January 28, 2014

La Braña 1 carried the very rare Y-DNA haplogroup C (possibly C6-V20)

La Braña 1 without makeup
(Check for the updates below, please).

The late Epipaleolithic forager from NW Iberia (previously discussed here) had the patrilineal haplogroup C6, found so far only very rarely among modern Europeans (Scozzari 2012). This, I must say, I know by the moment only from secondary sources (Eurogenes, Dienekes and a personal communication) because I have not been able yet to put my hands on the relevant paper and this key detail is not mentioned in the abstract.

Iñigo Olalde et al., Derived immune and ancestral pigmentation alleles in a 7,000-year-old Mesolithic European. Nature 2014. Pay per viewLINK [doi:10.1038/nature12960]

freely available supplementary materials.

Abstract

Ancient genomic sequences have started to reveal the origin and the demographic impact of farmers from the Neolithic period spreading into Europe1, 2, 3. The adoption of farming, stock breeding and sedentary societies during the Neolithic may have resulted in adaptive changes in genes associated with immunity and diet4. However, the limited data available from earlier hunter-gatherers preclude an understanding of the selective processes associated with this crucial transition to agriculture in recent human evolution. Here we sequence an approximately 7,000-year-old Mesolithic skeleton discovered at the La Braña-Arintero site in León, Spain, to retrieve a complete pre-agricultural European human genome. Analysis of this genome in the context of other ancient samples suggests the existence of a common ancient genomic signature across western and central Eurasia from the Upper Paleolithic to the Mesolithic. The La Braña individual carries ancestral alleles in several skin pigmentation genes, suggesting that the light skin of modern Europeans was not yet ubiquitous in Mesolithic times. Moreover, we provide evidence that a significant number of derived, putatively adaptive variants associated with pathogen resistance in modern Europeans were already present in this hunter-gatherer.


Relevance for the overall understanding of macro-haplogroup C

Until the discovery of this C6 lineage, there were some strong reasons to suspect that Y-DNA C may have coalesced already in SE Asia or, at least, very close to it, with its subclades forming by pairs a three pointed star with geographical center in that area: C1 and C3 in NE Asia (and America), C2 and C4 in Wallacea and Australasia and C5 and some rather homogeneous C* in India.

The discovery of this C6 lineage and its confirmation as a Paleolithic one in Europe (i.e. not a "recent" arrival from somewhere else) add phylogenetic weight to the Western geography of haplogroup C, one of two main subdivisions of the main non-African Y-DNA lineage CF. However we cannot yet reach to conclusions about the "exact" origins of C because the macro-lineage still awaits improvement of its phylogenetic structure at the basal levels.

In plain language: it is quite likely that C2 and C4 form a monophyletic clade and I would not be surprised at all if C1 and C3 do the same. But then it is also possible that C5 and the Indian C* and/or the European C6 also form their own distinct branches. It is even possible that some of these lineages are related across subcontinental regions, as was recently found within MNOPS (aka K(xLT)). So we need first to know how they relate with each other a the top phylogenetic level before we can rush to any conclusion. In any case the discovery of C6 adds some preliminary weight to the hypothesis of C coalescing when still in South Asia.


Pigmentation genetics

There have been some rush to conclusions on the pigmentation of this and another Western European hunter-gatherer based only on genetics. I think that some of the conclusions are most likely incorrect, at least to some extent, because they are based on a SNP which only weights ~15% on skin coloration.

Judging on the figures (freely accessible, it seems), La Braña 1 carried two pigmentation alleles of gene SLC45A2 now rare among Europeans (but common elsewhere, i.e. the ancestral variant):
  • rs16891982, which affects hair color (7x chances of black hair among Europeans)
  • rs1426654, which affects skin pigmentation to some degree (correlated with skin color in Indians, irrelevant among modern Europeans because of fixation, weights only ~15% in Cape Verdeans' skin coloration). 
Notice that while you can find online reconstructions that give La Braña 1 a very dark coloration, this is not necessarily the case at all but rather an oversimplistic  interpretation based only on one allele, allele that is not just dominant in West Asians and Europeans but also, for example, among Gujaratis, who are quite dark for European standards.
    It seems correct anyhow that this allele was only brought to Europe with Neolithic farmers (Stuttgart had it) but its alleged effect on pigmentation seems very much exaggerated.

    Fig. 4 from Beleza 2013 highlights that no single gene is decisive in skin pigmentation.

    It is probable anyhow that La Braña 1 had black hair.

    It is much more plausible that he had blue eyes because these are much more directly regulated by simple genetics.


    Continuity of immunity genetics

    La Braña 1 also had three immunity related alleles (derived variants) that have been retained at least to some extent by modern Europeans:
    • rs2745098 (PTX4)
    • rs11755393 (UHRF1BP1, related to lupus)
    • rs10421769 (GPATCH1)

    Comparison with global populations

    Fig. 5 (ED) offers various comparisons of La Braña 1 and Mal'ta 1 (from Siberia) with modern humans from around the World:

    Extended Data Figure 5: Pairwise outgroup f3 statistics.
    a, Sardinian versus Karitiana. b, Sardinian versus Han.
    c, La Braña 1 versus Mal’ta. d, Sardinian versus Mal’ta.
    e, La Braña 1 versus Karitiana. The solid line represents y = x.

    We can see in them that, La Braña 1 clusters well with modern Europeans, while Mal'ta instead strongly tends towards other Asians, often clustering with Pakistanis ("Central/South Asia" metapopulation).

    Maybe the most interesting graph is c, where we can see how the various populations deviate from the y=x line in the direction of La Braña (Europeans, West Asians) or Mal'ta (Native Americans particularly).


    Comparison with Neolithic samples and modern Europeans

    Extended Data Figure 4: Allele-sharing analysis.
    Each panel shows the allele-sharing of a particular Neolithic sample from refs 1 and 3 with La Braña 1 sample. The sample IDs are presented in the upper left of each panel (Ajv52, Ajv70, Ire8, Gok4 and Ötzi). In the upper right of each panel, the Pearson’s correlation coefficient is given with the associated P value.

    In all cases Swedes (SE), followed by Polish (PL), etc. share the greatest amount of alleles with La Braña 1, although I'm not sure if the differences are really that relevant (is really 69.3% significantly different from 68.7%?)

    In the vertical scale we can observe how the various populations tend more or less strongly towards various Neolithic samples (again with the same doubts about the significance of the differences). In the first row they are compared with Götland's Pitted Ware individuals (of plausible Eastern European origins: strong cultural connections with Dniepr-Don Neolithic). Here Central Europeans show the greatest affinity with Ajv52 and Ajv70 (Basques Bulgarians also score high). There are some differences in the case of individual Ire8, whose closest modern relatives seem to be the Dutch. Swedes only score high re. Ajv52 but low to the others, while Finns score neutral-to-low relative to all them.

    The lower row compares with to mainstream Neolithic samples: Gok4 was a Megalithic farmer from SW Sweden and Ötzi was a Chalcolithic shepherd from Southern Tirol. The Swedish farmer is best approached by the Dutch, followed by various West-Central Europeans, while Basques Bulgarians, Finns and Swedes score low here. In the case of Ötzi nobody scores particularly high (some tendency in Switzerland and nearby areas), while Finns score clearly low.

    And that's all I can say without direct access to the study. Enjoy.


    Update: I already got the paper (thanks again to the donor), I'll see to update as need be once I have time to read it. Minor urgent edits above in red (and slashed out text).


    Update (Jan 29): The supplementary data is freely available (LINK) but I could not find it earlier. Almost all the information is in it, including a long list, much longer than mentioned above, of the SNPs found in La Braña 1, compared to various modern population frequencies. I don't have time right now to dwell on it but I guess from a first read that I will have to amend some comments made on the issue of pigmentation above.

    Regarding the Y-DNA haplogroup, it is important to notice that its adscription withing haplogroup C seems very clear but its assignation to C6-V20 is more dubious because of the low quality of the genome. Only the V20 marker could be assigned, so the authors themselves are in doubt and wonder if it could alternatively be C* or C5, both with a South Asian affinity.

    In this sense I think it is worth noticing that the reference Y-DNA site ISOGG has recently revised the phylogeny of macro-haplogroup C and that they have already renamed C6-V20 as C1a2, making it a relative of the minor Japanese lineage earlier known as C1 (now renamed to C1a1), similarly South Asian C5-M356 has been renamed to C1b. So C1 is now perceived as a lineage that spans all Eurasia with an arguable South Asian centrality.

    Another (Papuan?) lineage once known as "C6" has long vanished from the phylogeny because of lack of plural samples, I understand.

    28 comments:

    1. The full paper is available online at Sribd if you put the title into Google. Someone uploaded it there, as they usually do.

      And yes the difference between 69.3% and 68.7% is significant, unless you don't think that the difference between, say, Italians and Poles is significant, in which case no.

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      1. I got it already, thanks. I'll read it tomorrow, I guess.

        "And yes the difference between 69.3% and 68.7% is significant, unless you don't think that the difference between, say, Italians and Poles is significant, in which case no".

        Well, the difference between Italians and Poles is tiny when compared with the big global picture, but guess we are splitting hairs, as they say, when trying to understand the rather subtle differences between Europeans, so fair enough.

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    2. "Notice that while you can find online reconstructions that give La Braña 1 a very dark coloration, this is not necessarily the case at all but rather an oversimplistic interpretation based only on one allele, allele that is not just dominant in West Asians and Europeans but also, for example, among Gujaratis, who are quite dark for European standards."

      Definitively, and i have stated as much, elsewhere, and I am just going to copy and paste:

      "Yes, of course - as I have mentioned in the past, this is a necessity since the Neolithic/ SE European/ West Asian variants don't really code for particularly light skin at all in the European context. And as you state, this causes a huge problem: if we don't know what those mutations/ locations are, we really have no clue what skin (or even hair) color Mesolithic Europeans had - but most likely it was not very dark or brown, at all."

      "We should also keep in mind that this single mutation [SLC24A5] does not make the skin particularly fair, on its own (it occurs at a significant fraction of people who have Mediterranean, dark Mediterranean, dark W Asian, or even dark S Asian/ African skin color). As I have mentioned before, one also needs to distinguish between static skin color and adaptable (over the seasons), and perhaps take into account subtle photochemistry effects on the molecular level that don't express themselves that simply and naively as "skin tone."

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    3. Maju wrote,

      "In plain language: it is quite likely that C2 and C4 form a monophyletic clade and I would not be surprised at all if C1 and C3 do the same. But then it is also possible that C5 and the Indian C* and/or the European C6 also form their own distinct branches. It is even possible that some of these lineages are related across subcontinental regions, as was recently found within MNOPS (aka K(xLT)). So we need first to know how they relate with each other a the top phylogenetic level before we can rush to any conclusion. In any case the discovery of C6 adds some preliminary weight to the hypothesis of C coalescing when still in South Asia."

      Former C1 (C-M8) and C3 (C-M217) definitely do not form a monophyletic clade relative to C-V20 (former C6) or C-M356 (former C5). In fact, the current ISOGG tree has C-M8 as C1a1-M8, C-V20 as C1a2-V20, and C-M356 as C1b-M356, all subclades of C1-F3393/Z1426. In other words, C-M8 and C-V20 form a monophyletic sister clade to C-M356, and all these three form a monophyletic clade relative to C-M217.

      The position of Australian C is unknown. There has been an unconfirmed report of an SNP linking C-M38 (former C2) with the group that subsumes C-M8, C-V20, and C-M356 as opposed to C-M217. Almost all known C* (most of which is from the Malay Archipelago, though some cases also have been found in India, Indochina, and China) seems to be close to the root of either C-M38 or Australian C, so it is possible (though not at all confirmed) that all these Y-chromosomes (Indian/SE Asian/Chinese C* + C2-M38 + Australian C) may form a "southern" clade opposed to "central"/"western" C1-F3393/Z1426 and "northern"/"eastern" C3-M217.

      Anyway, Indian examples of C*, which are very few and far between, seem to group closely with Southeast Asian C*, and they are certainly not particularly diverse in comparison to confirmed subclades, such as C3-M217 or C2-M38. I would say that the most likely region of origin of haplogroup C-M130 remains southeastern Asia (including e.g. China), with C3-M217 radiating mainly northeastward, C2-M38 and Australian C radiating southeastward, C* staying put (with some minor flow westward into India), and C1-F3393/Z1426 radiating probably northwestward to Central Asia (with one branch then moving southward into South Asia and another branch dispersing along an east-west axis, eventually yielding C-V20 in Mesolithic Europe and C-M8 in Mesolithic Japan).

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      1. My main point is that we do not know the exact phylogenetic position of each subclade and we must be cautious until the exact structure is discerned. However:

        1. Based on some STR old studies (no time right now to find them, sorry) C4 appeared as derived from C2 and that's why I suspect that they may be closely related.

        2. I have no particular genetic reason to think that C1 and C3 are closely related but their neighboring geography.

        3. Same for C5 and Tamil tribal C* (and maybe C6): only geography guides my speculation.

        I'll see if I can find the relevant study (from 2004?) and expand later on or tomorrow.

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    4. Maju a few things:

      The evidence that La Braña-1 belonged to haplogroup C6 is very thin, only due to one marker V20, and the authors think it could be due to "Possible post-mortem DNA damage". It's seem La Braña carried the ancestral state for V86, V182, V184, V222, so it is probably best to assume simply C*.

      Here is a link to the supplementary, look up Table-S9: http://www.nature.com/nature/journal/vaop/ncurrent/extref/nature12960-s1.pdf

      Also I don't think that BA from the PCA are Basques, instead I think BA stands for Bulgaria, I too thought those were Basques at first, but then noticed the following:

      "reference populations are the Finnish HapMap,FINHM and POPRES. Population labels with labelling of ref. 12 with theaddition of FI (Finns) or LFI (late-settlement Finns).
      [...]

      Continental terms make reference to each Omni population grouping as follows: Africans, Yoruba and Luyha; Asians, Chinese (Beijing, Denver, South, Dai), Japanese and Vietnamese; Europeans, Iberians, Tuscans, British, Finns andCEU; and IndianGujarati fromTexas.c ,Each panel shows PC1 and PC2 based on the PCA of one of the ancient samples with the merged POPRES. FINHM sample, before Procrustes transformation."


      So they are using the 1000 Genomes, which does include 7 Basques amongst the Iberian population, and the POBRES database which is where BA(Bulgaria) comes from. Notice their similarities with the YG(Yugoslavia) sample or RO(Romania).

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      1. Buff, a lot of misunderstandings, it seems. Why would BA be Bulgaria, why not BU or BG?

        I take note of what you say about C* and C6 and will try to update tomorrow. Thanks a lot, very informative.

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      2. In any case it makes better sense because I was quite perplex that Basques would show any relation towards Pitted Ware, really.

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    5. They did not test rs11636232 in HERC2, so I could not determine if his blue eyes are ht1 or ht2.
      http://1.bp.blogspot.com/-JKcfHEfIdRc/Ucl-P-4Sg2I/AAAAAAAAB_0/c-sm_XErh0I/s1600/eyecolorhtmap.png

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      1. Pity. The DNA is very degraded so maybe they could not detect it?

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    6. Another possibility imo, say for the sake of argument C developed wherever looking similar to modern South Indian HGs for example and then spread out east and west and when they hit more northern latitudes lost the selective pressure to stay dark or gained some selection pressure to get lighter what might be the quickest way that could come about? Some kind of partial depigmentation?

      http://media-cache-ec0.pinimg.com/236x/57/b2/70/57b2708d0d40295de096599768578969.jpg

      http://i.imgur.com/Ew5G7OL.jpg

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      1. In Europeans it seems that it is the TYR gene which correlates with freckles (and not with basic skin color), however in Indians it was reported as one of three genes correlated with skin color. It should not be what you seem to suggest in any case: that should rather be variants of TYR if anything.

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      2. http://en.wikipedia.org/wiki/IRF4

        "This gene is strongly associated with pigmentation: sensitivity of skin to sun exposure, freckles, blue eyes, and brown hair color."

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      3. IRF4 related to immune system so skin changes may have been secondary

        http://www.genome.gov/27555432

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      4. Proposed as such by a single recent PPV study. Seems tightly related to TYR anyhow: judging on the "graphical abstract", IRF4 would increase the gene expression of TYR.

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      5. Yes. La Brana had ASIP + TYRP1 + IRF4 and none of SLC24A5/SLC24A2/OCA2/KITLG

        If there is someone in Ireland or Scotland the same then we can see if they are are dark or very freckled.

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    7. Fig. 5 (ED) offers various comparisons of La Braña 1 and Mal'ta 1 (from Siberia) with modern humans from around the World

      Quick points about Fig 5 -

      Looking at the Central Asian samples in Fig 5 f3 charts a and b seem interesting to me.

      They all line up in terms of their affinity to Sardinians, the y axis on f3 charts A and B, in terms of having more than East Asians but less than Europeans.

      But 2 of the samples appear to have relatively elevated affinity to Karitiana and more so to Han Chinese, the x axis in chart a and b respectively. These may be Central Asians who have recent ancestry from East Asia, possibly Hazara and/or Uighur samples.

      However, these 2 samples aside, the other central Asian samples do not have a raised affinity to Han Chinese or Karitiana compared to Europeans.

      (see this link for a graphic explanation - http://i.imgur.com/Jsxi9hu.png)

      Mal'ta is similar in this respect, compared to La Brana. No real increased affinity to East Asians, but decreased affinity to Sardinians, plus the Mal'ta unique quality of a slight increase in sharing with Native Americans (who are nonetheless really distant from all other populations when an axis is ascertained against them, due to their high levels of drift). What Mal'ta really shares with Central Asians is a medium level of lacking affinity to Sardinians, but is otherwise not particularly like them.

      It would be interesting to see how Mal'ta fared in a f3 (Pathan, X: Yoruba) axis. It might sit with the Europeans (or not)...

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      1. All it says is that Mal'ta weights ~17 in Sardinian affinity and ~19 in Native American affinity. Previous studies have also underlined that MA1 is closer to Native Americans than to other Europeans, even those which have more "ANE" (i.e. MA1) affinity.

        Instead MA1 is still slightly closer to Sardinians than to Han (~16), probably underlining that it derives from the same West Eurasian common root, even if he might already have incorporated some East Asian genetics and even if that common West Eurasian root had only diverged from the general Eurasian pool not long before the Central Siberian ancient population diverged from the Western branch.

        I would not go crazy about all that: overreading autosomal genetics is a sure way to a new type of erudite quixotic madness, I fear. We need simpler and sturdier anchors like haploid genetics and archaeology.

        "It would be interesting to see how Mal'ta fared in a f3 (Pathan, X: Yoruba) axis. It might sit with the Europeans (or not)... "

        Not sure: judging on that graph, I'd say it'd tend to Pathans instead but he may also sit in the middle. I seriously doubt it would strongly cluster with Europeans: ancient Central Siberians ("ANE") were almost certainly not directly related to Europeans, not more (or at worse not much more) than to West Asians and such: they were a distinct third branch of the West Eurasian macro-population. More recent admixture has got them extinct (diluted to indistinctness) and has brought some of their genetics to some other populations such as NE Europeans and Northern West Asians but originally they were a distinct third branch with 99% certainty.

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    8. Not sure: judging on that graph, I'd say it'd tend to Pathans instead but he may also sit in the middle. I seriously doubt it would strongly cluster with Europeans

      My main point here in saying this is that,

      - if Europeans also sat in the middle on a f3 (Pathan, X: Yoruba) axis and so did Mal'ta

      - and the other axis involved in the comparison was say f3 (Han, X: Yoruba) axis, which we already know from this study Europeans are about the same as Mal'ta

      - then on a graph of those two axes Mal'ta would end up inevitably clustering with Europeans.

      But that wouldn't necessarily be that much of an indication of any overall similarity with Europeans.

      I would not go crazy about all that: overreading autosomal genetics is a sure way to a new type of erudite quixotic madness, I fear. We need simpler and sturdier anchors like haploid genetics and archaeology.

      Archaeology is important. As for haploid genetics I have to leave it to people for who that is a passion.

      Frankly any discussion I've ever read about haploid genetics more or less seems like incoherent nonsense to me. "This clade is here at this frequency here" "Oh yeah? Then how come this clade is here at this frequency then?", etc. There is no systematic, algorithmic methodology to make any sense of any of it, and interlocutors seem able to try and pretty much argue anything they fancy (seemingly often to suit some oddly motivated cultural or historical concept of vital importance to the peope making the argument). It doesn't even seem erudite.

      It's like reading tea leaves. I have more patience for that in archeology, where archaeological horizons and cultures are at least humanly interesting and worth learning about in their own right (compared to a trainspotter's guide-esque catalog of non-functionally different variants of mtdna and the y chromosome).

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      1. Haploid genetics' phylogenies are sound and straightforward, autosomal genetics on the other side are quite slippery because of all the recombination and drift happening along the long span of times we are pondering here. I don't say it's meaningless but I say that, understanding Native American haploid genetics one could infer something similar to Mal'ta. I for one was not surprised (rather vindicated) by its results, unlike so many others: totally to be expected judging only on yDNA Q and mtDNA X2 pylogenetically organized geography. Instead those giving too much emphasis to simple non-holistic interpretations of autosomal data were speculating about NA admixture in Europeans and what not (just because some modelings interpreted that, with some very oblique reason only).

        Haploid genetics give a quite sound mesh of reference, provided you ignore "molecular chronology" of course, which is almost invariably way off target. The problem is that the more complex the modeling or algorithm to apply, the more likely to be wrong. Simple straightforward anchors are not everything but they give a quite decent reference frame on what to fit the more elusive autosomal and other highly complex data analysis.

        The kind of discussion you mention seems a bit shallow for my understanding. At least I have always (I mean: since around 2006 or 2007) tried to use the phylogenies in their geographical context to reconstruct an approximation to human prehistory (with especial emphasis on Eurasia+ but also in Africa). If you are familiar with this blog you should know. For example: http://forwhattheywereweare.blogspot.com/2013/06/synthesis-of-early-colonization-of-asia.html

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      2. Actually I often see that kind of shallowness when interpreting (or rather over-interpreting) autosomal data. There are a lot of possible problems: highly drifted populations (which tend to form their own clusters just because they have an endogamous history), oversampling and undersampling of various groups (what can totally slant the results), misunderstanding affinity reflected in "components" as actual ancestral populations, speculating a lot on the direction of one's particular preferences (wishful thinking), etc.

        Anyhow well understood autosomal data is important: another complementary approach to population genetics but there must be a comprehensive holism of all possible approaches and autosomal data is often harder to analyze and understand properly. Generally I prefer to wait for several independent studies of this kind to converge onto something before I give it for granted (I've seen too many contradictory results to accept any single analysis as "the truth").

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    9. I find myself wondering what to make of the fact that, like Lochsbour, he has the combination of rather high cheekbones and noticeable browridge (the latter which you had commented in the case of Lochsbour as being a rare feature in Mesolithic Europeans). Btw, I have never heard of "C6"; is it the same thing that wikipedia is calling "C-V20"? Where else has it been found?

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    10. Oops, I see now from your Scozzari link, so V20 is C7; C6 is P55. Again, never heard of it. Interesting.

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    11. Wait. Wikipedia says, "C-P55 Found in the highlands of New Guinea[13]" ["^ Jump up to: a b Scheinfeldt, L.; Friedlaender, F; Friedlaender, J; Latham, K; Koki, G; Karafet, T; Hammer, M; Lorenz, J (2006). "Unexpected NRY Chromosome Variation in Northern Island Melanesia". Molecular Biology and Evolution 23 (8): 1628–41. doi:10.1093/molbev/msl028. PMID 16754639.]

      W.T.H.!

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      1. It is indeed Sozzari's C7-V20 (or so it seems, see Jean's objections above), which this paper (or at least everyone commenting on it) calls C6 and the latest ISOGG review describes as C1a2-V20. Last time I checked C-P55 had been dropped from the ISOGG tree, probably for lack of clear and plural samples.

        For what I see in the current ISOGG tree, it is being proposed as direct relative of Japanese rare lineage C1. Quite unexpected, I must say, but it seems not yet fully clear anyhow.

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    12. It seems to me that C6 is the remnant of the first migration of modern Homo sapiens to Europe. Those people probably had Y-haplogroups IJ and C* (that was later changed to C6).

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      1. Most likely there were two successive waves to Europe from West Asia ("first migrations", plural) in the early UP: the Aurignacoid one (possibly with several subgroups involved) and the Gravettian one, more homogeneous. Then there was most likely a long hiatus until the Neolithic without significant immigration to Europe (although some Europeans may have migrated outwards, to North Africa and parts of West Asia quite likely).

        As for which haplogroups they carried, some of them, judging on Mal'ta, where the same two waves are apparent via Altai, probably carried yDNA P (R1 in the West). It has yet to be attested with ancient DNA anyhow.

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    13. "So C1 is now perceived as a lineage that spans all Eurasia with an arguable South Asian centrality."

      That fits the model i have in my head:

      1) out of tropics into sub-tropical africa
      pause
      2) out of sub-tropical africa (still mainly coastal imo)
      pause
      3) out of india
      pause
      4) etc etc

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