December 26, 2013

Ancient European DNA and some debatable conclusions

There is a rather interesting paper still in preparation available online and causing some debate.

Iosif Lazaridis, Nick Patterson, Alissa Mittnik, et al., Ancient human genomes suggest three ancestral populations for present-day Europeans. BioArxiv 2013 (preprint). Freely accessibleLINK [doi:10.1101/001552]

Abstract

Analysis of ancient DNA can reveal historical events that are difficult to discern through study of present-day individuals. To investigate European population history around the time of the agricultural transition, we sequenced complete genomes from a ~7,500 year old early farmer from the Linearbandkeramik (LBK) culture from Stuttgart in Germany and an ~8,000 year old hunter-gatherer from the Loschbour rock shelter in Luxembourg. We also generated data from seven ~8,000 year old hunter-gatherers from Motala in Sweden. We compared these genomes and published ancient DNA to new data from 2,196 samples from 185 diverse populations to show that at least three ancestral groups contributed to present-day Europeans. The first are Ancient North Eurasians (ANE), who are more closely related to Upper Paleolithic Siberians than to any present-day population. The second are West European Hunter-Gatherers (WHG), related to the Loschbour individual, who contributed to all Europeans but not to Near Easterners. The third are Early European Farmers (EEF), related to the Stuttgart individual, who were mainly of Near Eastern origin but also harbored WHG-related ancestry. We model the deep relationships of these populations and show that about ~44% of the ancestry of EEF derived from a basal Eurasian lineage that split prior to the separation of other non-Africans.


Haploid DNA

The Lochsbour skull.
The prominent browridge
is very unusual for
Paleolithic Europeans.
The new European hunter-gatherer samples carried all Y-DNA I and mtDNA U5a and U2e.

More specifically, the hunter-gatherer mtDNA lineages are:
  • Lochsbour (Luxembourg): U5b1a
  • Motala (Sweden):
    • Motala 1 & 3: U5b1a
    • Motala 2 & 12: U2e1
    • Motala 4 & 6: U5a2d
    • Motala 9: U5a2
Additionally the Stuttgart Linear Pottery farmer (female) carried the mtDNA lineage T2c1d1.

The Y-DNA lineages are:
  • Lochsbour: I2a1b*(xI2a1b1, I2a1b2, I2a1b3)
  • Motala 2: I*(xI1, I2a2,I2a1b3)
  • Motala 3: I2*(xI2a1a, I2a2, I2b)
  • Motala 6: uncertain (L55+ would make it Q1a2a but L232- forces it out of Q1)
  • Motala 9: I*(xI1)
  • Motala 12: I2a1b*(xI2a1b1, I2a1b3)
These are with certainty the oldest Y-DNA sequences of Europe so far and the fact that all them fall within haplogroup I(xI1) supports the notion of this lineage being once common in the subcontinent, at least in some areas. Today I2 is most common in Sardinia, the NW Balcans (Croatia, Bosnia, Montenegro), North Germany and areas around Moldavia.

I2a1b (which may well be all them) is currently found (often in large frequencies) in the Balcans and Eastern Europe with some presence also in the eastern areas of Central Europe. It's relative I2a1a is most common in Sardinia with some presence in SW Europe, especially around the Pyrenees. I2a1 (probably I2a1a but not tested for the relevant SNPs) was also found, together with G2a, in a Chalcolithic population of the Treilles group (Languedoc) and seems to be somehow associated to Cardium Pottery Neolithic.

If you want my opinion, I'd think that I2a before Neolithic was dominant, like mtDNA U5 (and satellites U4 and U2e), in much of Central and Eastern Europe but probably not in SW Europe, where mtDNA U5 seems not so much hyper-dominant either, being instead quite secondary to haplogroup H (at least in Western Iberia). But we'll have to wait until geneticists manage to sequence Y-DNA in several SW European Paleolithic remains to be sure.



Autosomal DNA and derived speculations

Most of the study (incl. the must-read supplemental materials) deals however with the autosomal DNA of these and other hunter-gatherers, as well as of some Neolithic farmers from Central Europe and Italy (Ötzi) and their comparison with modern Europeans. 

To begin with, they generated a PCA plot of West Eurasians (with way too many pointless Bedouins and Jews, it must be said) and projected the ancient Europeans, as well as a whole bunch of Circum-Pacific peoples on it:


The result is a bit weird because, as you can see, the East Asians, Native Americans and Melanesians appear to fall way too close to the peoples of the Caucasus and Anatolia. This seems to be a distorting effect of the "projection" method, which forces the projected samples to align relative to a set of already defined parameters, in this case the West Eurasian (modern) PCA. 

So the projection basically formulates the question: if East Asians, etc. must be forcibly to be defined in West Eurasian (WEA) terms, what would they be? And then answers it as follows: Caucasian/Anatolian/Iranian peoples more or less (whatever the hidden reasons, which are not too clear).

Similarly, it is possible (but uncertain) that the ancient European and Siberian sequences show some of this kind of distortion. However I have found experimentally that the PCA's dimension 1 (but not the dimension 2, which corresponds largely to the Asian-specific distinctions) still correlates quite well with the results of other formal tests that the authors develop in the study and is therefore a valuable tool for visualization.

But this later. By the moment the PCA is asking and answering three or four questions by projecting ancient European and Siberian samples in the West Eurasian plot:
  • If ancient Siberians are forced to be defined in modern WEA terms, what would they be? Answer: roughly Mordvins (Afontova Gora 2) or intermediate between these and North Caucasus peoples (Mal'ta 1).
  • If ancient Scandinavian hunter-gatherers are forced in modern WEA terms, what would they be? Answer: extreme but closest (Skoglund) to Northern European peoples like Icelanders or Lithuanians.
  • If ancient Western European hunter-gatherers are forced in modern WEA terms, what would they be? Answer: extreme too but closest (La Braña 2) to SW European peoples like Basques and Southern French.
  • If ancient Neolithic/Chalcolithic farmers from around the Alps and Sweden are forced in modern WEA terms, what would they be? Answer: Canarians (next close: Sardinians, then Spaniards).

Whatever the case, there seems to be quite a bit of autosomal diversity among ancient Western hunter-gatherers, at the very least when compared with modern peoples. This makes some good sense because Europe was a big place already in Paleolithic times and must have harbored some notable diversity. Diversity that we may well find to grasp if we only sample people from the same areas once and again.

On the other hand, they seem to cluster in the same extreme periphery of the European cluster, opposed to the position of West Asians, and therefore suggesting that there has been some West Asian genetic flow into Europe since then (something we all assume, of course). 

Using Lochsbour as proxy for the WHG (Western hunter-gatherer) component, Mal'ta 1 as proxy for the ANE (ancient north Eurasian) one and Stuttgart as proxy for the EEF (early European farmer) one, they produce the following graph (to which I added an important note in gray):


The note in gray is mine: highlighting the contradictory position where the other Western hunter-gatherers may fall in because of assuming Lochsbour as valid proxy, when it is clearly very extreme. This was not tested in the study so it is inferred from the PC1, which seems to best approach the results of their formal tests in the WHG vs EEF axis, as well as those of the WHG vs Near East comparisons.

I tried to figure out how these formal tests are reflected, if at all in the PCA, mostly because the PCA is a much easier tool for comprehension, being so visual. Eventually I found that the dimension 1 (horizontal axis) is very close to the genetic distances measured by the formal tests (excepted those for the ANE component, obviously), allowing a visualization of some of the possible problems caused by their use of Lochsbour as only reference, without any control. Let's see it:

The same PCA as above with a few annotations in magenta and green

While not exactly, the slashed vertical magenta line (median in the dimension 1 between Lochsbour and Stuttgart) approximates quite well the WHG vs EEF values measured in the formal tests. Similarly, the slashed green axis (median in PC1 between Lochsbour and an good looking Bedouin) approximates to a great extent the less precise results of the formal tests the authors applied to guesstimate the West Asian and WHG ancestry of EEFs, which ranged between 60% and almost 100% West Asian (my line is much closer to the 60% value, which seems more reasonable). 

When I tried to find an alternative median WHG/West Asian line, using Braña 2 and the first non-Euro-drifted Turk I could spot (Anatolia is much more likely to be the direct source of West Asian ancestry in Europe than Bedouins), I got exactly the same result, so no need to plot any second option (two wrongs sometimes do make one right, it seems). But when I did the same with La Braña 2 and Stuttgart I got a genuine good-looking alternative median line, which is the slash-and-dot magenta axis.

This alternative line is probably a much more reasonable 50% WHG-EEF approximation in fact and goes right through Spain, what makes good sense for all I know.

Of course the ideal solution would be that someone performed good formal tests, similar to those done in the study, with Braña 2 and/or Skoglund, which should be more similar to the actual WHG ancestry of modern Europeans than the extremely divergent Lochsbour sequence. An obvious problem is that La Braña produced only very poor sequences but, well, use Skoglund instead or sample some Franco-Cantabrian or Iberian other Paleolithic remains.

Whatever the solution, I think that we do have a problem with the use of Lochsbour as only WHG proxy and that it demands some counter-testing. 

What about the ANE component? I do not dare to give any alternative opinion because I lack tools to counter-analyze it. What seems clear is that its influence on modern Europeans seems almost uniformly weak and that it can be ignored for the biggest part. As happens with the WHG, it's quite possible that the ANE would be enhanced if the sequence from Afontova Gora is used instead of that of Mal'ta but I can't foresee how much. 

Finally some speculative food-for-thought. Again using the visual tool of the PCA, I spotted some curiosities:

Speculative annotations on the PCA

Most notably it is apparent that the two WHG populations (Western and Scandinavian) are aligned in natural axes, which seem to act as clusters. Extending both (dotted lines) they converge at a point closest to some French, notably the only "French" that tends towards "Southern France" and Basques. So I wonder: is it possible that these two WHG cluster-lines represent derived ancient branches from an original population of SW France. We know that since the LGM, the area of Dordogne (Perigord) was like the megapolis of Paleolithic Europe, with population densities that must have been several times those of other areas. We know that this region was at the origin of both Solutrean and Magdalenian cultures and probably still played an important role in the Epipaleolithic period. 

So I do wonder: is that "knot" a mere artifact of a mediocre representation or is it something much more real? Only with due research in the Franco-Cantabrian region we will find out. 

112 comments:

  1. "On the other hand, they seem to cluster in the same extreme periphery of the European cluster, opposed to the position of West Asians, and therefore suggesting that there has been some West Asian genetic flow into Europe since then (something we all assume, of course). "

    It's not West Asian gene flow to Europe, it's North Pontic Steppe gene flow to both Europe and West Asia, along with R1. In other words, ANE + R1.

    This started during the Copper Age, and continued for a while. From the paper...

    "We observe a striking contrast between Europe west of the Caucasus and the Near East in degree of relatedness to WHG. In Europe, there is a much higher degree of allele sharing with Loschbour than with MA1, which we ascribe to the 60-80% WHG/(WHG+ANE) ratio in most Europeans that we report in SI12. In contrast the Near East has no appreciable WHG ancestry but some ANE ancestry, especially in the northern Caucasus."

    http://img12.imageshack.us/img12/3053/48ms.png

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    1. No. You seem to have completely misunderstood the matter (probably because of wishful thinking).

      The formal ANE scores are low across the board with a record high of 18.7% for Estonians and a record low of 4.1% among Sardinians. Most other populations are around 15%, except Basques which have a slightly lower score of 9.8%.

      What your quote says is that West Asians show no WHG admixture (logical!) but that they do show some ANE admixture also, notably North Caucasians.

      Mixing WHG with ANE is of no help because these components probably correspond to two different ancestral populations, even if their origins may have touched at some point. In essence the ANE component should correspond to Altaian UP (or at most Eastern Gravettian, i.e. last likely contact with Europe before the LGM) and the WHG must correspond to European specific later developments such as Solutrean and Magdalenian (or also Hamburgian-Ahrensburgian in the North).

      All them derive from ultimately West Asian origins (Aurignacoid cultures and later Gravettian originated all in West Asia) and that's why we all cluster as West Eurasians but beyond those pre-LGM common deep origins is where the differences between WHG, ANE and West Asian components begin.

      In the supplemental material it is formally tested the EEF affinities as function of Lochsbour and a Bedouin, with results that vary depending on the other control populations but ranking between slightly above 60% West Asian to almost 100% West Asian, so the WHG component in EEF is not greater than 40% most likely (the ANE component should be negligible, as happens with Sardinians but not directly tested for AFAIK).

      Then, as everyone can see in the simplified triangular plot Europeans align far away from the ANE component between the WHG and the EEF proxies. I question the validity of Lochsbour as good proxy for WHG (too drifted away probably) but otherwise I see no obvious errors in that triangular plot.

      The ANE component could be higher than spotted in the paper (AG2 is closer to Europeans than MA1) but I can't evaluate that with any clarity. The map you link to anyhow is just a measure or relative affinity in which we can see how Palestinians or Syrians score similar to Spaniards or peninsular Italians in ANE (MA-1) affinity, that would make them ~12% ANE, maybe a bit less but clearly above the Sardinian score of 4%, which is represented in black color.

      The right map illustrates the same kind of variance in the WHG-EEF axis and this one obviously excludes West Asians because they would be hyper EEF always (off the scale). But this map says nothing about ANE, only the left one does.

      I see no reason to relate the ANE component with R1 as you want to do, although I guess that some of its influence is indeed owed to Indoeuropean invasions and other flows from Eastern Europe (Uralics, Pitted Ware, antique-medieval slave trade and even the Huns if you wish). But alone it cannot explain R1b for example. I'd rather believe in the Neolithic origin hypothesis for R1b than in the Indoeuropean one, which is absolute nonsense. Now, some R1a correlation in Eastern-Central Europe and even Central Asia... that I can agree to (but not to a simplistic R1-IE correlation, which makes zero sense).

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    2. You correctly has insisted a lot in reading the supplemental material. Well the relevant data for Stuttgart's WHG-WestAsian admixture is discussed in SI-10. Quote:

      "The amount of Near Eastern admixture estimated for Stuttgart can be seen in Table S10.2 and range between 61-98% with estimates increasing as the amount of estimated African admixture in BedouinB increases".

      The near 60% West Asian admixture figures correspond all to an assumed Bedouin African admixture of 4.2%. For an assumed African admixture in Bedouins of 5.1%, the West Asian admixture estimates in EEF are around 70%. For an assumed African admixture in Bedouins of 7.2%, the West Asian component in EEF would be around 90%.

      Considering my independent estimates (using Braña 2 and a Turkish proxy on the PCA) I would think that the realistic figure would be c. 60%. But that also should imply, I believe, that my alternative line for the EEF-WHG half-way score is approximately correct.

      More in depth it may also imply that, even if Bedouin African admixture is effectively higher than the assumed 4.2%, the excess was anyhow present in the EEF population, as its Canarian affinity suggests (and the Neolithic lineage of African roots E1b-V13 may help to explain).

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    3. Uralics, Turks and other post-Copper Age easterners brought more Eastern Non-African (ENA) than ANE to Europe. That's why some Northeastern Europeans are outliers today.

      Post-Mesolithic ANE in Europe is linked to Indo-Europeans, and didn't come with any ENA. Moreover, basically all Near Eastern admixture in Europe, except among Sicilians, Maltese and Jews, can be explained by ancestry from EEF, not requiring additional gene flows from West Asia.

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    4. "Eastern Non-African (ENA)"

      ... is a mere catch-all term with no score anywhere. Some ENA like Amerindians are strongly related to MA-1 and therefore the more real ANE component. In principle they should be treated as various populations being just a convenience term for all non-Africans who are also non-WEA.

      We can't say nothing about such paraphyletic element in any case because it's measured nowhere.

      "Post-Mesolithic ANE in Europe is linked to Indo-Europeans"...

      Maybe but we do not know. We do not even know when it arrived to Europe, we do not know if it was already affecting WHGs, we only have some quite uniformly low scores for this component across the board with records in Uralic peoples (Estonians but also off-the-scale Finnish, Russians, Mordvins). We cannot say much about that other than it peaks in NE Europe, notably in Uralic peoples.

      You talk of this stuff as if it'd be as you say: "X is Y, A is B, Z happened in the W way" without no cautions and quite annoyingly without support of any kind. That way tentative opinions are ascended to the category of dogma.

      I don't like that at all. We can't debate that way. That's like a believer preaching religious dogma: no alternative opinions allowed, no need for evidence. Why do you do that? It's a mere propaganda trick: not serious scientific debate. I know you can do a lot better.

      "Moreover, basically all Near Eastern admixture in Europe, except among Sicilians, Maltese and Jews, can be explained by ancestry from EEF, not requiring additional gene flows from West Asia".

      This part may be correct, at least I think it makes some sense on first sight.

      But then: why EEFs don't lean towards Turkey: they should be some >60% West Asian, what means Anatolian (with lesser some other affinity) but their tendency is not towards Turkey at all but towards Palestine, Bedouins, Yemen... It's not possible that a mere 10-18% ANE influence can distort the modern European ancestry that much in the PCA, so maybe there's more to West Asianness than what you say.

      Rather than rushing to conclusions, as you seem so willing to do, I get lots of questions. And I think that difficult questions are much better than misleadingly easy answers. Explanations must be produced if the data allows for them but when there is room for doubt it's better to generally remain in the question zone and at most produce clearly tentative hypothesis and not oversimplifying unfounded dogmas.

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    5. There's no need to make this so complicated. The paper doesn't settle everything, because more ancient genomes are required for that. But it gives a very clear chronology of how the European gene pool formed, including two sets of outliers, in the Northeast and Southeast.

      WHG/ANE = Mesolithic foragers

      EEF = Neolithic farmers

      ANE/WHG = Indo-Europeans

      ENA/ANE = Uralics

      Finally, there was West Asian, North African and Sub-Saharan gene flow into Southern Europe.

      I don't see how you can argue with that if you read the paper and looked at all the supplementary data properly. It is what it is, and we'll soon see it confirmed beyond any doubt when more genomes get sequenced.

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    6. ANE people living in the Pontic-Caspian would integrate neatly with the Pontic-Caspian Indo-European hypothesis, so I can see why that would be compelling. But I don't really see any way to test that their meta-population was or wasn't also already in Bactria or other adjacent steppe regions prior to the Neolithic - it seems possible to me they already were given the presence of a component with ANE like distribution across West Asia in non-Indo European folk.

      One problem with the f3 statistics for Caucasian / West Asian people here is that the Stuttgart Woman, the reference for farmers who were migrating to that, is already quite mixed with West Eurasian / West European Hunter Gatherer clade, to the proportion of around 56% of her ancestry.

      Some of this may reflect a real pre-Neolithic or early Neolithic population fusion among Near Eastern "farmers" but much of it is probably due to expansion to Europe.

      So the problem is that this sets an artificial floor of how much West European Hunter Gatherer ancestry ancestry is in the West Asian populations.

      If using Stuttgart as a reference for early farmers and the early Near East, West European Hunter Gatherer ancestry in West Asian populations cannot go less than around 50% of their early farmer component!

      This is a real problem since most West Asian populations probably don't have ANY real West European Hunter Gatherer ancestry. They're instead more likely to be almost wholly mixes of Ancient North Eurasian and Basal Eurasian (or ancient Mediterranean, ancient Near Eastern, whatever this ghost population is best termed). As WEHG and ANE are similar this results in reduced estimates of ANE.

      If we look at an example of how this affects Lezgins, they're estimated to be 0.712 Near East using Stuttgart as a reference for Near East and 0.288 ANE.

      This would decompose to 0.313 "basal Eurasian" (or ancient Near Eastern), 0.399 West European Hunter Gatherer, and 0.288 ANE.

      However in reality, it seems likely to me based on admixture that they're instead closer to around 0.686 (0.399 + 0.288) ANE (perhaps with some slight WEHG fraction).

      Another example is why Ashkenazi Jews come out as the "closest" population to EEF at around 0.931 EEF and 0.069 ANE in the comparison panels as well, this despite the fact that we know that Sardinians are a much better proxy for EEF.

      In reality they're probably have more ANE ancestry, but using the mix of WEHG and BE that is EEF sets a level of an obligate WEHG ancestry that prevents their ANE being accurately represented.

      I may crosspost this to bioRxiv, to ask the study authors...

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    7. @David:

      "The paper (...) gives a very clear chronology of how the European gene pool formed, including two sets of outliers, in the Northeast and Southeast".

      As far as I can discern it is a mere speculative reconstruction based on nothing measurable nor measured.

      I'm not making this complicated: you are by appealing to non-measured nor measurable hypothetical components like "Basal Eurasian" and "Eastern non-African".

      "WHG/ANE = Mesolithic foragers"

      WHG: (some) European Epipaleolithic foragers

      ANE: Siberian Upper Paleolithic foragers

      Two different things, two different origins and two different effects across the board.

      And then there were also pre-Neolithic foragers in West Asia, believe me. And they were ancestrally related to the other two groups. They just became farmers at some point.

      "ANE/WHG = Indo-Europeans"

      We do not know for sure. Was there not even a slight element of EEF or West Asian or Uralic admixture in them? In fact we know nothing of where the ancestors of Samara culture originated and how the previous archaeological layers evolved, but we know that the Uralic/Oriental genetic element was already around Eastern Europe in the Epipaleolithic: not just in Karelia and such but also in the much more influential area of Ukraine. Those mtDNA C lineages must have arrived from the Far East (almost certainly in relation with yDNA N1).

      "ENA/ANE = Uralics"

      That's the worst simplification of all. Obviously Uralics are a complex issue and West of the Urals they must have been growingly Europanized since very early. It is notable that the most distinctive marker of Indoeuropean-like autosomal influence in Europe peaks in and around Finland. It may be an artifact or whatever but all analysis since the long gone times of Cavalli-Sforza show that. Is that ANE? Probably not because ANE is quite homogeneous in all European populations. Is it related? Surely but rather obliquely.

      The ENA pseudo-component is not measured anyhow.

      "I don't see how you can argue with that if you read the paper and looked at all the supplementary data properly."

      If I'm missing something, please point me to the relevant chapter and I'll check it. All I see is WHG (with the many doubts explained above), ANE (also with some doubts) and EEF. They also measured EEFs vs. BedouinB (and Lochsbour) but this comparison is full of issues.

      "It is what it is".

      It is not the way you describe it in many ways. I don't know how you arrived to this oversimplifying dogma but it is completely wrong, precisely because your insistence on exaggerating the certainty of what you claim to be the truth, which is proper of a preacher, not a scientist.

      I know you can do much better than that dogmatic junk and I have no idea why you always seem to fail into such oversimplifications.

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    8. @Matt:

      The WHG vs EEF measure obviously can only measure (at best) those two European ancestral components (with the caveats I mentioned before). EEF is not and should never be read as West Asian, of course.

      However, according to their estimates EEF (multiple consistent formal tests rel. to Bedouin B), they'd be 60-90% West Asian, depending on the African admixture assumed for Bedouins (although they failed to assume African admixture in EEF, which is somewhat likely considering the the importance of E1b-V13 in all this issue).

      I don't know where you get your figure of 56% but it is below of what the authors estimated in the formal tests.

      "If using Stuttgart as a reference for early farmers and the early Near East, West European Hunter Gatherer ancestry in West Asian populations cannot go less than around 50% of their early farmer component!"

      Let's assume this you say is correct (unsure), what's the problem with that? Don't you see that Europeans invariably cluster separately from West Asians?, quite neatly so. Don't you see that WHGs cluster with Europeans and opposite to West Asians? So at least some WHG must be there.

      "This is a real problem since most West Asian populations probably don't have ANY real West European Hunter Gatherer ancestry."

      I don't see why this is a problem: the study does not measure West Asians relative to their WHG component (or when it does, they fall off the margins, as expected - some Europeans do to, incidentally).

      "They're instead more likely to be almost wholly mixes of Ancient North Eurasian and Basal Eurasian".

      "Basal Eurasian" is a chimera! (Chimera: mythical critter made of many different parts). You and David (and some others for what I read in his blog's comments) are reading too much in the hypothetical reconstruction graph, which is almost certainly wrong.

      This we can't discuss on the basis of this study only but it's fairly obvious in non-African genetics overall. West Eurasians are derived from Asians from beyond the Hindu Kush, with at most a tenuous remnant of West Asian OoA survivals. All West Eurasian lineages (excepted N1 and E1b on the yDNA side and whatever correlates within mtDNA) derive from the general Asian lineages: F on the yDNA side, N (and some M) on the mtDNA one. Overall the West Eurasian ancestry looks South Asian and to some extent probably also SE Asian (ancient Oriental).

      That's the true Basal Eurasian thing and is essentially indistinct from the Other non-African polarity (although Native Americans have some West Eurasian ancestry of the ANE type and South Asians were not measured).

      Once the West Eurasian expansion began some 50 or more Ka ago, you can forget about Basal Eurasianness: it's all West Eurasian (with the N1 and E1b partial exceptions). There's nothing (but those two minor elements) that does not originate in Europe, West Asia or Central Asia-West Siberia.

      So how can you negate the West Asian partial ancestry of EEFs? I don't see that being possible at all and must imply some sort of fundamental misunderstanding on your side.

      I don't think you measure Lezgins or Ashkenazim in the WHG-ANE-EEF triangle: they should fall off the edge because they are not essentially European. Just draw the "triangle" on the PCA: you cannot effectively measure anything that falls too much outside it. Can you tell me in which table are they measured such things?, because I believe there lays part of the confusion.

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    9. An important clarification on the "Basal Eurasian" thing:

      In SI-10, where they ponder the West Asianness of EEF (Stuttgart), specifically in fig. S10.1 they use a highly technical concept of "Basal Eurasian", which is nothing but West Eurasian with some African admixture. It is very clear in the model and it should be interpreted not only as effect of the remnant OoA elements but very especially as effect of the E1b influence in the Thessalian Neolithic and derived populations.

      In other words, for a founder effect reason, the European Neolithic component was more African-leaning than most West Asians from past and present. For that very reason it may present some confusing affinity (or quasi-affinity) to Palestinian and Bedouin populations, which almost without doubt were intermediate (in the Epipaleolithic?) for this African component in the European Neolithic, most apparent in the yDNA lineage E1b-V13.

      It's not a too real different composition of Neolithic West Asia but actually a founder effect in Thessaly and by extension in all Europe (more diluted, of course).

      This is also apparent in EEFs clustering best with Canarians, who have a good deal of North African aboriginal admixture, more than any European has.

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  2. About the French dot approaching Basque and "South French" (actually 7 individuals from Béarn) dots, I suppose it matches the aberrant individual from the famous Lyon sample that most studies use.

    You can indeed see that one of the individuals from the Lyon sample possess a clear SW French genomic identity (on the following graph, much more loaded in the pink component and nearly lacking the yellow component) :

    Graph

    We clearly need regional French samples, both ancient and modern.

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    1. Looks like you're right, Heraus. Can you tell me if all the usual "French" sample is from Lyon? It's interesting to know that the "South French" sample must read as Bearn Gascons, because these invariably fall extremely close to Basques.

      "We clearly need regional French samples, both ancient and modern."

      ABSOLUTELY!

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  3. The HGDP French sample was indeed collected in Lyon. But after looking at it very extensively individual by individual in various admixture runs, and comparing it with Dodecad and Eurogenes samples, I am quite convinced that it's a good representation of French diversity in general, with perhaps a slight Rhodanian bend. There is indeed an obvious Gascon, but also an individual with recent Ashkenazi ancestry, several who cluster close to the British Isles and appear to be from the Channel coast (from Brittany to Dunkerque), as well as some more clearly mediterranean. Comparison with French Canadian samples, whose ancestry is overwhelmingly Western French also makes the Lyon sample seem a good representation of French variation.

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    1. Nice to know that for a fact, I wasn't sure if it was the same sample.

      I would not say that they reflect well French variation because for some reason they seem clearly scored northwards. This should be obvious when we compare this sample with the Bearn sample. They may reflect well the variation of Paris but what about Gascony, Brittany, etc.? France is a very large state which has played a key central role in European prehistory and history. It still does, although in the Ice Age it was much more important, of course.

      Something I often complain is that many samples from small ethnicities like Lithuanians, Slovaks, Basques, Sardinians, Finns, etc. may overshadow the real structure of Europe because the really big states, which hold the bulk of European population are seldom sampled regionally nor represented according to their real (or historically realistic) weight. You can't (shouldn't) just compare in most cases 20 French and 20 Lithuanians, because it's 70 million French and 3 million Lithuanians. You can compare 20 Lithuanians with 20 Basques but better to compare 20 French (representative of all the territory) with 20 or 40 Russians (also territorially representative) in order to get a realistic structure of Europe in the autosomal aspect.That's a very neglected aspect, very especially in the case of France.

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    2. Is there some reason, such as French specific privacy laws for the poor sampling in France?

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    3. I know mostly from our Gascon friend Heraus but not enough to discern the exact legal and bureaucratic obstacles. As you may know several countries such as India forbid the export of human DNA and this may be the core of the French legislation.

      But French red tape can be utterly annoying, more so as there are nationalist ideological reasons to oppose the "butchering" of the Republique on regions and ethnicities. French national ideology is quite extremist in opposition to all forms of difference: everyone is equally French, no peculiarities allowed or almost. The few studies performed have suggested a strong diversity within the Hexagon and that's something that many may feel threatens the artificial essence of France as unity. At least that's the impression I have.

      An example of utterly annoying red tape for no reason in France was the recent long struggle of organic farmers to get nettle purine legalized again as fertilizer. The state had arbitrarily decided to forbid it and was stubborn in that position for many years. It was eventually legalized again after a long environmentalist struggle but there seems to be no such impetus re. human population genetics.

      So researchers and private companies alike seem to face all kind of legal trouble but I don't really know the details in depth, just Heraus complaining all the time. Ask him.

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  4. I am working on some maps of Europe for ANE, WHG and EEF for David at the moment, hope that can add some clarity to the numbers.

    I find myself wondering if there is a possibility that the variation between WHG and ANE was taking the shape of a West/East cline across Northern Eurasia, rather than that of completely distinct entities with a break off point somewhere East of Europe.

    It seems like the pattern of Siberians moving west in waves is quite ancient, almost akin to the meteorology of the Siberian high pushing cold air out every winter. Could it be that the following happened :
    -As a result of the Chinese Neolithic, Hunter-Gatherer yDNA N carriers and associates are pushed North of Manchuria into Siberia by the advent of yDNA O agriculturalists on their lands.
    -Thus interbreeding with and displacing (perhaps due to some technological advantage) the local Siberian "ANE" towards Western Eurasia.
    -As the Neolithic revolution is taking over Europe, the new "ANE" are kept at bay in far Eastern Europe and the Northern Slopes of the Caucasus (where they gain admixture from agriculturalists, thus creating the West Asian component we usually see) and adopt pastoralism at the margins of neolithic civilisation on the Steppe.
    -Something (perhaps a harshening of winters, probably the same thing that has also pushed Huns, Xiongnu, Mongols and Turkics out of their homeland) pushes a subset of yDNA N (most likely Uralic-speaking) carriers West of the Ural. Again interbreeding with and displacing the ANE-WHG Hunter-Gatherers from the Northeast European forest belt.
    -That last event must have upset the demographic balance in the area and probably caused our ancestrally Siberian ANE (but now quite admixed by local HG and some trans-Caucasian agriculturalists) Steppe pastoralists to move West into Europe perhaps bringing with them IE languages...
    -Later the same pattern repeats with Huns, and Turkic speakers, spreading West, displacing and interbreeding with populations in their wake. If we compare the levels of Siberian admixtures in Turkey to the levels of "ANE" admixture in Europe, things seem quite comparable and can even make place for a more ancient "ANE"-like presence in Europe as part of a Eurasian paleolithic cline...

    It also explains the difference between the Caucasus and "Gedrosian" components we've seen regularly pop up. With Gedrosian being an admixture of Middle Eastern agriculturalists and "ANE"-like people from the Himalayan slopes (another place where Turkic speakers would later come to settle) and Caucasus being an admixture of Middle Eastern agriculturalists with drifted and WHG-admixed "ANE"-like people in the Caucasus.

    This leads me to think that contemporary Turks cannot be a good proxy for Neolithic Anatolians. I believe that what comes up as "West Asian" (as opposed to "Southern"/"Middle Eastern") in admixture tests in Turkey is the result of a more recent flow into Anatolia of ANE-admixed agriculturalists from the Caucasus (perhaps IE speaking, but also likely to speak Caucasus languages, depending on their level of ANE-admixture) back into Turkey (and incidently Greece).

    Although your idea of African-admixed yDNA E1b Thessalians being the cause of that "Southern" component as opposed to the "West Asian" component being much more ancient in West Asia is quite tempting. I am going to reflect on it, as it brings an opposite vision to the model I have in mind.

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    1. "I find myself wondering if there is a possibility that the variation between WHG and ANE was taking the shape of a West/East cline across Northern Eurasia, rather than that of completely distinct entities with a break off point somewhere East of Europe."

      Impossible to say without aDNA from Eastern Europe and the like. But in any case WHG and ANE are defined for being different clusters of Paleolithic sequences, so the distinction is clearly there.

      "It seems like the pattern of Siberians moving west in waves is quite ancient"...

      Epipaleolithic at most, I'd say. Earlier the flow seems rather from West Asia (and maybe in some cases also Europe) into Siberia. The only doubt I have is regarding Kostenki and its U2 sequence, which may indicate one such early minor flow from Central Asia.

      The yDNA N1 flow is almost certainly from Epipaleolithic onwards in the case of Europe and it came via the taiga area, which was earlier (Ice Age) almost certainly uninhabited. Rather than pushing peoples westwards it seems that the N1 clan went around the pre-existing peoples through the Far North. Their distribution does not suggest that they pushed others around, at least not often.

      It's simpler to think that most ANE advanced with Indoeuropeans but then, why don't we see a stronger gradation? Maybe the lower frequencies are just an artifact? Maybe the ANE component partly reflects the Eastern European influences in Siberia, such as Gravettian? In this last case it's possible that the basic levels of distributed European affinity are mostly reflection of the partial Europeanness of ANE peoples themselves. Can't say with any certainty.

      ... " contemporary Turks cannot be a good proxy for Neolithic Anatolians."

      Better than Bedouins for sure. Anatolia has suffered several invasions since the Neolithic but it was also always a densely populated area, so the impact of each of the invasions should be weak, although cummulative. However the invasions arrived from different corners, so the accumulation of weak effects should cancel each other to some extent.

      The Caucasus-Gedrosian component makes all the sense to me anyhow: it's Zagros Neolithic, as opposed to Eastern Mediterranean Neolithic (which came in two flavors: Palestinian and Anatolian). Today we'd say "Iranian" but then it was more like Elamites, Hurrians, Sumerians and such. West Asia was not homogeneous in the Neolithic but it had at least those three distinct regions: (Southern) Anatolia, Palestine (and some other Levant) and Zagros (with expansion to Mesopotamia), they met around Gobekli Tepe (although in origin this megalithic site was still Mesolithic). Then Elamites were clearly a different group than Sumerians and there were probably other ethnic nuances like the difference between PPNB and the CAPC. We can't risk oversimplifying Neolithic West Asia: it was a complex of several cultures. Some of those distinctions we still see today quite neatly, like the two quite apparent West Asian autosomal components (N & S), related to the geographical cline between yDNA J2 and J1, which is almost certainly from the Upper Paleolithic. They overlap but the duality persists.

      "Although your idea of African-admixed yDNA E1b Thessalians being the cause of that "Southern" component as opposed to the "West Asian" component being much more ancient in West Asia is quite tempting".

      Thanks. I find easier to think in terms of founder effect in this case, rather than imagining mass replacements in West Asia after Neolithic. After all lineages like yDNA J2 do not come from outside that region.

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    2. "The yDNA N1 flow is almost certainly from Epipaleolithic onwards in the case of Europe and it came via the taiga area, which was earlier (Ice Age) almost certainly uninhabited. Rather than pushing peoples westwards it seems that the N1 clan went around the pre-existing peoples through the Far North. Their distribution does not suggest that they pushed others around, at least not often. "

      That's fair enough. I guess if we accept the narrative that there was some movement of pIE speakers out of the steppe, there needs to be a reason they left, I don't believe they were somehow mystically inhabitated with a conquering spirit, something must have forced them out a bit, Uralic newcomers seemed like a good culprit . However climate change would be another very good culprit.

      "It's simpler to think that most ANE advanced with Indoeuropeans but then, why don't we see a stronger gradation? Maybe the lower frequencies are just an artifact? Maybe the ANE component partly reflects the Eastern European influences in Siberia, such as Gravettian? In this last case it's possible that the basic levels of distributed European affinity are mostly reflection of the partial Europeanness of ANE peoples themselves. Can't say with any certainty. "

      I am stuck with this question as well. I think ANE has been present in Northern, Central and Eastern Europe since at least Epipaleolithic times (as in the Motala samples) and the result of exchanges and continuum between WHG/Sauveterrian/Tardenoisian Atlantic refugia and a ANE/Gravettian Mammoth Steppe ; and if an ANE influx there has been since then, it must have been quite minimal.

      I took to the task of making some maps representing how I believe preNeolithic populations spread out and what was the stage at the dawn of the Neolithic revolution in the Middle East. But I realised this is a very difficult task, and resulted in a series of maps seriously lacking in subtlety. Still I thought I'd share them with some ample commentary to convey that subtlety which the maps lack. Please feel free to correct me, I am far less knowledgeable than you are on the topic.

      http://i43.tinypic.com/rj1bur.jpg
      This map attempts to represent a repartition of different more or less differentiated groups at about -35000BCE. The lack of precision in the assignments of mtDNA subclades (see U in particular) is aknowledged and waiting to be corrected. The haplogroups assigned are meant to be read as being that of the plurality/majority of people in the regions they are assigned to, they were even perhaps a minority but are mentioned as the only ones that survived from that specific location/population.
      -Africa in Maroon is populated by "paleo-African" HG groups, perhaps very differentiated from each other, but it's hard to infer to which extent from current data.
      -The Yellow group in the Middle East represents a hypothetical Basal Eurasian group that would have somehow re-expended in the region (as part of a wet cycle in the area), and mostly carrying an undifferentiated yDNA E with some more traditionally "West Eurasian" and Middle Eastern mitochondrial haplogroups.
      -Blue is quite straightforward, although quite hypothetical.
      -Grey, purple and teal are similarly very hypothetical. I am tempted to think the spread of P from somewhere in Northern India or Southeast Asia has displaced these groups towards the West, or at least eaten away at some of their territory.
      -The Brown section over India is quite self-explanatory, although I left the mitochondrial very unspecified (similarly with other places in Asia) because haplogroup diversity was already great in the area at that time.
      -I realise I let the Blue area spread far too far South for that time. I think Southeast Asia at that time actually had more groups transitional between the groups in India at that time and current Papuan groups (yDNA S and M).
      (to be followed).

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    3. http://i40.tinypic.com/zk5qip.jpg
      This map attempts to represent populations during the LGM.
      ppia = previously present in the area
      -The Middle East and Sahara had to be evacuated due to their drying up. Paleo-African hunter gatherers retreated to the South and the North. Where they were joined (taken over?) by formerly "Arabian" yDNA E carriers, creating new hybrid populations (resulting in Ibero-Maurusian in North Africa for instance).
      -Our blue "Aurignacian" people had definitely spread to Western Europe (although realistically, I should have already put them on Europe in the previous map, instead of only Anatolia), and were trailed by "Arabians" in Anatolia and Southeastern Europe where new hybrid populations appeared.
      -Most of the pink P (now Q and R) carriers were squeezed North by advancing Hindu Kush glaciers and Central Asian desert onto the Mammoth Tundra-Steppe, an ecosystem they would exploit as nomads.
      -Dark blue and purple again are supposition. A branch of R (R1b?) would have stayed South there.
      -I think Asia is also quite self-explanatory although I overestimated the spread of O-populations again.

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    4. http://i42.tinypic.com/2cid9b8.jpg
      This maps attempts to represent Eurasia at the dawn of the Neolithic Revolution in the Middle East after the retreat of the last Ice Age.
      -Maroon still represents paleo-African HGs.
      -Orange represents Neo-African populations, from which the Bantu expansion will later begin.
      -Warm Yellow represents North-African populations descended from the earlier Ibero-Maurusian.
      -Acid Green are other populations descending from yDNA E-carrying Basal Eurasians and "West Eurasian" mtDNA carrying populations in the Eastern Mediterranean basin (Natufians etc...). They will soon be involved in the Thessalian and Egyptian (and perhaps Levantine) Neolithics.
      -Blue represents the continuity of Aurignacian into Sauveterrian culture populations in Western Europe (WHG).
      -Purple represents a hybrid population (SHG) resulting from the meeting of the pink Mammoth Steppe ANE population and Hunter Gatherers from the Franco-Cantabrian Refugium.
      -Pink is rather self-explanatory. The Pink population would push remnants of the Purple population further into Western Europe during the Iron or Bronze Age, resulting in the current ANE pattern in Europe, probably bringing IE languages, likewise it would be squeezed south onto Central and South Asia.
      -Claret is there to represent the admixed population that went onto the Americas, although it's very likely they were already extinct in Eurasia by that time.
      -The Teal population of the Middle East is already quite dense and the result of admixture and back and forth movement (following game perhaps) around the area of previously present groups. I expect there to have been several foyers of dense populations with there own distinct language and drifted paternal lineage each. The Neolithic revolution would be sparked more or less simultaneously among a variety of these foyers. J2 and
      G2a rich foyers (located around current Armenia and Anatolia I would expect [PPNB?]) expended onto the Mediterranean and the Caucasus, mating with indigenous populations on their way creating EEF in the former (with WHG and E-bearing Thessalians and East Mediterranean Neolithic) and the "West Asian" component (with ANE) in the latter. Although on G2a bearers would have reached Europe the original J2 and G2a were likely from the same population, I believe, which haven't left any linguistic traces today (perhaps Hurro-Urartian, but I don't think so).
      J1 foyers would have been located further South in Mesopotamia and spread both North towards the Caucasus and the Arabic peninsula. Again, I wouldn't assign them a linguistic family, perhaps Sumerian, but definitely not Semitic (which came from the Levant later).
      I am tempted to associate the Zagros Neolithic both with another J2 foyer and a R1b foyer, of which a R1b group would have spread Westward through Anatolia towards the Danube, as well as some towards Central Asia, and perhaps spoke a now extinct proto-proto-proto-Vasconic like language.
      Likewise an L foyer somewhere in Mesopotamia expended East, but also North (L in the Caucasus).
      -As of Asia on the map, I am very unsure about the Northern yDNA C carriers, and they might well have been limited to a much smaller refugium along the coast and in Mongolia. The N group was ready to expend North into the Taiga. And O is most likely still overestimated...

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    5. Something that happened in the Chalcolithic, even where there was no copper metallurgy, was that society became more complicated than before, with social hierarchies and specialization and armed conflicts everywhere. In order to conceive the Chalcolithic one may think in pre-Columbian America, for example, although that was rather a late than an early phase of the Chalcolithic.

      In North America, for example, there were wealthy civilizations like the Mexicas, which attracted the greed and ambition of barbarians like the Nahuas (Aztecs) who arrived from less wealthy contexts but managed to take over. Similarly Karanovo-Gumelnita worked as a magnet for early Indoeuropeans (or Sumer for early Semites, etc.)

      It is true that there were other branches who arrived to less strikingly wealthy locations such as Altai or East Germany but it is also true that we cannot discern the detail of the politics of the time. On the contrary we have a quite decent knowledge on how the Mongols began expanding in all directions, implying the rise of an unusual statesman (Gengis Khan), unification of tribes and a militarist-imperialist policy that glued the polity. This is similar to what we know of Attila or, barring the peculiar religious element, what we know of Mohamed and successors. So it's easy to assume that there was: (1) political-military organization in the expansionist side (and almost certainly individual leaders whose names have been forgotten) and (2) weakness on the victim's side. The exact details we will never know but I think it's clear that there is no need to appeal to pressure from third parties or climatic elements as fundamental to Mongol or Islamic expansion, and therefore they are also not necessary for the prehistoric expansions of comparable character like IE, Semitic or Turkic ones.

      As for Uralic peoples, I don't think they were numerous nor politically organized enough to pressure the early IEs. Also these expanded in all directions (E, S and W), what means to my mind that all these avenues were open to their newly found warrior might.

      The ups and downs that we can discern in IE cultures probably relate to political cycles of unity and division, and maybe to individual leadership or lack of it.

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    6. "This map attempts to represent a repartition of different more or less differentiated groups at about -35000BCE".

      I can't agree with many elements. For example, yDNA P is almost certainly of South Asian origin, while mtDNA X must have coalesced further to the SW, because X1 is almost exclusively African (Egyptian). Similarly yDNA LT must correspond to Pakistan, H to Southern India, E to Africa only, while IJ soon split into I (Europe), J1 (Southern West Asia, projecting into Africa) and J2 (Northern West Asia). Back to P, R1a might have coalesced around the area you suggest (or maybe further North) but R1b almost certainly coalesced in West Asia senso stricto, with only one branch being present in Central Asia. Q coalesced either in Iran or neighboring regions of South Asia. Tibet was a frozen desert back then, etc.

      Also the date for most of those nodes is doubly old, of around 55-60 Ka ago.

      "Africa in Maroon is populated by "paleo-African" HG groups".

      Actually yDNA E had already taken over most of Africa, along mtDNA L2 and L3.

      "The Yellow group in the Middle East represents a hypothetical Basal Eurasian group".

      I do not believe in the "Basal Eurasian" concept. I think it hides actual African admixture in EEFs and Levant peoples. Said that, there is almost positively some "basal Eurasian" remnant in parts of Arabia Peninsula but only conserved in mtDNA. This remnant had no important influence in Northern West Asia nor Europe and even in Arabia it is pretty much residual. It was overcome by the Asian peoples who arrived in the earliest UP and who replaced also the Neanderthals who inhabited most of the West Eurasian region.

      "The Brown section over India is"...

      ... wrong in my understanding. That area had two subregions: N and S, separated by semi-arid areas in the Northern Deccan. The South was almost certainly dominated by yDNA H, while the North (Narmada-Ganges-Indus) was the boiling pot that spawned some of the most successful lineages: CF/F first, then K, then P. All that of course much earlier than 35 Ka BP, between 100 and 60 Ka ago in fact.

      "I realise I let the Blue area spread far too far South for that time."

      Not at all. By 35 Ka BP IJ was spread through all or most West Eurasia and also probably NE Africa (J1). Part of the problem with your concept is that you tend to imagine too short chronologies but very especially too tightly cut distinctive population clusters. Most likely by 35 Ka BP, most of those lineages were already very mixed with each other.

      Focusing in West Eurasia, I would expect c. 35 Ka BP that:
      → Europe: I (at the very least)
      → Levant: J1 (maybe also G)
      → Highland West Asia: J2, G
      → Central Asia (with emphasis in Siberia): Q1, R1a (also as minor lineages in West Asia, where they probably originated)
      → R1b somewhere in West Asia, don't know if already in Europe (it may have arrived with Gravettian only).
      → L in Northern Pakistan
      → T in Southern Pakistan with extension through Arabia and probably projecting to Africa

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    7. "This map attempts to represent populations during the LGM".

      I cannot agree in any way. For example:
      → If I was concentrated in SW Europe, why the origin of I appears to be in or near Ukraine?
      → Why on Earth would be Anatolia dominated by E?
      → etc.

      "This maps attempts to represent Eurasia at the dawn of the Neolithic Revolution in the Middle East after the retreat of the last Ice Age".

      I'm still perplex at many of your conclusions. At the very least R1a was not present in Central Europe yet. In Europe R1a almost certainly spread with Indoeuropeans (with maybe lesser exceptions). Also, you are assuming that R1b was still in mainland West Asia and was not part of the Greek Neolithic, what makes impossible its arrival to Western Europe in its last possible window (Neolithic).

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    8. Hey Maju!
      Thanks for the reality check. I realise trying to orchestrate and map some sort of all encompassing narrative with only vague dilettante knowledge of the field can lead to that type of mistakes you are pointing out...
      I very much agree with P originating in South Asia that said. Also I placed R1a early on in Central Europe to account for that early L664 clade. Your conclusions are otherwise very sound. I also agree that the tightly cut distinctive populations is unrealistic, which is an easy pitfall with that sort of mapping. I think that that tendancy to categorise, in many of us, is what leads to misunderstanding admixture results (in particular I think the variety of West Asian components we see...)

      I will have a good think about the idea of R1b as gravettian. To be honest R1b is a piece of the puzzle I can't really reconcile. Looking at the series of flaming hoops that someone like Maciamo from Eupedia makes the R1b chronology jump through, just so that he can associate them with Indo-Europeans, is something that I want to find a sound counter-narrative to.

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    9. "So it's easy to assume that there was: (1) political-military organization in the expansionist side (and almost certainly individual leaders whose names have been forgotten) and (2) weakness on the victim's side. The exact details we will never know but I think it's clear that there is no need to appeal to pressure from third parties or climatic elements as fundamental to Mongol or Islamic expansion, and therefore they are also not necessary for the prehistoric expansions of comparable character like IE, Semitic or Turkic ones. "

      I have to admit, I get wishy-washy wishful ideas that imperialism/expensionism is not intrinseque to human nature, that it can only be the result of hardship/pressure, get in the way of recognising what you point out here. On that note, I guess the history of colonisation (like here in Australia, or looking at the Americas) would be another parallel to what you say (although in a different technological context, of course).

      Would you say that that model of conquest you describe is one that can also be ascribed to R1b in Europe?

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    10. Oh and also :
      "If I was concentrated in SW Europe, why the origin of I appears to be in or near Ukraine?"
      I was trying to show that I originated in Western Asia and spread to Europe. Not sure where Ukraine comes in.

      "Why on Earth would be Anatolia dominated by E?" They would have been there before being pushed towards Greece by advancing groups from further East... I wonder how E could have gotten into Greece, would people have been able to sail there from the Levant in Paleolithic times?

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    11. "Looking at the series of flaming hoops that someone like Maciamo from Eupedia makes the R1b chronology jump through, just so that he can associate them with Indo-Europeans, is something that I want to find a sound counter-narrative to".

      In truth R1b looks anything but Indoeuropean. I mean: there is a bunch of R1b in Central Africa for example and never mind the absolute hegemony of this lineage among Basques, quite greater than among our IE neighbors (so IE = less R1b in SW Europe).

      So there are two options: Paleolithic or Neolithic. For what I know of Basque origins both could fit well. However the Neolithic aDNA has not yielded yet any R1b and the pattern of spread of its main subhaplogroup R1b-S116 strongly suggest a South France origin, which does not fit well with Neolithic. So I'm still leaning to a Paleolithic origin with re-expansion in Chalcolithic times, associated probably to the increase of mtDNA H in Central Europe in this period and decrease of this matrilineage in Portugal (maybe in the Bronze Age), so maybe it re-expanded with Megalithism but with ultimate Paleolithic origins in SW Europe.

      That's the narrative I can muster but in wait for more clarifying aDNA, of course.

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    12. "I have to admit, I get wishy-washy wishful ideas that imperialism/expensionism is not intrinseque to human nature, that it can only be the result of hardship/pressure, get in the way of recognising what you point out here".

      I don't feel able to detect the possible "natural hardships" at play but I have clear that population can grow fast easily, leading to internecine conflicts, leading in turn to exiles and leading in turn to export of that internecine conflict. These spontaneous problems can be aggravated by hierarchization of society and wealth-grabbing on few hands.

      In a modern example, one can look for environmental pressures in Africa or the Muslim World behind the ongoing emigration wave, but the main reasons are unchecked demographic growth and widespread poverty. These migrants are workers because of the current conditions but there is no reason why they couldn't become armed invading hordes in other conditions, more proper of the Metal Ages (without the overwhelming military-technological gap of today).

      In the Metal Ages, organized polities could not conquer extense open spaces like the steppe or the circum-Arabian semi-desert (unlike today), so in the best case they could fence off the pastoralists with "stick and carrot" and "divide and rule" policies and, in the worst case, they became their prey. When, for whatever reason, these pastoralist groups became unified (typically behind a charismatic statesman), they also became a major threat, more so if the sedentary polities were in bad shape.

      "Would you say that that model of conquest you describe is one that can also be ascribed to R1b in Europe?"

      I don't think so, for the reasons mentioned above.

      "I was trying to show that I originated in Western Asia and spread to Europe. Not sure where Ukraine comes in".

      Well, in my understanding that area has the greatest basal diversity for yDNA I, so I tend to imagine that it expanded from that region (possibly in various waves).

      Alternatively, the abundance of yDNA I (I2) in NW European HGs could suggest other centers for the expansion of this lineage but it cannot be related to West Asia in any case: I is clearly European since Paleolithic times.

      "They would have been there before being pushed towards Greece by advancing groups from further East"...

      Difficult to argue on light of the very low frequencies of E in Anatolia today.

      "I wonder how E could have gotten into Greece, would people have been able to sail there from the Levant in Paleolithic times?"

      I suspect so. They may have gone via a coastal route with no or very limited impact in Asia Minor but we know for a fact that the early European Farmers (Impressed-Cardium variant) were deep sea sailors and we know also that people were sailing to Cyprus since the Mesolithic, so an epic sailing journey is not out of the question either.

      If you look at the frequencies of yDNA E1b it is clear that there is a gap in Anatolia. So either you imply a mass replacement in that region or you assume that Asia Minor was essentially skipped by the people who arrived to Greece in the early Neolithic. They did not really need to skip it but they were never able to cause a founder effect in that peninsula in any case. However in the SW Balcans they were clearly successful, with secondary impact in much of Europe.

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  5. I might also make some maps to clarify the model I have in mind, writing it down might make it a bit obscure. I would love to hear what you think, or how it matches with archeological record.

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    1. Sure. I'd like to see those maps in any case.

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    2. Just saw them. Nice maps.

      I have some lesser concerns like moving Lyon to Central France (not the true data point) or the equally dubious ref. for England near York (shouldn't be further south, maybe in London?) Does somebody know if the HGDP Spanish sample is from Madrid or Valencia or what? Given the size of these states, different positioning to their true geography may introduce important errors in the clines.

      Also the lack of scale may lead to the wrong assumptions about the meanings. For example, in the ANE map Sardinia appears as "white", what may be understood as zero, when it is in fact 4.1%. In that same map the differences appear very much aggrandized: even if most populations range very close (12-18%) in absolute values, they are shown with very different colors. Personally I'd use just four gradients: <5%, 5-10%, 10-15% and 15-20%.

      Otherwise it seems a good reproduction of the apportions suggested in the paper, which, as I mentioned before I do not agree with too much, especially in the use of Lochsbour as proxy for WHG without further control testing. But those problems are from the paper, not yours.

      As for the color tripartite map, it really ends up looking bipartite. I would suggest using a two color WHG-EEF scale and marking the presence of minority ANE with some other means than a third color that ends up looking pretty much invisible, for example dotted lines (a single dotted line indicating the <10% ANE pockets should suffice).

      Self-reference for further discussion:
      → http://i40.tinypic.com/2dt8zmb.jpg (WHG)
      → http://i44.tinypic.com/9gzvyc.jpg (EEF)
      → http://i42.tinypic.com/bjd3sn.jpg (ANE)
      → http://i40.tinypic.com/9tlw76.jpg (color composite)

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    3. Sorry for answering so late. I've been on summer vacations away from internet for a while.

      I agree with all your suggestions. I've mostly attempted to place data points near the geographic centre of the countries they are supposed to represent, based on the assumption that the population sample is a good representation of said country. Which is something I still believe about the French hgdp sample. But I should be more rigorous and actually use the actual sampling locations.

      Sardinia isn't actually white, it's a very light grey, but your criticism is completely valid, it is not apparent, and a scale is definitely needed. I'll do that next time around.

      Also your suggestion for the composite map is one I will definitely take on board.

      I will probably try making new versions of these maps in the near future, taking your advices into account. But I might wait to see if new developments or new samples come up.

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    4. I am not sure if my answer to this was posted. But in case it wasn't, I was appreciative of the valid criticism, and will definitely take it on board next time I am making maps (hopefully when new data arises within this context).

      Delete
  6. "Rather than pushing peoples westwards it seems that the N1 clan went around the pre-existing peoples through the Far North."

    I think that they took different routes (N1b took a more northern route than N1c-E) and mixed with people carrying R1a and I1 lines. Within N1c there are at least two different flows to the west, earlier one and later one that affected only the Urals area. It is also possible that the Balkanic N is the earliest N in Europe that reached Europe first.

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  7. :)

    Someone needs to hand these authors a map of Europe during ice ages:

    http://www.ngdc.noaa.gov/mgg/topo/pictures/GLOBALsealeveldrop110m.jpg

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    Replies
    1. Marnie,

      Yes,
      As I have mentioned numerous times before, I would like someone to explain to me how SE Europeans could have been substantially different from W Anatolians, at the beginning of the Neolithic. It is virtually impossible.

      Delete
    2. Agree, especially since the Bosphorus is estimated to have opened only about 7,600 years ago.

      Delete
    3. What do you mean by "substantially different"? 8800 years ago (or 8500 with pottery) the evidence of agricultural settlements in Western Anatolia is as thin as a graphene layer (effectively zero). See: → http://context-database.uni-koeln.de/index.php

      Most relevant maps:
      → http://context-database.uni-koeln.de/img/P4B.jpg
      → http://context-database.uni-koeln.de/img/P4C.jpg

      There's a major gap between that area (Fertile Crescent) and Thessaly. It may be because of gaps in our knowledge but the gap is clearly there and recent research in Thrace and such has not changed the situation (Thessaly is still oldest farmer country of Europe and many hundred kilometers west of the nearest Anatolian (Cappadocian) site.

      It is very possible that Thessalian Neolithic was seeded by a coastal migration with peculiar founder effects (notably E1b-V13) which seem original from further south.

      The Ice Age sea levels do not matter, first because the Ice Age was long gone, second because with or without Bosporus/Dardanelles that area was always a very practicable crossing point but also a funnel.

      The problem is not what you can imagine from physical maps, but what we can understand from actual human socio-cultural geography.

      Delete
    4. "... the Bosphorus is estimated to have opened only about 7,600 years ago".

      Even if true (under some debate), it would not have affected the Aegean significantly nor the ability to cross between Anatolia and Thrace. The Marmara Sea already existed and crossing those extremely narrow straits can be done even with Neanderthal navigation skills very easily. It's not any kind of natural barrier at all and there is no archaeological evidence of people crossing through that path either.

      Delete
  8. I do think the LGM is big in this.

    imo

    1) Hyperborean parental population in a crescent shape from europe through north asia to siberia.

    2) Hyperboreans split into at least two child populations (and pushed south) by the ice, those child populations spreading out north again afterwards - i'll call them WHG and EHG.

    3) First farmers somewhere in eastern Anatolia
    3a) Small numbers of first farmers move north towards the steppe and mix with the EHG to eventually become the Kurgan guys.
    3b) Much larger numbers of first farmers move west into central Europe forming LBK mostly displacing the WHG to marginal terrain and the peripheries where they undergo a HG to herder transition.

    4) Something weakens the LBK leading to
    4a) firstly a herder over-run from the west into the center (funnelbeaker)
    4b) a second herder over-run this time from the east into the center (unetice)
    4c) bell beakers involved in this process as well but as a caste of miner/metalworkers like the african blacksmiths rather than as full displacing type populations

    5) All four populations WHG/EHG, Bell Beaker, LBK survivors merge in central europe to become the ancestors of Hallstadt, La Tene etc spreading out from there in multiple directions.

    Leading to
    Western Europe: WHG + first farmers + EHG (in indo-european form)
    Eastern Europe: EHG + first farmers
    South Eastern Europe: first farmers + WHG

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    Replies
    1. I don't understand much of what you say here, beginning with the totally misleading anachronistic mythical term "Hyperboreans".

      In Central Europe there must have been important Chalcolithic flows from the West in any case. Indoeuropeans alone cannot explain the current genetic pool. Maybe you should start thinking in other mythological terms: "Atlanteans". Just a guess.

      Delete
  9. "I don't understand much of what you say here"

    I think WHG and ANE will turn out to be WHG and EHG i.e. both descended from the same ANE population split by the ice.

    "In Central Europe there must have been important Chalcolithic flows from the West"

    That's the other part i'm saying.

    "Maybe you should start thinking in other mythological terms: "Atlanteans"."

    Yes, Funnelbeakers and Bell Beakers imo.

    ReplyDelete
    Replies
    1. "I think WHG and ANE will turn out to be WHG and EHG i.e. both descended from the same ANE population split by the ice."

      As I understand it, they both descend (in essence) from from the same West Asian population but ANE went to Central Asia and Siberia, and WHG ancestors went to Europe. Eastern Europe could have been settled from both origins but the main origin should be from Europe.

      Anyhow your hypothesis does not explain why the ANE affinity is homogeneous across the European and even all the West Eurasian geography, much more than the WHG one. No numerical data is provided for West Asia but, judging from the colors in the Extended Data Figure 6a, only some populations in Palestine and Yemen are at the relatively low levels of ANE affinity of Sardinians (when compared with EEF). Map EDF 6c shows that, when compared with Lochsbour, West Asians are extremely much more akin to ANE than Europeans (logical because their negative affinity is only rel. to Lochsbour WHG, not ANE).

      The study does not attempt (nor certainly can) study ancient admixture in West Asia (no local HG samples) but something that becomes clear in these visual analysis is that, while WHG defines Europe vs. West Asia (maps B and C), ANE and EEF affinities do not.

      We would probably need to make more extensive comparisons, with more ancient and modern samples (missing North Africa, Siberians), before we can reach to any well structured conclusion but, whatever the case for me it is clear that ANE has no particular preferential relation to Europe when considered vs. West Asia and hence it must be considered a population of general West Eurasian stock (with some Far East admixture probably), neither more akin to Europe nor to West Asia, but specifically Central Asian instead.

      "Yes, Funnelbeakers and Bell Beakers imo".

      Vale. I could not understand that in your previous comment.

      Delete
  10. "for me it is clear that ANE has no particular preferential relation to Europe when considered vs. West Asia"

    I agree. I think it will turn out to be ancestral over a very broad crescent shape stretching from Europe to Siberia and then became even more broad after that population was pushed south by the LGM.

    However what i'm guessing is ANE only *appears* to be more eastern because when the first farmers expanded out of their epicenter larger numbers (as a proportion) moved into west and central europe than the east.

    so
    Stage 1) ANE
    Stage 2) ANE -> WHG & EHG
    Stage 3a) WHG & lots of first farmers
    Stage 3b) EHG & far fewer first farmers

    (Quite possibly wrong but hopefully clearer at least.)

    ReplyDelete
    Replies
    1. WHG individuals seem less leaning to ANE than modern Eastern and even NW Europeans in the PCA, so if anything there's been some increase in the influence of ANE after the Epipaleolithic, not the opposite. ANE reflects an intrusive element in Europe just like the West Asian component (partly reflected in EEF).

      Additionally EEFs seem a simple West Asian + WHG mix, with negligible ANE relevance, so the excess ANE almost certainly arrived after Neolithic, what should be read as Indoeuropean influence. This seems corroborated by the fact that, other than Sardinians, Basques are the lowest scoring in ANE affinity (barely 10%, half that of Estonians).

      The question that seems to fuel your speculations may be the fact that Sardinians score even lower than Basques (less than half in fact) and that Sardinians seem a good proxy for EEFs. I'm not really sure how to explain this but either Basques have got some IE-like influence or EEFs scored particularly low in ANE affinity, as do for example Palestinians or Yemenis today. That's why the authors use the "Basal Eurasian" conjecture, I guess, but I'd rather explain it with North African admixture, something that seems very obvious in the fact that EEFs were vector for the spread of Y-DNA E1b-V13 in Europe.

      This "unexpected" African affinity of EEFs subtracts from basal ANE affinity, which should be otherwise relatively neutral for all West Eurasians, with lesser clines relative to geography or whatever. Alternatively or complementarily, some of ANE-affinity weight could reflect Gravettian roots, shared by all European HGs (West and East equally), as well as Siberian ones but whose impact in West Asia may have been less uniform.

      In other words: variable affinity with ANE does not need to reflect immigration from Siberia, it may reflect relations with peoples more or less akin to Siberian HGs since the beginning. Lower ANE affinity can for example be caused by greater North African affinity, while greater ANE affinity can be caused by mere Eastern European (Indoeuropean, Uralic) flows.

      "... hopefully clearer at least".

      I understand what you suggest but not why. There's no reason to think of flows from Siberia into Western Europe in the Paleolithic, none at all. Aurignacian and Gravettian have West Eurasian roots, while post-LGM cultures are all locally rooted in Western Europe. If there was some admixture with actual ANEs in the Paleolithic, it must have been an Eastern European phenomenon only (and Eastern Europe did not influence Western Europe back then).

      So I understand that the issue is best explained by "anti-ANE" African (or just Palestinian) extra affinity in EEFs and by later "pro-ANE" IE (and Uralic) migrations.

      Delete
    2. "There's no reason to think of flows from Siberia into Western Europe in the Paleolithic"

      Yeah i'm thinking the opposite. I think there were multiple routes out of Africa and one (not the main one) was Atlantic coastal: west coast of Africa - Iberia - Western Europe - Scandinavia - North Eurasia - Siberia in a crescent shape.

      I think that base population was split into two segments by the LGM (and pushed further south including into Caucasus / West Asia) and that some of the eastern half of that segmented original population came back west later as part of the Indo-European thing (but some of it was already there).

      Delete
    3. "I think there were multiple routes out of Africa"...

      Almost certainly not.

      "and one (not the main one) was Atlantic coastal: west coast of Africa - Iberia - Western Europe - Scandinavia - North Eurasia - Siberia in a crescent shape."

      Nonsense. Don't get offended but it's total unfounded nonsense. There are so many wrongs in that idea that I don't deem it worth explaining them.

      But I'll tell you something about maps: they are not the reality. For example Marnie's Ice Age map shows reasonably well the extension of the coasts in the LGM (not before nor after, just around that cold peak) but it does not show temperatures, ice shelves (covering most of the route you want to imagine) or Neanderthal presence.

      "I think that"..

      "I think" is not a good explanation, sorry. "I think" without factual support is not better than the "I believe" of some poor soul brainwashed by preachers.

      Science works the other way around: (1) let's see what we can get to know, (2) let's put that together in one or several most parsimonious and coherent explanations, (3) let's contrast them with the facts, (4) let's continue researching...

      There are absolutely no facts in support of your conjecture. You have provided not one, just "I think that"...

      Delete
    4. Hi Maju, Nothing to add about Paleolithic stuff (mad props to you for synthesizing and presenting so much Paleo information over the years - we are all still miles behind your knowledge and there are no substitutes).

      My one quibble is that yes, Palestinians and Yemenis score low in ANE relative to Europeans. But, these pops are much closer to ANE than they are to WHG, so there is some kind of relationship (maybe indirect somehow in terms of what's modeled in the paper?).

      Also, Iranians and Lezgins score especially high in ANE vs WHG ratio (like Palestinians/Yemenis but stronger), in addition to strong Z scores for direct MA1 admixture (unlike Palestinians/Yemenis).

      It's shown in the Extended Data Figure 6(C) at http://biorxiv.org/content/biorxiv/suppl/2013/12/23/001552.DC1/001552-1.pdf

      I agree that ANE looks like something non-European that came back into Europe and especially northern areas.

      Delete
    5. "... Palestinians and Yemenis score low in ANE relative to Europeans. But, these pops are much closer to ANE than they are to WHG, so there is some kind of relationship (maybe indirect somehow in terms of what's modeled in the paper?)".

      Good question. I think that it simply reflects relative affinity to either. As West Asians have no WHG ancestry, they score comparatively high in the opposite vector, which is similarly akin to all. So basically Europeans look "dark blue" (meaning strong WHG affinity rel. to ANE) and West Asians look "yellow" (meaning not strong affinity to either). Notice that the highest value is still negative (-2), so they are probably slightly closer to WHG than ANE in fact. Similarly all populations are closer to EEF than to ANE but more or less so (map A, same scale).

      Instead in map B there is positive score to WHG (+4), what means that some populations are more akin to WHG than to EEF.

      Delete
    6. "but it does not show temperatures, ice shelves (covering most of the route you want to imagine) or Neanderthal presence."

      People moved out of Africa before the LGM so the route I am imagining is before the things you mention blocked the route. The question then is what effect would the expansion of the ice have on that *pre-existing distribution* i.e. could a related population have been split into separate refuges and pushed south and secondly what re-expansion paths existed after the LGM peak when the sea levels rose.

      "Science works the other way around: (1) let's see what we can get to know"

      I accept that but an Atlantic coastal route which was largely restricted to the coast has the problem that the old coast is now underwater so any evidence would have to be looked for where the coast *was*.

      "There are absolutely no facts in support of your conjecture."

      Sardinia and Scotland. They don't support my conjecture but they don't support the ANE as Siberian either.

      #

      (On a completely separate note could the Slavic slave trade from the Crimea have effected the ANE proportions around the eastern med.)

      Delete
    7. "People moved out of Africa before the LGM so the route I am imagining is before the things you mention blocked the route".

      But then also the coasts would be similar to modern ones. Ice sheet and sea level are inversely correlated.

      ... "the problem that the old coast is now underwater so any evidence would have to be looked for where the coast *was*".

      Two major problems:

      1. As mentioned above, if there was no ice sheet, the coast was also higher, similar to modern, maybe even further inland.

      2. Why would people so strictly adhere to the coast. We should expect some groups traveling further inland or even the same groups moving camp seasonally or whatever. They must have left some sort of evidence in any case and nope.

      "Sardinia and Scotland. They don't support my conjecture but they don't support the ANE as Siberian either".

      ANE is Siberian by definition: it is defined by a Central Siberian specimen (MA-1) or at most by two (incl. AG-2, not really used in the tests). Their genetic influence in the West is very homogeneous (with very few exceptions all populations range ~15%, 3 points up/down) and that homogeneity includes West Asia or most of it.

      IMO at least part of that affinity is mere default affinity because of shared origin in Aurignacian/Gravettian. The "negative affinity" is what needs most explanation here and IMO this is best explained by (North) African influence or, if you wish, as the authors hint to albeit in imperfect manner, by a "Basal Eurasian" element (or Arabian/Palestinian OoA remnants, which would tend to Africa in any comparison).

      Speculatively it is possible that the Hamburgian-Ahrensburgian late UP population of the North Sea had slightly more ANE component than Magdalenian ones. That seems apparent from the position of Scandinavian Epipaleolithic samples in the PCA but that would be about it: less important variation because of ancient founder effects. The origin of Hamburgian is not well understood, so it's not impossible that they had some obscure Eastern influences.

      "... could the Slavic slave trade from the Crimea have effected the ANE proportions around the eastern med".

      You never know but I doubt it is a major element. It rather seems to me that the SE Europeans go secondary influences from Eastern Europe or West Asia after the initial Neolithic, slanting their affinity. Alternatively the SW European (Cardial) Neolithic had important founder effects affecting their overall ANE affinity (or maybe both).

      The reality is that we have too few references for deep judgment. We would need more ancient samples, preferably more relevant ones. All we have now is a generic draft and many open questions. It's better than nothing but clearly not the last word on the matter.

      Delete
    8. "ANE is Siberian by definition"

      Yes, my mistake. I meant "solely" Siberian.

      .

      "Why would people so strictly adhere to the coast."

      As climate zones and ecosystems are partially latitude driven then a population adapted to one latitude band that wants to expand north or south has to adapt to the different climate zone whereas a population who are already adapted to coastal food-getting require less adaptation to move north or south as long as they stick to the coast.

      I don't want to get too hung up on the OOA aspect. The main thing is i think it will be a general case that expansion along coasts will be faster than inland if the inland route crosses climate zones or obstacles because a coastal sea food getting population will have to adapt less.

      Delete
    9. Well, whatever the case, you are speculating with a totally invisible population for dozens of millennia, just because. I can't but dismiss that as the wildest baseless speculation, sorry.

      Delete
    10. "I can't but dismiss that as the wildest baseless speculation, sorry."

      That's okay. My background is one of creating as many speculations as possible and then trying to knock them down one by one. I appreciate it's not scientific.

      Delete
  11. More wild speculation and i apologize if it's annoying but one of the things that most struck me when i first started getting into this is how the R1b distribution looked like a blood splatter diagram with the spray pattern centered somewhere on an intersection south of Ireland and west of Brittany.

    If you drew a vertical line down from the center of Ireland and a horizontal line across from the center of Brittany the intersection would pretty much sit right on the LGM coastline.

    http://www.ngdc.noaa.gov/mgg/topo/pictures/GLOBALsealeveldrop110m.jpg

    ReplyDelete
  12. "So I wonder: is it possible that these two WHG cluster-lines represent derived ancient branches from an original population of SW France. We know that since the LGM, the area of Dordogne (Perigord) was like the megapolis of Paleolithic Europe, with population densities that must have been several times those of other areas."

    More "reindeer games", Luis.

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    Replies
    1. I explain here:

      http://linearpopulationmodel.blogspot.com/2014/01/mesolithic-western-european-hunter.html

      Delete
    2. That's a very complex speculation with many issues.

      First, the paper is PPV, so I do not know where and very especially WHEN the reindeer remains belong to in the "Eastern Cantabrian Region" (i.e. Southern Basque Country). If you know the dates, you don't provide them in the entry. I would imagine for what I know that the reindeers appeared in the coldest spell, i.e. around the LGM, not much before and not much later. In general the evidence rather supports other foods and totemic animals (first: deer, later: bison) as dominant, so the presence of reindeer in the LGM probably has little relationship with the Epipaleolithic remains discussed here and in your post.

      On the other hand it is possible that it holds some relationship between core Solutrean in the Franco-Cantabrian region in the LGM and apparent offshoots in Hungary and later Poland (i.e. maybe some FCR hunter-gatherers migrated to Central Europe before Magdalenian was consolidated and maybe these had some symbiotic relationship to reindeer or other tundra-like ecology.

      But since then what we see in the FCR and surrounding regions is:

      1. Development and expansion of Magdalenian (seemingly more oriented to aquatic resources and with important totemism of the bison).

      2. Development of early "Epimagdalenian" cultures such as Azilian (Laminar Microlithism).

      3. Possible migrations from roughly "Northern France" with the Geometric Microlithic industries (Tardenois).

      I would emphasize the latter because it may explain the alleged "Nordic" genetic affinity of La Braña, which in some aspects look immigrants (notably deer fang ornaments). Then the Neolithic arrived (essentially contemporary of La Braña site, so maybe they were being pushed around by Neolithic expansion as well).

      Finally I see no particular reason for your "transposition" of the axes, if anything the need of such transposition in your graph says that your conjecture is wrong, because you need to alter the data in order to reach to your conclusion, what, let's be frank, it's outright cheating. Suddenly, with no other explanation than your subjective need to arrive to certain conclusions (wishful thinking), you imagine axes-clusters that are not in the data but which you treat as if they were real. It doesn't make any sense.

      I'm not sure if I am onto something, as you say, with the issue of the axes (I clearly expressed that I was being highly speculative and had all kind of doubts), but certainly not what you imply after all those arbitrary "transpositions". You have taken my speculation and bastardized it beyond recognition. The axes converge where they do, not where you wish them to. If that convergence means anything at all or is just a meaningless artifact, I can't say at this point but what I'm certain is that your "transposed" axes only exist in your imagination.

      I feel a bit outraged, I must say.

      Delete
    3. Swimming Cows, Last Glacial Maximum Parisien Reindeer . . . what will she think of next?

      :)

      I added some references to my blog post. You can check them out if you would like. I did skim through them and it does appear that, more or less, the Paris Basin was continuously occupied by reindeer hunters during the Last Glacial Maximum.

      "On the other hand it is possible that it holds some relationship between core Solutrean in the Franco-Cantabrian region in the LGM and apparent offshoots in Hungary and later Poland (i.e. maybe some FCR hunter-gatherers migrated to Central Europe before Magdalenian was consolidated and maybe these had some symbiotic relationship to reindeer or other tundra-like ecology."

      A quick look at various papers seems to support this.

      Regarding bison, horses and other animals, its definitely possible. However, only reindeer hug and thrive on and near glaciers.

      "Finally I see no particular reason for your "transposition" of the axes, if anything the need of such transposition in your graph says that your conjecture is wrong, because you need to alter the data in order to reach to your conclusion, what, let's be frank, it's outright cheating."

      No. The ancient DNA data needs to be transposed because it hasn't been Neolithically shifted like most of the other samples. During the Neolithic, most of the other modern DNA samples have genetically shifted toward Greece and Anatolia. So to put the ancient DNA in the correct geographic position, it has to be "Neolithically transposed."

      I say most, because, probably, certain groups such as the Basques have shifted less. So maybe we have to transpose the Basques along with the ancient DNA. That would put the Basques in the correct geographic position. I'm not sure about this, but I think the shift away from Anatolia for certain genetically isolated groups should be pointed out.

      If that were the case, then with transposition, the Basques would begin to nicely line up with the La Brana-Loschbour axis. Luis, I always new you were meant to be a reindeer hunter.

      "You have taken my speculation and bastardized it beyond recognition."

      I haven't done anything, just drawn a few lines on the plot, tried to reconcile the ancient DNA with their actual geographic positions and recent archaeological findings.

      While you're being outraged, I'm going to try to read the details in some of these papers.

      Delete
    4. "... only reindeer hug and thrive on and near glaciers".

      Maybe but, other than mountain areas, there were no glaciers over here even in the LGM. In fact the usual maps of glaciation typically show the approx. max. extension of ice at the LGM, which reached to central England, all Poland and Germany East of the Elbe - also the Alps. Even in the Seine area there were no glaciers, although for what I know it was a arid steppe-tundra environment, not too good for human inhabitation, which concentrated in the loess steppe-tundra ones in the northern latitudes of Europe (Rhine-Danube and Dniepr-Don areas). Maybe the reindeer were particularly able to exploit such dry environments (and surely many others in that cold spell).

      I posted some good quality LGM ice cover maps in this entry: → http://forwhattheywereweare.blogspot.com/2012/03/ice-free-pockets-in-ice-age-scandinavia.html (originally from Don's Maps: http://www.donsmaps.com/icemaps.html).

      In pop culture the Ice Age looks like Antarctica but that's not an accurate notion at all. It should look more like, say, Russia, with greater dominance of an ecosystem now rare known as steppe-tundra (→ http://www.esd.ornl.gov/projects/qen/carbon11.html). This ecosystem was harsh but had variedly degrees of habitability judging from actual human presence:

      1. Dry steppe-tundra (Seine basin, Balcans): almost uninhabited
      2. Loess steppe-tundra (Rhine-Danub, Dniepr-Don): inhabited at low densities
      3. Mixed ecology (Franco-Cantabrian region): densely to very densely inhabited

      Additionally the Mediterranean was dominated by forests (European trees found refuge there), what apparently limited inhabitation, although this one was highest in the LGM at least in Southern Iberia (Gravetto-Solutrean).

      "No. The ancient DNA data needs to be transposed because it hasn't been Neolithically shifted like most of the other samples".

      What?! That's precisely why we can compare it with modern samples. Some are closer and others farther to either WHG or EEF. You get the modern samples and compare aDNA with these (PCA format) or you get some of the aDNA and compare moderns with them (formal tests). The results are similar for the WHG/EEF bipolarity in both procedures.

      "During the Neolithic, most of the other modern DNA samples have genetically shifted toward Greece and Anatolia."

      That is not what EEFs say. Something great about this study and the PCA in particular is that we do have actual ancient farmer DNA (several samples) and they look more like Canarians or Sardinians than anything Anatolian.

      In fact, when compared with WHGs and modern West Asians (no ancient autosomal DNA from West Asia yet), they do not tend to Anatolia but rather Palestinian/Bedouins. Why? The authors used the "Basal Eurasian" hypothesis to explain it but I actually think it has to do with the peculiarities of the Thessalian Neolithic and its African genetic affinity (most obvious in Y-DNA E1b-V13 but maybe also in G2a, which is important in parts of Palestine).

      Whichever explanation you prefer, EEFs do not tend to Anatolia, so the Anatolian tendency of modern SE Europeans (Balcans and Italy) needs of another post-Neolithic explanation (maybe a combo of Halaf-like Grey Pottery "Pelasgians" and Indoeuropean Chalcolithic invaders from Eastern Europe).

      "I say most, because, probably, certain groups such as the Basques have shifted less."

      Not just Basques but also Iberians, Sardinians, Gascons... everything SW European seems still strongly attached to that first EEF wave (with variable WHG admixture).

      To do what you want you would have to, first of all, shift EEFs, who do not show any "Anatolian tendency". But what you actually need to shift is your Anatolian origin paradigm for the European Neolithic, which is probably wrong, at least partly wrong.

      "I haven't done anything"...

      Oh yes, you did! You are just imagining things, very wildly so.

      Delete
    5. Here's an up to date map of Europe at the LGM:
      http://www.diercke.com/kartenansicht.xtp?artId=978-3-14-100790-9&stichwort=modern%20man&fs=1

      Regarding EEF, Sardinians, Iberians, Gascons, yes, the are somewhat isolated, like Basques.

      However, much of the Neolithic came from Anatolia and the Danube (not Sicily).

      "Oh yes, you did! You are just imagining things, very wildly so."

      Must be my hunter-gatherer ancestry, but in any case, I am going to read the reindeer papers.

      Delete
    6. The map looks good but it has no legend anywhere. This may lead to confusion. For example it is important to realize that the blue line going through Budapest and Cologne is not the ice sheet but the permafrost, the ice sheet is represented as a white area with a light blue contour. I'm not sure of what the other colors may mean, although the green zone is obviously the Mediterranean forest of the time.

      "Regarding EEF, Sardinians, Iberians, Gascons, yes, the are somewhat isolated, like Basques".

      They score high in the SW European component, what seems to translate as EEF affinity.

      "However, much of the Neolithic came from Anatolia and the Danube (not Sicily)".

      You don't know what you're talking about. The European Neolithic (barring Eastern Europe and some less important unclear cases) originated in Thessaly from unknown West Asian precursors. Then it split in two groups: one via Epirus to the Adriatic, Italy (incl. Sicily!) and SW Europe (including even parts of Central Europe: La Hoguette) known as Impressed-Cardium Pottery Culture and the other via Macedonia to the Northern Balcans (Red-White Painted Ware) and Central Europe (Linear Pottery Culture, LBK by its German acronym). They have absolutely no known precursor in Anatolia, although it's possible that they back-influenced some Hacilar and Biblos later on.

      I posted links to some reliable maps above:
      → http://context-database.uni-koeln.de/img/P4B.jpg
      → http://context-database.uni-koeln.de/img/P4C.jpg

      As of now the exact West Asian precursor(s) of Thessalian Neolithic (and therefore the bulk of European Neolithic) remains unknown. However the genetics of these peoples, carrying the lineages E1b-V13 and G2a, suggest a partial Palestinian connection, although there may also be some Anatolian element (G2a is also found in Anatolia but only very rarely E1b). This is very apparent in the PCA of this study, as well as in the formal tests performed on EEF aDNA, which suggest some African affinity (what is also apparent in their tight clustering with modern Canarians, who have ~20% North African ancestry).

      In my understanding this is best explained by Egyptian-like genetic influence via Palestine, possibly followed by some sort of coastal (or maybe even high seas) migration to Thessaly, far away from the known Neolithic centers of the time.

      Delete
    7. Luis, it doesn't really matter whether the transposition is done because of hypothesized admixture with Thessalians or Anatolians. The fact is that Loschbour man is buried in Luxembourg in front of his cave. He appears to be related to the La Brana hunter gatherers. (We don't really know yet. More ancient DNA would help.)

      We have to connect Loschbour Man's DNA with his cave. Therefore, we have to transpose the Loschbour-La Brana vector northward a bit. Maybe with more ancient DNA, including some from Dordogne reindeer hunters, we might be able to add to this picture.

      Regarding this:

      "As of now the exact West Asian precursor(s) of Thessalian Neolithic (and therefore the bulk of European Neolithic) remains unknown. However the genetics of these peoples, carrying the lineages E1b-V13 and G2a, suggest a partial Palestinian connection, although there may also be some Anatolian element (G2a is also found in Anatolia but only very rarely E1b)."

      I don't really agree with this. The connection of some parts of what is modern Greece to points east is obviously through Cyprus. However, regarding Epirus, I'm convinced that it is probably more connected to EEF than Cyprus.

      However, I am not going to debate this with you. My husband's parents are in fact Western Macedonian/Epirotes. I've looked into the matter extensively If they partly cluster with Canarians, it is because they cluster with North Africa, just like EEF. You know . . . the swimming Sicilian/Tunisian Ice Age cows.

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    8. How does it matter where Lochsbour man was buried or lived?

      "... it doesn't really matter whether the transposition is done because of hypothesized admixture with Thessalians or Anatolians."

      How can it not matter? We are seeing the EEFs genetic signature and, unlike the WHGs, they all look very similar among them. They do not tend to Anatolia.

      "The connection of some parts of what is modern Greece to points east is obviously through Cyprus".

      I don't see how it is "obvious" nor how modern Greeks can inform us of early Neolithic peoples better than EEFs. Greece suffered at least two other immigrant flows after Neolithic (and before recorded history): the Gray Pottery culture (Dimini-Vinca, apparently from the Upper Euphrates) and the properly Greek (IE) conquest in the Bronze Age. These or similar flows affected to all the Balcans and had echoes in Italy.

      "However, regarding Epirus, I'm convinced that it is probably more connected to EEF than Cyprus".

      Epirus is at the origin of Impressed-Cardium Pottery (after Otzaki). It has no relation I know of with the continental wave. The original knot was in Thessaly.

      "My husband's parents are in fact Western Macedonian/Epirotes."

      I don't see how this is relevant.

      " I've looked into the matter extensively If they partly cluster with Canarians, it is because they cluster with North Africa, just like EEF".

      Well, that was exactly what I am proposing all the time, at least for EEF (not sure about modern Epirotes: Albanians and Greeks do not cluster with Canarians nor EEFs).

      "You know . . . the swimming Sicilian/Tunisian Ice Age cows. "

      WTF?!

      No, I don't know and I suspect I do not want to know. Swimming Ice Age cows?!

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    9. I went back to the Dodecad Admixture data to try to figure this out.

      Have a look. It's on my blog.

      From looking at the Admixture bar graphs, I've come to several conclusions:

      1. The La Brana/Loschbour Hunter Gatherers probably had some Early European Farmer DNA. That's what is pulling their vector downward compared to the Motala Hunter Gatherer vector. If that's the case, then all of these ancient DNA samples have "Scandinavian Hunter Gatherer" ancestry. However the La Brana and Loschbour samples also have some "Early European Farmer" DNA.

      2. Basques and Sardinians are shifted downward and leftward compared most of the other European pops because they lack "West Asian" component ancestry.

      3. It does appear that Early European Farmers came from North Africa. I recently found a map (see the first map in the maps section of my blog) which indicates that the crossing could easily have been made from Tunisia to Sicily during the LGM. Gibraltar was also a very likely crossing point.

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    10. "1. The La Brana/Loschbour Hunter Gatherers probably had some Early European Farmer DNA."

      It's the other way around EEFs were a mixture of non-Europeans (West Asians but with some African tendency, it seems) and WHGs.

      "Basques and Sardinians are shifted downward and leftward compared most of the other European pops because they lack "West Asian" component ancestry".

      That cannot be true, certainly not for Sardinians. If you make a K=2 West Eurasian ADMIXTURE analysis (example: http://forwhattheywereweare.blogspot.com/2011/12/playing-around-with-admixture.html), you always get some West Asian affinity in every European population (almost zero in Lithuanians, followed by other Eastern Europeans, Basques and Orcadians, in this order). Sardinians however have more West Asian affinity than French (the same HGDP Lyon sample) and every other European, excepted Italians and some Balcanic peoples.

      Sardinians however have more North African affinity. I'm using this K=4 run which clearly dissects West Asian (red), European (purple), North African (green) and Siberian (light blue) components.

      Among Europeans only Sardinians and Spaniards have significant North African affinity (still quite low). In West Asia instead we see some noticeable differences, with Palestinians scoring highest for North African affinity, followed by Syrians. On the opposite extreme, Kurds, Turks and Caucasians score low in this aspect.

      "It does appear that Early European Farmers came from North Africa".

      I am not saying that. What I think is that they came from West Asia but carrying a relatively important North African affinity element (of Epipaleolithic origins). The best match for this are Palestinians, whose relative West Asian (red) and North African (green) components match very well those of Sardinians (after the necessary scaling).

      "I recently found a map (see the first map in the maps section of my blog) which indicates that the crossing could easily have been made from Tunisia to Sicily during the LGM".

      That map (which takes over my computer each time I look at it) is most probably incorrect for all I know, while it's true that Sicily was larger, it was not quite as large nor so close to Tunisia. The crossing was still quite a challenge compared with Gibraltar or the Bosporus/Dardanelles and in any case, there is no archaeological evidence of such crossing ever happening.

      Italy was intermediate between the Balcans and SW Europe in the Neolithic but the origins of European Neolithic lay in Greece, not in Africa, nor Italy nor even West Asia in any clear way (generically yes but not in any discernible level of detail).

      These first Greek farmers would probably be located a bit to the right of Stuttgart and Ötzi in the PCA, quite lower in the dimension 2 than modern Greeks. This means that most likely modern Greeks and Italians have been significantly affected by post-Neolithic flows, unlike Sardinians and Basques (and also to lesser extent Spaniards and Gascons).

      These post-Neolithic flows surely had two origins: (1) Grey Pottery tending to Turkey and (2) Indoeuropean (Kurgans) tending to Eastern Europe and ANE.

      ...

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    11. ...

      "Gibraltar was also a very likely crossing point".

      This one is archaeologically (and genetically) documented: (1) crossing of Iberians to North Africa (with major genetic impact, still apparent today) in the LGM (Oranian genesis), including a probable minor back-flow with greatest impact in West; (2) "bounce" of the Cardium Pottery migration in North Africa with plausible impact in West Iberia (alternative or complementary to #1). However the impact in Europe by this route is quite minor (even in West Iberia it does not exceed ~10% anywhere).

      In addition to these, there are still unclear issues on the origin and dating of the first(?) SW Iberian Neolithic (La Almagra pottery, earliest dolmens). Some like Zilhao argue that these are mere Epicardial, while Spanish scholars instead still think that the available data tells of older dates for La Almagra than for Cardial, at least in Andalusia. The oldest known dolmens of Portugal are also slightly older than Cardial. The impact of Cardial in all this area of South and SW Iberia was relatively minor and often found with locally rooted Epimagdalenian toolkits. All this might be related to North African Capsian Neolithic or maybe just to the adoption by local aborigines of Neolithic elements by "contact" influences rather than immigration. It's a very confusing issue in any case.

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    12. Clearly, more ancient DNA is needed to comb out some of these details.

      Delete
  13. Hi Luis,

    Gotta go to work, but will respond tonight.

    :)

    Marnie

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  14. Luis, could you test my -110 meter world map?

    If you hit +, you should be able to zoom in.

    - zooms you out.

    You should be able to pan with the arrow keys.

    Thank you. I would appreciate it as I would like to make sure that this map is easily readable.

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    Replies
    1. I don't know what exactly you want me to "test". I have problems loading such a heavy image but there are resources showing the actual sea depths, for example: http://cmtt.tori.org.tw/data/App_map/maplist.htm

      Try opening the Mediterranean Sea map (the gray line marks the 100 m depth, there are scales on top of the page).

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    2. Actually the gray line marks 200m. The blue one marks 50 m. and the black one 2000 m. All the rest has to be read with color gradation.

      So your map seems to reflect a 200m. depth or more, what is very unreal.

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    3. The map I have on my sight is 3.4 MB. Not huge, but admittedly, maybe a little slow to download.

      I've checked the map against other sources for sea level. I think the NOAA map might be a little aggressive in terms of stating how low the sea level fell. However, it approximately matches sea level data I've seen calculated for Japan, which was estimated independently and not using the Red Sea or ice core data.

      There are some other effects, such as glacial rebound, which make it difficult to calculate the sea level.

      Since we are interested here in a discussion about the Strait of Sicily, see another discussion here:

      http://www.maxrap.it/atlantide/geologia/canaleE.htm

      Delete
  15. OK. I'm sorry that the image is too big to load. I'll put a high def and low def version in the maps section of my blog.

    Thanks for telling me.

    ReplyDelete
  16. This comment has been removed by the author.

    ReplyDelete
  17. Davidksi wrote: "ENA/ANE = Uralics"

    And Maju replied: "That's the worst simplification of all. Obviously Uralics are a complex issue and West of the Urals they must have been growingly Europanized since very early. It is notable that the most distinctive marker of Indoeuropean-like autosomal influence in Europe peaks in and around Finland. It may be an artifact or whatever but all analysis since the long gone times of Cavalli-Sforza show that. Is that ANE? Probably not because ANE is quite homogeneous in all European populations. Is it related? Surely but rather obliquely. "

    I must say I agree with Maju here. The Siberian component in Northeastern Europe actually peaks in the Arctic region, not in the Uralic Volga-Ural urheimat. It is highest in Norweigan Saami, and it is also high along the whole Arctic coast, whether Slavic- or Uralic speaking. Siberian-ness is simply something that has been native to the region for quite a long time.

    There has probably been many influx of ENA into Eastern Europe, but I think it would be a huge sensation if any of them could actually be connected to any linguistic expansion. The most plausible I think is that the Uralics were similar to their PIE-speaking neighbors, since the families nearly form a genetic unit. Of course, there might not have been an expanding Uralic people, the family might have spread as a trade language among unrelated people also.

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    1. I must say that Arctic Russians are quite obviously of Uralic origins (just look at their genetics and also at the history of the Russian conquest of that area, very recent). So we are not so much in agreement in fact.

      What I think is that the ANE samples reflect in any case a pre-Uralic population of the central steppe, while Uralics actually spread (from a NE Asian origin) via the taiga after the LGM. Ancient DNA shows that while we can spot East Asian lineages in the taiga or even in the European steppe (Ukraine) since very early (Epipaleolithi or Neolithic), the Siberian steppe was apparently impervious to those advances until the Iron Age.

      "Uralics were similar to their PIE-speaking neighbors, since the families nearly form a genetic unit".

      Actually the most likely correct interpretation of the evidence behind the "Indo-Uralic" hypothesis is that early IEs and early Uralics were in close contact and exchanged words, rather than a phylogenetic common origin.

      In terms of population genetics they also appear rather distinct. Although there has been some intense admixture through the ages, the main effect was the "Europeanization" of Uralics because, as I explained elsewhere, a balanced and sustained exchange of population (mostly women) among two neighbor peoples along a S-N cline, causes much greater effect in those to the North, because these have much lower population densities. In a numerical example:

      Let Ne(north)=10 and Ne(south)=100. Let the generational population exchange be 1 in each direction. The effect in the Northern population is 10%, while in the southern population it is only of 1%. The effect is of course accumulative through generations. As most of this exchange was done via women (patrilocality), we can still find notable amounts of yDNA N1 among Uralics (rare among others) but their mtDNA and autosomal DNA has become very much European. In exchange non-Uralics of Eastern Europe (and Scandinavia to some extent too) have got some Uralic (East Asian and Siberian) influence in their genomes but to a much lesser extent. These almost certainly spread those elements to other parts of Europe in even more diluted amounts.

      It is difficult to discern what role plays ANE in all this. It was surely associated to some degree (peaks in Estonians and Hungarians) but its correlation with Uralics is not straightforward nor linear and there were almost certainly other previous influences affecting Europe somehow.

      Whatever the case ANE is relatively non-important in modern Europeans and at least two populations (Basques and Sardinians) can be explained without appealing to ANE (i.e. as simple bipolar mix of WHG and EEF). Assuming that the Basque 10% ANE affinity is basal (Sardinian "negative" affinity can be explained by other factors, such as greater distance to ANE of EEF), then the 13% ANE affinity of Iberians, etc. should be read as 15% "NE European" (IE) admixture maybe, while the 18% of Estonians should be read as extra 40% "NE European" (Uralic mostly in this case) admixture, relative to WHG (Lochsbour, which I suspect not the best proxy in fact).

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    2. "I must say that Arctic Russians are quite obviously of Uralic origins (just look at their genetics and also at the history of the Russian conquest of that area, very recent)."

      Expansion of Finns beyond Southern Finland is maybe even more recent. My point was, that the Siberian component is high throughout the Arctic region regardless of ethnicity and peaks along the Arctic coast. It seems to be an indigenous component of the region. I have really hard to see how anyone could argue that it migrated from the Volga-Ural region, where there is a different Siberian component which is also much smaller.

      "Actually the most likely correct interpretation of the evidence behind the "Indo-Uralic" hypothesis is that early IEs and early Uralics were in close contact and exchanged words, rather than a phylogenetic common origin."

      But a phylogenetic common origin cannot be ruled out either, since there is some shared core vocabulary, such as the word for "name". So is there any special reason to assume an Eastern origin?

      N in Eastern Europe could be the result of any influx, connecting it to the Uralic languages is just speculation. Besides I think it seems to old and widespread to be caused by some hypothetical Uralic expansion. For example it is very common here in Sweden, where there are no attested Uralic languages except Saami in the North. So I must ask again, is there any special reason to connect entry of N to the Uralic languages?

      I'm not making any conclusions myself either way, I just wonder why only one side of the scientific debate is seen as correct.

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    3. "My point was, that the Siberian component is high throughout the Arctic region regardless of ethnicity and peaks along the Arctic coast. It seems to be an indigenous component of the region. I have really hard to see how anyone could argue that it migrated from the Volga-Ural region, where there is a different Siberian component which is also much smaller".

      Well:

      1. Most of that area was under the Ice until "recently", so "indigenous" cannot be really old.

      2. Kristiina (a regular commenter at this blog with a keen interest on Uralic prehistory) thinks (if I understand her correctly) that there may have been two related but different migration patterns: one more strictly arctic and another through more temperate zones. But I'm uncertain how solid this is.

      3. Alternatively (or complementarily) the Uralic peoples (incl. Northern Russians, who are Uralic by genetics) may simply show a gradient of admixture with the peoples south of them, that way the most northernly populations would naturally preserve better the general ancestral stock, while the southermost ones would be much more admixed, even almost beyond recognition in many aspects.

      "But a phylogenetic common origin cannot be ruled out either"...

      That's a matter I don't feel able to discuss but for all I read most experts favor mere contact. Mere contact and more or less intense sporadic domination can cause major linguistic changes: Brahui only retains a 15% Dravidian vocabulary, English only 25% Anglosaxon, Basque is full of Latin/Romance loanwords for "core" concepts like "beech" or "mountain", while nearby romances have Basque loanwords (or substrate words) for also important concepts like "left" or "kid/small". That's not a valid argumentation alone.

      "So is there any special reason to assume an Eastern origin?"

      I do not assume any origin for the Uralic linguistic family (if that's what you mean) but I would not be surprised if an Eastern origin is someday proposed for Uralic. Other possibilities can also be valid. I just don't know.

      What is clear is that the Uralic macro-ethnicity originates in Late Paleolithic and Neolithic processes associated to the patrilocal expansion of yDNA N1 from NE Asia. This process can be compared (with a reverse geography in the W-E axis) to the older genesis of proto-Amerindians (in that case "lead" by patrilineage Q1 from Central Asia).

      "N in Eastern Europe could be the result of any influx, connecting it to the Uralic languages is just speculation".

      Speculation? It's clear like water. It does not necessarily mean that the proto-Uralic language originated in East Asia (it could also have been picked "on the road" for whatever reason) but it is a plausible hypothesis.

      "For example it is very common here in Sweden, where there are no attested Uralic languages except Saami in the North".

      I do not understand well this sentence (seems to be ill-constructed) but it seems to me that the Sámi were much more widespread in the past, occupying much of Scandinavia, although not the southernmost parts.

      "So I must ask again, is there any special reason to connect entry of N to the Uralic languages?"

      The main reason is that N (N1c and N1b) is almost only found among Uralic peoples in Europe, often at very high frequencies. The correlation should be obvious only on that. But also both categories (the ethnolinguistic "Uralic" and the patrilineal genetic "N1") have a "Far North" specialization, although that of yDNA N is nowadays also associated to some non-Uralic speakers (in East Asia only) - this however may have been caused by acculturation processes. In West Siberia and Europe N1 is very strictly correlated with Uralic peoples.

      "... I just wonder why only one side of the scientific debate is seen as correct".

      Which is the other side?

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    4. "1. Most of that area was under the Ice until "recently", so "indigenous" cannot be really old."

      Is that necessarily true?

      Say you have a landmass with three latitudinal ecozones: A = warm, B = middle, C = cold, in a sequence south to north, each ecozone with a different primary herd animal and consequently three different human populations adapted to those primary herd animals. When the ice expands it doesn't just overwrite band C and the people that lived on band C - it changes the extents of all three bands i.e. band A shrinks in the north, band B expands to the south and shrinks in the north, band C shrinks in the north and expands to the south. As the three bands expand and contract their prospective herd animals and human predators expand and contracy also. So the original indigenous population in band C might well still survive just pushed south a ways (and possibly admixing with B a bit in the contraction process). Then when the ice retreats the same population moves back where it was.

      There's no guarantee obviously but it seems at least possible that in some cases populations that were pushed back by the ice could just as easily have moved back again after the ice retreated and therefore effectively be continuous.

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    5. It's more complex than that. For example at the end of the Ice Age there was an expansion of forests and then there was a sudden cold spell (arguably caused by a meteorite falling on North America) known as Younger Dryas, which froze Ireland (again) in just months. Later the climate improved again...

      But, whatever the complexity of climatic change, we can track archaeological cultures and in general terms Epipaleolithic cultures derived from Magdalenian stayed in the Magdalenian zone, while most of the expansion northwards (in Western Europe) was led by people of Hamburgian culture instead (and maybe others), the people who already lived at the edge of the ice sheet, so to say. I understand that all this means that it's rather easy to adapt to warmer climates and also that, in general terms, the expansion in the newly defrosted areas was starred by people already specialized in the coldest econiche. This was surely the case of proto-Uralics in Eastern Eurasia and the overall tendency of their expansion (judging on the spread of yDNA N) was then westwards following the retreating ice or permafrost (remember that the main ice sheet in Eurasia was right on Fenno-Scandia).

      Another issue which I do not have fully clear is to which extent these proto-Uralic populations expanding westwards through the taiga may have genetically influenced the already existing populations further south in Eastern Europe. Recent aDNA from Ukraine found significant amounts of East Asian lineages in the Neolithic of that area, lineages that probably echo the proto-Uralic spread in the Far North.

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    6. You strongly suggest that the N1c expansion is due to Uralic languages, but does not seem to be the case. N1c predates Uralic languages in Europe by almost 2000 years.
      http://www.elisanet.fi/alkupera/N1c1.pdf
      As I said the Swedish branch might be 4500 years old, and cannot be explained Uralic expansion. The author above proposes Comb-Ceramic as a source of the N1c expansion, and well that is also speculation, but a lot more plausible.
      (Of course Uralic speakers might have helped further spreading N in an already N-rich terrirtory!)

      Core vocabulary can be borrowed yes, but PU and PIE have their respective Urheimat next to each other, which is I guess why so many support a common phylogenetic origin. I'm just saying, it is a possibility which also has to be taken into account.

      "Alternatively (or complementarily) the Uralic peoples (incl. Northern Russians, who are Uralic by genetics) may simply show a gradient of admixture with the peoples south of them, that way the most northernly populations would naturally preserve better the general ancestral stock, while the southermost ones would be much more admixed, even almost beyond recognition in many aspects."

      Finns, Saami and Estonians all stem from the same Baltic population, as evident by linguistics. Saami, who migrated the farthest North, have gotten a significant Siberian admixture, which Finns have gotten a little of in turn, as they migrated from Estonia to Finland. But the Estonians remained largely non-ENA. Does this not point to the fact that the "source" was non-ENA? Is it not more plausible?

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    7. I would say that there is some ample consensus on Comb Ceramic being directly related to Uralic expansion, so I do not really understand what you say. Are you suggesting that Uralic languages in Europe only have 2000 years of age? I would think instead that they are at least as old as Indoeuropean (c. 6000 years), if not even older.

      "Saami, who migrated the farthest North, have gotten a significant Siberian admixture, which Finns have gotten a little of in turn, as they migrated from Estonia to Finland".

      I don't dare to judge in detail the exact processes leading to the formation of these populations because there may be "internal" Uralic processes of displacement/partial replacement that I do not really understand. However in general terms, I'd say that the Finnic group overall must have originated with the arrival of Combed Pottery to the area. Maybe, as happened with Berbers in NW Africa, there were processes of linguistic homogenization and/or expansion that displaced older branches (I don't know in the case of Finnic) but the ultimate origin should be in Combed Pottery.

      "But the Estonians remained largely non-ENA."

      I'm not sure how the ENA component enters the Uralic equation but what is clear is that both Estonians and Hungarians, the only two Uralic-speaking populations considered in the relevant study, have record levels of the ENA component in Europe, even more than Russians. Said that, we must understand that ENA was a Southern Siberian component of West Eurasian affinity (origins) and that, therefore, it is fundamentally different from whatever East Asian elements brought by proto-Uralics (and/or whatever carriers of yDNA N1 you want to imagine) via the taiga (further North).

      Estonians and in general European Uralics have lost in all this time most of their East Asian (or Siberian) genetics (excepting the yDNA, almost certainly because of patrilocality), much as proto-Amerindians lost almost all their West Eurasian affinity by admixture with East Asians even if they keep the West Eurasian patrilineage Q1. I believe it is important to make this distinction between an obvious "exotic" signature in the patrilineage and the near-loss of this signature in other genetic aspects (mtDNA, autosomal): it tells us a lot about the Uralic genesis (or the proto-Amerindian one also).

      "Does this not point to the fact that the "source" was non-ENA? Is it not more plausible?"

      I'm not sure what you have in mind but I'm not saying that the proto-Uralic source was ENA, just that it seems that they had a more intense contact with ENA than other Europeans, almost certainly because of their Siberian journey from East Asia. It's possible that proto-IEs of the Volga (and maybe even in Ukraine) also had contacts with ENA on their own terms (and/or via the proto-Uralics), what seems apparent is that, in Europe, the level of ENA affinity is related to the inferred level of IE and Uralic influence (both).

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    8. PS.-

      I just wrote: "... in general terms, I'd say that the Finnic group overall must have originated with the arrival of Combed Pottery to the area".

      I must add that this does not detract from possible even older Uralic (or Uralic-like or proto-Uralic) arrivals to the Finnic area (as well as to other parts of Eastern Europe) because aDNA shows some presence of East Asian mtDNA (haplogroup C) since Epipaleolithic in the area of Karelia/NW Russia and since Neolithic in Ukraine.

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    9. There are two things I strongly disagree with you on:

      1. "I would say that there is some ample consensus on Comb Ceramic being directly related to Uralic expansion"

      First, all archaeologists were always sceptical about correlating Comb Ceramic with any languages, and second, any correlation between Comb Ceramic and Uralic languages have long ago been abandoned.

      2. "almost certainly because of their Siberian journey from East Asia"

      What points to a journey from East Asia in the first place? Why make such an assumption? Y-DNA cannot be an evidence, as it is never any evidence in itself, and N1c is also older in Europe than the Uralic languages. There is certainly no linguistic evidence either.

      Clearly there must have been some migration from East Asia to Europe, connected to N1c and perhaps the Comb-Ceramic culture, but the connection to Uralic languages is just pure speculation. In the absence of evidence, pre-proto-Uralic speakers could have come from pretty much anywhere.

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    10. Well, I cannot really understand why would you reject these correlations: it is obvious that Uralic human geography strongly correlates with yDNA N1 (N1c and N1b in essence) and with Combed Ceramic. I cannot accept this as a mere coincidence unless you'd present strong evidence (extraordinary claims require extraordinary evidence), which you do not. You don't even articulate any half-reasonable alternative hypothesis on how would "Uralic peoples" acquired so much yDNA N1 or how the N1 peoples acquired Uralic language, which you seem to consider unrelated for some unclear reason.

      Lacking all that, I'm merely applying Occam's Razor: the most parsimonious explanation is most likely the correct one. Occam's Razor is not "pure speculation" but a quite reasonable method producing the default hypothesis and the best possible theory barring strong evidence on an opposite direction (lacking so far) and a time machine to check the facts on the ground.

      I'm anyhow a bit surprised by the ferocity of your denial, which I had never before encountered. But in wait of an alternative hypothesis (not articulated) and sufficient evidence in support of it (not presented), I can only shrug off that animosity against the most parsimonious theory.

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    11. Additionally, as you seem to base all your ideas on mere linguistic materials, it is worth mentioning that proto-Uralic clearly has many proto-Indoeuropean loanwords. PIE only existed up to c. 6000 years ago, so proto-Uralic must be of about that same age or older (and not a ridiculously short 2K age). This fits well with the Comb Ware culture as vector of Uralic expansions, without exclusion of possible previous (proto-Uralic?) waves from a similar origin, which can be associated to the presence of mtDNA C in various parts of Eastern Europe since Epipaleolithic/Neolithic.

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    12. Also, I am out of curiosity reviewing the general outlook of linguistic theories on Uralic and possible upstream connections (hypothetical macro-families) and all I find other than Indo-Uralic (which is generally understood as mere sprachbund) are theories of connection with NE Asian languages: Ural-Altaic, Uralo-Siberian, Eskimo-Uralic and Uralic-Ykaghir.

      Of these the latter seems the most widely accepted one, with very good arguments. Citing from Wikipedia's quote of Collinder (1965): "The features common to Yukagir and Uralic are so numerous and so characteristic that they must be remainders of a primordial unity".

      I don't intend to dive too deep into the linguistic waters, much less to take any strong stand on the possible origins of proto-Uralic but it is clear that this set of more or less well informed hypothesis all look to the East and not to West Eurasia. I find particularly intriguing this table of pronoun correspondences between Uralic and Altaic languages, which seem outstanding, especially considering that they should not have been in direct contact before the Huns. Unless we accept the ultimate NE Asian origin of the proto-Uralic yDNA N1 carriers, something that ancient DNA seems to support (N was dominant in Northern China in the Neolithic at least) there is no explanation for all these "coincidences".

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    13. The similarities between PU and PIE are still much, much greater than between PU and any other reconstructed language. Also when it comes to pronouns! These similarities are also deeper, concerning core vocabulary and morphology. The similarities with Altaic languages are mostly typological and poor proof of a phylogenetic origin. That is definitly the mainstream opinion today. (The small Yukagir family is an exception here, it might be an old branch of Samoyedic which has migrated further East)

      About N1c, what can I say, it is older than the arrival of Uralic languages. Surely PU is old but Uralic languages are quite recent arrivals in Finland and the Baltic. And there are also the old Swedish and Polish branshes of N1c, clearly non-Uralic. N1c can, like I said, perhaps be connected to the Comb-Ceramic culture. Uralic languages then started spreading within that N1c context. In any case the Comb-Ceramic culture certainly arrived in Finland before the Finnish language, and I see no reason to assume that there was an unknown, dead branch of Uralic spoken there before. (The article I linked deals with this, although it is not scientific)

      This is all mainstream archeology in the Nordic countries as far as I can see, and if it's new to you, then perhaps it is because it is such a narrow field which is given little attention outside the Nordic countries.

      And again I am not "denying" anything - merely pointing out that there are several theories, some more likely than others.

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    14. "About N1c, what can I say, it is older than the arrival of Uralic languages."

      Estimates on the age of the Uralic family vary and I believe that the oldest ones (c. 7 Ka ago) are more realistic, considering the many PIE loanwords.

      "Uralic languages are quite recent arrivals in Finland and the Baltic".

      How do you know?

      "And there are also the old Swedish and Polish branshes of N1c"

      Not familiar with the matter but in any case they would seem to imply much less important numbers. It's probable that those lineages arrived from Eastern Europe in the case of Poland at least, while in Sweden it may be the same wave (either Pitted Ware or Kurgans) or incorporated from proto-Sámi, who probably occupied much of Scandinavia not so long ago.

      "I see no reason to assume that there was an unknown, dead branch of Uralic spoken there before".

      I'm speculating on this aspect of course (mostly extrapolating from what is believed to have happened with Berber languages of NW Africa) but, in any case, I see no other processes implicated in the spread of either Uralic languages nor yDNA N1. You contend with all this but you put forward no clear alternative theory, much less factual data that could support it.

      "The article I linked deals with this"...

      I must apologize here because I missed that link, sorry. Now that I look at it however, it is clear that the oldest distinct branches of N1c are the so called "Asian" and "Siberian" ones, so it should be obvious that N1c comes from that part of the World (also supported by upstream N presence). Downstream it splits into Central and Eastern European branches, so maybe it is correct that N1c-L550 has a distribution different from that of the much more common Finnic N1c-L132 but... how common is the Central European branch? Judging from the Spanish subclade, it must have almost negligible presence (mere erratics). How did this branch coalesce? I do not know but maybe it has something to do with Eastern Asian mtDNA in Ukraine and later waves (Pitted Ware first and Kurgans later) from there westward. Do these "Central" lineages have any presence among Uralics? I do not see that specified in the paper and is clearly a key point.

      In brief too many doubts and several reasonable possibilities that still connect N1c overall with Uralics (not just Finnics, of course).

      "This is all mainstream archeology in the Nordic countries as far as I can see, and if it's new to you, then perhaps it is because it is such a narrow field which is given little attention outside the Nordic countries".

      Possibly but I have as custom to discuss with many people and drink from many sources, including some Finnish ones, and I am all but familiar with your ideas, so it'd be interesting if you could expand on them with links and/or maybe some articles in your blog. I'm totally amiss about possible post-Comb Ceramic Finnic immigrations to that area. Anyhow there are often several competing theories, I'm at the very least familiar with some "popular" Spanish theories that are total nonsense (yet supported by some "respectable" academics), so I would appreciate a deep immersion on the matter, really.

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    15. Of course N has its origins in Siberia, that is an indisputable fact. Its spread to Europe is much older than the spread of Uralic languages, though. Here is for example a map by the same author of the spread of the Finnish language:
      http://www.elisanet.fi/alkupera/Suomensynty.html
      Just one linguist's summary, but within the mainstream science there should not be big differences.

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    16. N has its origins in East Asia (→ http://forwhattheywereweare.blogspot.com/2013/07/a-review-of-haplogroup-n-y-dna.html), most likely somewhere between Central and South China (Yunnan is a good candidate). Some branches of N1 may have coalesced already in Siberia (Late UP probably).

      As for your link, I do not find it convincing, nonetheless because it only seems to deal with Finnic (and not Uralic overall). Even if this author would be correct and Estonian-Finnish spread as he says, they must have replaced older Uralic languages in the area, maybe of the Sámi branch. No plausible archaeological record is associated with these purely linguistic speculations (linguistics is often plagued by arbitrariness and extreme uncertainty, so it needs of some material support).

      I ask again: which is the material culture, the archaeology that could be related to this speculation?

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    17. As for those similarities between Turkic and Uralic personal pronouns, we should compare Turkic, Uralic and IE personal pronous. Please see below:
      Turkish ben sen o biz siz onlar
      Mongolian bi chi ter bid ta ted
      Persian man to u (human)/ân (non-human) mâ šomā ânhâ
      Russian ja tyj on (m)/ ona (f) myj vyj oni
      Finnish minä sinä hän (human)/se (non-human) me te he (human)/ne (non-human)
      Hungarian én te ő mi ti ők
      Komi me тэ cijo mi ti najo
      Mansi amki naŋki takʷi ma:nki na:nki ta:nki
      Nenets məny pidər pida mənyaq pidraq pidoh

      Russian ”ja” looks different from the rest, ”vyj” as well. Finnish ”hän” and ”he” look different from the rest. Mansi and Nenets forms look different from other Uralic pronouns. The common forms seem to be: m/b for 1. person sing.&plural., t/s for 2. person sing&plural., n/t/s for 3. person sing.&plural.

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  18. Davidksi wrote: "ENA/ANE = Uralics"

    And Maju replied: "That's the worst simplification of all. Obviously Uralics are a complex issue and West of the Urals they must have been growingly Europanized since very early. It is notable that the most distinctive marker of Indoeuropean-like autosomal influence in Europe peaks in and around Finland. It may be an artifact or whatever but all analysis since the long gone times of Cavalli-Sforza show that. Is that ANE? Probably not because ANE is quite homogeneous in all European populations. Is it related? Surely but rather obliquely. "

    I must say I agree with Maju here. The Siberian component in Northeastern Europe actually peaks in the Arctic region, not in the Uralic Volga-Ural urheimat. It is highest in Norweigan Saami, and it is also high along the whole Arctic coast, whether Slavic- or Uralic speaking. Siberian-ness is simply something which has been native to the region for a long time. Obviously there has been many influx of ENA genes into Europe, but I think it would be a sensation if any could actually be connected to linguistic expansion.

    There are many opinions on the Uralic origins - the most extreme are at one end is that Uralic formed a genetic family with Indo-European (Indo-Uralic). The most extreme on the other end is that Uralic migrated from the East and only acquired later loan words from PIE. Most linguists argue for a compromise, however. Uralic was a group close to PIE with very early loan words.

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    1. "The Siberian component in Northeastern Europe actually peaks in the Arctic region, not in the Uralic Volga-Ural urheimat. It is highest in Norweigan Saami, and it is also high along the whole Arctic coast, whether Slavic- or Uralic speaking. Siberian-ness is simply something which has been native to the region for a long time. Obviously there has been many influx of ENA genes into Europe, but I think it would be a sensation if any could actually be connected to linguistic expansion."

      I think it's worth thinking in terms of climate zones / ecozones. If the steppe and far north was divided into mostly horizontal ecological slices there's no reason why a rapid east-west expansion in one slice should necessarily have the same effect in the other slices as people adapted for one slice aren't necessarily adapted for the other slices e.g. Mongols and Inuit.

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  19. Maju,

    What I meant above was that SE Europeans just before the Neolithic were very likely different from other Europeans, while there is little reason to believe they were much different from people in W Anatolia (excluding some clines, obviously).

    During the Ice Age, not just LGM, it was difficult for people to move between NW Italy and SE France (~San Remo), and it was also difficult to cross the Carpathians - both of which must have substantially limited gene flow. And we see the outcome of that culturally in the unified Epigravettian culture of Italy, Greece, and the Balkans up to the Black Sea (that was different from Magdalenien).

    Conversely, there was no geographic barrier between Thrace and NW Anatolia, and we know that the Greek islands were populated by very good seafarers that had trade with Anatolia (and probably the Northernmost Levant).

    A good example are the layers of the Franchthi Cave, which show a succession of imported and then locally-grown "crop" items, such as fruits, nuts, legumes, and finally cereal, from ~ SW Anatolia/ N Levant(?) (given their likely geographic origin). This exemplifies a longstanding maritime trade connection and an actual route for the introduction of agriculture, which does not necessitate migration of (a substantial number of) people.

    So, what I am saying is that there isn't a strong necessity that a lot of new genetic material came to Europe with the initial Neolithic, but that it (largely) was already there, in SE Europe (in particular, G2a).

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    1. "What I meant above was that SE Europeans just before the Neolithic were very likely different from other Europeans, while there is little reason to believe they were much different from people in W Anatolia (excluding some clines, obviously)".

      We know very little about the Paleolithic Balcans, apparently not the most desired place to live back then. It is reasonably to assume that it was sparsely populated, judging on the very low frequency of findings. Instead we do know something about the earliest Neolithic peoples from the Balcans (from indirect sources): that they carried yDNA G2a, E1b-V13 and also I2a. Of these lineages only G2a can be correlated with Anatolia and not too strongly (Anatolian frequencies are not higher than European ones and certainly much lower than among EEFs).

      Regarding the barriers you mention, they have some important "holes", and very much used for what archaeology can tell: (1) the Viennese corridor by the West and (2) the Wallachian plain by the East. The first one was clearly used in Westward direction by Aurignacoid and Gravettian cultures and in Eastward direction by Solutrean and Magdalenian ones, the latter was surely also used in both directions. The Carpathians themselves were probably also crossed from South to North in the Epipaleolithic (Central European Solutrean-like cultures).

      "Conversely, there was no geographic barrier between Thrace and NW Anatolia"...

      Maybe but we have no evidence of Late Paleolithic or Early Neolithic in either area.

      "... and we know that the Greek islands were populated by very good seafarers that had trade with Anatolia"...

      Yes but from a later period. Anatolian influences in the Cyclades and Crete are detectable, it seems, in the Chalcolithic (and may be related with the distinctive dominance of yDNA J2 in that area).

      "A good example are the layers of the Franchthi Cave, which show a succession of imported and then locally-grown "crop" items, such as fruits, nuts, legumes, and finally cereal"...

      But Franchthi is not in Thessaly but in the Peloponnese. It probably illustrates a somewhat different phenomenon to what happened in Thessaly, where Neolithic shows up in full and soon even with abundant pottery (at a time when pottery was still incipient in West Asia). It's very possible that Franchthi Neolithic was influenced by Thessaly rather than directly from Anatolia or other West Asian sources.

      "So, what I am saying is that there isn't a strong necessity that a lot of new genetic material came to Europe with the initial Neolithic, but that it (largely) was already there, in SE Europe (in particular, G2a)".

      I can't say if G2a and E1b-V13 arrived to Greece in the Mesolithic or in the Neolithic genesis but I think that is most likely that E1b arrived to Palestine only with Mesolithic because it is then when we can see some Egyptian influences in Kebaran and successor cultures, especially in the arid areas of the south. This process of the Levant probably originated the Semitic language family via Harifian, PPNC and the CAPC. Offshoots of this process must have arrived to Thessaly somehow (coastal migration?), otherwise it is very difficult to explain E1b-V13 in Neolithic Europe (with greatest impact in modern Greece and Albania).

      Delete
  20. "We know very little about the Paleolithic Balcans, apparently not the most desired place to live back then."

    I have no idea why you would say this. The Balkans, including the Adriatic/ Po plane and Greece, were one of the most desirable places to live during the Ice Age and continued to be so even during LGM. The Eastern Adriatic/Po plain received substantial rainfall and was mild, with easy access to interior hunting hills and planes. There is evidence of long-sustained summer hunting camps on the SW side of the Pannonian Basin, as expected from climate. Much of low-land Greece was temperate and received much more rainfall than the western portion of the Mediterranean. Similarly the-then widened transition to Anatolia.

    See, e.g.,

    http://www.jstor.org/stable/40960242

    As to the seafarers, my point was that sites such as the Franchthi Cave and early occupation of many relevant islands demonstrate this was a Paleolithic/ Mesolithic practice and success, long before pottery neolithic.

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    1. → http://forwhattheywereweare.blogspot.com/2010/10/revisiting-bocquet-appel-2005.html

      Judging on the archaeological record, the Balcans were nearly deserted in the Ice Ages. A reason may be that they were dominated by arid steppe-tundra (interior) and forest (coasts). The first one was among the less productive habitats and in general shows very low densities (another example is Northern France). The Mediterranean forests were probably more interesting but not the most desired environment either.

      Whatever the reasons, the Balcans were never desert but they were not densely populated either. The prime regions were in this order: 1. Franco-Cantabrian region, 2. Central Europe around Moravia, 3. Central-West Europe (Rhine), 4. Southern parts of Eastern Europe, 5. Iberian peninsula (non-Cantabrian) and 6. Italy. The order may change in some periods (for example Iberia became more important in the LGM and the Rhine area in the Late UP).

      Read the original paper, please, because it is a very important synthetic material:

      → http://www.evolhum.cnrs.fr/bocquet/jas2005.pdf

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    2. Agree.

      Look at the PCA plot that started this conversation, above.

      Greeks (more properly, in their own language, Hellenas) clearly have at least four influences:

      1. Asia Minor
      2. Cyprus
      3. Early Neolithic Farmers (probably by way of Sicily > Adriatic > Epiros > Mantineia
      4. Western European Hunter Gatherers > Pannonian Basin > Upper Macedonian Plain > diffused southward

      I agree that the Hellenic Republic was likely continuously inhabited from the Palaeolithic. However, given its temperate climate, and key location in the Mediterraean, it also continuously absorbed people from surrounding areas.

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    3. I can't agree with you however, Marnie.

      EEFs were an imprecise mixture of early Thessalian farmers and WHGs. Plain and simple. Modern mainland Greeks (Cretans and other islanders may have different origins) seem largely descended from EEFs or something similar to them (maybe less mixed with WHGs than the actual EEF samples from Germany and Italy), i.e. from early Thessalian farmers.

      In addition to that they show a marked Anatolian and East European shift, which may be attributed to secondary flows, not continuous but quite punctual and relatively easy to describe:

      1. Grey Ware migrants from Anatolia/Syria (Halaf related) → Dimini-Vinca (continuous till IE-Greek arrival in the Bronze Age, let's call them "Pelasgians").

      2. Indoeuropean Greeks (Bronze Age).

      Add to that whatever minor "continuous" flows but those are the two main ones. In fact modern Greeks still speak Greek, what says a lot about ethno-cultural continuity.

      These two inflows alone can explain the Greek drift from a reconstructed position just to the right of EEFs to their current position more to the top of the graph.

      Italy may have got less "Pelasgian" influence or EEFs were more mixed with WHGs in Italy/Germany than the original Thessalian farmer population. Maybe both (difficult to discern).

      Cyprus' position is more influenced by Greek immigration since the Bronze Age than for being a direct influence on Greece. Cyprus was settled originally from West Asia and it is still quite similar to Syria/Turkey in its genetic signature. However, like some Turks, they lean towards Europe because of European admixture (Greek in essence).

      In synthesis:

      1. Levant-1 + Egypt → Levant-2
      2. Levant-2 + SE European HGs → Early Thessalian Farmers (ETF)
      3. ETF + WHG → EEF
      4. Eastern European HG + some ANE → IE
      5. EEF + WHG + IE → most modern Europeans (variable apportions)
      6. ETF + "Halaf" + IE → modern Greeks & Albanians
      7. Ancient Cypriots + Greeks → modern Cypriots

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    4. PS- in the synthesis possible relationship between SE European HGs, Eastern European HGs and Western HGs (WHG) are not considered because of uncertainty but in general I would assume they were roughly the same stock.

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    5. We're discussing the above PCA plot, or something else?

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    6. Using it as reference (although in the Cypriot case I also recalled previous studies). I'm also considering the overall data from the paper, notably the various formal tests' results.

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    7. Had a careful look at what you are saying. Started typing out a long message but decided to delete it. I think you are making inferences beyond what the data can support.

      Returning to the subject of the reindeer, have you had a look at the latest post on:

      http://www.aggsbach.de/

      ?

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    8. "... you are making inferences beyond what the data can support".

      I don't think so. Maybe I'm filling in some blanks but is a good fit in the data.

      What I do think is that the data we have, including the choice of references and tests performed in this study, is clearly incomplete. Beside other things I have already mentioned, I just committed the likely error of assuming that "SE European HGs, Eastern European HGs and Western HGs (WHG) (...) were roughly the same stock". Probably I'm oversimplifying there because Solutreo-Magdalenian rooted WHGs and Gravettian-rooted SE and Eastern HGs (several groups) were separated in techno-cultural terms, and hence almost certainly in "stock" (genetics) for almost as long as they were separated from West Asians or Siberian "ANE". So we have yet two big question marks regarding Gravettian tradition European (and Siberian) HGs, something that I was indeed obviating to some extent in my previous comments.

      It is possible that ANE affinity actually just reflects generic Eastern European stock, but at frequencies different than those denoted by direct ANE scores. Lacking Eastern European aDNA samples, we cannot directly know.

      Similarly we lack direct data from ancient SE European (and West Asian) peoples. We can make some inferences but there is indeed some uncertainty.

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    9. "Returning to the subject of the reindeer"...

      Not sure what to say but mostly: "so what?" I really don't understand your obsession with reindeer: for what matters to us, or rather to UP hunter-gatherers, they are just a kind of available foodstuff.

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    10. "obsession with reindeer"

      :)

      I think you should look a little closer into this reindeer thing.

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    11. I don't know if Reindeer are significant but isn't any significance a proxy for a type of ecozone?

      Delete
  21. At work right now. Will respond more this evening (your morning).

    ReplyDelete

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