Uralians of the Sargat horizon

Many years ago, well before the start of the ancient DNA revolution, someone made the very clever inference that the N-Tat Y-chromosome marker was closely associated with the expansion of Uralic languages.

Since then, N-Tat has been renamed several times over, to the point that I no longer know what it's called, but the aforementioned inference has turned into a very solid consensus backed up by a wide range of studies focusing on modern and ancient DNA.

Nowadays, Y-haplogroup N-L1026, a subclade of N-Tat, is seen as the main genetic signal of the Uralic expansions, along, of course, with Nganasan-related genome-wide genetic ancestry.

A recent paper at Science Advances by Gnecchi-Ruscone et al. featured the first ever genome-wide samples from the Sargat horizon, which is an Iron Age archeological formation in western Siberia normally associated with the Ugric branch of the Uralic language family. Surprisingly, and disappointingly, the authors failed to investigate this widely accepted connection.

If we go by the Y-haplogroup classifications in the paper, which may or may not be the smart thing to do, at least two of the Sargat horizon males belong to N-L1026, and one also to the more derived N-Z1936 subclade, which has been found in the remains of Hungarian Conquerers from Medieval Hungary. Of course, Hungarian is an Ugric language generally thought to have been introduced into the Carpathian Basin by the Hungarian Conquerers who originally came from western Siberia.

That's probably enough to corroborate the association between the Sargat horizon and the spread of Ugric/Uralic languages, but let's also take a quick look at the autosomal DNA of these Sargat individuals. Firstly, here's a Principal Component Analysis (PCA), based on Global25 data and produced with the Vahaduo G25 Views online tool. The results are self-explanatory.

Interestingly, I can't get a decent statistical fit when I try to reproduce the four-way qpWave/qpAdm model done by Gnecchi-Ruscone et al., probably mostly because my right pops or outgroups are different. This suggests to me that there's something important missing in their model.

Sargat_IA
MNG_Khovsgol_LBA 0.203±0.045
RUS_Ekven_IA 0.183±0.044
RUS_Sintashta_MLBA 0.545±0.014
TKM_Gonur1_BA 0.068±0.013
chisq 16.805
tail prob 0.0186971
Full output

So how about if I replace RUS_Ekven_IA with kra001, the oldest Nganasan-like individual in the ancient DNA record (see here), and MNG_Khovsgol_LBA with KAZ_Mereke_MBA, to add a more local stream of ancestry?

Sargat_IA
KAZ_Mereke_MBA 0.135±0.017
kra001 0.301±0.007
RUS_Sintashta_MLBA 0.499±0.023
TKM_Gonur1_BA 0.066±0.015
chisq 8.872
tail prob 0.262001
Full output

That's a better statistical fit and also, I'd say, a more realistic model, at least in terms of distal ancestry proportions. Note that Nganasan-related ancestry makes up 30% of the genome-wide genetic structure of the Sargat samples, which again corroborates the view that Uralic languages were spoken within the Sargat horizon.

Update 28/04/21: This is the best qpAdm model that I could find for Sargat_IA, at least in terms of the chisq and tail prob. It shows that the Sargat population was in large part very similar to that of KAZ_Pazyryk_IA.

Sargat_IA
KAZ_Mereke_MBA 0.032±0.016
KAZ_Pazyryk_IA 0.698±0.016
RUS_Sintashta_MLBA 0.236±0.021
TKM_Gonur1_BA 0.034±0.014
chisq 2.023
tail prob 0.958561
Full output

It's missing kra001, because KAZ_Pazyryk_IA packs enough kra001-related ancestry for the job.

KAZ_Pazyryk_IA
KAZ_Mereke_MBA 0.144±0.018
kra001 0.429±0.008
RUS_Sintashta_MLBA 0.378±0.026
TKM_Gonur1_BA 0.049±0.018
chisq 8.899
tail prob 0.259983
Full output

The fact that KAZ_Pazyryk_IA can be modeled with significant kra001-related ancestry isn't surprising, considering that its territory was located in Siberia. However, my model doesn't necessarily prove that the Sargat population was largely or even partly of Pazyryk origin. Indeed, N-L1026 hasn't yet appeared in any Pazyryk remains.

See also...

The Uralic cline with kra001 - no projection this time

First taste of Early Medieval DNA from the Ural region

Hungarian Conquerors were rich in Y-haplogroup N

More on the association between Uralic expansions and Y-haplogroup N

It was always going to be this way

On the association between Uralic expansions and Y-haplogroup N

The Uralic cline with kra001 - no projection this time

A whole lot of nonsense was posted online, often by people who should've known better, after I claimed that kra001 was a solid proxy for a proto-Uralic genome (see here).

For those of you who still don't get it, below are three Principal Component Analysis (PCA) plots featuring Uralic speakers and other present-day Eurasians. Kra001 is also there. These graphs are based on genotype data not reprocessed Global25 data. The relevant datasheet is available here.

Compared to my previous PCA with kra001, here I included a bigger range of East Eurasian populations to help mitigate the effects of extreme genetic drift in some of the Siberian groups, at least on the first few Principal Components (PCs). Moreover, kra001 wasn't projected onto PCs computed with modern-day samples, so he was free to influence the outcome of the PCA.

Note the east to west clines made up largely of Uralic speaking groups on the first two plots. These plots are based on PCs 1/2 and 1 /3, respectively. The third plot, based on PCs 1/4, is more complex and thus more difficult to interpret, but it also manages to isolate many of the Uralic populations from the others.

The Uralic-specific clines do intersect with the clines and clusters formed by the other linguistic groups. However, based on the three plots, the Yeniseian-speaking Kets are the only Asian group that can plausibly be confused for Uralic speakers.

Importantly, apart from the Kets, kra001 is the only Asian individual who shifts his position on all three plots as if he were a Uralic speaker. This might well be a coincidence, and we'll never know what language was spoken by kra001, but it does suggest to me that his genome is a solid proxy for a proto-Uralic genome.

See also...

First taste of Early Medieval DNA from the Ural region

The BOO people: earliest Uralic speakers in the ancient DNA record?

Fresh off the sledge

Finally, a proto-Uralic genome

Obviously, genes don't speak languages, people do. But sometimes it's possible to associate a linguistic group with a very specific genetic signature.

A while ago many of us in the blogosphere spotted an uncanny connection between the Uralic language family, Y-haplogroup N-L1026 and Nganasan-like genome-wide genetic ancestry.

As a result, we expected a Nganasan-like population rich in N-L1026 to eventually appear in the ancient DNA record, probably somewhere in Siberia and in burials from a likely proto-Uralic archeological culture. This hasn't happened yet, but we now have direct evidence that such a population must have existed somewhere deep in Siberia as early as the Bronze Age.

Kra001, whose genome was published recently along with Kilinc et al., belongs to a pre-N-L1026 lineage and, at least in terms of genome-wide genetic structure, could well be from a population directly ancestral to present-day Nganasans. Of course, the Nganasan language is part of the Samoyedic branch of Uralic.

Below is a series of Principal Component Analyses (PCA) featuring kra001. He's labeled RUS_Krasnoyarsk_BA, after the location and age of his burial. Note the obvious Uralic cline running across the plots. That is, from west to east. Kra001 is positioned at the end of this cline very close to a small cluster of Nganasans. To see interactive versions of the plots, paste the Global25 coordinates here into the relevant field here.

Admittedly, there's no way of knowing whether this individual spoke proto-Uralic or not. Indeed, he may have spoken something totally unrelated. The important point is that the very specific genetic signature shared by almost all present-day Uralic speakers, except perhaps Hungarians, is now finally represented in the ancient DNA record. And I can reveal to you that we'll soon be seeing many more ancients very similar to kra001 in upcoming papers.

See also...

The Uralic cline with kra001 - no projection this time

The BOO people: earliest Uralic speakers in the ancient DNA record?

Fresh off the sledge

The great shift

Here's a Principal Component Analysis (PCA) featuring some of the ancients from the recent Saag et al. paper at Science Advances. To see an interactive version of the plot paste the Global25 coordinates here into the relevant field here.

Note that the Fatyanovo culture agropastoralists, who are rich in Y-haplogroup R1a and steppe ancestry, cluster with present-day Eastern Europeans. On the other hand, the Volosovo culture singleton sits near the European hunter-gatherer cline that no longer exists.

This Volosovo individual belongs to Y-haplogroup Q1a. However, most of the Volosovo males whose genomes are soon to be published belong to Y-haplogroup R1b.

Thus, in much of Eastern Europe during the Bronze Age, agropastoralists rich in R1a and steppe ancestry replaced hunter-gatherers rich in R1b and with no steppe ancestry. Of course, that's not where the story ends, but I'll get back to that later this year.

By the way, the relatively high coverage Fatyanovo Y-chromosome sequences are being analyzed at YFull. You can check out the results here.

See also...

That old chestnut: Northeast vs Northwest Euros

In the last comment thread reader Greg put forth this question:

David, when are you going to explain the genetic discrepancy between Northeastern and Northwestern Europeans? You know, the one that people believe is due to Baltic Hunter-Gatherer admixture, whereas you believe it is due to genetic drift? You ought to make a post about this issue at some point, because a lot of people are wondering what's causing the differences.

Well, Greg, this issue has been discussed to the proverbial death here and elsewhere. In fact, there were two posts and rather lengthy comment threads on the same topic at this blog just a few months ago. See here and here.

Nevertheless, it seems that a fair number of people are still befuddled, so I'm going to try to explain this one last time, as briefly as a I can using just a handful of f4-stats.

Admittedly, Northeast Europeans generally do pack higher levels of indigenous European hunter-gatherer ancestry than Northwest Europeans. This is especially true of Balts, who show more of this type of ancestry than even Scandinavians in practically every type of analysis.

The f4-stats below back this up unambiguously. Note the significantly positive (>3) Z scores, which suggest that Latvians and Lithuanians harbor more Baltic hunter-gatherer-related ancestry than Norwegians and Swedes.

Chimp Baltic_HG Norwegian Latvian 0.001301 7.114
Chimp Baltic_HG Swedish Latvian 0.001017 4.205
Chimp Baltic_HG Norwegian Lithuanian 0.001023 7.341
Chimp Baltic_HG Swedish Lithuanian 0.000763 3.408

Greg, I know what you're thinking: the naysayers are right! But wait, because there's a twist to this tale. Check out these f4-stats:

Chimp Baltic_HG Norwegian Belarusian 0.000265 1.934
Chimp Baltic_HG Swedish Belarusian 0.000152 0.7
Chimp Baltic_HG Norwegian Polish 6.4E-05 0.519
Chimp Baltic_HG Swedish Polish -0.000235 -1.074

Please note, Greg, that none of the Z scores reach significance, which means that these Northwest Europeans and Slavs are symmetrically related to Baltic_HG. They're also symmetrically related to other relevant ancient groups such as the Yamnaya steppe herders. This, of course, suggests that they harbor very similar levels of basically the same ancient genetic components.

Chimp Karelia_HG Norwegian Belarusian 0.000136 0.844
Chimp Karelia_HG Swedish Belarusian 7.9E-05 0.32
Chimp Karelia_HG Norwegian Polish -4.7E-05 -0.304
Chimp Karelia_HG Swedish Polish -0.000134 -0.54

Chimp Yamnaya_Samara Norwegian Belarusian -0.000134 -1.085
Chimp Yamnaya_Samara Swedish Belarusian -6.6E-05 -0.34
Chimp Yamnaya_Samara Norwegian Polish -0.000225 -1.995
Chimp Yamnaya_Samara Swedish Polish -0.000311 -1.574

Chimp Barcin_N Norwegian Belarusian -0.000335 -2.809
Chimp Barcin_N Swedish Belarusian -0.000284 -1.491
Chimp Barcin_N Norwegian Polish -0.000222 -2.057
Chimp Barcin_N Swedish Polish -0.000318 -1.662

Chimp Baikal_N Norwegian Belarusian 0.000186 1.3
Chimp Baikal_N Swedish Belarusian -7E-05 -0.33
Chimp Baikal_N Norwegian Polish -4.6E-05 -0.351
Chimp Baikal_N Swedish Polish -0.000477 -2.277

Interestingly, pairing up Ukrainians with English samples from Cornwall and Kent produces similar outcomes. But that's because most ancient ancestry proportions in Europe show a closer correlation with latitude than longitude.

Chimp Baltic_HG English_Cornwall Ukrainian 0.000282 2.242
Chimp Baltic_HG English_Kent Ukrainian 0.000225 1.748

Chimp Karelia_HG English_Cornwall Ukrainian 0.000323 2.175
Chimp Karelia_HG English_Kent Ukrainian 0.000239 1.634

Chimp Yamnaya_Samara English_Cornwall Ukrainian -6.6E-05 -0.569
Chimp Yamnaya_Samara English_Kent Ukrainian -0.000112 -0.977

Chimp Barcin_N English_Cornwall Ukrainian -0.000519 -4.641
Chimp Barcin_N English_Kent Ukrainian -0.000598 -5.232

Chimp Baikal_N English_Cornwall Ukrainian 0.000385 2.874
Chimp Baikal_N English_Kent Ukrainian 0.00036 2.836

Now, Greg, if at least in terms of genetic ancestry, Latvians, Lithuanians, Belarusians, Poles and Ukrainians all qualify as Northeast Europeans, then what makes them different, as a group, from Northwest Europeans? Do you believe that the key factor is admixture from Baltic hunter-gatherers? Or is it genetic drift?

Of course, considering all of the f4-stats above, logic dictates that it must be relatively recent genetic drift.

Keep in mind, however, that this only applies to Balto-Slavic speaking Northeast Europeans without significant Uralian ancestry. Overall, Uralic speakers have a more complex population history, and indeed genetic differences between them and Northwest Europeans are in large part due to somewhat different ancestry proportions and also Siberian admixture.

See also...

So who's the most (indigenous) European of us all?

Warriors from at least two different populations fought in the Tollense Valley battle

I can't get the genotype data from the Burger et al. paper. The lead authors, Joachim Burger and Daniel Wegmann, aren't replying to my emails.

But they were gracious enough to release the BAM files for each of their samples, and these files can be converted to genotype data. So I've included ten of the Tollense Valley warriors (DEU_Tollense_BA) in the Global25 datasheets (see here).

The claim in the paper that these warriors "represent an unstructured population" is absolutely false and extremely naive.

Below are a couple of Principal Component Analysis (PCA) plots produced with Vahaduo Global25 views. The samples are labeled according to their Y-chromosome haplogroups. To see interactive versions of the same plots, paste the Global25 coordinates from the text file here into the relevant fields here.

These warriors are not a single unstructured population, because they cover too much ground in the above plots for that to be possible. It's clear to me that they represent at least two different groups from Central Europe and surrounds.

Of course, this would be a lot easier to work out if Burger et al. cared to supply more information about each of the warriors, such as their attire, weapons, circumstances of death, and so on. It's a complete mystery to me why this wasn't included in the paper, and the authors are refusing to talk to me, so it's unlikely that I'll ever be able to get it from them.

In the absence of such crucial archeological and anthropological data, I don't want to speculate too much, and get overly creative, but here are a couple of possible scenarios to explain the ancient DNA results:

- this may have been a battle between two Central European armies, one rich in Y-haplogroup R1b and the other rich in Y-haplogroup I2a, as well as their allies or hired help, including warriors from Eastern Europe belonging to Y-haplogroup R1a

- or perhaps it was an invasion from the east by warriors rich in Y-haplogroup R1a, and it was a success, with the local armies, rich in Y-haplogroups R1b and I2a, losing the battle and suffering most of the casualties.

I'm sure that one day someone will attempt to undertake a decent multidisciplinary study of this epic battle, and we'll at least have a rough idea about what happened. Or not.

Citation...

Burger et al., Low Prevalence of Lactase Persistence in Bronze Age Europe Indicates Ongoing Strong Selection over the Last 3,000 Years, Current Biology, Available online 3 September 2020, https://doi.org/10.1016/j.cub.2020.08.033

See also...

Genetic and linguistic structure across space and time in Northern Europe

Low prevalence of lactase persistence in Bronze Age Europe (Burger et al. 2020)

Over at Current Biology at this LINK. Unfortunately, this is the long-awaited Tollense Valley battle paper. Despite the obvious presence of some very interesting genetic substructures among the Tollense Valley warriors (see here), the authors have the audacity to claim that these individuals represent a "single unstructured Central/Northern European population".

One of the warriors, labeled WEZ56, belongs to Y-haplogroup R1a and shows an exceedingly Balto-Slavic-like genome-wide genetic structure. But none of this is even mentioned in passing in the paper. Indeed, according to Burger at al., WEZ56 is best classified as belonging to R1, even though the R1a classification is quite secure based on the raw data that the authors posted online.

Be extremely wary of what you read in this paper, and anything else that these scientists have published in the past and will publish in the future. Below is the paper summary:

Lactase persistence (LP), the continued expression of lactase into adulthood, is the most strongly selected single gene trait over the last 10,000 years in multiple human populations. It has been posited that the primary allele causing LP among Eurasians, rs4988235-A [1], only rose to appreciable frequencies during the Bronze and Iron Ages [2, 3], long after humans started consuming milk from domesticated animals. This rapid rise has been attributed to an influx of people from the Pontic-Caspian steppe that began around 5,000 years ago [4, 5]. We investigate the spatiotemporal spread of LP through an analysis of 14 warriors from the Tollense Bronze Age battlefield in northern Germany (∼3,200 before present, BP), the oldest large-scale conflict site north of the Alps. Genetic data indicate that these individuals represent a single unstructured Central/Northern European population. We complemented these data with genotypes of 18 individuals from the Bronze Age site Mokrin in Serbia (∼4,100 to ∼3,700 BP) and 37 individuals from Eastern Europe and the Pontic-Caspian Steppe region, predating both Bronze Age sites (∼5,980 to ∼3,980 BP). We infer low LP in all three regions, i.e., in northern Germany and South-eastern and Eastern Europe, suggesting that the surge of rs4988235 in Central and Northern Europe was unlikely caused by Steppe expansions. We estimate a selection coefficient of 0.06 and conclude that the selection was ongoing in various parts of Europe over the last 3,000 years.

Burger et al., Low Prevalence of Lactase Persistence in Bronze Age Europe Indicates Ongoing Strong Selection over the Last 3,000 Years, Current Biology, Available online 3 September 2020, https://doi.org/10.1016/j.cub.2020.08.033

See also...

Warriors from at least two different populations fought in the Tollense Valley battle

The oldest R1a to date

My popular map of the oldest instances of Y-haplogroup R1a in the ancient DNA record has a new entry: PES001 from the recent Saag et al. preprint. PES001 comes from a burial site in what is now northwestern Russia and is dated to a whopping 10785–10626 calBCE.

Indeed, I'm not aware of any R1a samples older than PES001 among the treasure trove of thousands of ancient samples waiting to be published. So it's likely that this individual will remain the oldest member of our R1a clan for some years to come.

See also...

Y-haplogroup R1a and mental health

Like three peas in a pod

The mystery of the Sintashta people

Fatyanovo males were rich in Y-haplogroup R1a-Z93 (Saag et al. 2020 preprint)

I'd say that thanks to this preprint we're now a lot closer to solving the mystery of the Sintashta people. Over at bioRxiv at this LINK. From the preprint:

Transition from the Stone to the Bronze Age in Central and Western Europe was a period of major population movements originating from the Ponto-Caspian Steppe. Here, we report new genome-wide sequence data from 28 individuals from the territory north of this source area - from the under-studied Western part of present-day Russia, including Stone Age hunter-gatherers (10,800-4,250 cal BC) and Bronze Age farmers from the Corded Ware complex called Fatyanovo Culture (2,900-2,050 cal BC). We show that Eastern hunter-gatherer ancestry was present in Northwestern Russia already from around 10,000 BC. Furthermore, we see a clear change in ancestry with the arrival of farming - the Fatyanovo Culture individuals were genetically similar to other Corded Ware cultures, carrying a mixture of Steppe and European early farmer ancestry and thus likely originating from a fast migration towards the northeast from somewhere in the vicinity of modern-day Ukraine, which is the closest area where these ancestries coexisted from around 3,000 BC.

...

Interestingly, in all individuals for which the chrY hg could be determined with more depth (n=6), it was R1a2-Z93 (Table 1, Supplementary Data 2), a lineage now spread in Central and South Asia, rather than the R1a1-Z283 lineage that is common in Europe [38,39].

Saag et al., Genetic ancestry changes in Stone to Bronze Age transition in the East European plain, BioRxiv, Posted July 03, 2020, doi: https://doi.org/10.1101/2020.07.02.184507

See also...

Like three peas in a pod

Like three peas in a pod

One of the most interesting questions still waiting to be answered by ancient DNA is where exactly did the ancestors of the present-day European and South Asian bearers of Y-haplogroup R1a part their ways? Indeed, the answer to this question is likely to be informative about the place and time of the split between the Balto-Slavic and Indo-Iranian language families.

I was doing some reading today and discovered that the peoples associated with the Bronze Age Fatyanovo-Balanovo and Unetice archeological cultures shared strikingly similar metalwork, despite being separated by well over two thousand kilometers of forest and steppe. Apparently, this similarity is especially pronounced in the metalwork of the Unetice culture from what is now Slovakia (see Ancient Metallurgy in the USSR: The Early Metal Age, page 136).

S11953 is currently the only sample from Slovakia associated with the Unetice culture (Sirak et al. 2020). There are no Fatyanovo-Balanovo samples available yet. However, as far as I can tell, I0432 from Samara, Russia, should be a decent stand in (Mathieson et al. 2015).

Of course, both S11953 and I0432 belong to Y-haplogroup R1a. Moreover, S11953 belongs to a typically Balto-Slavic subclade of R1a, while I0432 belongs to a closely related subclade that is dominant nowadays among the Indo-Iranian speakers of Asia.

S11953 is younger than I0432, but this doesn't necessarily mean that his ancestors arrived in East Central Europe from deep in Russia during the Bronze Age. Indeed, the opposite is more likely to be true. That is, I0432 is probably the recent decedent of migrants from somewhere near the North Carpathians, because he shows elevated European Neolithic farmer ancestry compared to earlier ancients from the Samara region (see here).

Below is a Principal Component Analysis (PCA) showing how S11953 and I0432 compare to each other in the context of ancient West Eurasian genetic variation. Obviously, they're sitting in the same part of the plot, which suggests that they harbor very similar ratios of ancient genetic components and probably share relatively recent ancestry. The relevant PCA datasheet is available here.

I've also highlighted myself, Davidski, on the plot. That's because I share the same Balto-Slavic-specific subclade of R1a with S11953 and, in terms of overall ancestry, I'm similar to both S11953 and I0432. Moreover, I'm the speaker of Polish, which is a Balto-Slavic language. What are the chances that we're dealing here with a remarkable string of coincidences? Indeed, was the North Carpathian region perhaps the homeland of the language ancestral to both Balto-Slavic and Indo-Iranian?

However, please note that there's nothing unusual or remarkable about my ancestry. The vast majority of people of Central, Eastern and Northern European origin - that is, mostly the speakers of Balto-Slavic, Germanic and Celtic languages - would also land in this part of the plot.

See also...

On the doorstep of India

Y-haplogroup R1a and mental health

The mystery of the Sintashta people

An early Mitanni?

I've updated my Global25 datasheets with most of the ancients from the new Skourtanioti et al. paper. Here's a Principal Component Analysis (PCA) based on the data. It was produced with the Vahaduo PCA tools freely available here and the text file here.

Note that one of the Bronze Age females from Alalakh, labeled ALA019, appears to have ancestry from Turan and the Eurasian steppe. She may well have been a Mitanni of Indo-Aryan origin.

Interestingly, a Copper Age male from Arslantepe, ART038, belongs to Y-haplogroup R1b1a2 aka R1b-V1636. This is an unusual find, because R1b hasn't yet been reported in any Copper Age or earlier samples from outside of Europe and the Eurasian steppe.

As far as I can tell, this individual doesn't harbor any genome-wide ancestry from north of the Caucasus. However, R1b-V1636 is a rare lineage that is first attested in Eneolithic samples from the North Caucasus Piedmont steppe, so ART038's Y-chromosome might be the first evidence of the presence of steppe ancestry in Copper Age Anatolia.

I've also added most of the ancients from the new Agranat-Tamir et al. paper to the Gobal25 datasheets. The PCA below is based on the text file available here.

The Megiddo samples include a trio of interesting outliers dated to 1600-1500 BCE with significant ancestry from the steppe. One of these individuals is a male, I2189, who belongs to Y-haplogroup R and probably R1a. So he might also be of Indo-Aryan origin.

Another Megiddo male, S10768, belongs to R1b-M269 and probably shows a few per cent of steppe ancestry. I've already discussed how R1b and steppe ancestry may have ended up in the Bronze Age Near East in a couple of my previous posts:

R1b-M269 in the Bronze Age Levant

How did steppe ancestry spread into the Biblical-era Levant?

R-V1636: Eneolithic steppe > Kura-Araxes?

Big deal of 2019: ancient DNA confirms the link between Y-haplogroup N and Uralic expansions

The academic consensus is that Indo-European languages first spread into the Baltic region from the Eastern European steppes along with the Corded Ware culture (CWC) and its people during the Late Neolithic, well before the expansion of Uralic speakers into Fennoscandia and surrounds, probably from somewhere around the Ural Mountains.

On the other hand, the views that the Uralic language family is native to Northern Europe and/or closely associated with the CWC are fringe theories usually espoused by people not familiar with the topic or, unfortunately it has to be said, mentally unstable trolls.

The likely close relationship between the CWC expansion and the early spread of Indo-European languages was discussed in several papers in recent years (for instance, see here). This year, we saw the first ancient DNA paper focusing on the transition from the Bronze Age to the Iron Age in the East Baltic, including the likely first arrival of Uralic speech in what is now Estonia.

Published in Current Biology courtesy of Saag et al., the paper showed that the genetic structure of present-day East Baltic populations largely formed in the Iron Age (see here). It was during this time, the authors revealed, that the region experienced a sudden influx of Y-chromosome haplogroup N, which is today common in many Uralic speaking populations and often referred to as a Proto-Uralic marker. Little wonder then that Saag et al. linked this genetic shift in the East Baltic to the westward migrations of early Uralic speakers.

The table below, based on data from the Saag et al. paper, surely doesn't leave much to the imagination about what happened.

Unfortunately, I have to say that the genome-wide analysis in the paper was less informative than it could have been. The authors focused their attention on rather broad genetic components, and, as a result, missed an interesting fine scale distinction between their Bronze Age and Iron Age samples. The spatial maps below, based on my Global25 data for most of the ancients from Saag et al., show what I mean. The hotter the color the higher the genetic similarity between them and present-day West Eurasian populations.

Note that the Bronze Age (Baltic_EST_BA) samples are most similar to the Baltic-speaking, and thus also Indo-European-speaking, Latvians and Lithuanians, rather than the Uralic-speaking Estonians, even though they're from burial sites in Estonia. On the other hand, the Iron Age (Baltic_EST_IA) samples show strong similarity to a wider range of populations, including Estonians and many other Uralic-speaking groups.

See also...

It was always going to be this way

Fresh off the sledge

More on the association between Uralic expansions and Y-haplogroup N

Y-haplogroup R1a and mental health

I've updated my map of pre-Corded Ware culture R1a samples with a couple of new entries from Central and South Asia (the original is still here). However, before any of you get overly excited, please note that these samples aren't older than the Corded Ware culture. The reason I added them to my map is to counter the ongoing absurd claims online that South Asian R1a isn't derived from European R1a.

Just in case the map can't be viewed in all of its glory in some devices, here's what the fine print says:

The oldest example of R1a in ancient DNA from Central Asia is dated to 2132-1940 calBCE (ID I3770, Narasimhan 2019). Moreover, this sequence is closely related to much older R1a samples from Central, Eastern and Northern Europe, and phylogenetically nested within their diversity. Thus, it must surely represent a population expansion from Europe to Central Asia. Indeed, it's also associated with the Bronze Age Andronovo archeological culture, which is usually seen as an offshoot of the Corded Ware culture (CWC) of Late Neolithic Europe. The vast majority of present-day R1a lineages in Central Asia are closely related to that of I3770, and so must also ultimately derive from Europe.

The oldest instance of R1a in ancient DNA from South Asia is dated to just 1044-922 calBCE (ID I12457, Narasimhan 2019). This sequence, as well as the vast majority of present-day South Asian R1a lineages, are closely related to much older R1a samples from Central, Eastern and Northern Europe, and phylogenetically nested within their diversity. Thus, they must surely represent a population expansion from Europe to South Asia via Central Asia, in all likelihood during the Bronze Age. Even if R1a existed in South Asia before the Bronze Age, which is extremely unlikely, because it's found in samples from indigenous European hunter-gatherers, the vast majority of present-day R1a lineages in South Asia must be ultimately from Europe.

The idea that most, if not all, South Asian R1a is derived from European R1a seriously scares a lot of people. This is obvious in many online discussions on the topic. I suspect they're so frightened by it because, in their minds, it has the potential to encourage discrimination and even racism, perhaps by re-defining the colonization of much of the world by European nations in the recent past as the natural order of things?

In any case, clearly we're dealing with some sort of mass phobia here. I've got advice for those of you suffering from this problem: if you're honestly worried that the geographic provenance and expansion history of some Y-haplogroup is going to negatively impact on your life in any meaningful way, then it's time to find yourself a quality mental health professional. All the best with that.

See also...

The mystery of the Sintashta people

The Poltavka outlier

Yamnaya isn't from Iran just like R1a isn't from India

The PIE homeland controversy: August 2019 status report

Archeologist David Anthony has a new paper on the Indo-European homeland debate titled Archaeology, Genetics, and Language in the Steppes: A Comment on Bomhard. It's part of a series of articles dealing with Allan R. Bomhard's "Caucasian substrate hypothesis" in the latest edition of The Journal of Indo-European Studies. It's also available, without any restrictions, here.

Any thoughts? Feel free to share them in the comments below. Admittedly, I found this part somewhat puzzling (emphasis is mine):

It was the faint trace of WHG, perhaps 3% of whole Yamnaya genomes, that identified this admixture as coming from Europe, not the Caucasus, according to Wang et al. (2018). Colleagues in David Reich’s lab commented that this small fraction of WHG ancestry could have come from many different geographic places and populations.

I think that's highly optimistic. It really should be obvious by now thanks to archeological and ancient genomic data, including both uniparental and genome-wide variants, that the Yamnaya people were practically entirely derived from Eneolithic populations native to the Pontic-Caspian (PC) steppe. So, in all likelihood, this was also the source of their minor WHG ancestry.

Indeed, they clearly weren't some mishmash of geographically, culturally and genetically disparate groups that had just arrived in Eastern Europe, but the direct descendants of closely related and already significantly Yamnaya-like peoples associated with long-standing PC steppe archeological cultures such as Khvalynsk and Sredny Stog. I discussed this earlier this year, soon after the Wang et al. paper was published:

On Maykop ancestry in Yamnaya

I hope I'm wrong, but I get the feeling that the scientists at the Reich Lab are finding this difficult to accept, because it doesn't gel with their theory that archaic Proto-Indo-European (PIE) wasn't spoken on the PC steppe, but rather south of the Caucasus, and that late or rather nuclear PIE was introduced into the PC steppe by migrants from the Maykop culture who were somehow involved in the formation of the Yamnaya horizon.

Inexplicably, after citing Wang et al. on multiple occasions and arguing against any significant gene flow between Maykop and Yamnaya groups, Anthony fails to mention Steppe Maykop. But the Steppe Maykop people are an awesome argument against the idea that there was anything more than occasional mating between the Maykop and Yamnaya populations, because they were wedged between them, and yet clearly distinct from both, with a surprisingly high ratio of West Siberian forager-related ancestry (see here and here).

Despite all the talk lately about the potential cultural, linguistic and genetic ties between Maykop and Yamnaya, including claims that the latter possibly acquired its wagons from the former, my view is that the Steppe Maykop and Yamnaya wagon drivers may have competed with each other and eventually clashed in a big way. Indeed, take a look at what happens after Yamnaya burials rather suddenly replace those of Steppe Maykop just north of the Caucasus around 3,000 BCE.

Yamnaya_RUS_Caucasus
RUS_Progress_En_PG2001 0.808±0.058
RUS_Steppe_Maykop 0.000
UKR_Sredny_Stog_II_En_I6561 0.192±0.058
chisq 13.859
tail prob 0.383882
Full output

Yep, total population replacement with no significant gene flow between the two groups. Apparently, as far as I can tell, there's not even a hint that a few Steppe Maykop stragglers were incorporated into the ranks of the newcomers. Where did they go? Hard to say for now. Maybe they ran for the hills nearby?

Intriguingly, Anthony reveals a few details about new samples from three different Eneolithic steppe burial sites associated with the Khvalynsk culture:

The Reich lab now has whole-genome aDNA data from more than 30 individuals from three Eneolithic cemeteries in the Volga steppes between the cities of Saratov and Samara (Khlopkov Bugor, Khvalynsk, and Ekaterinovka), all dated around the middle of the fifth millennium BC.

...

Most of the males belonged to Y-chromosome haplogroup R1b1a, like almost all Yamnaya males, but Khvalynsk also had some minority Y-chromosome haplogroups (R1a, Q1a, J, I2a2) that do not appear or appear only rarely (I2a2) in Yamnaya graves.

As far as I can tell, he suggests that they'll be published in the forthcoming Narasimhan et al. paper. If so, it sounds like the paper will have many more ancient samples than its early preprint that was posted at bioRxiv last year.

For me the really fascinating thing in regards to these new samples is how scarce Y-haplogroup R1a appears to have been everywhere before the expansion by the putative Indo-European-speaking steppe ancestors of the Corded Ware culture (CWC) people. It's basically always outnumbered by other haplogroups wherever it's found prior to about 3,000 BCE, even on the PC steppe. But then, suddenly, its R1a-M417 subclade goes BOOM! And that's why I call it...

The beast among Y-haplogroups

At this stage, I'm not sure how to interpret the presence of Y-haplogroup J in the Khvalynsk population. It may or may not be important to the PIE homeland debate. Keep in mind that J is present in two foragers from Karelia and Popovo, northern Russia, dated to the Mesolithic period and with no obvious foreign ancestry. So it need not have arrived north of the Caspian as late as the Eneolithic with migrants rich in southern ancestry from the Caucasus or what is now Iran. In other words, for the time being, the steppe PIE homeland theory appears safe.

Update 20/12/2019: A note on Steppe Maykop

See also...

Is Yamnaya overrated?

The PIE homeland controversy: January 2019 status report

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

They mixed up Huns with Tocharians

I don't yet have the genomes from the recent Ning et al. paper on the Iron Age nomads from the Shirenzigou site in the eastern Tian Shan. But I do have most of the previously published data featured in the paper, including the Damgaard et al. 2018 Hun and Saka samples from the western Tian Shan.

After reading the Ning et al. paper between the lines and running a few analyses of my own, it's clear to me that most of the supposedly Tocharian-related Shirenzigou individuals actually share a very close relationship with the Tian Shan Huns, and indeed may have been their ancestors.

For instance, Ning et al. found that a large part of the ancestry of the Shirenzigou ancients could be modeled with the Tian Shan Huns, which was an anachronistic approach because the former are older than the latter. They also found that Ulchi-related ancestry was a key part of the genetic structure of eight out of the ten Shirenzigou individuals, and this likewise appears to be an important part of the genetic structure of the Tian Shan Huns.

Note the strong statistical fits in the Global25/nMonte and qpAdm mixture models below, respectively, which characterize these Huns as a two-way mixture between the Ulchi and the earlier Tian Shan Saka. And keep in mind that the Saka also harbor significant Ulchi-related ancestry.

Hun_Tian_Shan
Saka_Tian_Shan,92
Ulchi,8
distance%=1.2553

Hun_Tian_Shan
Saka_Tian_Shan 0.928±0.009
Ulchi 0.072±0.009
chisq 4.409
tail prob 0.992464
Full output

Moreover, the Shirenzigou males belong to Y-haplogroups Q1a and R1b (two instances of each), and they share the latter with one of the Tian Shan Huns. Judging by the data from the relevant BAM files, it's also possible that the Shirenzigou males share a very rare subclade of R1b with the Hun, defined by the PH155 mutation (see here). The Y-haplogroup assignments for the other Tian Shan Huns end at R and R1, but that's almost certainly due to missing data.

On the other hand, two Tian Shan Sakas belong to Y-haplogroup R1a but none to R1b, which fits with the pattern from currently available ancient DNA that R1a was more common than R1b in Saka-related groups, such as the Scythians and Sarmatians (see here).

This is all very interesting, because the Huns replaced the Saka in the western Tian Shan, and, considering their R1b and excess Ulchi-related ancestry, very likely moved into the region from the direction of Shirenzigou. Indeed, in my opinion a strong argument can now be made that the Iron Age population from the Shirenzigou region took part in the formation of the Hunnic confederacy.

So where does that leave the theory presented by Ning et al. that the Shirenzigou ancients may have been closely related, and perhaps even ancestral, to the Tocharians, simply because they packed a lot of Yamnaya-related and possibly proto-Tocharian Afanasievo ancestry, and were living close to the Tarim Basin, where Tocharian languages were subsequently first attested?

I'm not sure, but I now find it difficult to reconcile this theory with the fact that they were closely related, and probably ancestral, to the Tian Shan Huns. As far as I'm aware, Huns cannot be linked to Tocharians in any meaningful way.

Of course it's possible that different Afanasievo-derived groups were living in the Tarim Basin and surrounds, and, as some merged with new populations pushing into the region from the east and adopted non-Indo-European languages, others retained their Tocharian speech and eventually split into communities speaking Tocharian A, B and apparently also C (see here).

But this has to be demonstrated directly with ancient DNA from archeological sites where Tocharian languages were attested. Till then, I'll keep thinking that Ning et al. wrote a paper about Tocharians that really should've been a paper about Huns.

Here's a famous wall painting of Tocharian princes from the cave of the sixteen sword-bearers in the Tarim Basin, dated to 432–538 AD. They don't look like guys with a lot of Ulchi-related admixture to me, but I might be wrong. Feel free to let me know what you think in the comments below.

Update 08/17/2019: The Shirenzigou nomads are now in my dataset. Below are a few successful and not so successful qpAdm mixture models for them. Note that I tried to use a wide range of relevant "right pops", but also retain a lot of markers, specifically to be able to discriminate between different types of steppe and steppe-derived sources of gene flow (refer to the full output). Admittedly, the Shirenzigou nomads can be modeled with Afanasievo-related ancestry, but...

CHN_Shirenzigou_IA
KAZ_Botai 0.161±0.023
KAZ_Wusun 0.490±0.023
NPL_Mebrak_2125BP 0.349±0.019
chisq 5.793
tail prob 0.926172
Full output

CHN_Shirenzigou_IA
KAZ_Botai 0.143±0.022
NPL_Mebrak_2125BP 0.295±0.019
Saka_Tian_Shan 0.562±0.024
chisq 6.796
tail prob 0.870794
Full output

CHN_Shirenzigou_IA
KAZ_Botai 0.185±0.023
NPL_Mebrak_2125BP 0.428±0.021
RUS_Sintashta_MLBA 0.270±0.026
TJK_Sarazm_En 0.117±0.027
chisq 11.351
tail prob 0.414345
Full output

CHN_Shirenzigou_IA
KAZ_Botai 0.032±0.027
KAZ_Zevakinskiy_LBA 0.567±0.025
NPL_Mebrak_2125BP 0.401±0.019
chisq 15.157
tail prob 0.232961
Full output

CHN_Shirenzigou_IA
NPL_Mebrak_2125BP 0.452±0.031
RUS_Afanasievo 0.435±0.025
RUS_Okunevo_BA 0.114±0.049
chisq 19.808
tail prob 0.0708003
Full output

CHN_Shirenzigou_IA
NPL_Mebrak_2125BP 0.409±0.031
RUS_Okunevo_BA 0.173±0.050
Yamnaya_RUS_Caucasus 0.418±0.026
chisq 20.453
tail prob 0.0589872
Full output

CHN_Shirenzigou_IA
NPL_Mebrak_2125BP 0.464±0.033
RUS_Okunevo_BA 0.104±0.053
Yamnaya_RUS_Samara 0.432±0.027
chisq 27.189
tail prob 0.0072566
Full output

Both the Wusun and Saka are generally accepted to have been the speakers of Indo-Iranian languages. So it's possible that the Shirenzigou nomads were Indo-Iranian speakers too, or at least derived from such peoples.

Surprisingly, NPL_Mebrak_2125BP was the key to obtaining the best statistical fits. This is a trio of samples, roughly contemporaneous with the Shirenzigou nomads, from a burial site high up in the Himalayas in what is now Nepal (see here).

To be honest, I'm not quite sure why the Himalayan ancients work so well in my models. Perhaps they're just a really good proxy for an Iron Age population from the northern part of the Tibetan Plateau? By the way, most of the Shirenzigou nomads made it into the latest Global25 datasheets (see here).

See also...

Almost everything you ever wanted to know about the Xiaohe-Gumugou cemeteries

The mystery of the Sintashta people

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

Fresh off the sledge

As things stand, the closest individual to a Proto-Uralic speaker in the ancient DNA record is arguably 0LS10 from an Iron Age tarand grave in what is now Estonia. I say that because:

- isotopic data suggest that 0LS10 wasn't born where he died, and considering his elevated Siberian ancestry relative to earlier and most contemporaneous Baltic ancients, he was very likely a migrant to the Baltic region from the east

- the tarand grave tradition appears to be specifically a Finnic (west Uralic) phenomenon that probably spread from the Volga-Oka region, which is just west of where most people place the Proto-Uralic homeland

- 0LS10 belongs to Y-chromosome haplogroup N-L1026, a paternal marker that is especially closely associated with Uralic-speaking populations and probably only appeared in the East Baltic region during the transition from the Bronze Age to the Iron Age

You can find more background info about 0LS10 and other relevant samples in Saag et al. 2019 (see here). This is where he sits in my Principal Component Analyses (PCA) focusing on fine scale Northern European genetic diversity. The relevant datasheets are available here and here, respectively.

Note that 0LS10 doesn't cluster strongly with any ancient or modern populations. To investigate this in more detail I ran a series of two-way qpAdm analyses, testing tens of ancient individuals and populations as potential admixture sources. These two models stood out above the rest in terms of their statistical fits, chronology and overall plausibility.

Baltic_EST_IA_0LS10
Baltic_EST_BA 0.826±0.045
RUS_Sintashta_MLBA_o1 0.174±0.045
chisq 12.527
tail prob 0.564048
Full output

Baltic_EST_IA_0LS10
Baltic_EST_BA 0.683±0.102
RUS_Mezhovskaya 0.317±0.102
chisq 13.811
tail prob 0.463864
Full output

Please note that RUS_Sintashta_MLBA_o1 isn't representative of the Sintashta culture population as a whole. It's a group of the most extreme genetic outliers among the Sintashta samples, and they may or may not have been Uralic speakers (see here). Interestingly, the Mezhovskaya culture population is generally associated with the Ugric branch of the Uralic language family.

I was also able to closely replicate these results with the Global25/nMonte method; down to almost one per cent. However, the statistical fits (distances) are poor, probably because the reference populations aren't the real mixture sources. This is in line with the fact that their Y-haplogroups are Q1a, R1a and R1b, rather than any type of N.

Baltic_EST_IA:0LS10
Baltic_EST_BA,83.8
RUS_Sintashta_MLBA_o1,16.2
distance%=4.7955

Baltic_EST_IA:0LS10
Baltic_EST_BA,69.8
RUS_Mezhovskaya,30.2
distance%=3.5783

I do realize that two Bronze Age samples from Bolshoy Oleni Ostrov, Kola Peninsula, belong to N-L1026, but adding them to my mixture models doesn't help. Little wonder, because the Kola Peninsula lies within the Arctic Circle, and I'm pretty sure that 0LS10 and his N-L1026 came from somewhere just north of the mixture cline marked on the map below. Unfortunately, I can't test this directly yet due to the scarcity of ancient samples from this region.

See also...

It was always going to be this way

On the association between Uralic expansions and Y-haplogroup N

Uralic-specific genome-wide ancestry did make a signifcant impact in the East Baltic

Uralic-specific genome-wide ancestry did make a signifcant impact in the East Baltic

I've started analyzing the ancient genotype data from the recent Saag et al. paper on the expansion of Uralic languages and associated spread of Siberian ancestry into the East Baltic region. The paper is freely available here and the data are here.

I really like the paper, but I don't agree with the authors' claim that the appearance of Y-chromosome haplogroup N in what is now Estonia and surrounds during the Iron Age is "not matched by a clear shift in autosomal profiles". In my opinion it certainly is, and, as one would expect, it's a shift towards a genetic profile typical of western Uralic speakers.

I'd say that the easiest way to find this signal is with a Principal Component Analysis (PCA) focusing on fine scale genetic substructures within Northern Europe, like the one below. The relevant datasheet is available here.

Note that the East Baltic Iron Age samples, all from burial sites in what is now Estonia, appear to be peeling away from their Bronze Age predecessors and overlapping strongly with present-day Estonians, who are Uralic speakers. Indeed, the PCA suggests to me that the formation of the greater part of the present-day Estonian gene pool took place in the East Baltic during the transition from the Bronze Age to the Iron Age. That is, when Uralic languages are generally accepted to have arrived in the region from near the Ural Mountains in the east.

I was also able to closely replicate these outcomes with my Global25 data using the method described here. However, in this effort, present-day Estonians are clearly more western than the Estonian Iron Age samples (EST_IA), which might be due to the presence of low level Germanic ancestry in Estonia dating to the medieval period. The relevant datasheet is available here.

Interestingly, the Estonian Bronze Age samples (EST_BA) come from stone-cist graves which are widely hypothesized to have been introduced to the East Baltic from the Nordic Bronze Age civilization. I even recall reading a paper on the topic which claimed that the remains buried in such graves were those of Proto-Germanic-speaking Scandinavian migrants. Well, I haven't had a chance to study these samples in any great detail yet, but considering that in both of the PCA above they're overlapping strongly with Latvian Bronze Age samples (LVA_BA) and sitting far away from the nearest Scandinavians, I'd say they're probably of local stock from way back.

See also...

It was always going to be this way

On the association between Uralic expansions and Y-haplogroup N

Inferring the linguistic affinity of long dead and non-literate peoples: a multidisciplinary approach

It was always going to be this way

The native peoples of the East Baltic - Estonians, Latvians and Lithuanians - are genetically alike and their paternal gene pools are dominated by the same two Y-chromosome haplogroups: R1a and N3a.

Linguistically, however, Estonians are a world apart from Latvians and Lithuanians. That's because the Estonian language belongs to the Uralic language family, which has an obvious North Eurasian character. On the other hand, Latvian and Lithuanian are both classified as Indo-European languages, along with the vast majority of other European languages.

The Uralic and Indo-European language families may or may not descend from the same ancestral tongue, but even if they do, their relationship is very distant.

So how is it that Estonians came to speak a Uralic language? As far back as I can remember, the basic explanation accepted by most people was that Uralic speech arrived in what is now Estonia and neighboring Finland during the Bronze Age with migrants, or perhaps invaders, rich in N3a from somewhere around the Ural Mountains. Conversely, Latvians and Lithuanians were generally assumed to have retained the Indo-European speech of their R1a-rich forefathers from the Pontic-Caspian steppe, who colonized much of Eastern Europe north of the steppe during the Late Neolithic.

Ancient DNA has now uncannily corroborated these theories (for instance, see Mittnik et al. 2018 and, published today, Saag et al. 2019). All it took was a handful of samples from a few relevant sites. I think that's awesome; I love it when sensible, long-standing hypotheses are validated by cutting edge science.

I'll have a lot more to say about the spread of Uralic languages and Uralian genes to the East Baltic when I get my hands on the genotype data from the new Saag et al. paper. I also have a post coming soon about the Nordic Bronze Age. Stay tuned.

Update 10/05/2019: Uralic-specific genome-wide ancestry did make a signifcant impact in the East Baltic

See also...

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

Corded Ware people =/= Proto-Uralics (Tambets et al. 2018)

Inferring the linguistic affinity of long dead and non-literate peoples: a multidisciplinary approach

Map of pre-Corded Ware culture (>2900 BCE) instances of Y-haplogroup R1a (updated)

Below is a map showing the global distribution of Y-chromosome haplogroup R1a prior to the expansions of the R1a-rich Corded Ware culture (CWC) people and their descendants across Europe and Asia from around 2900 BCE. I'll be updating this map regularly and using it to help me narrow down the options for the place of origin of R1a, and also to counter the misinformation about this topic that has appeared in print and online over the years, including in many scientific publications and popular websites such as Wikipedia.

Incredibly, as far as I know, there are just six reliably called instances of R1a in the now ample Eurasian ancient DNA record dating to the pre-CWC period. To put this into perspective, consider that R1a is today the most common Y-haplogroup in much of Europe and Asia. How did that happen I wonder? However, please note that I chose to base the map only on samples sequenced with the capture and shotgun methods, rather than the PCR method, which is susceptible to producing contaminated results and no longer used in major ancient DNA studies.

See also...

Y-haplogroup R1a and mental health

The Poltavka outlier

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

Corded Ware people =/= Proto-Uralics (Tambets et al. 2018)

A new paper on the genetic structure of Uralic-speaking populations has appeared at Genome Biology (see here). It looks to me like the prelude to a forthcoming paleogenetics paper on the same topic that was discussed in the Estonian media recently (see here). Although not exactly ground breaking (because it basically argues what I've been saying at this blog for years, like here), it's a very nice effort all round and must be read by anyone with an interest in this topic. From the paper, emphasis is mine:

Background The genetic origins of Uralic speakers from across a vast territory in the temperate zone of North Eurasia have remained elusive. Previous studies have shown contrasting proportions of Eastern and Western Eurasian ancestry in their mitochondrial and Y chromosomal gene pools. While the maternal lineages reflect by and large the geographic background of a given Uralic-speaking population, the frequency of Y chromosomes of Eastern Eurasian origin is distinctively high among European Uralic speakers. The autosomal variation of Uralic speakers, however, has not yet been studied comprehensively.

Results: Here, we present a genome-wide analysis of 15 Uralic-speaking populations which cover all main groups of the linguistic family. We show that contemporary Uralic speakers are genetically very similar to their local geographical neighbours. However, when studying relationships among geographically distant populations, we find that most of the Uralic speakers and some of their neighbours share a genetic component of possibly Siberian origin. Additionally, we show that most Uralic speakers share significantly more genomic segments identity-by-descent with each other than with geographically equidistant speakers of other languages. We find that correlated genome-wide genetic and lexical distances among Uralic speakers suggest co-dispersion of genes and languages. Yet, we do not find long-range genetic ties between Estonians and Hungarians with their linguistic sisters that would distinguish them from their non-Uralic-speaking neighbours.

Conclusions: We show that most Uralic speakers share a distinct ancestry component of likely Siberian origin, which suggests that the spread of Uralic languages involved at least some demic component.

...

Recent aDNA studies have shown that extant European populations draw ancestry form three main migration waves during the Upper Palaeolithic, the Neolithic and Early Bronze Age [2, 3, 45]. The more detailed reconstructions concerning NE Europe up to the Corded Ware culture agree broadly with this scenario and reveal regional differences [65–67]. However, to explain the demographic history of extant NE European populations, we need to invoke a novel genetic component in Europe—the Siberian. The geographic distribution of the main part of this component is likely associated with the spread of Uralic speakers but gene flow from Siberian sources in historic and modern Uralic speakers has been more complex, as revealed also by a recent study of ancient DNA from Fennoscandia and Northwest Russia [68]. Thus, the Siberian component we introduce here is not the perfect but still the current best candidate for the genetic counterpart in the spread of Uralic languages.

Citation...

Tambets et al., Genes reveal traces of common recent demographic history for most of the Uralic-speaking populations, Genome Biology, (2018) 19:139 https://doi.org/10.1186/s13059-018-1522-1

See also...

Big deal of 2019: ancient DNA confirms the link between Y-haplogroup N and Uralic expansions