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Monday, October 20, 2014

PIE homeland update: paleogenomics supports the steppe hypothesis


Several people tweeted from Iosif Lazaridis' talk at the ASHG earlier today, which focused on ancient DNA from 65 Neolithic and Bronze Age Europeans. Here are a couple of the tweets that caught my eye:
There was an influx from north Eurasian steppe into Europe after advent of farming. Consistent w linguistic evidence.Link

Admixture shows multiway admixture among late Neolithic ancient samples. Yamnaya good source as 3rd ancestral reference.Link

So it seems that latest paleogenomics data support the linguists and archeologists who see the Proto-Indo-European (PIE) homeland on the Eastern European steppe. For some background on that, check out the videos here.

Razib also tweeted a few times from the talk, and as far as I can tell, his main point was that the Yamnaya samples showed affinity to the Ancient North Eurasian (ANE) proxy Mal'ta boy, but were also partly of Near Eastern origin, and indeed could be modeled as a 50/50 mixture between present-day Armenians and ancient Karelian hunter-gatherers. He also said that the ancient Karelians were classified as eastern hunter-gatherers (let's call them EHG for now), along with the hunter-gatherers from the Samara Valley, which probably means they carried a lot of ANE admixture.

Moreover, he added that Corded Ware genomes from late Neolithic Germany could be modeled as 75% Yamnaya, while another source from the talk revealed to me that they carried a minimum of 36% EHG.

All of this makes sense, considering that during the Neolithic much of present-day Ukraine west of the Dnieper was home to the Cucuteni-Trypillian farmers, probably of Near Eastern origin, while at the same time large groups of indigenous hunter-gatherers still foraged east of the Dnieper. Based on archeological data, it seems these two groups mixed at some point, becoming mobile pastoralists associated with the Yamnaya culture, and then expanded in all directions during the late Neolithic/early Bronze Age, potentially spreading Indo-European culture and languages as they went.

The Cucuteni-Trypillian farmers might well have been very similar to present-day Armenians, although probably without the 10-15% of ANE carried by them, which likely arrived in eastern Anatolia with the early Indo-Europeans from the steppe.

By the way, it's possible that the Karelian hunter-gatherers are the same samples as those featured in Der Sarkissian et al. 2013., where they were reported to carry mitochondrial (mtDNA) haplogroups C1 (3 instances), U2e (x2), U4 (x2), U5a and H.

Here's a spatial map from that study showing genetic distances between the ancient Karelian mtDNA and that of modern populations.


Der Sarkissian C, Balanovsky O, Brandt G, Khartanovich V, Buzhilova A, et al. (2013) Ancient DNA Reveals Prehistoric Gene-Flow from Siberia in the Complex Human Population History of North East Europe. PLoS Genet 9(2): e1003296. doi:10.1371/journal.pgen.1003296

See also...

Corded Ware Culture linked to the spread of ANE across Europe

Coming soon: genome-wide data from more than forty 3-9K year-old humans from the ancient Russian steppe

157 comments:

Shaikorth said...

MtDNA-wise the aUzPo sample is close to intermediate between Scandinavian PWC (which included the Ajvide samples) and modern Kets/Selkups.

http://www.plosgenetics.org/article/fetchObject.action?uri=info:doi/10.1371/journal.pgen.1003296.g002&representation=PNG_L

Grey said...

some points from the previous thread that fit better here:

@ryukendo

"the usage of the words EEF, when one is really trying to refer to Basal Eurasian, is confusing"

I think this may turn out to be critical.

Logically it seems to me there are two ways "basal" could have entered into the equation.

1) It could have entered Europe with the early farmers
2) It was already in Europe as part of WHG

Is it possible that the large proportion of y dna E1b in parts of north wales and the Balkans is the y dna component of the Basal remnant?

And if for example WHG existed all around the coasts and rivers around the Mediterrean, up the Atlantic and into the Baltic and EEF derived from the WHG in the near east i.e. EEF = WHG 2.0, then that would confuse the proportions with survivors of the original WHG population being mistaken for farmers.

This is a broader expansion of the Brit = Anglo + Welsh vs Brit = Anglo + Continent argument.

Grey said...

@Chad

"I'm sure there is some tripolye ancestry, but probably minimal if they need Armenians to cover the farmer ancestry."

Cucuteni farmers existed adjacent to PIE for a thousands year. It is the obvious source for both genes and loan words.

It's only because it disappeared (possibly *because* it was adjacent to PIE without any protective mountains) that people are casting around looking for alternative explanations e.g. Caucasus or Armenia

I think ancient Cucuteni dna is the obvious place to be looking.

.

Also if Armenians are a back flow after some kind of collapse event then if they are the closest fit it might be because they are the closest fit for Cucuteni i.e. they fit because of where they used to live not because of where they live now.

.

The big difference between celtic and Germanic imo is:
Celtic = R1b
Germanic = R1b + I1
i.e. same source population but two different routes with the part taking the northern route picking up a lot of northern mixture along the way.

.

If the WHG vs EHG (ANE) split is coastal/riverine vs deep interior then it would be mostly but not entirely north vs south or east vs west.

For example WHG along the coast of Lithuania or Black Sea might be further east than some EHG and Finno-Ugric EHG in the far north might be more west than a lot of WHG.

.

@Chad

"Karelians cant be mostly ANE, That would make Yamnaya over 32% ANE. Considering that all of us in Northern Europe are mostly Yamnaya, it puts ANE too high for modern pops."

Or...

conquest leading to a two layered population with male mediated gene flow from the upper layer to the lower layer leading eventually to a mixed autosomal population with y dna mostly from the upper layer.

barakobama said...

Matt,
"If Yamnaya is 43%NE, 35%WHG, 22% ANE, and Karelian is 70% WHG, 30% ANE, as in your figures, and Corded is 75% Yamnaya and 25% Karelian on top, then Corded would be 32.25% NE, 43% WHG, 24% ANE. Which *would* fit a far point on a European cline..."

You're miss interpreting the Tweets. Corded ware was 75% Yamna, which they inherited the 36% Karelian from. Their other 25% was something non-Yamna, supposedly non-eastern.

barakobama said...

"MtDNA-wise the aUzPo sample is close to intermediate between Scandinavian PWC (which included the Ajvide samples) and modern Kets/Selkups.

http://www.plosgenetics.org/article/fetchObject.action?uri=info:doi/10.1371/journal.pgen.1003296.g002&representation=PNG_L"

It's not very relabel testing the relationship between two small sample sets of weak coverage results. Just break down the mutations by hand, that's the best way to do it. Any test that tests their relatedness is going to probably be inaccurate.

barakobama said...

Chad,
"Karelians cant be mostly ANE, That would make Yamnaya over 32% ANE. Considering that all of us in Northern Europe are mostly Yamnaya, it puts ANE too high for modern pops."

When did they ever say this? This is an accurate conclusion considering 50/50 Karelian Armenian, but being majority Yamna is a big statement.

Davidski said...

No one's said that modern northern Europeans were mostly of Yamnaya origin.

What was said was that Corded Ware could be modeled as 75% Yamnaya, and at least 36% Mesolithic Karelian.

We don't know yet how much Corded Ware ancestry modern northern Euros have, and there are other groups to consider too, like obviously Bell Beakers, who also apparently had some ANE, but not as much as the Corded Ware folk.

Davidski said...

I've confirmed that the Karelians are indeed ancient hunter-gatherers from Karelia, and not modern Finns.

By the way, the Corded Ware samples can be modeled as 73% Yamnaya and 36% Karelian, with the Yamnaya the much better fit.

Davidski said...

OK, the CWC tops the list in terms of Yamnaya-like ancestry, at 73%, followed by East Baltic groups, namely Lithuanians and Estonians, and then Czechs and other North/Central/East Euros. No figures available at this stage.

Southern Europeans have much less of this ancestry, and Sardinians are apparently the least Yamnaya-like group on the continent. Makes sense.

Chad Rohlfsen said...

Okay, that makes more sense, David. Czechs are more Beaker, than Corded. Beakers from Moravia and Bavaria, entered Poland and NE Germany. I would like to know the dates of the Beaker sites that they compared. If it's not Karsdorf vs Kromsdorf, I don't think that is a fair comparison. If the Beaker sites that they go by are from a few hundred years after, then they should be less ANE than Corded Ware.
R1b and R1a are likely from the same source, both R, and both sharing may mtDNA lines. German Beaker mtDNA is a 100% match for Yamnaya. I don't think that it's safe to assume that one is much more ANE than the other. The difference being that Corded entered an area that had ANE, whereas Bell Beaker likely didn't at first. So, the difference may be minimal, say 25% vs 20%, or 20% vs 15%. For instance, Britain, is way more Beaker than Corded Ware, yet is almost as much ANE as the Baltic, yet still possibly possessing no ANE, prior to Beaker.

Ryu,
On H, in Beaker sites likely to have R1b

Kromsdorf 0/7
Quedlinburg 5/9 (H5,H4,H13,H1,H)
Benzingerode-Heimburg 1/3 (H1)
Rothenschirmbach 3/4 (H5,H3,H)
Alberstedt 2/2 (H,H3)
Eulau 1/3 (H)
Karsdorf 0/3
Damsbo (Denmark) 0/2

12/33 really driven by 2 sites.

Grey,
There was no Basal Eurasian hiding out in Wales during the Mesolithic. That's crazy talk. None in Loschbour, probably none in LaBrana, and none in Scandinavian hunters.




Chad Rohlfsen said...

David,
Excuse the slight misquote. Kromsdorf is 100% Yamnaya mtDNA.

la señora bibiloni said...

My theory: Cucuteni-Trypolye people spoke PIE, a local dialect stemming from local hunter-gatherers who adopted agriculture (receving some gene flow from Anatolia) and then expanded east (along the Black Sea coasts up to North Caucasus) and west (along the Danube). There was an Indo-European "corridor" north of the Black Sea and eventually its eastern portion developed satemization. When Yamna people started raiding them, IE speakers moved to the south of the Caucasus (Indo-Iranian); up the Danube (Proto Germanic); crossing the Carpathian Basin to the Adriatic coast (Proto Italo-Celtic) and south along the Black Sea (Greek, Phrygian, Hittite and anatolian languages). Corded Ware was the western frontier of the Yamnaya people, non IE, probably Uralic languages.

Grey said...

"There was no Basal Eurasian hiding out in Wales during the Mesolithic. That's crazy talk. None in Loschbour, probably none in LaBrana, and none in Scandinavian hunters."

Well maybe. I just find it interesting that there's 30% E1b tucked away in some remote Welsh mountain valleys. It could simply be the result of some odd migration history or it could be the few places where this pattern exists i.e. in parts of the Balkans, Iberia, Wales etc they are a surviving remnant of something that used to be widespread around the coasts.

There'll be an interesting story behind it either way.

Chad Rohlfsen said...

It's proabably just that someone with a Neolithic lineage outbred the other locals from a few hundred to a couple thousand years ago. It doesn't really change anything autosomal. It is the same false argument used by those who thought that there was greater Mesolithic survival in Scandinavia, because of I1. We now know that it probably came from LBK, and expanded during Funnelbeaker

Simon_W said...

Cucuteni-Triploye was probably unlike Armenians. I think they were similar to the EEF. Archeologically they had roots in LBK and Vinca. And craniometrically they were close to other EEF. The Armenian-like input in Yamnaya must have had more direct roots south of the Caucasus. After all, archeologists have shown that the custom of Kurgan building arose in the Leylatepe culture in Azerbaijan. The Kurgans there are older than those of Yamnaya. Moreover Yamnaya had a lot of mt-haplogroup T1 which wasn't that common in EEF. And an origin of the PIE south of the Caucasus would easily explain the Anatolian branch of IE, it's the one that split of earliest. Maybe at least this one didn't disperse from the Steppe.

Chad Rohlfsen said...

Ryu,
Just for comparison, Corded Ware is 13/41 for H. Not much lower than Bell Beaker 12/33.

Chad Rohlfsen said...

Simon,
I also find it unlikely that C-T played much of a role in the genetics of Yamnaya. C-T was mixed with mtDNA of Dnieper Don and other possible North Pontic hunters. I am not wanting to argue the linguistics part of this. I still think that given R1b's dominance in the Caucasus and West Asia, that it is likely the one that went into C-T, and then towards the Carpathians and Anatolia. Corded Beakers go from Khavalynsk to C-T, to Germany/Central Europe. Bell Beaker mtDNA in Kromsdorf is 100% like that in Russia and Bulgaria.

Grey said...

If the various Balkan cultures to the west of the Black Sea disappeared - for whatever reason or possibly multiple reasons at different times - then some of them may have moved to get away from whatever the problem was - to Armenia for example.

If Armenians have a chunk of ANE maybe they picked it up when they were lived adjacent to some ex mammoth hunters on the other side of the Black Sea and crossed the Black Sea to get away from some problem or other.

(I'm not saying this did happen just saying there are numerous examples from later steppe history of whole tribes moving away and going on a long trek to some new home.)

Grey said...

"Cucuteni-Triploye was probably unlike Armenians. I think they were similar to the EEF. Archeologically they had roots in LBK and Vinca. And craniometrically they were close to other EEF."

Would Armenians fit EEF + ex mammoth hunters?

Chad Rohlfsen said...

Grey,
I am saying that the R1b ancestors of Armenians came from the Balkans, which can explain them having less ANE than their neighbors. Some of it would've been lost during mixing with Balkan farmers.

Chad Rohlfsen said...

Armenians are way too Near Eastern. They have more Basal Eurasian than Stuttgart, and have 14% ANE. They are probably over 60% Basal, maybe a hair over 70%.

Chad Rohlfsen said...

David,
Razib said that they haven't done the y-DNA for Samara yet. It may be a while before the paper is done if they haven't started that process yet.

Chad Rohlfsen said...

http://www.livescience.com/48373-ancient-humans-lactose-tolerance.html

Everyone, check out this article!! Lots of new people are being tested. One is 13kya from the Caucasus.

Chad Rohlfsen said...

Ignore the projections, of course

Chad Rohlfsen said...

http://www.nature.com/ncomms/2014/141021/ncomms6257/fig_tab/ncomms6257_F5.html

several graphs available

Chad Rohlfsen said...

Two of the Neolithic men are C6 y-DNA.

Chad Rohlfsen said...

Our first J2a, is Bronze Age.

Chad Rohlfsen said...

The 13kya remains from the Caucasus, are coming out later. Bummer.

Chad Rohlfsen said...

David,
You should get your hands on these samples. The Baden and Lengyel samples look like they could have a dash of ANE, coming in with J. Still no R1b, prior to Steppe migrations.

Chad Rohlfsen said...

Check that. It's the Makó Culture and the Kyjatice Culture. Post Beaker.

Matt said...

Chad, thanks for the link to the Pinhasi paper.

http://www.nature.com/ncomms/2014/141021/ncomms6257/full/ncomms6257.html

To a certain extent, that pattern of early Southeast Europeans looking like Stuttgart, then looking like Northwest Europeans (seems close to Northern French) makes sense - Northern and West Europeans seem mostly like a mix of Corded Ware and Early Farmers in varying proportion, and that's seems like exactly what you'd find in Southeast Europe at the time after the Corded expansion.

Then you have an expansion that is associated with Iron Age cultures, of which my knowledge is spotty, but Aegean colonization is one (if late), etc which spread elements which are directly influenced by cultures who are probably somewhat like Yamanya, but perhaps more Near Eastern (closer to Armenian).

Although as you say, there is projection on the PC, it is perhaps more blurry than that, so the Bronze Agers may not have been as "Northwest European" as they appear etc .

This also suggests that ancient Danubian farmers are much more Stuttgart like than Armenian, unless they're being compressed towards Sardinia.

http://www.nature.com/ncomms/2014/141021/ncomms6257/fig_tab/ncomms6257_F3.html

The allele spectrum on the pigmentation suggests a shift to almost modern European light skin around the late Neolithic to Copper Age boundary (4490-2700 BC) *before* the Bronze Age likely Corded Ware influenced shift (around 2910 BC), lightest hair in the late Neolithic and Iron Age samples (rather than the more Northern European like Bronze Agers) and sporadic light eyes in the Neolithic. KO1, which seems essentially Mesolithic is unsurprisingly as light eyed as we would expect from La Brana).

It's a limited sample size but suggests modern European pigment has more to do with relatively recent selection (last 3000 years?) or selection within ancient farmers than ancient admixture or the Indo Europeans. Although, on top of that, it does seem that light eyes have some link to hunter gatherers, so at the least, they are maybe why that allele is around.

Seems like the Bronze Agers may have introduced lactase persistence in Southeast Europe as well.

Davidski said...

Chad,

You should ask Felix to get the 22x Bronze Age sample: BR2. I don't have the bandwidth, but that's the one to go for in this dataset.

Here it is...

http://www.ncbi.nlm.nih.gov/sra/SRX484080%5Baccn%5D

Davidski said...

By the way, I'm not assuming that Corded Ware were a lot more ANE and Yamnaya-like than Bell Beakers. This is what I've been told.

Davidski said...

Simon,

It's probably not very practical to model the Yamnaya people as a mixture of a modern people (ie. Armenians) who have non-trivial ANE ancestry, probably from the steppe to the north.

But I guess it's OK for now, until ancient DNA from Anatolia and the Caucasus becomes available.

Also, the problem with seeing the Yamnaya people as a straight 50/50 mixture of Eastern European hunter-gatherers and Caucasians is that it ignores the Neolithic farmers of Ukraine. Why would the Yamnaya population be a mixture of Near Easterners from the Near East, if there were Near Eastern-like people living in Ukraine at the time?

Chad Rohlfsen said...

Matt,
I think that only bell beaker was in Hungary. That would make the NW Europe thing make sense. I don't think corded was that far south.

Chad Rohlfsen said...

I don't think they were too near eastern like in ukraine. These Hungarian samples plot with otzi. They had to find something only found in the Caucasus and near east.

Chad Rohlfsen said...

Okay, I let him know.

Chad Rohlfsen said...

I did ask Razib, about the Bell Beaker, this morning. He said that there was no mention made about R1b or ANE.

spagetiMeatball said...

If Yamnaya can be modelled as 50% armenian than does that show us conclusively how Proto-kartvelian influences might have come to indo-european? We're assuming that the western ukrainian farmers spoke those languages of course, even after leaving the norther middle east a few thousand years earlier.

But that might finally settle all those arguments, and linguists won't have to do any ridiculous triangulations anymore to account for the influence of disparate languages.

Grey said...

It seems odd to me that if those Balkan farming cultures had the earliest script and earliest coppersmithing etc that the direction of travel would all be inwards rather than at least some of it outwards.

Davidski said...

Chad,

The reason those Neolithic samples land near Oetzi is because they have some WHG ancestry above what is found in the Near East, and no ANE.

Classifying Near Easterners based on ANE only works for modern people, not ancient people. So a 50/50 modern Armenian/ancient Karelian mixture is not 50/50 ancient Near Eastern/ancient Karelian, but probably something like 40/60, at best.

Also, Yamnaya-related and Eastern Hunter-Gatherer ancestry estimates were given at the talk for Corded Ware, Bell Beakers and modern Europeans. The Bell Beakers were in the modern range, but lower than Czechs.

sM,

Yes, it looks like the linguists were right, and we're likely to find Caucasus-specific mtDNA among Yamnaya samples, probably from proto-Kartvelians.

But I just don't see why Yamnaya people would be 50/50 proto-Kartvelians from the Caucasus and proto-IE barbarians from the steppe, when we have a third element there in Ukraine already which persisted well into the Yamnaya period.

Tesmos said...

David how much cash do you need for enough bandwidth? I can donate some if you want.

barakobama said...

"Here it is...

http://www.ncbi.nlm.nih.gov/sra/SRX484080%5Baccn%5D"

I can't fucking believe it!! It's probably a sneak preview of what we'll learn happened in Germany from the Neolithic-bronze age. Hopefully Felix can work with a bronze age and Neolithic Hungarian, because the change was probably caused by mostly Yamna-descended Indo Europeans. It seems little has changed in Europe since the bronze age, it was the last major step.

Davidski said...

Tesmos,

I'm in Australia, so I don't think I can actually upgrade to anything that will allow me to download so many gigs. The internet infrastructure here is quite poor.

Chad Rohlfsen said...

I'm not sure about the Armenian thing. I guess we'll see. Who was the source on the bell beakers? Do you know which sites and dates?

Davidski said...

I spoke to a guy who was at the talk and took notes.

The Bell Beakers tested were the same as those in Brandt et al. 2013, from the Mittelelbe-Saale region in Germany. The dates for those are 2500-2050 BC.

I don't have any more details about them, but the impression I get is that they were very similar to modern Central Europeans, while the Corded Ware were clearly different, with higher levels of the Yamnaya and Eastern HG stuff.

Chad Rohlfsen said...

Okay. Maybe these Hungarians will represent bell beakers input. I'd prefer Britain, since they had many more beaker input, but it's a start. Does he know their approximate EEF/WHG ?

barakobama said...

"OK, the CWC tops the list in terms of Yamnaya-like ancestry, at 73%, followed by East Baltic groups, namely Lithuanians and Estonians, and then Czechs and other North/Central/East Euros. No figures available at this stage."

Assuming Laz thinks all of Corded ware and modern European ANE comes from Yamna, this means Yamna at the very very very least had 28% ANE. Because Lithuanians(Estonians have east Asian admixture) who are said to have the most Yamna-ancestry today have 20.17%(from Eurogenes), and Corded ware had more. 20.17/73=27.63. I'm sure they had over 30%.

This also means Karelian hunter gatherers were probably 50% or more ANE. Assuming Yamna can literally be fit as 50/50 Karelian-Armenian, and Armenian's 14% ANE contributes 7% ANE to Yamna.

There's alot of assumptions in there but it seems we now have a basic idea how much ANE Karelian hunter gatherers had and how much Yamna had.

NE Europeans should be 50% Yamna or more(assuming all ANE is Yamna-descended), which is a big surprise to me. Their total Yamna ancestry actually matches pretty nicely with the amount of R1a-Z282 in eastern Europe, so male-only replacement must be false.

They might keep mentioning specifically Czechs after Balts, because Czechs are the only other NE Europeans they have. As far as I know there's nothing special about them their just mostly Polsih-like, with some extra southeastern and western admixture.

Davidski said...

I don't have any figures, apart from the 73% Yamnaya and 36% low bound Karelian hunter-gatherer admixtures in the Corded Ware samples.

But the impression I have is that the WHG/ANE model is now outdated for Europe, and a new one has taken its place, featuring WHG, SHG and EHG meta-populations (SHG = Scandinavian hunter-gatherers and EHG = eastern hunter-gathers).

Also, I'd say that EHG were mostly ANE.

Chad Rohlfsen said...

NE, NC Europeans likely already had ANE from motala and Karelian types. It could still be 25%.

Chad Rohlfsen said...

David,
Any chance you could give that guy my email address for some info on bell beakers? If you would please, it's chadrohlfsen@hotmail.com thanks!

Davidski said...

Apparently Lazaridis touched on the subject of Scandinavian hunter-gatherers being a potential source of ANE in modern Europeans, but their WHG is way too high for that, and that's covered in the last paper anyway.

The eastern hunter-gatherers appear to be the main source, as far as I can gather from all the info.

Chad Rohlfsen said...

It can be a partial source. Obviously if Yamnaya is similar to modern Northern Europeans in EEF, then there has to be a 50-60% survival of the Mesolithic. So Scandinavia could be 40%EEF, 50%WHG, and 10%ANE. If they are then 1/3-1/2 Yamnaya, with much lower WHG, and higher ANE, then it all evens out.

Chad Rohlfsen said...

If the Motala people are part Eastern Hunter, then Eastern hunters will look like a majority source.

barakobama said...

"featuring WHG, SHG and EHG meta-populations (SHG = Scandinavian hunter-gatherers and EHG = eastern hunter-gathers)."

Trying to break down percentages of ancestry from each groups is like trying to break down French vs German vs English ancestry in an American. SHG and EHG are a mixture of WHG and ANE. It just happens the only non-western Mesolithic genomes come from eastern Europe and Sweden(southern). Who knows what was happening in southeast Europe and west Asia.

BTW, I'm not surprised Karelian hunter gatherers were something like 50% ANE and similar to Samara ones, while southern Swedish were only 19%. Because it do be the main ancestors of people as far northeast as Karelia. But it totally makes sense ANE-heavy hgs could spread throughout east-central Europe and into Scandinavia.

Chad Rohlfsen said...

Just to elaborate more...

At 50% replacement, that makes Yamnaya maybe 45%EEF, 31%WHG, 24%ANE

At 33% 48%EEF, 23%WHG, 29%ANE.

That's just if you make them like modern North Central Euros.

Chad Rohlfsen said...

In terms of EEF, that is

ryukendo kendow said...

@ Davidski
Agree that EHG is mostly, if not all ANE.

Overall this is a wonderful development. And congratulations on David's PIE frankenstein. It was almost right on the spot. Hurray for power of internet and such, and also to ppl like Chad, Balaji, Sein, AT and many more. I think it would be quite accurate to say that we as a community here reached something like this conclusion quite some time ago.

Years ago we finally settled the neolithic diffusion/replacement issue. Now we have a fitting conclusion to over a decade of gene blogging and years of ruckus over the PIE homeland. Turns out PIE had inputs from both NE Eur/steppe, and West Asia/ME, and so both theories are right in a way.

Now we are left with a few questions, filling up the gaps basically:
1) the affiliation of the bell beakers.
2) the affiliation of the celts.

What strikes me as interesting and quite mysterious is that Irish, Orcadian, Iron-age britons, and such pull the NE Eur-SW Eur cline outward, resulting in a 'Northwestern bulge', and that this seems to be associated with elevated WHG, with WHG genomes placing NW of modern day variation in PCAs.

There seems to be an independent pole of variation pulling Europeans toward the NW, which might explain higher-than-expected WHG in N Euro in general.

Maybe BB?

It also seems likely that modern eastern Europeans are higher in WHG that corded ware were, so this is a widespread phenomenon. If this came from mesolithic survival, this immediately raises the question of why PIE chose to marry farmers at a vastly lower rate than the HGs, or kill farmers at a higher rate.

It is also almost certain that Slavic/European-autosome domination of the Pontic steppe is late, and raises the question of where the Slavs originated. I am willing to bet that this came from Kurganized HGs/forest Neolithics, which then expanded with ag later, raising the WHG throughout Eastern Europe.

3) Eurasian influence in Africa, and African influence in Middle East

4) Basal Eurasian.






barakobama said...

"Apparently Lazaridis touched on the subject of Scandinavian hunter-gatherers being a potential source of ANE in modern Europeans, but their WHG is way too high for that, and that's covered in the last paper anyway.

The eastern hunter-gatherers appear to be the main source, as far as I can gather from all the info."

There's a simple conclusion everyone should have on this topic, until future evidence says otherwise. All hunter gatherers in Luxembourg and father west were ~100% WHG, the same is probably true for hunter gatherers a little father east and probably in southeast Europe(Maybe there was another form of ancestry).

Karelian and Samara hunter gatherer(or all hunter gatherers in Russia) were mostly ANE, maybe over 60%, put probably not 70%. The rest of their ancestry was probably WHG. Southern Swedish hunter gatherers were a mix of EHG and WHG, and around 19% ANE and 81% WHG. ANE ancestry probably increased in hunter gatherers as you move east from Sweden.

Most modern Europeans ANE comes from Yamna-Indo European's EHG ancestors. But it's possible that hunter gatherers from Scandinavia and east of there also contributed ANE to modern Europeans. West Asians certainly contributed ANE ancestry to some Europeans, especially southeastern and Italian.

Alot of the ANE in west and south-central Asia likely doesn't come directly from WHG or PIE, because they have to little WHG.

barakobama said...

"Agree that EHG is mostly, if not all ANE.

Overall this is a wonderful development. And congratulations on David's PIE frankenstein. It was almost right on the spot. Hurray for power of internet and such, and also to ppl like Chad, Balaji, Sein, AT and many more. I think it would be quite accurate to say that we as a community here reached something like this conclusion quite some time ago."

WHat?! We still don't know what the WHG-ANE ratio of ENG was, it's anything from 50%-100% at this point. So, we don't know how accurate Davidski's Frankenstein is.

Davidski's obviously pushing for EHG to be something around 100% ANE, because I don't think he has any sources saying Bell Beaker was similar to North-central Europeans while Corded ware was differnt because of much more ANE. My guess is EHG was at least over 60% ANE.

We know Corded ware is estimated to be 73% Yamna, so unless they were something very differnt from modern northern Europeans, EHG was probably well under 100% ANE.

barakobama said...

Ryukendo, most of us mostly listened to what more educated people wrote and made common sense conclusions.

Chad Rohlfsen said...

Karelian doesn't work at more than 60%ANE. Not for that kind of input into Lithuanians. That might be high. That would mean that Lithuanians would be in the Single digits for ANE, prior to Yamnaya. 40-50%ANE for the Karelians works best.

ryukendo kendow said...

@ Barak
50%-100% ANE is precisely "mostly or all ANE". You don't disagree, so I have no idea what you are getting so worked up over.

Let's just wait for the results. In the meantime, let's not make assumptions over people's level of education shall we?

Chad Rohlfsen said...

David,

What is the input of Yamnaya, into Lithuanians? Any idea? I can get pretty damn close that way..

Chad Rohlfsen said...

If Lithuanians are 2/3 Yamnaya, then Karelians are in the 40-50% ANE range. If it's half, 60-70%ANE.

barakobama said...

"Turns out PIE had inputs from both NE Eur/steppe, and West Asia/ME, and so both theories are right in a way."

It seems there's now great evidence IE in NE Europe came from Yamna-steppe. The same is true for Indo-Iranian languages, while the rest are unconfirmed and up to debate. Also, the authors of this upcoming paper believe Yamna was PIE, so if anything people who argued for a steppe origin are getting backed up by genetics. Yamna were admixed between near easterns and Mesolithic people of the region, which was expected considering what has been discovered with ancient DNA from further west. Whether PIE and or its ancestor originated in near eastern+natives of Russia or one of the two is up to debate.

ryukendo kendow said...

@ Chad Rolfsen
Its highly unlikely that Lithuanians are 2/3 Yamnaya. Corded Ware are already 75% Yamnaya only, for that figure to become 0.66 in NE Euro, the replacement has to be almost 90% by Corded Ware.

We already know that there is very little WHG in Central Asia+South Asia, and these areas did not get PIE input passed through Corded Ware, so there is no room for an epistemological 'cushion' either. All this implies that the WHG in Yamnaya could not have been very substantial.

@ Chad
There is no doubt about the Steppe origin of PIE culture. I was referring to the autosomal composition of PIE, over which a lot of saliva had been spilled.

barakobama said...

"Karelian doesn't work at more than 60%ANE. Not for that kind of input into Lithuanians. That might be high. That would mean that Lithuanians would be in the Single digits for ANE, prior to Yamnaya. 40-50%ANE for the Karelians works best."

We can't judge whether it works at more than 60%, unless one of us have convincing insight from our own research or the authors. No one said Lithuanians are whatever % Yamna. It could be as low as 40% for all we know. Why'd they say CW was 36% Karelian, if they already said they were 73% Yamna(math says they must have been ~36%)? Maybe Karelian was close to 100% ANE, and that's why.

My point is we don't know and our speculation is almost as valuable as pure guessing. I mostly agree with you, and Balts are probably mostly Yamna.

People have to start asking questions about Yamna and or EHG ANE percentage!!!!!!!!!

barakobama said...

"We already know that there is very little WHG in Central Asia+South Asia, and these areas did not get PIE input passed through Corded Ware, so there is no room for an epistemological 'cushion' either. All this implies that the WHG in Yamnaya could not have been very substantial."

Do you know of data from central Asians? Excluding their Turkic-admixture we should get something close to a Sycthian, right? Pre-IE South Asians may have had more ANE than Yamna. Like Laz said we can't know conclusively the Yamna influence on south Asia without knowing who the pre-IE people were. I agree Yamna probably didn't have a whole lot of WHG, but I'm sure i was higher than Stuttgart and maybe higher than Gok2.

By taking a quick glance at the new paper with ancient Hungarian genomes, it's easy to see there was continuum throughout the Neolithic with Sardinia-like farmers and a sudden change in the bronze age. The bronze age Hungarians obviously have more WHG than Neolithic ones, and overall not a whole lot of ANE. Maybe Yamna did have more WHG than Sardinian-like farmers.

Chad Rohlfsen said...

Go ahead and call me crazy here...

Perhaps someone can come up with a scenario to explain the timeframes and spread of the languages.

I don't think it's possible to make Corded Ware, the only thing that is Indo-European.

I think the only other thing we can do is make later Bell Beaker and any predecessor, IE too. If Bell Beaker is Celto-Italic-Germanic.. those Beakers mix in Northern Poland, Northern Germany, and Denmark, become Germanic speakers.. mixing with Corded Ware extensively.. blurring the line between Slavic and Celtic related, for Germanic. Balto-Slavic is 2800BCE, and fits Corded Ware very well.. and could seriously influence Germanic. Armenian is from 2800BCE... Armenian didn't come from Corded Ware, and pretty certain it wasn't Catacomb.

Several languages are older than Corded Ware. We can't have people speaking Italic, Celtic, Germanic, Armenian, and Balto-Slavic in the same place.

I know that Bell Beaker is different than Corded WAre, but if R1b is on the Southern and Western Steppes, then it should be more like the Armenian part, and R1a, being more forest tribes, would be closer to the Karelians than Armenians.

R1b goes to the Balkans and becomes like a Balkan/North Caucasus mix, and R1a goes to the Baltic, where it becomes a hyper NE European.

I just don't see so many unrelated languages, crossing centum-satem, all coming from the same place.

Speaking a similar language at 6000BCE leaves a lot of time to diverge genetically. Especially with the movements of farming. Anyone caught near the Near East, became more Near East like.

Chad Rohlfsen said...

Anyway, I picture Yamnaya being like Europe. More Near Eastern in the South, more hunter in the North. Southern groups may be 60-70%EEF, Northern Tribes near the forest and above closer to 30-40%EEF.

ryukendo kendow said...

@ Chad
I find that quite plausible actually.

@ Barak
Sardinian and EEF are approx 50% WHG, and any assertion that PIE were higher than that is simply contradicted by the evidence.

You need to accept that for most of prehistory, population flows have been into Europe, and not out of Europe. No need for nativist sentiment here.

Whatever process that raised WHG in Europe later has to be independent from the original PIE invasion, and could be due to later Iron Age IE movements as Chad suggests.

ryukendo kendow said...

So far, neolithic individuals found in Europe have been Sardinian-like, not Bedouin- or Armenian- like. Maju's assertion that all neolithic in Europe derives from a single population in the Med is supported by the fact that everywhere we look we find the same 50/50 WHG/Basal mix in neolithic indiv, or mixes of that with more WHG, including that sardinian-like individual in the Balkans sometime earlier.

The fact that Basal ancestry is found in Yamnaya without correspondingly high levels of WHG more or less precludes a European source of Basal ancestry in Yamnaya IMO.

Chad Rohlfsen said...

Barak, it looks like the neolithic replaced less than 50% in Norhern Europe. When they became pastoral, they had a boom, and raised WHG, further south. IE speakers had to lower the WHG, in order to make room for their higher EEF and ANE. WHG couldn't be more than was already in Northern Europe. That leaves no room for an "Armenian/Karelian" mix. Look at Southern Europeans with more ANE than WHG.

barakobama said...

"Sardinian and EEF are approx 50% WHG, and any assertion that PIE were higher than that is simply contradicted by the evidence.
You need to accept that for most of prehistory, population flows have been into Europe, and not out of Europe. No need for nativist sentiment here.

Whatever process that raised WHG in Europe later has to be independent from the original PIE invasion, and could be due to later Iron Age IE movements as Chad suggests."

I apologize for a little biased in my reasoning.

I’m not crazy for thinking Yamna had a big chunk WHG. Is it random northern Europeans have more ANE and WHG(which correlate together), and are said to have the most Yamna-ancestry. Is it random WHG randomly rose in Hungary along with ANE? I’m not crazy for thinking this all can’t be coincidence. I don’t think Yamna was very WHG, I’m thinking well under 30%(in laz terms), but certainly not 0%. No one has come up with a plausible theory yet how WHG rose in Europe right around when Indo Europeans arrived. It’s only been speculation how it happened without Neolithic farmers and Indo Europeans.
If EHG is 40% WHG, then Yamna should be at least over 10% in Laz terms. ANE K7's ENF and WHG scores are inconsistent with Laz’s WHG-EEF scores, Davidski said the other 6 are just meant to take out non-ANE. Stuttgart/EEF scored 56% ENF, therefore Estonians should be scoring ~18% but they scored well under 5%. EHG had to of been 20% or less for Yamna to have about 0% in Laz terms.

Davidski said...

Modern Armenians are not a pure Neolithic population.

They definitely have ANE, probably WHG, and maybe other admixtures.

Chad Rohlfsen said...

Armenians aren't like Stuttgart, but their 86% near eastern has more basal than Stuttgart.

Davidski said...

The point is that if we take out all of their presumably non-early Neolithic components, then we'd basically end up with Stuttgart's ancestors.

Chad Rohlfsen said...

I see what you're saying. Close to Stuttgart, but looking at WHG in the region, makes me think that CT was at least 20-30% WHG if not way more, considering Yamnaya came in with little WHG. If ANE was the issue, they could've just used MA-1. They needed some more Basal, if you ask me. That would give them 100%ANE. It's not like they didn't have the data.

Chad Rohlfsen said...

Yamnaya at 30%ANE, Corded around 20-25% makes more sense to me. Even at 33% for Northern Europeans, that's 10% ANE. 55-66% of the total, for all of us. Still fits the bill.

Chad Rohlfsen said...

And it doesn't get messy for Lithuanians and other Balts. It leaves room for them to have 10%ANE survival from the Mesolithic, which makes sense.

Davidski said...

That's probably pretty close.

Chad Rohlfsen said...

Bell Beaker being similar to Central Europeans, leads me to think of an origin to the SE of Germany. If it was Iberian, it would be more farmer. If it were West Asian, it would be more Farmer. R1b out of the Western Steppes looks more likely, if they make people like modern Central Europeans. R1a must've been the forest tribes. Even modern distribution looks like that.

Davidski said...

The Bell Beakers being like modern Central Europeans is just me paraphrasing though. I could be wrong.

Chad Rohlfsen said...

Hungarian Bronze samples look relatively close to where they fit now between Gokhem hunters and Otzi.

barakobama said...

Has anyone emailed Reich?

Chad Rohlfsen said...

Felix is going to town on the samples. He is starting with NE-4, it appears.

Nirjhar007 said...

@David
''So it seems that latest paleogenomics data support the linguists and archeologists who see the Proto-Indo-European (PIE) homeland on the Eastern European steppe.''
It is absolutely idiotic to draw any conclusions from the data we have at the moment! we don't have any clear data from Y-DNA and Component wise so having a cue from previous assumptions/Planning is day dreaming.
About Steppe hypothesis it is a complete failure for SC And Central Asia if we dig deep on things! so we need a model which works both for Europe and Central Asia and The Near East a Valid start of that has been made here from every aspect of Studies(Archaeology,Genetics,Linguistics etc) by an academic scholar-
http://new-indology.blogspot.in/2014/10/can-we-finally-identify-real-cradle-of.html
Also from the Data we have WHG component is missing in notable proportions from SC Asian and C asian populations aka Very Very Weak but ENF and ANE type of ancestry have good scores and at last we need aDNA from SC Asia and Central Asia which we don't know what has kept from us buy any means....

Chad Rohlfsen said...

David,
Do you think crossing a Lithuanian or Estonian, with a Lezgin, could give us close to 50%EEF, 20% WHG, 30%ANE? Maybe throw in a few snps from AG-2, to be safe.

Chad Rohlfsen said...

55/15/30 May be safer, if they needed extra Basal.

Chad Rohlfsen said...

I think that these Bronze Age Hungarians are going to be around 40/45/15.

Chad Rohlfsen said...

Felix has the snps from NE-4 available.

https://drive.google.com/folderview?id=0B49499_WVp2fUXlRc1BYaU84bWc&usp=sharing#list

Chad Rohlfsen said...

Check that. I forgot about the hunter, in the mix. They may be 55/30/15.

Chad Rohlfsen said...

David,
BR2 could be related to Urnfield.

ZeGrammarNazi said...

The culture that BR2 belonged to, Kyjatice, is supposedly an amalgamation of the Piliny, Lausitz, and Urnfield cultures.

Grey said...

@barak

"No one has come up with a plausible theory yet how WHG rose in Europe right around when Indo Europeans arrived."

ex mammoth hunters
-> Yamnaya (ANE) steppe raiders

Yamnaya mounted aristocracy (ANE) conquer an adjacent WHG 1.0 or WHG 2.0 population creating a two layer population: upper layer ANE, lower layer WHG

that combined but not yet mixed population then expands as a combined population

that would be one easy explanation

(of which there are many historical examples most of which involve a population with a raider advantage of some kind whether horses or ships)

Grey said...

@Chad

"it looks like the neolithic replaced less than 50% in Northern Europe"

There is no reason to assume that when pop A mixed with pop B it was completely even 50:50 other than as a useful starting point. In fact I'd say it would be the least likely option.

A good current example is south America with a very high percentage of euro descended adna but very high native mdna due to continuous waves of mostly male colonists.

Similarly the difference in outcomes between north and south America due to the difference between mostly single males versus mostly families.

Grey said...

"that would be one easy explanation"

quoting myself, not sure the description was very clear so

Stage 1) population A north of the Black sea and population B west of the Black sea.

(pop A is too weak to conquer B but has a raiding advantage so raids)

Stage 2) pop B is sick of being raided so parts of it break away and go on a volkwanderung in multiple different directions

Stage 3) The displacement of part of pop B makes them weak enough for pop A to conquer outright leading to a *combined* not yet mixed A/B population with A as the aristocracy and B as the peasantry.

Stage 4) At some later date this combined A/B population expands west also as a second wave

conclusion:
two waves, a B wave and a later A/B wave

this would fit the eastern expansion of PIE having less WHG

Simon_W said...

These maps are from a talk by Mallory:

http://justpaste.it/ho39

The first one shows the eastward expansion of CT. It went up to the Dnieper.
The second map shows the extent of the early Yamnaya culture. It extended very far to the east. CT at best influenced its western fringe. Furthermore the Caucasus and the Transcaucasus are close to the middle of the east-west extent of Yamnaya.

Simon_W said...

What concerns Balkan or even Steppe ancestry in Anatolia, this needn't be ascribed to the Anatolian branch of IE, because at a later stage tribes with definite Balkan origin invaded Anatolia: The Phrygians (close relatives of the Greeks) and Thracian tribes like the Bithyni. And according to Herodotus also the Armenians were from the Balkans.

Grey said...

http://www.csmonitor.com/Science/2014/1022/Archaeologists-find-6-000-year-old-temple-in-Ukraine

barakobama said...

I emailed Reich asking how much ANE ancestry is estimated for the Karelian hunter gatherers and the Yamna people. I also, asked how much Yamna-related ancestry Lithuanians or another European ethnic group have.

He said they haven't been able to find a good method to predict ANE ancestry in the Karelian hunter gatherers and the Yamna people. He also said the Karelian and Samara "obviously have lots of ANE ancestry"

I think his response confirms EHG is not 100% ANE. All we know is they had over 50% and under 100%.

I think the reason Laz said Corded ware had about 75% YAmna ancestry, and 36% Karelian, low bound(or something like that), is because they predicted how much Karelian ancestry Corded ware had like with Yamna, and doubled CW's Karelian score to get a Yamna score.

Chad Rohlfsen said...

Ask if he's done tree mix, the same way he figured the ma-1 into native Americans. Maybe he'll release it to David, to play around with.

Chad Rohlfsen said...

Or however they figured the ANE in motala. You'd think it would be the same process.

Davidski said...

It sounds like the ancient Karelians and even Yamnayans had a bit of ENA admixture. Even a small amount, say, more than Estonians, would mean that it's not possible to use the same formulas to estimate ANE as for Motala12.

Chad Rohlfsen said...

Well, that makes sense of yDNA N, in Hungary. Maybe that's how I got that curious East Asian on my Eurogenes....

ANE 14.46%
ASE 1.19%
WHG-UHG 64.33%
East_Eurasian 1.44%
West_African 0.34%
East_African 0.13%
ENF 18.10%

I always get Siberian, for my East Eurasian. Maybe it's not noise.

Chad Rohlfsen said...

David,
I have 6 chromosomes that are between 3.8-6.9% East Eurasian. Do you get similar results?

barakobama said...

"It sounds like the ancient Karelians and even Yamnayans had a bit of ENA admixture. Even a small amount, say, more than Estonians, would mean that it's not possible to use the same formulas to estimate ANE as for Motala12."

I wouldn't be surprised either considering the east Asian mtDNA everyone keeps ignoring in Karelia. Have a tinkle of east Asian ancestry isn't new to me because all ready have it from native American ancestry.

Davidski said...

Yes, I have a few spots of Siberian on my chromosome paintings at GEDmatch, and one spot of less than 0.1% East Asian at 23andMe. It's on the telomere, which is usually a highly conserved area.

jackson_montgomery_devoni said...

@Chad Rohlfsen,

You wrote...''David,
You should get your hands on these samples. The Baden and Lengyel samples look like they could have a dash of ANE, coming in with J. Still no R1b, prior to Steppe migrations.''

You think that the ANE component may have arrived in these ancient samples with Y-DNA haplogroup J? I thought J would have come with early pre-ANE West Asian/Caucasus like people who would have been most EEF or ENF like.

Chad Rohlfsen said...

There's too much ANE in the Caucasus and northern west Asia for it to all be because of ie's or yamnaya. There had to be prior spill over for them to have more than east Europe. The lack of WHG today, makes me think it was pretty Karelian like around the northern Caucasus and maybe it was Armenian like, in the South Caucasus, prior to IE.

barakobama said...

Reich could have simply told me the percentage of Yamna-related ancestry in Lithuanians, which would give a good idea what their ANE was, what EHG ANE is, and how Yamna ancestry modern people have. I don't understand why he didn't. I've asked him a few questions and in every answer he avoids answering the question, and tells me common sense.

jackson_montgomery_devoni said...

The source of J in Europe is most likely Near Eastern like though still I think not ANE really.

Nirjhar007 said...

@Obama
''He said they haven't been able to find a good method to predict ANE ancestry in the Karelian hunter gatherers and the Yamna people. He also said the Karelian and Samara "obviously have lots of ANE ancestry" ''
Seriously? no Y-DNA data?? Obviously the Samara data is so far disappointing...
Happy Diwali To All BTW.....

Chad Rohlfsen said...

David,
We should find El Portalon, and run him through the y-test and aDNA. I have a good feeling about him.

Chad Rohlfsen said...

Find him, for Felix to get it ready, at least.

barakobama said...

Felix finished his Analysis of Hinxton 5. She supposedly died in her 100's and her parents were half-siblings!!!!!!!!!!!!!!!!!!!!!!!!

http://www.fc.id.au/2014/10/hinxton-2-analysis.html

I'm pretty sure all but one of the Hinxton's parents were either 1st or 2nd cousins. They all seem to have died of old age, except maybe the one who died at 55.

She seems to be one of the Anglo Saxons, or at least of mixed ancestry, and very unlike Hinxton-4.

Chad Rohlfsen said...

David,
I have a strange and unrelated request for a couple of f3, z-score comparisons, if you're up for it.

Davidski said...

What kind of f3 comparisons? Why strange?

Chad Rohlfsen said...

I'm not sure if you've tried this before, but it's regarding Oetzi, again.

His culture is supposedly a mix Central, Balkan, and Remedello. It would also help to see if there is any kind of mix coming in, at all.

I was thinking of running him, as a few mixes, and comparing it to numbers from people that are just a little different. To see if the numbers show the same significance.

Something like

Stuttgart Loschbour Oetzi
Stuttgart Storvar11 Oetzi
Stuttgart MA-1 Oetzi

Then, for comparison
French Loschbour English
French Storvar11 English
French MA-1 English

I am wondering if he is 3-5% of both WHG and or ANE. He is always shifted towards Central Europe, in relation to Stuttgart. If the scores are more significant than the English, it might be the case. The difference between the French and English is about equal to what I'm looking for.

Maybe you've already tried this, I'm not sure.

Chad Rohlfsen said...

I say this because, if the Northern Europeans are much more Yamnaya than Southern Europeans, they shouldn't be that close in ANE. Either we've got a completely different kind of group, West of Yamnaya, that first brought in a bit of ANE, or their calculations of Yamnaya input are off, because Tuscans are way too close to the English and Danish in ANE, for it to be a major difference. Especially, if both have some ANE, before Yamnaya ancestry inserts itself.

The Western Steppes my have been a Modern Balkan/Central Euro mixed type. That's kind of what I'm thinking.

Chad Rohlfsen said...

In other words, it makes Yamnaya input look even, across the board. It makes sense more, if Northeast/North Central Europeans are more of the Samara type, and Southern and Western Europeans with a "Kemi Oba", type of ancestor. More Balkan related, I guess. More Dnieper Don, than Karelian. Obviously there wasn't anyone pushing 50% ANE in Ukraine, right next to CT.

Grey said...

"It sounds like the ancient Karelians and even Yamnayans had a bit of ENA admixture."

I think that makes a lot of sense. If you imagine western eurasia as a rectangle made up of an L shaped WHG distribution around the western coasts and two east-west rectangular blocks making up the rest of the rectangle**, an upper ENA layer in the furthest north and an ANE layer in the middle, then the ANE/ENA border would run all along the northern edge of the ANE zone not just the eastern edge.

(**With maybe small regions of surviving neanderthal hybrids tucked away in the remote mountains.)

.

@jackson

"I thought J would have come with early pre-ANE West Asian/Caucasus like people who would have been most EEF or ENF like."

Thing is as the farmers expanded they may have picked up and carried along elements of groups they absorbed along the way like glaciers picking up rocks so J may be associated with EEF but is it EEF itself?

barakobama said...

Chad, Italians and Balkans have a big chunk of post-Neolithic west Asian ancestry. So alot of the ANE in Tuscans, come from west Asia not Yamna. While in southwestern and northern European's ANE most likely comes from Yamna.

Iberians are probably basically the same as NW Europeans, just they have less WHG, more EEF(mostly from Stuttgart-like), and less Yamna(so something like 7% ANE, which is less than half as much). Laz and Reich may have considered west Asian-ANE in Europe, but I doubt it. Their statement still stands though, the ANE difference between N Europe and SE Europe may not be big but there's still a difference. When they say northern European, they may be mostly referring to NE Europeans. There isn't a ginormous difference in WHG-EEF between NW Europeans and Iberians either.

Chad Rohlfsen said...

There's a pretty good difference from Iberians to NW Europeans. Irish 45%, Scottish 42%, English 50%, Spanish 80%, Basque 60%. Basques are only like 7-8% of the population of Spain, they're hardly representative. Southern French are 69% EEF.

As far as getting a lot of ANE from West Asia, I am not so sure of that. Remedello Beaker, was just to the North, there are Beaker sites in Tuscany, plus there were plenty of Celtic tribes around there.

barakobama said...

I emailed Reich again, but this time about WHG-affinity of EHG, Yamna, and Corded ware.

I asked specifically asked if EHG has less WHG-affinity than northern Europeans, if Yamna has less than southern Europeans, and if Corded ware had more than Yamna and how are they compared to modern Europeans.

If he simply says EHG had less or more WHG-affinity than northern Europeans, that gives a hige clue as to how much ANE, how the Yamna were, and how Yamna-related ancestry modern Indo European-speakers have.

Matt said...

Re: good question why the Motala12 model doesn't work for Karelian HGs.

I do wonder if it is because of ENA why they couldn't add an extra admixture edge between ENA and Karelian HGs. Too much complexity to the model? That wouldn't seem inherently the case but they may not have wanted to get into models with that many edges. Or is there another reason ENA mixture affects estimation?

I was wondering if the reason that ANE estimation was difficult would be because Karelians actually might have their ancestry from a group which was (distantly) within clade to ANE relative to WHG. So you have divergence WHG from ancestors of ANE+EHG, then divergence between ANE and EHG. Not sure how that would affect the models.

It should be doable to see if the K HGs share an excess of drift with ENA relative to WHG and ANE, so that can be tested.

Davidski said...

Apparently they can't accurately estimate ANE admixture in South Asians because of their Onge-related ancestry.

So it seems that an extra Eurasian component can complicate matters to the point where nothing that is currently available will work.

For instance, as far as I can tell, if the Karelians are WHG/ANE/ENA, then none of the f4 formulas listed in the last paper will work for them, just like they don't work for present-day Finns, although their ancestry is even more complex because they have EEF.

ryukendo kendow said...

Wouldn't the position of Karelian in a pca with an ANE cluster (Mal'ta+AG2) and a WHG cluster (la brana+loschour) and say, dai give us at least a ballpark estimate? The first eigenvector will be dominated by w. Vs e. Eurasian (WHG+ANE vs ENA) and the second by ANE vs WHG, and if Karelian has no basal, we can see where it falls in the triangle.

Not a formal test, but I don't think we will be in the dark completely. No idea what the authors of the paper would do--a west eurasia pca would be useless--but this seems like such an obvious thing to try that I'll be surprised if they did not do this.

jackson_montgomery_devoni said...

Grey,

That is a good question I do not know really if J is associated with true EEF or early Neolithic farmer ancestry because G seems to be a better fit for that overall.

Chad Rohlfsen said...

What if there's something "Ust-ishim" like, still out there, on the steppes. Maybe that relic is what is getting thrown into south Asians, for Northern Europeans, and makes it impossible to figure the ANE. Throw on some real east Eurasian, and it's kind of a messy tree.

Chad Rohlfsen said...

Of course, I guess East Asian, mixing with ANE, and or Basal, could make something South Asian like.

Matt said...

Davidski, OK, I wasn't sure whether the reason Finns weren't modelled was simply to dispense with testing complex models while demonstrating the simple ones, or if it was a technical limitation. Likewise for South Asia, whether it was a problem of a lack of ancient dna, and avoiding complex models.

Obama, if you're in communication with Reich, you could ask if Karelian hunter gatherers are as distant as MA-1 and Loschbour from East Asians.

That would should tell us in a single statistic if they were ENA admixed, assuming current population models.

ryukendo kendow said...

@ Barak
A good question to ask too would be where they fall between ANE and WHG, and how much closer they are to one than the other.

barakobama said...

Reich's response:

"I’m sorry, this is too complicated to explain by email! We are working on this, and will hopefully have a paper on it ready to submit some time in the next few months."

He doesn't want to answer any questions it seems. First he avoided it and now he's basically saying he won't even if he knows the answer, or because he and his team aren't 100% sure what the answer is yet.

Maybe we'll get the paper by Christmas. I'd suggest preparing for it.

Davidski said...

His reply is understandable, considering that he probably doesn't want to say anything now that will come back to haunt his career, and/or wants to reveal all the really powerful info in a formal publication in a major journal.

We'll just have to wait.

Davidski said...

Report from Laz's talk...

"By examining admixture levels in these groups, they found that an early European farmer split off from the rest of the European farmers early on and mixed with eastern European hunter-gatherers to form the Yamnaya population, which lived on the Steppes, Lazaridis said. Meanwhile, a middle Neolithic population mixed with Yamnaya to form the late Neolithic Corded Ware population of central Europe."

http://cdnwww.genomeweb.com/sequencing/ashg-panelists-discuss-new-insights-european-population-structure

That sounds like the Cucuteni-Trypillian farmers.

Grey said...

@Davidski

"That sounds like the Cucuteni-Trypillian farmers."

Yeah, I think there's some obvious human psychology in this. Because Cucuteni disappeared people naturally look for clues elsewhere but the Cucuteni disappearing is the biggest clue of all. It's the dog that isn't barking.

.

@Jackson

"because G seems to be a better fit for that overall"

That was my thinking too. I have a similar thought about the I1 in northern Euros i.e. it was a group that amalgamated with the expanding R1 and got carried along with them like rocks in a glacier.

Chad Rohlfsen said...

Ct is too late David. It probably affected the sw tribes, way later, but not samara. Samara was Neolithic before ct ever existed.

Davidski said...

Cochran on Corded Ware again...

http://westhunt.wordpress.com/2014/10/26/yamna-and-corded-ware/

barakobama said...

BTW, it seems they estimated CW was 73% Yamna by designing a test on how much Karelian hg they were(36%) and then doubled the score. Now it makes sense that Reich said they haven't been able to estimate ANE in EHG, so they are currently treating it as a population to estimate Yamna-ancestry.

Davidski said...

Cochran on the early Indo-Europeans.

http://westhunt.wordpress.com/2014/10/27/centum-and-satem-2/

barakobama said...

What's the big deal about Cochran?

Davidski said...

Nothing. Might be interesting to see what people are thinking elsewhere though.

Mike Thomas said...

Cochrans spins even more BS than the usual Kurganist day-dreamer ! Wat a crock if shit. "Lactose tolerance helped them expand" .
Good one . Philosopher

Davidski said...

Ironically, Cochran looks like he fits better in the Blatterhole cave than on the Russian steppe.

But anyway, here's some news about a 6,000-year-old Trypillian temple in Ukraine.

http://www.archaeology.org/news/2650-141023-ukraine-trypillian-temple

Davidski said...

The Cucuteni-Tripolye migration into Central Ukraine began 4150 B.C.E.

http://www.chronikajournal.com/resources/Harper.pdf

That's basically the same period when typically Neolithic mtDNA haplogroups appear near the Samara.

"Samara experienced major population turnovers over time: early samples (>6000 years) belong primarily to mtDNA haplogroups U4 and U5, typical of European hunter-gatherers but later ones include haplogroups W, H, T, I, K, J."

http://eurogenes.blogspot.com.au/2014/06/coming-soon-genome-wide-data-from-more.html

Grey said...

"Lactose tolerance helped them expand"

It might have helped them expand dramatically in the northern forest zone where crops weren't very effective.

so instead of direct invasion maybe it's

1) migration to under exploited niche
2) population explosion due to LP
3) conflict with LBK to the south.

Simon_W said...

The quote by Ciara Curtin does sound as if the Armenian-like ancestry in Yamnaya was from CT. Because where else should EEF ancestry have come from? Surely not from Western Asia. But if it was basically EEF ancestry that mixed with EHG, why isn't it simply labelled EEF in the ASHG presentation? So maybe she got something wrong. Moreover it seems quite unlikely that CT, which was neither particularly early nor particularly southeastern geographically, was really Armenian-like. I would suppose that they had similar levels of WHG as the other EEF. Then there's the problem that the Dnieper is far from Samara. And the elevated incidence of mt-haplogroup T1 and the presence of haplogroup I, both atypical for the central European EEF.

Archeologically, Yamnaya is linked with the Maikop culture, which was earlier:
http://dienekes.blogspot.com/2013/06/the-maikop-singularity.html
And Maikop in turn is linked with the Kura valley and Lake Urmia area south of the Caucasus, close to modern-day Armenia:
http://dienekes.blogspot.com/2013/05/origins-of-maykop-phenomenon.html
This area is close to the location of the PIE homeland according to Gamkrelidse and Ivanov. It's important to note that from a purely linguistic point of view there is no unequivocal consensus placing the PIE homeland on the Pontic-Caspian Steppe. To the contrary, linguists have put forward many different theories regarding the homeland, and in particular, afaik it's nearly impossible to decide between the northern Pontic area and central Europe, using purely linguistic arguments. The reason why the Steppe theory has got more credit among linguists is because they were influenced by Kurganist archeologists. And vice versa. I'm not a linguist, so I don't have the expertise to evaluate the linguistic arguments by Gamkrelidse and Ivanov. That means I can't simply dismiss their view either.

Interestingly, according to the second link above, the Kura valley and Lake Urmia area received important input from the Iranian plateau. This might explain how R1b spread from there via the Northeastern Caucasus to southern Yamnaya.

It's also noteworthy how distinct the Anatolian branch of IE is from the other IE languages, see the net here:
http://dienekes.blogspot.com/2011/11/splits-or-waves-trees-or-webs.html

If it was the language that stayed behind and got, as a consequence, split off from the relatives by the massive Caucasus chain, then that might explain the big difference. In contrast, if it was just the forefront of a very early expansion wave, I'd expect more something like a continuum to others.

Simon_W said...

And by the way, Brandt, Haak et al. 2013 show also a special relationship of the Unetice mt-DNA with that above mentioned hotspot in Azerbaijan and surrounds, in the genetic distance map. The Corded Ware map is similar, but a bit less specific, including a larger area.

Davidski said...

We would've heard something by now if there was a late significant contribution from south Caucasian populations to the steppe groups.

The data they have includes full genome sequences and full mtDNA sequences, so it'd be pretty easy to spot a south Caucasian link.

Also, the theories that ANE-relatd lineages, like Y-HG R, moved up from the Near East to the steppe come from a time before we knew ANE existed.

And Anatolian really isn't native to Anatolia, despite the name.

Simon_W said...

But they mentioned Armenians as proxies for part of the ancestry of the Yamnaya population. Armenians are not exactly a Caucasus population, they needn't be one, but they are from northern West Asia, and I guess their European admixture is fairly weak.

Regarding the ANE related lineages, I'm not sure what to think of them atm. I can't believe that the strong ANE we see in the entire Caucasus and in Turks, Armenians and Iranians is all from the Pontic-Caspian steppe. If MA1-related ancestry spread westwards along the Eurasian steppe, why not westwards through Iran as well? Moreover the South Asian component is also related with MA1 and they've got R2 there.

Hittite and related languages were not native to the places where they were spoken in historical times, that's right. But where exactly they had come from isn't known for sure, it's all speculation. An origin to the east definitely cannot be ruled out. (And the places I mentioned in my above comment are not in Anatolia anyway.)

I should also add that the theory of Gamkrelidse and Ivanov mustn't be conflated with Colin Renfrew's Anatolian farmer theory. The exact timing and location of the homeland differs, and Renfrew's not a linguist - his theory was never accepted in the linguistic community because it was completely at odds with the linguistic evidence.

Simon_W said...

Well I was thinking of the PCA projections in Laz et al. But I have to admit that if we consider the shared drift stats, then Iranians etc look considerably less MA1-related than most Europeans.

Still, I think some of the ANE admixture entered Western Asia earlier than the people from the Eurasian steppe, and I think it's very likely that this was associated with the spread of R1b1b (found in Anatolia) and R1b1c (found in the Levant and Africa). Related with the Eurasian steppe populations is probably just R1b1a and subclades.

Simon_W said...

Regarding the early beginning of the Neolithic in Samara that Chad mentioned, that refers to the Elshanka culture. Which is usually considered Neolithic by the local archeologists. But by Neolithic they primarily seem to mean pottery making. So it's not really the full Neolithic package with agriculture. It's comparable to Ertebölle in Northern Europe, hunter-gatherers with pottery. And according to some archeologists the introduction of pottery making needn't involve a lot of immigration. Indeed, the Neolithic pottery making in Samara started at least 9000 BP or earlier. And as the abstract of the Yamnaya paper tells us: Prior to 6000 BP the mtDNA was mostly U4 and U5, hunter-gatherer haplogroups. So the change after 6000 BP wasn't associated with the Neolithisation. Rather with Kurganism I suspect.

Simon_W said...

Some comparisons regarding the MDLP World-22 West_Asian component:

NE7 has 0%
CO1 has 0%
Gok2 has 0%

These are what Europeans were like before eastern hunter-gatherer descended groups brought in ANE, together with R1a and R1b.

But ANE admixture didn't necessarily entail West_Asian admixture, as the example of the Estonians shows: Estonians have more ANE than any other European population (18.7%), yet they have only 0.7% West_Asian. But the same can also be observed on the Basques, even though their ANE is modest, 9.6%. Yet their West_Asian admixture is much lower than this and amounts to only 0.2%. Hence the West_Asian component isn't merely a function of ANE admixture.

That was clear anyway because the expansion of y-haplogroup J2 was real, and J2 correlates well with West_Asian admixture.

Now, present-day Basques are 0.2% West_Asian, Estonians are 0.7% West_Asian, and Sardinians 0.8%. Indo-European populations (other than Sardinians) all have more: Latvians have at least 1.1%, Lithuanians have at least 1.2%, Swedes have at least 1.2%, Norwegians have 2.8%, Brits have 4.8% and Portuguese have 5%.

Even though the pre-ANE Europeans had 0% and ANE itself didn't raise West_Asian admixture.
I'd say it's speaking in favour of a West Asian origin of the PIEs.

Simon_W said...

Perhaps the West_Asian component looks like a function of ANE admixture at first glance because many of the groups who brought in ANE had West_Asian admixture, but on closer examination it turned out they are not necessarily linked.

I'm open again to the Celtic from the West theory, though Italo-Celtic from the West might perhaps be more sensible.

Arch Hades said...

"OK, the CWC tops the list in terms of Yamnaya-like ancestry, at 73%, followed by East Baltic groups, namely Lithuanians and Estonians, and then Czechs and other North/Central/East Euros. No figures available at this stage.

Southern Europeans have much less of this ancestry, and Sardinians are apparently the least Yamnaya-like group on the continent. Makes sense"


So this study will offer a statistical breakdown of modern Eurasian/European populations with Yamna like ancestry? If so I can't wait.