The genetic history of Ice Age Europe

I wish I would have been there to see the expression on his face, if he has seen it yet. I bet he was jumping up and down, screaming..
It's nice to see he was proven right, about finding ancient R1b in Italy.Not many were as bold as he was in his prediction. Already you could see when the R1b-V88 ancient Sardinia results came in on Yfull that R1b had a ancient presence in Italy.

When you think of the abuse he took over the years from certain people, people who don't even have the good manners to acknowledge that he was at least correct that it would be found in Paleolithic Italy. He just had faith in his analysis, and stuck to it. It's rather remarkable, I think.

Now, the question is how long was it there, where did it come from, and where did it go.
 
If you look at the R1b tree, I posted you will see that R1b-Z2103=Z2106=Z2108/9=Z2110/CTS7822 are all dated to 6100+/YBP
https://www.yfull.com/tree/R1b/
Gioiello- Belongs to R1b- Z2110*
My branch is same R1b-9219+ found in Eastern Europe and Digor Ossetians.
There are not to many of our R1b branch in your map. There is no variance in our branches of R1b. in Iraq/Iran;and they are just as old as R1b Z2103+
Oh, I know what you mean. Thanks for this tree btw. I see it for the first time.

But some things doesn't make sense to me. This tree is not complete or full mistakes. If Z2109* = 6200 YBP, while Poltavka = 4800 YBP (on this map), then Poltavka can't be exactly the same as Z2109* (6200 YBP), because there is a gap of almost 1500 years in between, right?
And therefore I don't see any direct links between Poltavka and Pathan. This tree is saying that Pathan mutated directly from Z2109 (which is 6200 YBP) and not from Poltavka. Doesn't make any sense either, because Pathan is much younger than Z2109*.


Further, I don't think Z2110* is from Poltavka, but it is actually native to Yamnaya. When Z2110* mutated Poltavka didn't exist yet (according to this tree).


It seems that different subtypes of R1b migrated into the Yamnaya Horizon at the same time. Not only 1 subtype but a couple of cousin subtypes of R1b.

From this map I got that your branch is from Z2110* and that Z2110* is a downstrem of Z2109*. After Z2109* really settled down in the Yamnaya Horizon it became really native to that region and started to change/mutate. Your branch is native to East Europe.
I don't know anything about Digor Ossetians, but if it is true what you're saying (and I believe you) then your subtype could be also part of the 'European' Scythians who brought R1b to the Ossetians.
 
Fire Haired14;479447]I doubt it because WHG, who they think might have come from SE Europe/Near East, was essentially 100% mtDNA U
.

I didn't mean that "H" got to Europe that early. I meant that this particular warming period may have been the time when populations started to increase, and new lineages arose.

As to "H" in Europe, are any of these samples the same ones from Spain that were supposedly tested previously and found to to "H"? If that's the case, then it's sort of case closed, isn't it? MtDna "H" in Europe would be Neolithic, except for that stray from the northeast which probably arrived from the Caucasus.

Speaking of mtDna. They found mtDna "M" in this era too, and U6. So, U6 is probably a back migration to North Africa and just got very lucky there. As for "M", I don't know why the authors say it doesn't exist in Europe. It does; I know it can be found in southern Italy. Now, without all sorts of detailed sub-clade analysis it's impossible to know or speculate when that mtDna "M" arrived there, but I think we should know by now that Italy harbors some very ancient haplogroups.
That makes a lot of sense. Even our 11,000 year old genome from the Southern tip of Italy is a typical WHG person. So, how could WHG be from Spain if it was so far SouthEast so long ago? Doesn't make sense. Our oldest examples of WHG so far are from Northern Italy(14ko),Switzerland(13ko), and France(13ko). 19,000 year old girl from Spain looks like a mixture of WHG and people who lived in Belgium 30,000 years ago. She did clearly have lots of WHG-affinity, but wasn't pure WHG.

I believe most of the 15,000 year old samples from Germany and Belgium are just like that Spanish lady, then suddenly after 15,000 ears ago everyone became WHG. But I'll have to check the paper to see if that's what happened. If that is the case, then WHG was a newcomer maybe from the SouthEast.

I'm not sure if that's the case. They seem to be hedging their bets by saying it could be sub-stucture in Europe.

This all leaves the role of "Basal Eurasian" still murky for me. If this Villabruna like group(which includes La Brana and Loschbour), came from the southeast via Italy, and that's why it's "related" to Near Eastern populations according to the authors, why does it have no Basal Eurasian? That would mean there was no Basal Eurasian in Anatolia in 14,000 BC? So when did it arrive and from where?* They don't talk about the farmers of Anatolia, but they mention that CHG, which is much older, was about 30% Basal Eurasian. So how would that have worked? A non-Basal Eurasian Near Eastern group of hunter-gatherers leaves for Europe from Anatolia. Then, somehow Basal Eurasian arrives in Anatolia? How could it have come from the south if it was already present earlier in the Caucasus?

I have to re-read the paper tomorrow as well, because I don't understand how the re-population of Europe after the LGM by people who were part GoyetQ116-1 as well as part Villabruna could have led to all these analyses showing WHG admixture.

Also, what the heck yDna did they carry?
 
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I cannot see how he was correct. Gioiello was only concerned about his particular subclade of R1b found commonly in the North of Italy. There are hundreds of subclades of R1b in Europe, and elsewhere. Finding one subclade in Europe among ancient Europeans does not prove Gioiello right. Personally his whole argument about R1b in Europe and Italy in ancient times was based purely on his own jingoistic bias and his own haplogroup.
 
I think that R1b-Z2103 is from the Maykop culture and Indo-Europized the Yamnaya Horizon. By the time when R1b-Z2103 migrated into the Yamnaya Horizon (5500 YBP), the Y-DNA hg. R1a-Z283 & R1a-Z93 already existed.

"Indo-Europeanized", even though all the elite graves are R1b? It doesn't appear that R1b is ancient in the Caucasus, several studies have already arrived at this conclusion. ie: http://digitalcommons.wayne.edu/humbiol_preprints/54/
 
"Indo-Europeanized", even though all the elite graves are R1b? It doesn't appear that R1b is ancient in the Caucasus, several studies have already arrived at this conclusion. ie: http://digitalcommons.wayne.edu/humbiol_preprints/54/

Everybody had me convinced it wasn't ancient in Europe, either, so who knows where it was. As someone else mentioned, R1b people are apparently the restless sort.
 
I've added mtDNA/Y DNA results to my spreadsheet with the list of samples. I also labelled each according to the cluster they were assigned using F3-stats.

New Paleo European Genomes

The ~30,000 year old Italian and Central European samples form a cluster. They're closer to WHG than Kostinki and 30,000 ear old Belgium are. Their Y DNA is C1a and IJKH(inlu. confirmed I). Their mtDNA is U2*, pre-U5*, U8*, U2'3'4'7'8'9*, and M. Looks like a good canidate for the ancestor of WHG.
 
The Conclusions section:

"We show that the population history of pre-Neolithic Europe was
complex in several respects.

First, at least some of the initial modern
humans to appear in Eurasia, exemplified by Ust’-Ishim and Oase1, failed to contribute appreciably to the current European gene pool.Only from around 37,000 years ago do all the European individuals analysed share ancestry with present-day Europeans.

Second, from
the time of Kostenki14 about 37,000 years ago until the time of the Villabruna Cluster about 14,000 years ago, all individuals seem to derive from a single ancestral population with no evidence of substantial genetic influx from elsewhere. It is interesting that during this time, the Mal’ta Cluster is not represented in any of the individuals we sampled from Europe.

Thus, while individuals assigned to the
Gravettian cultural complex in Europe are associated with the Věstonice Cluster, there is no genetic connection between them and the Mal’ta1 individual in Siberia, despite the fact that Venus figurines are associated with both. This suggests that if this similarity is not a coincidence, it reflects diffusion of ideas rather than movements of people.

Third,
we find that GoyetQ116-1 derives from a different deep branch of the European founder population than the Věstonice Cluster which became predominant in many places in Europe between 34,000 and 26,000 years ago including at Goyet. GoyetQ116-1 is chronologically associated with the Aurignacian cultural complex. Thus, the subsequent spread of the Věstonice Cluster shows that the diffusion of the Gravettian cultural complex was mediated at least in part by population movements.

Fourth, the population represented by GoyetQ116-1 did not disappear,
as its descendants became widespread again after ~19,000 years ago in the El Mirón Cluster when we detect them in Iberia. The El MirónCluster is associated with the Magdalenian culture and may represent a post-Glacial Maximum expansion from southwestern European refugia.
Thanks for posting Angela. It means a lot to me and others lacking time to read whole paper.

Fifth, beginning with the Villabruna Cluster at least ~14,000
years ago, all European individuals analysed show an affinity to the Near East. This correlates in time to the Bølling-Allerød interstadial, the first significant warming period after the Glacial Maximum.

Archaeologically, it correlates with cultural transitions within the Epigravettian in southern Europe and the Magdalenian-to-Azilian transition in western Europe. Thus, the appearance of the Villabruna Cluster may reflect migrations or population shifts within Europe at the end of the Ice Age, an observation that is also consistent with the evidence of mitochondrial DNA turnover. One scenario that couldexplain 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.
There was a give away when Anatolian farmer genome was published last year. They contained WHG admixture even before entering Europe. I imagined some sort of hunter gatherer "refugium" in Anatolia, and later repopulation throughout Europe. Now it becomes more obvious. I think someone back then proposed name change to Anatolian HG from WHG.
Sixth, within
the Villabruna Cluster, some, but not all, individuals have an affinity to east Asians.
Now, this is something new, isn't it? Do they say anything about African admixture showing in some Mesolithic hunter gatherers?

An important direction for future work will be to generate
similar ancient DNA data from southeastern Europe and the Near East to arrive at a more complete picture of the Upper Palaeolithic population history of western Eurasia."
Yes, yes, yes! Though it might take another couple of years for a complete research. They should include North Africa too.
This possible movement into Europe from the Near East connected to the Bølling-Allerød interstadial warm period may correlate with the sudden expansion of certain mtDna lineages leading to "H".
I always thought it was Mesolithic expansion. Probably together with Anatolian WHG. No wonder they could have brought some R1b from over there too.

Also, it's interesting that they're pointing to the Herc2 blue eyes mutation as coming from the southeast. Didn't an old paper trace its expansion to somewhere around the Black Sea about 10,000 years ago?

If the refugia for the Villabruna cluster was in Europe and not West Asia, could it have been in and around the now submerged Great Adriatic plain?
Blue eye Anatolian Hunter Gatherer. Another twist of the story. Nice.



 
I think the main conclusion of the paper is the identification of the 5 clusters.
How reliable this identification is, the future will show.
 

Archaeologically, it correlates with cultural transitions within the Epigravettian in southern Europe and the Magdalenian-to-Azilian transition in western Europe. Thus, the appearance of the Villabruna Cluster may reflect migrations or population shifts within Europe at the end of the Ice Age, an observation that is also consistent with the evidence of mitochondrial DNA turnover. One scenario that couldexplain 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.

So far we know that:

- Aurignacian people in central and western Europe belonged to mt-haplogroups M and R* and Y-haplogroup C1a, while contemporary people in Romania and Russia belonged to U*, U2 and U6 and Y-haplogroup CT and C1b.

- Gravettian people belonged to mt-haplogroups M, U*, U2, U5 and U8c, and Y-haplogroups BT, CT, C1a2, F, IJK and I.

- Magdalenian people belonged to mt-haplogroup U5b and U8a and Y-haplogroups HIJK and I.

- One Epigravettian person from northern Italy belonged to mt-haplogroup U5b2b and Y-haplogroup R1b1*.

- One Azalian sample belonged to mt-haplogroup U5b1h and Y-haplogroup I2.

- Epipaleolithic samples from France and Germany belonged to mt-haplogroup U5b1 and U5b2 and Y-haplogroup I.


Note that six years ago I attempted to retrace the original mtDNA of Proto-Indo Europeans and I concluded that the earliest lineages associated with Neolithic R1b cattle herders a few millennia before the Yamna period were U5, J1b1a and V. I explained that R1b originated in Central Asia and spread around the Caspian Sea, northward to Russia and southward to Iran and Kurdistan. Some tribes that had ended up in northern Mesopotamia eventually domesticated cattle. J1b1a was picked up in the northern Msopotamia before R1b cattle herders split in two groups, one that migrated across the Caucasus to the Pontic-Caspian Steppe (R1b-M269) and one that migrated south to the Levant and Africa (R1b-V88). So it is not surprising to see that the mitochondrial haplogroup associated with an even earlier R1b hunter-gatherer should be either U5 or V.

U5 is a very old haplogroup that could be up to 50,000 years old by some estimates. Yet U5 is completely absent from the Aurignacian and the oldest U5 sample to date is a Gravettian dating from 30,000 years ago. Since Gravettian brought Y-haplogroup I to Europe from the Middle East or Southeast Europe, the origins of U5 would also appear to be more eastern. It is now known that the But given the Gravettian culture xtended as far east as Romania and Bugaria and possibily even Russia. Gravettian were big game hunters and included groups of mammoth hunters like the Ancient North Asians (Mal'ta boy) who belonged to Y-haplogroup R. Given the paucity of U5 in the Middle East today, it would make more sense if U5 originated in Eastern Europe and/or western Siberian with big game hunters, belonging both to Y-haplogroup I and R.

In Paleolithic Europe we see a sharp increase in mt-haplogroup U5 from about 15,000 ybp. This also corresponds to the disapperance of older mitochondrial (M, N, R, U*, U8) haplogroups from the gene pool and the overwhelming replacement of old Y-DNA lineages (BT, CT, C1a, F, IJK) by R1a and R1b in eastern Europe and Y-DNA I in the rest of Europe during the Epipaleolithic and the Mesolithic.

In other words, the Bronze Age invasion from the Steppe was not the first major population replacement that originated in Eastern Europe. It looks like an Epipaleolithic wave of big game hunters belonging mostly to Y-DNA I2 and mtDNA U5 swept over Europe from the Steppe.

So the Gravettian may have originated in Southeast Europe and sent waves of Y-haplogroup I to both western and eastern Europe. But the latter came back 15,000 to 20,000 years later as (mostly) Y-haplogroup I2 and mt-haplogroup U5 and replaced most of the older male and female lineages in Europe. A few R1b may have ended up with them, which is why an R1b1* showed up in Villabruna.
 
When was R1b-Z2103 Indo-Europeanized then and by which subtypes of R1a? Because at the time of Yamnaya culture, where Z2103 is found, there was already a split between R1a-Z283 and R1a-Z93.

If R1b-Z2103 was Indo-Europeanized around the Yamnaya Horizon then it has to be by R1a-Z93. But this whole story is way to one-dimensional and to simple.

I'm sure something else happened. What we know for sure is that Yamnaya culture was Indo-European and that R1b-Z2103 was part of that culture. Yamnaya is the OLDEST Indo-European culture in Europe.

Yes, by R1a-Z93 (and possibly by other R1a)

IMO R1b-M269 / R1b-L23 were the guys who invented metallurgy. Later as we know L23 split into two branches.

In my opinion R1b-L51 went directly to Western Europe without "visiting" the Steppe on their way there.

They brought metallurgy to Western Europe. And R1b-Z2103 brought metallurgy and "Teal" ancestry to the Steppe.

Metallurgy was invented most likely in the Middle East - that's where both L51 and Z2103 expanded from.

Yamnaya people were a mixture of these two:

Smiths - R1b-Z2103
Commoners - R1a

That's why we see Z2103 in kurgans, in which smiths (who were at the same time chieftains) - not commoners - were buried.

Men buried in kurgans was not the whole population, but only "shamans" (and metallurgy was their "magic") or chieftains.

===========================


Where is Yamnaya L23(xL51) and especially L23(xL11) in western Europe today? There is none.

Once you cross the Czech-German border or the Slovene-Italian border, almost all of R1b becomes L11.

The last countries with significant (over 5%) frequency of L23(xL11) are Slovenia, Czech Rep. and Poland.

Frequency of L23(xL51) in Germany is only around 0.5% (see: Natalie Myres 2010).

It is clear that Bell Beaker expanded West-->East and encountered CWC in Rhineland.

CWC was R1a with a minority of R1b-L23+ (but actual L23/Z2103, not L51 or L11).

Non-L11 subclades of R1b-M343 (including Non-M269 & Non-L23):

R1b-M343(xL11):

Poland (n=202) - 6,44%
Slovenia (n=102) - 5,9%
Czech Rep. (n=87) - 5,7%
Slovakia (n=276) - 5,0%

Germany (n=321) - 1,83% (including Non-M269)

And now only R1b-M269+, L23+ which is actually R1b-L23(xL51):

R1b-L23(xL51):

Czech Rep. (n=87) - 5,7%
Poland (n=202) - 5,44%
Slovenia (n=102) - 3,9%
Slovakia (n=276) - 3,6%

Germany (n=321) - 0,62%

This data is from Myres 2010:

http://www.nature.com/ejhg/journal/v19/n1/suppinfo/ejhg2010146s1.html

So in a sample of 321 Germans, only 2 had L23(xL51).

And Germany is in Central Europe, not Western Europe.

Go to Western Europe and you will find hardly any L23.

R1b-L11 expanded West->East, in the opposite direction to how R1b-L23(xL11) expanded...
 
"Indo-Europeanized", even though all the elite graves are R1b?

In France and in Italy Germanic elites were "Latinized" (started speaking Romance languages), not the other way around.

What we see in kurgan graves are smiths - who were at the same time shamans and/or chieftains - community leaders.

They were not the majority of the population. This refers also to cultures in which kurgans were full of R1a (like Srubna).

Simply in all of those Steppe cultures, kurgans were burials of ruling clans / leading families, not of "average Joes".
 
R1b was IMO native to the Near East and there were numerous successive emigrations of R1b from that area.

Villabruna represents one of the earliest migrations of R1b from the Near East to South Europe. Later other waves came. R1b migrated both to the Steppe (Z2103 was "intrusive" in Yamna - perhaps came from south of the Caucasus with Maykop?). Z2103 was not the majority of Yamna males - but they brought the "magic" of metallurgy, and became "shamans" / "chieftains". That's why we see them overrepresented in kurgans.

R1b-L51 in my opinion expanded directly from the Middle East to Western Europe, bringing Copper Age to Iberia (ATP3).
 
I don't think that there was ever such a thing as "replacement of R1b by R1a in the Steppe after Yamna".

Yamna was probably R1b-Z2103 minority (but "smiths-shamans") + R1a majority (but "commoners").

And what we see later on, is the loss of high status by Z2103 and R1a men becoming "chieftains".

However, Z2103 continued to exist as a minority lineage and went to India - as Parasar wrote:

parasar said:
R1b:
1. Oldest R1b on the P297 line found in Italy ~14000ybp
2. Oldest R1b on the V88 line found in Spain ~7000ybp.
3. Oldest R1b on the M269 line potentially found in Spain ~5500ybp
4. Oldest R1b on the M73 line found in Samara ~7600ybp

Lets see how this comports with modern DNA evidence from India.
1. L278xL389 lines are present in India - trace amounts (eg. N93357 Joshi P25+, L21-, L23-, L51-, M18-, M222-, M269-, M335-, M73-, P297-, P312-, U106-, U152-, V88-, L389- & 267597 Raza Varanasi R-M343 https://www.familytreedna.com/public/r1b1asterisk/default.aspx?section=ysnp) - perhaps remnants of early R1b moving west along with other early R/Q lines (incl. R1a and R2xM124).
2. V88 completely absent in India because V88 splits in the Mediterranean region and does not move east.
3. M269 present in present in India beyond trace amounts - of the Yamna type - likely enters India with R1a-Z94 lines.
4. M73 - trace or absent in India.

What he wrote does not contradict my theory about Maykop smiths with "M269 of the Yamna type" (= Z2103 and L23*).

But let's add also information about L51 in India:

5. L51 completely absent in India because L51 splits from L23 in the Mediterranean region and does not move east.

There might be some L21 or U106 though - due to British occupation of India in recent history.
 
BBC article about this study, with photos of some skulls - including Villabruna's:

http://www.bbc.com/news/science-environment-36150502

"This 14,000-year-old individual from Villabruna, Italy, lived at a time when the climate was warming up":


_89538772_89538771.jpg

Is it sure this interesting skull is this famous Y-R1b's one? If true have you a profile (lateral) picture?
 
My current assertion is that M269 & L23 were responsible for spreading early metallurgy around Western Eurasia.

It does not require the presence of M269/L23 already in Neolithic Europe.

It also does not imply that L23 originally spoke PIE, because we know well that metallurgy was not invented by PIE people.

The PIE community only adopted metallurgy from Non-IE people who were spreading it.

Villabruna is a proof that R1b expanded from the Near East in several succesive waves between Upper Paleolithic and Metal Ages.

EEF autosomal DNA is still more abundant in Western Europe (especially South-Western, but in fact also in North-Western) than is Steppe autosomal DNA. By contrast, G2a haplogroup is very rare in that region. This implies that initial EEF population (G2a-heavy) got extinct, and was replaced by another wave of Near Eastern immigrants - bringing with them similar EEF admixture, but different Y-DNA.

They were R1b folks spreading metallurgy as well as EEF and "Teal" ("Armenian") admixtures.

================

RISE 413 from Middle Bronze Age Armenia was R1b-P297* according to Sergey Malyshev.

He lived very late, but it is possible that P297 (and M269) was native to areas south of Caucasus:

RISE413.png
 
my conclusions so far :

Aurignacian was C1a2 - same branche as La Brana (C-V20xV86)
Gravettian in central and western Europe developped after arrival of I and arrival of J in eastern Europe (north of the Black Sea) around 35 ka
Vestonice - Gravettian cluster 26-34 ka is a mixture of C1a2 and I
Mal'ta cluster 24-17 ka is P1,Q,R
El Miron - Magdalenian cluster 19-14 ka is I and mtDNA U8a
Satsurblia - Epigravettian cluster 13 - 10 ka is J ; Epigravettian originated in Europe but expanded to Transcaucasia & Armenia 18 - 19 ka
Villabruna cluster with microliths and during forestation of Europe was triggered by immigrants from the Near East 14 ka, one of them was R1b pré-P297 but not ancestral to R1b M269 neither R1b M73 ; the main component is I, with few C1a2
the paleolithic European C1a2 is extinct, what is left in Europe today is neolithic C1a2 with origins in the Levant (C-V86)
I seems to have partly dissapeared during neolitisation but expanded again at the onset of the bronze age, while C1a2 remained marginal

PS : Aurignacian C1a2 had higher Neanderthal admixture than I who arrived in Europe 35 ka
 

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