The genetic history of Ice Age Europe

If that's the case, as I pointed out early in the thread, there's a Basal Eurasian problem. Villabruna doesn't have any Basal Eurasian. Did WHG leave the Near East before it arrived? CHG is 13,300 before present, yes? It already had about 32% Basal Eurasian according to this paper. How could Villabruna, dated 14,000 YBP have avoided it? Isn't that cutting it a little close?

Villabruna cannot be Near Eastern without Basal Eurasian (ENF) admxiture. And according to David's PCA chart, Villabruna clusters perfectly with other Epipaleolithic and Mesolithic Western Europeans (WHG). Villabruna is as remote from CHG as from ENF, but is typical WHG without clustering with older Paleolithic European samples.

In other words, the Villabruna R1b1 is a typical Mesolithic European hunter-gatherer related to mtDNA U5 and Y-DNA I2 rich populations. It confirms exactly my suspicion that this R1b1 lineage was originally a EHG and was integrated (probably any centuries or millennia earlier) into U5+I2 tribes in Eastern Europe before U5+I2 moved west and replaced Aurignacian and Gravettian lineages in central and western Europe. This is the first major wave from the Steppe I was referring to, which took place about 10,000 years before Yamna.

 
response to random comments from the thread

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Neanderthals

If they were specifically adapted for anything it would be the ice age so I think the idea that their dna was necessarily junk is very stupid. It seems far more likely they had dna that was very useful to survive the ice but gradually became less so over time as the climate warmed up - for example a warming higher metabolism and greater physical size and strength that could only be supported by megafauna size calorie packages.

This obviously makes more sense than the idea that people who survived the Ice Age were some kind of genetic waste dump.

I think the clear desire to play down the role of Neanderthals is because the base Out of Africa theory is wrong and humans needed that archaic DNA to adapt to the new environments - at least at the time - and that idea triggers a PC reaction.

could be wrong but proof may come when there's enough dna to calculate the rate at which it dropped - if slow at the beginning and fast later it might support the cost/benefit of neanderthal dna being climate dependent.

(edit: i always get a bit cranky about people attacking neanderthals as for some reason it always makes me feel people are attacking my relatives lol. if too cranky sounding, apologies)

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Ukraine as a refugia

We know humans can survive from marine resources further north than otherwise (e.g Eskimo) so it seems to me that would apply during the LGM as well, so not the land part of Ukraine per se but the parts around the Black Sea maybe.

Similarly the various now sunken lands seem like the best candidates for other refugia for the same reason, marine resources less affected by the colder climate
- adriatic
- aegean
- Doggerland
- Franco-Cantabria

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I've never understood why at least some OoA along the west coast of Africa over to Iberia is always discounted.

I don't believe it never happened - especially when the sea level was lowest.

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WHG / R1b etc from the middle east feels like a repeat of the media flurry to report European replacement by neolithic farmers from Turkey - it's connected to current politics imo and partly acts as a distortion - although having the possibility in the paper guarantees publicity so can't blame them.

The WHG part could work if "Basal Eurasian" came from the CHG region and the CHG repopulated the middle east after the WHG spread west.

(which is possible)

but again i'd take the near east aspect with a pinch of salt first and wait to see if it stands up after the flurry has moved on.

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With the Italian find I've gone back to thinking Doggerland/Cantabria/Iberia may be more to do with this than seemed possible recently so WHG (maybe coming in from the east) mixing with the mystery ancient population from the Atlantic coast.

Be interesting to hear Maju's take as I think this was always his thing until the lack of evidence seemed overwhelming.

Yamnaya R1b and SW Europe R1b - separate chain of events?

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edit - missed one

i don't think east asian ancestry implies near east origin at all

pottery was apparently east asian first and seems to have arrived in the west via the steppe route

which if you look at the physical geography makes sense imo - before sailing the steppe route looks dramatically easier
 
An interesting thing about the origin of the R1b guy, taking into account that i don't master admixtures, is that having not basal Euroasian but having Near Eastern... a steppe route is not tenable as the Euroasian split might be somewhere in Central Asia (as the Indian subcontinent is not the place), so as to have Near Eastern DNA but lacking a share with East Asian, the origin or route left is before the splitting area of Euroasian, and that is Anatolia, Levant; also the Caucasus would be possible and would give some reasonable cause: maybe the Euroasian formed north of it as the genetic contact with the south is more difficult to keep. But Central Asian deserts also are a good place to lose genetic contact.
 
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Greying Wanderer;479639]response to random comments from the thread

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Neanderthals

If they were specifically adapted for anything it would be the ice age so I think the idea that their dna was necessarily junk is very stupid. It seems far more likely they had dna that was very useful to survive the ice but gradually became less so over time as the climate warmed up - for example a warming higher metabolism and greater physical size and strength that could only be supported by megafauna size calorie packages.

This obviously makes more sense than the idea that people who survived the Ice Age were some kind of genetic waste dump.

I think the clear desire to play down the role of Neanderthals is because the base Out of Africa theory is wrong and humans needed that archaic DNA to adapt to the new environments - at least at the time - and that idea triggers a PC reaction.

could be wrong but proof may come when there's enough dna to calculate the rate at which it dropped - if slow at the beginning and fast later it might support the cost/benefit of neanderthal dna being climate dependent.

(edit: i always get a bit cranky about people attacking neanderthals as for some reason it always makes me feel people are attacking my relatives lol. if too cranky sounding, apologies)

They're not saying it was all "junk". :) They're saying the stuff that helps for cold climates we've kept and the stuff that isn't helpful is being culled over time. That seems to me to be exactly how natural selection works. I don't see how the fact that there is Neanderthal depletion or decrease of genes over time at certain specific loci relating to function can be explained away.

As for taking it personally, I try never to take discussions like this personally, or at least I try never to let my personal feelings one way or another affect my analysis or the logic of the argument. That may partly be because I spent my work life having to do that. Also, it's just too long ago, and too small a part of our dna, even if I'm half sort of "quasi-Tuscan", and last time I checked Tuscans have a good healthy dose of Neanderthal dna. :) My concern is for the modern humans affected adversely by this dna.
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Ukraine as a refugia

We know humans can survive from marine resources further north than otherwise (e.g Eskimo) so it seems to me that would apply during the LGM as well, so not the land part of Ukraine per se but the parts around the Black Sea maybe.

Similarly the various now sunken lands seem like the best candidates for other refugia for the same reason, marine resources less affected by the colder climate
- adriatic
- aegean
- Doggerland
- Franco-Cantabria

I agree in general, although the terrain, flora, fauna, and the temperature in certain of these areas might have still been relatively more conducive to long term survival than in others.

WHG / R1b etc from the middle east feels like a repeat of the media flurry to report European replacement by neolithic farmers from Turkey - it's connected to current politics imo and partly acts as a distortion - although having the possibility in the paper guarantees publicity so can't blame them.

The WHG part could work if "Basal Eurasian" came from the CHG region and the CHG repopulated the middle east after the WHG spread west.

(which is possible)

but again i'd take the near east aspect with a pinch of salt first and wait to see if it stands up after the flurry has moved on.

I'm also taking a wait and see attitude, as indeed are the authors of the study if their claims in the paper are taken at face value. We need similarly old samples from far eastern Europe and the Near East (including around the Caucasus).

As for your quasi-conspiracy theory, I sincerely doubt it. A Chinese researcher now working in Beijing wants to push a Near East to Europe population movement in the absence of data? You also forget that indeed farmers coming from the Near East did largely replace the WHG, or perhaps we should now call them "Villabrunians" in large parts of Europe. All of these groups, including the Aurignacians, originally came from elsewhere. Using modern, cultural-political designations to assign "identity" to 14,000 year old humans doesn't make any sense to me. Do you think some hunter-gatherer band following their food sources suddenly became "Near Eastern" if they moved from Greece to Turkey, or "European" if they moved in the other direction? The same would apply to Mal'ta descendants too, btw.

i don't think east asian ancestry implies near east origin at all

pottery was apparently east asian first and seems to have arrived in the west via the steppe route

which if you look at the physical geography makes sense imo - before sailing the steppe route looks dramatically easier

The authors specifically said that these were two separate population movements. The East Asian gene flow is completely separate from the possible "Near Eastern" one.

Again, here's the direct link:
http://genetics.med.harvard.edu/reich/Reich_Lab/Welcome_files/FuQ_nature17993.pdf

A relevant quote:
"Second, we detect an excess of allele sharingwith east Asians in a subset of Villabruna Cluster individuals—beginning with an ~13,000-year-old individual from Switzerland—asrevealed by significant statistics of the form D(Test1, Test2; Han, Mbuti)(Fig. 4b and Extended Data Fig. 3). For example, Han Chinese sharemore alleles with two Villabruna Cluster individuals (Loschbour andLaBrana1) than they do with Kostenki14, as reflected in significantlynegative statistics of the form D(Kostenki14, Loschbour/LaBrana1; Han,Mbuti)4. This statistic was originally interpreted as evidence of BasalEurasian ancestry in Kostenki14. However, because this statistic is consistentwith zero when Han is replaced with Ust’-Ishim, these findingscannot be driven by Basal Eurasian ancestry (as we discuss earlier),and must instead be driven by gene flow between populations relatedto east Asians and the ancestors of some Europeans."

The Villabrunian samples which show this affinity are Bichon and Loschbour, not Villabruna himself.

Interestingly, the highest level of East Asian is in Mal'ta, if I remember correctly. So, the anthropologists might have been correct to see "Mongolian" features in the remains.
 
In my opinion, with what I understand, this route for R1b, looks most probable with known data. How does it work for you?

Look at all the Variance between Sardinia and the Steppe! It looks like to me R1b from the Middle East is becoming an impossible pipe dream.



What about autosomally? Would you venture a guess if Maikop will plot with Iranians, modern Caucasus, or Steppe?
Maybe it is possible, I don’t know. But I doubt it. All the major academic studies of the recent times on DNA are concluding all the same that R1* went into the Iranian Plateau.


Very simple. They would be not very different from that Bronze Age fella from Armenia. But a lttle bit more with Steppes ancestry, because Mayko was bordering the Steppes. So, there was a some kind of minor gene flow from the Steppes into the Maykop. But culturally Maykop was very West Asian in nature. And it was in Northern Caucasus. Yamnaya was genetically heavily influenced by folks from Maykop. And there was also a lot West Asian auDNA in the Yamnaya Horizon. Maykop would have twice more (2x) West Asian auDNA than the Yamnaya Horizon.

Modern WEST Iranians and modern Caucasians are very close and similar to each other. Modern Caucasians have a little bit more Steppes ancestry because they are closer to the Steppes. While West Iranians have a little bit SouthWest (Semitic) ancestry, because West Asians live closer to the Semites. The closest people to West Iranians are actually South Caucasians.
 
There were distinct populations in Ice age Europe(besides WHG and EHG). Here are the conclusions by the authors based on treemix and admixturegraph.


Conclusion
>Paleo Euros as clade opposed to MA1
>Vestonice is mostly Kostinki with minor Goyet.
>El Miron is mostly Goyet with minor Villabruna.’
>Loschbour is mostly Villabruna with minor Goyet.


Vestonice: 30,000 years old Czech Republic(Central Europe).
Kostinki: 36,000 years old Russia(Western egde).
Goyet: 30,000 years old Beligium.
El Miron: 20,000 years old Spain.
Villabruna: 14,000 years old Northern Italy.
Loschbour: 8,000 years old Luxembourg.


Goyet is 35.000 years old. It makes him Aurignacian and nearly comtemporary to Kostenki 14
 
So, while admixture isn't going to show this "Near Eastern" affinity, formal stats do show it. What's being lost in the renewed R1a vs R1b wars as to who is the most Indo-European of them all, is that the authors leave open two possibilities: one that the Villabruna set was in Europe as early as the "Gravettian" group, and just represents substructure in Europe, and one that they arrived in Europe somewhere around 14000 YBP from the southeast, through Greece perhaps.

If the former was true, why all this Near Eastern "affinity", unless there was some inflow that changed them slightly perhaps?

Kostenki14 shows in a lot of formal stats as well as ADMIXTURE a clear Middle-Eastern signal. In f3 stats for instance Sardinia is high among other WHG. It shows in all kind of D-stats. However, the paper makes a point of showing no Basal Eurasian admixture was shown in Kostenki14. Then what is that signal? It could be part of WHG was another branch from Kostenki14 in which, other than the rest, a number of SNPs are available that drifted away in other decendants of Kostenki14. Suppose that WHG added to Middle-Eastern DNA, then we might come up with this result. Anatolian ENF showed WHG admixture, so there was contact.

Maybe they are just being really cautious. I guess we need to stay tuned.

It doesn't seem then that Magdalenian expansions left all that large a trace; most "European hunter-gatherer" ancestry is WHG or related to the Villabruna group.

Twitter-Iosif Lazaridis (one of the co-authors)

"It seems that there was a hunter-gatherer "reset" ~14,000 years ago that left only the "Villabruna cluster" as the inheritors of Europe..."

I don't know. El Miron shows a Villabruna signal. But that could very well be shared drift, which would mean that the typical GoyetQ116 signal in all of us may be just 5-8%, but the total inheritance may be a tad larger. And yes, I'm biased as that gives me the "we have always been here" tingling sense ;)
 
@Angela

Yes, I was a bit annoyed at the time from reading a lot of comments on various sites so I'll wait for a bit.

The main thing is

Twitter-Iosif Lazaridis (one of the co-authors)

"It seems that there was a hunter-gatherer "reset" ~14,000 years ago that left only the "Villabruna cluster" as the inheritors of Europe..."


(Epoch:)I don't know. El Miron shows a Villabruna signal. But that could very well be shared drift, which would mean that the typical GoyetQ116 signal in all of us may be just 5-8%, but the total inheritance may be a tad larger. And yes, I'm biased as that gives me the "we have always been here" tingling sense ;)

I've always felt (from rock climbing years ago) that there was (small amounts of) something odd and old in pockets along the Atlantic coast so the Gotye thing is a big deal to me but seemed to be drowned out by the media's usual spin.

But yes, best to be calm :)
 
An interesting thing about the origin of the R1b guy, taking into account that i don't master admixtures, is that having not basal Euroasian but having Near Eastern... a steppe route is not tenable as the Euroasian split might be somewhere in Central Asia (as the Indian subcontinent is not the place), so as to have Near Eastern DNA but lacking a share with East Asian, the origin or route left is before the splitting area of Euroasian, and that is Anatolia, Levant; also the Caucasus would be possible and would give some reasonable cause: maybe the Euroasian formed north of it as the genetic contact with the south is more difficult to keep. But Central Asian deserts also are a good place to lose genetic contact.

I'm not sure this will be easy to explain without a diagram - which i suck at anyway - but...


I mostly view these things geographically with population split into ecozones so the way i think the data *might* fit is to imagine three zones


1) mammoth steppe zone extending in a crescent all the way from Doggerland to Siberia (ydna R)


2) mountain/hill zone along the edge of the steppe in the east including Caucasus/Altai (CHG, basal)


3) southerly zone (WHG)


something like


SSSSSSSSSSSS
WWWWCCCCCC
WWWWWWWW

(where S = mammoth steppe, W = WHG, C = CHG)


so the mammoth steppe people bordering the WHG people in the west but separated from them by CHG people further east


this way the italian dude could be
- autosomal WHG (whether from near east or not)
- ydna mammoth steppe
- east asian connection from steppe dudes (if EA connection was via steppe bypassing CHG)
- no CHG so no basal


This model would require the near east to have had a CHG expansion later to explain the basal there now.
 
How come? To my understanding, it is quite possible that R1b started expansion from Kurdistan area, or anywhere in 2k km radius. And who knows as original cow herders or blacksmiths?
Ancestor's with R1b who had copper on Steppe- plotted in 3D. 1000years+ R1b copper on Steppe.
SVP58/I0444 buried with a blunt mace 48 cm long, 767 g in weight, cast/annealed and made of pure copper, like most Yamnaya metal objects.
SVP35-4700-4000BC R1b1 M415 Imported copper. https://www.youtube.com/watch?v=QapUGZ0ObjA 11:50
SVP58/I0444 3300-2700 BC R1b1a2a2 (Z2103)CTS-1078/Z2103,L150+,M415+



http://www.open-genomes.org/analysis/PCA/Eurogenes_Ice_Age_Eurasians_PC_plot_1-2-3.html & Eurogenes Ice Age run.
R1b copper1.png
R1b2 Yamnaya copper mace..pngkvhalynsk and Yamnaya R1b.jpg

In distance, the 2 R1b samples with copper were relatively close.
 
Would not be more easy or economic to explain the actual East Asian genes as the result of admixture of old Paleolithics with recent ones from Near East? As somehow seems to display some guys of Villabruna cluster.

In fact for each Spanish or Italian sample i would do x100 as this could be the demographic difference among tundra dwellers and refugia dwellers, and as the refugia ecosystem was spreading north then the refugia dwellers may follow it. Mammoth hunters would be few and out of water by then, as an Eskimo in the Italian Riviera.
 
haplogroup-r-migration-map.jpg



In my opinion, with what I understand, this route for R1b, looks most probable with known data. How does it work for you?



Possible, but with more and more samples it is looking less likely. However I'd be willing to give you more than benefit of doubt.
.



This is what startled me a couple of weeks ago, when I realized what Gioiello was trying to bring to everyones attention. The age and variance of R1b V-88 of places,in remote Island- Cagliari, Sardinia! Nigeria, Saudi, Kuwait are all down stream. R1b-V88 is generally considered quite old[formed 17200 ybp, TMRCA 10200 ybpinfo]. Same with R1b-M73 found almost exclusive on Steppe. BTW some Egyptian samples have never been made officially public, King Tut for example.
https://www.yfull.com/tree/R-V88/

We also have more variance.
Previously I've pointed out, that some of the most basal lineages of L51 can be found in Sardinia:

Sardinian_L51.png

Look at all the Variance between Sardinia and the Steppe! It looks like to me R1b from the Middle East is becoming an impossible pipe dream.



What about autosomally? Would you venture a guess if Maikop will plot with Iranians, modern Caucasus, or Steppe?

Do not use Ftdna map...........it does not even cover the Birth of R haplogroup in south-east Asia on the Karafet papers of 2014 and 2015
 
Would not be more easy or economic to explain the actual East Asian genes as the result of admixture of old Paleolithics with recent ones from Near East? As somehow seems to display some guys of Villabruna cluster.

In fact for each Spanish or Italian sample i would do x100 as this could be the demographic difference among tundra dwellers and refugia dwellers, and as the refugia ecosystem was spreading north then the refugia dwellers may follow it. Mammoth hunters would be few and out of water by then, as an Eskimo in the Italian Riviera.

Well (IIRC) pottery supposedly first developed in East Asia and the middle east got it *later* than the steppe which hints that it spread via the steppe route. Maybe I am remembering it wrong.
 
asgjb6.png


Previous studies assumed that Kostenki14 was the oldest European man with Basal Eurasian ancestry, who survived the Ice Age, but Fu et al. (2016) put it more accurately. Kostenki14 was likely to be a mixed-race individual between populations related to East Asians (hg C) and the ancestors of Europeans (hg U2). This new study also found that some of the Villabruna Cluster individuals such as Loschbour and LaBrana1 are closely associated with Han Chinese but not all Villabruna individuals have an affinity with East Asians. The ancestors of Loschbour and LaBrana1 in the Villabruna Cluster may have interbred with ancient populations related to East Asians as well, which was why excess allele sharing with East Asians was detected. Moreover, the ‘Villabruna Cluster’ is composed of 15 post-Last Glacial Maximum individuals from 14,000–7,000 years ago. Of these fifteen Villabruna samples, older twelve samples belonged Haplogroup R or R1 and haplogroups R1b and R1b1 only appear in more recent three samples, which are dated around 7,000 years ago (Table S4.2. Details of Y haplogroup SNPs in Villabruna Cluster samples).

So the R individuals turn up towards the end of the mesolithic and are not found in the older samples. Another indication for me that they do came from Southeast and combined with the Dstat results the conclusion of the paper makes sense.
 
Good points. After all R1b wasn't found among acquired HG Y DNA by early farmers in Anatolia or Hungary. Either it was a rare mix into WHG, or lost tribe who wandered in Italy and died there, or perhaps contamination?
the thing is I don't believe it came from Anatolia as claimed many times in the past I think those Villabruna individuals got some of their ancestry via the Iranian Plateau and came through the Caucasus. This "Near Eastern ancestry" what they call is probably something that resembles CHG and not EEF.
 
the thing is I don't believe it came from Anatolia as claimed many times in the past I think those Villabruna individuals got some of their ancestry via the Iranian Plateau and came through the Caucasus. This "Near Eastern ancestry" what they call is probably something that resembles CHG and not EEF.
The mystery is that there was no R1b found till late Neolithic/Bronze Age in Anatolia and Balkans. If there was a serious movement of R1b HGs from Iranian Plateau or else, they got swallowed by WHGs, leaving just trace of Near Eastern. It is rather unlikely that WHG comes from Iranian Plateau, but possibly just R1b haplotype.
I'm thinking that WHG picked some R1b in refugium in Anatolia from other HGs close by, let it be R1b of Iranian Plateau or Kurdistan.
 
The mystery is that there was no R1b found till late Neolithic/Bronze Age in Anatolia and Balkans. If there was a serious movement of R1b HGs from Iranian Plateau or else, they got swallowed by WHGs, leaving just trace of Near Eastern. It is rather unlikely that WHG comes from Iranian Plateau, but possibly just R1b haplotype.
I'm thinking that WHG picked some R1b in refugium in Anatolia from other HGs close by, let it be R1b of Iranian Plateau or Kurdistan.


I don't think it is unlikely. I also don't understand how some poeple think "this is impossible" because they believe yDNA I is all European component and backmigrated to West Asia.

If welook at the yDNA tree and take into account some studies which came out a few years ago we will see that it makes quite frankly sense.

I is the brother clade of J and both must have their ancestor evolved in the same region. This can be the Caucasus or the Iranian Plateau. But I think the Caucasus or South_Central Asia is a pass away. Than we have the very first IJ* samples on the Iranian Plateau- which indicates that at least the ancestor of both I and J must have lived in close proximity or maybe both Haplogroups diverged even there.

Now the Iranian Plateau is connected to South and Central Asia by land and this means ancient samples from there and indeed also modern samples show some affinities to East Asians.

Now the Steppes could also be the source but if the Steppes were the sources the authors wouldn't mention something very West Asian in their DNA that shows up in Dstats.

I assume the authors know already more than they presented us.
 
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Razib Khans opinion on this. Interesting allot of his points agree with my opinion.

The map and chart above is from The genetic history of Ice Age Europe, a new paper in Nature from the Reich lab (the new data has been posted). It illustrates probably the major finding of the paper, using a ~40,000 year paleogenetic transect of 51 ancient DNA samples the authors conclude that there have been at least three major population turnovers/disruptions across Pleistocene Europe. These correspond to three genetic clusters that they’ve identified in their data; the El Mirón, Věstonice, and Villabruna groups. Respectively they are the Magdalenian, Gravettian, and Epigravettian/Azilian cultures. There are also stray individuals which are harder to place, but signal other turnovers. An individual from Goyet that dates to 37,000 years ago and was presumably of the Aurignacian culture, and is somewhat sui generis. But, unlike the ~40,000 year old sample from Romania, and the ~45,000 year old Siberian, Goyet is ancestral to some later Europeans.
The figure to the left is one interpretation of their results. It shows that the Goyet sample contributed substantial ancestry to the Magdalenian culture which flourished nearly 20,000 years later. But, Goyet did not contribute substantial ancestry to the Gravettian culture, which succeeded it! Rather, the Vestonice cluster which represents the Gravettians has only marginal admixture from other Pleistocene Europeans, but a notable affinity to the Konsteki sample. Intriguingly, Goyet-like ancestry can be found in the Loschbour hunter-gatherer from the Holocene.
This suggests that I was wrong in one of my predictions: I had assumed that most European hunter-gatherer ancestry dates to the Gravettian at the earliest. This is wrong. In fact, this paper suggests minimal legacy of Gravettian peoples as represented by the Vestonice cluster. Rather, earlier peoples have left their mark on modern Europeans via the Holocene “Western Hunter-Gatherers” (WHG) who mixed with incoming farmers.
There’s more. Most of Loschbour’s ancestry is not from Goyet-like groups. Rather, it is from a population with affinities to the Villabruna culture. This to some extent vindicates another prediction I made: that most European hunter-gatherer ancestry would probably date to groups which became established after the Last Glacial Maximum 20,000 years ago. That was right, as Villabruna-like affinity seems to be the dominant signal in the Mesolithic hunter-gatherers.
700px-Weichsel-Wu%CC%88rm-Glaciation-300x243.png
Some of the patterns above are perplexing. So at this point, I think I want to drop a conjecture which I think can be inferred from this paper, but probably will have to be explored with future results and analysis: the Villabruna cluster ~14,000 years is a product of a massive expansion of a hunter-gatherer population from the Middle East. The original papers which posited that “Early European Farmers” (EEF) were admixtures between “Basal Eurasians” (BEu) and WHG, at 40% to 60% proportions, were somewhat misleading I suspect. Rather, WHG, the Mesolithic hunter-gatherers of Europe, derive predominantly from an expansion of Middle Eastern hunter-gatherers which had larger populations in the wake of the grueling climatic regime of the Last Glacial Maximum. The WHG in EEF was not European hunter-gatherer at all, but local Middle Eastern hunter-gatherer.

The further affinities of Villabruna make a likely exotic origin obvious. As noted in the paper a Near Eastern, but not BEu, affinity of European hunter-gatherers emerges specifically with Villabruna, ~14,000 years ago. And, some individuals in this cluster likely exhibit admixture from a population related to modern East Asians. This gene flow is independent of the Middle Eastern gene flow, though I suspect that the Middle Eastern gene flow is simply an expansion of hunter-gatherers from that region, with some absorption of the local substrate. There are other explanations for why this affinity might exist (read the supplements), but other papers have indicated the possibility of this relationship, so it is probably the most likely. The Middle Eastern origin of Villabruna makes more sense of the relationship between it and “Caucasus Hunter-Gatherers” (CHG). Geographically they would have been positioned near each other.

These West Eurasian clusters form a very deep clade with the Ma’lta North Eurasian population as an outgroup, with these nested together with East Eurasians, Amerindians, and Oceanians, in comparison to BEu. But if you take a look at the tree, and consider the chronology, it seems that modern Eurasians diversified into several distinct lineages over the course of 5-10 thousand years after the Out of Africa event. Individuals on the ~40,000 year time limit are no more related to all Eurasian groups, perhaps because their lineage went extinct. Ma’lta and the North Eurasians seem to have diverged from other West Eurasians very soon after these two diverged from East Eurasians; there just isn’t that much time to allow for this, but it did happen.
By about 30,000 years ago many of the pieces were in place. Much of the demographic change we see subsequent involve a set of operations to mix and match basic elements. The patchiness and segregation of these populations is probably why ancient DNA, itself spotty and poor and seeing the landscape with precision, assigns all of Europe to a particular cluster at a particular time. There were clearly other peoples, but they are not always at accessible archaeological sights, or perhaps they had retreated into the forests as a marginal folk?
There are many other interesting aspects of this paper, such as the Neanderthal admixture. But I’ll save that for another day….
 
Razib Khans comment on a question.

What exactly is that relationship? Don’t CHG have a lot of Basal Eurasian? Yet the Villabruna didn’t have any. But the EEF had Villabruna ancestry and Basal Eurasian? But not the exact same Basal Eurasian as in CHG (diverged by more than 20,000 years)?

the villabruna/el miron share drift with CHG via the NO-basal eurasian. and i believe the issue

So the affinity these Villabruna samples show resemble the non BE ancestry of CHG more than the EHG ancestry. Another reason for me to assume that some R samples will turn up in further CHG samplings. And the East Eurasian Han like ancestry is from a different event/source which is also atypical for EHG as well CHG but support a region in close range to East Eurasian ancestry. Something tells me this might possibly even be South_Central Asia.
 
Razib Khans opinion on this. Interesting allot of his points agree with my opinion.
There’s more. Most of Loschbour’s ancestry is not from Goyet-like groups. Rather, it is from a population with affinities to the Villabruna culture. This to some extent vindicates another prediction I made: that most European hunter-gatherer ancestry would probably date to groups which became established after the Last Glacial Maximum 20,000 years ago. That was right, as Villabruna-like affinity seems to be the dominant signal in the Mesolithic hunter-gatherers.
Well, yes and no. I believe most of us suspected that population of Europe grew back from HGs coming from refugium. We didn't expect them to be so dramatically different than before. I imagined that cold weather pushed them into refugium in first place, but this is not true (except just handful of them). They didn't go into refugium, they've froze and died out instead. Just like Neanderthals.
In this case the refugium is not really a refugium. Europe was repopulated by HGs who lived in the South like South Balkans and Anatolia, even before LGM. Well, the last statement is a bit of supposition, but should be close to the truth.
 

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