Updates to ASoSaH: new maps, updated PCA, and added newest research papers

steppe-ancestry-cut

The title says it all. I have used some free time to update the series A Song of Sheep and Horses:

I basically added information from the latest papers published, which (luckily enough for me) haven’t been too many, and I have added images to illustrate certain sections.

I have updated the PCAs by including North Caucasus samples from Wang et al. (2018), whose position I could only infer for older versions from previously published PCA graphs.

pca-steppe-eneolithic-early
PCA of ancient and modern Eurasian samples. Early Eneolithic admixture events in the steppe drawn.

I have also added to the supplementary materials the “Tip of the Iceberg” R1b tree by Mike Walsh from the FTDNA R1b group, with permission, because some relevant genetic sections are centered on the evolution of R1b lineages, and the reader can get easily lost with so many subclades.

I have also updated maps, including some of the Y-DNA ones, and managed to finish two new maps I was working on, and I added them to the supplementary materials and to the menu above:

One on Yamna kurgans in Hungary, coupled with contemporaneous sites of Baden-Boleráz or Kostolac cultures:

burials-yamnaya-hungary
Map of attested Yamnaya pit-grave burials in the Hungarian plains; superimposed in shades of blue are common areas covered by floods before the extensive controls imposed in the 19th century; in orange, cumulative thickness of sand, unfavourable loamy sand layer. Marked are settlements/findings of Boleráz (ca. 3500 BC on), Baden (until ca. 2800 BC), Kostolac (precise dates unknown), and Yamna kurgans (from ca. 3100/3000 BC on).

Another one on Steppe ancestry expansion, with a tentative distribution of “steppe ancestry” divided into that of Sredni Stog/Corded Ware origin vs. that of Repin/Yamna origin, a difference that has been known for quite some time already.

It is tentative because there hasn’t been any professional study or amateur attempt to date to differentiate both “steppe ancestries” in Yamna, and especially in Bell Beakers. So much for the call of professional geneticists since 2018 (see here and here) and archaeologists since 2017 (see e.g. here and here) to distinguish fine-scale population structure to be able to follow neighbouring populations which expanded with different archaeological (and thus ethnolinguistic) groups.

steppe-ancestry-corded-ware
Tentative map of fine-scale population structure during steppe-related expansions (ca. 3500–2000 BC), including Repin–Yamna–Bell Beaker/Balkans and Sredni Stog–Corded Ware groups. Data based on published samples and pairwise comparisons tested to date. Notice that the potential admixture of expanding Repin/Early Yamna settlers in the North Pontic area with the late Sredni Stog population (and thus Sredni Stog-related ancestry in Yamna) has been omitted for simplicity purposes, assuming thus a homogeneous Yamna vs. Corded Ware ancestry.

I think both maps are especially important today, given the current Nordicist reactionary trends arguing (yet again) for an origin of Indo-Europeans in The North™, now based on the Fearsome Tisza River hypothesis, on cephalic index values, and a few pairwise comparisons – i.e. an absolutely no-nonsense approach to the Indo-European question (LOL). At least I get to relax and sit this year out just observing how other people bury themselves and their beloved “steppe ancestry=IE” under so many new pet theories…

NOTE. Not that there is anything wrong with a northern origin of North-West Indo-European from a linguistic point of view, as I commented recently – after all, a Corded Ware origin would roughly fit the linguistic guesstimates, unlike the proposed ancestral origins in Anatolia or India. The problem is that, like many other fringe theories, it is today just based on tradition, or (even worse) ethnic, political, or personal desires, and it doesn’t make sense when all findings from disciplines involved in the Indo-European and Uralic questions are combined.

steppe-ancestry-modern-populations
Simple ancestry percentages in modern populations. Recent image by Iain Mathieson 2019 (min. 5.57). A simplistic “Steppe ancestry” defining Indo-European speakers…? Sure.

Within 20 or 30 years, when genetic genealogists (or amateur geneticists, or however you want to call them) ask why we had the opportunity since 2015 to sample as many Hungarian Yamnaya individuals as possible and we didn’t, when it is clear that the number of unscathed kurgans is diminishing every year (from an estimated 4,000 in the 20th century, of the original tens of thousands, to less than 1,500 today) the answer will not be “because this or that archaeologist or linguist was a dilettante or a charlatan‘, as they usually describe academics they dislike.

It will be precisely because the very same genetic genealogists – supposedly interested today in the origin of R1b-L151 and/or genetic marker associated with North-West Indo-Europeans – are obsessed with finding them anywhere else but for Hungary, and prefer to use their money and time to play with a few statistical tools within a biased framework of flawed assumptions and study designs, obtaining absurd results and accepting far-fetched interpretations of them, to be told exactly what they want to hear: be it the Franco-Cantabrian homeland, the Dutch or Moravian Beaker from CWC homeland, the Maykop homeland, or the Moon homeland.

Poetic justice this heritage destruction, whose indirect causes will remain written in Internet archives for everyone to see, as a good lesson for future generations.

ASoSaH Reread (I): Y-DNA haplogroups among Indo-Europeans (apart from R1b-L23)

eneolithic-early-admixture-steppe-ancestry

Given my reduced free time in these months, I have decided to keep updating the text on Indo-European and Uralic migrations and/or this blog, simultaneously or alternatively, to make the most out of the time I can dedicate to this. I will add the different ‘A Song of Sheep and Horses (ASoSaH) reread’ posts to the original post announcing the books. I would be especially interested in comments and corrections to the book chapters rather than the posts, but any comments are welcome (including in the forum, where comments are more likely to stick).

This is mainly a reread of iv.2. Indo-Anatolians and vi.1. Disintegrating Indo-Europeans.

Indo-Anatolians and Late Indo-Europeans

I have often written about R1b-L23 as the majority haplogroup among Late Proto-Indo-Europeans (see my predictions for 2018 and my summary of 2018), but always expected other haplogroups to pop up somewhere along the way, in Khvalynsk, in Repin, in Yamna, and in Bell Beakers (see e.g. the post on common fallacies of R1a/IE-fans).

Luckily enough – for those of us who want precise answers to our previous infinite models of Indo-European language expansions (viz. GAC-associated expansion, IE-speaking Old Europe, Anatolian homeland, Iran homeland, Maykop as Proto-Anatolian, Palaeolithic Continuity Theory, Celtic in the Atlantic façade, etc.) – the situation has been more clear-cut than expected: it turns out that, especially during population expansions, acute Y-chromosome bottlenecks were very common in the past, at least until the Iron Age.

Khvalynsk and Repin-Yamna expansions were no different, and that seems quite natural in hindsight, given the strong familial ties and aversion to foreigners proper of the Late Proto-Indo-European society and culture – probably not really that different from other contemporary societies, like the neighbouring Late Proto-Uralic or Trypillian ones.

y-dna-khvalynsk
Y-DNA samples from Khvalynsk and neighbouring cultures. See full version here.

Y-DNA haplogroups

During the expansion of early Khvalynsk, the most likely Indo-Anatolian culture, the society of the Don-Volga area was probably made up of different lineages including R1b-V1636, R1b-M269, R1a-YP1272, Q1a-M25, and I2a-L699 (and possibly some R1b-V88?), a variability possibly greater than that of the contemporary north Pontic area, probably a sign of this region being a sink of different east and west migrations from steppe and forest areas.

During its expansion, the Khvalynsk society saw its haplogroup variability reduced, as evidenced by the succeeding expansive Repin culture:

Afanasevo, representing Pre-Tocharian (the earliest Late PIE dialect to branch off), expanded with R1b-L23 – especially R1b-Z2103 – lineages, while early Yamna expanded with R1b-L23 and I2a-L699 lineages, which suggests that these are the main haplogroups that survived the Y-DNA bottleneck undergone during the Khvalynsk expansion, and especially later during the late Repin expansion. Nevertheless, other old haplogroups might still pop up during the Repin and early Yamna period, such as the R1b-V1636 sample from Yamna in the Northern Caucasus.

It is still unclear if R1b-L23 sister clade R1b-PF7562 (formed ca. 4400 BC, TMRCA ca. 3400 BC), prevalent among modern Albanians, expanded with Yamna migrants, or if it was part of an earlier expansion of R1b-M269 into the Balkans, and represent thus Indo-Anatolian speakers who later hitchhiked the expansion of the Late PIE language from the north or west Pontic area. The early TMRCA seems to suggest an association with Repin (and therefore Yamna), rather than later movements in the Balkans.

chalcolithic-early-y-dna
Y-DNA samples from Yamnaya and neighbouring cultures. See full version here.

‘Yamnaya’ or ‘steppe’ ancestry?

After the early years when population genetics relied mainly on modern Y-DNA haplogroups, geneticists and amateurs have been recently playing around with testing “ancestry percentages”, based on newly developed free statistical tools, which offer obviously just one among many types of data to achieve a proper interpretation of the past.

Today we have quite a lot Y-DNA haplogroups reported for ancient samples of more recent prehistoric periods, and they seem to offer (at least since the 2015 papers, but more evidently since the 2018 papers on Bell Beakers and Europeans, Corded Ware, or Fennoscandia among others) the most straightforward interpretation of all results published in population genomics research.

NOTE. The finding of a specific type of ancestry in one isolated 40,000-year-old sample from Tianyuan can offer very interesting information on potential population movements to the region. However, the identification of ethnolinguistic communities and their migrations among neighbouring groups in Neolithic or Bronze Age groups is evidently not that simple.

PCA-caucasus-steppe-all
Yamnaya (Indo-European peoples) and their evolution in the steppes, together with North Pontic (eventually Uralic) peoples.Notice how little Indo-European ancestry changes from Khvalynsk (Indo-Anatolian) to Yamna Hungary (North-West Indo-Europeans) Image modified from Wang et al. (2018). See more on the evolution of “steppe ancestry”.

It is becoming more and more clear with each paper that the true “Yamnaya ancestry” – not the originally described one – was in fact associated with Indo-Europeans (see more on the very Yamnaya-like Yamna Hungary and early East Bell Beaker R1b samples, all of quite similar ancestry and PCA cluster before their further admixture with EEF- and CWC-like groups).

The so-called “steppe ancestry”, on the other hand, reflects the contribution of a Northern Caucasus-related ancestry to expanding Khvalynsk settlers, who spread through the steppes more than a thousand years before the expansion of Late Proto-Indo-Europeans with late Repin, and can thus be found among different groups related to the Pontic-Caspian steppes (see more on the emergence and evolution of “steppe ancestry”).

In fact, after the Yamna/Indo-European and Corded Ware/Uralic expansions, it is more likely to find “steppe ancestry” to the north and east in territories traditionally associated with Uralic languages, whereas to the south and west – i.e. in territories traditionally associated with Indo-European languages – it is more likely to find “EEF ancestry” with diminished “steppe ancestry”, among peoples patrilineally descended from Yamna settlers.

Y-DNA haplogroups, the only uniparental markers (see exceptions in mtDNA inheritance) – unlike ancestry percentages based on the comparison of a few samples and flawed study designs – do not admix, do not change, and therefore they do not lend themselves to infinite pet theories (see e.g. what David Reich has to say about R1b-P312 in Iberia directly derived from Yamna migrants in spite of their predominant EEF ancestry): their cultural continuity can only be challenged with carefully threaded linguistic, archaeological, and genetic data.

Related

Mongolian tribes cluster with East Asians, closely related to the Japanese

mongolian-sampling

New paper behind paywall Whole-genome sequencing of 175 Mongolians uncovers population-specific genetic architecture and gene flow throughout North and East Asia, by Bai et al Nature Genetics (2018).

Interesting excerpts (emphasis mine):

Genome sequencing, variant calling, and construction of the Mongolian reference panel. We collected peripheral blood with informed consent from 175 Mongolian individuals representing six distinct tribes/regions in northern China and Mongolia, including the Abaga, Khalkha, Oirat, Buryat, Sonid, and Horchin tribes.

mongolians-pca
Population genetic structure. a, PCA of Mongolian individuals and 1000G samples. Mongolians fill a large, less characterized gap between Admixed/Native Americans and other East Asians in the 1000G project. b, PCA of Mongolians and East Asians of 1000G. The abbreviations of EAS populations were used from reference 11.

The fixation index (FST) was used to estimate pairwise genetic differentiation among our Mongolian samples and 26 modern human populations selected from 1000G (…) the Mongolian tribes cluster with East Asian groups. The Mongolian populations show the smallest differentiation from the CHB, and FST values increase relative to the magnitude of geographical separation. The Buryat are the most differentiated tribe compared with other East Asians (1.82–2.97%), while the Horchin are the least (0.25–1.35%). All tribes are closer to the Japanese (JPT) than the CHS with the exception of the Horchin. Among the tribes, the Abaga, Khalkha, Oirat, and Sonid show the least differentiation from one another (FST < 0.15%)

A PCA places the Mongolians in close genetic proximity to a group of North Asian Siberians, including Altaians, Tuvinians, Evenki, and Yakut, indicating that the Mongolian whole-genome variation panel could be a better proxy for these groups than any populations currently in the 1000G panel

The most common Y-chromosome haplogroups are from the C3 sublineage (41.67%), including C3c (29.17%) and C3b (12.50%), followed by haplogroup O (23.61%), and haplogroup N (18.06%) (…) While haplogroups C and O are primarily restricted to Asia, haplogroup N is present at high frequency in Finns (60.5%), at low frequency in non-Mongolian East Asians (< 1%), and virtually absent throughout the remainder of European and African samples in 1000G

Comparison with Finns

ibd-sharing-mongolian-finns
Distribution of D-values from D-test under the model of [EAS, Mongolians, X, chimpanzee], where X represents the test population and chimpanzee serves as an outgroup. The positive D-value (Z > 3) indicates that the test population (X) is closer to Mongolians than to EAS. The whiskers correspond to range, and the dots to individual data points, box limits are the upper and lower quartiles. The n in each boxplot is 30. All abbreviations of populations in the figure were used from reference 11.

Of the populations included in our study, Mongolians share the second-highest level of IBD with the Finnish people (FIN), behind only Northern Han Chinese (CHB). While Mongolians share more IBD with Europeans (EUR) as a whole compared with other non-EAS people (Fig. 4b), removal of Finns from the Europeans drops the level of sharing to as low as that with South Asians (SAS) or Admixed American (AMR).

There is considerable geographic separation between modern-day Mongolians and Europe. The positive D-statistic that reveal gene flow between Mongolians and Europeans (Fig. 4c), and the high degree of IBD sharing with Finnish people in particular suggest that complex admixture may have occurred throughout northeastern Europe and Siberia. To see whether Mongolians represent the ethnic group in East Asia with the highest level of gene flow with Finnish people, we calculated a D-statistic for each set of populations [Mongolians, X, FIN, Yoruba (YRI)], where X represents a population from Siberia or Northern Canada. Most of the populations reveal an imbalance in allele frequencies that suggests gene flow with Finns (D >0, Z >3), but the greatest imbalance is observed between Siberians/Northern Canadians and Finnish, rather than between Mongolians and Finns. This pattern indicates that northern Asian populations interacted across large geographic ranges.

migration-finns-nganasan
6 migration events, from the supplementary materials.

I guess the 1000G does not have northern Eurasian groups, because the IBD map and values would be lightening up with Palaeo-Siberian peoples

Related

“Steppe ancestry” step by step: Khvalynsk, Sredni Stog, Repin, Yamna, Corded Ware

dzudzuana_pca-large

Wang et al. (2018) is obviously a game changer in many aspects. I have already written about the upcoming Yamna Hungary samples, about the new Steppe_Eneolithic and Caucasus Eneolithic keystones, and about the upcoming Greece Neolithic samples with steppe ancestry.

An interesting aspect of the paper, hidden among so many relevant details, is a clearer picture of how the so-called Yamnaya or steppe ancestry evolved from Samara hunter-gatherers to Yamna nomadic pastoralists, and how this ancestry appeared among Proto-Corded Ware populations.

anatolia-neolithic-steppe-eneolithic
Image modified from Wang et al. (2018). Marked are in orange: equivalent Steppe_Maykop ADMIXTURE; in red, approximate limit of Anatolia_Neolithic ancestry found in Yamna populations; in blue, Corded Ware-related groups. “Modelling results for the Steppe and Caucasus cluster. Admixture proportions based on (temporally and geographically) distal and proximal models, showing additional Anatolian farmer-related ancestry in Steppe groups as well as additional gene flow from the south in some of the Steppe groups as well as the Caucasus groups.”

Please note: arrows of “ancestry movement” in the following PCAs do not necessarily represent physical population movements, or even ethnolinguistic change. To avoid misinterpretations, I have depicted arrows with Y-DNA haplogroup migrations to represent the most likely true ethnolinguistic movements. Admixture graphics shown are from Wang et al. (2018), and also (the K12) from Mathieson et al. (2018).

1. Samara to Early Khvalynsk

The so-called steppe ancestry was born during the Khvalynsk expansion through the steppes, probably through exogamy of expanding elite clans (eventually all R1b-M269 lineages) originally of Samara_HG ancestry. The nearest group to the ANE-like ghost population with which Samara hunter-gatherers admixed is represented by the Steppe_Eneolithic / Steppe_Maykop cluster (from the Northern Caucasus Piedmont).

Steppe_Eneolithic samples, of R1b1 lineages, are probably expanded Khvalynsk peoples, showing thus a proximate ancestry of an Early Eneolithic ghost population of the Northern Caucasus. Steppe_Maykop samples represent a later replacement of this Steppe_Eneolithic population – and/or a similar population with further contribution of ANE-like ancestry – in the area some 1,000 years later.

PCA-caucasus-steppe-samara

This is what Steppe_Maykop looks like, different from Steppe_Eneolithic:

steppe-maykop-admixture

NOTE. This admixture shows how different Steppe_Maykop is from Steppe_Eneolithic, but in the different supervised ADMIXTURE graphics below Maykop_Eneolithic is roughly equivalent to Eneolithic_Steppe (see orange arrow in ADMIXTURE graphic above). This is useful for a simplified analysis, but actual differences between Khvalynsk, Sredni Stog, Afanasevo, Yamna and Corded Ware are probably underestimated in the analyses below, and will become clearer in the future when more ancestral hunter-gatherer populations are added to the analysis.

2. Early Khvalynsk expansion

We have direct data of Khvalynsk-Novodanilovka-like populations thanks to Khvalynsk and Steppe_Eneolithic samples (although I’ve used the latter above to represent the ghost Caucasus population with which Samara_HG admixed).

We also have indirect data. First, there is the PCA with outliers:

PCA-khvalynsk-steppe

Second, we have data from north Pontic Ukraine_Eneolithic samples (see next section).

Third, there is the continuity of late Repin / Afanasevo with Steppe_Eneolithic (see below).

3. Proto-Corded Ware expansion

It is unclear if R1a-M459 subclades were continuously in the steppe and resurged after the Khvalynsk expansion, or (the most likely option) they came from the forested region of the Upper Dnieper area, possibly from previous expansions there with hunter-gatherer pottery.

Supporting the latter is the millennia-long continuity of R1b-V88 and I2a2 subclades in the north Pontic Mesolithic, Neolithic, and Early Eneolithic Sredni Stog culture, until ca. 4500 BC (and even later, during the second half).

Only at the end of the Early Eneolithic with the disappearance of Novodanilovka (and beginning of the steppe ‘hiatus’ of Rassamakin) is R1a to be found in Ukraine again (after disappearing from the record some 2,000 years earlier), related to complex population movements in the north Pontic area.

NOTE. In the PCA, a tentative position of Novodanilovka closer to Anatolia_Neolithic / Dzudzuana ancestry is selected, based on the apparent cline formed by Ukraine_Eneolithic samples, and on the position and ancestry of Sredni Stog, Yamna, and Corded Ware later. A good alternative would be to place Novodanilovka still closer to the Balkan outliers (i.e. Suvorovo), and a source closer to EHG as the ancestry driven by the migration of R1a-M417.

PCA-sredni-stog-steppe

The first sample with steppe ancestry appears only after 4250 BC in the forest-steppe, centuries after the samples with steppe ancestry from the Northern Caucasus and the Balkans, which points to exogamy of expanding R1a-M417 lineages with the remnants of the Novodanilovka population.

steppe-ancestry-admixture-sredni-stog

4. Repin / Early Yamna expansion

We don’t have direct data on early Repin settlers. But we do have a very close representative: Afanasevo, a population we know comes directly from the Repin/late Khvalynsk expansion ca. 3500/3300 BC (just before the emergence of Early Yamna), and which shows fully Steppe_Eneolithic-like ancestry.

afanasevo-admixture

Compared to this eastern Repin expansion that gave Afanasevo, the late Repin expansion to the west ca. 3300 BC that gave rise to the Yamna culture was one of colonization, evidenced by the admixture with north Pontic (Sredni Stog-like) populations, no doubt through exogamy:

PCA-repin-yamna

This admixture is also found (in lesser proportion) in east Yamna groups, which supports the high mobility and exogamy practices among western and eastern Yamna clans, not only with locals:

yamnaya-admixture

5. Corded Ware

Corded Ware represents a quite homogeneous expansion of a late Sredni Stog population, compatible with the traditional location of Proto-Corded Ware peoples in the steppe-forest/forest zone of the Dnieper-Dniester region.

PCA-latvia-ln-steppe

We don’t have a comparison with Ukraine_Eneolithic or Corded Ware samples in Wang et al. (2018), but we do have proximate sources for Abashevo, when compared to the Poltavka population (with which it admixed in the Volga-Ural steppes): Sintashta, Potapovka, Srubna (with further Abashevo contribution), and Andronovo:

sintashta-poltavka-andronovo-admixture

The two CWC outliers from the Baltic show what I thought was an admixture with Yamna. However, given the previous mixture of Eneolithic_Steppe in north Pontic steppe-forest populations, this elevated “steppe ancestry” found in Baltic_LN (similar to west Yamna) seems rather an admixture of Baltic sub-Neolithic peoples with a north Pontic Eneolithic_Steppe-like population. Late Repin settlers also admixed with a similar population during its colonization of the north Pontic area, hence the Baltic_LN – west Yamna similarities.

NOTE. A direct admixture with west Yamna populations through exogamy by the ancestors of this Baltic population cannot be ruled out yet (without direct access to more samples), though, because of the contacts of Corded Ware with west Yamna settlers in the forest-steppe regions.

steppe-ancestry-admixture-latvia

A similar case is found in the Yamna outlier from Mednikarovo south of the Danube. It would be absurd to think that Yamna from the Balkans comes from Corded Ware (or vice versa), just because the former is closer in the PCA to the latter than other Yamna samples. The same error is also found e.g. in the Corded Ware → Bell Beaker theory, because of their proximity in the PCA and their shared “steppe ancestry”. All those theories have been proven already wrong.

NOTE. A similar fallacy is found in potential Sintashta→Mycenaean connections, where we should distinguish statistically that result from an East/West Yamna + Balkans_BA admixture. In fact, genetic links of Mycenaeans with west Yamna settlers prove this (there are some related analyses in Anthrogenica, but the site is down at this moment). To try to relate these two populations (separated more than 1,000 years before Sintashta) is like comparing ancient populations to modern ones, without the intermediate samples to trace the real anthropological trail of what is found…Pure numbers and wishful thinking.

Conclusion

Yamna and Corded Ware show a similar “steppe ancestry” due to convergence. I have said so many times (see e.g. here). This was clear long ago, just by looking at the Y-chromosome bottlenecks that differentiate them – and Tomenable noticed this difference in ADMIXTURE from the supplementary materials in Mathieson et al. (2017), well before Wang et al. (2018).

This different stock stems from (1) completely different ancestral populations + (2) different, long-lasting Y-chromosome bottlenecks. Their similarities come from the two neighbouring cultures admixing with similar populations.

If all this does not mean anything, and each lab was going to support some pre-selected archaeological theories from the 1960s or the 1980s, coupled with outdated linguistic models no matter what – Anthony’s model + Ringe’s glottochronological tree of the early 2000s in the Reich Lab; and worse, Kristiansen’s CWC-IE + Germano-Slavonic models of the 1940s in the Copenhagen group – , I have to repeat my question again:

What’s (so much published) ancient DNA useful for, exactly?

See also

Related

Waves of Palaeolithic ANE ancestry driven by P subclades; new CWC-like Finnish Iron Age

New preprint The population history of northeastern Siberia since the Pleistocene, by Sikora et al. bioRxiv (2018).

Interesting excerpts (emphasis mine; most internal references removed):

ANE ancestry

The earliest, most secure archaeological evidence of human occupation of the region comes from the artefact-rich, high-latitude (~70° N) Yana RHS site dated to ~31.6 kya (…)

The Yana RHS human remains represent the earliest direct evidence of human presence in northeastern Siberia, a population we refer to as “Ancient North Siberians” (ANS). Both Yana RHS individuals were unrelated males, and belong to mitochondrial haplogroup U, predominant among ancient West Eurasian hunter-gatherers, and to Y chromosome haplogroup P1, ancestral to haplogroups Q and R, which are widespread among present-day Eurasians and Native Americans.

Symmetry tests using f4 statistics reject tree-like clade relationships with both Early West Eurasians (EWE; Sunghir) and Early East Asians (EEA; Tianyuan); however, Yana is genetically closer to EWE, despite its geographic location in northeastern Siberia

Using admixture graphs (qpGraph) and outgroup-based estimation of mixture proportions (qpAdm), we find that Yana can be modelled as EWE with ~25% contribution from EEA

Among all ancient individuals, Yana shares the most genetic drift with Mal’ta, and f4 statistics show that Mal’ta shares more alleles with Yana than with EWE (e.g. f4(Mbuti,Mal’ta;Sunghir,Yana) = 0.0019, Z = 3.99). Mal’ta and Yana also exhibit a similar pattern of genetic affinities to both EWE and EEA, consistent with previous studies.The ANE lineage can thus be considered a descendant of the ANS lineage, demonstrating that by 31.6 kya early representatives of this lineage were widespread across northern Eurasia, including far northeastern Siberia.

siberian-samples-haplogroup

Ancient Palaeosiberian

(…) the 9.8 kya Kolyma1 individual, representing a group we term “Ancient Paleosiberians” (AP). Our results indicate that AP are derived from a first major genetic shift observed in the region. Principal component analysis (PCA), outgroup f3-statistics and mtDNA and Y chromosome haplogroups (G1b and Q1a1a, respectively) demonstrate a close affinity between AP and present-day Koryaks, Itelmen and Chukchis, as well as with Native Americans.

For both AP and Native Americans, ANS ancestry appears more closely related to Mal’ta than Yana, therefore rejecting a direct contribution of Yana to later AP or Native American groups.

Lake Baikal Neolithic – Bronze Age

(…) the newly reported genomes from Ust’Belaya and recently published neighbouring Neolithic and Bronze Age sites show a succession of three distinct genetic ancestries over a ~6 ky time span. The earliest individuals show predominantly East Asian ancestry, closely related to the ancient individuals from DGC. In the early Bronze Age (BA), we observe a resurgence of AP ancestry (up to ~50% ancestry fraction), as well as influence of West Eurasian Steppe ANE ancestry represented by the early BA individuals from Afanasievo in the Altai region (~10%) This is consistent with previous reports of gene flow from an unknown ANE-related source into Lake Baikal hunter-gatherers.

Our results suggest a southward expansion of AP as a possible source, which is also consistent with the replacement of Y chromosome lineages observed at Lake Baikal, from predominantly haplogroup N in the Neolithic to haplogroup Q in the BA. Finally, the most recent individual from Ust’Belaya, dated to ~600 years ago, falls along the Neosiberian cline, similar to the ~760 year-old ‘Young Yana’ individual from northeastern Siberia, demonstrating the widespread distribution of Neosiberian ancestry in the most recent epoch.

finnish_ia_palaeosiberian
Genetic structure of ancient northeast Siberians. PCA of ancient individuals projected onto a set of modern Eurasian and American individuals. Abbreviations in group labels: UP – Upper Palaeolithic; LP – Late Palaeolithic; M – Mesolithic; EN – Early Neolithic; MN – Middle Neolithic; LN – Late Neolithic; EBA – Early Bronze Age; LBA – Late Bronze Age; IA – Iron Age; PE – Paleoeskimo; MED – Medieval

Finland Saami

At the western edge of northern Eurasia, genetic and strontium isotope data from ancient individuals at the Levänluhta site documents the presence of Saami ancestry in Southern Finland in the Late Holocene 1.5 kya. This ancestry component is currently limited to the northern fringes of the region, mirroring the pattern observed for AP ancestry in northeastern Siberia. However, while the ancient Saami individuals harbour East Asian ancestry, we find that this is better modelled by DGC rather than AP, suggesting that AP influence was likely restricted to the eastern side of the Urals. Comparison of ancient Finns and Saami with their present-day counterparts reveals additional gene flow over the past 1.6 kya, with evidence for West Eurasian admixture into modern Saami. The ancient Finn from Levänluhta shows lower Siberian ancestry than modern Finns .

EDIT (27 OCT 2018): By comparing the three, I see these are samples published already (at least two) in Lamnidis et al. (2018), but here with added (1) specific radiocarbon dates, (2) comparison with Neosiberian populations and (3) strontium isotope analyses.

Finnish_IA (ca. 350 AD) is probably a Saami-speaking individual, just like the Saami_IA with newly reported radiocarbon dates from Levänluhta ca. 400-600 AD (since Fennic peoples were then likely around the Gulf of Finland).

The conflicting strontium isotope data on marine dietary resources on certain samples from the supplementary material hint at possible external origin of the diet of some of the previously reported (and possibly one newly reported) Saami Iron Age individuals, from some 25-30 km. to the northwest through the river up to hundreds of km. to the southwest of Levänluhta (i.e. the whole coast of the Bothnian Sea). It is unclear why they would prefer an origin of the dietary source in southern Baltic regions instead of some km. to the west, though, unless that’s what they want to propose based on the sample’s admixture…

The coast of the Bothnian Sea (=the northern part of the Baltic Sea, between Sweden and Finland) lay only 25-30 km to the northwest, and accessible to the Iron Age people of the Levänluhta region via the Kyrönjoki river. (…) For individual JA2065/DA236, the low 87Sr/86Sr value (0.71078) would imply an exceptionally heavy reliance on Baltic Sea resources. The δ13C and δ15N values of the individual are near comparable (especially considering within-Baltic latitudinal gradients in δ13C; Torniainen et al. 2017) to the δ13C and δ15N values of a Middle Neolithic population on the Baltic island of Gotland (Eriksson, 2004) interpreted to have subsisted primarily on seals.

These new data on the samples give us some more information than what we already had, because the early date of Finnish_IA implies that there was few East Asian admixture (if any at all) in west Finland during the Roman Iron Age, which pushes still farther forward in time the expected appearance of Siberian ancestry among Saamic (first) and Fennic populations (later). It is unclear whether this East Asian ancestry found in Finnish_IA is actually related to DGC, or it is rather related to the ENA-like ancestry found already in Baltic hunter-gatherers (i.e. in some EHG samples from Karelia), for which Baikal_EN is a good proxy in Lazaridis et al. (2018).

Since Bronze Age and Iron Age samples from Estonia show more Baltic_HG drift compared to Corded Ware samples, it is likely that this supposedly DGC-related ancestry (here considered part of the ‘Siberian ancestry’) is actually an EHG-related ENA component of north-east European hunter-gatherers, with whom Finno-Saamic peoples admixed during the expansion of the Corded Ware culture into Finland.

The paper finds thus increased (probably the actual) Siberian ancestry in modern Finns compared to this Iron Age Saami individual. Coupled with the later Saami Iron Age samples, from between one to three centuries later – showing the start of Siberian ancestry influx – , we can begin to establish when the expansion of Siberian ancestry happened in central Finland, and thus quite likely when the Saami began to expand to the north and east and admix with Palaeo-Laplandic peoples.

siberian-population-expansions
Admixture modelling using qpAdm. Maps showing locations and ancestry proportions of ancient (left) and modern (right) groups.

One sample of haplogroup N1a1a1a1a4a1-M1982, Yana_MED, is found in the Arctic region (north-eastern Yakutia) ca. 1100 AD. Since it is derived from N1a1a1a1a-L392, it might be a surprise for some to find it in a clearly non-Uralic speaking environment at the same time other subclades of this haplogroup were admixing in the west with well-established Finno-Saamic, Volga-Finnic, Ugric, and Samoyedic populations…

On the growing doubts that these data – contradicting the CWC=IE theory – are creating among geneticists (from the supplementary materials):

NOTE. This paper comes from the Copenhagen group, also signed by Kristiansen, one of today’s strongest supporters of this connection

The Proto-Saami language evolved in southern Finland and Karelia in the Early Iron Age, an area now host to Finnish and the closely related Karelian, but with Saami toponyms showing that the latter two languages are intrusive here (Saarikivi 2004). Saami-speaking populations are thought to have retreated to Lapland during the Middle Iron Age (300–800 AD), where it diverged into the modern Saami dialects. Genetically, the northward retreat of the Saami language correlates with the documented decrease of Saami ancestry in Southern Finland between the Iron Age and the modern period (cf. Lamnidis et al. 2018).

On the way to Lapland, the Saami replaced at least two linguistically obscure groups. This can be inferred from 1) an influx of non-Uralic loanwords into Proto-Saami in the Finnish Lakeland area, and 2) an influx of non-Uralic, non-Germanic words into Saami dialects in Lapland (Aikio 2012). Both of these borrowing events imply contact with non-Saami-speaking groups, e.g. non-Uralic-speaking hunter-gatherers that may have left a genetic and linguistic footprint on modern Saami populations.

The linguistic prehistory of Finland thus does not allow for a straightforward interpretation of the genetic data. The detection of East Asian ancestry in the genetically Saami individual is indicative of a population movement from the east (cf. Lamnidis et al. 2018, Rootsi et al. 2007), one that given the affinities with the ~7.6 ky old individuals from the Devil’s Gate Cave may have been a western extension of the Neosiberian turnover. However, it remains unclear whether this gene flow should be associated with the arrival of Uralic speakers, thus providing further support for a Uralic homeland in Eastern Eurasia, or with an earlier immigration of pre-Uralic, so-called “Paleo-Lakelandic” groups.

I think the genetic interpretation is already straightforward, though. We had a sneak peek at how this late admixture with non-Uralians (mainly Palaeo-Lakelandic and Palaeo-Laplandic peoples from Lovozero and related asbestos ware cultures) is going to unfold among expanding Saami-speaking populations thanks to Lamnidis et al. (2018):

saamic-lovozero-pca
PCA plot of 113 Modern Eurasian populations, with individuals from this study projected on the principal components. Uralic speakers are highlighted in light purple. Image modified from Lamnidis et al. (2018)

Also, still no trace of R1a in far East Asia (reported as M17 ca. 5300 BC near Lake Baikal by Moussa et al. 2016), so I still have doubts about my previous assessment that R1a split into M17 (and thus also M417) in Siberia, with those expanding hunter-gatherer pottery.

Related

The Tungusic Ulchi population probably linked to haplogroup C2b1a

ulchi-marital

New paper (behind paywall) Demographic and Genetic Portraits of the Ulchi Population, by Balanovska et al. Russian Journal of Genetics (2018) 54(10):1245–1253.

Interesting excerpts (emphasis mine):

Marital structure. The intensity of interethnic marriages puts the existence of the Ulchi population at risk. The colorful ethnic composition of the Ulchi settlements is reflected in the marriage structure [see featured image]. We found that the proportion of single-ethnic marriages of the Ulchi is on average 51%. The greatest number of such marriages takes place in the village of Bulava. Marriages of Ulchi with Russians are in second place. Marriages with indigenous peoples of the Far East, Nanais, Nivkhs, Evenks, and others, are in third place. Thus, almost half of the Ulchi marriages are with representatives of other nationalities. Such a significant level of interethnic mixing makes it possible to talk about intense processes of assimilation of this indigenous people and puts to the forefront the problem of loss of the unique gene pool of the Ulchi.

Haplogroup C (its branch M48) was genotyped for its five subbranches with markers M86, B470, F13686, B93, and the marker at position 16645386 (GRCh37), which was found by our team for the first time. Variant B93 is rare in the Ulchi, and 14 samples (that is, more than a quarter of the entire gene pool of the Ulchi, Fig. 2) belong to M86 and its subvariants. Therefore, we genotyped STR markers of C-M86 carriers for the Ulchi and neighboring Amur populations and analyzed the relationships of detected haplotypes on the phylogenetic network (Fig. 3, STR haplotypes are available from authors upon request).

(…) On the network, different clusters are associated with different populations: most Mongols belong to F13686, all Evenks of the Amur River region with this haplogroup form a subcluster within F13686, and part of Upper Nanais is the basis of cluster B470.

ulchi-y-chromosome
Frequencies of haplogroups of Y chromosome in the Ulchi population. The nomenclature of haplogroups is given according to [9]. Markers that are not in bold type were not typed, but are ancestral for these nodes.

An estimate of the age of the entire haplogroup C-F12355 obtained from the data of genome-wide sequencing of seven specimens is 2400 ± 500 years (O.P. Balanovsky, unpublished data). That is, the common ancestor of all the studied representatives of various peoples with this haplogroup lived not so long ago, the first millennium BC. The formation time of cluster F13686 is somewhat later: 1990 ± 600 years.

(…) obvious traces of the interaction of the gene pool of the Ulchi with neighboring and remote peoples of the Far East and Central Asia in the time range of the last one to three thousand years were revealed. This shows that the results of work [4] on the similarity of the gene pool of the ancient (age of 7500 years) Neolithic genomes of the Amur River region to the Ulchi probably indicate not the uniqueness of the Ulchi, but the fact that this ancient gene pool was preserved in a vast circle of populations of the Far East interwoven with gene flows both with each other and, to a lesser extent, with populations of Central Asia.

The expansion of C2b1a2a-M86 (among many basal C2-M217 samples) is thus possibly associated with the spread of Tungusic, which puts C2b1a at the root of the Micro-Altaic expansion, with a formation date ca. 12700 BC, TMRCA 12500 BC (and not only Mongolian). This shows that Micro-Altaic is connected with a local population which shows a clear continuity since at least 3500 BC. This, however, tells us little about the origin of the language.

See also the recent ISBA presentation on the Houtaomuga site, Neolithic transition in Northeast Asia; and also Bronze Age population dynamics and rise of dairy pastoralism in Mongolia, Impact of colonization in north-eastern Siberia

That leaves the ancestral N lineages found among Far East Asians as Palaeo-Siberian in origin, and their late expansions to the west not particularly linked with any of the known Palaeo-Siberian ethnolinguistic groups, let alone a supposed “Uralo-Altaic” language…

Related

Resurge of local populations in the final Corded Ware culture period from Poland

poland-kujawy

Open access A genomic Neolithic time transect of hunter-farmer admixture in central Poland, by Fernandes et al. Scientific Reports (2018).

Interesting excerpts (emphasis mine, stylistic changes):

Most mtDNA lineages found are characteristic of the early Neolithic farmers in south-eastern and central Europe of the Starčevo-Kőrös-Criş and LBK cultures. Haplogroups N1a, T2, J, K, and V, which are found in the Neolithic BKG, TRB, GAC and Early Bronze Age samples, are part of the mitochondrial ‘Neolithic package’ (which also includes haplogroups HV, V, and W) that was introduced to Europe with farmers migrating from Anatolia at the onset of the Neolithic17,31.

A noteworthy proportion of Mesolithic haplogroup U5 is also found among the individuals of the current study. The proportion of haplogroup U5 already present in the earliest of the analysed Neolithic groups from the examined area differs from the expected pattern of diversity of mtDNA lineages based on a previous archaeological view and on the aDNA findings from the neighbouring regions which were settled by post-Linear farmers similar to BKG at that time. A large proportion of Mesolithic haplogroups in late-Danubian farmers in Kuyavia was also shown in previous studies concerning BKG samples based on mtDNA only, although these frequencies were derived on the basis of very small sample sizes.

y-dna-poland

A significant genetic influence of HG populations persisted in this region at least until the Eneolithic/Early Bronze Age period, when steppe migrants arrived to central Europe. The presence of two outliers from the middle and late phases of the BKG in Kuyavia associated with typical Neolithic burial contexts provides evidence that hunter-farmer contacts were not restricted to the final period of this culture and were marked by various episodes of interaction between two societies with distinct cultural and subsistence differences.

The identification of both mitochondrial and Y-chromosome haplogroup lineages of Mesolithic provenance (U5 and I, respectively) in the BKG support the theory that both male and female hunter-gatherers became part of these Neolithic agricultural societies, as has been reported for similar cases from the Carpathian Basin, and the Balkans. The identification of an individual with WHG affinity, dated to ca. 4300 BCE, in a Middle Neolithic context within a BKG settlement, provides direct evidence for the regional existence of HG enclaves that persisted and coexisted at least for over 1000 years, from the arrival of the LBK farmers ca. 5400 BCE until ca. 4300 BCE, in proximity with Neolithic settlements, but without admixing with their inhabitants.

poland-pca
Principal component analysis with modern populations greyed out on the background (top), ADMIXTURE results with K = 10 with samples from this study amplified (bottom).

The analysis of two Late Neolithic cultures, the GAC and CWC, shows that steppe ancestry was present only among the CWC individuals analysed, and that the single GAC individual had more WHG ancestry than previous local Neolithic individuals. (…) The CWC’s affinity to WHG, however, contrasts with results from published CWC individuals that identified steppe ancestry related to Yamnaya as the major contributor to the CWC genomes, while here we report also substantial contributions from WHG that could relate to the late persistence of pockets of WHG populations, as supported by the admixture results of N42 and the finding of the 4300-year-old N22 HG individual. These results agree with archaeological theories that suggest that the CWC interaction with incoming steppe cultures was complex and that it varied by region.

Some comments

About the analyzed CWC samples, it is remarkable that, even though they are somehow related to each other, they do not form a tight cluster. Also, their Y-DNA (I2a), and this:

When compared to previously published CWC data, our CWC group (not individuals) is genetically significantly closer to WHG than to steppe individuals (Z = −4.898), a result which is in contrast with those for CWC from Germany (Z = 2.336), Estonia (Z = 0.555), and Latvia (Z = 1.553).

ancestry-proportions-poland
Ancestry proportions based on qpAdm. Visual representation of the main results presented in Supplementary Table S5. Populations from this study marked with an asterisk. Values and populations in brackets show the nested model results marked in green in Supplementary Table S5.

Włodarczak (2017) talks about the CWC period in Poland after ca. 2600 BC as a time of emergence of an allochthnous population, marked by the rare graves of this area, showing infiltrations initially mainly from Lesser Poland, and later (after 2500 BC) from the western Baltic zone.

Since forest sub-Neolithic populations would have probably given more EHG to the typical CWC population, these samples support the resurge of ‘local’ pockets of GAC- or TRB-like groups with more WHG (and also Levant_Neolithic) ancestry.

The known presence of I2a2a1b lineages in GAC groups in Poland also supports this interpretation, and the subsistence of such pockets of pre-steppe-like populations is also seen with the same or similar lineages appearing in comparable ‘resurge’ events in Central Europe, e.g. in samples from the Únětice and Tumulus culture.

About the Bronze Age sample, we have at last official confirmation of haplogroup R1a1a (sadly no subclade*) at the very beginning of the Trzciniec period – in a region between western (Iwno) and eastern (Strzyżów) groups related to Mierzanowice – , which has to be put in relation with the samples from the final Trzciniec period in the Baltic published in Mittnik et al. (2018).

EDIT (8 OCT 2018): More specific subclades have been published, including a R1a-Z280 lineage for the Bronze Age sample (see spreadsheet).

This confirms the early resurge of R1a-Z645 (probably R1a-Z282) lineages at the core of the developing East European Bronze Age, a province of the European Bronze Age that emerged from evolving Bell Beaker groups in Poland.

bell-beakers-poland-kujawy
Arrival of Bell Beakers in Poland after ca. 2400 BC, and their origin in other BBC centres (Czebreszuk and Szmyt 2011).

I don’t have any hope that the Balto-Slavic evolution through BBC Poland → Mierzanowice/Iwno → Trzciniec → Lusatian cultures is going to be confirmed any time soon, until we have a complete trail of samples to follow all the way to historic Slavs of the Prague culture. However, I do think that the current data on central-east Europe – and the recent data we are receiving from north-east Europe and the Iranian steppes, at odds with the Indo-Slavonic alternative – supports this model.

I guess that, in the end, similar to how the Yamna vs. Corded Ware question is being solved, the real route of expansion of Proto-Balto-Slavic (supposedly spoken ca. 1500-1000 BC) is probably going to be decided by the expansion of either R1a-M458 (from the west) or R1a-Z280 lineages (from the east), because the limited precision of genetic data and analyses available today are going to show ‘modern Slavic’-like populations from the whole eastern half of Europe for the past 4,000 years…

Related

Corded Ware—Uralic (II): Finno-Permic and the expansion of N-L392/Siberian ancestry

finno-ugric-samoyedic

This is the second of four posts on the Corded Ware—Uralic identification:

I read from time to time that “we have not sampled Uralic speakers yet”, and “we are waiting to see when Uralic-speaking peoples are sampled”. Are we, though?

Proto-language homelands are based on linguistic data, such as guesstimates for dialectal evolution, loanwords and phonetic changes for language contacts, toponymy for ancient territories, etc. depending on the available information. The trace is then followed back, using available archaeological data, from the known historic speakers and territory to the appropriate potential prehistoric cultures. Only then can genetic analyses help us clarify the precise prehistoric population movements that better fit the models.

uralic-language-family
The traditional family tree of the Uralic branches. Kallio (2014)

The linguistic homeland

We thought – using linguistic guesstimates and fitting prehistoric cultures and their expansion – that Yamna was the Late Proto-Indo-European culture, so when Yamna was sampled, we had Late Proto-Indo-Europeans sampled. Simple deduction.

We thought that north-eastern Europe was a Uralic-speaking area during the Neolithic:

  • For those supporting a western continuity (and assuming CWC was Indo-European), the language was present at least since the Comb Ware culture, potentially since the Mesolithic.
  • For those supporting a late introduction into Finland, Uralic expanded the latest with Abashevo-related movements after its incorporation of Volosovo and related hunter-gatherers.

The expansion to the east must have happened through progressive infiltrations with Seima-Turbino / Andronovo-related expansions.

uralic-time-space
Some datings for the traditional proto-stages from Uralic to Finnic. Kallio (2014).

Finding the linguistic homeland going backwards can be described today as follows:

I. Proto-Fennic homeland

Based on the number of Baltic loanwords, not attested in the more eastern Uralic branches (and reaching only partially Mordvinic), the following can be said about western Finno-Permic languages (Junttila 2014):

The Volga-Kama Basin lies still too far east to be included in a list of possible contact locations. Instead, we could look for the contact area somewhere between Estonia in the west and the surroundings of Moscow in the east, a zone with evidence of Uralic settlement in the north and Baltic on the south side.

The only linguistically well-grounded version of the Stone Age continuation theory was presented by Mikko Korhonen in 1976. Its validity, however, became heavily threatened when Koivulehto 1983a-b proved the existence of a Late Proto-Indo-European or Pre-Baltic loanword layer in Saami, Finnic, and Mordvinic. Since this layer must precede the Baltic one and it was presumably acquired in the Baltic Sea region, Koivulehto posited it on the horizon of the Battle Axe period. This forces a later dating for the Baltic–Finnic contacts.

Today the Battle Axe culture is dated at 3200 to 3000 BC, a period far too remote to correspond linguistically with Proto-Baltic (Kallio 1998a).

Since the Baltic contacts began at a very initial phase of Proto-Finnic, the language must have been relatively uniform at that time. Hence, if we consider that the layer of Baltic loanwords may have spread over the Gulf of Finland at that time, we could also insist that the whole of the Proto-Finnic language did so.

migration-theory
Prehistoric Balts as the southern neighbours of Proto-Finnic speakers. 1 = The approximated area of Proto-Uralic. 2 = The approximated area of Finnic during the Iron Age. 3 = The area of ancient Baltic hydronyms. 4 = The area of Baltic languages in about 1200 AD. 5 = The problem: When did Uralic expand westwards and when did it meet Baltic? Junntila (2012).

II. Proto-Finno-Saamic homeland

The evidence of continued Palaeo-Germanic loanwords (from Pre- to Proto-Germanic stages) is certainly the most important data to locate the Finno-Saamic homeland, and from there backwards into the true Uralic homeland. Following Kallio (2017):

(…) the loanword evidence furthermore suggests that the ancestors of Finnic and Saamic had at least phonologically remained very close to Proto-Uralic as late as the Bronze Age (ca. 1700–500 BC). In particular, certain loanwords, whose Baltic and Germanic sources point to the first millennium BC, after all go back to the Finno-Saamic proto-stage, which is phonologically almost identical to the Uralic proto-stage (see especially the table in Sammallahti 1998: 198–202). This being the case, Dahl’s wave model could perhaps have some use in Uralic linguistics, too.

The presence of Pre-Germanic loanwords points rather to the centuries around the turn of the 2nd – 1st millennium BC or earlier. Proto-Germanic words must have been borrowed before the end of Germanic influence in the eastern Baltic at the beginning of the Iron Age, which sets a clear terminus ante quem ca. 800 BC.

The arrival of Bell Beaker peoples in Scandinavia ca. 2350 BC, heralding the formation of the Dagger Period, as well as the development of Pre-Germanic in common with Finnic-like populations point to the late 3rd / early 2nd millennium BC as the first time of close interaction through the Baltic region.

III. Proto-Uralic homeland

(…) the earliest Indo-European loanwords in the Uralic languages (…) show that Proto-Uralic cannot have been spoken much earlier than Proto-Indo-European dated about 3500 BC (Koivulehto 2001: 235, 257). As the same loanword evidence naturally also shows that the Uralic and Indo-European homelands were not located far from one another, the Uralic homeland can most likely be located in the Middle and Upper Volga region, right north of the Indo-European homeland*. From the beginning of the Subneolithic period about 5900 BC onwards, this region was an important innovation centre, from where several cultural waves spread to the Finnish Gulf area, such as the Sperrings Ware wave about 4900 BC, the Combed Ware wave about 3900 BC, and the Netted Ware wave about 1900 BC (Carpelan & Parpola 2001: 78–90).

The mainstream position is nowadays trying to hold together the traditional views of Corded Ware as Indo-European, and a Uralic Fennoscandia during the Bronze Age.

The following is an example of how this “Volosovo/Forest Zone hunter-gatherer theory” of Uralic origins looks like, as a ‘mixture’ of cultures and languages that benefits from the lack of genetic data for certain regions and periods (taken from Parpola 2018):

asbestos-ware
The extent of Typical Comb Ware (TCW), Asbestos- and Organic-tempered Wares (AOW) and Volosovo and Garino-Bor cultures; areas with deposits of native copper in Karelia and copperbearing sandstone in Volga-Kama-area are marked dark gray (after Zhuravlev 1977; Krajnov 1987; Nagovitsyn 1987; Chernykh 1992; Carpelan 1999; Zhul´nikov 1999). From Nordqvist et al. (2012).

The Corded Ware (or Battle Axe) culture intruded into the Eastern Baltic and coastal Finland already around 3100 BCE. The continuity hypothesis maintains that the early Proto-Finnic speakers of the coastal regions, who had come to Finland in the 4th millennium BCE with the Comb-Pitted Ware, coexisted with the Corded Ware newcomers, gradually adopting their pastoral culture and with it a number of NW-IE loanwords, but assimilating the immigrants linguistically.

The fusion of the Corded Ware and the local Comb-Pitted Ware culture resulted into the formation of the Kiukais culture (c. 2300–1500) of southwestern Finland, which around 2300 received some cultural impulses from Estonia, manifested in the appearance of the Western Textile Ceramic (which is different from the more easterly Textile Ceramic or Netted Ware, and which is first attested in Estonia c. 2700 BCE, cf. Kriiska & Tvauri 2007: 88), and supposed to have been accompanied by an influx of loanwords coming from Proto-Baltic. At the same time, the Kiukais culture is supposed to have spread the custom of burying chiefs in stone cairns to Estonia.

The coming of the Corded Ware people and their assimilation created a cultural and supposedly also a linguistic split in Finland, which the continuity hypothesis has interpreted to mean dividing Proto-Saami-Finnic unity into its two branches. Baltic Finnic, or simply Finnic, would have emerged in the coastal regions of Finland and in the northern East Baltic, while preforms of Saami would have been spoken in the inland parts of Finland.

The Nordic Bronze Age culture, correlated above with early Proto-Germanic, exerted a strong influence upon coastal Finland and Estonia 1600–700 BCE. Due to this, the Kiukais culture was transformed into the culture of Paimio ceramics (c. 1600–700 BCE), later continued by Morby ceramics (c. 700 BCE – 200 CE). The assumption is that clear cultural continuity was accompanied by linguistic continuity. Having assimilated the language of the Germanic traders and relatively few settlers of the Bronze Age, the language of coastal Finland is assumed to have reached the stage of Proto-Finnish at the beginning of the Christian era. In Estonia, the Paimio ceramics have a close counterpart in the contemporaneous Asva ceramics.

Eastern homelands?

I will not comment on Siberian or Central Asian homeland proposals, because they are obviously not mainstream, still less today when we know that Uralic was certainly in contact with Proto-Indo-European, and then with Pre- and Proto-Indo-Iranian, as supported even by the Copenhagen group in Damgaard et al. (2018).

This is what Kallio (2017) has to say about the agendas behind such proposals:

Interestingly, the only Uralicists who generally reject the Central Russian homeland are the Russian ones who prefer the Siberian homeland instead. Some Russians even advocate that the Central Russian homeland is only due to Finnish nationalism or, as one of them put it a bit more tactfully, “the political and ideological situation in Finland in the first decades of the 20th century” (Napolskikh 1995: 4).

Still, some Finns (and especially those who also belong to the “school who wants it large and wants it early”) simultaneously advocate that exactly the same Central Russian homeland is due to Finnlandisierung (Wiik 2001: 466).

Hence, for those of you willing to learn about fringe theories not related to North-Eastern Europe, you also have then the large and early version of the Uralic homeland, with Wiik’s Palaeolithic continuity of Uralic peoples spread over all of eastern and central Europe (hence EHG and R1a included):

atlantic-finnic-theory
Palaeolithic boat peoples and Finno-Ugric. Source

These fringe Finnish theories look a lot like the Corded Ware expansion… Better not go the Russian or Finnish nationalist ways? Agreed then, let’s discuss only rational proposals based on current data.

The archaeological homeland

For a detailed account of the Corded Ware expansion with Battle Axe, Fatyanovo-Balanovo, and Abashevo groups into the area, you can read my recent post on the origin of R1a-Z645.

1. Textile ceramics

During the 2nd millennium BC, textile impressions appear in pottery as a feature across a wide region, from the Baltic area through the Volga to the Urals, in communities that evolve from late Corded Ware groups without much external influence.

While it has been held that this style represents a north-west expansion from the Volga region (with the “Netted Ware” expansion), there are actually at least two original textile styles, one (earlier) in the Gulf of Finland, common in the Kiukainen pottery, which evolves into the Textile ware culture proper, and another which seems to have an origin in the Middle Volga region to the south-east.

The Netted ware culture is the one that apparently expands into inner Finland – a region not densely occupied by Corded Ware groups until then. There are, however, no clear boundaries between groups of both styles; textile impressions can be easily copied without much interaction or population movement; and the oldest textile ornamentation appeared on the Gulf of Finland. Hence the tradition of naming all as groups of Textile ceramics.

textile-ware-cultures
Maximum distribution of Textile ceramics during the Bronze Age (ca. 2000-800 BC). Asbestos-tempered ware lies to the north (and is also continued in western Fennoscandia).

The fact that different adjacent groups from the Gulf of Finland and Forest Zone share similar patterns making it very difficult to differentiate between ‘Netted Ware’ or ‘Textile Ware’ groups points to:

  • close cultural connections that are maintained through the Gulf of Finland and the Forest Zone after the evolution of late Corded Ware groups; and
  • no gross population movements in the original Battle Axe / Fatyanovo regions, except for the expansion of Netted Ware to inner Finland, Karelia, and the east, where the scattered Battle Axe finds and worsening climatic conditions suggest most CWC settlements disappeared at the end of the 3rd millennium BC and recovered only later.

NOTE. This lack of population movement – or at least significant replacement by external, non-CWC groups – is confirmed in genetic investigation by continuity of CWC-related lineages (see below).

The technology present in Textile ceramics is in clear contrast to local traditions of sub-Neolithic Lovozero and Pasvik cultures of asbestos-tempered pottery to the north and east, which point to a different tradition of knowledge and learning network – showing partial continuity with previous asbestos ware, since these territories host the main sources of asbestos. We have to assume that these cultures of northern and eastern Fennoscandia represent Palaeo-European (eventually also Palaeo-Siberian) groups clearly differentiated from the south.

The Chirkovo culture (ca. 1800-700 BC) forms on the middle Volga – at roughly the same time as Netted Ware formed to the west – from the fusion of Abashevo and Balanovo elites on Volosovo territory, and is also related (like Abashevo) to materials of the Seima-Turbino phenomenon.

Bronze Age ethnolinguistic groups

In the Gulf of Finland, Kiukainen evolves into the Paimio ceramics (in Finland) — Asva Ware (in Estonia) culture, which lasts from ca. 1600 to ca. 700 BC, probably representing an evolving Finno-Saamic community, while the Netted Ware from inner Finland (the Sarsa and Tomitsa groups) and the groups from the Forest Zone possibly represent a Volga-Finnic community.

NOTE. Nevertheless, the boundaries between Textile ceramic groups are far from clear, and inner Finland Netted Ware groups seem to follow a history different from Netted Ware groups from the Middle and Upper Volga, hence they could possibly be identified as an evolving Pre-Saamic community.

Based on language contacts, with Early Baltic – Early Finnic contacts starting during the Iron Age (ca. 500 BC onwards), this is a potential picture of the situation at the end of this period, when Germanic influence on the coast starts to fade, and Lusatian culture influence is stronger:

aikio-finnic-saamic
The linguistic situation in Lapland and the northern Baltic Sea Area in the Early Iron Age prior to the expansion of Saami languages; the locations of the language groups are schematic. The black line indicates the distribution of Saami languages in the 19th century, and the gray line their approximate maximal distribution before the expansion of Finnic. Aikio (2012)

The whole Finno-Permic community remains thus in close contact, allowing for the complicated picture that Kallio mentions as potentially showing Dahl’s wave model for Uralic languages.

Genetic data shows a uniform picture of these communities, with exclusively CWC-derived ancestry and haplogroups. So in Mittnik et al. (2018) all Baltic samples show R1a-Z645 subclades, while the recent session on Estonian populations in ISBA 8 (see programme in PDF) clearly states that:

[Of the 24 Bronze Age samples from stone-cist graves] all 18 Bronze Age males belong to R1a.

Regarding non-Uralic substrates found in Saami, supposedly absorbed during the expansion to the north (and thus representing languages spoken in northern Fennoscandia during the Bronze Age) this is what Aikio (2012) has to say:

The Saami substrate in the Finnish dialects thus reveals that also Lakeland Saami languages had a large number of vocabulary items of obscure origin. Most likely many of these words were substrate in Lakeland Saami, too, and ultimately derive from languages spoken in the region before Saami. In some cases the loan origin of these words is obvious due to their secondary Proto-Saami vowel combinations such as *ā–ë in *kāvë ‘bend; small bay’ and *šāpšë ‘whitefish’. This substrate can be called ‘Palaeo-Lakelandic’, in contrast to the ‘Palaeo-Laplandic’ substrate that is prominent in the lexicon of Lapland Saami. As the Lakeland Saami languages became extinct and only fragments of their lexicon can be reconstructed via elements preserved in Finnish place-names and dialectal vocabulary, we are not in a position to actually study the features of this Palaeo-Lakelandic substrate. Its existence, however, appears evident from the material above.

If we wanted to speculate further, based on the data we have now, it is very likely that two opposing groups will be found in the region:

A) The central Finnish group, in this hypothesis the Palaeo-Lakelandic group, made up of the descendants of the Mesolithic pioneers of the Komsa and Suomusjärvi cultures, and thus mainly Baltic HG / Scandinavian HG ancestry and haplogroups I / R1b(xM269) (see more on Scandinavian HG).

siberian-ancestry-map
Frequency map of the so-called ‘Siberian’ component. From Tambets et al. (2018).

B) Lapland and Kola were probably also inhabited by similar Mesolithic populations, until it was eventually assimilated by expanding Siberian groups (of Siberian ancestry and N1c-L392 lineages) from the east – entering the region likely through the Kola peninsula – , forming the Palaeo-Laplandic group, which was in turn later replaced by expanding Proto-Saamic groups.

Siberian ancestry appears first in Fennoscandia at Bolshoy Oleni Ostrov ca. 1520 BC, with haplogroup N1c-L392 (2 samples, BOO002 and BOO004), and with Siberian ancestry. This is their likely movement in north-eastern Europe, from Lamnidis et al (2018):

The large Siberian component in the Bolshoy individuals from the Kola Peninsula provides the earliest direct genetic evidence for an eastern migration into this region. Such contact is well documented in archaeology, with the introduction of asbestos-mixed Lovozero ceramics during the second millenium BC, and the spread of even-based arrowheads in Lapland from 1,900 BCE. Additionally, the nearest counterparts of Vardøy ceramics, appearing in the area around 1,600-1,300 BCE, can be found on the Taymyr peninsula, much further to the east. Finally, the Imiyakhtakhskaya culture from Yakutia spread to the Kola Peninsula during the same period.

saamic-lovozero-pca
PCA plot of 113 Modern Eurasian populations, with individuals from this study projected on the principal components. Uralic speakers are highlighted in light purple. Image modified from Lamnidis et al. (2018)

Obviously, these groups of asbestos-tempered ware are not connected to the Uralic expansion. From the same paper:

The fact that the Siberian genetic component is consistently shared among Uralic-speaking populations, with the exceptions of Hungarians and the non-Uralic speaking Russians, would make it tempting to equate this component with the spread of Uralic languages in the area. However, such a model may be overly simplistic. First, the presence of the Siberian component on the Kola Peninsula at ca. 4000 yBP predates most linguistic estimates of the spread of Uralic languages to the area. Second, as shown in our analyses, the admixture patterns found in historic and modern Uralic speakers are complex and in fact inconsistent with a single admixture event. Therefore, even if the Siberian genetic component partly spread alongside Uralic languages, it likely presented only an addition to populations carrying this component from earlier.

2. The Early Iron Age

The Ananino culture appears in the Vyatka-Kama area, famed for its metallurgy, with traditions similar to the North Pontic area, by this time developing Pre-Sauromatian traditions. It expanded to the north in the first half of the first millennium BC, remaining in contact with the steppes, as shown by the ‘Scythian’ nature of its material culture.

NOTE. The Ananino culture can be later followed through its zoomorphic styles into Iron Age Pjanoborskoi and Gljadenovskoi cultures, later to Ural-Siberian Middle Age cultures – Itkuska, Ust’-Poluiska, Kulaiska cultures –, which in turn can be related as prototypes of medieval Permian styles.

ananino-culture-homeland
Territory of (early and maximum) Ananino material culture. Vasilyev (2002).

At the same time as the Ananino culture begins to expand ca. 1000 BC, the Netted Ware tradition from the middle Oka expanded eastwards into the Oka-Vyatka interfluve of the middle Volga region, until then occupied by the Chirkovo culture. Eventually the Akozino or Akhmylovo group (ca. 800-300 BC) emerged from the area, showing a strong cultural influence from the Ananino culture, by that time already expanding into the Cis-Urals region.

The Akozino culture remains nevertheless linked to the western Forest Zone traditions, with long-ranging influences from as far as the Lusatian culture in Poland (in metallurgical techniques), which at this point is also closely related with cultures from Scandinavia (read more on genetics of the Tollense Valley).

malar-celts-ananino
Mälar celts and molds for casting (a) and the main distribution area (в) of Mälar-type celts of the Mälar type in the Volga-Kama region (according to Kuzminykh 1983: figure 92) and Scandinavia (according to Baudou 1960: Karte 10); Ananino celts and molds for casting (б) and the main distribution area (г) of the distribution of the celts of the Ananino type in the Volga-Kama area (according to Kuzminykh 1983: figure 9); dagger of Ananino type (д).Map from (Yushkova 2010)

Different materials from Akozino reach Fennoscandia late, at the end of the Bronze Age and beginning of the Early Iron Age, precisely when the influence of the Nordic Bronze Age culture on the Gulf of Finland was declining.

This is a period when Textile ceramic cultures in north-eastern Europe evolve into well-armed chiefdom-based groups, with each chiefdom including thousands or tens of thousands, with the main settlements being hill forts, and those in Fennoscandia starting ca. 1000-400 BC.

Mälar-type celts and Ananino-type celts appear simultaneously in Fennoscandia and the Forest Zone, with higher concentrations in south-eastern Sweden (Mälaren) and the Volga-Kama region, supporting the existence of a revived international trade network.

akozino-malar-axes-fennoscandia
Distribution of the Akozino-Mälar axes according to Sergej V. Kuz’minykh (1996: 8, Abb. 2).

The Paimio—Asva Ware culture evolves (ca. 700-200 BC) into the Morby (in Finland) — Ilmandu syle (in Estonia, Latvia, and Mälaren) culture. The old Paimio—Asva tradition continues side by side with the new one, showing a clear technical continuity with it, but with ornamentation compared to the Early Iron Age cultures of the Upper Volga area. This new south-eastern influence is seen especially in:

  • Akozino-Mälar axes (ca. 800-500 BC): introduced into the Baltic area in so great numbers – especially south-western Finland, the Åland islands, and the Mälaren area of eastern Sweden – that it is believed to be accompanied by a movement of warrior-traders of the Akozino-Akhmylovo culture, following the waterways that Vikings used more than a thousand years later. Rather than imports, they represent a copy made with local iron sources.
  • Tarand graves (ca. 500 BC – AD 400): these ‘mortuary houses’ appear in the coastal areas of northern and western Estonia and the islands, at the same time as similar graves in south-western Finland, eastern Sweden, northern Latvia and Courland. Similar burials are found in Akozino-Akhmylovo, with grave goods also from the upper and middle Volga region, while grave goods show continuity with Textile ware.

The use of asbestos increases in mainland Finnish wares with Kjelmøy Ware (ca. 700 BC – AD 300), which replaced the Lovozero Ware; and in the east in inner Finland and Karelia with the Luukonsaari and Sirnihta wares (ca. 700-500 BC – AD 200), where they replaced the previous Sarsa-Tomitsa ceramics.

The Gorodets culture appears during the Scythian period in the forest-steppe zone north and west of the Volga, shows fortified settlements, and there are documented incursions of Gorodets iron makers into the Samara valley, evidenced by deposits of their typical pottery and a bloom or iron in the region.

Iron Age ethnolinguistic groups

According to (Koryakova and Epimakhov 2007):

It is commonly accepted by archaeology, ethnography, and linguistics that the ancestors of the Permian peoples (the Udmurts, Komi-Permians, and Komi-Zyryans) left the sites of Ananyino cultural intercommunity.

NOTE. For more information on the Late Metal Ages and Early Medieval situation of Finno-Ugric languages, see e.g. South-eastern contact area of Finnic languages in the light of onomastics (Rahkonen 2013).

finno-saamic-mordvin
Yakhr-, -khra, yedr-, -dra and yer-/yar, -er(o), -or(o) names of lakes in Central and North Russia and the possible boundary of the proto-language words *jäkra/ä and *järka/ä. Rahkonen (2011)

Certain innovations shared between Proto-Fennic (identified with the Gulf of Finland) and Proto-Mordvinic (from the Gorodets culture) point to their close contact before the Proto-Fennic expansion, and thus to the identification of Gorodets as Proto-Mordvinic, hence Akozino as Volgaic (Parpola 2018):

  • the 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 *kümmen.
  • The presence of loanwords of non-Uralic origin, related to farming and trees, potentially Palaeo-European in nature (hence possibly from Siberian influence in north-eastern Europe).
ananino-textile-ware-cultures
Map of archaeological cultures in north-eastern Europe ca. 8th-3rd centuries BC. [The Mid-Volga Akozino group not depicted] Shaded area represents the Ananino cultural-historical society. Purple area show likely zones of predominant Siberian ancestry and N1c-L392 lineages. Blue areas likely zones of predominant CWC ancestry and R1a-Z645 lineages. Fading purple arrows represent likely stepped movements of haplogroup N1c-L392 for centuries (Siberian → Ananino → Akozino → Fennoscandia), found eventually in tarand graves. Blue arrows represent eventual expansions of Fennic and (partially displaced) Saamic. Modified image from Vasilyev (2002).

The introduction of a strongly hierarchical chiefdom system can quickly change the pre-existing social order and lead to a major genetic shift within generations, without a radical change in languages, as shown in Sintashta-Potapovka compared to the preceding Poltavka society (read more about Sintashta).

Fortified settlements in the region represented in part visiting warrior-traders settled through matrimonial relationships with local chiefs, eager to get access to coveted goods and become members of a distribution network that could guarantee them even military assistance. Such a system is also seen synchronously in other cultures of the region, like the Nordic Bronze Age and Lusatian cultures (Parpola 2013).

The most likely situation is that N1c subclades were incorporated from the Circum-Artic region during the Anonino (Permic) expansion to the north, later emerged during the formation of the Akozino group (Volgaic, under Anonino influence), and these subclades in turn infiltrated among the warrior traders that spread all over Fennoscandia and the eastern Baltic (mainly among Fennic, Saamic, Germanic, and Balto-Slavic peoples), during the age of hill forts, creating alliances partially based on exogamy strategies (Parpola 2013).

Over the course of these events, no language change is necessary in any of the cultures involved, since the centre of gravity is on the expanding culture incorporating new lineages:

  • first on the Middle Volga, when Ananino expands to the north, incorporatinig N1c lineages from the Circum-Artic region.
  • then with the expansion of the Akozino-Akhmylovo culture into Ananino territory, admixing with part of its population;
  • then on the Baltic region, when materials are imported from Akozino into Fennoscandia and the eastern Baltic (and vice versa), with local cultures being infiltrated by foreign (Akozino) warrior-traders and their materials;
  • and later with the different population movements that led eventually to a greater or lesser relevance of N1c in modern Finno-Permic populations.

To argue that this infiltration and later expansion of lineages changed the language in one culture in one of these events seems unlikely. To use this argument of “opposite movement of ethnic and language change” for different successive events, and only on selected regions and cultures (and not those where the greatest genetic and cultural impact is seen, like e.g. Sweden for Akozino materials) is illogical.

NOTE. Notice how I write here about “infiltration” and “lineages”, not “migration” or “populations”. To understand that, see below the next section on autosomal studies to compare Bronze Age, Iron Age, Medieval and Modern Estonians, and see how little the population of Estonia (homeland of Proto-Fennic and partially of Proto-Finno-Saamic) has changed since the Corded Ware migrations, suggesting genetic continuity and thus mostly close inter-regional and intra-regional contacts in the Forest Zone, hence a very limited impact of the absorbed N1c lineages (originally at some point incorporated from the Circum-Artic region). You can also check on the most recent assessment of R1a vs. N1c in modern Uralic populations.

Iron Age and later populations

From the session on Estonian samples on ISBA 8, by Tambets et al.:

[Of the 13 samples from the Iron Age tarand-graves] We found that the Iron Age individuals do in fact carry chrY hg N3 (…) Furthermore, based on their autosomal data, all of the studied individuals appear closer to hunter-gatherers and modern Estonians than Estonian CWC individuals do.

EDIT (16 OCT) A recent abstract with Saag as main author (Tambets second) cites 3 out of 5 sampled Iron Age individuals as having haplogroup N3.

EDIT (28 OCT): Notice also the appearance of N1a1a1a1a1a1a1-L1025 in Lithuania (ca. 300 AD), from Damgaard (Nature 2018); the N1c sample of the Krivichi Pskov Long Barrows culture (ca. 8th-10th c. AD), and N1a1a1a1a1a1a7-Y4341 among late Vikings from Sigtuna (ca. 10th-12th c. AD) in Krzewinska (2018).

estonian-pca
PCA of Estonian samples from the Bronze Age, Iron Age and Medieval times. Tambets et al. (2018, upcoming).

Looking at the plot, the genetic inflow marking the change from the Bronze Age to the Iron Age looks like an obvious expansion of nearby peoples with CWC-related ancestry, i.e. likely from the south-east, near the Middle Volga, where influence of steppe peoples is greater (hence likely Akozino) into a Proto-Fennic population already admixed (since the arrival of Corded Ware groups) with Comb Ware-like populations.

All of these groups were probably R1a-Z645 (likely R1a-Z283) since the expansion of Corded Ware peoples, with an introduction of some N1c lineages precisely during this Iron Age period. This infiltration of N1c-L392 with Akozino is obviously not directly related to Siberian cultures, given what we know about the autosomal description of Estonian samples.

Rather, N1c-L392 lineages were likely part of the incoming (Volgaic) Akozino warrior-traders, who settled among developing chiefdoms based on hill fort settlements of cultures all over the Baltic area, and began to appear thus in some of the new tarand graves associated with the Iron Age in north-eastern Europe.f

A good way to look at this is to realize that no new cluster appears compared to the data we already have from Baltic LN and BA samples from Mittnik et al. (2018), so the Estonian BA and IA clusters must be located (in a proper PCA) in the cline from Pit-Comb Ware culture through Baltic BA to Corded Ware groups:

baltic-samples
PCA and ADMIXTURE analysis reflecting three time periods in Northern European prehistory. a Principal components analysis of 1012 present-day West Eurasians (grey points, modern Baltic populations in dark grey) with 294 projected published ancient and 38 ancient North European samples introduced in this study (marked with a red outline). Population labels of modern West Eurasians are given in Supplementary Fig. 7 and a zoomed-in version of the European Late Neolithic and Bronze Age samples is provided in Supplementary Fig. 8. b Ancestral components in ancient individuals estimated by ADMIXTURE (k = 11)

This genetic continuity from Corded Ware (the most likely Proto-Uralic homeland) to the Proto-Fennic and Proto-Saamic communities in the Gulf of Finland correlates very well with the known conservatism of Finno-Saamic phonology, quite similar to Finno-Ugric, and both to Proto-Uralic (Kallio 2017): The most isolated region after the expansion of Corded Ware peoples, the Gulf of Finland, shielded against migrations for almost 1,500 years, is then the most conservative – until the arrival of Akozino influence.

NOTE. This has its parallel in the phonetic conservatism of Celtic or Italic compared to Finno-Ugric-influenced Germanic, Balto-Slavic, or Indo-Iranian.

Only later would certain regions (like Finland or Lappland) suffer Y-DNA bottlenecks and further admixture events associated with population displacements and expansions, such as the spread of Fennic peoples from their Estonian homeland (evidenced by the earlier separation of South Estonian) to the north and east:

diversification-finnic
The Finnic family tree. Kallio (2014).

The initial Proto-Fennic expansion was probably coupled with the expansion of Proto-Saami to the north, with the Kjelmøy Ware absorbing the Siberian population of Lovozero Ware, and potentially in inner Finland and Karelia with the Luukonsaari and Sirnihta wares (Carpelan and Parpola 2017).

This Proto-Saami population expansion from the mainland to the north, admixing with Lovozero-related peoples, is clearly reflected in the late Iron Age Saamic samples from Levänluhta (ca. 400-800 AD), as a shift (of 2 out of 3 samples) to Siberian-like ancestry from their original CWC_Baltic-like situation (see PCA from Lamnidis et al. 2018 above).

Also, Volgaic and Permic populations from inner Finland and the Forest Zone to the Cis-Urals and Circum-Artic regions probably incorporate Siberian ancestry and N1c-L392 lineages during these and later population movements, while the westernmost populations – Estonian, Mordvinic – remain less admixed (see PCA from Tambets et al. 2018 below).

We also have data of N1c-L392 in Nordic territory in the Middle Ages, proving its likely strong presence in the Mälaren area since the Iron Age, with the arrival of Akozino warrior traders. Similarly, it is found among Balto-Slavic groups along the eastern Baltic area. Obviously, no language change is seen in Nordic Bronze Age and Lusatian territory, and none is expected in Estonian or Finnish territory, either.

Therefore, no “N1c-L392 + Siberian ancestry” can be seen expanding Finno-Ugric dialects, but rather different infiltrations and population movements with limited effects on ancestry and Y-DNA composition, depending on the specific period and region.

estonians-hungarians-mordvinian
Selection of the PCA, with the group of Estonians, Mordovians, and Hungarians selected. See Tambets et al. (2018) for more information.

An issue never resolved

Because N1c-L392 subclades & Siberian ancestry, which appear in different proportions and with different origins among some modern Uralic peoples, do not appear in cultures supposed to host Uralic-speaking populations until the Iron Age, people keep looking into any direction to find the ‘true’ homeland of those ‘Uralic N1c peoples’? Kind of a full circular reasoning, anyone? The same is valid for R1a & steppe ancestry being followed for ‘Indo-Europeans’, or R1b-P312 & Neolithic farmer ancestry being traced for ‘Basques’, because of their distribution in modern populations.

I understand the caution of many pointing to the need to wait and see how samples after 2000 BC are like, in every single period, from the middle and upper Volga, Kama, southern Finland, and the Forest Zone between Fennoscandia and the steppe. It’s like waiting to see how people from Western Yamna and the Carpathian Basin after 3000 BC look like, to fill in what is lacking between East Yamna and Bell Beakers, and then between them and every single Late PIE dialect.

But the answer for Yamna-Bell Beaker-Poltavka peoples during the Late PIE expansion is always going to be “R1b-L23, but with R1a-Z645 nearby” (we already have a pretty good idea about that); and the answer for the Forest Zone and northern Cis- and Trans-Urals area – during the time when Uralic languages are known to have already been spoken there – is always going to be “R1a-Z645, but with haplogroup N nearby”, as is already clear from the data on the eastern Baltic region.

So, without a previously proposed model as to where those amateurs expressing concern about ‘not having enough data’ expect to find those ‘Uralic peoples’, all this waiting for the right data looks more like a waiting for N1c and Siberian ancestry to pop up somewhere in the historic Uralic-speaking area, to be able to say “There! A Uralic-speaking male!”. Not a very reasonable framework to deal with prehistoric peoples and their languages, I should think.

But, for those who want to do that, let me break the news to you already:

ananino-culture-balto-slavic
First N1c – Finno-Ugric person arrives in Estonia to teach Finno-Saamic to Balto-Slavic peoples.

And here it is, an appropriate fantasy description of the ethnolinguistic groups from the region. You are welcome:

  • During the Bronze Age, late Corded Ware groups evolve as the western Textile ware Fennic Balto-Slavic group in the Gulf of Finland; the Netted Ware Saamic Balto-Slavic group of inner Finland; the south Netted Ware / Akozino Volgaic Balto-Slavic groups of the Middle Volga; and the Anonino Permic Balto-Slavic group in the north-eastern Forest Zone; all developing still in close contact with each other, allowing for common traits to permeate dialects.
  • These Balto-Slavic groups would then incorporate west of the Urals during and after the Iron Age (ca. 800-500 BC first, and also later during their expansion to the north) limited ancestry and lineages from eastern European hunter-gatherer groups of Palaeo-European Fennic and Palaeo-Siberian Volgaic and Permic languages from the Circum-Artic region, but they adopted nevertheless the language of the newcomers in every single infiltration of N1c lineages and/or admixture with Siberian ancestry. Oh and don’t forget the Saamic peoples from central Sweden, of course, the famous N1c-L392 ‘Rurikid’ lineages expanding Saamic to the north and replacing Proto-Germanic…

The current model for those obsessed with modern Y-DNA is, therefore, that expanding Neolithic, Bronze Age and Iron Age cultures from north-eastern Europe adopted the languages of certain lineages originally from sub-Neolithic (Scandinavian and Siberian) hunter-gatherer populations of the Circum-Artic region; lineages that these cultures incorporated unevenly during their expansions. Hmmmm… Sounds like an inverse Western movie, where expanding Americans end up speaking Apache, and the eastern coast speaks Spanish until Italian migrants arrive and make everyone speak English… or something. A logic, no-nonsense approach to ethnolinguistic identification.

I kid you not, this is the kind of models we are going to see very soon. In 2018 and 2019, with ancient DNA able to confirm or reject archaeological hypotheses based on linguistic data, people will keep instead creating new pet theories to support preconceived ideas based on the Y-DNA prevalent among modern populations. That is, information available in the 2000s.

So what’s (so much published) ancient DNA useful for, exactly?

[Next post on the subject: Corded Ware—Uralic (III): Seima-Turbino and the Ugric and Samoyedic expansion]

See also

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