Hungarian Yamnaya predictions

About ten thousand ancient burial mounds still stand in the Carpathian Basin and surrounds. Many of these kurgans or tumuli show direct archeological links with the highly mobile Yamnaya culture of the Pontic-Caspian steppe to the east, and may have been built by Yamnaya migrants.

The testing of ancient DNA from the remains in these burials is important, because the results are likely to be informative about the profound genetic, cultural and linguistic changes that took place in what is now Hungary and the Balkans during the Copper and Bronze Ages.

But, alas, probably to the disappointment of some readers, my great prediction is that they're not going to be overly relevant to what happened at this time in Northern and Western Europe, and won't upend the current consensus that the Corded Ware culture (CWC) was the main vector for the spread of steppe ancestry and Indo-European languages into these parts of the continent.

The important thing to understand about the Yamnaya expansion into the Carpathian Basin is that it mostly stopped at the Tisza River. It's true that some archeological cultures west of the Tisza, such as Mako and Vucedol, do show fairly strong Yamnaya influences, but they can't be regarded as part of the Yamnaya colonization of Central Europe. Below is a slightly modified map from Heyd 2011 to illustrate my point.

In fact, four early Yamnaya period samples from one of the few kurgans west of the Tisza have already been published along with the Olalde et al. 2018 paper on the Bell Beaker culture (BBC). And one of these samples, labeled I5117, even represents a male buried in a Yamnaya-like pose. But this is how three of these individuals cluster in my Principal Component Analysis (PCA) of ancient West Eurasian genetic variation.

They sit firmly among other Copper Age and Neolithic samples from west of the Pontic-Caspian steppe. In other words, they show practically zero Yamnaya-related or steppe ancestry. Moreover, both of the males belong to Y-haplogroup G2a-L91, which is yet to be found in any samples from the Copper and Bronze Age steppe.

That's not to suggest, however, that the spread of the Yamnaya culture into the Carpathian Basin was a cultural process with little or no genetic impact. It probably wasn't, because five samples labeled "Yamnaya Hungary" were featured in the Wang et al. 2018 preprint on the genetic prehistory of the Greater Caucasus, and judging by their PCA and ADMIXTURE results (in the figure below from the said preprint) they're not very different from most other Yamnaya samples, such as those from far to the east in Kalmykia or Samara.

But the point I'm making is that not every one of the ten thousand kurgans and tumuli in the Carpathian Basin and surrounds was built by newcomers from the steppe, and, thus, my other prediction is that a fair proportion of the Yamnaya-related burial mounds, especially west of the Tisza, might contain remains without any steppe ancestry.

As far as I know, the Y-haplogroups of the aforementioned Yamnaya Hungary samples haven't yet been reported anywhere. But there are three ancients in the Mathieson et al. 2018 paper on the genetic prehistory of southeastern Europe that are probably highly informative about what we can expect in this context, because based on their archeology and ancestry, they're likely to be closely related to the Hungarian Yamnaya population. They are:

Balkans_BronzeAge I2165: Y-hg I2a-L699 3020-2895 calBCE

Vucedol_Croatia I3499: Y-hg R1b-Z2103 2884-2666 calBCE

Yamnaya_Bulgaria Bul4: Y-hg I2a-L699 3012-2900 calBCE

That's not much to work with, you might say. Perhaps, but keep in mind that R1b-Z2103 has now been reported in Yamnaya samples from Ciscaucasia, Kalmykia, and Samara, and I2a-L699 in a Yamnaya singleton from Kalmykia. Thus, a lot of outcomes are still possible, but some are more likely than others. So I'm expecting most Hungarian Yamnaya males to belong to R1b-Z2103 and I2a-L699, or perhaps even the other way around!

However, in line with my great prediction, I don't expect to see any R1a-M417 or R1b-L51, the two most common Y-halogroups among present-day Europeans living north and west of the Balkans. And I think that if these markers do actually show up, then they'll be represented by nowadays rare or even extinct lineages that aren't very important to the peopling of Europe. Any thoughts? Feel free to share them in the comments.

See also...

Hungarian Yamnaya > Bell Beakers?

Single Grave > Bell Beakers

Dutch Beakers: like no other Beakers

Dutch Beakers: like no other Beakers

In my last two blog posts I tried to explain why the so called Bell Beakers of prehistoric Europe cannot be confidently derived in any significant way from the Yamnaya population of the Carpathian Basin, and are more likely to have been an offshoot, in varying degrees, of the Single Grave or Corded Ware people of the Lower Rhine region (see here and here).

To help drive my message home, below is a series of new Principal Component Analysis (PCA) plots that illustrate the unique position of Dutch Beakers from the Lower Rhine relative to the Corded Ware population of Germany and all the other Beaker groups sampled to date. The relevant datasheet is available here.

The Dutch Beakers don't exactly sit between the Corded Ware and the other Beaker samples, but generally at the apex of their clusters, suggesting to me that they're not a mixture between Corded Ware and one or more of the other Beaker groups, but rather, as per my recent argumentation, a genetically homogeneous, relatively unique and thus long-standing Corded Ware-related population that may have contributed significant gene flow to the other Beaker groups.

Please note also that all of these outcomes can be confirmed with various types of formal statistics. I know this because I've done it.

See also...

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

Single Grave > Bell Beakers

I've been studying in detail the genetic substructures within the Bell Beaker population with formal statistics and Principal Component Analyses (PCA). As far as I can see, among the two most homogeneous, and thus least likely to be recently admixed, Beaker groups are the Dutch Beakers and also the Dutch and British Beaker males belonging to Y-haplogroup R1b-P312. This, of course, makes good sense, because both the Dutch and British Beakers are so called Rhenish Beakers.

The results are also in line with the observation that the Dutch Beakers are the quintessential Beakers in terms of physique, with three quarters or more sporting exceedingly brachycephalic, planoccipital skulls (like this).

Moreover, these two Beaker groups are among the most Yamnaya-like Beakers, with almost as much Yamnaya-related ancestry as the Corded Ware culture samples from Germany (~60% vs ~70%). As a result, in my PCA of ancient West Eurasian genetic variation the Dutch Beakers form a more or less continuous, west to east cline with these and other Corded Ware individuals that runs all the way to the Yamnaya cluster.

In the same PCA, the R1b-P312 Dutch and British Beaker males form a tight cluster at the apex of a Beaker cline that stretches to European Neolithic groups with no steppe ancestry. The only Beaker who is positioned clearly east of the Dutch/British R1b-P312 Beaker cluster is from Hungary, and in all likelihood he harbors recent Yamnaya ancestry because his Y-haplogroup is the Yamnaya-specific R1b-Z2103.

These findings potentially have important implications for the origins of the Dutch Beakers and the Beakers who dominated much of Central and Western Europe during the Late Neolithic and Bronze Age, and these are:

- the Dutch Beakers are unlikely to be the result of a recent migration from afar into what is now The Netherlands and surrounds, but rather the descendants, by and large, of the earlier local Single Grave (and thus Corded Ware) populations

- the R1b-P312 lineages in the Dutch and British Beakers probably derive from Single Grave R1b-P312, which suggests that R1b-P312 was common among some clans within the Corded Ware culture

- the spread of most of the Yamnaya-related or steppe ancestry and quintessential Beaker physique across the Beaker world and into Western Europe can probably be blamed on the massive expansions of Beakers from what is now The Netherlands and surrounds (ie. the Lower Rhine region)

- late Yamnaya groups contributed some ancestry to eastern Beaker groups, such as those in the Carpathian Basin, but the Dutch Beakers acquired their high level of Yamnaya-related ancestry from their Single Grave predecessors, who, in turn, acquired it from their proto-Corded Ware ancestors from the steppe.

Admittedly, I find the discussion about the origin of the Bell Beaker cultural package somewhat confusing. For all I know, it might have come from Iberia, the Carpathian Basin, or even North Africa. But this post isn't about that, it's about the homeland of the classic Beaker warrior male, with his R1b-P312, Corded Ware-like genome-wide genetic structure and brachycephalic skull. I'm almost certain now that this was the Lower Rhine region.

See also...

Hungarian Yamnaya > Bell Beakers?

Dutch Beakers: like no other Beakers

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

Hungarian Yamnaya > Bell Beakers?

Ever since the publication of the Olalde et al. Beaker paper (see here), there's been a lot of talk online about Hungarian Yamnaya as the most likely source of the Yamnaya-related, R1b-P312-rich northern Bell Beakers who went on to dominate much of Central and Western Europe during the Late Neolithic and Bronze Age.

Certainly, this is still possible, and we might find out soon if it's true because several Hungarian Yamnaya samples are apparently about to be published. But I wouldn't bet the proverbial farm on it just yet.

The most Yamnaya-like Beaker in the Olalde et al. dataset and ancient DNA record to date is from the Szigetszentmiklós burial site, which is indeed in present-day Hungary. But this individual, labeled I2787, is dated to just 2457–2201 calBCE, which isn't an early date for a Beaker and probably a couple hundred years past the proto-Beaker time frame.

Moreover, he belongs to Y-haplogroup R1b-Z2103, a paternal marker most closely associated in the ancient DNA record with eastern Yamnaya groups. And he doesn't exactly look like a classic northern Beaker, because he doesn't have a brachycephalic head with an exceedingly flat occiput (like this).

So I'd say that this is either an acculturated Beaker of recent Yamnaya origin, or perhaps the son of a Yamnaya father and Beaker mother. Below are several qpAdm mixture models that I ran to explore the latter possibility. They look very solid.

Beaker_Hungary_I2787
Beaker_Bavaria 0.442±0.045
Yamnaya_Samara 0.558±0.045
chisq 8.562
tail prob 0.73982
Full output

Beaker_Hungary_I2787
Beaker_Czech 0.441±0.045
Yamnaya_Samara 0.559±0.045
chisq 10.009
tail prob 0.61513
Full output

Beaker_Hungary_I2787
Beaker_The_Netherlands 0.576±0.062
Yamnaya_Samara 0.424±0.062
chisq 11.469
tail prob 0.489238
Full output

The idea that I2787 is a Beaker with recent Yamnaya ancestry isn't an original one. It was put forth very eloquently and convincingly months ago by the Bell Beaker Blogger himself:

Szigetszentmiklós Cemetery (Santa's Six Foot Elves)

I2786 is another Beaker male from the Szigetszentmiklós site who shows excess Yamnaya-related ancestry compared to most other Beakers. Again, it's likely that this individual harbors recent Yamnaya ancestry, because his Y-haplogroup is I2a-M223, which has been recorded in eastern Yamnaya alongside R1b-Z2103.

So my gut feeling for now is that Hungarian Yamnaya samples will mostly belong to Y-haplogroups R1b-Z2103 and I2a-M223, rather than R1b-P312, and thus they won't fit the bill in any obvious way as the population that may have given rise to northern Beakers.

One of the oldest individuals in the ancient DNA record belonging to R1b-P312 is I5748, a Beaker dated to 2579–2233 calBCE from the Oostwoud-Tuithoorn burial site in what is now West Frisia, The Netherlands.

Interestingly, this part of Northwestern Europe was home to the Single Grave population shortly before I5748 was alive. And the Single Grave culture is a variant of the Corded Ware culture. So can anyone tell me if there's any evidence that I5748 and his kind were relative newcomers to West Frisia, from, say, somewhere in the direction of the Carpathian Basin? If not, then what are the chances that northern Beakers are by and large the descendants of the Single Grave people?

In fact, there's not much difference in terms of genome-wide genetic structure between the Beakers from the Oostwoud-Tuithoorn site and Corded Ware people from what is now Germany. The Principal Component Analysis (PCA) below illustrates this well. But, you might say, Corded Ware males by and large belong to Y-haplogroup R1a-M417. Yep, but this doesn't mean that R1b-P312 wasn't common in some Single Grave clans.

At this stage, I don't have a clue where the northern Beakers might have come from, and unfortunately I don't have any inside information about the Y-haplogroups of Hungarian Yamnaya. I don't even know if any Single Grave samples are being analyzed. But I'll leave you with this map from a recent paper by French archeologist and Beaker expert Olivier Lemercier (see here). To me it suggests rather strongly that northern Beakers developed from the synthesis of Corded Ware newcomers to Western Europe and indigenous Western Europeans. As far as I can tell, that's what the paper basically argues as well.

See also...

Single Grave > Bell Beakers

Dutch Beakers: like no other Beakers

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

PIE Urheimat poll: two or three options left

If we let ancient DNA dictate the terms in the Proto-Indo-European (PIE) homeland debate ahead of historical linguistics and archeology, then, as far as I can see, there are two or three realistic options for the location of the said homeland. Here they are, in order of my own preference:

1) The Don-Caspian steppe around 4,300 BCE (see here). The ancestors of the Hittites and other Anatolian speakers also came from this homeland and entered Anatolia via the Balkans (or, less likely, the Caucasus) in fairly small groups sometime between 4,000 and 2,000 BCE. A lot of samples from Bronze Age Anatolia are needed to confirm or debunk the presence of steppe ancestry there.

2) The eastern Balkans during the peak of the ostentatious Copper Age in the region. Proto-Indo-European developed in the wealthy Chalcolithic communities of the western Black Sea coast and quickly spread both into the steppes and Anatolia via elite and trade contacts, and thus with minimal gene flow. Proto-Indo-European minus Anatolian, or PNIE, then spread from Eastern Europe during the Bronze Age with the mass migrations of the Yamnaya and closely related populations. A lot of samples from Chalcolithic western Anatolia are needed to confirm or debunk that people moved from the Balkans into Anatolia at this time.

3) Transcaucasia and/or nearby around 10,000 BCE. Proto-Indo-European, or rather Indo-Hittite, is much older than generally accepted, and came from the Epipaleolithic northern Near East. It was introduced into the steppes by foragers of the so called Caucasus Hunter-Gatherer (CHG) type, where it eventually became Proto-Indo-European minus Anatolian, or PNIE. Proto-Anatolian was spoken by closely related CHG-like foragers who stayed in the northern Near East.

Admittedly, that last theory is way out there, and at the moment, has about as much chance of being accepted by most historical linguists as Out-of-India. But the one advantage that is has over the other two proposals is that it doesn't need any additional sampling of ancient DNA.

I'll probably get grilled in the comments as to why I didn't include a proposal with the Maykop culture as the PIE community, or at least the Indo-Europeanizing agent in the steppe. Honestly, after seeing the ancient DNA from a wide range of Maykop remains courtesy of Wang et al., I think the chances that Maykop was an Indo-European-speaking culture are low. Indeed, both the Maykop genome-wide data and uniparental markers scream "Northwest Caucasian" to me.

Also, if the Caucasus was the PIE homeland, or even a major expansion point for early Indo-European languages, then considering its widely accepted status as a linguistic hotspot and refuge, it's fair to expect that it should still harbor at least one highly diverged Indo-European language. Is there any evidence that it ever did?

Below is an interactive poll. Please vote for one of the three options and feel free to let us know in the comments why you made the choice that you did. I might add more options to the poll if compelling reasons are given in the comments to do so.

PIE Urheimat poll

Don-Caspian steppe around 4,300 BCE

Chalcolithic eastern Balkans

Epipaleolithic northern Near East

Created with PollMaker

See also...

Yamnaya: home-grown

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

On the doorstep of India

The PIE homeland controversy: January 2019 status report

Last year, the preprint that claimed to have presented archaeogenetic data that opened up the possibility of the Proto-Indo-European (PIE) homeland being located south of the Caucasus was, ironically, also the preprint that considerably strengthened my confidence that the said homeland was actually located north of the Caucasus.

Of course, I'm talking about the Wang et al. manuscript at bioRxiv, which is apparently soon to be published as a peer-reviewed paper in Nature Communications (see here).

It'll be fascinating to observe if and how the peer-review process has impacted on the preprint, and especially its conclusion. My impression was that the authors seemed pretty sure that the Maykop people gave rise to the Yamnaya culture, or at least Indo-Europeanized it. But, as far as I saw, the archaeogenetic data didn't bear this out at all, and instead showed a lack of any direct, recent and meaningful genetic relationship between Maykop and Yamnaya (see here). Was this also picked up by the peer reviewers? We shall see.

Moreover, there was some exceedingly interesting fine print in the manuscript's supplementary information:

Complementary to the southern [Darkveti-Meshoko] Eneolithic component, a northern component started to expand between 4300 and 4100 calBCE manifested in low burial mounds with inhumations densely packed in bright red ochre. Burial sites of this type, like the investigated sites of Progress and Vonyuchka, are found in the Don-Caspian steppe [10], but they are related to a much larger supra-regional network linking elites of the steppe zone between the Balkans and the Caspian Sea [16]. These groups introduced the so-called kurgan, a specific type of burial monument, which soon spread across the entire steppe zone.

Always read the fine print, they say. And they're right. Imagine if I only read the preprint's conclusion and missed this little gem; I'd probably think that the PIE homeland was located south of the Caucasus rather than on the Don-Caspian steppe.

Wow, proto-kurgans with inhumations densely packed in bright red ochre? A supra-regional network linking the elites of the steppe all way from the Balkans to the Caspian Sea? An expansionist culture? And, as evidenced by the ancient DNA from the Progress and Vonyuchka sites, a people who may well have been in large part ancestral to the Yamnaya, Corded Ware and Andronovo populations, that have been identified based on archeological and historical linguistics data as the main vectors for the spread of Indo-European languages as far as Iberia in the west and the Indian subcontinent in the east.

I wonder if the authors actually asked themselves who these people may have been, before so haphazardly turning to Maykop and, ultimately, the Near East, as the likely sources of the Yamnaya culture? To me they look like the Proto-Indo-Europeans and true antecedents of Yamnaya.

So as things stand, my pick for the PIE homeland is firmly the Don-Caspian steppe. And I genuinely thank Wang et al., and indeed the Max-Planck-Institut für Menschheitsgeschichte (aka MPI-SHH), for their assistance.

But, you might ask, what about the Hittites? Yes, I realize that no one apart from me and a few of my readers here can find any steppe ancestry in the so called Hittite genomes published to date. However, consider this: if the PIE homeland really was on the steppe, and a dense sampling strategy of Hittite era Anatolia fails to turn up any unambiguous steppe ancestry in at least a few individuals, then there has to be an explanation for it. But let's wait and see what a dense sampling strategy of Hittite era Anatolia actually reveals before we go that far.

See also...

The PIE homeland controversy: August 2019 status report

Yamnaya: home-grown

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

R-V1636: Eneolithic steppe > Kura-Araxes?

Ancient samples from the Wang et al. preprint on the genetic prehistory of the Greater Caucasus are now available as BAM files at the European Nucleotide Archive (see here). I've requested the genotype data from the authors and I'm eagerly awaiting their response.

But various online genetic genealogy communities are already studying in detail the Y-chromosome data from the BAM files. One interesting outcome is that both of the Eneolithic steppe males, PG2001 and PG2004, apparently belong to Y-haplogroup R-V1636 (see here). This extremely rare subclade of R1b has apparently also been found in an ancient individual from what is now Armenia associated with the Kura-Araxes culture: Armenia_EBA I1635.

Importantly, the Eneolithic steppe males are dated to 4336-4047 calBCE, and don't show any recent genome-wide ancestry from south of the Caucasus, while the Kura-Araxes individual is dated to just 2619-2465 calBCE.

It'll be interesting to see whether Armenia_EBA I163 shows any genome-wide admixture from north of the Caucasus when I can test this with these new Wang et al. Eneolithic samples from the southernmost steppes. But in any case, if the R-V1636 link between the Eneolithic steppe and Kura-Araxes is real, then this is more evidence of migrations from the steppe across the Greater Caucasus into the Near East during the Eneolithic and/or Bronze Age.

Such population movements could potentially explain the appearance of Hittite and other closely related Indo-European languages in Anatolia during the Bronze Age.

See also...

Steppe ancestry in Chalcolithic Transcaucasia (aka Armenia_ChL explained)

Likely Yamnaya incursion(s) into Northwestern Iran

A potentially violent end to the Kura-Araxes Culture (Alizadeh et al. 2018)

The Hajji Firuz fiasco

The specter of Hajji_Firuz_ChL I2327 still haunts us. Judging by some recent comments that I've seen here and elsewhere, it seems that a good number of confused souls haven't yet given up hope that this ancient sample represents a Near Eastern population ancestral to the Yamnaya people of Early Bronze Age Eastern Europe. But the chances of this are slim to none, because...

- Hajji_Firuz_ChL I2327 is the only (supposedly) pre-Yamnaya individual to date in the now ample West Asian ancient DNA record who belongs to Y-haplogroup R1b-Z2103, R1b-M269, or even R1b, which has got to tell you something about the reliability of his early dating

- overall the genome-wide structure of Hajji_Firuz_ChL I2327 most certainly doesn't fit the profile of the Near Eastern-related half of the Yamnaya genotype

- in fact, practically every analysis that I've run with Hajji_Firuz_ChL I2327 suggests that he harbors Yamnaya or Yamnaya-related genome-wide ancestry, which makes sense considering his Yamnaya-specific Y-haplogroup, don't you think?

Heck, even if Hajji_Firuz_ChL I2327 is more or less accurately dated, and really was alive during the Chalcolithic period, then considering the points I've made above, the only honest explanation for his presence that early in what is now Iran is that there was a migration of a Yamnaya-like people from the steppes across the Caucasus during the Chalcolithic.

I already wrote a post on this topic back in April, and a lengthy discussion ensued, with most of the commentators agreeing with my stance. But my efforts haven't had much of an impact outside of this blog. It's possible that my post was too confusing, so I went back today and rewrote it, also adding new stats and mixture models to help me drive home my point. Here's the link...

Likely Yamnaya incursion(s) into Northwestern Iran

Of course, no matter how strong my arguments are, many people will choose to disagree with me nevertheless and believe what they want to believe, because this is such an emotional issue for them. I don't want to get into the details about that here, but suffice to say that it's imperative for many people, particularly those of Near Eastern and Southern European backgrounds, that the origin of Yamnaya is somehow, by hook or by crook, put south of the Caucasus. I'm not kidding. It's a pointless cause though, especially now considering all of the new ancient DNA data from Eastern Europe that make this scenario about as likely as Out-of-India (see here).

See also...

Yamnaya: home-grown

Big deal of 2018: Yamnaya not related to Maykop

Ahead of the pack

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

Italians are interesting people (Raveane et al. 2018 preprint)

Over at bioRxiv at this LINK. As far as I can see from skimming through the preprint, it's a thorough effort with conclusions that make good sense. But, needless to say, it'll be very useful to plug the dataset from this study into the Global25 to see what these new Italian samples are really made of. Here's the abstract, emphasis is mine.

European populations display low genetic diversity as the result of long term blending of the small number of ancient founding ancestries. However it is still unclear how the combination of ancient ancestries related to early European foragers, Neolithic farmers and Bronze Age nomadic pastoralists can fully explain genetic variation across Europe. Populations in natural crossroads like the Italian peninsula are expected to recapitulate the overall continental diversity, but to date have been systematically understudied. Here we characterised the ancestry profiles of modern-day Italian populations using a genome-wide dataset representative of modern and ancient samples from across Italy, Europe and the rest of the world. Italian genomes captured several ancient signatures, including a non-steppe related substantial ancestry contribution ultimately from the Caucasus. Differences in ancestry composition as the result of migration and admixture generated in Italy the largest degree of population structure detected so far in the continent and shaped the amount of Neanderthal DNA present in modern-day populations.

Raveane et al., Population structure of modern-day Italians reveals patterns of ancient and archaic ancestries in Southern Europe, Posted December 13, 2018, bioRxiv, doi: https://doi.org/10.1101/494898

See also...

Greeks in a Longobard cemetery

Migration of the Bell Beakers—but not from Iberia (Olalde et al. 2018)

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

How did Y-haplogroup N1c get to Bolshoy Oleni Ostrov?

Y-haplogroup N1c probably entered Europe from Siberia during the Bronze Age or the Eneolithic period. It first appears in the European ancient DNA record in two samples from a burial site at Bolshoy Oleni Ostrov, in the Kola Peninsula, dated to 1523±87 calBCE (see here). These individuals also harbor significant Siberian genome-wide ancestry, but it's possible that this is in large part a coincidence, and that N1c spread into the Kola Peninsula from the south in a population of overwhelmingly European ancestry.

Crazy, huh? Not really. Consider the qpAdm mixture models below for BOO002 and BOO004, the two males from the Bolshoy Oleni Ostrov site belonging to N1c, and BOO006, a female and the most Siberian-admixed individual from the same site. Although BOO002 and BOO004 show a lot of Nganasan-related and thus Siberian ancestry, they also require significant input from a source closely related to Baltic_BA, a fully European Bronze Age population from the East Baltic region. On the other hand, BOO006 doesn't need Baltic_BA for a successful model.

BOO002_&_BOO004
Baltic_BA 0.124±0.029
EHG 0.406±0.032
Nganasan 0.469±0.017
chisq 10.847
tail prob 0.286316
Full output

BOO006
Baltic_BA 0.065±0.043
EHG 0.265±0.084
Nganasan 0.517±0.033
West_Siberia_N 0.152±0.074
chisq 8.847
tail prob 0.355397
Full output

BOO006
EHG 0.367±0.049
Nganasan 0.544±0.031
West_Siberia_N 0.089±0.063
chisq 9.878
tail prob 0.360451
Full output

Keep in mind that N1c is very common in the East Baltic today in populations with minimal Siberian genome-wide ancestry. Indeed, Latvians and Lithuanians can often be modeled with no Siberian input. Thus, it's likely that by the time N1c arrived in the East Baltic, probably during the Late Bronze Age or Early Iron Age, it did so with populations with heavily diluted Siberian genome-wide ancestry. Such groups may also have taken N1c north of the Baltic and into the Kola Peninsula.

See also...

On the trail of the Proto-Uralic speakers (work in progress)