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Friday, June 26, 2015

Genetic substructures among Late Neolithic/Bronze Age Scandinavians


I may have discovered an interesting pattern in the Allentoft et al. data. It seems that during the Late Neolithic/Bronze Age, Scandinavia was populated by two somewhat different populations; one characterized by Y-Chromosome haplogroup R1b and a genome-wide genetic structure typical of present-day Northwestern Europeans, and another by Y-Chromosome haplogroup R1a and a relatively more eastern genome-wide genetic profile.

Below are two Principal Component Analyses (PCA), both featuring ancient Swedish genomes classified as part of the Late Neolithic Battle-Axe archeological culture. However, the first sample clusters near present-day Norwegians and belongs to Y-haplogroup R1b-U106, which is nowadays typically known as a Germanic paternal marker. On the other hand, the second sample clusters among present-day Russians and Mordovians, from all the way near the Volga, and belongs to Y-haplogroup R1a-Z645, which very likely expanded from Eastern Europe during the Late Neolithic.




Here's another example of basically the same thing, but this time with two ancient genomes from Denmark. If you're having trouble finding the ancient samples, download the PDF files and type their IDs in the PDF search field.



Coincidence? Probably not, but we obviously need more samples to confirm these results and establish that there is indeed a pattern.

Citation...

Allentoft et al., Bronze Age population dynamics, selection, and the formation of Eurasian genetic structure, Nature 522, 167–172 (11 June 2015) doi:10.1038/nature14507

111 comments:

Mike Thomas said...

Dave
Do we know what Y groups the neolithic samples belonged to (eg Gokhem etc) - if tested ?

Davidski said...

Gokhem is a female. I don't think we have any decent male genomes from Middle Neolithic Scandinavia. But they'll probably turn out to be I2 and G when they do come.

Mike Thomas said...

Yep that's what I thought

Rokus said...

No, this certainly isn't any coincidence. Sarauw (2007):
'Another point of view is put forward by Vandkilde (1996, 296; 2005, 20). According to her, the Danish metal objects, supplies, and technology originated from north-western Europe, and she sees this as an indication of particularly tight bonds between the Veluwe area at the Lower Rhine and northern Jutland. This certainly seems a very convincing interpretation.'

The Beaker culture in Jutland (2350–1950 BC) was intrusive as well as influential. Flint resources were especially distributed in Norway and western Sweden. They also maintained a separate identity:
'People of northern Jutland probably did not feel any cohesion or social identity with people elsewhere in Europe or Denmark apart from areas where tight matrimonial connections existed.'

Referring to the Swedish RISE98 sample (Nordic LN, R1b1a2a1a1-M405/S21/U106*):
In the Nederlands R1b-U106 is diverse and there it also has its highest frequencies. I have been hypothesizing for years that my own Z18 derives from the 2200 BC Bell Beaker site Molenaarsgraaf, derived from Veluwe Bell Beaker, that is just 10 km from the origin of my surname. Being on top of riverdunes that originate from last stage of the Weichselien iceage, habitation since this period was never interrupted by floods. Veluwe beakers have some especially striking similarities with the westgroup of Funnelbeaker, so I still wait for some verification on this part as well.

Helgenes50 said...

The two first samples
with AmiK8 and Steppe

AmiK8

RISE98

0.72% Amerindian
0.01% Siberian
60.08% Euro_HG
0.57% Oceanian
1.07% Sub-Saharan
0.00% Southeast_Asian
22.19% LBK
15.37% South-Central_Asian

RISE94

0.18% Amerindian
0.00% Siberian
57.39% Euro_HG
0.00% Oceanian
0.17% Sub-Saharan
0.00% Southeast_Asian
23.43% LBK
18.83% South-Central_Asian

Steppe

RISE98

23.85% Near_Eastern
0.00% East_Asian
0.01% Siberian
0.33% Oceanian
41.47% WHG-UHG
1.01% Sub-Saharan
2.71% Hindu_Kush
30.18% Steppe
0.45% Amerindian
0.00% Southeast_Asian

RISE94

22.93% Near_Eastern
0.00% East_Asian
0.00% Siberian
0.00% Oceanian
37.07% WHG-UHG
0.36% Sub-Saharan
7.12% Hindu_Kush
32.52% Steppe
0.01% Amerindian
0.00% Southeast_Asian

Alberto said...

So similarly as with Yamnaya-CW-BB, we see this still not well understood phenomenon:

- Yamnaya being R1b and very eastern
- Supposedly derived from Yamnaya, but being R1a, CW people being more western
- Supposedly derived from CW, but again R1b, BB folk being even more western

Isn't it a strange pattern? And it repeats again with these Nordic samples.

Could be just coincidence: a founder effect of an R1a Yamnaya (though we still didn't find any), giving rise to CW. Then a founder effect of an R1b CW (which were minority there) giving rise to BB (and Nordic LN/BA). But it's not very parsimonious. There might be a better explanation once we get more samples.

Chad Rohlfsen said...

Rokus,
I predicted a while back that U106 will be from Barbed Wire Beaker, and some U106 as pre-Saxon. I got blasted for it, but I still think it's possible. It won't be of TRB origin though.

Mike Thomas said...

Alberto
I know you're not a y-Haplogroup -centric kind of guy, but what is your working hypothesis?

Davidski said...

Helgenes50,

Thanks, those test results back up my PCA. Good to see.

Btw, that K8 test is the ASI K8. Kurd made a mistake and called it AMI, and now everyone's calling it that. Haha.

Helgenes50 said...

David,

Ok, I'll rectify the mistake next time

Helgenes50 said...

About AsiK8 ( ex Ami). What is exactly the difference between the South-Central_Asian and the K9 Teal, we know that this one, in K8, is roughly composed of 37% ANE and 58/59% of NE.
And in the case of the South-Central_Asian ?

Davidski said...

They're very similar, but "Teal" peaks in the Caucasus, among Georgians. From memory, South Central Asian peaks among the Kalash.

truth said...

So what happened to the farmers in Sweden during the Bronze Age ? Did they already melted with the hunter-gatherers

Helgenes50 said...

@ David

Thanks

Fanty said...

Hmm.
So much for the decades old idea of
Modern Skandinavians = TBK (non- IE)+ Corded Ware (IE) mixed.

Those Bronce Age R1b guys are already like modern Skandinavians and the R1a are not.
Could it be by chance that all the R1b guys are mixed with locals and the R1a guys are not? -.-

Davidski said...

Neither of them are mixed with the locals yet all that much IMO. I think what we're seeing here is the entry of R1a and R1b into Scandinavia from different parts of continental Europe and/or via somewhat different routes.

Alberto said...

@Mike

Without more data is hard say. But if I had to try to guess something that made sense, one possibility could be:

- We have Yamnaya as purely R1b (for now, at least). My best guess (especially after seeing Armenian Bronze Age R1b samples), is that R1b was not a main EHG marker, but R1a was.
- If R1b entered the steppe with the Armenian-like population, EHG could have been pushed to some extent into CW territory, forming a proto-CW population (EHG + MN).
- Yamnaya population moved rapidly to the west, passing through the CW territory and having some genetic and cultural impact (CW proper), but finally mostly settling in Central Europe.
- Central Europe was more populated by MN people, who were at the time receiving the Bell Beaker influence from the (South?) West. Yamnaya people mixed with these early MN Bell Beakers, expanding rapidly throughout all of Western Europe.

Of course it's purely speculative, but it helps to make some things fit a bit better:
- CW having 70-75% Yamnaya input makes little sense with the haplogroups of each culture. Besides, CW is equally close to EHG as Yamnaya, but much closer to MN. This means it had some EHG input independent from Yamnaya. The fact that CW is archaeologically distinct from Yamnaya also doesn't support such a massive population migration. The influence (both genetic and cultural) was probably quite more moderate.
- Yamnaya people moving fast to the west and mixing with a larger MN population from Central Europe makes sense both with them being exclusively R1b and autosomally. These early BB are some 35-40% Yamnaya, and from there it diluted more (not much more when going to more scarcely populated areas like the British Islands, but quite a lot when going to Iberia).

What's still missing would be how to fit the other mysterious population that appeared in Hungary. They look like EN+Motala. And they seem to have had an important impact in modern European populations (it's difficult to NE Europeans as just mostly CW).

Some D stats of ancient populations' affinity to EHG (vs. Loschbour):
Loschbour EHG Yamnaya Chimp -0.0354 -6.545 341312
Loschbour EHG Corded_Ware_LN Chimp -0.0189 -3.32 341342
Loschbour EHG Unetice_EBA Chimp 0.0019 0.337 339167
Loschbour EHG Bell_Beaker_LN Chimp 0.012 2.154 338507
Loschbour EHG HungaryGamba_BR2 Chimp 0.0236 3.292 341755
Loschbour EHG Germany_MN Chimp 0.0314 5.105 316966
Loschbour EHG HungaryGamba_BR1 Chimp 0.0371 4.504 173274
Loschbour EHG Spain_MN Chimp 0.0401 6.949 335016

Some modern populations:
Loschbour EHG Russian Chimp 0.0101 2.062 341763
Loschbour EHG Czech Chimp 0.0159 3.111 341763
Loschbour EHG Belarusian Chimp 0.0171 3.413 341763
Loschbour EHG Bergamo Chimp 0.0178 3.563 341763
Loschbour EHG Lithuanian Chimp 0.0189 3.687 341763
Loschbour EHG Scottish Chimp 0.0202 3.771 341763
Loschbour EHG Spanish Chimp 0.0235 4.812 341763
Loschbour EHG Basque Chimp 0.0296 5.891 341763
Loschbour EHG Sardinian Chimp 0.0316 6.338 341763

Tesmos said...

Davidski, how about the I1 samples from Scandinavia? I wonder where they cluster.

Davidski said...

Only RISE175 has enough markers for this type of test.

Interesting result. Looks like I1a came from a WHG-rich pop.

https://drive.google.com/file/d/0B9o3EYTdM8lQWUNIeHV2QWpOSDA/view?usp=sharing

Matt said...

Any telling patterns on Eurogenes K15?

Simon_W said...

IMHO it's completely obvious that this is merely a matter of different ages.

This is zero evidence for the idea that R1a and R1b originated in different LNBA populations.

Because, consider this:

The R1b Swede RISE98 dates to 2150 BC.

The R1a Swede RISE94 dates to 2550 BC.

400 years are enough time to mix with the locals and to become more northwest European.

The R1b Dane RISE276 dates to ridiculously recent 670 BC.

The R1a Dane RISE61 dates to 2670 BC.

Please, these can't be compared!

Those who still think that R1b somehow had a different origin are barking up the wrong tree.

Simon_W said...

@ David

What's the source for RISE61 having R1a-Z284? To me that's particularly interesting, because I happen to be positive for that SNP.

According to Jean Manco's ancestraljourneys site he has R1a1a1b (S224/Z645), the source is indicated as "personal communication from author + aditional info from KSDA (Anthrogenica)".

Well, I'm not sure if KSDA from Anthrogenica is a reliable source.

Alberto said...

@Simon_W

I really don't know how it happened, but you have to admit that it's strange. We have 12/13 Yamnaya samples being R1b, the other one being I2a. Accepting that CW is 73% Yamnaya when CW is mostly (80-90%?) R1a is odd. And then accepting Bell Beakers being 50% CW, but 100% R1b is also odd.

Could all be just luck. But it's some pretty damn luck that's needed for that to happen.

When both ancient and modern DNA show R1a and R1b in different populations, why not just prefer to think that they were actually 2 different populations?

Simon_W said...

It might be useful to make a PCA of RISE1, the R1b1 Corded male from Poland. He dates to 2720 BC, so he belongs to the earlier members of the Corded Ware.

I guess he will come out rather eastern. But I'm also sure this will not convince the R1b from the west and the R1b from Hungary crowd. :D

@ Alberto

I guess R1a was more common towards the north/northwest, R1b more to the south/east. I would suggest that the origin of the Corded people was pure EHG who had been merely acculturated by Yamnaya influence, if (IF) they were not so damn similar to Yamnaya on the autosomes. They also have a considerable chunk of that Caucasus/West Asian admixture, which is also evidenced by their mtDNA, and this must have come from the south, via Yamnaya. They are only a little more EHG than Yamnaya is. At least that's true for the Corded Ware that entered central Europe and Sweden. The Estonian Corded Ware was different, with no West_Asian in K15.

And as I said in another thread, I would surmise that the Corded people were not completely homogenized in their yDNA proportions, and that there were pockets of R1b rather than single individuals in every village. I think that idea makes sense. If the Corded Ware R1b from Poland happened to have had many descendants and relatives on the paternal line, then there was a pocket of R1b there. And one of these pockets adopted the Bell Beaker package, which gave them apparently some advantage. In any case, the Bell Beaker people may have been fashionable and wealthy merchants and warriors and quite successful with women.

Archeological and genetic evidence favours this view, because: The Iberian R1b EEF turned out to be R1b-V88, and thus he's not at all a candidate for being ancestral to R1b-M269. And the Corded Ware had R1b1 at a very early date in central Poland. Thus, neither late nor particularly western and certainly predating the arrival of Bell Beakers in central Europe. And there was R1b-U106 in Late Neolithic Sweden, in a Corded Ware derived context. This R1b there predated the arrival of tin Bronze by centuries, and Sweden wasn't part of the Bell Beaker complex either. And moreover R1b-U106 is a close relative of R1b-P312, it's his brother clade. And last but not least the Yamnaya input in Hungary appears to have been of a rather different kind than Corded Ware and Samara Yamnaya, more western and without the West_Asian admixture, thus not like a typical R1a/R1b population, and indeed, although the Hungarian Yamnaya seems to have had quite some impact on Bronze Age Hungary, no R1b has turned up there so far.

bellbeakerblogger said...

David, makes sense. You may see the equal and opposite phenomenon among Beakers in Scotland clustering in the same hula-hoop with CW.

Grey said...

If you look at the bio-region map you can see the Atlantic coast bio-region covers the west half of Denmark and has small patches along the west coast of Norway.

http://www.eea.europa.eu/data-and-maps/figures/biogeographical-regions-in-europe-1/map_2-1_biogeographical-regions.eps/image_original

(There's also copper along the west coast of Norway and in the mountains.)

Grey said...

The bio-region thing may be relevant elsewhere too.

The division of Anatolia into three bio-regions may have been significant.

Also you can see by looking at the Mediterranean bio-region any farming that started in one part of it might be able to spread east-west quite rapidly but north or south of that zone maybe not so easily.

For example the parts of Anatolia that are/were part of the Mediterranean bio-region may have been easier to spread to than the parts labeled Anatolian or Black Sea.

Grey said...

Also the Hungarian plain is an exception to itself so it might not be surprising if it had a unique history.

Grey said...

Too many posts in a row - apols but

if R1b was an artisan minority within CW that might explain CW while also providing a reason why R1b types were running ahead of the main advance in such a way they could reap the benefit of founder effects along the Atlantic coast.

the question then would be how did that artisan association come about?

one option would be they happened to live on the Kargaly copper field.

another would be the first copper workers started elsewhere but were drawn to the Kargaly copper field via the Caucasus or Black Sea and the association with the locals started that way.

(if you look at the Samara bend it's plausibly the perfect spot for a trading post between the Black Sea and Kargaly)

either way - if this idea is correct or partly so - once the association was made you should see a correlation between R1b, steppe ancestry, red hair and metal working pretty much everywhere not just Europe - the difference being most places they stayed a small minority and only became a majority along the Atlantic coast cos founder effects.

probably wrong but fun all the same

Fanty said...

@David:
Yeah maybe they came from different routes. But my question was not "why are they different" but rather "why are the R1b people already "final version" while the R1a people look like as if their grandfathers might have been born in the steppe or something.

Modern Scandinavians have quiet large amounts of I1, R1b and R1a (>20% in Norwegians or Icelanders for example) If those R1b people can already pass as final, modern Scandinavians, where is the contribution of the I1 and R1a people to the modern Scandinavian genepool?

Or other:
Arent those R1b guys a little too western for so early or had they have a 500 years vacaction in France? ^^

Krefter said...

This pattern has been expected for a long time, because the R1a and R1b in Scandinavians obviously come from differnt sources.

@Fanty,
"If those R1b people can already pass as final, modern Scandinavians, where is the contribution of the I1 and R1a people to the modern Scandinavian genepool?"

The R1b-U106 guy probably had already ancestry from the Corded Ware-like R1a-rich people of Scandinavia. The Scandinavian paternal and autosomal gene pool had probably formed by circa 2200 BC, after a Western(more EEF and WHG) R1b-U106 and I1(?) pop mixed with an Eastern R1a(especially Z284 pop).

The I1 one way or another derives from pre-IE EEF/WHG Central-North Euros, because it was found in Neolithic Hungary. We don't have Y results for any of the Neolithic Swedes who lived about 1,000 years before the Late Neolithic I1 Swedish guy. Geneticker got I2 for Gok4 while Felix couldn't find a haplogroup(a relevant one at least). Modern I1 may mostly derive from a founder effect that occurred after Gok4.

Alberto said...

@Simon_W

Ok, I was not going for anything too radical, like R1b coming from Spain, or even a Hungarian route. Nor did I say that CW were pure EHGs acculturated by Yamnaya. I was just trying to find an alternative explanation to the "just by chance" complete turnover in haplogroups from Yamnaya to CW and from CW to Bell Beaker. But it's all too speculative, so I'll better wait for more data.

Unrelated: I was checking f3 stats from Allentoft et al. and while CW have some significant ones:

baCw hunterW baArm -0.0127780596 -10.1662437033
baCw neolHu baAfan -0.0083291953 -7.8082142527

Sintashta doesn't:
baSin neolC baOku -0.004324988 -2.4322022003
baSin hunterW baArm -0.0025766798 -1.7605615921
baSin neolC baAfan -0.0014268257 -1.0886681421

Still wondering what's the difference between Sintashta and CW...Anyone has any clue already about it?

Mike Thomas said...

Alberto

You have long been commenting on the role of Motala-like input jn Hungary. Given the large finds of (unofficially at least) I2a2a there could have something to do with this: an eastern admixed WHG group in East Central Europe.

Davidski said...

Simon, you're right, RISE61 is just R1a-Z645.

Rokus said...

'And there was R1b-U106 in Late Neolithic Sweden, in a Corded Ware derived context. This R1b there predated the arrival of tin Bronze by centuries, and Sweden wasn't part of the Bell Beaker complex either.'
As stated above, Sweden received flint resources from their Bell Beaker neighbors in Denmark. These neighbors were acquainted with metal objects and technology that originated from north-western Europe. To link the Bell Beaker complex exclusively with tin bronze is a strawman.

'I would surmise that the Corded people were not completely homogenized in their yDNA proportions, and that there were pockets of R1b rather than single individuals in every village. I think that idea makes sense.'

The assumption that R1b-individuals must be fully Corded Ware and eastern, notwithstanding the evidence stated above, is wrong. Tight matrimonial connections existed with neighboring Bell Beaker people in the west. Moreover, CW 'not completely homogenized'?? I understand that Bell Beaker R1b(xYamnaya), Yamnaya(xR1a) and CW(xR1b-L151) are three huge problems being worthy such Kurganist tricks, though please stick to the facts if making sense is indeed the purpose.

Krefter said...

@Davidski,

What about the Allentoft samples from Montenegro? Can you get anything useful out of them? Do they have ANE, who are they closest to, etc.?

Chad Rohlfsen said...

Barbed-wire Beakers were in Sweden.

Tesmos said...

R1b-U106/Rise98's grave/burial was not typical Bell Beaker but not typical CWC or Battle Axe either. So we can't exclude the possibility that he was part of an another population.

Alberto said...

@Mike

Yes, those samples from Hungary (at that time and place) are really exceptional in the sense that they don't seem to match with anything else that we know. IIRC, BR1 plotted between Basques and French, but north from them both. Basically that high WHG with a relatively small amount of ANE could only come from some Motala-like population + EN (or alternatively, from a pure WHG + Armenian). The most significant f3 stats for the new BA Hungary samples from Allentoft et al (hunterN = SHG):

baHu hunterW baArm -0.0111574792 -8.6858766695
baHu hunterN neolHu -0.0087444106 -7.1650028411

Where did they come from is a mystery. I2a could have existed anywhere around Europe, I would guess. But where could we find pure WHG/SHG at that time? Western Yamnaya or Northern Europe, seem more likely than other places. Maybe this population was related to CT? Or did it come from the north somehow? I honestly don't have any clue that points to any specific origin.

However, modern North Europeans are clearly shifter North West of CW, in the direction of this population (and also shifter towards WHG while CW was much more EHG). So a similar population to these Hungarians seems to have played a role in producing modern North Europeans.

Davidski said...

Krefter,

I can just barely run RISE595. But the result is interesting. I'd describe it as southeastern European before the major influx of West Asian admixture into the region during the Iron Age and medieval period.

https://drive.google.com/file/d/0B9o3EYTdM8lQc2FXb1NsYTRPWkE/view?usp=sharing

Rokus said...

According to Vandkilde (2005) barbed wire decoration in Southern Scandinavia characterises the period 2460-1990 BC, thus correlating with Nordic LN I and older. According to the shape the tradition is most likely derived from the Single Grave culture ( a Corded Ware), though part of the decoration may point at a Bell Beaker hybrid.
But: "The subject is difficult due to the fact that Late Neolithic pottery is insufficiently studied, and so far chronological groupings are not distinguishable."
Barbed Wire in the northern Netherlands comes after a period of Veluwe Style and Epimaritime Beakers. This tradition is not necessarily equal to Scandinavian Barbed Wire. The Dutch kind reaches Britain as one of the seven intrusive Beaker cultures, never referred to as Corded Ware. In the Netherlands archeologists recognize the existence of 'Corded Ware' but they don't make the difference between Corded Ware and Bell Beaker: like in Britain it's all Beaker people.
BTW, there is also some Mediterranean Barbed Wire, probably a degeneration of some other kind of Beaker-related culture.

Mike Thomas said...

David

"southeastern European before the major influx of West Asian admixture into the region during the Iron Age and medieval period."

Which influx are you referring to here ?

Davidski said...

The influx that created the "bridge" between Europe and the Near East on West Eurasian PCA. This bridge didn't exist when baMon was alive, and it shows up in shared IBD between some Europeans and Near Easterners. See here.

http://eurogenes.blogspot.com.au/2013/06/recent-gene-flow-from-africa-and-near.html

Only southeastern Europeans share IBD with Near Easterners. So their West Asian admix, or at least a large chunk of it, isn't from Yamnaya. If it was from Yamnaya, it would be too old to pick up via shared IBD with Near Easterners.

Mike Thomas said...

This influx was at the Bronze Age, then, at the latest. There are no documented Intrusions from Iron Age, much less the Medieval period, unless you;re suggesting Slavs came from Anatolia ?

Mike Thomas said...

And you never know. Z2103 in SEE might have come via West Asia (at least in part), as did other rarer subclades of M269 (like PF7558).

Simon_W said...

@ Fanty

It's the age of the samples, as I've just explained. The R1a dudes are early and still rather pure, the R1b guys by chance are later and more admixed with the locals. If RISE1, who had R1b but who was also early, was plotted on a PCA I'm extremely sure he would come out eastern like the early R1a dudes. This difference between the R1a and the R1b guys is just non-evidence. In the same way I could test a Swedish R1a guy from 700 BC and I'm sure he'll come out rather modern and northwest European. This would surprise no one, so why is it surprising that an R1b guy from 670 BC is modern and northwestern, while an R1a guy from 2670 BC isn't?

@ Krefter

That was just Genetiker's nonsensical theory. And of course it's perpetuated by people like Rokus who deny an eastern origin of R1b and who want to have the PIE originating in central or even western Europe. No it's not obvious to me that R1b and R1a in Scandinavians had different sources.

Frank said...

First of all, the IBD study uses Qataris, who do not carry west asian admixture. Then, there is also this:

http://www.isabs.hr/registration2013/__abstract_review/index.php?what=review&do=view&id_program=20&id_topic=54&id_abstract=427

Chad, in a previous discussion, said how there is dicontinuity in late neolithic greece as well, in the aegean palatial civilizations presentation. And most of that IBD sharing in the balkans is indeed in greece also.

Given how Allensoft also puts RISE595 more east than K8 and the effect of the slavic migrations in the balkans, post-bronze age west asian influx seems completely unlikely.

Would be quite hard to dillute the biggest eastern mediterannean population (greeks) in that time period. Plus, most of their slaves came from scythia, dacia and the coast of asia minor.

Simon_W said...

@ Rokus

I wouldn't link the Bell Beaker complex with tin Bronze because tin Bronze was still very unusual in the Bell Beaker complex, that's why it's classified as late Chalcolithic in German speaking countries. Strange you didn't know this. The reason why I spoke if tin Bronze was that RISE98 is partly classified as „Bronze Age Scandinavia“ in Allentoft et al, instead of „Corded Ware“ or „late Neolithic“, and this might make some people believe that the Bronze connection explains the R1b, but it doesn't.

Trade between neighbouring groups is a widespread phenomenon, and maybe there were even matrimonial connections, occasionally, why not. But you can't build theories that way, you can't explain disagreeable evidence away by making matrimonial connections with neigbours the default assumption. You still have to admit that basically Sweden wasn't Bell Beaker area. Even in Denmark the Bell Beaker presence was far from pervasive, there was most of all a Bell Beaker center in Northern Jutland. And if you want to derive Swedish R1b-U106 from trade contacts with Bell Beakers in Denmark you're close to circular reasoning: R1b must be from trade contacts with Bell Beakers, and this proves that all R1b is from Bell Beakers.

I never said that R1b people must be fully eastern on the autosomes, what a nonsensical idea. Autosomes and yDNA are inherited independently. And my assumption that Corded Ware wasn't fully homogenized is rather natural, if you'd think about it without bias.

I agree that Bell Beaker R1b(xYamnaya), but so what? The Yamnaya sample we have is still small. And I'm sure, if one day a Yamnaya male with R1b-L151 is found, you'd still not be convinced because he doesn't have L11... And CW(xR1b-L151) I disagree. There is RISE98 having R1b-L151.

>>In the Netherlands archeologists recognize the existence of 'Corded Ware' but they don't make the difference between Corded Ware and Bell Beaker: like in Britain it's all Beaker people. <<

Seems to be the most reasonable way to deal with it.

Davidski said...

Slavs obviously didn't come from Anatolia, but there were regular population movements between Europe and West Asia within the Roman Empire and Greece. Much of Greece was of course until recently in Anatolia.

We can also add to that the Muslim and Ottoman incursions into Europe, and it's easy to see why some southeastern Europeans fall into that space between the baMon and Cypriots on West Eurasian PCA.

Davidski said...

Frank,

That PCA wasn't done with the K8. It's based on genotype data.

And the reason Allentoft put RISE595 further east is because they projected their ancient samples on the plot. Projection causes the PCA space to shrink, which pushes the test samples into the middle of the plot. You can ask them if you don't believe me.

You'll see what I'm talking about when we get high coverage genome-wide data from southeastern Europe.

Mike Thomas said...

David:

"We can also add to that the Muslim and Ottoman incursions into Europe, and it's easy to see why some southeastern Europeans fall into that space between the baMon and Cypriots on West Eurasian PCA."

I don't think that this made much of an impact. In fact, I beleive the truth is the opposite to your theory. During the Neolithic to Bronze Age, SEE and Anatolia were closer than today, and this was reversed during the Middle Ages by Slavic invasions, etc.

Frank said...

David,

"You'll see what I'm talking about when we get high coverage genome-wide data from southeastern Europe. "

Is there an ETA for these?

Davidski said...

During the late Bronze Age there were two parallel clines made up of European and Near Eastern groups, both caused by massive migrations from the steppe, but not overlapping.

Population movements from West Asia to southeastern Europe during and after the late Bronze Age produced the slight overlap between these clines that we can see now on West Eurasian PCA.

Davidski said...

Frank, I have no idea what their ETA is, but I do know that lots of data is on the way, including some juicy Bronze Age stuff from SE Europe.

Mike Thomas said...

Dave
Do you know if any ancient data coming from British Isles any time soon, apart from those still unpublished Hinxton samples.

Simon_W said...

Some additional thoughts regarding the sceptical question raised by Alberto, that is, why is it reasonable to assume that central/western European R1a and R1b originated in the same population, the Corded Ware IMO? And in particular, why did R1b not come from Hungary?

Since R1b-L11 hasn't been found in Yamnaya, some people like Rokus might suggest that it's in reality from local, central/west European HG. I would object that it's at least very curious that R1b(xV88) wasn't found in all EN and MN samples, who surely had their fair share of WHG admixture. And that it turned up for the first time exactly at the time where there was a major influx of eastern autosomal DNA, in a population that's characterized as more than 70% of non-local and eastern origin.

Then, R1b-U106 has been found in Battle Axe derived Sweden, and R1b-P312 in Bell Beakers in Germany. Their common ancestor is R1b-L11. But look where R1b-L11(xU106, P312) is found at present: http://jpst.it/zOZP
Not around Hungary, rather around Corded Ware area.

Then, the argument is that R1b-P312 must have had a different origin, because in some modern populations it's extremely common and R1a at the same time very rare. However, at least R1b-U106 seems to have originated in the Corded Ware, and yet it's very common in some populations who have less than 10% R1a, so why should the same thing be impossible for R1b-P312?

Finally, the modern distribution of R1b frequency, with the highest incidence on the western fringe of Europe, and successively decreasing incidence towards the east, clearly suggests that there was a major expansion of R1b, followed by subsequent migrations from central Europe which supplanted the R1b again. Since Hungary was something like the capital of the Bronze Age world, as Mike Thomas put it, and since it was poor in R1b, expansions starting from there are probably the culprit.

This ties in perfectly with what Alberto just wrote:

>>However, modern North Europeans are clearly shifter North West of CW, in the direction of this population (and also shifter towards WHG while CW was much more EHG). So a similar population to these Hungarians seems to have played a role in producing modern North Europeans.<<

Simon_W said...

Wow, the Bronze Age Montenegro sample is truly amazing! Clusters amidst modern Spaniards. And that was Late Bronze Age, from deep southeastern Europe!

Simon_W said...

I think Cyprus, Greece, Sicily and parts of mainland Italy were already different at that date and will fall inbetween the European and the Near Eastern cline. It's surely Bronze Age related for the most part, it just didn't expand much to the north. Much less than many of us, including me, had expected.

Rokus said...

@Simon_W
'The reason why I spoke if tin Bronze was that RISE98 is partly classified as „Bronze Age Scandinavia“ in Allentoft et al, instead of „Corded Ware“ or „late Neolithic“, and this might make some people believe that the Bronze connection explains the R1b, but it doesn't.'

Please read Allentoft (2015) Supplementary Table 1: RISE98 was dated Battle Axe/Nordic LN, ie. 2275-2032 calBC.
LN=Late Neolithic.

'Trade between neighbouring groups is a widespread phenomenon, and maybe there were even matrimonial connections, occasionally, why not. But you can't build theories that way, you can't explain disagreeable evidence away by making matrimonial connections with neigbours the default assumption.'

This phenomenon was explicitly mentioned by Sarauw (2007) to apply for Sweden in relation with the Bell Beaker horizon in Jutland. The truth of this can now be verified by the autosomal affinity of RISE98 (R1b-U106) with NW Europe, while the R1a sample mentioned above has an eastern affiliation.

' And CW(xR1b-L151) I disagree. There is RISE98 having R1b-L151.'
Autosomally, RISE98 rather approximates the Beaker populations of NW Europe than Corded Ware/Battle Axe. Please don't make things up.

Simon_W said...

@ Rokus

Please read Allentoft (2015) Supplementary Table 9: RISE98 is listed as "baSca". In case you don't understand, this means Bronze Age Scandinavia.

The Swedish R1a sample you're referring to is 400 years older than RISE98, hence closer to the time when R1a and R1b came in from the east. If RISE1 was tested, who also has R1b but who lived around 2720 BC, he'll most certainly plot like the other Corded Ware samples, with an eastern affiliation. In spite of his R1b!

I don't make things up, RISE98 may possibly have had ancestors in northern Jutland and the Netherlands, but this is far from obvious and certain.

I would suggest a PCA to be made with RISE431. He was very late Corded Ware, had R1a and lived around 2170 BC. THEN you'll have a fair comparison. Let's see if he turns out as eastern as the early Corded specimens, or if he isn't rather northwest European-like like his R1b coevals.

And BTW, I forgot to reply to your assertion in your previous post, that R1a hasn't been found in Yamnaya. This is correct, but does it mean that R1a is from central/western Europe? No, because R1a has been found in eastern European hunter-gatherers. It has not been found in WHG, nor in SHG. Nor in any EN or MN farmer. And "coincidentally" the Corded people had strong EHG ancestry.

Simon_W said...

@ David

Is a PCA with either RISE1 or with RISE431 feasible?

Frank said...

David,

I looked that that IBD study you posted. In the pdf provided, the only balkan samples are 2 greek ones, both of which are actualy greek jews, not actual greeks, and taken from Atzmon G. et al.

The map provided seemed very weird to me, given how I remember from dodecad that greeks outside of anatolia didn't share any IBD with near easterners, not even turks.

Davidski said...

Simon,

RISE1

https://drive.google.com/file/d/0B9o3EYTdM8lQWTBuOUxWRXFQc2M/view?usp=sharing

RISE431

https://drive.google.com/file/d/0B9o3EYTdM8lQSWlVbFBNZjJUS28/view?usp=sharing

Frank,

I'm getting the feeling that you'll fall off your chair when you finally see a comparison of Bronze Age Italians and Greeks and their modern counterparts. But this isn't my area of interest, so let's leave it at that for now.

Mike,

I don't know of any ancient genomes from the UK apart from those advertised already.

Krefter said...

@Simon,
"This would surprise no one, so why is it surprising that an R1b guy from 670 BC is modern and northwestern, while an R1a guy from 2670 BC isn't?"

The R1b-U106 guy is from 2000 BC.

Krefter said...

Davidski,

Can you post ADMIXTURE results for the Bronze age Montenegrons?

Grey said...

"Even in Denmark the Bell Beaker presence was far from pervasive, there was most of all a Bell Beaker center in Northern Jutland."

Minor thing but IIRC that flint mining site in northern Jutland goes back to Atlantic Megalith times which to me adds to the idea that BB spread along the existing trade networks (or derived out of Atlantic Megalith for those who think that) so may have been influenced by it at least in some places if not all.

Alberto said...

One thing I find interesting about this debate in which more modern samples move North West (whether R1a or R1b), plus the Montenegro sample, plus BA Hungarians, etc... is how much things have changed in so few time.

It was less than a year ago that I was debating here about the pre-BA European population regarding the Lazaridis et al. 2014 model. There was a general consensus that all Europeans by the late neolithic were like Sardinians, and that's why the model of modern Europeans as Sardinian + EHG proposed in the paper kind of worked.

I opposed such model, because I highly doubted that Neolithic Farmers represented the whole of the European population, but just some unknown percentage of it. So I thought that it was a Caucasus-like population who brought ANE to Europe and that they pulled the population South East, not North East (as the paper implied).

Well, it was not a Caucasus-like population. Just half Caucasus-like (though when adding J2 migrations it's probably closer to it than not). But in any case it's now clear that LN Europeans were (when averaged) very northern and that BA and later migrations pulled them very definitely South East.

Now no one seems surprised about this. We see it normal to find such high levels of WHG ancestry within Europe and no one remembers the "Neolithic Europeans = Sardinians" model. I guess we learn fast, and forget fast.

Roy King said...

@Davidski
The Serbian Vinca Neolithic sample: NG21-10 has approximately 10% Steppe admixture from your analysis. It is circa 4600 BCE. He also has an enhanced WHG in comparision to BAR100, the Late Neolithic NW Anatolian sample. Do you think that the local hunter gatherers in Serbia and by extension SE Europe were a partial source of this 10% Steppe component?
The Montenegro sample from LBA clusters with contemporary Spaniards as you showed us. Could a small, preIE steppe component have occupied the Balkans?

Matt said...

For the outgroup f3 statistics from Allentoft, for modern people, the Bronze Age Montenegro sample goes from highest to least:

Scottish, Pais_Vasco, French_South, Icelandic, Lithuanian, Orcadian, Norwegian, Ukrainian, Croatian, Czech, English

So it's believable on that basis that it would sit in the Western European general area.

Alberto: There was a general consensus that all Europeans by the late neolithic were like Sardinians, and that's why the model of modern Europeans as Sardinian + EHG proposed in the paper kind of worked.

Well, kind of.

The Laziridis model, based on the two clines, was always of a little dubious utility in South Italy and South East Europe, even by Laziridis, as suggested by them being much more off cline for Sardinians / Basques ->Lithuanians than other Europeans (NW, NE, Central, SW).

It's not something that we talked about a lot at the time, because the interest was on how the WHG resurgence and ANE arrival happened, more than the 1/5 of Europe where it didn't work so well. But I'm sure if we went back to the comments around that time. I'm pretty sure David at least would've said something about later migrations effecting SE Europe. The Laziridis paper also may have commented on that topic.

If we never talked about it, then that's interesting, but I'm not sure that's how I remember that.

Alberto: But in any case it's now clear that LN Europeans were (when averaged) very northern and that BA and later migrations pulled them very definitely South East.

Not so sure about this. After all, I think many Eastern European groups share more drift with EHG + WHG than LN groups, for instance.

According to the outgroup f3 stats from Allentoft for instance, even English and French share more absolute drift with hunterW (WHG) and hunter(N) than either Corded Ware or Bell Beaker or Unetice.

And this can't be through Sardinian or Basque like admixture only, because those populations have either comparable or less shared drift with hunterW (Neolithic North European people like TRB are a better idea, but still fall short of being enough).

The stats you posted up for D(Loschbour, EHG; Pop, Chimp) also bear this out.

In terms of pull to South East we may be able to say Italy and Greece probably changed, perhaps with the Roman Empire or perhaps earlier, and some of the changes via Italy may have been mediated into the wider Roman Empire (England, France, Spain, South Germany).

Krefter said...

All Corded Ware from Estonia-Germany-Sweden-Poland will probably come out very Eastern(over 20% ANE). It'll be interesting to see how Allentoft's Unetice and Bronze age Lithuanian turn out, because somehow East Europeans were pulled West. They're far closer to Bell beaker and Unetice than to Corded Ware.

The high amount of R1a-Z283 makes me doubt there was heavy intermarriage with R1b-rich Bell beaker-types or I1/R1b rich Scandinavians. Corded Ware mixing with people who had a ton of WHG and little ENF and ANE is probably what happened.

Grey said...

Alberto

"Now no one seems surprised about this. We see it normal to find such high levels of WHG ancestry within Europe and no one remembers the "Neolithic Europeans = Sardinians" model. I guess we learn fast, and forget fast."

Well that may be partly because people didn't really believe it as although that's what the data showed if HGs had been displaced into the mountains and swamps the data would still be correct but misleading.

Fanty said...

""Scottish, Pais_Vasco, French_South, Icelandic, Lithuanian, Orcadian, Norwegian, Ukrainian, Croatian, Czech, English

So it's believable on that basis that it would sit in the Western European general area."

Deleted my comment of this beeing "northern" not "western" area. Because I missed Pais_Vasco, French_South....

Now I would say it roughly mimmicks the "R" area of Europe. Not the west.

Fanty said...

"The R1b-U106 guy is from 2000 BC."

Yeah. So he lives like 1000 years after the suposed U106 superfather (some claimed possible ages of 5000-7000 years). And 1000 years before the advent of what archeology accepts as "Germanic" culture.

Alberto said...

@Matt

"The Laziridis model, based on the two clines, was always of a little dubious utility in South Italy and South East Europe,"

Yes, that's why I also said "kind of" worked. If you go back to the paper, you'll see that South Europeans needed a 100% pure ANE population to fit in their model, while ANE was decreasing and WHG increasing as you went north. I remember joking about this ANE population arriving by boat to the Mediterranean islands and from there expanding north while losing ANE and picking up WHG.

"Not so sure about this [pulling populations South East]. After all, I think many Eastern European groups share more drift with EHG + WHG than LN groups, for instance.

According to the outgroup f3 stats from Allentoft for instance, even English and French share more absolute drift with hunterW (WHG) and hunter(N) than either Corded Ware or Bell Beaker or Unetice
"

Yes, exactly. What I meant is that the populations from Europe from before the irruption of ANE/R1a/b where more northern (and western, obviously) than what they are after the irruption of ANE.

If we take Yamnaya as the main source of ANE (after all CW is modelled as 73% Yamnaya and 27% Germany_MN, though it might have some extra EHG too), what Yamnaya did was pulling the pre-Yamnaya populations to the South East.

If English are 50% Yamnaya, and Yamnaya was 35% WHG, then the pre-Yamnaya population of England had to about 55% WHG (because modern English are 45% WHG). So the Yamnaya (and derived R1b) migrations pulled them South East.

Or did I misunderstand what you were trying to say?

Rokus said...

'Please read Allentoft (2015) Supplementary Table 9: RISE98 is listed as "baSca". In case you don't understand, this means Bronze Age Scandinavia.'

Hahahah!! Table 9 merely groups some samples together for facilitating D-tests and f3-statistics. The authors must have some good reasons for designing a baSca group that beside the Late Neolithic RISE98 sample also includes RISE61, a Middle Neolithic Danish R1a that is quite a bit older than your RISE94. The latter is grouped as baCW! Bronze Age?? Ask the authors why they did this, it's irrelevant for this discussion. The true archeological grouping can be found in table 1.

Why didn't you ask to run the tests on RISE71 and RISE97 for having your local mixture hypothesis tested? And RISE61 for having an idea how these locals might have looked like?

RISE97: Nordic LN (Sweden) 2025-1885 calBC F. - grouped as baSca
RISE71: Nordic LN (Denmark) 2196-2023 calBC F. - grouped as baSca
RISE61: Nordic MN (Denmark) 2851-2492 calBC M. -grouped as baSca, YDNA R1a1a1

'R1a has been found in eastern European hunter-gatherers.'
These people didn't have a steppe culture!

Matt said...

All I'm commenting on is:

"But in any case it's now clear that LN Europeans were (when averaged) very northern and that BA and later migrations pulled them very definitely South East."

But if

a) Corded Ware and Bell Beaker and Unetice, who are Late Neolithic, share less drift with WHG and SHG than modern English and Germans from the same areas

(e.g. outgroup f3 for hunterW from Allentoft et al:
Lithuanian - 0.2874445, Belorussian - 0.283517, Basque - 0.2831854, English - 0.281927, French - 0.280115, Corded Ware - 0.278850, Bell Beaker - 0.277947, Unetice - 0.277678, Spanish - 0.276669.

outgroup f3 for hunterN:
Lithuanian - 0.284864, Belorussian - 0.281184, English - 0.279784, Basque - 0.278751, French - 0.277673, Bell Beaker - 0.276923, Unetice - 0.276690, Corded Ware - 0.276554, Spanish - 0.273753)

b) WHG and SHG define "north"

then I'm not sure that it's correct to say later migrations have pulled LN Europeans south east, to get to present day populations.

Does that make sense? I was understanding you on the statement that LN Europeans = Corded Ware and Bell Beaker, and that you were stating that later Europeans are just like them but shifted away from WHG / SHG towards the Near East.

If drift with WHG and SHG is "north", and Late Neolithic = Corded Ware, Unetice and Bell Beaker, then post-Bronze and Iron Age Europeans are in a sense by the f3 stat "north" of late Neolithic ones from the same area. Although they might also share more drift with Near Eastern people as well though.

Alberto said...

That RISE61 sample is interesting. He's labelled as Nordic MN B, so it seems to be pre-CW, from ca. 2675 BC. Yet he has a modern R1a subclade. Would be interesting to see how he looks like.

Alberto said...

@Matt

Sorry, I made you confused by using the term LN. I should have said MN, or specify that I referred to the pre-Yamnaya intrusion.

Because the main point was that in Lazaridis et al. 2014 model, Neolithic Europeans were more or less like Sardinians, and then some EHGs from the steppe shifted them North East (this can be easily visualized in the "fateful triangle" from the previous post).

But the reality that we are seeing now is that Neolithic Europeans (when averaged, and not just assuming that only farmers existed), where actually pretty northern (maybe like 55% WHG, 45% ENF), and that the LN/BA migrations shifted them South East (which is what I was saying back then and why I thought it was a Caucasus-like population original from Central Asia who brought ANE to Europe, and not some Steppe HGs).

Angantyr said...

@Alberto

Nordic MN B means either CW derived Single Grave/Battle Axe, or Pitted Ware. So RISE61 is most likely CW.

Mike Thomas said...

@ Prof King

"Montenegro sample from LBA clusters with contemporary Spaniards as you showed us. Could a small, preIE steppe component have occupied the Balkans?"

If not mere artefact, it is theoretically feasible that some EHG form the steppe had already trickled it's way to balkans from before the Yamnaya perios

Davidski said...

Alberto,

There's a very well defined cline on West Eurasian PCA that runs from what is now Sardinia to the Volga. This has always looked like the main trajectory of the admixture event that took place across most of Europe during the Bronze Age, even before we had ancient DNA.

Now that we have ancient DNA, we can confirm that this is indeed what it means. If you look at the PCA below with Esperstedt_MN and a Corded Ware sample from Germany, they fall at the two extreme ends of this cline.

https://drive.google.com/file/d/0B9o3EYTdM8lQb1NROWh4aWsxMkE/view?usp=sharing

https://drive.google.com/file/d/0B9o3EYTdM8lQeTdoeG9BNXd4eEk/view?usp=sharing

What this shows us is that most Europeans during the Late Neolithic/Early Bronze Age were very similar to Esperstedt_MN, and most of the newcomers to Corded Ware.

Btw, RISE61 looks like an early Corded Ware sample, but its dating is iffy because of the so called reservoir effect.

Roy,

I don't think NG21-10 has any steppe ancestry. In the K8 it's straight up WHG/ENF, with inflated WHG.

Other tests with more Ks might show all sorts of other things, but they're probably wrong considering the small amount of markers available.

Alberto said...

@David

"What this shows us is that most Europeans during the Late Neolithic/Early Bronze Age were very similar to Esperstedt_MN, and most of the newcomers to Corded Ware."

Not really. My comment was triggered by Simon_W's probably correct observations about the age of the 2 first samples in this post. One plots around CW, and the other clearly to the North West. And Simon said it's because he had more time to mix with locals (with who else could he mix to change that way?). Now, if we imagine that he is a 50/50 mix of the CW types and the locals, you can imagine how northern the locals were. Certainly nowhere near Esperstedt_MN.

CW itself is modelled as 73% Yamnaya and 27% Esperstedt_MN. Yamnaya was 35% WHG and Esperstedt_MN some 45% (IIRC). So CW already shifted North West by mixing with locals. But it's still 41% WHG, about Spanish level.

But in any case Yamnaya is the new population that came to Europe. We have Estonians or Lithuanians as 50% Yamnaya (according to Haak et al.), and they are some 52% WHG. With Yamnaya being 35% WHG, it means that the other half was about 69% WHG. It won't plot with Esperstedt_MN.

We have BA Hungarians with very high WHG too. They don't plot with Sardinians either. The Vinca sample won't plot with Sardinians if the results are legit. The BA Montenegro plots with Spaniards, and he already got some ANE/Yamnaya/West_Asian admixture (without it, he would move NW of Basques).

And just do math for any North European population. The pre-Yamnaya average was some 55% WHG. Esperstedt_MN and EEFs in general were only a part of the local population. Maybe half of it. The other half plotted very north to make the average about 55% WHG.

So the shift was definitely South East. Even only counting Yamnaya and leaving aside migrations from West Asia. Counting those it becomes much more obvious in southern Europe.

Frank said...

Roy,

in the Live reports from AAPA 2015 topic there was mention of discontinuity in greece between the early and late neolithic, with the mtdna pointing out to a migration from anatolia.

http://eurogenes.blogspot.de/2015/03/live-reports-from-aapa-2015.html

There was also an abstract about a sample from Kumteppe in Anatolia during the LN that had a "component found mainly in modern populations from the Near and Middle East and Northern Africa, and to a much smaller degree, in some Neolithic European farmers". This could be a southwest asian-like admixture. P192-1 from Iron Age Thrace also scored a hefty proportion of southwest asian in K=6.

This could be what is pulling the balkans southeast and could have been in the southern part of the penisula since the late neolithic, reinforced during the late bronze age and then expanded northwards into the montenegrin highlands.

To me, this makes the most sense.

Matt said...

Alberto: And just do math for any North European population. The pre-Yamnaya average was some 55% WHG. Esperstedt_MN and EEFs in general were only a part of the local population. Maybe half of it. The other half plotted very north to make the average about 55% WHG.

But why not just assume a pure(ish) WHG(ish) population was still around, in low numbers, and then continued HG resurgence after CW arrived on the scene, reproducing and contributing to the farmer community at much greater levels than expected?

It's what seems to have happened during the Middle Neolithic, WHG seemingly expanding demographically in farming communities at the expense of Near Eastern farmer ancestry*, so would the process suddenly end with the arrival of CW?

It also helps explain why the Spanish and German MN samples have a pretty consistent WHG profile - that's all the WHG admixture that had happened by then, and more happened in the future.

*as far as we can tell from burials.

Davidski said...

Alberto,

The shift by and large was certainly not southeast. It was east/northeast.

You're confusing local exceptions from the rule, in which the late survival of unusually WHG-rich groups pushed up the level of WHG above the Esperstedt_MN/Corded Ware cline, with the rule.

Mike Thomas said...

Hhm apparently another Y Hg N1a from Polgar, Hungary (acc to Felix).

Unknown said...

Rokus wrote:
"BTW, there is also some Mediterranean Barbed Wire, probably a degeneration of some other kind of Beaker-related culture."

Just FYI. Olivier Lemercier, who just recently lectured the Reich Haravrd group on Beaker has this to say about barbed-wire pottery (it may come from Italy or the Balkans)
"The appearance of barbed-wire pottery...corresponds to a new cycle where the former privileged relationship with the Iberian Peninsula disappears. Although some elements, such as the lithic industry, remain anchored in local traditions, the pottery shows a strong influence from the Italic area, both in vessel shapes and in the style of decoration. If the origin of the barbed-wire decoration style is perhaps to be sought in the north-western Balkans (Lemercier 2004b), the vessel forms are more reminiscent of central and north-Italian Beakers. The assemblages known from southern France at this time belong to a material culture that has a strong Beaker ascendency, both local and Italic."

Krefter said...

A bunch of new mtDNA and Y DNA from Pre-Historic Hungary. R1b as old or older than the oldest German Beaker R1b was found. People have been suspected R1b from Hungary in this time period. It's probably the ancestors of Western European R1b-L11.

http://bellbeakerblogger.blogspot.com/2015/06/2-vucedol-period-are-r1b.html

Mike Thomas said...

It's not that new
Same stuff from her prev paper, more or less

Alberto said...

@Matt

Well, it could be partly also due to what you say. A smaller number of mostly WHG suddenly growing in numbers. But the phenomenon seemed to have happened immediately, so I doubt they were such a small number. If a mass migration from a population that was 35% WHG (Yamnaya) by the time they reached Germany they were up to 48% WHG (Bell Beakers, some 40% Yamnaya) it means that the locals outnumbered the mass migrants, and that they had some 61% WHG on average. We are seeing this phenomenon in Germany, in Sweden, in Estonia, in Hungary and basically everywhere from where we have samples. Why would a mass migration choose to specifically mix with the very small numbers of "Northerners" before expanding and outnumbering the populations with a more southern profile? First if the number of northerners were small it wouldn't add much to the mass migrants. And then it seems a bit fo a stretch in realistic terms.

@David

It couldn't be an exception to the rule. Otherwise North Europeans would be like North Italians or Bulgarians. It was the average.
2
Which means that Neolithic Europeans in the "Fateful Triangle" were a cline that started around Sardinians, from there went north to close to Loschbour and then east to somewhere near Motala (a bit south west from it, probably). When you compare that cline to modern Europeans, the movement is towards the Caucasus (South East). All the cases we're seeing confirm this pattern.

Davidski said...

Alberto, is there any reason you can't see that major cline of European genetic diversity running from Sardinia to the German Corded Ware?

Alberto said...

@David

I'm not blind, I can see that. But I'm also not easily deceived by incomplete plots with missing populations.

This is simple maths or, if that's hard for you, you can just visualize it in your triangle. Choose your preferred North European populations (i.e, Poles). If we assume that they are about 50% Yamnaya, then draw a line from where (on average) Yamnaya plots to where (on average) Poles plot. Then continue that line in the same direction and in equal distance. That will tell you were the pre-Yamnaya locals plotted (on average). Is it close to Sardinia?

Why are those locals not plotted in your triangle so you can see the cline? Because for some reason we're missing those samples. But we know that they must have existed, unless modern Poles got input from people coming from Mars.

Davidski said...

Poles are easily modeled as mostly Corded Ware and Middle Neolithic farmer with maybe some extra HG. That's what both genome-wide and uniparental data show.

I don't know why you're going on about the Caucasus? There's nothing pointing there; no clines, just empty spaces, and no uniparental markers. You might as well be talking about Mars.

Alberto said...

"Poles are easily modeled as mostly Corded Ware and Middle Neolithic farmer with maybe some extra HG. That's what both genome-wide and uniparental data show."

Fine. But when you plot all the MN + HG that's needed to get modern Poles by mixing them with Yamnaya, that population will plot where I explained above. Not near Sardinia. If you want, I'll give you the numbers and you can plot it, maybe that way you can see it more easily.

If we do that with all the populations, and if we also count West Asian migrations, we can get a complete plot of Neolithic Europeans. It will look like I explained above.

Simon_W said...

@ David

Thanks for the PCAs. RISE1 looks rather eastern, as I had expected. But RISE431 looks eastern, too, not what I had expected. Hence the Scandinavians were affected by a different source of admixture.

Simon_W said...

@ Krefter

Yes sure, the R1b-U106 Swede is roughly 2000 BC. But the R1b Dane is much later.

Simon_W said...

@ Rokus

Indeed the groupings in Supplementary Table 9 do not always make sense. And I never claimed otherwise. I merely had said that RISE98 lived at a time before tin bronze became common in Scandinavia, and this just in case some less informed readers might think his R1b must be linked with the people who had brought the bronze. Of course this isn't relevant to our discussion, you've just misinterpreted it as a strawman.

I didn't ask to run tests on RISE71 and RISE97 because I wanted to show that the autosomal change observed by Davidski was rather correlated with sample age than with different yDNA, and as females they don't have yDNA. And RISE61 is just another early R1a. You're wrong if you think he's an example for what the locals have been like. Nordic MN B is Single Grave culture, not TRB. Single Grave = Corded Ware related.

As for the EHG with R1a, yes they were not found on the steppe. But at least they came from the east, and from people who were autosomally very similar to the R1b EHG on the steppe.

Simon_W said...

@ Alberto

I may have been wrong speaking about local influence, I hadn't analyzed the data thoroughly, I was just thinking the late Neolithic Scandinavians were more Gok2 admixed and "Atlantic", hence more Mediterranean than the early Corded people. Certainly I didn't want to suggest that the locals or Northwest European MN farmers were strongly WHG, much more than Sardinians. But the question how and why the Scandinavians changed after the Corded Ware arrival needs further investigation.

Simon_W said...

@ Alberto

The Vinca sample is really an exception so far. Of course there may have been some other areas with elevated WHG admixture, but we cannot say they were the rule. One big nucleus of elevated WHG ancestry was certainly the Baltic, and the main reason for this is that the LBK derived farming cultures didn't spread there. The people of the Baltic stayed hunter-gatherers until the arrival of the Corded Ware. And check the WHG level of the Estonian Battle Axe sample from Allentoft. It's much higher than the WHG level in German Corded people or the Yamnaya. And then there is the apparently rather western Yamnaya in Hungary which caused the Bronze Age Hungarians especially from the Vatya culture to have inflated WHG levels. Actually wouldn't admixture with Vatya-like people largely suffice to explain the extra WHG in most modern Europeans?

Simon_W said...

And the Montenegro sample is late Bronze Age, by that time I would surely expect it to be Indo-European. That it clusters with Spaniards (who are to a large extent descended from IE Celtiberians) and not with, say, modern Greeks or Albanians, suggests that most if not all of the Balkan IEs are steppe derived, and not directly from West Asia. That's actually an important finding.

Alberto said...

@Simon_W

Yes, sorry for kind of attributing to you that affirmation. I Know you didn't mean specifically that local people had more WHG or anything. But what you said is just the obvious reasoning. People came from outside and looked one way. Then after mixing with more locals they looked different.

And it's me who is saying that the way they look different is having more WHG ancestry. And it's me who therefor concludes that the locals had very high WHG ancestry.

The reason why it's not an exception is rather obvious when looking at the numbers. If we leave CW aside (because CW is basically Yamnaya + locals), and we just take Yamnaya as the non-local population:

Yamnaya: 35% WHG
Germany_MN: 45% WHG
Bell Beakers: 48% WHG
Modern Poles: 48% WHG

If you mix 50/50 Yamnaya with Germany_MN you will get a population with 40% WHG (like modern Spaniards). So the locals were not just MN, but there was also WHG. How much? Easy to calculate:

(Yamnaya + Locals)/2 = 48
(35 + X)/2 = 48
X = (48 * 2) - 35 = 61

Locals = 61% WHG (84% Germany_MN + 16% WHG)

When you plot a population that is 61% WHG and 39% ENF, it just won't plot with Sardinians. The most northern populations we have today (Lithuanians, Estonians) are some 50-52% WHG. So a population with 61% WHG is extremely northern.

This was the average population of locals who gave birth to Bell Beakers and modern Poles alike (and by extension all northern Europeans), after mixing with Yamnaya types. Not an exception of one sample.

Mike Thomas said...

@ SImon W

"That (MOntenegro) clusters with Spaniards (who are to a large extent descended from IE Celtiberians) and not with, say, modern Greeks or Albanians, suggests that most if not all of the Balkan IEs are steppe derived, and not directly from West Asia. That's actually an important finding"

Im not sure that follows. Modern greeks and Albanians might be different to groups 4000 years ago.

Simon_W said...

@ Alberto

I think we all agree that there was additional WHG admixture. But I would suggest that most of this was from western Yamnaya in Hungary and from the Baltic. RISE479 from Vatya is 83.8% WHG-UHG in ANE K7. RISE00 from Estonian Battle Axe is 74.5% WHG-UHG in the same calculator.

@ Mike

Yes, I didn't want to imply that they were the same 4000 years ago. But it's factual that modern Greeks and Albanians have substantial West Asian admixture and fall on the bridge between the clines David has spoken about. And if late Bronze Age Montenegrins had become IE because of West Asian admixture, they would be more similar to modern Greeks and Albanians than to modern Spaniards.

Simon_W said...

Regarding the autosomal change in Scandinavians, I've made an Eurogenes K15 comparison between the early R1a Corded Swede RISE94 and the later R1b Swede RISE98. The latter deviates from the former mostly in that he has:
+5% East_Euro
-10% West_Asian
+7% Baltic
-12% Atlantic
+8% West_Med

Compare how the Estonian Battle Axe RISE00 deviates from RISE94:
+2% East_Euro
-10% West_Asian
+10% Baltic

So, in most components the later Swedish Corded Ware with R1b-U106 deviates from the early Swedish Corded Ware with R1a in the same direction as the Estonian Battle Axe does. And wouldn't it make sense that they communicated and mixed with each other across the Baltic? They belonged to the same culture after all. On the other hand the increase of West_Med in RISE98 cannot be explained by admixture with the Baltic Corded Ware. But this is easily explained by admixture with Gökhem-like local farmers.

Alberto said...

@Simon

Yes, those places are likely to have contributed a good amount of WHG ancestry (though in the case of West Ukraine is still very speculative).

But going back to my original point, the change in our views (and model) from less than a year ago is very evident. Haak et al. first proved that there were HG groups around Europe along the Neolithic, because farmers were picking up WHG ancestry with time, and MN were more like Basques than like Sardinians. We don't know exactly when the process finished, but it seems that it was in this LN/BA period when it did, by the irruption of a 3rd population who probably changed the previous status quo. The rise in WHG ancestry is sharp at this time (the extra WHG you mentioned), so it seems it was when all 3 groups mixed into a (mostly) final one.

Haak et al. also proved that Yamnaya people were half EHG and half "Armenian-like". This is another big leap in the narrative.

I know it's obvious that the old model doesn't exist anymore and we all threw it away for a new and improved one, but your comment above made me realise how little surprised we are now about saying, when comparing two samples and one is more North Western, that he probably mixed with locals. If you said that just last December people would have thought you didn't know what you were talking about (because locals equated to Sardinians and the newcomers were the northern guys). Good to look back and see how fast we're moving.

Matt said...

Alberto: But the phenomenon seemed to have happened immediately, so I doubt they were such a small number.

I'm not sure it did happen immediately. As I say, it seems that modern people are share more drift with WHG than LN / EBA people, who work very well as MN plus Yamnaya indicating it happened slowly, over time.

I know this is different from the ANE-WHG-ENF K8 by David, and the IBS-genotype PCA he has run, but looking at a PCA based on the f3 stats from Allentoft et al:

http://i.imgur.com/RfmIuqs.jpg

All the positions are similar to where you'd expect them to, a double cline, *except* that the LN / EBA are less close to SHG / WHG.

Maybe we have enough evidence to say 100% whether the WHG were a lot more resurgent / persistent or more successful vs pre-IE farmers than the German MN, North MN and Spain MN suggest or whether there was more resurgence after the Indo-Europeans. Unsampled populations could always surprise.

Alberto said...

@Matt

I would think that the IBS based PCAs and K8 admixture runs are more accurate in plotting populations that the F3 stats from the Allentoft paper. But either way, whether faster or slower, it doesn't change the basic premise.

LBK_EN was some 30% WHG. Esperstedt_MN was some 45% WHG. So there in Germany there were farmers who were gradually mixing with some other populations along the Neolithic that had very high WHG ancestry. Why don't we have samples of those populations? Probably there are harder to find, and people are just not looking for them in random places when they have Neolithic settlements to uncover.

Then going to the LN/BA, we have Yamnaya types entering Europe. They were about 35% WHG. Bell Beakers or modern Poles are around 48% WHG. So again we need a 3rd population in the mix to match the numbers (Yamnaya + MN won't do it).

The numbers can't lie. The high WHG ancestry in modern North Europeans was within Europe. It was not lost during the Neolithic and then brought back by the R1a/R1b people (these people when they entered Europe were very Eastern, but not significantly northern). WHG ancestry was mostly related to I1/I2, which is to be expected when WHG/SHG had those haplogroups.

These that is rather obvious now, was extremely controversial only 6 months ago. And maybe 6 months from now new finding will again completely change our views. This field moves very fast, which is very exciting. And it's fun to try to guess, but quite risky (I for one have been surprised about many findings, for example R1b in Yamnaya was not what I predicted, or Europeans in Sintashta, etc...)

Mike Thomas said...

Of course they're there Alberto. BB & CWC are (so far) all R1b and R1a, yet we get later Uneticians and urnfielders being I2c, I2a, etc. So they obviuosly didn;t pop out of the ground, but havent been sample yet. But this is not anyones fault. naturally scholars sample 'most diagnostic' of types, esp when going specifically after CWC and BB samples.

Matt said...

Alberto: I would think that the IBS based PCAs and K8 admixture runs are more accurate in plotting populations that the F3 stats from the Allentoft paper.

I'm not so sure either way. On the one hand, the f3 stat should be actually how related a population is to another population. It's not a composite compromise representing that population produced by any algorithm (like ADMIXTURE, which on occasions finds WHG to be 100% of a EuroHG cluster which SHG is also 100% of). On the other hand, I don't know exactly what can affect f3 stats (drift, etc.) There are similar basic structures and also differences in all of these models.

The high WHG ancestry in modern North Europeans was within Europe. It was not lost during the Neolithic and then brought back by the R1a/R1b people

Sure, I agree with the basic point that there was probably more HG ancestry within Europe somewhere at either the dawn of the Bronze Age or later - the questions being whether it was within a lot of people, or a few who became disproportionately successful, and whether it had already been introduced into farming communities somewhere or was still in a reserve of HG. And those are unanswerable as of now.

Either way, yes, contra the simplest version of the thesis that varying ratios of EN plus a single hunter population with a fixed ratio of WHG:ANE, with more sampling the reality is more complex.

Mike Thomas said...

Davidski

How do the Scandinavian I1's look ? Where do they plot ?

batman said...

Scandinavia seems to very close to the bifurication area of I1/I2:

https://genetiker.wordpress.com/2015/08/16/more-y-snp-calls-from-stone-age-sweden/