The genomic history of southeastern Europe-Mathiesen et al

[Angela]

So BR1 with little or no steppe, and BR2 which can be modeled with Anatolia Ch or Armenia Ch instead of steppe? Does anyone have d based stats for these samples, or Admixture based on actual ancient samples? We'd also need some with Anatolia Ch etc in place of steppe and the fits for all of them.

To be clear here, we're talking about possible CHG from the Balkans reaching up to Bronze Age Hungary, perhaps with a big new infusion of WHG. This is not Bell Beaker.

 

[Angela]

@ LeBrok, Bicicleur,

We were talking about the composition of Yamnaya:


"Iosif Lazaridis (Broad) said...
It's great to see the data already being analyzed and I hope it will be useful in your analyses!

I just wanted to leave a brief comment that the model of Steppe_EMBA as a mixture of EHG+CHG is rejected (Table S7.11), while that of EHG+Iran_ChL is not. Note that in Table S7.11 we are modeling Steppe_EMBA and the references with respect to 13 outgroup populations (the set O9ALNW), not all of which are included in the TreeMix graph.

It is possible for some models to succeed with a particular set of outgroups (both EHG+CHG and EHG+Iran_ChL are feasible with only the O9 set of outgroups; Table S7.10), but for some of them to be rejected when additional outgroups are introduced (Table S7.11). As we mention further down, that doesn't mean there is no CHG-related ancestry in Steppe_EMBA as we can model it as a 3-way mixture involving CHG as one of the sources. What it does mean, however, is that CHG+EHG cannot be the only sources, as this model is rejected (Table S7.11). A further test of our overall model is that when we withhold Iran_ChL as a source, and infer mixture proportions by intersecting the EHG->Steppe_EMBA and Levant_N+Levant_BA clines (p. 134), we get fairly reasonable agreement (mixture proportions).

We try to be cautious in our interpretation of the admixture models, because of three factors: (i) we don't know the geographical extent of populations like "CHG" or "Iran_ChL" so admixture from Iran_ChL does not imply admixture from geographical Iran or CHG from the geographical Caucasus, (ii) we do not have samples from many places and it's very likely that slightly different mixtures than the sampled populations existed elsewhere, (iii) it is possible that the actual history of admixture may be more complex than the simplest parsimonious models identified by the analysis.

Overall, our admixture analysis rejects several possible models (such as EHG+CHG) and thus puts constraints on what may have happened, and also proposes some models that are more resilient to rejection (such as EHG+Iran_ChL+CHG). But, by no means should these be regarded as the final word or unique solutions, but rather as one possible way that the data can be modeled."

I don't think they've found the specific population (s), or they hadn't at that time.

In the subject paper Mathiesen uses CHG + Iran Neolithic as the formulation for the group that moved into Anatolia and then into the Balkans in the Chalcolithic and then even more so in the Bronze Age. Now that may be the population only for this migration westward. The Reich Lab formulation for the group that mixed with the EHG to create Yamnaya may still be where Lazaridis left it, may be closer to this Mathiesen formulation, or may have changed based on samples they've discovered but not released. I just don't know.

Ed. Just X out the last paragraph. Mathiesen is saying the combination that went into the Balkans is the same as what went onto the steppe.

From the paper:

"In eastern Europe we document the appearance of CHG/Iranian Neolithic ancestry north of the Black Sea, and its eventual extension as far north as the Baltic. In some ways, this expansion parallels the expansion of Anatolian farmer ancestry into western Europe although it is less dramatic, and several thousand years later. These expansions set up the two, largely separate, populations in western and eastern Europe that would come together in the Final Neolithic and Early Bronze Age to form the ancestry of present-day Europe."

I would think Lazaridis would agree, but we'll have to wait for his new paper. Maybe they've found a new and slightly different Iranian farmer population.

 

[LeBrok]

@ LeBrok, Bicicleur,

We were talking about the composition of Yamnaya:


"Iosif Lazaridis (Broad) said...
It's great to see the data already being analyzed and I hope it will be useful in your analyses!

I just wanted to leave a brief comment that the model of Steppe_EMBA as a mixture of EHG+CHG is rejected (Table S7.11), while that of EHG+Iran_ChL is not. Note that in Table S7.11 we are modeling Steppe_EMBA and the references with respect to 13 outgroup populations (the set O9ALNW), not all of which are included in the TreeMix graph.

It is possible for some models to succeed with a particular set of outgroups (both EHG+CHG and EHG+Iran_ChL are feasible with only the O9 set of outgroups; Table S7.10), but for some of them to be rejected when additional outgroups are introduced (Table S7.11). As we mention further down, that doesn't mean there is no CHG-related ancestry in Steppe_EMBA as we can model it as a 3-way mixture involving CHG as one of the sources. What it does mean, however, is that CHG+EHG cannot be the only sources, as this model is rejected (Table S7.11). A further test of our overall model is that when we withhold Iran_ChL as a source, and infer mixture proportions by intersecting the EHG->Steppe_EMBA and Levant_N+Levant_BA clines (p. 134), we get fairly reasonable agreement (mixture proportions).

We try to be cautious in our interpretation of the admixture models, because of three factors: (i) we don't know the geographical extent of populations like "CHG" or "Iran_ChL" so admixture from Iran_ChL does not imply admixture from geographical Iran or CHG from the geographical Caucasus, (ii) we do not have samples from many places and it's very likely that slightly different mixtures than the sampled populations existed elsewhere, (iii) it is possible that the actual history of admixture may be more complex than the simplest parsimonious models identified by the analysis.

Overall, our admixture analysis rejects several possible models (such as EHG+CHG) and thus puts constraints on what may have happened, and also proposes some models that are more resilient to rejection (such as EHG+Iran_ChL+CHG). But, by no means should these be regarded as the final word or unique solutions, but rather as one possible way that the data can be modeled."

I don't think they've found the specific population (s), or they hadn't at that time.

In the subject paper Mathiesen uses CHG + Iran Neolithic as the formulation for the group that moved into Anatolia and then into the Balkans in the Chalcolithic and then even more so in the Bronze Age. Now that may be the population only for this migration westward. The Reich Lab formulation for the group that mixed with the EHG to create Yamnaya may still be where Lazaridis left it, may be closer to this Mathiesen formulation, or may have changed based on samples they've discovered but not released. I just don't know.

We've learned a lot but we have a long way to go.

Thanks Angela for posting it! I'm happy that not only I rejected EHG-CHG model for Yamnaya. I can live with Iran Chl, as a source, though I like Iran Neolithic better for my Yamnaya model. I wonder if they've tried modeled it with IN and was rejected?
It is a nail to the coffin of cultural spread of farming, as many guys (farmer haters) dream of.
Confusion is understandable because CHG and Iran Farmer are closely related, though one would think this should be recognized by researchers more easily than for us. More confusion will continue when calculating Steppe influence in BA Balkans, due to inflow of CHG/Iranian Farmer genes through Anatolia, which will be confused with part of BA Steppe admixture also containing Iranian Farmer.

 

[Angela]

Thanks Angela for posting it! I'm happy that not only I rejected EHG-CHG model for Yamnaya. I can live with Iran Chl, as a source, though I like Iran Neolithic better for my Yamnaya model. I wonder if they've tried modeled it with IN and was rejected?
It is a nail to the coffin of cultural spread of farming, as many guys (farmer haters) dream of.
Confusion is understandable because CHG and Iran Farmer are closely related, though one would think this should be recognized by researchers more easily than for us. More confusion will continue when calculating Steppe influence in BA Balkans, due to inflow of CHG/Iranian Farmer genes through Anatolia, which will be confused with part of BA Steppe admixture also containing Iranian Farmer.

I was just in the process of editing the post above. You may be right about it being Iranian farmer for the Reich Lab. Mathiesen is saying the combination that went into the Balkans is the same as what went onto the steppe.

"In eastern Europe we document the appearance of CHG/Iranian Neolithic ancestry north of the Black Sea, and its eventual extension as far north as the Baltic. In some ways, this expansion parallels the expansion of Anatolian farmer ancestry into western Europe although it is less dramatic, and several thousand years later. These expansions set up the two, largely separate, populations in western and eastern Europe that would come together in the Final Neolithic and Early Bronze Age to form the ancestry of present-day Europe."

I would think Lazaridis would agree, but we'll have to wait for his new paper. Maybe they've found a new and slightly different Iranian farmer population.

 

[Promenade]

Mesolithic SHG from Stora Forvar, StF11, was I1 as well.

Do you have a source for this? Looking at the Skoglund study it seems they only tested his mtdna.

 

[LeBrok]

I was just in the process of editing the post above. You may be right about it being Iranian farmer for the Reich Lab. Mathiesen is saying the combination that went into the Balkans is the same as what went onto the steppe.

"In eastern Europe we document the appearance of CHG/Iranian Neolithic ancestry north of the Black Sea, and its eventual extension as far north as the Baltic. In some ways, this expansion parallels the expansion of Anatolian farmer ancestry into western Europe although it is less dramatic, and several thousand years later. These expansions set up the two, largely separate, populations in western and eastern Europe that would come together in the Final Neolithic and Early Bronze Age to form the ancestry of present-day Europe."

I would think Lazaridis would agree, but we'll have to wait for his new paper. Maybe they've found a new and slightly different Iranian farmer population.

The combination might be the same. I think the devil will be in figuring out the proportions and ratios of admixtures. Instead of clear pictures, the plot thickens.

 

[holderlin]

Yes you are right. Now I also see that Vovnigi and Derevka are the most likely candidates for PIE.
These people from the Vovnigi and the Dereivka probably ancestors of all the existing Indo-European peoples.

CW and BB (geneticly and partly culturaly in BB case) probably come from there. And even the historical Hittites, which some researchers derive from the Sredniy Stog. And languages we are speaking now, began to disintegrate precisely there and in those times. The rest of the vast territory of possible PIE languages ​​(as the eastern yamnaya), did not have offspring of historical IE speakers. Therefore, we can not say in what language they spoke, although their cultural characteristics were identical to IE.

The source of these burials Vovnigi and Derevka is somewhere on the Popovo-Zvejnieki mesolite line.
Apparently, there began process of absorbing EEF genes, which are presented in CW.
Probably, they took women as trophies during the devastating raids on the neighboring Tripolye and the killing of their population.

Therefore, the people of Corded Ware:
-the presence of EEF genes
-pronounced horizontal profiling of the face in the upper part (more pronounced than in their Mesolithic ancestors)
-prevalence farmer-Middle Eastern mtDNa in all descendants of CW in Europe.
- burial form as "tower of silence" among the Iranians, has its roots in the Tripolye and possibly introduced by the Trypollian women in Indo-European culture (although, it could be just borrowing)

Also worth noting, from here probably there is an old confusion among paleoanthroologists, who indicates that people of CWC are often metrically similar to the Mediterranean.

Looks like it to me, as far as we can tell at this point.

Interesting connections. I had only read about the "Mediterranean" types in Sredny Stog and CWC, which of course supported the Balkan interface etc.

I do think that Yamnaya is eventually Indo-Iranian, at least around Samara and further East. Indic speakers (ANI) are descended Yamnaya (or afanesevo), but some Iranian speakers are descended from Yamnaya, and others are descended from MLBA_steppe. So we know that Yamnaya, at least in the East was eventually speaking Aryan, which diverged into various Iranian dialects that may or may not have taken on a LBA_steppe structure from Western sources.

But as you say, it does look like the vast majority of IE languages can be traced back to this initial Steppe/Balkans interface.

So this was a dynamic process. Something like this:

We have some form of PIE by the Eneolithic on the steppe (Iron Gates, Baltic?) during which time we see an influx of CHG. Anatolian is formed shortly after this time and begins to separate in the West, maybe in Ukraine, maybe in the Balkans proper, who knows, but early on in the West. We see continual influx of steppe admixture from the East while it receives an influx of CHG, which eventually shows its influence especially in the metallurgical styles. In the West Italo-Celtic separates. The developing bronze age steppe begins to envelop the Baltic and continually pushes against the Balkans diluting or pushing out WHG/Iron gates, which yields Yamnaya Ukraine. Germanic separates in the West. The BA steppe begins to speak Indo-Iranian which expands East while the West begins to speak Balto-slavic. Indo-Iranians in the West who are adjacent to the proto-corded ware/Yamnaya Ukraine Balto-slavic speakers are continually mixing with proto-corded ware/Yamnaya Ukraine while maintaining their linguistic identity. This is ongoing as Corded Ware develops in the West and Poltovka develops in the East. Eventually you have Iranian speaking LBA_steppe that looks an awful lot like corded ware. During this time Greek separates in the West.

Something like that.

 

[Angela]

I was just in the process of editing the post above. You may be right about it being Iranian farmer for the Reich Lab. Mathiesen is saying the combination that went into the Balkans is the same as what went onto the steppe.

"In eastern Europe we document the appearance of CHG/Iranian Neolithic ancestry north of the Black Sea, and its eventual extension as far north as the Baltic. In some ways, this expansion parallels the expansion of Anatolian farmer ancestry into western Europe although it is less dramatic, and several thousand years later. These expansions set up the two, largely separate, populations in western and eastern Europe that would come together in the Final Neolithic and Early Bronze Age to form the ancestry of present-day Europe."

I would think Lazaridis would agree, but we'll have to wait for his new paper. Maybe they've found a new and slightly different Iranian farmer population.

Eurogenes has always strongly objected to the CHG/Iran Chalcolithic formulation for Yamnaya. He came up with a EHG/CHG/some kind of Near Eastern farmer? combination that supposedly worked. He's also not happy with this CHG/IRANIAN farmer formulation. One of his objections apparently is that IR farmer has Basal Eurasian and that...

"Mesolithic and Neolithic Eastern Europeans rich in R1a and R1b don't carry any Iran_N ancestry, because they have 0% Basal Eurasian admixture."

I must be missing something, because if they have CHG and we know CHG has lots of Basal, how can they not have Basal?

 

[holderlin]

Eurogenes has always strongly objected to the CHG/Iran Chalcolithic formulation for Yamnaya. He came up with a EHG/CHG/some kind of Near Eastern farmer? combination that supposedly worked. He's also not happy with this CHG/IRANIAN farmer formulation. One of his objections apparently is that IR farmer has Basal Eurasian and that...

"Mesolithic and Neolithic Eastern Europeans rich in R1a and R1b don't carry any Iran_N ancestry, because they have 0% Basal Eurasian admixture."

I must be missing something, because if they have CHG and we know CHG has lots of Basal, how can they not have Basal?

I think some of this confusion is the fact that EHG itself seems to have admixture from something (Baloch) that's ancestral to Iranian Neolithic presumably from MA-1 or AG. It confuses calculators because Iranian Neolithic is like 1/2 Baloch and 1/2 Kotias (CHG). It could be that the Baloch in EHG doesn't share basal eurasian with CHG? I dunno.

 

[Sile]

The highest EHG+WHG percentage belonging to the Early Balkan Neolithic is a T1a1a individual from Malak Preslavets with 35%
The lowest EHG+WHG from Malak Preslavets is a G2a2b2a individual with 0%

This show extreme differences among individuals from a same settlement in a same time period.

I read that Malak Preslavets is on the danube river , the period when it was inhabited was from 6100BC - 4800BC . they found many many bird skeletons as the area was a bird migrational nesting place.

The Thracian took it over in the bronze and iron age and renamed with ....unsure what and it was a boat making place

The Roman built a fort on or near this town called Candidiana

From most of what I read on the net, it seems a place where steppe people met anatolian people over many centuries............maybe it was a trading place ........my opinion.

Interesting that the T1a1 youth ( young teenager is what I was told ) had the 35%

 

[bicicleur 2]

Thanks Angela for posting it! I'm happy that not only I rejected EHG-CHG model for Yamnaya. I can live with Iran Chl, as a source, though I like Iran Neolithic better for my Yamnaya model. I wonder if they've tried modeled it with IN and was rejected?
It is a nail to the coffin of cultural spread of farming, as many guys (farmer haters) dream of.
Confusion is understandable because CHG and Iran Farmer are closely related, though one would think this should be recognized by researchers more easily than for us. More confusion will continue when calculating Steppe influence in BA Balkans, due to inflow of CHG/Iranian Farmer genes through Anatolia, which will be confused with part of BA Steppe admixture also containing Iranian Farmer.

Do I understand correctly that Mathieson distinguishes Iran Neo from Iran Ch in his admix analyses in this paper?
To simplify, Iran Neo + CHG went west into Anatolia & the Balkans, while Iran Ch + CHG crossed the Caucasus into the steppe.
I would think, if it is Iran Ch + CHG the expansion should have initiated later than 7.5 ka, the onset of the copper age. IMO Khvalynsk and Dnjepr Donets witness these first movements into the steppe.
Pure CHG to expand into the steppe or already populated areas elsewhere would seem unlikely and I've always understood that this was a simplification of the reality.
It would be very interesting if we had Maykop DNA to compare with this Iran Ch.
And I wonder about the Kumptepe samples : do they match Iran Neo + CHG?

It is very good that Laziridis reminds us once again about the restrictions of admixture models, which always simplifies the reality which may be hidden beyond the limits of the model.

 

[Apsurdistan]

Do you have a source for this? Looking at the Skoglund study it seems they only tested his mtdna.

Hah, the elusive I1. Again it seems like this haplogroup so far unless they start to find some ancient dna... is either very young, or really old and was isolated from pretty much everybody for a long, long time.

 

[Alpenjager]

I read that Malak Preslavets is on the danube river , the period when it was inhabited was from 6100BC - 4800BC . they found many many bird skeletons as the area was a bird migrational nesting place.

The Thracian took it over in the bronze and iron age and renamed with ....unsure what and it was a boat making place

The Roman built a fort on or near this town called Candidiana

From most of what I read on the net, it seems a place where steppe people met anatolian people over many centuries............maybe it was a trading place ........my opinion.

Interesting that the T1a1 youth ( young teenager is what I was told ) had the 35%

I think he was younger than teenager, according to his photo. Sadly, the upper half of the adult individual disappeared.

 

[MarkoZ]

I must be missing something, because if they have CHG and we know CHG has lots of Basal, how can they not have Basal?

I'm not entirely sure how accurate this is given that the 'Basal Eurasian' component seems to be a bit speculative still (at least when it comes to the exact details), but didn't Lazaridis point out that the Swedish Motala cluster already had significantly more BE than the WHGs?

At least that seems to be the case here (~10% in the Motala cluster):

I'd think that the more eastern-shifted hunters carried even more of the BE component. In the Samara HG it's 17-18% BE already.

 

[LeBrok]

I think some of this confusion is the fact that EHG itself seems to have admixture from something (Baloch) that's ancestral to Iranian Neolithic presumably from MA-1 or AG. It confuses calculators because Iranian Neolithic is like 1/2 Baloch and 1/2 Kotias (CHG). It could be that the Baloch in EHG doesn't share basal eurasian with CHG? I dunno.

Yes, EHG and Iranian Neolithic/CHG have ancient connection, (one common ancestor), through Baloch/Gedrosia, which could have happen right after LGM. They never met since, till Neolithic. Caucasian admixture, so high in CHG and IN, showed up rather late in the area. It either came from farther south, or was developed in CHG during long isolation and deep bottlenecking, perhaps secluded in one of Caucasian valleys. That's why it didn't have time or a chance to penetrate into Steppe in any decent numbers, till Neolithic of course. Making it very obvious that it came with Iranian Farmers/Herders rather than CHG hunter gatherers.
If it comes to basal eurasian, I'm lost with this ghost.

 

[berun]

To look at the possibility to assign to CHG the origin of PIE I have tried to get some essential info from the messy data about the orange admixture (steppe component) found in the Balkan samples:

- 3 women labeled as Peloponnese_Neolithic buried in a cave display some 5-15% steppe. Date provided is 5000-3200. Some daggers were found, and that is just against the Neolithic concept... a true Neolithic woman buried in other cave of the Peloponnese (I2318) dates of the Final Neolithic (4000 BC) and has 0% steppe.
- I2215 has a 10% steppe, is the unique individual buried in a Cris Culture cemetery with steppe component (1 of 9). Date 5600 BC.
-I4918 has a 5% steppe, this woman was found in a Chalcolithic burial, but in the exel file this sample is labeled as Starcevo_EN and dated to 6000-5300. Another nonsense.
- I2181 has a 25% steppe, was found in a chalcolithic burial and it's dated to 5600-3000... a long timespam, but in the excel it's dated to 4500 and his Y-DNA is R.
- Varna samples have different percentages of steppe admixture, a sample Y-DNA is R, the necropolis is dated to 4600-4300.
- the Trypillie samples found in Verteba cave are assigned to the Cucuteni-Tripolie culture, but such culture had unkown burial techniques so they must be outliers. They show some 5% steppe component, but I1927 had 30%, it is considered an outlier dated to 3600-2900, his Y-DNA was G2a.
- The Bronze Age samples (3200-1200 BC) display the steppe component in diverse percentages, except I2510; the case is that even if such sample is labeled as Bronze Age, in the supp info it is labeled as Chalcolithic and dated by 2800 BC, with Y-DNA G2a, and being his tomb disturbed by an Early Bronze Age pit...

 

[berun]

My bet: herders from actual Kurdistan spread towards the Balkans around 4500 after learning advanced metalurgy from their Mesopotamian neighbours.

 

[Alpenjager]

The sample I2532 (Cotatcu) reported as "Young Female" is wrongly linked to G2a2b2b in the Supplementary Table 1.

 

[berun]

With so much losely treated data, it's difficult to get a picture from that, by that it's allways better to check by oneself, that could change what the paper states about intermitent steppe occurrence in the Balkans

 

[halfalp]

Whats the difference between CHG / Iran_Neolithic and CHG / Iran_Chl ? If neolithic iranians went to steppe, so maybe CHG / iran_Chl is more like Steppe_EHG / Iran Neolithic with a reflux in iranian in chalcolithic / bronze age.