Mediterranean Europe major source of European Neolithic?

[MarkoZ]

It is exactly in the admixture model Fu et al you pasted in this response. That models Satsurblia as 32% Basal Eurasian and 68% Mal'ta related ancestry. As there is no Basal in WHG, which I get from this paper, then the Middle-Eastern affinity that WHG shares with CHG must be ANE. But that is basically means that the admixture came from the north.

As has been discussed here multiple times, Mal'ta boy is very diverged from that parent population which contributed to Kotias-Satsurbalia. There is no indication whatsoever that this component came from the north. South-Central Asia via Iran would perhaps be a more reasonable explanation.

Fu et al stays clear of proposing a migration from the Middle-East.

They put said admixture on either a migrating Near Eastern population or a population with that affinity that was hiding in the Balkans.

Exactly the time we see the Middle-Eastern affinity for the first time. In the red lady of El Miron, who has been modeled as 1/3 (or 2/3 in the pasted admixture graph) of WHG and lived during the LGM. So my argument stands.

No your point doesn't stand. Hunters could have walked along the northern shores of the Mediterranean without encountering desertic tundra let alone the ice fields. There was no barrier impeding gene flow whatsoever.

 

[epoch]

As has been discussed here multiple times, Mal'ta boy is very diverged from that parent population which contributed to Kotias-Satsurbalia. There is no indication whatsoever that this component came from the north. South-Central Asia via Iran would perhaps be a more reasonable explanation.

But the diagram that you posted yourself models Satsurblia that way. Have a look at it.

They put said admixture on either a migrating Near Eastern population or a population with that affinity that was hiding in the Balkans.

Let me rephrase that: The Fu paper stays clear of pushing for M.E. immigration

Let's quote the exact text:

The Satsurblia Cluster individuals from the Caucasus dating to ~13,000–10,000 years ago share more alleles with the Villabruna Cluster individuals than they do with earlier Europeans, indicating that they are related to the population that contributed new alleles to people in the Villabruna Cluster, although they cannot be the direct source of the gene flow. One reason for this is that the Satsurblia Cluster carries large amounts of Basal Eurasian ancestry while Villabruna Cluster individuals do not (Supplementary Information section 12; Extended Data Fig. 4). One possible explanation for the sudden drawing together of the ancestry of Europe and the Near East at this time is long-distance migrations from the Near East into Europe. However, a plausible alternative is population structure, whereby Upper Palaeolithic Europe harboured multiple groups that differed in their relationship to the Near East, with the balance shifting among groups as a result of demographic changes after the Glacial Maximum

(No word of the Balkans in the paper though)

No your point doesn't stand. Hunters could have walked along the northern shores of the Mediterranean without encountering desertic tundra let alone the ice fields. There was no barrier impeding gene flow whatsoever.

But then the bottle neck wouldn't have occurred. Because of exactly that gene flow. You can't have the cake and eat it on this. Either gene flow or extreme bottle neck.

 

[MarkoZ]

But the diagram that you posted yourself models Satsurblia that way. Have a look at it.

You don't seem to understand: there's no indication that the population that coalesced with the hypothetical BE component came from the north.

Let me rephrase that: The Fu paper stays clear of pushing for M.E. immigration

Let's quote the exact text:



(No word of the Balkans in the paper though)

From the paper:

One scenario that could explain these patterns is a population expansion from southeastern European or west Asian refugia after the Glacial Maximum, drawing together the genetic ancestry of Europe and the Near East. Sixth, within the Villabruna Cluster, some, but not all, individuals have an affinity to east Asians.

But then the bottle neck wouldn't have occurred. Because of exactly that gene flow. You can't have the cake and eat it on this. Either gene flow or extreme bottle neck.

That's just ridiculous. A bottleneck tells us nothing about the gene flow that occurred before and after this particular event. If a very mixed population persists during a cataclysmic event and while other populations are culled, that's still a bottleneck. That's particularly true when most models that yield LGM & Late Glacial bottlenecks are primarily based on the study of mtDNA.

EDIT: I just noticed that we've veered off of your original hypothesis of Aurignacian genetic structure, so let's just cut it here.

 

[epoch]

You don't seem to understand: there's no indication that the population that coalesced with the hypothetical BE component came from the north.

So Fu et al were wrong when they present a model with 32% Basal en 68% Malta? Malta being the prime example of a population called Ancient North-Eurasians? I mean, you posted that graph and I don't see it modeled as admixed with anything else but Basal (split of before Ust Ishim) and something from which came from the same source as Malta did.

Now you may have the opinion that this is not the case and maybe that's right. But this graph is not your piece of evidence for it. May you should provide links which make the case for it.

From the paper:

Fair enough, that constitutes the Balkans and Anatolia

That's just ridiculous. A bottleneck tells us nothing about the gene flow that occurred before and after this particular event. If a very mixed population persists during a cataclysmic event and while other populations are culled, that's still a bottleneck. That's particularly true when most models that yield LGM & Late Glacial bottlenecks are primarily based on the study of mtDNA.

From Lararidis 2014: "
Through comparison of Loschbour’s two chromosomes we find that this low diversity is not due to recent inbreeding but instead due to a population bottleneck in this individual’s more distant ancestors (Extended Data Fig. 2)."

EDIT: I just noticed that we've veered off of your original hypothesis of Aurignacian genetic structure, so let's just cut it here.

But this is exactly why I came up with that idea. Migration from M.E. is virtually impossible because of the Basal issue. So we have two possibilities left:

1) S.E. Europe. However, a number of odd things make it less likely candidate to host the original admixture which was partly ancestral to WHG. First, we have a number of adjacent samples from 30.000 + something years ago, from Romania. They show no affinity to WHG (see Fu et al). We know that during the Mesolithic Greek HG's were of a different mtDNA: K1b.

2) Italy and South-East France. Both El Miron and Paglicci71 are mtDNA U5b, which is said to be a tracer for WHG/Villabruna cluster (Krause) and are situated at the end of LGM. El Miron has part WHG admixture and has the affinities that WHG exposes at well.

Now what culture could that have been? Perhaps the only one not sampled in Fu et al, the Solutreans. But they can't have been Gravettians since that would mean WHG would nicely and sweetly model as Gravettians. Hence something pre-Gravettian must have survived from before. And that makes it Aurignacian.

Now, completely independent from that we read that Aurignacian toolkits from the same area that the Solutrean came look like an intrusive population in the Levant some 33.000 years ago.

Now the latter is just an idea I have. But the case against M.E admixture is simple: No Basal.

 

[epoch]

You don't seem to understand: there's no indication that the population that coalesced with the hypothetical BE component came from the north.

There is actually quite some indication. Lazaridis et al 2016 models CHG as Basal + EHG.

We can model CHG as a mixture of Iran_N and different European hunter-gatherer populations (Table S7.7), with an estimate of 71.6±6.0% Iran_N, 7.0±3.8% WHG, 21.4±7.7% EHG.

Table 7.6 shows that the differences between CHG and meso Iran doesn't favour any of the WHG's or EHG's. That means it may very well be old.

No strong evidence that allele frequency differences between CHG and Iran_N are strongly associated with a particular ancient West Eurasian hunter-gatherer population

The paper states that Mesolithic Iran had EHG/ANE admixture:

involving the EHG and the related Eneolithic Steppe population also from eastern Europe groups than the Neolithic. Tentatively,this might suggest that the pre-Neolithic population of Iran had an affinity to the EHG/Ancient North Eurasians that was diluted during the Neolithic, although the lack of negative f4-statistics does not allow us to discern what is the source of this dilution.

It models Anatolian as Levantine + extra WHG.

So it's not that nobody but me suggests something old akin to WHG or EHG admixted into the M.E. somehow.

 

[MarkoZ]

So Fu et al were wrong when they present a model with 32% Basal en 68% Malta? Malta being the prime example of a population called Ancient North-Eurasians? I mean, you posted that graph and I don't see it modeled as admixed with anything else but Basal (split of before Ust Ishim) and something from which came from the same source as Malta did.

Now you may have the opinion that this is not the case and maybe that's right. But this graph is not your piece of evidence for it. May you should provide links which make the case for it.

The point is that the Near Eastern ancestry of the Villabruna cluster isn't related to the West Eurasian branch. This same branch coalesced with another population to give birth to the Mal'ta population, which is not ancestral the population that converged with Basal Eurasian to form Satsurbalia-Kotias and related specimens.

Through comparison of Loschbour’s two chromosomes we find that this low diversity is not due to recent inbreeding but instead due to a population bottleneck in this individual’s more distant ancestors (Extended Data Fig. 2)."

I guess we all agree that the Near Eastern admixture appeared before the Mesolithic.

But this is exactly why I came up with that idea. Migration from M.E. is virtually impossible because of the Basal issue. So we have two possibilities left:

I see no reason to believe that Basal Eurasian was spread far and wide in West Asia before and even during the Iranian Zarzian and Levantine Kebaran phases respectively. The skeletal evidence suggests that typically West Eurasian skeletons predominate before a shift into a rather African ('Mediterranean') direction occurs with the latter two cultures. See for example Hershkovitz 1995 on Ohalo II, or Stock 2005 on Wadi Mataha F-81.

Be that as it may ( I guess one can only speculate due to the lack of relevant samples), the Basal Eurasian issue is yet less relevant in the case of an Upper Paleolithic population that emanated from Central Asia or thereabouts. One doesn't have to trek through the Arabian deserts to get to Europe from Iran or Turkmenistan.

1) S.E. Europe. However, a number of odd things make it less likely candidate to host the original admixture which was partly ancestral to WHG. First, we have a number of adjacent samples from 30.000 + something years ago, from Romania. They show no affinity to WHG (see Fu et al). We know that during the Mesolithic Greek HG's were of a different mtDNA: K1b.

Didn't you cite the Mueirii samples as evidence of backmigration just a few posts earlier? In any case, we're not talking about the main ancestry of the Villabruna cluster (which is on the West Eurasian branch), but the Near Eastern influence that affected those populations. Mesolithic and early Upper Paleolithic samples aren't really relevant in this case.

Now what culture could that have been? Perhaps the only one not sampled in Fu et al, the Solutreans. But they can't have been Gravettians since that would mean WHG would nicely and sweetly model as Gravettians. Hence something pre-Gravettian must have survived from before. And that makes it Aurignacian.

That would make sense if the only thing that made the Villabruna cluster divergent was internal West Eurasian structure (as seems to be the case with, for example, Gravettian differentation with respect to the Aurignacian) but with Y-DNA P1 and a non-West-Eurasian aDNA component we're clearly talking about foreign influence.

Now the latter is just an idea I have. But the case against M.E admixture is simple: No Basal.

Perhaps the problem lies in the terminology: what's the maximum extent of the Middle East for you? Does it include Iran-Armenia-Anatolia?

 

[MarkoZ]

So it's not that nobody but me suggests something old akin to WHG or EHG admixted into the M.E. somehow.

Hold on, we're in agreement that both the Middle East and Europe were affected by the same population movements of the West Eurasian branch? Of course there's old West Eurasian ancestry everywhere in the Middle East - it just didn't get there from Europe

 

[LeBrok]

Relationship of WHG and CHG seems to be through NE Euro component only. NE Euro developed in Europe and NW Asia only. However the complete lack of Med admixture in CHG is telling us that WHG wasn't the source of NE Euro in CHG. More likely the contact was with EHG instead, or most likely with central asian h-g type, because of surprising 1% of Siberian admixture in CHG. EHG had substantial American admixture but 0 Siberian.

M325047	KO1, I-L68	M551062
Hungarian, Tiszaszőlős-Domaháza	7.7 kya		Kotias CHG
Run time	9.43		Run time	15.3
S-Indian	-		S-Indian	0.78
Baloch	-		Baloch	34.39
Caucasian	-		Caucasian	50.95
NE-Euro	80.37		NE-Euro	5.89
SE-Asian	-		SE-Asian	0.88
Siberian	-		Siberian	0.95
NE-Asian	-		NE-Asian	-
Papuan	0.53		Papuan	0.11
American	-		American	-
Beringian	-		Beringian	0.16
Mediterranean	18.59		Mediterranean	-
SW-Asian	-		SW-Asian	-
San	-		San	-
E-African	-		E-African	-
Pygmy	-		Pygmy	-
W-African	0.5		W-African	5.89

 

[LeBrok]

There is much stronger connection of WHG to Anatolia and Levant through Mediterranean admixture. We are yet to find h-g who was the source of it.

M325047	KO1, I-L68	M041601	Merged		M936428	I0709		EN Jordan, .Ain Ghazal	7.5k BC
Hungarian, Tiszaszőlős-Domaháza	7.7 kya		Natufian			Anatolian EF		M632231	I1707
Run time	9.43		Run time	6.39		Run time	12.82		Run time	3.31
S-Indian	-		S-Indian	-		S-Indian	-		S-Indian	-
Baloch	-		Baloch	-		Baloch	-		Baloch	-
Caucasian	-		Caucasian	13.98		Caucasian	36.48		Caucasian	27.79
NE-Euro	80.37		NE-Euro	-		NE-Euro	0.96		NE-Euro	2.52
SE-Asian	-		SE-Asian	-		SE-Asian	0.08		SE-Asian	-
Siberian	-		Siberian	-		Siberian	-		Siberian	-
NE-Asian	-		NE-Asian	-		NE-Asian	-		NE-Asian	-
Papuan	0.53		Papuan	0.68		Papuan	-		Papuan	-
American	-		American	-		American	-		American	-
Beringian	-		Beringian	-		Beringian	-		Beringian	-
Mediterranean	18.59		Mediterranean	27.39		Mediterranean	48.79		Mediterranean	34.48
SW-Asian	-		SW-Asian	53.62		SW-Asian	13.49		SW-Asian	34.6
San	-		San	-		San	-		San	0.51
E-African	-		E-African	4.33		E-African	-		E-African	-
Pygmy	-		Pygmy	-		Pygmy	-		Pygmy	-
W-African	0.5		W-African	-		W-African	0.16		W-African	0.1

 

[bicicleur 2]

There is much stronger connection of WHG to Anatolia and Levant through Mediterranean admixture. We are yet to find h-g who was the source of it.

we know that the origin of the European farmers was (G2a2) central Anatolian, not (Natufian) Levant and not Zagros (Iran Neo)
therefore the Mediterranean in EN Jordan may be to some extent derived from admixture with Central Anatolians

 

[Sile]

we know that the origin of the European farmers was (G2a2) central Anatolian, not (Natufian) Levant and not Zagros (Iran Neo)
therefore the Mediterranean in EN Jordan may be to some extent derived from admixture with Central Anatolians

All Natufian samples have caucasian % ...........this fits with all Ydna haplogroups from K origins are north of the Zargos mountains

 

[LeBrok]

we know that the origin of the European farmers was (G2a2) central Anatolian, not (Natufian) Levant and not Zagros (Iran Neo)
therefore the Mediterranean in EN Jordan may be to some extent derived from admixture with Central Anatolians

We need to find Central Anatolian HG.

 

[epoch]

The point is that the Near Eastern ancestry of the Villabruna cluster isn't related to the West Eurasian branch. This same branch coalesced with another population to give birth to the Mal'ta population, which is not ancestral the population that converged with Basal Eurasian to form Satsurbalia-Kotias and related specimens.

These are three assumptions that need at least some evidence. You assume:

1) Near Eastern ancestry of the Villabruna cluster isn't related to the West Eurasian branch
2) This same branch coalesced with another population to give birth to the Mal'ta population
3) Mal'ta population is not ancestral the population that converged with Basal Eurasian to form Satsurbalia-Kotias and related specimens

I don't see any of these three assumptions proven.

I see no reason to believe that Basal Eurasian was spread far and wide in West Asia before and even during the Iranian Zarzian and Levantine Kebaran phases respectively. The skeletal evidence suggests that typically West Eurasian skeletons predominate before a shift into a rather African ('Mediterranean') direction occurs with the latter two cultures. See for example Hershkovitz 1995 on Ohalo II, or Stock 2005 on Wadi Mataha F-81.

Mesolithic Hotu samples already showed basal. Upper Paleolithic CHG (13,300 years old) had it. Are you suggesting these were islands of Basal? Or that 13.000 years ago Basal was in the Caucasus but not Iran?

Be that as it may ( I guess one can only speculate due to the lack of relevant samples), the Basal Eurasian issue is yet less relevant in the case of an Upper Paleolithic population that emanated from Central Asia or thereabouts. One doesn't have to trek through the Arabian deserts to get to Europe from Iran or Turkmenistan.

But what makes you think it went that way?

Didn't you cite the Mueirii samples as evidence of backmigration just a few posts earlier? In any case, we're not talking about the main ancestry of the Villabruna cluster (which is on the West Eurasian branch), but the Near Eastern influence that affected those populations. Mesolithic and early Upper Paleolithic samples aren't really relevant in this case.



That would make sense if the only thing that made the Villabruna cluster divergent was internal West Eurasian structure (as seems to be the case with, for example, Gravettian differentation with respect to the Aurignacian) but with Y-DNA P1 and a non-West-Eurasian aDNA component we're clearly talking about foreign influence.

Y-DNA R1b1b could have been coming from ANE as well. It is prevalent among populations which all show ANE admixture. WHGa has been modeled as partly ANE by Lazaridis 2016 and Davidski. KO1's mtDNA can also be found in Afontova Gora 3, so contact with ANE seems logical. Han has been modeled a partly ANE so that explains the tad Asian affinity. No need for a bypass via Iran here.

Perhaps the problem lies in the terminology: what's the maximum extent of the Middle East for you? Does it include Iran-Armenia-Anatolia?

 

[MarkoZ]

These are three assumptions that need at least some evidence. You assume:

1) Near Eastern ancestry of the Villabruna cluster isn't related to the West Eurasian branch
2) This same branch coalesced with another population to give birth to the Mal'ta population
3) Mal'ta population is not ancestral the population that converged with Basal Eurasian to form Satsurbalia-Kotias and related specimens

I don't see any of these three assumptions proven.

I don't assume these things, these are literally in the tree.

Edit: I think those results were replicated by the authors of the recent Baltic paper, with the exception that they sought to model the Karelian hunters. I'll see if I can find it.

Mesolithic Hotu samples already showed basal. Upper Paleolithic CHG (13,300 years old) had it. Are you suggesting these were islands of Basal? Or that 13.000 years ago Basal was in the Caucasus but not Iran?

But what makes you think it went that way?

Apart from the skeletal evidence, we can narrow down the timeframe of Basal Eurasian dispersal by look at the European evidence. The beginning of the Gravettian in sees Middle Eastern Ahmarian-Lagaman-Dabba blade industries appearing in along the Danubian and in the North Caucasus. I think that we can say with near-certitude that these dynamics brought at least some gene flow into the aforementioned areas. Since we do not see a subsequent increase in Basal Eurasian affinity, I guess we can safely assume that 30k-22k BC is the terminus post quem for this hypothetical component in Jordan-Israel and vicinity.

 

[epoch]

I don't assume these things, these are literally in the tree.

Not the way you interpret it a number of posts ago. That lineage is clearly ANE.

As has been discussed here multiple times, Mal'ta boy is very diverged from that parent population which contributed to Kotias-Satsurbalia. There is no indication whatsoever that this component came from the north. South-Central Asia via Iran would perhaps be a more reasonable explanation.

Apart from the skeletal evidence, we can narrow down the timeframe of Basal Eurasian dispersal by look at the European evidence. The beginning of the Gravettian in sees Middle Eastern Ahmarian-Lagaman-Dabba blade industries appearing in along the Danubian and in the North Caucasus. I think that we can say with near-certitude that these dynamics brought at least some gene flow into the aforementioned areas. Since we do not see a subsequent increase in Basal Eurasian affinity, I think we can safely say that 30k-22k BC is the terminus post quem for this affinity in Jordan-Israel and vicinity.

I read more papers claiming an Ahmarian origin for the Gravettian. But we have a large amount of Gravettian DNA and it shows no M.E. affinity. I think we can say with near-certitude that this fact disproves the theory.

https://paleo.revues.org/607#tocto1n2

 

[MarkoZ]

Not the way you interpret it a number of posts ago. That lineage is clearly ANE.

ANE was coined for the Mal'ta boy, but it doesn't lead to Satsurbalia-Kotias in Fu's tree. In fact it is highly diverged from the source population.

I read more papers claiming an Ahmarian origin for the Gravettian. But we have a large amount of Gravettian DNA and it shows no M.E. affinity. I think we can say with near-certitude that this fact disproves the theory.

https://paleo.revues.org/607#tocto1n2

That's exactly the point. Middle East until 22k BC = fully or mostly West Eurasian.

How frustrating. Am I being trolled?

 

[epoch]

ANE was coined for the Mal'ta boy, but it doesn't lead to Satsurbalia-Kotias in Fu's tree. In fact it is highly diverged from the source population.

But the exact same way is how they show Goyet116's relation to El Miron and Kostenki14's relation to Vestonice16, both of which are considered ancestral in the text.

That's exactly the point. Middle East until 22k BC = fully or mostly West Eurasian.

I am really making an effort to see your point. Let's assume that at 22k BC something fully Eurasian was in Iran. From what I read the UP culture also shows connection to the Aurignacians. Then, 22k BC, something Basal Eurasian comes in and introduces it admixture. 3000 years later an affinity to the Middle-East pops up in an European population without any Basal. What are you suggesting? Displaced pre-Basal Iranians trekked to Europe during LGM? Before LGM? Or complete replacement in Europe by Iranian refugees? But when?

Furthermore, the fact that WHG (and CHG) went through a bottle neck is not just established via mtDNA, it is established via RoH. Check the CHG paper. So I still think my argument stands.

 

[MarkoZ]

I am really making an effort to see your point. Let's assume that at 22k BC something fully Eurasian was in Iran. From what I read the UP culture also shows connection to the Aurignacians.

Not sure what you mean by that - Upper Paleolithic sites aren't really that remarkable, and of course within Eurasia they all ultimately derive from a common Asian source, so some broad similarities can be identified everyhwere. Though in all likelihood Europe would have been little more than a sink, with some possible exceptions like North-West Africa where gene-flow from Europe must have occurred at some point.

Then, 22k BC, something Basal Eurasian comes in and introduces it admixture. 3000 years later an affinity to the Middle-East pops up in an European population without any Basal. What are you suggesting? Displaced pre-Basal Iranians trekked to Europe during LGM? Before LGM? Or complete replacement in Europe by Iranian refugees? But when?

Without ancient samples this is impossible to say. I think Villabruna's Y-DNA makes a relatively late introduction more likely. There's also the peculiar finding that Villabruna's skeletal morphology is most similar to present-day North Africans, which makes him an outlier among increasingly temperate-adapted European hunters:

In conclusion, the results of bivariate analysessuggest that Villabruna 1 had a relatively slenderphysique, intermediate in its nature between thetropically-adapted constitution characteristic ofEUP and the temperate-adapted structure of contemporaryLUP people. When compared withmodern samples, the specimen shows highestsimilarities with North Africans populations. Aspreviously mentioned, the ponderal index is theonly parameter in disagreement with this generalpattern. In order to evaluate the importance of thelatter characteristic in a generally differently orientedmorphocomplex, a multivariate analysis ofVillabruna 1’s body proportions was carried out.

http://www.isita-org.com/jass/Contents/2008 vol86/09_Vercelotti.pdf

That fact that he retains a tropically-adapted skeleton makes me doubt that the majority of his ancestors were in Europe throughout the LGM. But I guess the evidence isn't strong enough to completely rule out the possibility of a Mediterranean population in the Balkans, Italy or one of the islands which already had said 'West Asian' affinity. In the latter scenario Villabruna's Y-DNA would be quite difficult to explain, too.

Furthermore, the fact that WHG (and CHG) went through a bottle neck is not just established via mtDNA, it is established via RoH. Check the CHG paper. So I still think my argument stands.

The internal relatedness of Mesolithic WHG populations isn't in dispute. CHG is another such population which derives from highly diverse ancestors and became very homogeneous for whatever reason.

 

[epoch]

Not sure what you mean by that - Upper Paleolithic sites aren't really that remarkable

Which undermines your earlier remark on Ahmarian influences in Gravettian.

, and of course within Eurasia they all ultimately derive from a common Asian source, so some broad similarities can be identified everyhwere. Though in all likelihood Europe would have been little more than a sink, with some possible exceptions like North-West Africa where gene-flow from Europe must have occurred at some point.

Both the origin of mtDNA U6 and the resurgeance of GoyetQ116 in Magdalenian point to something more complex that that sink. It may need to be reconsidered.

Without ancient samples this is impossible to say. I think Villabruna's Y-DNA makes a relatively late introduction more likely. There's also the peculiar finding that Villabruna's skeletal morphology is most similar to present-day North Africans, which makes him an outlier among increasingly temperate-adapted European hunters:

True, although comparison of a 14k yo old sample with present day North Africans is not very meaningful.

http://www.isita-org.com/jass/Contents/2008 vol86/09_Vercelotti.pdf

That fact that he retains a tropically-adapted skeleton makes me doubt that the majority of his ancestors were in Europe throughout the LGM. But I guess the evidence isn't strong enough to completely rule out the possibility of a Mediterranean population in the Balkans, Italy or one of the islands which already had said 'West Asian' affinity. In the latter scenario Villabruna's Y-DNA would be quite difficult to explain, too.

That article itself also claims that Villabruna lies nicely inbetween Early Upper Paleolithics and Late Upper Paleolithics

These observations are confirmed by the ratios between limb bones and skeletal trunk height, which place Villabruna 1 near to the tropically-adapted EUP morphotype, among fossil series, and close to North Africans, among recent samples. Therefore, limb bones/trunk proportions suggest that Villabruna 1 retains warm-climate adaptations characterizing EUP populations more than his contemporaries (Holliday, 1997).

If you take a look at the PCA's it isn't all that dramatic and may very well be on the tail of normal variation.

The internal relatedness of Mesolithic WHG populations isn't in dispute. CHG is another such population which derives from highly diverse ancestors and became very homogeneous for whatever reason.

Well, it was established with Loschbour so you'd have to figure out a way of admixture before the bottle neck but after LGM. LGM being the best, even only good candidate for the bottle neck as we know population of humans dropped and stayed low during it.

It simply doesn't fit IMHO.

 

[MOESAN]

I doubt today NorthAfrican population is homogenous for limbs!!! legends