Fortunately, the current obsession with simplifying ancestry components into three or four general, atemporal groups, and the common use of the same ones across labs, make it very simple to merge data and map them.
Corded Ware ancestry
There is no doubt about the prevalent ancestry among Uralic-speaking peoples. A map isn’t needed to realize that, because ancient and modern data – like those recently summarized in Jeong et al. (2019) – prove it. But maps sure help visualize their intricate relationship better:
Edit (29/7/2019): Here is the full Steppe_MLBA ancestry map, including Steppe_MLBA (vs. Indus Periphery vs. Onge) in modern South Asian populations from Narasimhan et al. (2018), apart from the ‘Srubnaya component’ in North Eurasian populations. ‘Dummy’ variables (with 0% ancestry) have been included to the south and east of the map to avoid weird interpolations of Steppe_MLBA into Africa and East Asia.
Anatolia Neolithic ancestry
Also interesting are the patterns of non-CWC-related ancestry, in particular the apparent wedge created by expanding East Slavs, which seems to reflect the intrusion of central(-eastern) European ancestry into Finno-Permic territory.
The cline(s) between WHG, EHG, ANE, Nganasan, and Baikal HG are also simplified when some of them excluded, in this case EHG, represented thus in part by WHG, and in part by more eastern ancestries (see below).
Arctic, Tundra or Forest-steppe?
Data on Nganasan-related vs. ANE vs. Baikal HG/Ulchi-related ancestry is difficult to map properly, because both ancestry components are usually reported as mutually exclusive, when they are in fact clearly related in an ancestral cline formed by different ancient North Eurasian populations from Siberia.
When it comes to ascertaining the origin of the multiple CWC-related clines among Uralic-speaking peoples, the question is thus how to properly distinguish the proportions of WHG-, EHG-, Nganasan-, ANE or BaikalHG-related ancestral components in North Eurasia, i.e. how did each dialectal group admix with regional groups which formed part of these clines east and west of the Urals.
The truth is, one ought to test specific ancient samples for each “Siberian” ancestry found in the different Uralic dialectal groups, but the simplistic “Siberian” label somehow gets a pass in many papers (see a recent example).
Below qpAdm results with best fits for Ulchi ancestry, Afontova Gora 3 ancestry, and Nganasan ancestry, but some populations show good fits for both and with similar proportions, so selecting one necessarily simplifies the distribution of both.
A simplistic Iran Chalcolithic-related ancestry is also seen in the Altaic cline(s) which (like Corded Ware ancestry) expanded from Central Asia into Europe – apart from its historical distribution south of the Caucasus:
The first question I imagine some would like to know is: what about other models? Do they show the same results? Here is the simplistic combination of ancestry components published in Damgaard et al. (2018) for the same or similar populations:
NOTE. As you can see, their selection of EHG vs. WHG vs. Nganasan vs. Natufian vs. Clovis of is of little use, but corroborate the results from other papers, and show some interesting patterns in combination with those above.
Baikal HG ancestry
Ancient North Eurasians
Once the modern situation is clear, relevant questions are, for example, whether EHG-, WHG-, ANE, Nganasan-, and/or Baikal HG-related meta-populationsexpanded or became integrated into Uralic-speaking territories.
When did these admixture/migration events happen?
How did the ancient distribution or expansion of Palaeo-Arctic, Baikalic, and/or Altaic peoples affect the current distribution of the so-called “Siberian” ancestry, and of hg. N1a, in each specific population?
NOTE. A little excursus is necessary, because the calculated repetition of a hypothetic opposition “N1a vs. R1a” doesn’t make this dichotomy real:
There was not a single ethnolinguistic community represented by hg. R1a after the initial expansion of Eastern Corded Ware groups, or by hg. N1a-L392 after its initial expansion in Siberia:
Different subclades became incorporated in different ways into Bronze Age and Iron Age communities, most of which without an ethnolinguistic change. For example, N1a subclades became incorporated into North Eurasian populations of different languages, reaching Uralic- and Indo-European-speaking territories of north-eastern Europe during the late Iron Age, at a time when their ancestral origin or language in Siberia was impossible to ascertain. Just like the mix found among Proto-Germanic peoples (R1b, R1a, and I1)* or among Slavic peoples (I2a, E1b, R1a)*, the mix of many Uralic groups showing specific percentages of R1a, N1a, or Q subclades* reflect more or less recent admixture or acculturation events with little impact on their languages.
*other typically northern and eastern European haplogroups are also represented in early Germanic (N1a, I2, E1b, J, G2), Slavic (I1, G2, J) and Finno-Permic (I1, R1b, J) peoples.
The problem with mapping the ancestry of the available sampling of ancient populations is that we lack proper temporal and regional transects. The maps that follow include cultures roughly divided into either “Bronze Age” or “Iron Age” groups, although the difference between samples may span up to 2,000 years.
NOTE. Rough estimates for more external groups (viz. Sweden Battle Axe/Gotland_A for the NW, Srubna from the North Pontic area for the SW, Arctic/Nganasan for the NE, and Baikal EBA/”Ulchi-like” for the SE) have been included to offer a wider interpolated area using data already known.
Similar to modern populations, the selection of best fit “Siberian” ancestry between Baikal HG vs. Nganasan, both potentially ± ANE (AG3), is an oversimplification that needs to be addressed in future papers.
NOTE. The samples from Levänluhta are centuries older than those from Estonia (and Ingria), and those from Chalmny Varre are modern ones, so this region has to be read as a south-west to north-east distribution from the Iron Age to modern times.
Baikal HG-like ancestry
The fact that this Baltic N1a-VL29 branch belongs in a group together with typically Avar N1a-B197 supports the Altaic origin of the parent group, which is possibly related to the expansion of Baikalic ancestry and Iron Age nomads:
The dilution of Nganasan-like ancestry in an Arctic region featuring “Siberian” ancestry and hg. N1a-L392 at least since the Bronze Age supports the integration of hg. N1a-Z1934, sister clade of Ugric N1a-Z1936, into populations west and east of the Urals with the expansion of Uralic languages to the north into the Tundra region (see here).
The integration of N1a-Z1934 lineages into Finnic-speaking peoples after their migration to the north and east, and the displacement or acculturation of Saami from their ancestral homeland, coinciding with known genetic bottlenecks among Finns, is yet another proof of this evolution:
Similarly, WHG ancestry doesn’t seem to be related to important population movements throughout the Bronze Age, which excludes the multiple North Eurasian populations that will be found along the clines formed by WHG, EHG, ANE, Nganasan, Baikal HG ancestry as forming part of the Uralic ethnogenesis, although they may be relevant to follow later regional movements of specific populations.
It seems natural that people used to look at maps of haplogroup distribution from the 2000s, coupled with modern language distributions, and would try to interpret them in a certain way, reaching thus the wrong conclusions whose consequences are especially visible today when ancient DNA keeps contradicting them.
The evolution of each specific region and cultural group of North Eurasia is far from being clear. However, the general trend speaks clearly in favour of an ancient, Bronze Age distribution of North Eurasian ancestry and haplogroups that have decreased, diluted, or become incorporated into expanding Uralians of Corded Ware ancestry, occasionally spreading with inter-regional expansions of local groups.
Given the relatively recent push of Altaic and Indo-European languages into ancestral Uralic-speaking territories, only the ancient Corded Ware expansion remains compatible with the spread of Uralic languages into their historical distribution.
As I said 6 months ago, 2019 is a tough year to write a blog, because this was going to be a complex regional election year and therefore a time of political promises, hence tenure offers too. Now the preliminary offers have been made, elections have passed, but the timing has slightly shifted toward 2020. So I may have the time, but not really any benefit of dedicating too much effort to the blog, and a lot of potential benefit of dedicating any time to evaluable scientific work.
On the other hand, I saw some potential benefit for publishing texts with ISBNs, hence the updates to the text and the preparation of these printed copies of the books, just in case. While Spain’s accreditation agency has some hard rules for becoming a tenured professor, especially for medical associates (whose years of professional experience are almost worthless compared to published peer-reviewed papers), it is quite flexible in assessing one’s merits.
However, regional and/or autonomous entities are not, and need an official identifier and preferably printed versions to evaluate publications, such as an ISBN for books. I took thus some time about a month ago to update the texts and supplementary materials, to publish a printed copy of the books with Amazon. The first copies have arrived, and they look good.
Corrections and Additions
I have changed the names and order of the books, as I intended for the first publication – as some of you may have noticed when the linguistic book was referred to as the third volume in some parts. In the first concept I just wanted to emphasize that the linguistic work had priority over the rest. Now the whole series and the linguistic volume don’t share the same name, and I hope this added clarity is for the better, despite the linguistic volume being the third one.
I have changed the nomenclature for Uralic dialects, as I said recently. I haven’t really modified anything deeper than that, because – unlike adding new information from population genomics – this would require for me to do a thorough research of the most recent publications of Uralic comparative grammar, and I just can’t begin with that right now.
Anyway, the use of terms like Finno-Ugric or Finno-Samic is as correct now for the reconstructed forms as it was before the change in nomenclature.
The most interesting recent genetic data has come from Iberia and the Mediterranean. Lacking direct data from the Italian Peninsula (and thus from the emergence of the Etruscan and Rhaetian ethnolinguistic community), it is becoming clearer how some quite early waves of Indo-Europeans and non-Indo-Europeans expanded and shrank – at least in West Iberia, West Mediterranean, and France.
Some of the main updates to the text have been made to the sections on Finno-Ugric populations, because some interesting new genetic data (especially Y-DNA) have been published in the past months. This is especially true for Baltic Finns and for Ugric populations.
Consequently, and somehow unsurprisingly, the Balto-Slavic section has been affected by this; e.g. by the identification of Early Slavs likely with central-eastern populations dominated by (at least some subclades of) hg. I2a-L621 and E1b-V13.
I have updated some cultural borders in the prehistoric maps, and the maps with Y-DNA and mtDNA. I have also added one new version of the Early Bronze age map, to better reflect the most likely location of Indo-European languages in the Early European Bronze Age.
As those in software programming will understand, major changes in the files that are used for maps and graphics come with an increasing risk of additional errors, so I would not be surprised if some major ones would be found (I already spotted three of them). Feel free to communicate these errors in any way you see fit.
I have selected more conservative SNPs in certain controversial cases.
I have also deleted most SNP-related footnotes and replaced them with the marking of each individual tentative SNP, leaving only those footnotes that give important specific information, because:
My way of referencing tentative SNP authors did not make it clear which samples were tentative, if there were more than one.
It was probably not necessary to see four names repeated 100 times over.
Often I don’t really know if the person I have listed as author of the SNP call is the true author – unless I saw the full SNP data posted directly – or just someone who reposted the results.
Sometimes there are more than one author of SNPs for a certain sample, but I might have added just one for all.
For a centralized file to host the names of those responsible for the unofficial/tentative SNPs used in the text – and to correct them if necessary -, readers will be eventually able to use Phylogeographer‘s tool for ancient Y-DNA, for which they use (partly) the same data I compiled, adding Y-Full‘s nomenclature and references. You can see another map tool in ArcGIS.
NOTE. As I say in the text, if the final working map tool does not deliver the names, I will publish another supplementary table to the text, listing all tentative SNPs with their respective author(s).
If you are interested in ancient Y-DNA and you want to help develop comprehensive and precise maps of ancient Y-DNA and mtDNA haplogroups, you can contact Hunter Provyn at Phylogeographer.com. You can also find more about phylogeography projects at Iain McDonald’s website.
I previously used certain samples prepared by amateurs from BAM files (like Botai, Okunevo, or Hittites), and the results were obviously less than satisfactory – hence my criticism of the lack of publication of prepared files by the most famous labs, especially the Copenhagen group.
Fortunately for all of us, most published datasets are free, so we don’t have to reinvent the wheel. I criticized genetic labs for not releasing all data, so now it is time for praise, at least for one of them: thank you to all responsible at the Reich Lab for this great merged dataset, which includes samples from other labs.
NOTE. I would like to make my tiny contribution here, for beginners interested in working with these files, so I will update – whenever I have time – the “How To” sections of this blog for PCAs, PCA3d, and ADMIXTURE.
For unsupervised ADMIXTURE in the maps, a K=5 is selected based on the CV, giving a kind of visual WHG : NWAN : CHG/IN : EHG : ENA, but with Steppe ancestry “in between”. Higher K gave worse CV, which I guess depends on the many ancient and modern samples selected (and on the fact that many samples are repeated from different sources in my files, because I did not have time to filter them all individually).
I found some interesting component shared by Central European populations in K=7 to K=9 (from CEU Bell Beakers to Denmark LN to Hungarian EBA to Iberia BA, in a sort of “CEU BBC ancestry” potentially related to North-West Indo-Europeans), but still, I prefer to go for a theoretically more correct visualization instead of cherry-picking the ‘best-looking’ results.
Since I made fun of the search for “Siberian ancestry” in coloured components in Tambets et al. 2018, I have to be consistent and preferred to avoid doing the same here…
In the first publication (in January) and subsequent minor revisions until March, I trusted analyses and ancestry estimates reported by amateurs in 2018, which I used for the text adding my own interpretations. Most of them have been refuted in papers from 2019, as you probably know if you have followed this blog (see very recent examples here, here, or here), compelling me to delete or change them again, and again, and again. I don’t have experience from previous years, although the current pattern must have been evidently repeated many times over, or else we would be still talking about such previous analyses as being confirmed today…
I wanted to be one step ahead of peer-reviewed publications in the books, but I prefer now to go for something safe in the book series, rather than having one potentially interesting prediction – which may or may not be right – and ten huge mistakes that I would have helped to endlessly redistribute among my readers (online and now in print) based on some cherry-picked pairwise comparisons. This is especially true when predictions of “Steppe“- and/or “Siberian“-related ancestry have been published, which, for some reason, seem to go horribly wrong most of the time.
I am sure whole books can be written about why and how this happened (and how this is going to keep happening), based on psychology and sociology, but the reasons are irrelevant, and that would be a futile effort; like writing books about glottochronology and its intermittent popularity due to misunderstood scientist trends. The most efficient way to deal with this problem is to avoid such information altogether, because – as you can see in the current revised text – they wouldn’t really add anything essential to the content of these books, anyway.
The recent study of Estonian Late Bronze Age/Iron Age samples has shown, as expected, large genetic continuity of Corded Ware populations in the East Baltic area, where West Uralic is known to have been spoken since at least the Early Bronze Age.
The most interesting news was that, unexpectedly for many, the impact of “Siberian ancestry” (whatever that actually means) was small, slow, and gradual, with slight increases found up to the Middle Ages, compatible with multiple contact events in north-eastern Europe. Haplogroup N became prevalent among Finnic populations only through late bottlenecks, as research of modern populations have long suggested, and as ancient DNA research hinted since at least 2015.
I risked to correlate the arrival of chiefs from the south-west with the infiltration of N1c-VL29 subclades during the transition to the Iron Age, coupled with that minimal “Siberian” ancestry (see e.g. here and here). Now we know that the penetration of this non-CW ancestry started, as predicted, in the Iron Age; that it was highly variable in the few samples where it appeared, with ca. 1-4%, while most Iron Age individuals show 0%; and that it was not especially linked to individuals of N1c-Vl29 lineages.
It is also basically confirmed, based on the (ancient and Modern Swedish) N1c-L550 subclades found among Iron Age Estonians, that N1c-VL29 lineages and the so-called “Siberian” ancestry will be found simultaneously around the Baltic coastal areas, and that different lineages must have suffered later founder effects among Finns, which suggests that these alliances through exogamy brought exactly as much language change in Sweden, Lithuania, or Poland, as they did in the East Baltic region…
On the other hand, the paper has also shown a potential movement of Corded Ware-derived peoples, if the change from LBA to IA samples is meaningful; in fact, even more Corded Ware-like than Baltic and Estonian BA populations. The exact origin of that movement is difficult to pinpoint, and it may not be related to the arrival of Akozino warrior-traders from the south-east, since theirs seems to be a minor impact proper of elites in a chiefdom system around the Baltic.
Also suggesting a potential movement is the ‘southern’ shift observed in the West and East Baltic areas, likely showing the arrival of Proto-East Baltic speakers (such as the Trzciniec outlier), as we have already discussed in this blog. The unexpected increase in Corded Ware-like ancestry in the Eastern Baltic, coupled with the expected large continuity of hg. R1a-Z283 in the homeland of Balto-Finnic expansions, gives even more support to the known complex system of exogamy along the Baltic coasts, and offers another potential reason for the rise of Baltic-speaking territories in the West Baltic: elite domination.
It is nevertheless important to understand that, even among the most “genetic continuous” regions like Estonia, not a single population in Europe is heir of some ancestral, immutable people. Not in terms of haplogroups, and not in terms of admixture. Balto-Finnic speakers, however continuous they might seem (e.g. in Southern Estonians) aren’t an exception.
With the currently available tools – linguistics, archaeology, and now genetics -, I don’t think there is any argument to date to question the direct connection of the Late Proto-Uralic expansion with allEastern Corded Ware groups (i.e. Battle Axe, Fatyanovo-Balanovo, and Abashevo), and thus at least with the unifying A-horizon of Corded Ware and the bottlenecks under R1a-Z645.
NOTE. The only out-group among Corded Ware cultures is the Single Grave culture. It appears to be an early Corded Ware offshoot, reflected in their non-unitary cultural traits (distinct from later unifying waves), in their varied patrilineal clans, and in the short-lasting cultural effect in northern Europe before their complete demise under pressure of expanding Yamna/Bell Beaker peoples from the Danube. The culture’s minimal (if any) effects on succeeding peoples might be seen mostly in the (mainly phonetic) Uralic substrate found in Balto-Slavic – although this may also stem from a more eastern influence, close to the Baltic – and in the contacts of Celtic with Uralic. The huge time depth between this early hypothetic Uralic layer in northern Europe and the emergence of peoples inhabiting these territories in recorded history have no doubt been erroneously interpreted as a lack of Uralic presence in the area.
1) That connection was evident in the Yamna – CWC differences in archaeology, and especially later, with at least Fatyanovo-Balanovo and Abashevo representing the obvious replacement of the Volosovo culture before further expansions of CWC-related groups west and east of the Urals.
The mythical millennia-long continuity of Volosovo hunter-gatherers, including centuries among Corded Ware peoples, as expected lately by the Copenhagen group (and anyone who doesn’t want to question the 1960s association of Indo-European with CWC) must be rejected today in population genomics, as the recent studies of ancient and modern populations show, and as ancient DNA from the region will confirm.
2) In linguistics, the survival of Volosovo as The Uralic-speaking culture was also hardly believable. From Kallio (2015):
While we can say at least something about Uralic substrates in Northeastern Europe, non-Uralic substrates cannot at all easily be identified, because of multiple language shifts, viz. first from non-Uralic to Uralic and then from Uralic to Russian. Yet the Soviet Uralicist Boris Serebrennikov (1956, 1959) argued that there are some non-Uralic substrate toponyms in the Volga-Oka region, but his idea was never taken seriously in the west (cf. Sauvageot 1958), and it pretty soon also sank into oblivion in Russia, even though it can still occasionally pop up there in non-onomastic circles (cf. Napolskikh 1995: 18–19). However, not all the hypotheses on non-Uralic substrates in Northeastern Europe should be rejected (see e.g. Helimski 2001b).
Helimski (2001) argues for a non-Uralic topo-hydronomy in Northern Russia, whose population may have kept their languages up to the Common Era despite the Corded Ware expansion, which is in line with the survival of some non-Indo-European languages everywhere in Europe after the expansion of Yamna and its offshoots:
It should be borne in mind that these [Uralic] hydronyms reached us mainly through Northern Russian and, accordingly, with a tendency to phonetic-morphological adaptation and unification (for river names it is “natural” to be, like the word ‘river’ itself, feminine and to end in -a). Taking into account this circumstance, it may turn out to be non-useless for etymological identification of at least some of the hydronyms on the Finno-Ugric basis.
On the other hand, I wouldn’t exclude the possibility that some parts of this large geographical area were never (completely) Finno-Ugric. The population that created the most important part of the hydronymy of the Russian North could be finally pushed aside or assimilated only at the end of the 1st – beginning of the 2nd millennium AD, during the Russian colonization, retaining the memory of the White-Eyed Chude in its own memory.
NOTE. For more on this non-IE substrate in (especially West) Uralic, see e.g. Zhivlov (2015),
The same non-Uralic substrate is most likely behind most of the shared traits by Mordvinic and Balto-Finnic (see below).
3) In genetics, I don’t think the picture could get any clearer. I don’t know what “Steppe ancestry = Indo-European” proponents expected from 2019, if they expected anything at all (I haven’t seen any coherent model, proposal, or prediction for a long time now), but I doubt the recent results are compatible with any of their implied expectations.
Notice, from the PCA above, how this Baltic Late Neolithic group shows actually a shift from Sredni Stog (see PCA with Sredni Stog) towards typical Khvalynsk-Urals-related ancestry, i.e. populations from eastern European forested regions, derived from hunter-gatherer pottery groups, as I have proposed for a very long time, since the first time a Baltic LN “outlier” appeared. It’s amazing how some amateurs can find 0.1% of any Siberian outlier’s ancestry among Uralians 4,000 years later, but fail to see the direct connection here. The esoteric uses of qpAdm, I guess…
Especially noticeable is the extra WHG-like ancestry and corresponding shift, seen especially marked in late Polish CWC samples, but also in Baltic CWC and especially in one Sweden Battle Axe sample, all of them shifting apparently closer to Pitted Ware and SHG. While that may have been interpreted as an in situ admixture in Scandinavia before, the late Polish CWC samples show likely a resurgence of local populations, so we can assume that both shifts (to SHG- and EHG-like populations) of available CWC samples around the Baltic are clearly part of the WHG:EHG continuum that will be found in the eastern European sub-Neolithic cultures, from Narva to Volosovo.
This WHG-related ancestry is clearly predominant in groups with which Battle Axe peoples admixed, based on the shift towards Pitted Ware, which – I can only guess based on modern Volga Finns – is different from the shift we will see in Netted Ware, more towards the Khvalynsk-Urals cluster. This is in line with the expansion of Battle Axe eastward through coastal areas (West to East Baltic and Finland into Sweden), while Fatyanovo peoples probably emerged from a slightly different route, but also a northern one, if one is to follow archaological similarities and their chronology.
During the Iron Age, the only peoples that probably shifted strongly (based on modern populations) are West Baltic ones, getting closer to the available Late Trzciniec samples, and even closer to the Trzciniec outlier, i.e. away from the earlier Eastern Corded Ware cluster, and towards Central European groups like Czech EBA or Poland EBA, both of them clearly derived from Bell Beakers, but also admixed with (and thus shifted toward) CW-like populations.
If one looks carefully at the previous PCA on Bronze Age populations, and the next one on Iron Age clusters, it is evident that adding the Swedish LN outlier to East Baltic BA (both strongly related to Battle Axe populations) essentially gives us the continuity of East Baltic BA into the Iron Age. This cluster is continued also in two outliers from Sigtuna, a Viking town close to the Gulf of Finland, known to be an important trading site, 1,500 years later. Not much of a change around the Gulf of Finland, then:
Based on the two simplistic Uralic clines one might see described (among the many that certainly existed, from Corded Ware to different Eurasian populations), and just like BOO was for some months fashionable as “Samic”, some may be tempted to say that certain Sintashta or Srubna outliers close to the Urals mark the True Uralic™ peoples. Because, of course they do. Ghost haplogroup N and stuff. And Corded Ware never ever Uralic. Because Gimbutas, and my IE R1a grandfather.
NOTE. Funny thing here: there might be Corded Ware, Iranian, Slavic, Germanic, etc… outliers or out-groups, and they might form the widest genetic clusters ever seen, but they are all of one language, because archaeology and linguistics; however, one “outlier” (also, put your own definition of “outlier” here, let’s say 1% of whatever, and strontium isotope potentially from 100 km away) ca. 600 BC in the Baltic who (surprise!) happens to show hg. N, and he signals the first incoming True Uralic™ speaker from wherever… It won’t be the first or the last time some people resort to “the complexity of Uralic-speaking peoples” in ancestry, just to look for “hg. N = Uralic” like crazy. You only need common sense to understand that this is not how this works. Amateur genomics can’t get more embarrassing than the current “let’s look for ‘Siberian ancestry’ in every individual of haplogroup N” trend. Or maybe it can, and it will, but I can’t see it yet.
If one were to insist on looking for ‘foreign’ contributions among Iron Age Estonians, though, I think one should also check out first archaeology, and then the PC3 (or, more graphically, a 3D plot), to understand what might be happening with the many Uralic clines derived from Corded Ware, before starting to play around with bioinformatic tools to discover a teeny tiny 1% admixture of the wrong population, and rushing to build far-fetched narratives. Apparently, one of the different clines formed roughly between southern (steppe – forest-steppe) and northern (tundra-taiga) populations in Uralians is also seen in some Iron Age Estonian individuals – especially in some late samples from Ingria…This is not my main interest, so I will leave this here for others to keep wasting their time chasing the white whale of the 0.5% of True Uralic™ ancestry in ancient Baltic samples of hg. N.
An exclusive Volga-Kama homeland for Disintegrating Uralic?
Since I don’t believe in macro-regions of largely continuous ethnolinguistic communities, as I have often said about Slavic (naively associated with prehistoric tribes of Eastern Europe) or Germanic (absurdly considered to be represented by Battle Axe), it is difficult for me to believe that Battle Axe-derived cultures remained of the same Finno-Samic dialects since the Corded Ware expansion…unless we live in Westeros, where everything happens “for thousands of years”.
I have to admit, then, that the now prevalent identification among Uralicists has become quite attractive:
Fatyanovo-Balanovo as Finno-Permic:
Fatyanovo/Netted Ware with West Uralic (also called Finno-Mordvinic).
Balanovo/Chirkovo-Kazan with Central Uralic (Mari-Permic).
Abashevo, into the Andronovo-like Horizon through the Seima-Turbino phenomenon, with East Uralic (also Ugro-Samoyedic).
Exactly like the identification of Yamna Hungary – Bell Beaker transition as the North-West Indo-European homeland, it gives us simplicity and small and late ethnolinguistic communities, away from the traditionally overused big and early language territories.
This late homeland would be supported, among others, by:
The presence of Indo-Iranian loanwords in Finno-Permic and Ugric (probably also in Samoyedic, either lost, or – much more likely – underresearched), compatible with the immediate contact between Abashevo – Sintashta-Potapovka-Filatovka and Fatyanovo-Balanovo.
The supposed expansion of Netted Ware from Fatyanovo to the north-west, which may be explained as the split and expansion of Balto-Finnic and Samic ca. 1900 BC.
A longer-lasting Finno-Permic (West+Central Uralic) community contrasting with the early separation of East Uralic.
The compatibility of this late expansion with the late expansion of Pre-Germanic from Denmark with the Dagger Period, and of Balto-Slavic with Trzciniec, which puts all three dialects reaching the Baltic Sea in the EBA.
NOTE. I meant to update the linguistic text to include the most recently favoured phylogenetic tree of Uralic languages after Häkkinen (2007, 2009, 2014), which has very quickly become the new normal among Uralicists, but I don’t think I will have enough time to review the necessary papers for that. I am rushing to publish a printed edition, so the text will wind up being a mixture of “traditional” (meaning, basically, pre-2010s) description of Uralic dialects but using modern divisions; say, “West Uralic” instead of “Finno-Samic”. By the way, I am still amazed that none of my reader-haters (or any online user discussing Uralic migrations, for that matter) have come up with the questions that the new division pose, and it supports my suspicion about the complete lack of interest in linguistics of most (a)DNA fans, except for the occasional use of old and free PDFs Googled to support new narratives invented expressly for some qpAdm results…
Problems with this Parpola-Carpelan’s (2012-2018) interpretation include:
The differentiation between Fennoscandian Textile Ceramics vs. Netted Ware, which is not warranted in archaeology. The assumption that Netted Ware expanded to the Baltic Sea (as Kallio does, following the traditional view) is thus weak, and it was probably a question of cultural contacts coupled with short-distance population movements/exchange in both directions (from the Baltic to the Volga and vice versa). In fact, the culture division relies on some fairly common and technically simple ornamentation patterns, widespread all over northern Europe, even before the Corded Ware expansion, and it is very difficult to separate certain neighboring Textile Ceramics from Netted Ware groups in southern Finland (i.e. Sarsa-Tomitsa groups).
The strict and radical direction described for the Netted Ware by Carpelan, as an eastward and northward expansion, within a very short time frame (ca. 1900-1800 BC), based on few radiocarbon dates, which seems to me like a very risky assumption. We know how this kind of descriptions of direction of culture expansion based on radiocarbon dates has turned out in much more complex “packages”, like the Bell Beaker culture… In fact, the earliest dates for Textile Ware are from the East Baltic, earlier than those of Netted Ware.
The assumption that Balto-Finnic traits shared with Mordvinic are a) late and b) meaningful for dialectalization of two closely related dialects, when it is clear that both dialects separated quite early. Phonologically Finnic is more conservative, morphologically less so, and the shared traits include a handful of non-Uralic substrate words which can’t be traced to a single common source, hence they were adopted when both languages had already separated… All in all, Finnic – Mordvinic correspondances are not even close to Italo-Celtic ones, which is clearly fully incompatible with a proposal of a Finnic separation from Mordvinic coinciding with the LBA-IA transition.
Especially problematic for Parpola’s model is the lack of genetic impact in Bronze Age or Iron Age Estonians, not reaching a significant level under any possible statistical threshold – which I am sure was quite disappointing for some of my readers -, but is in line with major archaeological continuity of groups the from region, only disturbed in cultural (and Y-chromosome) terms by the expansion of Akozino warrior-traders all over the Baltic Sea. Any proposed population movement will be very difficult to support in genetics, given the Corded Ware-derived populations that we will see in both regions, and the continued Baltic-Volga contacts since the Corded Ware expansion.
Problems with an interpretation of such a small impact in population genomics includes the similarly weak impacts and haplogroup infiltrations that can be seen among populations basically everywhere in Eurasia, during any given period, and much greater genetic impacts that are supposed to be (or that were certainly) followed by ethnolinguistic continuity.
The Battle Axe question
From Kallio (2015), about choosing a tentative homeland for Proto-Uralic:
(…) linguistically uniform Proto-Uralic would have been spoken in the Volga-Oka region until the mid-third millennium BC when the Proto-Uralic-speaking area would have expanded to the Volga-Kama region as well. By the end of the same millennium, this expansion would have led to the earliest dialectal splits within Uralic into Finno-Mordvin, Mari-Permic, and Ugro-Samoyed. The splitting up of these three soon followed during the early second millennium BC when the Uralic-speaking area finally stretched from the Baltic Sea in the west to the Altai mountains in the east. Indeed, no matter where Proto-Uralic was spoken, the branching into the nine well-attested subgroups (viz. Finnic, Saami, Mordvin, Mari, Permic, Hungarian, Mansi, Khanty, and Samoyed) must have taken less than a millennium, because their shared phonological and morphosyntactic isoglosses are rather limited (see Salminen 2002). The traditional view that all this branching would have taken several millennia violates everything linguistic typology teaches us about the rate of language change.
The basic problem of this identification of Fatyanovo-Balanovo as West-Central Uralic and Abashevo as East Uralic is the nature of the Battle Axe culture, including the Bronze Age East Baltic and Gulf of Finland area. Even if it is accepted that Fatyanovo-Balanovo represented all Western groups, Battle Axe must have represented West Uralic-like dialects.
The ethnolinguistic identification of Battle Axe depends ultimately on the nature of contacts of Fatyanovo/Netted Ware with Battle Axe/Textile Ceramics. If both groups were close and interacted profusely, as it seems, it doesn’t seem granted that we will be able to distinguish a close Para-West Uralic dialect of Scandinavia from the actual expanding Balto-Finnic and Samic dialects, if they were actually linked to the Netted Ware expansion. Also from Kallio (2015):
No doubt the most convincing substrate theory has recently been put forward by the Saami Uralicist Ante Aikio (2004), who has not only rehabilitated but also improved the old idea of a non-Uralic substrate in Saami. His study shows that there were still non-Uralic languages spoken in Northern Fennoscandia as recently as the first millennium AD. Most of all, they were not only genetically non-Uralic but also typologically non-Uralic-looking, bearing a closer resemblance to the so-called Palaeo-European substrates (for which see e.g. Schrijver 2001; Vennemann 2003).
In comparison, the case of Finnic is much more difficult. The fact that Proto-Uralic was not spoken in the East Baltic region means that this area must have originally been non-Uralic-speaking, but so far the evidence for a non-Uralic substrate in Finnic has consisted of appellatives and proper names with no etymology (cf. Ariste 1971; Saarikivi 2004a). Contrary to the proposed substrate words in Saami, those in Finnic show no structural non-Uralisms, as if they had indeed been borrowed from some genetically related or at least typologically similar languages, as I suggested above. Also none of them is more recent than the Middle Proto-Finnic stage, which makes them at least two millennia old. All this agrees with archaeological evidence discussed earlier that the Uralicization of the East Baltic region occurred during the Bronze Age (ca. 1900–500 BC).
The discussion of the paper continues with an unsuccessful attempt to find a hypothetical ancient Indo-European substrate that Kallio believes must be associated with the expansion of Corded Ware, in line with the traditional belief. For example, the often mentioned – almost folk etymology-like, unsurprisingly popular among amateurs – ‘Neva’ as derived from IE “young” is logically rejected…Unlike Parpola, Kallio’s view seems to be confident that Netted Ware (as Textile Ware) expanded into the East Baltic, on both sides of the Gulf of Finland, already during the Bronze Age.
As it has become apparent in population genomics, none of them was right, and Textile Ceramics will essentially show – like Netted Ware – a large genetic continuity of Corded Ware peoples in the whole north-eastern European forest zone – despite small regional population movements, obviously -, which necessarily implies that the whole Corded Ware culture – and not only Fatyanovo-Balanovo and Abashevo – were Uralic-speaking territories.
The similarities in terms of culture and Y-DNA bottlenecks between Battle Axe and Fatyanovo-Balanovo also imply that the linguistic differences between these groups were probably not many, and became strongly divided only after their territorial division. Continued contacts between Battle Axe- and Fatyanovo-derived groups can explain the proposed contacts (Finnic with Samic, Finnic with Mordvinic) after their linguistic-but-not-physical separation.
Battle Axe spoke “Para-Balto-Finnic”?
The Balto-Finnic-speaking nature of Battle Axe is thus supported by:
The lack of non-Uralic substrates in Balto-Finnic territory (Kallio 2015).
The early separation of Samic and Finnic from Mordvinic, and the virtual identity of Proto-West-Uralic and Proto-Uralic, which suggests that Proto-Uralic spread fast (Parpola 2012).
The scarce non-Uralic topo-hydronymy in the East Baltic and around the Gulf of Finland (Saarikivi 2004), comparable to that on the Upper Volga region.
The strong influence of a Balto-Finnic-like substrate on Pre-Germanic (or, in Kallio’s opinion, the same Scandinavian substrate influencing both Germanic and Balto-Finnic at the same time), and the continued influence of Balto-Finnic on Proto-Baltic and Proto-Slavic.
The continued influence of Corded Ware-derived groups in central-east Sweden in Finland and the East Baltic in terms of agricultural innovations appearing in the LBA, compatible with Schrijver’s proposal of intermediate Germanic-shifted Balto-Finnic groups and Balto-Finnic groups influenced by their pronunciation.
The intense Palaeo-Germanic and late Balto-Slavic / early Proto-Baltic superstrate on Balto-Finnic, which place all three dialects around the Baltic Sea since the Early Bronze Age.
The easy replacement of a hypothetic Para-Balto-Finnic dialect by incoming Proto-Balto-Finnic-speaking peoples (say, with textile ceramics), without much linguistic impact.
In fact, the continuous contacts of the East Baltic with the Volga, and especially the close interaction with Akozino warrior-traders just before the Tarand-grave period, could be the actual origin of the recent (if any) Finnic-Mordvinic connections that need to be traced back to the LBA-IA (maybe here the number ‘ten’), since most of them can be related to a Pit-Comb Ware culture substrate and earlier contacts through the forest zone, which Samic (due to its early split and presence to the north of the Gulf of Finland during the BA) does not share. In fact, some of them can be traced back to Balto-Finnic first…
These are the most often mentioned, in order of descending relevance for a shared ancient community:
Noun paradigms and the form and function of individual cases.
The geminate *mm (foreign to Proto-Uralic before the development of Fennic under Germanic influence) and other non-Uralic consonant clusters.
The change of numeral *luka ‘ten’ with (non-Uralic) *kümmen.
The presence of loanwords of non-Uralic origin, related to farming and trees, potentially Palaeo-European in nature.
It’s not only a question of quantity. Are these shared Mordvinic – Balto-Finnic traits really more relevant than, say, those between Italo-Celtic, which are supposed to have formed a community for a very short period at the end of the 3rd millennium around the Alps? Are these traits even sufficient to propose a common early Mordvinic-Finnic group within West Uralic, rather than loose Mordvinic – Balto-Finnic contacts, i.e. contacts between East Baltic (Textile Ceramics) and Volga-Kama (Netted Ware)?
Based on the alternative (Kallio’s) view of continued contacts between Textile Ceramics groups, even without knowing anything about linguistics, you can guess that Parpola is spinning very thin when assuming that these changes suggest that Balto-Finnic may have expanded with Akozino warrior-traders, separating thus ca. 800 BC from Mordvinic…
Genetic findings now clearly help dismiss any meaningful population impact in the LBA-IA transition, although any linguist can obviously argue for linguistic change in spite of major genetic continuity. But then we are stuck in the pre-ancient DNA era, so what’s ancient DNA for.
Genetic continuity = language continuity?
In the end, it’s very difficult to say how much language continuity there is around Estonia since the arrival of Corded Ware peoples. Looking at Modern Estonians, they have been clearly influenced by recent contacts with Baltic- and Germanic-speaking peoples clustering to the south-west in the PCA. They seem to have also received contacts from north(-east)ern peoples, likely from Finland, evidenced by their shifts toward the modern Estonian cluster during and after the Middle Ages, with a slight increase in Siberian ancestry and N1c subclades associated with Lovozero Ware. How much language change did these contacts bring? Maybe an expansion of Gulf of Finland Finnic (Northern Estonian) over Inland Finnic (Southern Estonian) and Gulf of Riga Finnic (Livonian)? Difficult to know, exactly, but, in the traditional view of Balto-Finnic dialectal distribution among Uralicists like Kallio, possibly no change at all.
So, if the obvious changes in the Estonia_MA cluster relative to Estonia_IA cluster and Estonia_Modern relative to Estonia_MA do not represent radical language change…Why would Estonia_IA represent a change relative to Estonia_BA, when it is statistically basically the same? Or Estonia_BA relative to CWC_Baltic? Because of the infiltration of haplogroup N1c around the whole Baltic? Because of the occasional 1% “Siberian” ancestry in some non-locals of varied haplogroups across the whole Baltic area?
In spite of all this, the amount of special pleading we are seeing among openly Nordicist amateurs when discussing the Uralic homeland relative to the Indo-European question in genetics has become a matter of plain willful ignorance. Like the living corpses of the Anatolian homeland, the Armenian homeland, the OIT proponents, or the nativist Basque R1b association, the personal involvement in the revival of “R1a=Indo-European” and “N=Uralic” trends is just painful to watch.
[Next post in this line, if I manage to make time for it: “Genetic (dis)continuity in Central Europe“. Let’s see if early Balts and early Slavs, as well as Germanic peoples, show a cluster closer to Danubian EBA (viz. Maros), Hungary-Balkans BA, and Urnfield-related samples than their predecessors in their areas, i.e. away from East Corded Ware groups… If you want, you can enjoy for the moment the new PCAs I could get done and the tentative map of languages in the Early Bronze Age, that will probably give you the right idea about early Indo-European and Uralic population movements]
In this study, we present new genomic data from Estonian Late Bronze Age stone-cist graves (1200–400 BC) (EstBA) and Pre-Roman Iron Age tarand cemeteries (800/500 BC–50 AD) (EstIA). The cultural background of stone-cist graves indicates strong connections both to the west and the east [20, 21]. The Iron Age (IA) tarands have been proposed to mirror “houses of the dead” found among Uralic peoples of the Volga-Kama region .
(…) The 33 individuals included 15 from EstBA, 6 from EstIA, 5 from Pre-Roman to Roman Iron Age Ingria (500 BC–450 AD) (IngIA), and 7 from Middle Age Estonia (1200–1600 AD) (EstMA) and yielded endogenous DNA ∼4%–88%, average genomic coverages ∼0.017–0.734×, and contamination estimates <4% (Table S1). We analyzed the data in the context of modern and other ancient individuals, including from Neolithic Estonia .
We identified chrY hgs for 30 male individuals (Tables 1 and S2; STAR Methods). All 16 successfully haplogrouped EstBA males belonged to hg R1a, showing no change from the CWC period, when this was also the only chrY lineage detected in the Eastern Baltic [11, 13, 30, 31]. Three EstIA and two IngIA individuals also belonged to hg R1a, but three EstIA males belonged to hg N3a, the earliest so far observed in the Eastern Baltic. Three EstMA individuals belonged to hg N3a, two to hg R1a, and one to hg J2b. ChrY lineages found in the Baltic Sea region before the CWC belong to hgs I, R1b, R1a5, and Q [10, 11, 12, 13, 17, 32]. Thus, it appears that these lineages were substantially replaced in the Eastern Baltic by hg R1a [10, 11, 12, 13], most likely through steppe migrations from the east [30, 31]. (…) Our results enable us to conclude that, although the expansion time for R1a1 and N3a3′5 in Eastern Europe is similar , hg N3a likely reached Estonia or at least became comparably frequent to modern Estonia  only during the BA-IA transition.
A clear shift toward West Eurasian hunter-gatherers is visible between European LN and BA (including Baltic CWC) and EstBA individuals, the latter clustering together with Latvian and Lithuanian BA individuals . EstIA, IngIA, and EstMA individuals project between BA individuals and modern Estonians, partially overlapping with both.
(…) EstBA individuals are clearly distinguishable from Estonian CWC individuals as the former have more of the blue component most frequent in WHGs and less of the brown and yellow components maximized in Caucasus hunter-gatherers and modern Khanty, respectively. The individuals of EstBA, EstIA, IngIA, EstMA, and modern Estonia are quite similar to each other on average, indicating that the relatively high proportion of WHG ancestry in modern Eastern Baltic populations compared to other present-day Europeans  traces back to the BA.
When comparing Estonian CWC and EstBA using autosomal outgroup f3 and Patterson’s D statistics (Table S3), the latter is more similar to other Baltic BA populations, to Baltic IA and Middle Age (MA) populations, and also to populations similar to WHGs and Scandinavian hunter-gatherers (SHGs), but not to Estonian CCC (Figures 2A and S2A; Data S1). The increase in WHG or SHG ancestry could be connected to western influences seen in material culture [20, 21] and facilitated by a decline in local population after the CCC-CWC period . A slight trend of bigger similarity of Estonian CWC to forest or steppe zone populations and of EstBA to European early farmer populations can also be seen.
(…) When comparing to modern populations, Estonian CWC is slightly more similar to Caucasus individuals but EstBA to Baltic populations and Finnic speakers (Figure 2B; Data S1). Outgroup f3 and D statistics do not reveal apparent differences when comparing EstBA to EstIA, EstIA to IngIA, and EstIA to EstMA (Data S1).
These results highlight how uniparental and autosomal data can lead to different demographic inferences—the genetic change between CWC and BA not seen in uniparental lineages is clear in autosomal data and the appearance of chrY hg N in the IA is not matched by a clear shift in autosomal profiles.
EstBA individuals have no Nganasan-related ancestry and EstIA, IngIA, and EstMA individuals on average have 2% or 4% (Figure 3; Data S1). The differentiation remains when using BA or IA Fennoscandian populations  instead of Nganasans (Data S1). Notably, the proportion of Nganasan-related ancestry varies between 0% and 12% among sampled EstIA, IngIA, and EstMA individuals (Data S1), which may suggest its relatively recent admixture into the target population. Moreover, two individuals from Kunda (0LS10 and V10) have the highest proportions of Nganasan ancestry among EstIA (6% and 8%), one of them has chrY hg N3a, and isotopic analysis suggests neither individual being born in Kunda .
About these two males from Tarand-graves, ‘foreign’ to Kunda:
0LS10: Male from tarand III (burial 9; TÜ 1325: L777), age 17–25 years . He had a fragment of a sheep/goat bone and ceramics as grave goods. This burial has two radiocarbon dates: 2430 ± 35 BP (Poz-10801; 760–400 cal BC) and 2530 ± 41 BP (UBA-26114; 800–530 cal BC) . According to the isotopic analysis, the person was not born in the vicinity of Kunda; his place of birth is still unknown (but south-western Finland and Sweden are excluded) . Sampled tooth r P1.
V10: Male from tarand XI (burial 24; TÜ 1325: L1925), age 25–35 years , date 2484 ± 40 BP (UBA-26115; 790–430 cal BC) . He had a few potsherds near the skull. Likewise, this person was not locally born . Sampled tooth l P1.
The paper shows thus:
Major continuity of ancestry from Corded Ware to modern Estonians, with only slight changes in different periods. In fact, one of the best fits for the Late Bronze Age ancestry is Gyvakarai1, one of the Corded Ware “outliers” described as “closer to Yamna”, which I already said may be closer to Sredni Stog/EHG populations instead. Another interesting take is that the change from Bronze Age to Iron Age corresponds to an increase in Baltic Corded Ware-related ancestry, rather than being driven by Siberian ancestry.
A Volosovo-related migration of hg. N1c with Netted Ware into the area seems to be discarded, based on the full replacement of paternal lines and continuity of R1a-Z283. It is only during the Tarand-grave period when a system of chiefdoms (spread from Ananyino/Akozino) brings haplogroup N1c to the Gulf of Finland. During the Iron Age, the proportion of paternal lineages is still clearly in favour of R1a (50% in the coast, 100% in Ostrobothnia), which indicates a gradual replacement led by elites, likely because of the incorporation of Akozino warrior-traders spreading all over the Baltic, bringing the described shared Mordvinic traits in Fennic.
The arrival of Akozino warrior-traders (bringing N1c and R1a lineages) was probably linked to this minimal “Nganasan-like” ancestry of some samples in the transition to the Iron Age. This arrival is supported by samples 0LS10 (the earliest hg. N1c) and V10 (of hg. R1a), both dated to ca. 800-400 BC, with V10 showing the highest “Nganasan-like” ancestry with 4.8%, both of them neighbouring samples showing 0%. This variable admixture among local and foreign paternal lineages might support the described social system of family alliances with intermarriages. In fact, a medieval sample, 0LS03_1 (hg. R1a) also shows a recent “Nganasan-like” ancestry, which probably points to the integration of different Arctic-related ancestry components among Modern Estonians, in this case related to Finnish expansions and thus integration of Levänluhta-related ancestry, as per the supplementary data.
NOTE. Such minimal proportions of “Nganasan-like” ancestry evidence the process of admixture of Volga Finns in Akozino territory through their close interactions with Permians of Ananyino, who in turn acquired this Palaeo-Arctic admixture most likely during the expansion of the linguistic community to hunter-gatherer territories, to the north of the Cis-Urals. This process of stepped infiltration and expansion without language change is not dissimilar to the one seen among Indo-Iranians and Balto-Slavs of hg. R1b, or Vasconic speakers of hg. I2a, although in the case of Baltic Finns of hg. R1a the process of infiltration and expansion of hg. N1c is much less dramatic, with no radical replacement anywhere before the huge bottlenecks observable in Finns.
The expansion of haplogroup N1c among Finnic populations, as we are going to see in samples from the Middle Ages such as Luistari, is the consequence of late founder effects after huge bottlenecks expected based on the analysis of modern populations. The expansion of N1c-VL29 is different in origin from that of N1c-Z1936 among Samic (later integrated into Finnish populations), most likely from the east and originally associated with Lovozero Ware.
In spite of all this, the conclusion of the paper is (surprise!) that Siberian ancestry and hg. N heralded the arrival of Finnic to the Gulf of Finland in the Iron Age… However, this conclusion is supposedly* supported, not by their previous papers, but by a recent phylogenetic study by Honkola et al. (2013), which doesn’t actually argue for such a late ‘arrival’: it argues for the split of Balto-Finnic around 1500 BC.
NOTE. I say ‘supposedly’ because Kristiina Tambets, for example, has been following the link of Uralic with haplogroup N since the 2000s, so this is not some conclusion they just happened to misread from some random paper they Googled. In those initial assessments, she argued that the “ancient homeland” of the Tat C mutation suggested that Finno-Ugrians were in Fennoscandia before Indo-Europeans. Apparently, since haplogroup N appears later and from the east, it is now more important to follow this haplogroup than what is established in archaeology and linguistics.
Even in the referred paper, this split is considered an in situ development, since the phylogenetic study takes the information – among others – 1) from Parpola and Carpelan, who consider Netted Ware, a culture derived from Fatyanovo/Abashevo and Volosovo, as the culprit of the Finno-Ugric expansion; and 2) from Kallio (2006), who clearly states that Proto-Balto-Finnic (like Proto-Finno-Samic) was spoken around the Gulf of Finland during the Bronze Age. Both of them set the terminus ante quem of the language presence in the Baltic ca. 1900 BC.
Anyways, as a consequence of geneticists keeping these untenable pre-ancient DNA haplogroup-based arguments today, I expect to see this “Finnic” language expansion also described for the Western Baltic, Scandinavia or northern Europe, when this same proportion of hg. N1c and “Nganasan” ancestry is observed in Iron Age samples around the Baltic Sea. The nativist trends that this domination of “Finns” all over Northern Europe 2,500 years ago will create will be even more fun to read than the current ones…
EDIT (10 May 2019) How I see the reaction of many to ancient DNA, in keeping their old theories:
So, the Bronze Age results for Iberia I2a, Yamna/BBC R1b, Baikalic/Palaeo-Arctic N1c, and now Corded Ware/Fennoscandia R1a (https://t.co/AKPnqpQHsb …), mean that the simplistic associations of haplogroup-language by geneticists during the pre-ancient DNA era was *right*? Mmmm… pic.twitter.com/pq9tde2QiU
Interesting excerpts, referring mainly to Uralic peoples (emphasis mine):
A model-based clustering analysis using ADMIXTURE shows a similar pattern (Fig. 2b and Supplementary Fig. 3). Overall, the proportions of ancestry components associated with Eastern or Western Eurasians are well correlated with longitude in inner Eurasians (Fig. 3). Notable outliers include known historical migrants such as Kalmyks, Nogais and Dungans. The Uralic- and Yeniseian-speaking populations, as well as Russians from multiple locations, derive most of their Eastern Eurasian ancestry from a component most enriched in Nganasans, while Turkic/Mongolic speakers have this component together with another component most enriched in populations from the Russian Far East, such as Ulchi and Nivkh (Supplementary Fig. 3). Turkic/Mongolic speakers comprising the bottom-most cline have a distinct Western Eurasian ancestry profile: they have a high proportion of a component most enriched in Mesolithic Caucasus hunter-gatherers and Neolithic Iranians and frequently harbour another component enriched in present-day South Asians (Supplementary Fig. 4). Based on the PCA and ADMIXTURE results, we heuristically assigned inner Eurasians to three clines: the ‘forest-tundra’ cline includes Russians and all Uralic and Yeniseian speakers; the ‘steppe-forest’ cline includes Turkic- and Mongolic-speaking populations from the Volga and Altai–Sayan regions and Southern Siberia; and the ‘southern steppe’ cline includes the rest of the populations.
For the forest-tundra populations, the Nganasan + Srubnaya model is adequate only for the two Volga region populations, Udmurts and Besermyans (Fig. 5 and Supplementary Table 8).
For the other populations west of the Urals, six from the northeastern corner of Europe are modelled with additional Mesolithic Western European hunter-gatherer (WHG) contribution (8.2–11.4%; Supplementary Table 8), while the rest need both WHG and early Neolithic European farmers (LBK_EN; Supplementary Table 2). Nganasan-related ancestry substantially contributes to their gene pools and cannot be removed from the model without a significant decrease in the model fit (4.1–29.0% contribution; χ2 P ≤ 1.68 × 10−5; Supplementary Table 8).
NOTE. It doesn’t seem like Hungarians can be easily modelled with Nganasan ancestry, though…
For the 4 populations east of the Urals (Enets, Selkups, Kets and Mansi), for which the above models are not adequate, Nganasan + Srubnaya + AG3 provides a good fit (χ2 P ≥ 0.018; Fig. 5 and Supplementary Table 8). Using early Bronze Age populations from the Baikal Lake region (‘Baikal_EBA’; Supplementary Table 2) as a reference instead of Nganasan, the two-way model of Baikal_EBA + Srubnaya provides a reasonable fit (χ2 P ≥ 0.016; Supplementary Table 8) and the three-way model of Baikal_EBA + Srubnaya + AG3 is adequate but with negative AG3 contribution for Enets and Mansi (χ2 P ≥ 0.460; Supplementary Table 8).
Bronze/Iron Age populations from Southern Siberia also show a similar ancestry composition with high ANE affinity (Supplementary Table 9). The additional ANE contribution beyond the Nganasan + Srubnaya model suggests a legacy from ANE-ancestry-rich clines before the Late Bronze Age.
Even among the earliest available inner Eurasian genomes, east–west connectivity is evident. These, too, form a longitudinal cline, characterized by the easterly increase of a distinct ancestry, labelled Ancient North Eurasian (ANE), lowest in western European hunter-gatherers (WHG) and highest in Palaeolithic Siberians from the Baikal region. Flow-through from this ANE cline is seen in steppe populations until at least the Bronze Age, including the world’s earliest known horse herders — the Botai. However, this is eroded over time by migration from west and east, following agricultural adoption on the continental peripheries (Fig. 1b,c).
Strikingly, Jeong et al. model the modern upper steppe cline as a simple two-way mixture between western Late Bronze Age herders and Northeast Asians (Fig. 1c), with no detectable residue from the older ANE cline. They propose modern steppe peoples were established mainly through migrations post-dating the Bronze Age, a sequence for which has been recently outlined using ancient genomes. In contrast, they confirm a substantial ANE legacy in modern Siberians of the northernmost cline, a pattern mirrored in excesses of WHG ancestry west of the Urals (Fig. 1b). This marks the inhospitable biome as a reservoir for older lineages, an indication that longstanding barriers to latitudinal movement may indeed be at work, reducing the penetrance of gene flows further south along the steppe.
Given the findings as reported in the paper, I think it should be much easier to describe different subclines in the “northernmost cline” than in the much more recent “Turkic/Mongolic cline”, which is nevertheless subdivided in this paper in two clines. As an example, there are at least two obvious clines with “Nganasan-related meta-populations” among Uralians, which converge in a common Steppe MLBA (i.e. Corded Ware) ancestry – one with Palaeo-Laplandic peoples, and another one with different Palaeo-Siberian populations:
The inclusion of certain Eurasian groups (or lack thereof) in the PCA doesn’t help to distinguish these subclines visually, and I guess the tiny “Naganasan-related” ancestral components found in some western populations (e.g. the famous ~5% among Estonians) probably don’t lend themselves easily to further subdivisions. Notice, nevertheless, the different components of the Eastern Eurasian source populations among Finno-Ugrians:
Also remarkable is the lack of comparison of Uralic populations with other neighbouring ones, since the described Uralic-like ancestry of Russians was already known, and is most likely due to the recent acculturation of Uralic-speaking peoples in the cradle of Russians, right before their eastward expansions.
A comparison of Estonians and Finns with Balts, Scandinavians, and Eastern Europeans would have been more informative for the division of the different so-called “Nganasan-like meta-populations”, and to ascertain which one of these ancestral peoples along the ancient WHG:ANE cline could actually be connected (if at all) to the Cis-Urals.
Because, after all, based on linguistics and archaeology, geneticists are not supposed to be looking for populations from the North Asian Arctic region, for “Siberian ancestry”, or for haplogroup N1c – despite previous works by their peers – , but for the Bronze Age Volga-Kama region…
Second in popularity for the expansion of haplogroup N1a-L392 (ca. 4400 BC) is, apparently, the association with Turkic, and by extension with Micro-Altaic, after the Uralic link preferred in Europe; at least among certain eastern researchers.
According to the views of a number of authoritative researchers, the Yakut ethnos was formed in the territory of Yakutia as a result of the mixing of people from the south and the autochthonous population .
These three major Sakha paternal lineages may have also arrived in Yakutia at different times and/ or from different places and/or with a difference in several generations instead, or perhaps Y-chromosomal STR mutations may have taken place in situ in Yakutia. Nevertheless, the immediate common ancestor(s) from the Asian Steppe of these three most prevalent Sakha Y-chromosomal STR haplotypes possibly lived during the prominence of the Turkic Khaganates, hence the near-perfect matches observed across a wide range of Eurasian geography, including as far as from Cyprus in the West to Liaoning, China in the East, then Middle Lena in the North and Afghanistan in the South (Table 3 and Figure 5). There may also be haplotypes closely-related to ‘the dominant Elley line’ among Karakalpaks, Uzbeks and Tajiks, however, limitations in the loci coverage for the available dataset (only eight Y-chromosomal STR loci) precludes further conclusions on this matter .
According to the results presented here, very similar Y-STR haplotypes to that of the original Elley line were found in the west: Afghanistan and northern Cyprus, and in the east: Liaoning Province, China and Ulaanbaator, Northern Mongolia. In the case of the dominant Omogoy line, very closely matching haplotypes differing by a single mutational step were found in the city of Chifen of the Jirin Province, China. The widest range of similar haplotypes was found for the Yakut haplotype Unknown: In Mongolia, China and South Korea. For instance, haplotypes differing by a single step mutation were found in Northern Mongolia (Khalk, Darhad, Uryankhai populations), Ulaanbaator (Khalk) and in the province of Jirin, China (Han population).
Notably, Tat-C-bearing Y-chromosomes were also observed in ancient DNA samples from the 2700-3000 years-old Upper Xiajiadian culture in Inner Mongolia, as well as those from the Serteya II site at the Upper Dvina region in Russia and the ‘Devichyi gory’ culture of long barrow burials at the Nevel’sky district of Pskovsky region in Russia. A 14-loci Y-chromosomal STR median-joining network of the most prevalent Sakha haplotypes and a Tat-C-bearing haplotype from one of the ancient DNA samples recovered from the Upper Xiajiadian culture in Inner Mongolia (DSQ04) revealed that the contemporary Sakha haplotype ‘Xuo’ (Table 2, Haplotype ID “Xuo”) classified as that of ‘the Xiongnu clan’ in our current study, was the closest to the ancient Xiongnu haplotype (Figure 6). TMRCA estimate for this 14-loci Y-chromosomal STR network was 4357 ± 1038 years or 2341 ± 1038 BCE, which correlated well with the Upper Xiajiadian culture that was dated to the Late Bronze Age (700-1000 BCE).
Also, a simple look at the TMRCA and modern distribution was enough to hypothesize long ago the lack of connection of N1c-L392 with Altaic or Uralic peoples. From Ilumäe et al. (2016):
Previous research has shown that Y chromosomes of the Turkic-speaking Yakuts (Sakha) belong overwhelmingly to hg N3 (formerly N1c1). We found that nearly all of the more than 150 genotyped Yakut N3 Y chromosomes belong to the N3a2-M2118 clade, just as in the Turkic-speaking Dolgans and the linguistically distant Tungusic-speaking Evenks and Evens living in Yakutia (Table S2). Hence, the N3a2 patrilineage is a prime example of a male population of broad central Siberian ancestry that is not intrinsic to any linguistically defined group of people. Moreover, the deepest branch of hg N3a2 is represented by a Lebanese and a Chinese sample. This finding agrees with the sequence data from Hallast et al., where one Turkish Y chromosome was also assigned to the same sub-clade. Interestingly, N3a2 was also found in one Bhutan individual who represents a separate sub-lineage in the clade. These findings show that although N3a2 reflects a recent strong founder effect primarily in central Siberia (Yakutia, Sakha), the sub-clade has a much wider distribution area with incidental occurrences in the Near East and South Asia.
The most striking aspect of the phylogeography of hg N is the spread of the N3a3’6-CTS6967 lineages. Considering the three geographically most distant populations in our study—Chukchi, Buryats, and Lithuanians—it is remarkable to find that about half of the Y chromosome pool of each consists of hg N3 and that they share the same sub-clade N3a3’6. The fractionation of N3a3’6 into the four sub-clades that cover such an extraordinarily wide area occurred in the mid-Holocene, about 5.0 kya (95% CI = 4.4–5.7 kya). It is hard to pinpoint the precise region where the split of these lineages occurred. It could have happened somewhere in the middle of their geographic spread around the Urals or further east in West Siberia, where current regional diversity of hg N sub-lineages is the highest (Figure 1B). Yet, it is evident that the spread of the newly arisen sub-clades of N3a3’6 in opposing directions happened very quickly. Today, it unites the East Baltic, East Fennoscandia, Buryatia, Mongolia, and Chukotka-Kamchatka (Beringian) Eurasian regions, which are separated from each other by approximately 5,000–6,700 km by air. N3a3’6 has high frequencies in the patrilineal pools of populations belonging to the Altaic, Uralic, several Indo-European, and Chukotko-Kamchatkan language families. There is no generally agreed, time-resolved linguistic tree that unites these linguistic phyla. Yet, their split is almost certainly at least several millennia older than the rather recent expansion signal of the N3a3’6 sub-clade, suggesting that its spread had little to do with linguistic affinities of men carrying the N3a3’6 lineages.
It was thus clear long ago that N1c-L392 lineages must have expanded explosively in the 5th millennium through Northern Eurasia, probably from a region to the north of Lake Baikal, and that this expansion – and succeeding ones through Northern Eurasia – may not be associated to any known language group until well into the common era.
There remain ongoing discussions about the origins of the ethnic Russian population. The ancestors of ethnic Russians were among the Slavic tribes that separated from the early Indo-European Group, which included ancestors of modern Slavic, Germanic and Baltic speakers, who appeared in the northeastern part of Europe ca. 1,500 years ago. Slavs were found in the central part of Eastern Europe, where they came in direct contact with (and likely assimilation of) the populations speaking Uralic (Volga-Finnish and Baltic- Finnish), and also Baltic languages [11–13]. In the following centuries, Slavs interacted with the Iranian-Persian, Turkic and Scandinavian peoples, all of which in succession may have contributed to the current pattern of genome diversity across the different parts of Russia. At the end of the Middle Ages and in the early modern period, there occurred a division of the East Slavic unity into Russians, Ukrainians and Belarusians. It was the Russians who drove the colonization movement to the East, although other Slavic, Turkic and Finnish peoples took part in this movement, as the eastward migrations brought them to the Ural Mountains and further into Siberia, the Far East, and Alaska. During that interval, the Russians encountered the Finns, Ugrians, and Samoyeds speakers in the Urals, but also the Turkic, Mongolian and Tungus speakers of Siberia. Finally, in the great expanse between the Altai Mountains on the border with Mongolia, and the Bering Strait, they encountered paleo-Asiatic groups that may be genetically closest to the ancestors of the Native Americans. Today’s complex patchwork of human diversity in Russia has continued to be augmented by modern migrations from the Caucasus, and from Central Asia, as modern economic migrations take shape.
In the current study, we annotated whole genome sequences of individuals currently living on the territory of Russia and identifying themselves as ethnic Russian or as members of a named ethnic minority (Fig. 1). We analyzed genetic variation in three modern populations of Russia (ethnic Russians from Pskov and Novgorod regions and ethnic Yakut from the Sakha Republic), and compared them to the recently released genome sequences collected from 52 indigenous Russian populations. The incidence of function-altering mutations was explored by identifying known variants and novel variants and their allele frequencies relative to variation in adjacent European, East Asian and South Asian populations. Genomic variation was further used to estimate genetic distance and relationships, historic gene flow and barriers to gene flow, the extent of population admixture, historic population contractions, and linkage disequilibrium patterns. Lastly, we present demographic models estimating historic founder events within Russia, and a preliminary HapMap of ethnic Russians from the European part of Russia and Yakuts from eastern Siberia.
The collection of identified SNPs was used to inspect quantitative distinctions among 264 individuals from across Eurasia (Fig. 1) using Principal Component Analysis (PCA) (Fig. 2). The first and the second eigenvectors of the PCA plot are associated with longitude and latitude, respectively, of the sample locations and accurately separate Eurasian populations according to geographic origin. East European samples cluster near Pskov and Novgorod samples, which fall between northern Russians, Finno-Ugric peoples (Karelian, Finns, Veps etc.), and other Northeastern European peoples (Swedes, Central Russians, Estonian, Latvians, Lithuanians, and Ukrainians) (Fig. 2b). Yakut individuals map into the Siberian sample cluster as expected (Fig. 2a). To obtain an extended view of population relationships, we performed a maximum likelihood-based estimation of ancestry and population structure using ADMIXTURE (Fig. 2c). The Novgorod and Pskov populations show similar profiles with their Northeastern European ancestors while the Yakut ethnic group showed mixed ancestry similar to the Buryat and Mongolian groups.
Possible admixture sources of the Genome Russia populations were addressed more formally by calculating F3 statistics, which is an allele frequency-based measure, allowing to test if a target population can be modeled as a mixture of two source populations . Results showed that Yakut individuals are best modeled as an admixture of Evens or Evenks with various European populations (Supplemental Table S4). Pskov and Novgorod showed admixture of European with Siberian or Finno-Ugric populations, with Lithuanian and Latvian populations being the dominant European sources for Pskov samples.
So, Russians expanding in the Middle Ages as acculturaded Finno-Volgaic peoples.
A new paper (behind paywall) offers insight into the prevalent presence of R1a-Z93 among eastern Scytho-Siberian groups (most likely including Samoyedic speakers in the forest-steppes), and a new hint to the westward expansion of haplogroups Q and N (probably coupled with the so-called “Siberian ancestry”) from the east with different groups of Iron Age steppe nomads:
From an archeological and historical point of view, the term “Scythians” refers to Iron Age nomadic or seminomadic populations characterized by the presence of three types of artifacts in male burials: typical weapons, specific horse harnesses and items decorated in the so-called “Animal Style”. This complex of goods has been termed the “Scythian triad” and was considered to be characteristic of nomadic groups belonging to the “Scythian World” (Yablonsky 2001). This “Scythian World” includes both the Classic (or European) Scythians from the North Pontic region (7th–3th century BC) and the Southern Siberian (or Asian) populations of the Scythian period (also called Scytho-Siberians). These include, among others, the Sakas from Kazakhstan, the Tagar population from the Minusinsk Basin (Republic of Khakassia), the Aldy-Bel population from Tuva (Russian Federation) and the Pazyryk and Sagly cultures from the Altai Mountains.
In this work, we first aim to address the question of the familial and social organization of Scytho-Siberian groups by studying the genetic relationship of 29 individuals from the Aldy-Bel and Sagly cultures using autosomal STRs. (…) were obtained from 5 archeological sites located in the valley of the Eerbek river in Tuva Republic, Russia (Fig. 1). All the mounds of this archeological site were excavated but DNA samples were not collected from all of them. 14C dates mainly fall within the Hallstatt radiocarbon calibration plateau (ca. 800–400 cal BC) where the chronological resolution is poor. Only one date falls on an earlier segment of calibration curve: Le 9817–2650 ± 25 BP, i.e. 843–792 cal BC with a probability of 94.3% (using the OxCal v4.3.2 program). This sample (Bai-Dag 8, Kurgan 1, grave 10) is not from one of the graves studied but was used to date the kurgan as a whole.
Y-chromosome haplogroups were first assigned using the ISOGG 2018 nomenclature. In order to improve the precision of haplogroup definition, we also analyzed a set of Y-chromosome SNP (Supplementary Table 2). Nine samples belonged to the R1a-M513 haplogroup (defined by marker M513) and two of these nine samples were characterized as belonging to the R1a1a1b2-Z93 haplogroup or one of its subclades. Six samples belonged to the Q1b1a-L54 haplogroup and five of these six samples belonged to the Q1b1a3-L330 subclade. One sample belonged to the N-M231 haplogroup.
The distribution of these haplogroups in the population must be confronted with the prevalence of kinship among the samples. Although five individuals belonged to haplogroup Q1b1a3-L330, three of them (ARZ-T18, ARZ-T19 and ARZ-T20) were paternally related (Fig. 2). It must, therefore, be considered that haplogroup Q1b1a3-L330 is present in three independent instances (given that the remaining two instances exhibit no close familial relationship with other samples or one another). All five were buried on the Eki-Ottug 1 archaeological site (although in two different kurgans).
In the same way, although two groups, of two and three individuals, shared haplotypes belonging to the R1a-M513 haplogroup, these groups likely include a father/son pair (ARZ-T2 and ARZ-T12). Therefore, among nine R1a-M513 men, we found six independent haplotypes, one being present in two independent instances. All R1a-M513 haplotypes, however, including those attributed to the R1a1a1b2-Z93 subclade, only differed by one-step mutations, across 5 loci at most. All R1a-M513 individuals were buried on the same site, Eki-Ottug 2, in a single Kurgan.
Haplogroup R1a-M173 was previously reported for 6 Scytho-Siberian individuals from the Tagar culture (Keyser et al. 2009) and one Altaian Scytho-Siberian from the Sebÿstei site (Ricaut et al. 2004a), whereas haplogroup R1a1a1b2-Z93 (or R1a1a1b-S224) was described for one Scythian from Samara (Mathieson et al. 2015) and two Scytho-Siberians from Berel and the Tuva Republic (Unterländer et al. 2017). On the contrary, North Pontic Scythians were found to belong to the R1b1a1a2 haplogroup (Krzewińska et al. 2018), showing a distinction between the two groups of Scythians. (…) The absence of R1b lineages in the Scytho-Siberian individuals tested so far and their presence in the North Pontic Scythians suggest that these 2 groups had a completely different paternal lineage makeup with nearly no gene flow from male carriers between them.
The seven other male individuals studied in this work were found to carry Eastern Eurasian Y haplogroups Q1b1a and one of its subclades (n = 6) and N (n = 1). Haplogroup Q1b1a-L54 was previously described in four males from the Bronze Age in the Altai Mountains (Hollard et al. 2014, 2018) and was clearly associated with Siberian populations (Regueiro et al. 2013).
The N-M231 haplogroup emerged from haplogroup K in Southern Asia around 21,000 years BCE, maybe in Southern China (Shi et al. 2013; Ilumäe et al. 2016). Previous studies attested to its presence in samples from Neolithic and Bronze Age in China (Li et al. 2011; Cui et al. 2013). Waves of northwestern expansion of this haplogroup are described as beginning during the Paleolithic period (Derenko et al. 2006; Shi et al. 2013) but traces of this expansion in archeological samples were reported only in two Scytho-Siberian males from the Altai (Pilipenko et al. 2015).
The sample of haplogroup N comes from the Aldy-Bel culture (ARZ-T15), from the Eerbek site, but has no radiocarbon date. All Q1b-L330 samples come from the Sagly culture, and three are paternally related. The other Q1b-L54 sample is from other tombs in one kurgan at Aldy Bel.