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Ancient Fennoscandian genomes reveal origin and spread of Siberian ancestry in Europe

So, it seems the Saami were probably there first, speaking an unknown language, adopted a Finno-Ugric language from another group, and to some extent mixed with people coming north from the direction of Estonia also speaking a Finno Ugric language? Those people and I1 people from Scandinavia pushed the poor Saami far north?

Interestingly, years ago when I looked up what we know of the history of the Saami, before genetic analysis, I read that the Saami were a European population who had spoken a different language previously. Some words and placenames originated in the previous language. Most probably a language isolate. A migration from the east had imposed a different language on the Saami. Then scientists had come along and named the language and the language group for the people where they first found it. So the Saami language and language group is not actually indigenous to the Saami people. Or so the story went back then, before all this new information. The migration that replaced the language was said to have been about 500 AD, which is much later than this new estimate. Although fairly recent research indicates humanity went extinct, or very near extinct in northern Scandinavia in the 540s so it would be good timing. There would have been more than one migration I expect.

Anyway, the Saami being pushed further north is fairly well attested local history. Its not been a case of Saami territories and Non-Saami ones, though. More like timeshare. We're at the edge of agricultural viability and in a warmer climate the agriculturalists expanded. Cooler periods meant the reindeer herders strategy paid off better and they expanded, while agricultural terriotries contracted. Thats not been the pattern for the last few centuries though because the tech edge of the agriculturalists have grown larger.
 
The case can be seen looking at Baltic_CCC showing 20% Yamna. It's a nonsense from an steppist point of view.

I've been saying since the first Haak paper came out that I thought the models over-estimated the amount of actual steppe "Yamnaya" genetic material initially moving into the far northeast, and ancient dna seems to be proving that correct. There have been a lot of changes in the genomes of this area since then, with movement from more "Yamnaya" heavy and also EEF heavy areas. There are also the "Siberian" like migrations, which contain not only ANE like, but also East Asian like material, or perhaps we should say that the "Siberian" component is on a cline between West Eurasian and East Eurasian?

I just think that the initial models were a bit simplistic and that with more ancient dna they have to be refined.

Also, ADMIXTURE doesn't work that well for those parts of Europe with more than just the three major ancient population groups. The more complicated the population history, the easier it is to confound it.


Likewise, the Basques are an issue in terms of the standard steppe theory. They carry downstream R1b, but it doesn't look to me like what was added in the Bronze Age was the standard Yamnaya "mix". There's too little CHG or, in these charts, Iran Neo in them. One possible explanation is that this could be because various streams of migration had various admixtures. It's a mistake, imo, to imagine that this was a simplistic movement of one homogeneous group. These weren't "national" entities in the way we think of them now, and even those "national" entities aren't homogeneous.
 
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I've been saying since the first Haak paper came out that I thought the models over-estimated the amount of actual steppe "Yamnaya" genetic material initially moving into the far northeast, and ancient dna seems to be proving that correct. There have been a lot of changes in the genomes of this area since then, with movement from more "Yamnaya" heavy and also EEF heavy areas. There are also the "Siberian" like migrations, which contain not only ANE like, but also East Asian like material, or perhaps we should say that the "Siberian" component is on a cline between West Eurasian and East Eurasian?

I just think that the initial models were a bit simplistic and that with more ancient dna they have to be refined.

Also, ADMIXTURE doesn't work that well for those parts of Europe with more than just the three major ancient population groups. The more complicated the population history, the easier it is to confound it.


Likewise, the Basques are an issue in terms of the standard steppe theory. They carry downstream R1b, but it doesn't look to me like what was added in the Bronze Age was the standard Yamnaya "mix". There's too little CHG or, in these charts, Iran Neo in them. One possible explanation is that this could be because various streams of migration had various admixtures. It's a mistake, imo, to imagine that this was a simplistic movement of one homogeneous group. These weren't "national" entities in the way we think of them now, and even those "national" entities aren't homogeneous.

Just to clarify, perhaps some of the changes we see between the two sets of ADMIXTURE results is simply that when there is no Nganasan sample, the algorithm "dumps" those alleles into the closest sample set. This often happens in ADMIXTURE from what I can see. It's clear in the beautifully done Supplement to Haak, done, I think, by Lazaridis, when, as new reference sets are added, the fits get better, and the percentages of the major components change.
 
ok, I see, but then the problem is with the Yamna share (EHG + CHG) of British and Frenchs, which is not covering so all the EHG share so that it is lumped by the Siberian component (?). If so it clearly points that IE was not coming from Yamna but from another place with extra EHG, from Kievan Rus area I guess.

Not necessarily. Even in the absence of EHG in that model the extra "eastern" HG ancestry that gets assigned to the Siberian component is very minor. Since nobody claims that steppe ancestry and IE arrived in Northwestern Europe directly with the Yamna people, but with people admixed with Yamnaya-like (not necessarily the average Yamnaya used as a proxy), the Indo-Europeanized cultures with heavy Yamna-like ancestry, but also a lot of the substrate ancestry (which must have included some extra EHG-like ancestry in Scandinavia and Northeast Europe), may have brought subsequent waves of IE languages, some of which came with some tiny extra EHG and even some downright Siberian ancestry, especially via the massive and attested Germanic migrations of tribes originating in Scandinavia and the North Sea coast. The extra EHG is so low that I do not think it changes much in the traditional perspectives of immigration and IE origins.

One should also remember that most of the steppe EBA samples that were studied came from the eastern portion of the Yamnaya horizon, and CWC for instançe may have come from a more western or northern layer of "Yamnayized" culture that was rich in R1a instead of R1b. It is possible in my opinion that there was genetic structure within the Pontic-Caspian region, Yamnaya was a cultural and probably linguistic phenomenon, not a totally homogeneous population, at least after its expansion from the Balkans to the Volga. I think it is likeky that some tribes, especiallY in the forest-steppe zone, were richer in EHG, so they would come off in admixture analyses as "Yamnaya + extra EHG ".
 
Thanks, Jovialis. I was getting extremely bored with this discipline. :)

I'll read it carefully later. Just looking at the admixture graph, I was immediately struck by the changes in the Northeast over time. Also, it clearly tells the same story as all analyses of the genomes of modern Europeans do. In western and central and certainly in southern Europe, although they don't show the latter, if you consider that perhaps forty percent of Yamnaya is "southern" the WHG and EHG portion is very much a minority one.

When I look at the distributions of WHG and EHG I'm drown in doubts. Saami with no EHG, only WHG, compared to Bolshoi Oleni Ostrov, and with some serious input of Yamnaya in Saami, supposed to be come from East with Finnic languages, at least for a part.
There is no more any paper using the same basis as the others for admixture, it's so puzzling... I 've lostmy latin and my celtic!
 
Not necessarily. Even in the absence of EHG in that model the extra "eastern" HG ancestry that gets assigned to the Siberian component is very minor. Since nobody claims that steppe ancestry and IE arrived in Northwestern Europe directly with the Yamna people, but with people admixed with Yamnaya-like (not necessarily the average Yamnaya used as a proxy), the Indo-Europeanized cultures with heavy Yamna-like ancestry, but also a lot of the substrate ancestry (which must have included some extra EHG-like ancestry in Scandinavia and Northeast Europe), may have brought subsequent waves of IE languages, some of which came with some tiny extra EHG and even some downright Siberian ancestry, especially via the massive and attested Germanic migrations of tribes originating in Scandinavia and the North Sea coast. The extra EHG is so low that I do not think it changes much in the traditional perspectives of immigration and IE origins.

One should also remember that most of the steppe EBA samples that were studied came from the eastern portion of the Yamnaya horizon, and CWC for instançe may have come from a more western or northern layer of "Yamnayized" culture that was rich in R1a instead of R1b. It is possible in my opinion that there was genetic structure within the Pontic-Caspian region, Yamnaya was a cultural and probably linguistic phenomenon, not a totally homogeneous population, at least after its expansion from the Balkans to the Volga. I think it is likeky that some tribes, especiallY in the forest-steppe zone, were richer in EHG, so they would come off in admixture analyses as "Yamnaya + extra EHG ".

I'm not in total agreement here, but I've always held that the R1a Corded Ware group was a "similar" population autosomally to Yamnaya which adopted the Yamnaya "culture". I'm not sure about the language. Unfortunately, there are no genes one can track for that. :)
 
When I look at the distributions of WHG and EHG I'm drown in doubts. Saami with no EHG, only WHG, compared to Bolshoi Oleni Ostrov, and with some serious input of Yamnaya in Saami, supposed to be come from East with Finnic languages, at least for a part.
There is no more any paper using the same basis as the others for admixture, it's so puzzling... I 've lostmy latin and my celtic!

I don't see any major contradictions at all. If you go back to the Supplement in Haak which I referenced above, this is not that different. The "Siberian" addition is explored there, as are the Saami if I remember correctly. After all, the Reich Lab paper and this one are both to some extent Haak papers.
 
So, it seems the Saami were probably there first, speaking an unknown language, adopted a Finno-Ugric language from another group, and to some extent mixed with people coming north from the direction of Estonia also speaking a Finno Ugric language? Those people and I1 people from Scandinavia pushed the poor Saami far north?

"The Finno-Ugric branch of the Uralic language family, to which both Saami and Finnish languages belong, has diverged from other Uralic languages no earlier than 4000–5000 years ago, when Finland was already inhabited by speakers of a language today unknown. Linguistic evidence shows that Saami languages were spoken in Finland prior to the arrival of the early Finnish language and have dominated the whole of the Finnish region before 1000 CE16,17,18. Particularly, southern Ostrobothnia, where Levänluhta is located, has been suggested through place names to harbour a southern Saami dialect until the late first millennium19, when early Finnish took over as the dominant language20. Historical sources note Lapps living in the parishes of central Finland still in the 1500s21. It is, however, unclear whether all of them spoke Saami, or if some of them were Finns who had changed their subsistence strategy from agriculture to hunting and fishing. There are also documents of intermarriage, although many of the indigenous people retreated to the north (see ref. 22 and references therein). Ancestors of present-day Finnish speakers possibly migrated from northern Estonia, to which Finns still remain linguistically close, and displaced but also admixed with the local population of Finland, the likely ancestors of today’s Saami speakers23.?

"
Our results suggest that a new genetic component with strong Siberian affinity first arrived in Europe at least 3500 years ago, as observed in our oldest analysed individuals from northern Russia. These results describe the gene pool of modern north-eastern Europeans in general, and of the speakers of Uralic languages in particular, as the result of multiple admixture events between Eastern and Western sources since the first appearance of this ancestry component. "

So, could we infer that the Siberian/East Asian component came with the Finno-Ugric speakers?

"
Ten of the eleven ancient individuals from this study fall on this Uralic cline, with the exception of one individual from Levänluhta (ID JK2065, here named Levänluhta_B), who instead is projected closer to modern Lithuanian, Norwegian and Icelandic populations. Specifically, two Levänluhta individuals and the two historical Saami from Russia are projected very close to the two previously published modern Saami (Saami.DG)32 and the new Saami shotgun genome generated in this study (as well as the previously published genome of the same individual, here labelled Saami (WGA)1), suggesting genetic continuity in the north from the Iron Age to modern-day Saami populations. In contrast, the six ancient individuals from Bolshoy are projected much further towards East Asian populations, and fall to an intermediate position along the Uralic cline and close to modern-day Mansi."

"
Interestingly, results from uniparentally-inherited markers (mtDNA and Y chromosome) as well as certain phenotypic SNPs also show Siberian signals in Bolshoy: mtDNA haplogroups Z1, C4 and D4, common in modern Siberia24,34,35 are represented by the individuals BOO002, BOO004 and BOO006, respectively (confirming previous findings24), whereas the Y-chromosomal haplotype N1c1a1a (N-L392) is represented by the individuals BOO002 and BOO004. Haplogroup N1c, to which this haplotype belongs, is the major Y-chromosomal lineage in modern north-east Europe and European Russia. It is especially prevalent in Uralic speakers, comprising for example as much as 54% of eastern Finnish male lineages today36. Notably, this is the earliest known occurrence of Y-haplogroup N1c in Fennoscandia. Additionally, within the Bolshoy population, we observe the derived allele of rs3827760 in the EDAR gene, which is found in near-fixation in East Asian and Native American populations today, but is extremely rare elsewhere37, and has been linked to phenotypes related to tooth shape38 and hair morphology39 (Supplementary Data 2). Scandinavian hunter-gatherers from Motala in Sweden have also been found to carry haplotypes associated with this allele4. Finally, in the Bolshoy individuals we also see high frequencies of haplotypes associated with diets rich in poly-unsaturated fatty acids, in the FADS genes4,40,41."

"Our results show that all of the test populations are indeed admixed, with the most negative values arising when Nganasan are used as the Siberian source (Supplementary Data 3). Among the European sources, Lithuanians gave the most negative results for Estonians, Russians and Mordovians. For modern Hungarians, the European source giving most negative results was French, while both Bolshoy and Nganasan gave equally negative results when used as the Siberian source. With Finns as test, modern Icelanders were the European source giving most negative statistics. Finally, Icelanders and Nganasan used as the European and Siberian sources, respectively, yielded the most negative result for the present-day Saami as a Test. This result is still non-significantly negative, either due to the low number of modern Saami individuals in our dataset (n = 3), or due to post-admixture drift in modern Saami. A high degree of population-specific drift can affect f3-statistics and result in less negative and even positive values42"



Not the first time this correlation of Hungarians and the French has shown up, to the incomprehension of some of our posters.

"
In contrast to previous models for European populations using three streams of ancestry2,3, we found that some populations modelled here require two additional components: a component related to modern Nganasans, as discussed above, and additional EHG ancestry, not explained by Yamnaya (who have been shown to contain large amounts of EHG ancestry themselves3). "

Not most, however.

"
While the Siberian genetic component presented here has been previously described in modern-day populations from the region1,3,9,10, we gain further insights into its temporal depth. Our data suggest that this fourth genetic component found in modern-day north-eastern Europeans arrived in the area before 3500 yBP. It was introduced in the population ancestral to Bolshoy Oleni Ostrov individuals 4000 years ago at latest, as illustrated by ALDER dating using the ancient genome-wide data from the Bolshoy samples. The upper bound for the introduction of this component is harder to estimate. The component is absent in the Karelian hunter-gatherers (EHG)3 dated to 8300–7200 yBP as well as Mesolithic and Neolithic populations from the Baltics from 8300 yBP and 7100–5000 yBP respectively8. While this suggests an upper bound of 5,000 yBP for the arrival of this Siberian ancestry, we cannot exclude the possibility of its presence even earlier, yet restricted to more northern regions, as suggested by its absence in populations in the Baltics during the Bronze Age. Furthermore, our study presents the earliest occurrence of the Y-chromosomal haplogroup N1c in Fennoscandia. N1c is common among modern Uralic speakers, and has also been detected in Hungarian individuals dating to the 10th century44, yet it is absent in all published Mesolithic genomes from Karelia and the Baltics3,8,45,49.The large Nganasan-related component in the Bolshoy individuals from the Kola Peninsula provides the earliest direct genetic evidence for an eastern migration into this region. Such contact is well documented in archaeology, with the introduction of asbestos-mixed Lovozero ceramics during the second millennium BC50, and the spread of even-based arrowheads in Lapland from 1900 BCE51,52. Additionally, the nearest counterparts of Vardøy ceramics, appearing in the area around 1,600-1,300 BCE, can be found on the Taymyr peninsula, much further to the East51,52. Finally, the Imiyakhtakhskaya culture from Yakutia spread to the Kola Peninsula during the same period24,53. Contacts between Siberia and Europe are also recognised in linguistics. The fact that the Nganasan-related genetic component is consistently shared among Uralic-speaking populations, with the exceptions of absence in Hungarians and presence in the non-Uralic speaking Russians, makes it tempting to equate this genetic component with the spread of Uralic languages in the area. However, such a model may be overly simplistic. First, the presence of the Siberian component on the Kola Peninsula at ca. 3500 yBP predates most linguistic estimates of the spread of extant Uralic languages to the area54. Second, as shown in our analyses, the admixture patterns found in historic and modern Uralic speakers are complex and in fact inconsistent with a single admixture event. Therefore, even if the Siberian genetic component partly spread alongside Uralic languages, some Siberian ancestry may have been already present in the area from earlier admixture events."

EDAR may have come with a bit of prior admixture quite early, but then disappeared with admixture with western and southern genomes.


For those interested in reading tea leaves:

"
The steppe ancestry component within modern Europeans (green), which is associated with the Yamnaya population, is maximised in ancient Iranian populations and to a lesser extent Caucasus hunter-gatherers (CHG). This ancestry component is also present in modern Armenians from the Caucasus, Bedouins from the Arabian Peninsula and South Asian populations."


I think it is really unlikely that this Bronze Age movement into Finland represents the arrival of the Saami. Saami and Finnic are estimated by the most recent and credible dating estimas made by linguists to have a common origin probably less than 3000 years ago. Finno-Ugric itself probably dates to less than 4000 years ago. Besides the really close connections between Saami and Finnish at least until the early Iron Age, Saami also shows a strong non-Finnic substrate that has been called Laplandic. Some have proposed it was the sole remnant of the Mesolithic languages of Scandinavia, but what if it was a later arrival? Anyway that language does not seem to have been Uralic. Then perhaps just like Finnish replaced Saami in most of Finland a former Uralic language was pushed aside by Saami.
 
I think it is really unlikely that this Bronze Age movement into Finland represents the arrival of the Saami. Saami and Finnic are estimated by the most recent and credible dating estimas made by linguists to have a common origin probably less than 3000 years ago. Finno-Ugric itself probably dates to less than 4000 years ago. Besides the really close connections between Saami and Finnish at least until the early Iron Age, Saami also shows a strong non-Finnic substrate that has been called Laplandic. Some have proposed it was the sole remnant of the Mesolithic languages of Scandinavia, but what if it was a later arrival? Anyway that language does not seem to have been Uralic. Then perhaps just like Finnish replaced Saami in most of Finland a former Uralic language was pushed aside by Saami.

Sorry, where do you see that the Saami arrived in Finland in the Bronze Age?
 
Abstract

European population history has been shaped by migrations of people, and their subsequent admixture. Recently, ancient DNA has brought new insights into European migration events linked to the advent of agriculture, and possibly to the spread of Indo-European languages. However, little is known about the ancient population history of north-eastern Europe, in particular about populations speaking Uralic languages, such as Finns and Saami. Here we analyse ancient genomic data from 11 individuals from Finland and north-western Russia. We show that the genetic makeup of northern Europe was shaped by migrations from Siberia that began at least 3500 years ago. This Siberian ancestry was subsequently admixed into many modern populations in the region, particularly into populations speaking Uralic languages today. Additionally, we show that ancestors of modern Saami inhabited a larger territory during the Iron Age, which adds to the historical and linguistic information about the population history of Finland.


https://www.nature.com/articles/s41467-018-07483-5

cKG1UoH.png

Am I the only one who's surprised by Finns being modeled as 10% Nganasan?
I mean Nganasans look like Eskimos. Prior to this study, the estimates were more conservative, up to 5% from what I noticed.
 
I've been saying since the first Haak paper came out that I thought the models over-estimated the amount of actual steppe "Yamnaya" genetic material initially moving into the far northeast, and ancient dna seems to be proving that correct. There have been a lot of changes in the genomes of this area since then, with movement from more "Yamnaya" heavy and also EEF heavy areas. There are also the "Siberian" like migrations, which contain not only ANE like, but also East Asian like material, or perhaps we should say that the "Siberian" component is on a cline between West Eurasian and East Eurasian?

I just think that the initial models were a bit simplistic and that with more ancient dna they have to be refined.

Also, ADMIXTURE doesn't work that well for those parts of Europe with more than just the three major ancient population groups. The more complicated the population history, the easier it is to confound it.


Likewise, the Basques are an issue in terms of the standard steppe theory. They carry downstream R1b, but it doesn't look to me like what was added in the Bronze Age was the standard Yamnaya "mix". There's too little CHG or, in these charts, Iran Neo in them. One possible explanation is that this could be because various streams of migration had various admixtures. It's a mistake, imo, to imagine that this was a simplistic movement of one homogeneous group. These weren't "national" entities in the way we think of them now, and even those "national" entities aren't homogeneous.

I was reading Aasko Parpola's very "novel" take on the PIE issue and though I disagree with him in most of the conclusions (though not about the fundamental archaeological/historical premises) he caused me to remind a part of the PIE expansion story that may be a little underestimated: the "steppization" of the late phase of Cucuteni-Tripolye and its subsequent demise and possible dispersal in the Post-Tripolye societies, presumably Indo-Europeanized by that time. Maybe there was much more to the Late PIE speaking area than the "classic" Yamnaya mix in the very specific 50-60% EHG 40-50% proportions that are often used as "the" steppe ancestry per se. It is plausible that LPIE first expanded for some centuries absorbing and significantly but only partially changing the genetic landscape of some of its later extensions. The sourde of steppe ancestry in CWC for instance does not look like actual Yamnaya (which is more based on samples east of the Don than west of it). Apart from the prevalence of R1a-M417 the specific mix of ancestral admixtures IIRC was not the same, though it was pretty similar. The case of the Basques (and potentially the mystery of L51) could be solved if the steppe ancestry in them came from a source much richer in EEF and with more EHG than CHG (more similar to Neolithic Ukraine, let us say), possibly in the Post-Tripolye territory near the Carpathians. Do you know if there are aDNA samples fully analyzed from that region right after the Yamnaya expansion?
 
Sorry, where do you see that the Saami arrived in Finland in the Bronze Age?

Oh okay, I probably misunderstood you. Were you asserting that the Saami were genetically the pre-Uralic inhabitants of Fennoscandia/Karelia who mixed with and adopted some Finno-Ugric language only to be later pushed northward by the Finns? I thought you were implying the pre-Saami Finnic language itself had come to Finland during the Bronze Age when you hypothesized if the Siberian/East Asian ancestry had been brought by Finno-Ugric speakers.
 
When I look at the distributions of WHG and EHG I'm drown in doubts. Saami with no EHG, only WHG, compared to Bolshoi Oleni Ostrov, and with some serious input of Yamnaya in Saami, supposed to be come from East with Finnic languages, at least for a part.
There is no more any paper using the same basis as the others for admixture, it's so puzzling... I 've lostmy latin and my celtic!

I also think the results look very confusing especially for more northeastern populations in thosecadmixture charts.some really weird and unexpected results (no extra EHG in Northeast Europe modern populations is a bit hard to swallow when the Siberian and even extra WHG are there). I guess there must be some unwitting issue in these models, though I am not knowledgeable enough to point it out. I noticed that the vast majority of the samples with EHG lack any Yamnaya at all and vice-versa, but that may be just a coincidence...

The Yamnaya admixture looks inflated in comparison with EHG in the Saami and other modern and ancient NE populations, but I do not think a good chunk of Yamnaya is that surprising. The Uralic and even more certainly the Finni-Ugric homeland were probably close to those of PIE and Indo-Iranian and Balto-Slavic territories, and they later absorbed and mixed with CWC populations (and Germanic in Scandinavia). So even if at their origin they were not rich in Yamnaya admixture at all they certainly acquired a lot of Yamnaya-like admixture over the millennia.
 
Am I the only one who's surprised by Finns being modeled as 10% Nganasan?
I mean Nganasans look like Eskimos. Prior to this study, the estimates were more conservative, up to 5% from what I noticed.

No, I'm not surprised. If I remember correctly I think Haak et al put it at around 7%.

Ten percent of that ancestry wouldn't be obvious, especially because people draw conclusions mainly by pigmentation, and the Finns are very fair. Even given that, many Finns do look quite different from other Scandinavians to me, although part of that is also the large percentage of WHG I think.
 
There "is" EHG in modern populations of Northeast Europe; it's just that there is no "extra" amount. From looking at the ADMIXTURE chart including Yamnaya, it looks to me that the "extra" ancestry present in the modern northeast is Nganasan and WHG, which is exactly what Haak said. Either they or one of the other early papers also said that it was unlikely SHG (which showed movement of EHG into the area) contributed much to subsequent populations.

As I pointed out above, there have been changes in the area since. I think it may be as simple as that.
 
There "is" EHG in modern populations of Northeast Europe; it's just that there is no "extra" amount. From looking at the ADMIXTURE chart including Yamnaya, it looks to me that the "extra" ancestry present in the modern northeast is Nganasan and WHG, which is exactly what Haak said. Either they or one of the other early papers also said that it was unlikely SHG (which showed movement of EHG into the area) contributed much to subsequent populations.

As I pointed out above, there have been changes in the area since. I think it may be as simple as that.

I should have checked the charts more carefully. These academics model European HG ancestry differently, and that may be the cause of some of our issues with the percentages. They don't seem to see much of a difference between WHG, SHG, and EHG, with WHG and SHG, in particular, being almost identical except for a smidgen of LBK like ancestry in the WHG. EHG is different in having about 10% Amerindian, i.e. something ANE heavy, and from 5-10% Iran Neo. The latter has come up in other recent papers, and would explain that Karelia J. They were perhaps slightly to the east? Or it may be that they were in the area but the "minority" ancestry is being assigned to Iran Neo and Nganasan?
 
I also think the results look very confusing especially for more northeastern populations in thosecadmixture charts.some really weird and unexpected results (no extra EHG in Northeast Europe modern populations is a bit hard to swallow when the Siberian and even extra WHG are there). I guess there must be some unwitting issue in these models, though I am not knowledgeable enough to point it out. I noticed that the vast majority of the samples with EHG lack any Yamnaya at all and vice-versa, but that may be just a coincidence...

The Yamnaya admixture looks inflated in comparison with EHG in the Saami and other modern and ancient NE populations, but I do not think a good chunk of Yamnaya is that surprising. The Uralic and even more certainly the Finni-Ugric homeland were probably close to those of PIE and Indo-Iranian and Balto-Slavic territories, and they later absorbed and mixed with CWC populations (and Germanic in Scandinavia). So even if at their origin they were not rich in Yamnaya admixture at all they certainly acquired a lot of Yamnaya-like admixture over the millennia.

It's not the presence of some taste of Yamnaya that amazed me but the HUGE weight of it and the total absence of EHG, what is purely incredible. Angela answered me, mentioning an HAAK supplement I have not red yet. I would have prefered having all the diverse admixtures results at hand at the same time. It ressembles a labyrinth!
That said, I saw so different %'s in admixtures concerning Yamnaya and its EHG and CHGlike components (and other old pops) that I 'm loosing some confidence in auDNA surveys precision.
 
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