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Ancient European DNA and some debatable conclusions

26 Dec
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 responses to “Ancient European DNA and some debatable conclusions

  1. Maju

    January 8, 2014 at 9:32 am

    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.

     
  2. Marnie

    January 8, 2014 at 4:35 pm

    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.

     
  3. Maju

    January 8, 2014 at 7:46 pm

    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?!

     
  4. Marnie

    January 9, 2014 at 9:34 am

    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.

     
  5. Maju

    January 9, 2014 at 10:25 am

    “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.

     
  6. Maju

    January 9, 2014 at 10:25 am

    “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.

     
  7. Marnie

    January 9, 2014 at 3:29 pm

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

     
  8. Grey

    January 9, 2014 at 4:02 pm

    “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.

     
  9. Marnie

    January 10, 2014 at 6:45 am

    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.

     
  10. Maju

    January 10, 2014 at 9:28 am

    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).

     
  11. Marnie

    January 10, 2014 at 9:30 am

    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.

     
  12. Maju

    January 10, 2014 at 9:31 am

    This comment has been removed by the author.

     
  13. Maju

    January 10, 2014 at 9:34 am

    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.

     
  14. Gui S

    January 11, 2014 at 11:25 pm

    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.

     
  15. Gui S

    January 11, 2014 at 11:30 pm

    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).

     
  16. Gui S

    January 12, 2014 at 12:17 am

    “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).

     
  17. Gui S

    January 12, 2014 at 1:03 am

    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.

     
  18. Gui S

    January 12, 2014 at 1:03 am

    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…

     
  19. Marnie

    January 12, 2014 at 4:10 am

    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

     
  20. Anders

    January 12, 2014 at 8:46 am

    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.

     
  21. Maju

    January 12, 2014 at 9:03 am

    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.

     
  22. Anders

    January 12, 2014 at 9:17 am

    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.

     
  23. Maju

    January 12, 2014 at 9:33 am

    “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

     
  24. Maju

    January 12, 2014 at 9:41 am

    “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).

     
  25. Maju

    January 12, 2014 at 10:20 am

    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).

     
  26. eurologist

    January 12, 2014 at 2:03 pm

    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).

     
  27. Gui S

    January 12, 2014 at 3:05 pm

    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.

     
  28. Gui S

    January 12, 2014 at 3:20 pm

    “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?

     
  29. Gui S

    January 12, 2014 at 3:26 pm

    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?

     
  30. Maju

    January 12, 2014 at 5:17 pm

    “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).

     
  31. Maju

    January 12, 2014 at 5:31 pm

    “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.

     
  32. Maju

    January 12, 2014 at 5:57 pm

    “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.

     
  33. eurologist

    January 12, 2014 at 6:01 pm

    “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.

     
  34. Maju

    January 12, 2014 at 6:39 pm

    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

     
  35. Marnie

    January 12, 2014 at 7:00 pm

    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.

     
  36. Maju

    January 12, 2014 at 8:38 pm

    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

     
  37. Maju

    January 12, 2014 at 8:42 pm

    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.

     
  38. Marnie

    January 13, 2014 at 2:58 am

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

     
  39. Maju

    January 13, 2014 at 8:38 am

    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.

     
  40. Marnie

    January 13, 2014 at 7:56 pm

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

     
  41. Marnie

    January 14, 2014 at 7:02 am

    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/

    ?

     
  42. Maju

    January 14, 2014 at 10:36 am

    “… 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.

     
  43. Maju

    January 14, 2014 at 10:46 am

    “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.

     
  44. Anders

    January 14, 2014 at 11:21 am

    “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.

     
  45. Maju

    January 14, 2014 at 12:30 pm

    “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?

     
  46. Marnie

    January 14, 2014 at 8:11 pm

    “obsession with reindeer”

    🙂

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

     
  47. Grey

    January 15, 2014 at 5:56 pm

    “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.

     
  48. Grey

    January 15, 2014 at 6:47 pm

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

     
  49. Grey

    January 15, 2014 at 7:07 pm

    “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.

     
  50. Maju

    January 15, 2014 at 9:55 pm

    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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