August 13, 2014

Chalcolithic mtDNA from Atapuerca still in the Neolithic range

Bell Beaker Blogger points me to this latest study on ancient mtDNA from the Center-North Iberian Peninsula, including 20 samples from a Chalcolithic site (without Bell Beaker apparently) that clearly shows continuity with mainline Neolithic (Cardium but also similar to Central European Linear Pottery Culture, both sharing the same Thessalian ultimate origins).

Daniel Gómez Sánchez, Iñigo Olalde et al., Mitochondrial DNA from El Mirador Cave (Atapuerca, Spain) Reveals the Heterogeneity of Chalcolithic Populations. PLoS ONE 2014. Open accessLINK [doi:10.1371/journal.pone.0105105]

Abstract

Previous mitochondrial DNA analyses on ancient European remains have suggested that the current distribution of haplogroup H was modeled by the expansion of the Bell Beaker culture (ca 4,500–4,050 years BP) out of Iberia during the Chalcolithic period. However, little is known on the genetic composition of contemporaneous Iberian populations that do not carry the archaeological tool kit defining this culture. Here we have retrieved mitochondrial DNA (mtDNA) sequences from 19 individuals from a Chalcolithic sample from El Mirador cave in Spain, dated to 4,760–4,200 years BP and we have analyzed the haplogroup composition in the context of modern and ancient populations. Regarding extant African, Asian and European populations, El Mirador shows affinities with Near Eastern groups. In different analyses with other ancient samples, El Mirador clusters with Middle and Late Neolithic populations from Germany, belonging to the Rössen, the Salzmünde and the Baalberge archaeological cultures but not with contemporaneous Bell Beakers. Our analyses support the existence of a common genetic signal between Western and Central Europe during the Middle and Late Neolithic and points to a heterogeneous genetic landscape among Chalcolithic groups.

The results show intense similitude with Catalan Neolithic and Languedoc's Chalcolithic but also with Central European Neolithic. They contrast instead with Portuguese and Basque Neolithic, as well as with Central European Bell Beaker, all them much higher in haplogroup H and, in the Basque case, also in U (being the most modern-like of all ancient mtDNA pools known before the Bronze Age in Europe).

Annotated version of fig. 2
Figure 2. Mitochondrial DNA haplogroup frequency for 21 ancient European samples.
This study: El Mirador (MIR). Published prehistoric cultures [21]: Hunter-gatherer central (HGC), Linear Pottery culture (LBK), Rössen culture (RSC), Schöningen group (SCG), Baalberge culture (BAC), Salzmünde culture (SMC), Bernburg culture (BEC), Corded Ware culture (CWC), Bell Beaker culture (BBC), Unetice culture (UC), Funnel Beaker culture (FBC), Pitted Ware culture (PWC), Hunter-Gatherer south (HGS), (Epi) Cardial (CAR), Neolithic Portugal (NPO), Neolithic Basque Country and Navarre (NBQ), Treilles culture (TRE), Hunter-gatherer east (HGE), Bronze Age Siberia (BAS), Bronze Age Kazakhstan (BAK).

Notice please that the above column for Central European Bell Beaker (BBC) includes the more than dubiously attributed Kromsdorft site, which has a totally different genetic signature. In my previous analysis of European ancient DNA evolution, I treated them separately and I still think that it is much more correct to do it that way.

Notice also that one sample from Mirador was sequenced for autosomal DNA by Evangelia Dasakali, producing an Italian-like sequence, roughly in line with other early European farmers, excepted the Atlantic ones. See here.

So what we see is a "wedge" of Mediterranean or Neolithic ancestry probably penetrating along the Ebro river up to Atapuerca and contrasting with Atlantic Iberian (Basque and Portuguese) ancestry, more modern-like or even "hyper-modern" (by contrast with mainline farmers) in the case of Portugal. This same contrast exists with Central European Bell Beaker sites and also (in the autosomal DNA aspect) with Megalithic farmers from Southern Sweden. 

None of those "modernizing" tendencies can be found instead among Eastern European Neolithic peoples nor among early Indoeuropeans of Central Europe such as those of the Unetice culture. So the overall conclusion can only be that there was in the Chalcolithic (and to some extent Neolithic) a duality of ancestries between the Mainline or Mediterranean (but also Danubian) Neolithic and a more "modern" Atlantic Neolithic, which is related to the Megalithic and Bell Beaker cultural phenomena (and in North-Central Europe also to Funnelbeaker). 

There are still a lot of dark spots in our understanding of how this "modernization" or "Westernization" of the genetic pool happened but, in general terms, it seems to imply a sizable (albeit somewhat irregular) demographic flow from Atlantic Europe into the areas previously occupied by Mainline Neolithic populations of partial West Asian affinity. This is apparent in both mtDNA as in autosomal DNA. 

These sequences from El Mirador (Atapuerca) only underline this phenomenon and the fact that in the Chalcolithic, some 4500 years ago, this process of "modernization" of the European genetic pool was still incomplete.

52 comments:

  1. Maju, these results may also have something to do with the idea that there were two 'poles' of Neolithicozation in Iberia - a northern (Danubian) one derived from LBK and a second , southern (Impressed & Cardial) pole which met in Central Iberia then underwent complex inrermixtures

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    1. Simply put: there is no LBK whatsoever in Iberia nor Southern France. What these and other similar results indicate is that the main two Neolithic waves of Europe (Impressed-Cardium and Painted-Linear) were product of a unique meta-ethnicity, with the same genetics, the same origins (Thessaly) and quite probably the same linguistic family (which I believe should be Vasconic). Their genetic heritage is better preserved in Southern Europe though, although here also there was some sort of back-flow from the Atlantic carrying greater frequencies of aboriginal genetics (and later, in the Late Bronze and Iron ages from Central Europe, carrying the Indoeuropean languages).

      I can't imagine where you could get the idea that LBK ever made it past the Loire. It simply didn't - instead Cardium-like La Hoguette culture (and/or other locally specific groups) overlaps with LBK in the areas of Northern France, West Germany and Belgium, making the wider area around the Rhine not so strictly Danubian but a crossroads of both traditions, as well as some innovative local ones, particularly in Belgium.

      Relatively good generic map of early Neolithic in West-Central Europe (not so good re. La Hoguette and other local Neolithic groups though)
      Map showing overlap of LBK and other non-LBK Neolithic groups in the Rhine area
      Decent map of La Hoguette sites (dots) with core, very approximative, Danubian/Cardium areas shaded in grey

      In any event there was absolutely no LBK in Iberia nor Southern France. The explanation to genetic similitudes between LBK and Cardium early farmers must be another: namely a shared origin in Thessaly.

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    2. Uh... first map link is broken. It is this one:

      Relatively good generic map of early Neolithic in West-Central Europe (not so good re. La Hoguette and other local Neolithic groups though)

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  2. Do you think this "modernization" could have come from NW Africa in the Upper Paleolithic? IE a non-EEF non-WHG population. Would explain a few things I think, especially if it brought R1b with it.

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    1. No. For many reasons, both genetic and archaeological.

      The kind of genetic flow is not at all related to NW Africa, certainly not by origin. While mtDNA H is present in NW Africa it clearly derives from SW Europe and not the other way around (almost certainly Solutrean→Oranian flow). Y-DNA R1b is very rare in NW Africa and the immediate origin of the European clades is clearly in Western Europe, while upstream it comes from West Asia, via the Balcans. Autosomally we don't see either NW African components in Iberia or Europe but the opposite is true (much of the NW African genetic pool seems to originate directly in Europe). I can give references but I'm tired now. Please ask later.

      Archaeologically there's also no obvious Africa → Europe influence. Again, if anything, the opposite would be true (Megalithism, some Bell Beaker).

      The source(s) must be in Atlantic Europe.

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    2. I thought the phylogeny contradicted an Atlantic origin for even the European R1b subclades. Do you think this modernization was a different wave than whatever spread R1b?

      And any thoughts on how R1b got its present distribution in both Western Europe and in Cameroon?

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    3. → http://leherensuge.blogspot.com/2010/08/r1b1b2a1-is-almost-unique-of-west.html

      R1b-S116 quite clearly stems from South France (where it seems most basally diverse), while R1b-U106 may have spread from around the Rhine basin (highest frequency in Netherlands, although unsure about where the greatest basal diversity is).

      Their common precursor (rather rare in "asterisk" variants) R1b-M412 may have spread from Central Europe and/or SW Europe. The ancestor of this one R1b-M269 may have spread from either the Balcans or West Asia.

      Only upstream of this node we enter the wider R1b of extra-European distribution, including the Central Asian/Bashkir and the Afro-Mediterranean clades, as well as some other that are ill studied AFAIK.

      So for me:

      1. R1b (as a whole) originated in West Asia, spreading in various directions.
      2. R1b-M269 (Euro-Asian clade), spread from either West Asia or the Balcans.
      3. R1b-M412 (European clade), spread from either Central or SW Europe.
      4a. R1b-S116 (SW European clade) spread from the Franco-Cantabrian Region.
      4b. R1b-U106 (NW European clade) spread from possibly the Rhine basin or the North Sea area (although an Upper Danube origin is not discarded for all I know).

      Particularly step #4 dissuades me from contemplating a recent origin, as well as some chronological estimates, that would make M412 and S116 Magdalenian, while R1b as a whole would be slightly older than Gravettian.

      As for the Afro-Mediterranean R1b-V88 haplogroup, it has no particular relation with the mainstream European R1b, being another branch of the early R1b spread from West Asia almost certainly. I've discussed it HERE. I would think that V88 as such spread from West Asia, with a branch reaching the Nile (Sudan particularly) and, from there, spreading to Central Africa in the context of the Chadic expansion (Neolithic?) Other branches reached Italy and Sardinia-Corsica, etc. It's not impossible however that it may have spread from Africa, much as E1b did.

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  3. LBK was not in southern France but in Northern.
    thirty years ago, the Norman population was supposed to be LBK, essentially, and that from the Neolithic to the Norman period.
    As Norman ( Cotentin peninsula), I am shifted west, I have a lot of Atlantic component and I am a pretty good match for Basques and southwest French( in K15 eurogenes)
    that is difficult to know without ancient samples, but for you Normandy is more Long barrows or more LBK ?

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    1. The Neolithic of Northern France, Belgium and much of West Germany presents a good deal of complexity: LBK elements are there for sure but also La Hoguette (Cardium-related) and Limburg (local) cultures, as well as Megalithism, etc.

      Anyhow the LBK genetic pool was clearly not modern: further demographic processes took place that heavily modified it in the Chalcolithic (and maybe also later). With due caution, I would think that the Megalithic phenomenon was largely demographic and "westernized" the rather "oriental" LBK (and Cardium) genetic pool.

      As for Normandy, High Normandy (west) was mostly within the same cultural region as Brittany and nearby West French areas (Armorica in historical terminology). Instead Low Normandy was in the LBK/Limburg/La Hoguette area. But as I just said, the cultural and demographic landscape suffered major ulterior shifts, so these differences were surely blurred later on, particularly as the Artenac culture (Megalithist archers from Dordogne) spread over all West France and Belgium. This may well explain (at least partly) what you say about being "a pretty good match for Basques and southwest French( in K15 eurogenes)".

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  4. Thank you for your speed reply

    We read a lot about the long barrows and more about bell beakers, but little about the Artenac culture, I remember you wrote that they stopped the advance LBK, you think they had a significant contribution in terms of demographics in Nothern France, to be honest I had never heard of them before reading your blog.

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    1. There's no much info online on Artenacian culture (although there is no doubt more in French, search "culture artenacien"). I know from books however that they were an important late Chalcolithic force which took over the whole West of France (also Belgium), replacing the previous cultures, notably the Seine-Oise-Marne one, which in turn had displaced the ancient Armorican Megalithic culture. They are often understood to be direct precursors of the historical Aquitani (i.e. Basques).

      Its characteristics were AFAIK abundance of arrows (often described as an "archers' culture") and a dolmenic Megalithic funerary fashion much more modest in size than what was styled by Bretons (I would say that a much less hierarchic society therefore, maybe by loss or weakening of the proto-druidism that may have persisted in Great Britain instead). The former core Seine-Oise-Marne area between the Seine and the Rhine (i.e. historical Belgium) had a slightly different facies, maybe reflecting its different, more loosely "Danubian" substrate. In any case that was about the time when the last remains of any Danubian (LBK) legacy were erased in all Europe because Artenacian is contemporary with Corded Ware and Vucedol.

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  5. "I would think that V88 as such spread from West Asia, with a branch reaching the Nile (Sudan particularly) and, from there, spreading to Central Africa in the context of the Chadic expansion (Neolithic?) Other branches reached Italy and Sardinia-Corsica, etc. It's not impossible however that it may have spread from Africa, much as E1b did. "

    Many HLA alleles and haplotypes which are shared between Sudan and Italy + Sardinia/Corsica are found in other African populations but not really very much in West Asia if at all. I can provide numerous data to support this.

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    1. HLA immunity alleles are not reliable ancestry markers because they are subject to strong selective pressures. Italy for example was one of the areas of Europe most strongly affected by malaria (once known as "Roman fever") and therefore also has unusually high frequencies of thalassemia, a kind of anemia that is an effective anti-malarian protection, which has anyhow closest relation to what is found in Greece, Southern Anatolia, Cyprus and Palestine. It is perfectly possible that the HLA markers were selected for the very same reasons (malaria) or similar ones. Although it suggests some kind of relatedness, being markers subject to strong positive selection subject to environmental pressures, this relatedness may be hair-thin (standard introgression between two otherwise almost totally distinct populations).

      R1b-V88 is found in an arch between Sardinia/Italy and the Eastern half of the Sudanese corridor, with clear presence in West Asia. Considering the overall spread of R1b (centered in West Asia) and the geographic pattern of the subclade, I'd say that a West Asian origin is at the very least quite parsimonious. Direct Sudan-Italy relations are most unlikely in any case, so the Eastern Mediterranean must have acted as corridor at the very least (but I lean towards origin).

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  6. The malaria protective HLA allele is B*53. [See: http://ghr.nlm.nih.gov/gene/HLA-B ]
    This is in fact at only a very low frequency in Corsica and Sardinia [both precisely 0.05%].
    The alleles and haplotypes which I am referring to play no protective role with regards to malaria.
    One example:

    B*58:01 in Europe

    Sardinia 6.40% [Peak European frequency - not evidence for selection, probably founder effect]
    Corsica 4.50%
    Switzerland Basel 4.21%
    Italy Rome 3.50%
    Italy North Pop.3 3.30%
    Switzerland Geneva Pop.2 3.26%
    Portugal Castelo Branco 2.80%
    Portugal Leiria 2.60%
    Croatia Pop.2 2.50%
    Portugal Santarem 2.50%
    Portugal Setubal 2.50%
    Portugal South Pop.2 2.50%
    Greece Pop.8 2.41%
    Switzerland Lugano 2.40%
    Portugal Evora 2.40%
    Portugal Portalegre 2.40%
    Portugal Braganca 2.30%
    Portugal Faro 2.30%
    Portugal Vila Real 2.30%
    Spain Ibiza 2.30%
    Switzerland Luzern 2.26%
    etc. etc.

    Lowest Euro frequencies:-
    Belgium 0.50%
    England North West 0.50%
    Serbia Pop.2 0.50%
    Wales 0.50%
    British (a) 0.50%
    British (b) 0.48%
    England Liverpool 0.45%
    Ireland South 0.40%
    Ireland South Pop.2 0.36%
    Ireland Northern 0.30%
    England Newcastle 0.25%
    Serbia 0.20%
    Sweden Uppsala 0.15%
    Scotland Orkney 0.00%
    Spain Gipuzkoa Basque 0.00%
    etc. etc.

    Where might it have entered from?

    Israel Arab Druze 1.40%
    Jordan Amman 1.40%

    Sudan Mixed 4.50%
    Tunisia 4.00%
    Tunisia Ghannouch 3.70%
    Morocco Settat Chaouya 3.40%
    Tunisia Pop.3 3.40%
    Mali Bandiagara 2.20%
    Sudan East Rashaida 1.86%
    Morocco Nador Metalsa Pop.2 1.40%
    [Searching for data from Libya and Egypt]

    Kenya Nandi 10.00%
    Kenya 8.00%
    Guinea Bissau 7.80%
    Kenya Luo 7.00%
    Senegal Niokholo Mandenka 6.90%
    Burkina Faso Fulani 6.10%
    Uganda Kampala Pop.2 6.00%
    Burkina Faso Mossi 5.70%
    Cameroon Yaoundé 5.40%
    Cameroon Bamileke 5.20%
    Sao Tome Island Angolar 4.70%
    Zimbabwe Harare Shona 4.40%
    Burkina Faso Rimaibe 4.30%
    Ghana Ga-Adangbe 4.20%
    Cameroon Bakola Pygmy 4.08%
    South Africa Natal Zulu 4.00%
    Cape Verde SE Islands 4.00%
    Uganda Kampala 4.00%
    Cameroon Beti 3.70%
    Cape Verde NW Islands 3.20%
    etc. etc.

    Source: http://allelefrequencies.net/

    There are several other HLA alleles and haplotypes in the Sardinia and Corsica + Italy populations which follow a similar pattern, again none of the alleles in question is involved with malaria protection. HLA should not be dismissed out of hand, if one learns which alleles have disease protective or susceptibility associations [they have been studied very extensively over the last 40 years and there is a huge amount of information and data available].

    There seems to be a reluctance on many people's part to consider data which may conflict with the existing paradigms. Many people also seem reluctant to accept the possibility of prehistoric migrations directly across the Mediterranean from Africa into Europe despite data which suggests this. I would be interested to know why. Looking at HLA data [involving alleles with no known disease protective role] I can see evidence of a number of these.

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    1. It may have been selected for whatever any other reason (plague epidemics for example) or maybe some aspects of malaria defense are not well known yet. In any case it is clearly an immunity marker subject to strong selective pressures and therefore not really informative re. ancestry. Frequencies do not matter either because in order for an adaptive marker to be selected for in the long run (consider repeated recombination and selection) only one ancestor is needed (regardless of the overall population). A low frequency allele here can become a high frequency one there, after lesser migration from here to there, just because it does work.

      That's why neutral markers (or those believed to be more neutral) are preferred when researching ancestry. Among them haploid lineages are the safest ones because they do not recombine at all (although they can still provide advantages in theory).

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    2. In any case your data for West Asia is very limited: you have not provided any data for most of the region and the two samples provided are almost certainly not too representative (particularly not the Druze, who have undergone dramatic endogamy-caused bottlenecks and may be partly of non-local origin).

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    3. Also the frequencies you provided for B*53 seem incorrect (per http://allelefrequencies.net/):
      → Gaza Palestinians 2.4%
      → Jordan 2.5%
      → Italy: 0.0-2.5%

      Actually per this source, Italians seem to have lower frequencies than Native peoples from the Southern Levant. The largest frequency is in North Pavia pop. 2 (2.5%), which should fall to much lower when averaged with pop. 1 (0.9%). The second largest frequency is from the historically super-cosmopolitan city of Rome (2.0%).

      Nothing to see here...

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    4. The All Italy sample (n=159,311) shows a frequency for this allele of just 0.9%!!!

      So...

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  7. Hi Maju, thanks for your reply.

    Ok, rather than the alleles I'll provide some full haplotypes.

    Sardinia
    #1 haplotype: A*30-B*18-C*05 [12.50%]
    [shared with Iberians, Basques, Balearic Islanders, Corsicans, S.French, N.Italians, Swiss, North West Africans, Senegalese, Tuaregs, N.Cameroon, Sudanese, + slightly different variant in Kenya Luo]
    #2 haplotype: A*02-B*58:01-C*07 [4.90%]
    [shared with Iberians, Italians, Swiss, North West Africans, Cameroonians, African Americans, Sudanese, Kenya Nandi, Kenya Luo]

    There are several haplotypes found in Sardinia which are shared with West Asians, however these are not.

    "That's why neutral markers (or those believed to be more neutral) are preferred when researching ancestry."

    Use of HLA as an ancestry marker is not limited to armchair geneticists or anthropologists like myself. Right now the University of Geneva's Department of Genetics & Evolution's AGP Lab [Laboratory of Anthropology, Genetics and Peopling history] is using them quite extensively in some research projects funded by the Swiss National Science Foundation:-

    "The AGP lab is involved in many research projects on human genetic evolution with the objective to understand better the dispersal of modern humans around the world and the mechanisms of their biological differentiations. Information from other disciplines, namely archaeology, paleoanthropology and linguistics, are also used to investigate the relationships between biological and cultural evolution. The team principally works in silico by developing computer tools and performing data analysis. One of the main polymorphisms used is HLA, but other genetic markers are also analyzed (mtDNA, Y chromosome, nuclear STRs, etc). Since several years, a big effort is devoted to the improvement of the HLA characterization of human populations from a methodological point of view and the maintenance of large databases." http://ua.unige.ch/fr/agp/

    "Also the frequencies you provided for B*53 seem incorrect"

    I didn't provide any frequencies for B*53! [that's the malaria protective allele].
    The frequencies I provided were only for the B*58:01 allele, I provided no data on B*53 whatsoever.

    To summarise, I can see geneflow into Sardinia and Corsica from three directions looking at HLA: Europe; N.Africa/Sahara; and West Asia. The gene flow from Africa impacts the mainland of Europe to some degree also. My issue is that a lot of people want to bury their head in the sand over this. Eurogenes blog deleted the entire 'Ancient Middle Easterners (AME)' post from 10th August, when this had been discussed in the comments section. I don't know if this is coincidental or not.

    Incidentally I regularly read your blog and would like to say keep up the good work. It is very well researched and interesting and informative. It's also refreshing to have a genetics and anthropology blogger who is from the left of the political spectrum rather than the right.

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    1. I didn't provide any frequencies for B*53! [that's the malaria protective allele].
      The frequencies I provided were only for the B*58:01 allele,


      My bad, sorry.

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    2. Hi!
      If you are interested....
      http://new-indology.blogspot.in/2014/10/can-we-finally-identify-real-cradle-of.html

      Delete
  8. I thought I'd mention I'm enjoying you destroying Squiggles on Dienekes' blog right now.

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    1. Me? Destroying what? I don't think so, unless you're talking from something of the past. I'm almost totally oblivious to the blogosphere and have been so for many months now.

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    2. Yah, it was an old post apparently. Linked to on Razib Khan's blog, I mistook it for a new one. A discussion about genetic diversity and structure in Africa.

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  9. Dear Maju,
    I need your clinical analysis on the newly developing projection that Bell Beaker culture might have its origins from northern Africa, David and Dienekes Disagree-
    http://cuni.academia.edu/JanTurek
    Happy 2015! BTW.

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    1. What paper are you specifically thinking of? I have discussed some of that in Bell Beaker Blogger's commentaries. What he argued for was that the characteristic clay found in many bell beakers, which is reddish, "must" come from North Africa. But that on its own says little about the origins of BB as such, only about where the materials were imported from. If, as archaeology seems to show nowadays, BB originated in Iberia (possibly Portugal), then importing clay from nearby Morocco or Algeria would be easy task, considering that the trade routes were already open since long earlier.

      In order to establish a North African origin of BB, we would need a more or less clear chronology of the BB phenomenon in North Africa. In fact BB is rare in this region and mostly limited to areas just across the Strait of Gibraltar in Arif country (aka North Morocco). This says little in favor of a North African origin of the phenomenon, but if you have some information I'm missing, please share.

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    2. Hi!, Turek mainly like here he suggests-
      ''it Should be borne
      in mind that the ornaments, design and in decoration of the Bell Beaker pottery techniques are
      based on decoration motives of the Saharan Late Neolithic pottery (Camps-Fabrere 1966). Finds
      of decorated ceramics from the western Moroccan burial sites of El- kiffen and Skirrat were probably the archaeotypes of the Bell Beaker Ornamentation(Case 2004; Turek, 2008; 2012a).''
      https://www.academia.edu/7174464/From_Copper_to_Bronze._Cultural_and_Social_Transformations_at_the_Turnof_the_3rd_2nd_Millennia_B.C._in_Central_Europe._Gewidmet_PhDr._V%C3%A1clav_Moucha_CSc._anl%C3%A4sslich_seines_80._Geburtstages._Herausgegeben_von_Martin_Bartelheim_Jaroslav_Pe%C5%A1ka_Jan_Turek
      There is also this paper on Bell Beaker:
      Jan Turek 2014: Social and symbolic foundations of the Beaker Phenomenon. Besse M. (ed.): Around the Petit-Chasseur Site in Sion (Valais, Switzerland) and New Approaches to the Bell Beaker Culture Proceedings of the International Conference (Sion, Switzerland – October 27th – 30th 2011), 285-293.
      Then we also have R1b in Europe and it is also there in Africa so if the actual origin is in Africa then , we can give an answer to the strange concentration of R1b in western Europe, although it is clearly a West Asian Hg: it spread in western Europe through Africa and then Iberia?, thanks to the great phenomenon of the Bell Beaker. The next step is to investigate the Afro-Asiatic elements not only in Basque and Sardinian, but also in English or Celtic languages, a topic that has a long tradition and a revival at present....

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    3. I have yet to read the paper but on R1b, the origins cannot be in Africa. Let's see:
      → K and P originated in SE Asia
      → P1 and R originated in South Asia (R1 maybe too)
      → R1b clearly originates in West Asia, with major branches expanding to Africa (part of R1b-V88 mainly), Europe (R1b-M269 and another part of R1b-V88) and North-Central Eurasia (Chuvash-Uyghur clade). There's no point in treating R1b as a single haplogroup unless you're talking of the Early Upper Paleolithic, when it almost certainly branched out.

      The African R1b seems mostly V88 (although the Nile area requires deeper study as of now). This lineage has "brothers" in West Asia/Europe, along the Northeast Mediterranean arch, peaking in Sardinia. The African specific branch(es) probably expanded to the Nile with the Early UP (~ early LSA) along with J1. These lineages later expanded inside Africa (NW Africa and Horn for J1, Central Africa mostly for R1b-V88, tightly associated with Chadic, Afroasiatic, speakers). NW Africa also has a minor amount of West European type R1b but this one clearly arrived from Europe, along mtDNA haplogroups H1, H3, H4, H7 and probably also V (surely in the Oranian/Iberomaurusian genesis, tightly related to South Iberian Gravetto-Solutrean, which is the first known UP of NW Africa, replacing Aterian). Later the Capsian (Afroasiatic, proto-Berber) expansion from the Nile rearranged the Y-DNA landscape in favor of E1b and J1 but had a weaker effect in the mtDNA pool.

      "The next step is to investigate the Afro-Asiatic elements not only in Basque and Sardinian, but also in English or Celtic languages, a topic that has a long tradition and a revival at present...."

      The only "Afroasiatic" (Semitic in fact) element in these languages (insular Celtic and possibly residually in English) is a grammatical feature that is actually widespread in West Asia, not only among Semitic languages. It may well be a fluke but, if it indicates something real, it can't so easily be attributed to Semitic (not the same as Afroasiatic, the part is not the whole) but surely can be thought of as a substrate element in (some) West Asian languages that, for whatever reason, managed to reach Far NW Europe (and nowhere else in the subcontinent).

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    4. PS- Hypothetically we could consider, as Venneman does (but with a huge deal of negative criticisms), that there was a proto-Semitic ("Semiditic") element inside the mainline European Neolithic wave that, for random reasons, only managed to establish or survive in the Western Islands (it's not a Celtic nor Germanic trait but something very specific to the Islands). This is very hard to explain unless we imagine that "Danubians" were "Semiditic" speakers and "Cardial/Megalithic" farmers were Vasconic speakers. But a simpler explanation is to consider the trait as mere fluke with too much wishful-thinking overinterpretation. Another possibility is that the trait is not specifically proto-Semitic but, as I said before, a West Asian archaic trait also preserved in other languages like Turkish, being hence also in Semitic a substrate feature of some sort.

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    5. I quote from Turek's paper:

      "The introduction of the Bell Beaker package did not cause discontinuity of the previous cultural development. Just the opposite, in many ways we can recognize a notable continuity with the previous Corded Ware Period".

      This continuity is not only apparent in Central Europe (where it explains the persistence of recently established Indoeuropean language, largely with Corded Ware) but also and even more clearly in West and SW Europe (where Vasconic languages were then probably mainstream). There may be localized exceptions but in general terms BB is a development that denotes ethno-cultural continuity everywhere, but with a cosmopolitan slant that is surely heir of Dolmenic Megalithism (which survives for long along BB West of the Rhine/North Sea line).

      "We cannot assume that the spread of Bell Beaker material culture into peripheral and neighbouring areas was also associated with the spread of the original Beaker ideology".

      Ditto.

      "... it is more than clear that the prehistoric European population, although its mobility was limited, had fairly clear ideas about the various forms of religion and material culture in different and often very remote regions of Europe and beyond".

      Turek therefore considers BB to be an "ideological", probably religious, phenomenon. This fits generally quite well with the general facts of cultural continuity with the previous ethno-cultural substrate. Some people suspect that there was an influential trader sect behind this intriguing phenomenon and I could easily subscribe that idea at least tentatively.

      What is quite clear is that, with Corded Ware and Artenac expansion c. 2400 BCE, an ethno-cultural boundary was set for around a whole millennium at the Rhine (Indoeuropeans to the East, Vasconics to the West). BB crosses this boundary but does not seem to alter the etno-cultural substrate at all. The genetic influence that some people attributes to BB must be surely reconsidered in terms of Megalithism, which had a much stronger demographic influence in most regions when we consider artifact density. However only further research of aDNA can confirm this "Megalithic model" of European demo-genesis.

      I can't find much re. Africa. Turek's paper is basically on Central Europe. What he says is:

      "The North African Bell Beaker inds outside the territory of Morocco are still very scarce (...) the center of the Bell Beaker occurrence in North Africa was along the Atlantic coast of Morocco"...

      That's about it. In brief: some of the Iberian BB spread out to Arif (North Morocco) and very rarely some of this reached out to other areas in Algeria and Tunisia. There's nothing in this suggesting a North African origin of BB.

      Arif is very easy to reach from Andalusia or vice versa. The narrowest point (Strait of Gibraltar) has a width of just 13 km and I've seen (on TV) people crossing it on a mere surf table in a matter of hours (especial permission from the authorities was required, of course, but that modern issue was not such in the Chalcolithic). Trade with Africa existed since much earlier in the Chalcolithic, as shown by at least ostrich egg objects (while elephant ivory could come to Iberia from Africa or Asia, ostrich eggshell could only come from Africa). Also NW Africa was more intensely influenced by the late Dolmenic-Megalithic expansion in the Western Mediterranean (which also affected Italy and approximates as a whole the extent of the "Atlantean empire" described by Plato, reaching to Tyrsenia and Libya).

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  10. I don't even bother to know about BB. clear to me tough is that lots of H came from paleolithic Sicily to North Africa like V and then crossed Gibraltar.
    not H17'27 which You said was Sungir and want to believe You

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    1. This would need some sort of demonstration. As of now, I know of a couple of major papers on North African mtDNA H and both suggest quite strongly that North African mtDNA H is derived from Iberian (or French in the case of H7).

      See: http://leherensuge.blogspot.com/2009/04/brief-review-of-recent-mtdna-h-info.html (lists and links all major mtDNA H papers back in the day).

      Cherni 2008 concludes that Tunisian mtDNA H pool is Iberian-derived. Enafaa 2009 has a very detailed account (table 1 and additional file 1) of H sub-haplogroups in North Africa, supporting the notion of H1, H3, H4 and H7 (basically all mtDNA H in North Africa) arriving there from SW Europe.

      This could almost only happen in the middle Upper Paleolithic (Solutrean → Oranian). I guess you can imagine a Megalithic scenario but, if so, where is the Y-DNA? The most parsimonious explanation is that it was largely wiped out by Capsian expansion in the Epipaleolithic and Early Neolithic. In any case North African H clearly comes from SW Europe, and this is plausibly also the case of V.

      Neither H1, nor H3 nor H4 have meaningful West Asian or Balcanic presence. H7 does however and this may be a case open for discussion but H7 only makes up 5% of all North African H and has a Tunisian centrality. In contrast, H1 and H4 have a clearly Moroccan centrality, while H3 is more ambiguous (but outside NW Africa it's only found commonly in SW Europe).

      Other H subclades in NW Africa only account for <2% each, having probably various origins. H* is more important (26% of all H in North Africa) and may have partial Tunisian centrality (Morocco is also important), what may allow for diverse origins (being a paragroup, it's likely it has various origins).

      Can you detail the subhaplogroups of H in Sicily? Judging on mainland Italy's data, it should be mostly H* and H1. H1 clearly is centered in NA towards Morocco, so only H* and H7 can be thought as Tunisian-centered and of these H* is the only one I can relate to Italy.

      Archaeologically speaking there's no particular relationship between Italy and NW Africa. The strait of Pantellaria is extremely wide (almost not any strait, even with Ice Age sea levels) and I have never read of any particular paleo-cultural connection between Tunisia and Sicily before the Phoenicians. So I'm very much skeptic about your claims, sorry.

      As for V, it surely deserves new studies but it seems very plausible that it has some sort of Eastern Pyrenean (Catalan) centrality. How did it end up in both Sápmi and Tunisia/Kabylia? Can't say for sure but the most parsimonious model is to make it a lesser associate of the mtDNA H of SW European centrality. In other words: it arrived to Tunisia with H7 and other H from a SW European origin most likely.

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    2. Hola! thanks for your conclusions BTW I didn't say that R1b originated in Africa But said West Asia as the Origin area:) but i will wait for the N African aDNA to see if R1b was present there from Neolithic or not and i have to more deeply investigate on the' Afro-Asiatic elements' in celtic,basque etc.
      Here a new news on N African DNA-
      http://dispatchesfromturtleisland.blogspot.in/2015/01/new-north-african-dna-data.html

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  11. Dear Maju,
    I wonder what will be your opinion on this genetic research based on 5 Core World Autosomal Components including Basque and South Indian!!
    http://www.dnatribes.com/dnatribes-digest-2014-07-02.pdf
    What do you say??

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    1. What do I say? Long live Greater Basqueland, from the Sahara to the Artic, from the Atlantic Ocean to the Arabian Sea! I'm just making the preparations to be crowned pharaoh, shah, emperor and tsar of such fundamental nation, where everybody will have to speak Basque and worship Mari and Sugaar, and then declare war against everybody else, mwahahahaha! XD

      Seriously now: it's an over-simplifying approach typical of that commercial DNA testing company. It has some pros and some cons but in any case it is oversimplifying. This is particularly true in the case of Africa, where a huge diversity, basal to that of all Humankind, is being hidden under a catch-all category.

      Another example can be the Mesoamerican reference. What would happen if instead of Mesoamerican Natives they would have picked Kets (from West Siberia), who are surely the closest living population to Mal'ta and related paleosiberian genetics? Basically the NEA component would be stronger in Europe, and possibly other places, while Native Americans would show up as a mix of NEA and East Asian.

      What happened to Australasian Aborigines also? They are definitively not any simple mix of Indian and East Asian but a distinct population (or several) on their own right.

      The problem is that when you get references you can always express anyone as product of those references. If we took Nigerians, Tanzanians and Angolans as references, we could also define anyone as a mix of those three populations, even if it's obviously a rather unrealistic exercise.

      So take with a pinch of salt. It has some pedagogic value for people with limited or no knowledge of population genetics, and I can even agree that, for this purpose, it is relatively adequate, but for a deeper, more nuanced approach it is clearly unsatisfactory.

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  12. Maju,
    What happened you stopped the Discussion In New Indology? It was going good.
    BTW here a new paper on modern Asian Y-DNA -
    http://dienekes.blogspot.in/2015/01/y-chromosome-super-fathers.html
    Do you think what Dienekes say on the STR based date that is Given on Indian R1a-Z93 to be practical? Please tell us.
    I also have a request though don't know will be valuable or not: Can you run a K=5 admixture analysis from EEHG,EFF,West Asian,ANE and Ancestral South Indian on South Asian or Asian populations? Do you think that is doable as an experimental basis Maju?....

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    1. I've kept commenting in NI, maybe the comments have not been approved yet or you are not loading the new comments ("load more") or whatever.

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    2. Balaresque is not credible. She is a hyper-recentist who does a very shallow analysis. I got very upset at her R1b paper years ago because it was ridiculously biased in all senses. I posted a somewhat angry comment at PLoS ONE back in the day, calling her approach pseudoscience and cheap (or something like that), which gained me a reprimand by the administrators - so I stopped commenting altogether (anyways the comments section has since been scrapped, I believe).

      I wouldn't believe anything Balaresque says, really. She's clearly a one sided extremist within the field.

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    3. "Can you run a K=5 admixture analysis from EEHG,EFF,West Asian,ANE and Ancestral South Indian on South Asian or Asian populations? Do you think that is doable as an experimental basis Maju?...."

      You could do that, the tools are freely available online.

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    4. Thank you!
      ''You could do that, the tools are freely available online.''
      Please you are the expert in that field:) if you can run the admixture just give us a google doc after doing that....

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    5. I'm not any "expert" and I'm actually dropping my blogging activity altogether (because I'm getting old and grumpy and tired and bored). I believe in DIY and that everyone is potentially a "leader", and in this case the tools are freely available. I explained how did I learn to in my first attempt to play with ADMIXTURE: http://forwhattheywereweare.blogspot.com/2011/12/playing-around-with-admixture.html , which redirects to Razib's relevant post: http://blogs.discovermagazine.com/gnxp/2011/03/analyzing-ancestry-with-admixture-step-by-step/

      So if you're interested, please DIY: learn, analyze and publish. I'll be glad to discuss the results surely.

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  13. Maju, I guess this work by Palisto is impressive right?-
    http://kurdishdna.blogspot.in/2015/01/balaresque-et-al-2015-r1a.html
    egun ona!

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    1. In general I consider his work quite interesting, yes. I haven't read that one though.

      Delete
  14. ==Both R1a-R1b Spread from Yamnaya To Europe aka Indo-European movement?==
    Maju what are you doing! come....
    http://eurogenes.blogspot.in/2015/02/david-reichs-oxford-seminar.html

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    1. I read what David wrote and there was nothing of interest in it: "I'm hoping that, it's likely that"... rumorology!

      The update is even more clarifying: "Jean Manco says"... Mrs. Wishful Thinking, Mrs Facts Denier says? Whatever! Don't waste my time until there's clear data please.

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    2. Actually the most interesting part is that the only R1b within the Euro-clade is from Spain. I quote Krefter: "The Iberian and German BB might have R1b. Iberian R1b might even come out P312. "

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    3. What about the South Asian Ancestry detected in K=8,K=15 on Yamnaya samples?.
      The non-Euro R1b appears to be from Maykop am i right or wrong Maju?.

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    4. Hah, came here to mention this and was beaten to the punch. R1b in early Neolithic Spain!

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    5. Also I found this interesting from the supplementary material:

      "A different detail of Fig. S8.6 is that the ANE ancestry in Karelia_HG is derived from the branch of “Ancient_North_Eurasian” that goes into the Karitiana Native Americans, rather than the MA1 branch."

      Gratifying to see this supported by data.

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    6. Looks like there will be a lot to discuss once all that it's published. Right now I feel like people know more than I do but it's not clear what part is real and what is just rumor. In any case what you guys are leaking seems within my expectations.

      "The non-Euro R1b appears to be from Maykop am i right or wrong Maju?"

      I know nothing. Awaiting publication.

      "R1b in early Neolithic Spain!"

      That's the kind of finding I felt was needed to shut up the "Indoeuropeanists". Now we can begin to discuss Megalithism, Bell Beaker and the legend of Atlantis.

      "I found this interesting from the supplementary material"...

      Is there anything published already?

      Delete

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