January 25, 2014

Andalusian mtDNA highlights W-E differences

The issue of W-E genetic differences in Iberia has been discussed before in this blog by guest author Argiedude on the grounds of Y-DNA, showing that roughly the Western third of Iberia is distinct from the rest, most notably because of its higher presence of North African lineage E1b-M81. However the differences also appear in the mtDNA, even if they may be a bit more subtle because they cross with a N-S gradient of sorts.

A new study focused on the matrilineages of two characteristic Andalusian provinces, Granada in the East and Huelva in the West, underscores that this difference is very real.

Candela L. Hernández et al., Human maternal heritage in Andalusia (Spain): its composition reveals high internal complexity and distinctive influences of mtDNA haplogroups U6 and L in the western and eastern side of region. BMC Genetics 2014. Open accessLINK [doi:10.1186/1471-2156-15-11]

Abstract (provisional)

Background

The archeology and history of the ancient Mediterranean have shown that this sea has been a permeable obstacle to human migration. Multiple cultural exchanges around the Mediterranean have taken place with presumably population admixtures. A gravitational territory of those migrations has been the Iberian Peninsula. Here we present a comprehensive analysis of the maternal gene pool, by means of control region sequencing and PCR-RFLP typing, of autochthonous Andalusians originating from the coastal provinces of Huelva and Granada, located respectively in the west and the east of the region.

Results

The mtDNA haplogroup composition of these two southern Spanish populations has revealed a wide spectrum of haplogroups from different geographical origins. The registered frequencies of Eurasian markers, together with the high incidence and diversification of African maternal lineages (15% of the total mitochondrial variability) among Huelva Andalusians when compared to its eastwards relatives of Granada and other Iberian populations, constitute relevant findings unknown up-to-date on the characteristics of mtDNA within Andalusia that testifies a female population substructure. Therefore, Andalusia must not be considered a single, unique population.

Conclusions

The maternal legacy among Andalusians reflects distinctive local histories, pointing out the role of the westernmost territory of Peninsular Spain as a noticeable recipient of multiple and diverse human migrations. The obtained results underline the necessity of further research on genetic relationships in both sides of the western Mediterranean, using carefully collected samples from autochthonous individuals. Many studies have focused on recent North African gene flow towards Iberia, yet scientific attention should be now directed to thoroughly study the introduction of European genes in northwest Africa across the sea, in order to determine its magnitude, timescale and methods, and to compare them to those terrestrial movements from eastern Africa and southwestern Asia. 

Naturally the most noteworthy data is the frequency of mtDNA haplogroups in the two sampled populations:


A map comparing this data with some other areas of Iberia (mostly the West) is also provided:

Figure 3 - mtDNA haplogroup profiles registered in some populations of the Iberian Peninsula. The two Andalusian subpopulations studied here are marked with a red arrow. Codes are as in Additional file 3.

What makes Onubenses (the inhabitants of Huelva, ancient Onuba, West Andalusia) peculiar in relation to their Granadino neighbors is their lower frequency of H (notably H*, H3 and H5), along with their higher frequencies of K1, U3a and several North African related haplogroups (U6, L1b and L2). The peculiarities of Granadinos (notably the high H5 frequency) are not so notable in comparison.

Some of these Onubense peculiarities are reproduced in other parts of West Iberia, notably the low frequencies of H (but not at all in the NW corner), the presence of U6 (notably in North Portugal and also, not shown here, among the Maragatos of the León-Galicia border area) and to some extent the elevated frequency of K and some L(xM,N) lineages (varied localized frequencies).

Much of this seems best explained by ancient flows from NW Africa (flows which may be Neolithic, Paleolithic or from the Metal Ages but hardly related to Phoenician or Muslim colonization, which had no W-E gradient whatsoever) but I have some qualms about the quick identification by the authors of the origins of Onubense high K1 in that area. At the very least it must be noticed that, unlike the other African markers, K1 is much more common towards the Eastern parts of NW Africa and is also found at similarly high frequencies in many parts of Europe, such as France and Central Europe.

K1 was first spotted in Iberian ancient DNA in the early Neolithic (Los Cascajos, Navarre), and was an important Neolithic lineage through Europe. While I can't discard the North African suggested origin, I think that other possibilities are at least as likely.

On U6, the presence of the very rare U6c, one of two basal lineages of U6, only found previously in 5 Moroccans, 10 Canarians and in one Italian, reinforces the idea of U6 expanding from West to East (against what most conclusions suggest on the most unclear grounds) with a most likely Moroccan origin. In turn this raises the question on how pre-U6 arrived to Morocco, especially as the Aurignacoid Dabban industries now seem to never have gone further West than Cyrenaica, opening the possibility of this lineage having arrived to NW Africa via Europe, where we know that U in general was common in the Upper Paleolithic, and which clearly influenced North Africa at the Oranian (aka Iberomaurusian) cultural genesis (LGM) via Morocco (Taforalt and other sites).

On the L(xM,N) lineages, it is worth mentioning that Cerezo 2012 claimed that some of them may be pre-Neolithic in Europe, especially L1b1a variants, which are widespread at low frequencies through Europe with an Iberian centrality.

The overall picture is maybe best visualized by the Hierarchical Cluster Analysis:

Figure 5 - Hierarchical Cluster Analysis (HCA) of 53 populations based on their mtDNAdiversity. The haplogroups used here are marked with arrows (vectors). Populations are indicated with numbers as in Additional file 3.

Cluster 1 is particularly noticeable for its high frequency of mtDNA H, being composed by mostly Iberian samples (along with South Germans and some Sicilians). Granada, as well as nearby Córdoba, sit here (even if with a tendency towards the more mainstream European Cluster 2). Huelva is not too remarkable when compared with other European samples in the mtDNA HCA, although it does show some deviation towards NW Africa, clustering closest to Canarians.

In spite of the good frequency of North African samples, it must be noted that the horizontal axis, in essence contrasting Europe vs. West Asia, has a weight of almost 50%, while the vertical axis, contrasting these two vs. North Africans (and the Sámi) only weights 13% (axis are almost never of the same relevance in PCA-like graphs, even if they are presented as such).

In synthesis: NW Africa had some minor but significant genetic influence in the West Iberian third, long before the Muslim period, and this mtDNA study ratifies these distinctions in the particular case of Andalusia, adding some rich detail of data.

Also, as the authors underline in their discussion, the issue of European genetic influence in North Africa still requires some serious investigation.

26 comments:

  1. Any information about from where the Granada samples come from? I mean if it´s from a coastal location, or not?

    Thanks!

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    1. Granada province does not have much coast, you know. Anyhow the locations of the samples are in fig. 1, listed in the legend as: 1: El Repilado, 2: Aracena, 3: El Cerro del Andévalo, 4: La Puebla de Guzmán, 5:Valverde del Camino, 6: Villablanca, 7: Niebla, 8: Huéscar, 9: Baza, 10: Montefrío, 11: Loja, 12: Alhama de Granada, 13: Órgiva. Nos. 1-7 are from Huelva and nos. 8-13 from Granada, only one of which is from the coast (13: Órgiva). In Huelva also the samples are from the interior in essence.

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    2. Coastal locations were avoided due to their intense population flux and resulting high degree of admixture (which goes against our aim of using autochthonous individuals). Órgiva is located in the Alpujarras, on the southern slope of Sierra Nevada mountain range, and technically can't be considered as coastal (although the sea is not too far away in a straight line). Hope it helped, and thanks a lot for reading.

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    3. Thanks to you for passing by and commenting, Guillermo. I must say that your study impressed me for its refreshing insight. I have read and sometimes discussed many other studies on Iberian genetics before and often simplistic "historical" explanations or lack of depth in the analysis have been disappointing. So I was typically forced to take the data and ignore much of the discussion. This is not at all your case: you are "joining the dots" very cleverly (and your data is also very informative, of course).

      So thanks to you and your colleagues. Keep up the good work.

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  2. What do you think about the presence of the Y-DNA E1b1b1, in Iberia?

    It´s due to the Moor influence?

    I don´t think that Moors left a significant genetic impact in Iberia, but I would like to know your opinion, on this.

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    1. Not the Moors. And the reason is that the Y-DNA pool of NW Africa is in essence E1b-M81 + J1, and we see almost no J1 in Iberia (mostly in East Andalusia, not much anyhow), instead E1b-M81 is rather common in places like Galicia, Asturias or Cantabria where the Muslim presence was effectively zero, it is as common in those areas as in Southern Portugal or Western Andalusia, if not more. The only cline we see is W-E: the Western third has it, the other two thirds of the peninsula do not (or at very low frequencies). The same happens with mtDNA lineages typical from North Africa like U6 or more rarely L(xM,N).

      In essence, in the Western third of the peninsula there is less than 10% of North African ancestry but is not from the Muslim period but necessarily older. How old? I have two alternative theories:

      1. It is a backflow from the Solutrean migration to NW Africa, triggering the Iberomaurusian or Oranian culture. That could explain its presence as far North as Asturias and Cantabria because the Solutrean facies of Asturias is related to Portugal rather than Aquitaine and the subsequent Magdalenian period has a Cantabrian-Asturian facies as opposed to another Basque facies (so we can imagine the Paleo-Asturians migrating to Cantabria in that period).

      2. If we don't like "paleo-theories", then we can always resort to Neolithic founder effects. I find a bit more difficult to explain the impact in Cantabria-Asturias with this model though but the information available is blurry enough to allow for it, I guess.

      I don't think there are later windows for this phenomenon but who knows?

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    2. PS- Anyhow there is also a less important presence of E1b-V13, whose origins are almost without doubt Neolithic. This lineage came via Greece, not directly from NW Africa, where it almost doesn't exist (instead it's very common in Greece, Albania, Kosovo, and the Asian coasts of the Eastern Mediterranean - it has also been detected in a Cardium Pottery early farmer from Catalonia).

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  3. Thanks for comment.

    U6: I have read somewhere that it was found in some Iberian hunter-gatherers and it has a diversity peak in Iberia.

    L: It´s not common in Iberia, and according to Cerezo et al, L1b is seemingly the most frequent one in Iberia (it is actually a «European specific subclade») and may have entered in this peninsula during the Epipaleolithic period.

    So maybe it´s safe to presume that there was some connection between both sides of Gibraltar and that Iberian hunters-gatherers should have had an impact in Northwest Africa.

    However, it seems also probable that Iberia and Northwest Africa, have had also some Neolithic contact, like it´s apparent in Huelva. Though this substrate probably has also some shared pre-neolithic ancestry.

    Interestingly William. W. Howells craniometric data set, presents some Muge skulls as having some skull close affinities with Afalou and Teviec. Afalou individuals are said to have survived the Neolithic transition period as the Muge ones. I don´t know about the Teviec individuals, though.

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    1. "U6: I have read somewhere that it was found in some Iberian hunter-gatherers and it has a diversity peak in Iberia."

      You may have read it to "old me", either in my old blog or in comments in some other blogs (sounds like me years ago). However the reality is that:

      1. The U(xU5) found in Portugal does not look at all like U6 - but the data lacks enough good quality to confirm.

      2. U6 seems to reach the peak of its basal diversity in Morocco and only afterwards in Iberia. It's a matter of a small lineage known as U6c but, together with its greater (and quite diverse) distribution in North Africa, including Canary Islands (where the sub-lineages are related to Morocco, not Iberia), the scales lean to NW Africa quite clearly. Another question is how did U6 reach Morocco from an ultimate U origin in West Asia (via NE Africa or maybe via Europe?) for which we lack clear answers, but the founder effect centered in Morocco is very apparent in any case, as it is its association in geography with Y-DNA E1b-M81 (also centered in Morocco) in the West Iberian case, both associated to smaller mtDNA L(xM,N) lineages of necessary African origin.

      "L: It´s not common in Iberia, and according to Cerezo et al, L1b is seemingly the most frequent one in Iberia (it is actually a «European specific subclade») and may have entered in this peninsula during the Epipaleolithic period".

      Yes (I'd rather think Solutrean period but whatever, it could be Neolithic too). But the case is that its distribution is similar to that of U6 and Y-DNE E1b-M81. There's also some L3 for what I know, again in the same Western Third area of concentration. So the three groups of lineages clearly imply the same migratory process, the same founder effect from almost certainly modern Morocco.

      "So maybe it´s safe to presume that there was some connection between both sides of Gibraltar and that Iberian hunters-gatherers should have had an impact in Northwest Africa".

      Absolutely the mtDNA lineages of SW European origin in NW Africa make up a 30-35% of the pool, what is very large. Instead Y-DNA lineages are residual (although I was surprisingly important in the Guanche mummies and I recently read that Moroccan Berbers, not so much Arabs, harbor relatively high frequencies of R1b). This seems to imply that the Oranian (aka Iberomaurusian) culture had important partial roots in Southern Iberia but that the later Capsian one (ultimately original from Upper Egypt/Nubia) wiped much of their legacy particularly on the Y-DNA side. Nevertheless the role of E1b-M81, seemingly too old and specific of NW Africa, is not clear enough. I'd even dare to adventure that Oranian was spread by mostly E-M81 guys with mtDNA H girls, what is a more complex founder effect than a mere migration from Iberia.

      ...

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    2. ...

      "Interestingly William. W. Howells craniometric data set, presents some Muge skulls as having some skull close affinities with Afalou and Teviec. Afalou individuals are said to have survived the Neolithic transition period as the Muge ones. I don´t know about the Teviec individuals, though".

      About this I hope that we will we get more useful archaeo-genetic information soon (see: https://sites.google.com/site/pinhasierc/home/samples). Teviec is AFAIK Megalithic from Brittany but the mtDNA sequence was deemed "melánge" (mixed and hence useless for identification) by Lacan 2011. Some have argued it could be HV(xH, HV0), what is not too informative either.

      But it is indeed possible IMO that there was a stronger survival of Paleolithic people and genetics in the Atlantic area, including Portugal, parts of the Basque Country and maybe also West France. Their ancestry would anyhow, for all we know, get mixed with the newcomer Neolithic one, this one dominantly, and a more "westernized" variant of the Neolithic peoples (say Basque vs. Sardinians, approximatively) became very important in the Chalcolithic (Megalithism, possibly also Bell Beaker), redefining to some extent the European genetic pool towards Western HG typologies. But the exact extent and nature of this "wooble" still awaits clarification.

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  4. Thanks a lot for your input!
    I have read about a L3 that diverged from the African branch, at least, since the Mesolithic period, found in Cantabria region (L3f1b6).
    So this makes sense, when put along with the Epipaleolithic entry of L1b in Iberia.

    Your new interpretation about the U6 presence in Iberia is also very interesting. Was it really found in some iberian hunter-gatherer remains? I find it odd to imagine entering in Iberia, only on the Neolithic period. The same for E1b.

    There are some authors, like João Zilhão (whom I actually find controversial sometimes), that theorize that some predominantly hunter-gatherer communities in Iberia, adopted caprid farming so that this was (regionally) mainly a cultural shift. I assume that some authors believe that hunters may have traded with new comers and survived well longer that originally thought. I find that this may be true, mostly because it was supported with skeletal evidence. We also already know about the already proven horse domestication event in Iberia (very likely followed later by the local aurochs domestication as well). Seemingly the first event led to the second event, according to some authors. And having these animals domesticated, Ancient Iberians, would have great advantage on their side. Several Mediterranean ancient cultures were known to acquire great amounts of horses coming from the Iberian Peninsula. Seemingly cattle was negotiated too.
    Well, either way, and going earlier in time again, I think that the isolation of these hunter communities didn´t lasted forever and other elements, arrived later and contributed to the local gene pool. What I think that´s relevant here, is that may have happened more recently than we usually assume. For instance, some hunter-gatherer fossil affinities (more specifically Combe Capelle, Predmost, Muge, etc...) with some modern iberians, have been established by different authors. So the blend between hunters and farmers, was probably never completed in Iberia and you can see stereotypical examples of both.

    More to come.

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    1. What I don't see is any archaeological evidence that suggests an Epipaleolithic exchange: it must be older (middle Upper Paleolithic) or more recent (Neolithic). Considering that "molecular clock" guesstimates usually seem quite more recent than they should (because of scholastic inertia or "academic conservatism"), I'd say it's something from the Solutrean period.

      I have also argued my case in favor of a Solutrean-Magdalenian impact of "Africanized" West Iberian genetics in Asturias (Solutrean) and Cantabria (Magdalenian), because that's the structure of archaeological facies in the Cantabrian Strip: (1) the Asturian Solutrean is related to that of Portugal and Salamanca (and not directly to Aquitaine) and (2) the Cantabrian Magdalenian forms a unified Western facies with the Asturian one in contrast with the Basque Magdalenian. As the Basque Country (and Old Castile too) lack the African genetic signature, this seems a quite plausible scenario.

      As for the rest, I agree with some details (local horse domestication) and disagree with others (local aurochs domestication) based on genetic evidence mostly. Aurochs were in essence domesticated in Asia Minor and there's nothing particularly "wild" in breeds like the toro de lidia. There was surely some minor European auroch admixture but it did not likely took place in Iberia but further East (Starcevo culture surely, also some indications in Italian cattle).

      On the other hand olives were quite possibly first "domesticated" in Iberia, which is still today the first global producer.

      About the anthropometric typologies, I distrust them because individual variability is vast and misleading. I'd rather see good autosomal genetic studies, which are in SW Europe sadly absent.

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    2. I have found this in your blog:

      «I think that at the least in Italy and Iberia, local aurochs did had a role on the formation of some local breeds.

      For example, in Portugal, and regarding Y-DNA, a study found that native cattle that was sampled contained 7 haplotypes (H2Y1, H3Y1, H5Y1, H7Y2, H8Y2, H10Y2, and H12Y2) found only in these breeds and not in any Middle Eastern or North African domestic cattle.

      «The presence in Portuguese breeds of Y1 patrilines, also found in aurochs, could represent more ancient local haplotypes.»


      I got this Y-DNA information in «Y Chromosome Haplotype Analysis in Portuguese Cattle Breeds Using SNPs and STRs».

      Regarding the status of Y1, is now supported that it´s indeed related with the European aurochs. So the endemic haplotype cases, present in some Portuguese breeds, very likely represent the introgression of European bull male aurochs.
      The endemic Y2 cases aren´t also out of equation (because iberian aurochs also had this haplogroup), but it are still being evaluated.
      Another interesting coincidence, is that there´s a Portuguese breed (Arouquesa) that combines an endemic patrilinear Y2 with the very rare matrilinear Q (which is atrongly suggested on this new study and in previous studies, to come from the aurochs cow).
      So both aurochs bull and cow, very likely, were involved on the formation of some Portuguese breeds.



      The study is here:

      pendientedemigracion.ucm.es/info/prodanim/html/JP_Web_archivos/64.pdf

      I just merely did comparisons, by using the current knowledge about the local cattle genetics and the results of this new study. Actually it´s surprising that the authors
      aren´t aware of this matches, but it´s normal, because Portugal usually is a less well studied country.

      Seemingly wild horses were also included in domestic stock breeds in Iberia:

      http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0018194


      The taurine matrilinear T3 situation is still being evaluated (because it´s also present in some local breeds), to differentiate the lineages until a deeper level, because it was found in both South European aurochs (Italy and Iberian peninsula, at least) and domesticated cattle from the Middle East.

      It seems clear anyway, that both Middle Eastern domestic cattle/Middle Eastern aurochs and South European aurochs, weren´t very distant genetically speaking.

      The North African aurochs, in my opinion, is a «recent» derivation from the Middle Eastern aurochs, on which the Sinai was used as stepping stone, during more wet periods.
      It´s possible that some North African cattle was exported to some European countries, but their influence doesn´t seem to be significative overall. At least and until now, this may sound realistic, I think.»

      I would add that the long thought to be North African mtDNA T1 was found to have far more diversity in Iberian Peninsula that in North Africa and that there are only very few lineages shared. Anyway, I wouldn´t go after this T1, specifically. The post above sums it up.

      PS1: Iberian Fighting cattle do have surely something ancient about them (like unique Y1 haplotypes, that very probably were acquired with the European bull aurochs influence).

      PS2: Iberians must have domesticated more plants and animals (partially the Iberian Black Pig breed, at least), but I would need more time, to expand that subject. Your interesting olive tree domestication hypothesis is based on what?

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    3. You wrote: «I have also argued my case in favor of a Solutrean-Magdalenian impact of "Africanized" West Iberian genetics in Asturias (Solutrean) and Cantabria (Magdalenian), because that's the structure of archaeological facies in the Cantabrian Strip: (1) the Asturian Solutrean is related to that of Portugal and Salamanca (and not directly to Aquitaine) and (2) the Cantabrian Magdalenian forms a unified Western facies with the Asturian one in contrast with the Basque Magdalenian. As the Basque Country (and Old Castile too) lack the African genetic signature, this seems a quite plausible scenario.»

      Ok, and I definitely agree with that.

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    4. Fair enough: you are confronting my forgotten opinion (based on something, it seems) against my current opinion (based only in my ability to forget, it seems), so I have to concede on the issue of aurochs, I guess.

      It also seems that you have done your homework... unlike my lazy self.

      I had more clear the issue of horses because mtDNA strongly suggests a second domestication center in Northern Iberia and archaeology shows many sites in the South consuming large amounts of horse meat in the Early Chalcolithic, the putative period of steppe domestication. Search for "horse genetics" and "bovine genetic" labels because there are some interesting materials in this blog, just not my main focus.

      Re. pig, it's possible. But archaeologists can't often discern what is a pig and what a wild boar because in the past all pigs were like black Iberian more or less, not so different from wild boars. So it's also possible that it's just a remnant of a more primitive type of pigs, rather than wild boar admixed ones. Only detailed genetic analysis can help here (and I know of none).

      Re. olives. Are you minimally familiar with the issue of La Almagra pottery and possible early Neolithic in South Iberia? In quick synthesis: according to some Spanish scholars this pottery (found in Andalusia and Southern Portugal) is pre-Cardial (by a few centuries only) and arrived from an unknown source with the full agrarian package (cereals, lentils, etc.) but without domestic animals (only rabbit was found typically but unclear if domestic or wild), also very large amounts of olive seeds were found. In recent times Portuguese scholars, notably Zilhao, have questioned this and argued for an epi-Cardial nature and chronology of La Almagra pottery at least in Portugal, but many Spanish researchers seem to stick to C14 apparent older dates. I don't know the full detail of the debate and its current state but in any case La Almagra pottery is early Neolithic and had lots of olive seeds, so maybe the olive tree was "discovered" in Southern Iberia. I know of no earlier date for olive consumption and hence semi-domestication of the tree, which is anyhow a tree, and a very slow growing one on top of that, and hence does not seem to allow for much "domestication" properly speaking, just some generic gardening, largely trans-generational planning of orchards and traditions on collection of the fruits and their consumption and processing.

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    5. Regarding the Iberian Black pig origin:

      http://www.ncbi.nlm.nih.gov/pubmed/22221023


      That olive tree domestication hypothesis sounds reasonable to me.
      I wasn´t familiar with it and thanks a lot for the insight.

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    6. It's a PPV study but, from the abstract alone, I don't understand how internal pig genetic distance (low in all cases) relates to local incorporation of wild boar. I also do not understand the assumption about Iberian pigs being imported from Central Europe: why? Iberian pigs should have arrived like all or most of the Neolithic package by boat from Greece, via Italy and Occitania. That's the Cardium Pottery Neolithic paradigm as I understand it: there's no relation with Central Europe initially except for some overlap at the Rhine area (assuming La Hoguette is Epicardial and not something else) and a common origin in Greece.

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  5. Guanches were also said to be composed by early farmers, hunters and an intermediate type, skeletally speaking. I´m still very skeptical regarding a R1b connection with the Epipaleolithic period, when thinking on entry dates in either Iberia or Northwest Africa. I would love to find otherwise though. But I think that there are probably other ill studied Y-DNA haplogroups in Iberia, that maybe indeed from this period. R1b may have spread in more recent times (still pre-roman), coming from the Caucasus (skeletons were proto-europid) and may have absorved more genes, before arriving at Iberia.

    Moors curiously weren´t also radically different subracially speaking when compared with some Guanches (said to be skeletally of low stature, Caucasoid and either Mediterranean or Cromagnon). I don´t think that Mediterranean necessarily excludes Cromagnon, because some hunter-gatherers found at Dolni Vestonice were actually Gracile Mediterraneans and even in Muge, Palestinian and Afalou assemblages, you can find similar examples. Maybe what the author means with Mediterranean here, it´s about a farmer type coming from West Asian or something also these lines (different from the West Mediterranean element and/or Epipaleolithic type). But modern North Africans, are different from Moors, with the notable exception of Berbers.

    And yes, I find the Megalithic culture a clear signal of local ancient input, in both culture and genetics. An Westernized variant of the European Neolithic, with a strong ancient local input..
    Its expansion contour maybe it´s not new though: an expansion from the iberian refugee may have taken place, right after the Ice age. And who knows what have happened in earlier cold phases. And there are also at least one shared human phenotype that unite with great accuracy what are thought to be some Paleolithic survivor remnants: from some locations in Norway, Sweden, Wales, Dordgone, North Iberia and Sardinia, there are groups of people with the very long, very high and narrow heads, strong brows, big cheek bones and rather broad noses of certain Upper Paleolithic skulls. their likeness to early skulls (especially Combe Capelle and Predmost skulls) supports the view that they are survivals.(I have «copy and paste» the features description, so sum it up faster).
    There are also some haplogroup connections between these different populations.

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    1. "I´m still very skeptical regarding a R1b connection with the Epipaleolithic period, when thinking on entry dates in either Iberia or Northwest Africa".

      Only more (quality) ancient DNA data can clarify this point. I see no reason to reject even a Gravettian or Aurignacian arrival personally: "molecular clock" is pseudoscience and is dramatically hindered by systematic errors which authors copy from each other in astonishing scholastic inertia, more proper of bureaucracies than of dynamic scientific institutions.

      Height is a highly variable aspect and is much more affected by nutrition and health, especially in early age, than by genetics. Young Galicians are the highest of Spain but old Galicians are the shortest ones, go figure! Other anthropometric aspects are surely also variable, at least to some extent. Epigenetics, nurture, does matter.

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    2. Ok, fair enough, at least you made me think that there´s still a open possibility that R1b might have entered before.

      I agree, regarding the nutrition and health subjects.

      One note about the Upper Paleolithic type described above that I failed to clarify: regarding Scandinavia it was found in inland Norway and mid-north-Sweden. And regarding Wales it was found in Plinlymon to be more specific.

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  6. I think that many people have been over-generalizing too much on that WHG element and in La Brana, Motala, Loschbourg, etc... Actually these are «recent» hunters. For one reason I do understand it: there is complete DNA data from them. But I don´t think that this would lead us to the answers that some want about the Epipaleolithic period, like who´s related with the hunter-gatherers, the diversity of human phenotypes back then, etc... Looking at the local hunter-gatherer fossil diversity and the different haplogroups that come out of it, I think that our view about what would be a true WHG definition, it still very far from complete. A Russian (Kostenki) hunter-gatherer with about 36.000 years old, did demonstrated that what have been said to be «farmer» genes until recently, occurred already back then. The so-called Basal Eurasian element, appeared. And this is maybe just the tip of the iceberg. I wonder where the same authors would place European hunter-gatherers with mtDNA like L3, L1b, or H (like that fantastic finding that you posted recently). Not even speaking about skull, left femora or dental trait affinities and so on.

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    1. They are "recent" HGs but (excepting maybe Motala) of Magdalenian roots. I agree that they are probably not the most representative possible samples (those should come from SW France, etc.) but they are what we have and they do show some patterns that we cannot just ignore, even if I am somewhat skeptical about the generality of all their markers, particularly the Y-DNA (too few samples so far).

      Kostenki does not have "farmer genes" that's an anachronistic interpretation: what happens is that he is old enough (Gravettian) to be relatively undifferentiated relative to post-LGM populations, let alone Holocene ones. So it may look like he "has" this or those ancestry when looked at the matter shallowly but he only has a distributed affinity (he's much older than the compared populations) because he was still too close to the common ancestors of those various populations (early West Eurasian HGs). Much of the same problem happens when looking at Mal'ta boy and the so-called "ANE" component: a great deal of the apparent affinity may not be because of paleo-Siberian ancestry but because of the very fact that the LGM paleo-Siberian population Ma1 represents was still not sufficiently differentiated relative to other populations of the same origin. Drift takes time to consolidate and "populations" are mostly made up of that unique drift specific to them. Of course smaller more inbred populations show stronger differences (examples: Druzes, Finns, Kalash, most Jewish populations, etc., or the Hadza in an African context - still very diverse for an Eurasian context though).

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    2. Well, yes, I still agree that those «recent» HGs are important (you can see that written on my post). It´s just that could be hidding other HG horizons, mostly when some articles appear with to label/overgeneralize them as West Eurasian HGs. I would be really careful with this generalizations and the conclusions that could be taken from it (like excluding other possible HGs groups).

      I know what you mean about Kostenki and I´m sorry if I wasn´t more specific. When I meant that he had «farmer» genes that have been (wrongly) excluded as pre-neolithic, I mean just that, not that he had only farmer genes (and not all farmer genes are pre-neolithic actually). He´s clearly from a period when some the main ancestral components that gave origin to many of the modern Europeans (like the so-called «proto» farmer genes), weren´t separated yet.

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    3. You wrote: «I have also argued my case in favor of a Solutrean-Magdalenian impact of "Africanized" West Iberian genetics in Asturias (Solutrean) and Cantabria (Magdalenian), because that's the structure of archaeological facies in the Cantabrian Strip: (1) the Asturian Solutrean is related to that of Portugal and Salamanca (and not directly to Aquitaine) and (2) the Cantabrian Magdalenian forms a unified Western facies with the Asturian one in contrast with the Basque Magdalenian. As the Basque Country (and Old Castile too) lack the African genetic signature, this seems a quite plausible scenario.»

      Maybe we could designate this West Iberian HG component as Solutrean? The «african» component maybe is Ibero-Maurusian instead (I think that Ibero-Maurusian denomination is more accurate than Oranian). And I wonder if in all Upper Paleolithic fossil assemblages of Asturias/Cantabria/Portugal/Salamanca we see this influence (or in most of it). Then this could lead us to a pattern (maybe a careful comparison between their cultural items would help as well).

      So those specific L mtDNA lineages may have entered in Iberia even before than I thought (that´s the more plausive scenario to me)...



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    4. "He´s clearly from a period when some the main ancestral components that gave origin to many of the modern Europeans (like the so-called «proto» farmer genes), weren´t separated yet."

      I'd use the term "defined" rather than "separated". Otherwise yes.

      "Maybe we could designate this West Iberian HG component as Solutrean?"

      Well, on one side my Solutrean-Iberomaurusian theory is not yet strongly confirmed and a later origin (Neolithic) can also be argued for, as Cardial seems to "bounce" in Arif (North Morocco) before heading to SW Iberia. Also "Solutrean" "senso strictissimo" only affected two sites (near Valencia) in the non-Cantabria Iberia, all the rest should be considered Gravetto-Solutrean, beginning in the SE and then heading to the West (with likely impact in NW Africa: Iberomaurusian genesis) and finally to Asturias. It's a very peculiar "hybrid" type of Solutrean that of most of Iberia, one that seems to feed more on the previous Gravettian layer in fact, and later maybe also incorporates ideas from NW Africa such as tanged points (Aterian conceptual influence?)

      As for the name of the culture, personally I'd call it Taforaltian... but I have to go with the historical canons and use what people understand and can check in the literature. Iberomaurusian was rejected because it had implications about its origin that once upon a time were questioned. Today it seems clear not just that relation but also its NW origin within North Africa (older dates) but for several decades this was challenged in favor of a NE African origin instead, hence Oranian, as many sites exist around this city and contrasts with Capsian, named also for another town (not site): Gafsa (ancient Capsa) in Tunisia. IMO the North African nomenclature is a bit crazy but it's the one being used, so whatever.

      "The «african» component maybe is Ibero-Maurusian instead"...

      In my theory, the African element should be pre-IM, Aterian or something that we don't know, for example an undocumented extension West of the Dabban Industry of Cyrenaica (Aurignacoid). I'd personally doubt it's strictly Aterian because at least (mtDNA) U6 is a backflow from Eurasia and even (Y-DNA) E1b-M81 looks like it should be a bit more recent than the OoA chronology (c. 125-100 Ka BP) that Aterian has. Also in my autosomal analysis I could only detect a very tiny "Aterian" component (?), mostly concentrated in Southern Morocco, what doesn't fit well with E1b-M81 prevalence. So I'd suspect a "Neo-Aterian" thing with some undefined Aurignacoid influence from Cyrenaica or Egypt but archaeology so far has not detected anything technically UP before Oranian that I know of in all the NW African region.

      "And I wonder if in all Upper Paleolithic fossil assemblages of Asturias/Cantabria/Portugal/Salamanca we see this influence (or in most of it)".

      Not in any obvious way that I know of but there's not recent explicit comparisons. What I have pondered is if the appearance of tanged and winged points in Southern Iberian Solutrean in general (including and typically the Levante groups) is a conceptual import from Aterian, after "Solutreanization" (i.e. much more refined). AFAIK there are no tanged nor winged points in all Europe before that but I don't know every detail.

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  7. Errata: Read «Dordogne» instead of «Dordgone», read «to sum it up faster» instead of «so sum it up faster». Maybe there are some more typos, but these ones are now evident to me.

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