians, so no, he wasn't alone or an exception or a lost hunter from the steppe.

    - Big surprise to not see R1a in Yamnaya. Go figure why.

    Regarding if R1b in Yamnaya was native or brought by the Armenian-like population, nothing conclusive. Both options are possible. If there's anything that this paper has shown about R1b is that it was very widespread since very early. It was in the steppe, and in Spain. It could be in Central Asia too (not related to any Basal Eurasian population from the Near East, but to an ANE population more to the east, that later moved west).

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  194. "The Armenian plateau hypothesis gains in plausibility by the fact that we have discovered evidence of admixture in the ancestry of Yamnaya steppe pastoralists, including gene flow
    from a population of Near Eastern ancestry for which Armenians today appear to be a reasonable surrogate "

    Unless the ancestors of the Armenians moved to where they are now to get away from the horse dudes.

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  195. Perhaps the R1b in Spain is V88 and related a North African/Mediterranean Cardial Neolithic migration. L91 is also found in North Africa and Egypt.

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