January 11, 2011

Mitochondrial DNA of West Europe

Maria Lluïsa, of NeanderFollia[cat], points me to this interesting paper:


I am unsure about the access status of this particular paper, but at the moment of writing this, it is openly available online as advance online publication

Most interesting is surely the quite important database (mtDNA) covering most of West and Central Europe, from the Northern parts of Spain to Denmark and Poland (including many populations from France), available in the supp. material table S3.


Some fun with bidimensional representation

Let's begin the linearized Fst distances in a bidimensional graph (colors are mine, see below):

(click to enlarge)
In order to clarify a bit all that nightmare of acronyms, I marked Basque and Bearnese samples in red, those from the Paleolithic Franco-Cantabrian region in orange and those from the Rhine-Danube region in blue. The larger orange dot FCT is Perigord (Dordogne), the most important single district of Paleolithic Europe. 

You can maybe guess some of the simpler acronyms such as ENG (England) or AUS (Austria), I won't list them here (they are in the supp. material) but I must say for clarity that all beginning with F are from France, initial G means Germany and initial I means Ireland. All the others have unique areal acronyms and those ending in MI mean "miscellanea". 

Not much apparent structure is observed: basically there is a big blob in the middle (albeit divided in two not too well defined subclusters, I drew a dotted line to mark this internal division) and then four very isolated samples that actually describe the two axis of the graph. 

These "polar" samples are the following ones:
  • ALA is Araba (Álava in Spanish), a Southern Basque province mostly looking to the Upper Ebro. It defines the positive polarity of the first dimension and its main characteristic is to have 80% R-CRS (surely all H).
  • PAS is Valle del Pas (Cantabria), a mountainous district famous for its soft cheese biscuits, its religious architecture and the frequent visits of geneticists... to their archived data (a pity because they miss the delicious quesadas). They define the negative pole of the first dimension. They are particularly high in haplogroup V (24%) but low in CRS (27%). They are also high in U5 (13%), I (6%) and T2b (8%).
  • GUI is Gipuzkoa (Guipúzcoa in Spanish spelling). Their most salient characteristic, as far as I can tell, is an unusual high frequency of H2a (22% within H, 12% of all). They also have rather high frequencies of V (12%) and U5 (17%) and are quite to very low in the Neolithic clades (J, T, K, W, X).
  • COR is Cornwall. Their most notable characteristic is very high J (20%).
It is curious that all the first dimension of West and Central European diversity is synthesized in a line of some 50 Km or maybe a little more just SW of where I live. The authors seem to agree:

Northern Iberia appears to be microgeographically differentiated. Excluding the highly divergent Pasiego isolate (Maca-Meyer et al., 2003), there are also significant differences between the Basque provinces or between Catalonia at the north-eastern edge and Galicia-Asturias in the northwest. In fact, when these results were represented in an multidimensional scaling plot (Figure 2), the Pasiegos and Spanish Basques from Guipuzcoa and Alava were the most outstanding outliers, also followed by samples from Catalonia and Galicia, the French Basque sample and the British samples from Cornwall and Wales.

Even if we decide to disdain the extreme Araban and Pasiego samples, the next in line marking this polarity are Bearnese/North Basques (FSW) and Catalans (CAT), what implies a somewhat longer line from ESE to WNW along the Pyrenees. Somewhat different in the direction and distance but not too much in the geographical regions involved.

The second dimension is  quite different, defining a S-N axis along the Atlantic coasts of France, between Cornwall and Gipuzkoa. After ignoring the outlier poles, we can redefine this second dichotomy as being between West Ireland and Cantabria or something like that (same axis mostly, though moved to the West a bit).

The two main clusters show tendencies in these two axes: one (low, left) tends to Cornwall and the Pasiegos (or Catalans if you wish), while the other (up, right) tends to Basques (Arabans and Gipuzkoans specifically). The clusters are not too well defined anyhow and there is also a smaller third cluster formed by Biscayans, Provenzals and Cantabrians, which stands between the Pasiegos and Gipuzkoa (what, excepting the Provenzals, makes almost perfect geographic sense).

I have tried to represent the findings from the Fst graph in maps:

1. The polarity axes: red is dimension 1 (dotted line after replacing Pasiegos by Catalans) and blue is dimension 2:


It is... curious, right?

2. The clusters: blue and red are the populations in each of the two main clusters, marked with stars the four "polar" or outlier populations, with colors representing the cluster they are closer to. Green is the third minor cluster. Magenta are two populations (Switzerland and Morbihan) for which two different samples exist and each falls in a different cluster. I ignored the "miscellanea" samples.


If we are to hypothesize a Franco-Cantabrian origin to some of this duality (which is surely more complex than just that), we'd see that northern Franco-Cantabrian populations (Dordogne, Herault and Lyonais) fall in the blue cluster, together with several Atlantic populations from France, Scotland and Ireland, the Danubian fraction of  Central Europe and the Mediterranean fraction of Iberia. 

The red cluster instead looks more specifically Atlantic, with Basques/Bearnois and Asturians being the only ones from the Franco-Cantabrian region and otherwise being concentrated towards the North-West.

The lesser green cluster is totally Franco-Cantabrian but should represent a peculiar intermediate alchemy rather than a distinctive ancestral group, I suspect. Not that the other larger clusters are safely any representation of shared ancestry necessarily either but at least of some intriguing coincidence in their alchemy that seems to ask for further exploration.


Some intriguing details of specific haplogroups

The authors seem to take a critical stand, based in previous work, on the Franco-Cantabrian or even Catalan origin of haplogroup V. Different papers have offered strikingly different results on the frequency of this clade specially in Catalans (earlier claimed to be 24%,  now just 3%) and Gipuzkoan Basques (initially said to be 20%, now more like half that amount). García and colleagues seem to hint that V may have a southern Iberian origin after all:

Diversity values for V are significantly higher in Southern Iberia than in the Cornice (P<0.05). Excluding Scandinavia, the lowest diversities are found in Northern Africa and the Iberian northeast.

I say that this would totally fit in my model of important Ibero-African contacts in the context of the Last Glacial Maximum and the genesis of Oranian culture in North Africa. It is also consistent with the known fact that North African mtDNA H (sister of V, together making up c. 30% of North African mtDNA) is of Iberian or otherwise SW European derivation. 

There is also some mention to HV4, with a novel sublineage, HV4a1, of apparent origin in the Cantabrian strip (also found in one Italian and one continental European). Other HV4 sublineages are Eastern Mediterranean however, with its closest relative HV4a2, being found in Jordan and Egypt. 

In regards to H, it is worth mentioning the general high frequencies in the Cantabrian strip and specially high frequency of H6a among Cantabrians (12%), however it lacks diversity. 

H7 is confirmed as being most frequent in NE Iberia and SE France, it is one of the four H subclades with significant presence in North Africa as well. The authors however yield to the Mediterranean origins temptation, claiming presence in West Asia that is actually quite anecdotal (4/253 per Enafaa 2009). Excepting Catalonia, H7 is rare in Iberia but it is quite common in France instead, where it is largely concentrated (Álvarez-Iglesias 2009). The newly revealed presence in Catalonia offers a plausible origin for its North African presence in the context of the LGM transmediterranean contacts, which would be quite parsimonious considering that in general all mtDNA H in the region is of Iberian origin (Cherni 2008). But whatever. 

A key haplogroup however is H1, the largest H sublineage. In this aspect the authors find surprising heterogeneity. While the highest frequencies are in the Cantabrian strip, the highest diversity seems to be in the Mediterranean area (Italy and Balcans). Next in line come Scandinavia (Finns included), NE Europe and North Central Europe, all three tied at the same value (9.4). Paragroup H1(xH1a,H1b) appears to have also greatest diversity in Italy-Balcans, followed Scandinavia and then the Western Islands and NE Iberia (Catalonia and Aragon). H1a is clearly most diverse in NE Europe and North-Central Europe. H1b, a smaller scattered lineage, is most diverse in the southern Iberian Peninsula. Within H1:

FST pairwise comparisons based on haplotype frequencies detected unexpected heterogeneity. France showed close affinities with only the nearby north-east Iberian sample. In addition, the Scandinavians seem to be very different from north-central Europeans, showing more affinities to Slavs.

H3, the second largest H sublineage probably, is most diverse in North-Central Europe, in spite of being much more common in SW Europe (3-8%).

All this suggests that haplogroup H spread to SW Europe from Central Europe and not the other way around. At least H1b has been detected in Epipalelolithic Portugal (Chandler 2005, revised sequence assignment by me) establishing a maximum date for this spread. I would therefore think that mtDNA H subclades expanded at the latest with the Gravettian wave because there are no more cultural flows towards SW Europe with that origin before the late Bronze Age (or the late Chalcolithic if you wish to consider Bell Beaker - not me).

It is quite surprising anyhow to find such relative low nucleotide diversity levels for these so abundant clades not just in the smaller NW and NE Iberian regions but specially in France, while South Iberia generally shows greater diversity instead. The results raise more questions than provide answers in this sense.

Recently described haplogroups H1r and H1t were found to exist among Basques. H1t seems to be an Iberian-exclusive clade, while H1r seems instead continental (found in one French and one "European", as well as one Basque now). 

In partial contrast haplogroup K has highest diversity in NE Iberia. However the differences are not too large for all Europe except the NE, where diversity is very low. 

T2b has highest diversity in Northern Iberia (both NW and NE), followed by North Africa.

Excluding two too low in number, the greatest diversity of W is in Balcans-Italy, followed by Iberia (all three regions).


Franco-Cantabrian post-Glacial expansion?

Apparently not. This seems the main conclusion from this paper and, on light of the perplexing diversity values, even for H3, I have to agree. H arrived here from Central Europe and probably Italy, already diversified to a large extent, and did not move much after that. This arrival probably happened in the Gravettian or Aurignacian periods.

It is however still possible that a male-biased expansion happened with Magdalenian, as suggested by the patterns of R1b1b1a1a2, the most common R1b sublineage, at least in Europe, which seems to have most of its phylogenetic diversity (safer than nucleotide diversity, which is the one analyzed by García et al. in the mtDNA) around the Pyrenees (see here). However a local Central-North European component is still evident in its smaller brother haplogroup R1b1b1a1a1.

Another caution is that no meaningful sampling of French mtDNA has been undertaken since 2004 (Dubut et al., data recycled for this paper) and, considering the many errors and flukes happening in other cases without enough second and third samplings, it is very possible that a lot is still hiding in that area, so important in European prehistory.

19 comments:

  1. The idea of mtDNA selection was rejected until recently I believe

    Do you think mtDNA been selected for more or less than is currently believed ? Do you think cold weather or agriculture could have resulted in some older mtDNA disappearing altogether.

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  2. Great work indeed. About mtDNA HV4a I think having demonstrated (I know you don’t like this word, but science is made by demonstrations, proofs, confutations) on “WorlFamilies”, examining the mitochondrial data (FGS) of “NealtheRed” (Downing), who has an Italian mother, and of “Humanist” (an Assyrian whose surname I can’t reveal), that its origin is in Italy. Its expansion is at the level of the Younger Dryas, as I have always said for the masculine counterpart R1b1b2.
    The paper falsifies the theory of the Cantabrian refuge and underlines, beyond France, the Mediterranean peninsulas. But why don’t say: Italy?

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  3. The blue clusters show something of an affinity for places that have the strongest Celtic influences. I'm not necessarily suggesting that this is the timing of the cluster differences, similarly that they are a fit for the cultural geography.

    The red clusters show something of an affinity for areas Germanic influences, either linguistically, or via Atlantic Viking raids, although this also has problems since Viking raids should create Y-DNA rather than mtDNA clusters.

    The instances of greater diversity in a number of cases in Southern Iberian than in the Franco-Cantabrian region, suggests that maybe LGM refugees in Western Europe went as far South as they possibly could because every place else was too damn cold. Perhaps Europe wasn't very habitable further North than 42 degree North Latitude then.

    The study also seems to support more gene flow from Iberia to North Africa than in the other direction.

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  4. @Ken:

    "Do you think mtDNA been selected for more or less than is currently believed ?"

    I do not have a solid opinion but I think that rather not. Selection may be at play at new lineages (pruning them before they can expand, in addition to drift) but whole established haplogroups such as N1a or I? It does not make any sense.

    However I might be wrong.

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  5. @Gioello: I can't really discuss HV4a. The paper proposes a Western (Cantabrian strip) origin for HV4a1 and seems to accept an Eastern one for the rest HV4, including HV4a2, only reported in Egypt and Jordan in this paper. That's all I can say, and being a minor lineage, I imagine that the subject remains open.

    The case looks somewhat similar to that of that U8 anyhow, which, as you know, has a Basque sublineage and then U8b'K which has greater likelihood of Eastern Mediterranean origins and an Italian connection. It's difficult to say, really, and maybe best is just to understand them as part of the colonization of Europe, with ultimate origins in West Asia in all variant theories.

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  6. @Andrew:

    "The blue clusters show something of an affinity for places that have the strongest Celtic influences".

    I also considered that but the parallel collapses from France to the South in any case. The clusters are not too relevant anyhow: they are weak (fuzzy). Also I'd expect more Celtic-specific genetic input in England than in Ireland or Scotland, honestly. One thing is what language you keep today and another thing is where the Celts arrived first and most likely made the greatest impact. Similarly in France, I'd expect more "Celtic blood" in the North than in the South, etc.

    "The red clusters show something of an affinity for areas Germanic influences, either linguistically, or via Atlantic Viking raids, although this also has problems since Viking raids should create Y-DNA rather than mtDNA clusters".

    Yes, it's very fuzzy. If anything, the Red cluster seems to mean greater H (and maybe greater U as well). Arduous confirmation would be needed but that's my impression so far.

    It is possible that they simply indicate greater/lesser Neolithic impact or, alternatively/complementarily two somewhat distinct (but related) waves or vectors in the European colonization as such.

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  7. Forgot:

    "Perhaps Europe wasn't very habitable further North than 42 degree North Latitude then".

    That's not real. I have already directed you in the past to Bocquet-Appel 2007, which is just one of many papers and books that could tell you that the Franco-Cantabrian region was densely populated all the time, including the LGM. Also in general greatest diversity is to Central Europe, which is above 45 degrees (the latitude of Dordogne).

    More real is that there was never a total depopulation of Central Europe and that at least part of the lineages present today have been there since Aurignacian or Gravettian times.

    "LGM refugees in Western Europe went as far South as they possibly could because every place else was too damn cold".

    This may explain Gravettian expansion, specially in Iberia proper. As you may know, Gravettian in SW Europe is a late arrival (like 22 Ka, while in Central Europe and Italy is like 28 Ka) and almost overlaps with Solutrean, which seems a reaction against this techno-culture, often described as "intrusive" in the Franco-Cantabrian region. In Iberia proper, however, Gravettian made a very strong impact (Aurignacian colonization was sparse) and recycled Solutrean that here is the "intrusive" element into a quite unique Gravetto-Solutrean.

    But it's not like the northern areas really got depopulated, certainly not the FC region nor Moravia (where the evidence is clear) and I'd dare say that other pockets must have persisted, re-expanding when climate improved. It's damn cold in Lapland but people live there anyhow (and they would be able with UP tech as well). Where they do not live is (only) in glaciated areas, such as Central Greenland. This ice cap extended then to Central-South England, Denmark, Mecklemburg, most of Poland, etc. So it's reasonable to think that some people lived in the Netherlands or even the southern coasts of England and Doggerland, even if we lack evidence today.

    There are even theories that propose that Aurignacian survived over there (some evidence seems to exist, though not sure how much or how good) and this influenced the Aurignacian renaissance we call Magdalenian.

    In this sense we might argue that the blue and red clusters might represent Gravettian and Aurignacian waves respectively. It sounds better than "Celtic" to my ears, certainly. But it is just an speculation so far.

    "The study also seems to support more gene flow from Iberia to North Africa than in the other direction".

    That is what I think as well. However the characteristic Y-DNA of those contacts seems to have receded in North Africa more than in Europe. I'd expect that 30% European (H and V) mtDNA in North Africa should correlate with at least 30% European Y-DNA once. But today it is little more than 5% (albeit 17% in Guanche mummies). Instead E1b1 in Europe is in good apportion with what can be considered North African mtDNA (U6 and L(xM,N)).

    So Capsian culture in North Africa must have implied some sort of male-biased replacement but I can't detect anything of the like in Europe instead.

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  8. "[T]he characteristic Y-DNA of those contacts seems to have receded in North Africa more than in Europe."

    This makes sense to me. The arrival of camels and then of horses and associated Berber and then Muslim technological and cultural influences would provide a perfect situation for North African and/or Middle Eastern men from outside the area of Iberian genetic influence to sweep in and displace local men in the influenced area, much as the Indo-Europeans, Turks, Mongols and Yaoyi did in much of Eurasia and Japan.

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  9. Andrew: you make a lot of assumptions: "The arrival of camels and then of horses and associated Berber and then Muslim technological and cultural influences would provide a perfect situation for North African and/or Middle Eastern men from outside the area of Iberian genetic influence to sweep in and displace local men in the influenced area".

    All this tells me more about your mentality and your historical fetishes than about what really happened, specially because all is out of the hat, nothing has any particular second thought or greater depth to it.

    The main problem seems obvious: the camel arrived happened via Egypt and was spread by the pre-existent Berbers: the arrival of the camel supposes no apparent change, maybe economical but not linguistic or ethnic (except maybe in the Sahara but that's a very specific place with very low density and peculiar peoples).

    The other main problem is that there is no meaningful Arabian Y-DNA in North Africa: please compare Arabia Peninsula Y-DNA or your usual Bedouin Y-DNA with North Africa: there is not J2 in North Africa (except at low levels in Tunisia and there it is much more likely to be Phoenician in fact).

    So you imagine how it ought to be first and then try to make the facts, the data, fit in that script.

    But I do not have those fantasies, those preconceptions (maybe I have others but camels and "orientalism" don't seem to work with me), and I know some facts instead, so your fantasy does not fit with what I know and does not even feel interesting at all in an emotional aspect either. So I cannot share your enthusiasm for camels or Arabs being much involved in this change either.

    It leaves me so cold and yawning that I'm going to sleep right away. And I really mean it.

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  10. Maju : "But it's not like the northern areas really got depopulated, certainly not the FC region nor Moravia (where the evidence is clear) and I'd dare say that other pockets must have persisted, re-expanding when climate improved. It's damn cold in Lapland but people live there anyhow (and they would be able with UP tech as well). "

    Also my opinion.

    This kind of things supports it IMO (indirectly) :

    http://www.thelocal.se/23546/20091129/

    "People lived in the Torne River Valley on the border with Sweden and Finland some 11,000 years ago, an important new archaeological find has shown."

    "The settlement, found near Pajala in the far north of Sweden, are the oldest known find in the county of Norrbotten, according to the archaeologist Olof Östlund."

    "pajala" on Google maps.

    And must have been pretty cold in Sunghir too, ~25,000 yrs ago.

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  11. "The main problem seems obvious: the camel arrived happened via Egypt and was spread by the pre-existent Berbers: the arrival of the camel supposes no apparent change, maybe economical but not linguistic or ethnic (except maybe in the Sahara but that's a very specific place with very low density and peculiar peoples).

    The other main problem is that there is no meaningful Arabian Y-DNA in North Africa"

    My conjecture was, intentionally, more narrowly phrased than you are reading it, because I am aware that there isn't much Arabian Y-DNA in North Africa and don't have evidence of Berbers being replaced by ethnically non-Berbers. Instead, I specificly refrained from suggesting that it was the Arabs themselves who demically brought the profound cultural changes that Islamization brought about, or there was replacement of North African populations by non-North Africans.

    My assumption was that once upon a time there were ethnically homogeneous Berbers, but that this changed with Iberian admixture. After that point in time you have Northwest African Berbers (call them proto-Moroccans and proto-Algerians if you will) who have significant Iberian DNA, and "near Egyptian" Berbers (call them proto-Tunsians and proto-Libyans and proto-West Egyptians perhaps) who don't. It may be a gradual cline that is significant simply because the distances are so great in the region, or it may be due to a clear divide caused by the Atlas Moutains, the result is the same either way.

    The "near Egyptian" Berbers get camels, horses, Islam, and who knows what else, in multiple waves of new technology via Egypt (mostly early Neolithic and Islamic respectively, but possibly other waves in between as well), make their way West and with their newly superior technology displace the Iberian admixed Berber men with Berber men who aren't very Iberian admixed. Power is sexy and technology confers survival advantages for the whole family. Voila! A lower proportion of European Y-DNA in North Africa.

    Intra-ethic conflict of this kind by the first to get new technology has been seen in modern times, for example, in the New Zealand Wars in which some Maori tribes with guns ravaged those that lacked them.

    There is a fundamental flaw with this scenario only if there is no population genetic cline from West to East in North Africa among Berbers, and this seems unlikely, particularly in the pre-camel, pre-horse, pre-domesticated animal era. In that era one would expect far more localized gene pools than existed once riding animals appeared.

    I have good company in thinking that the camel and domesticated animals had a profound cultural impact and confer great selective advantage.

    It so happens that I personally think that it is more likely that there was linguistic shift to Berber from a non-Afro-Asiatic language around the time that the camel arrived in Berber history (although probably not much demic replacement, because North Africa may not have been too attractive as a place for newcomers to settle at the time). But, that doesn't have much relevance to the population genetic effect of that kind of scenario.

    It is beyond dispute, of course, that there was significant linguistic shift in connection with Islamization - and linguistic shift basically never happens without other significant cultural or political changes as well. People don't drop their languages because they're bored and want to try something new.

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  12. it's not like the northern areas really got depopulated.

    The Garcia paper states right at the start that the LGM "almost depopulated central and northern areas of Europe."

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  13. @Wagg: good point, because that's pretty much the heart of Lappland precisely.

    ...

    @Andrew:

    "My assumption was that once upon a time there were ethnically homogeneous Berbers, but that this changed with Iberian admixture. After that point in time you have Northwest African Berbers (call them proto-Moroccans and proto-Algerians if you will) who have significant Iberian DNA, and "near Egyptian" Berbers (call them proto-Tunsians and proto-Libyans and proto-West Egyptians perhaps) who don't. It may be a gradual cline that is significant simply because the distances are so great in the region, or it may be due to a clear divide caused by the Atlas Moutains, the result is the same either way".

    That does make sense, and probably reflects what happened in Capsian times. But has nothing to do with Islam or Romans, not that I can see. If it has anything to do with the introduction of camel, I'd like to see a relevant paper or at least a recount of relevant archaeological evidence (nothing so far). I'd imagine that horse was introduced from the West in Chalcolithic times or Bronze Age the latest anyhow.

    But is anyhow a much more beautiful model than just vagenesses: Capsian would in this model be just the beginning of a more or less sustained East to West flow that gradually reduced the European ancestry, specially in the Y-DNA side. I can tentatively agree with that, however let's not forget that Morocco, Algeria(coasts) and Tunisia (coasts) can harbor much larger populations than all Lybia can. This is a major issue when one posits that a tiny nomadic group from a semi-desert area replaced a huge sedentary group, like Turkmens "colonizing" Anatolia (what obviously never happened but in minimal amounts).

    In general the more clear archaeological evidence of an East to West flow is Capsian culture (from Nubia rather tan Lybia), however what you say makes some sense as reinforcement of this process. Much as Metal Ages Mediterranean flows surely reinforced (in some cases dramatically I understand) the early Neolithic colonizations. Yet these are the ones which made most of the impact probably.

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  14. "It is beyond dispute, of course, that there was significant linguistic shift in connection with Islamization - and linguistic shift basically never happens without other significant cultural or political changes as well. People don't drop their languages because they're bored and want to try something new".

    Obviously Arabic was imposed on the conquered peoples and consolidated via Islam. The same happened with Latin and so on.

    Nobody denies that there was a socio-political change with Islam in North Africa, what I reject is any major colonization from West Asia or Egypt at that point. But the "Arabs" from some point on are more likely to be descendants of Vandals, Romans and Carthaginians (along with, of course, some denationalized Berbers themselves and a few drops of genuine immigrant soldiers and top administrators). It is these cosmopolitan, often urban groups, which quickly assimilate in the new order, while the rural, mountain, ones often retained a Berber identity instead.

    "The Garcia paper states right at the start that the LGM "almost depopulated central and northern areas of Europe.""

    That is not what they conclude or can be concluded from mtDNA diversity: they reject recolonization from the SW and instead continuity in most lineages seems dominant. "Almost" is anyhow not the same as "totally". And I doubt Central-North Europe would retain such a high apparent diversity for all lineages if it was depopulated in any radical way. What the paper seems to support for most lineages is a Central European origin, which can only be understood in the context of European colonization.

    This has been my main model since I discovered the super-star of mtDNA H, but I did not expect this to extend so radically to almost all clades. However, if H1, H3, H4, H7 and V were to be in South Iberia in the LGM, so they could jump to North Africa, it is obvious they had to be consolidated by then and not to be signatures of "post-glacial expansion". So yeah, we need more depth in the analysis of H1 and the other many H subclades, as well as U5, V, etc. if we are going to clarify if there is any mtDNA signal of such glacial re-expansion or if this is restricted to Y-DNA (male biased migration like in North Africa - but less exaggerated maybe).

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  15. The Jordanian HV4a2 and I, unlike the Egyptian, appear to share transitions at 16287 and 16311 as well. I am accession # HM228422 (Assyrian) on GenBank. The transition at 16287, in addition to the Jordanian and I, is also shared by men with similar haplotypes on SMGF. One is a fellow Assyrian, and the other, I believe, is either an Iraqi Kurd or Iraqi Arab from northern Iraq.

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  16. Your haplogroup (HV4a2) is clearly a Middle Eastern clade. Probably it has been differentiated there, and there is an Assyrian/Iranian etc. clade different from the Egyptian one. But your clades are anyway derived. The Ancestor clade is HV4 and I think have demonstrated on Worldfamilies that, even though there is an Assyrian (Humanist), the Italian samples are more ancient and at the origin. The other more ancient are the Russian ones: then this haplogroup was born in Europe, either Eastern or Western one. Malyarchuk has demonstrated that many Western European haplogroups peopled Eastern Europe about 15,000 years ago and HV4 could be one of them.

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  17. It is very likely that QYV9Y (16168T, 263G, 309.1C, 309.2C, 315.1C), found on SMGF and put on Mitosearch, from Italy, is an ancient HV4* like AY738941, even though we'd need an FGS for being sure.

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  19. This paper showed up again in the debate here.

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