Volosovo hunter-gatherers started to disappear earlier than previously believed

volosovo-corded-ware

Recent paper (behind paywall) Marmot incisors and bear tooth pendants in Volosovo hunter-gatherer burials. New radiocarbon and stable isotope data from the Sakhtysh complex, Upper-Volga region, by Macānea, Nordqvist, and Kostyleva, J. Archaeol. Sci. (2019) 26:101908.

Interesting excerpts (emphasis mine):

The Sakhtysh micro-region is located in the Volga-Oka interfluve, along the headwaters of the Koyka River in the Ivanovo Region, central European Russia (Fig. 1). The area has evidence of human habitation from the Early Mesolithic to the Iron Age, and includes altogether 11 long-term and seasonal settlements (Sakhtysh I–II, IIa, III–IV, VII–XI, XIV) and four artefact scatters (sites V–VI, XII–XIII), in addition to which burials have been detected at five sites (I–II, IIa, VII, VIII) (Kostyleva and Utkin, 2010). The locations have been known since the 1930s and intensively studied since the 1960s under the leadership of D.A. Kraynov, M.G. Zhilin, E.L. Kostyleva, and A.V. Utkin.

Sakhtysh II and IIa are the most extensively studied sites of the complex, with ca. 1500m2 and around 800m2 excavated, respectively. The burial grounds at both sites are considered as fully investigated.

volosovo-sakhtysh-dates
AMS datings from the sites Sakhtysh II and IIa. Sampled contexts are given in parentheses (burial/hoard), “crust” indicates samples of charred organic
residues on pottery from cultural layer. For data, see Tables 1–2.

Sakhtysh chronology

The AMS dates do not support the previously proposed phasing of the Sakhtysh burials to early (4750–4375 BP/3600–3000 cal BCE), late (or developed; 4375–4000 BP/3000–2500 cal BCE), and final (4000–3750 BP/2500–2200 cal BCE): the early and late burials at Sakhtysh IIa do not stand out as two separate groups, and also the burials and hoards from Sakhtysh II, connected to the final phase, are temporally overlapping with these. Neither the use sequence, where the settlement and burial phases are non-overlapping and also complementary between the sites (Kostyleva and Utkin, 2010, 2014), finds support in the present material.

The AMS datings indicate that the Volosovo people started to bury their dead at Sakhtysh IIa after 3700 cal BCE; dates earlier than this may be affected by FRE or suffer from mixed contexts and poor quality of dates. The present data questions the interpretation that the Sakhtysh IIa cemetery was used without interruptions between 4800 and 4080 BP (Kostyleva and Utkin, 2010), i.e. for a millennium between 3550 and 2600 cal BCE. The AMS dates rather suggest a use period of some centuries only around the mid-4th millennium cal BCE, tentatively 3650–3400 cal BCE. This would also be more realistic considering the number of burials at the site.

volosovo-sakhtysh-sites
The core area of Volosovo culture (after Kraynov, 1987) and the sites of the Sakhtysh complex (after Kostyleva and Utkin, 2010). Eurasian map base made with Natural Earth. Illustration: K. Nordqvist.

Volosovo chronology

The absolute dating of Volosovo culture was for a long time hampered by the small number of radiocarbon dates (see Kraynov, 1987). Today,>100 datings connected with it can be found in literature (Korolev and Shalapinin, 2010; Chernykh et al., 2011; Nikitin, 2012; Mosin et al., 2014). Unfortunately, the available dates do not form solid grounds for dating the cultural phenomenon, as many of them have quality-related issues, large measurement errors, and ambiguous cultural or physical contexts. Consequently, particular datings may be connected to different cultural phases by different scholars. Finally, a large part of the newly-published datings are obtained through direct dating of potsherds (Kovaliukh and Skripkin, 2007; Zaitseva et al., 2009), and therefore, their cogency must be faced with reservation (see Van der Plicht et al., 2016; Dolbunova et al., 2017).

The datings connected with Volosovo cover a wide time range between ca. 5500 BP (4400 cal BCE) and ca. 3700 BP (2100 cal BCE). However, datings from secure contexts, with good quality (error ca. 50 years or below) and no probable FRE, place the beginning of Volosovo culture to the first half of the 4th millennium cal BCE, around 3700–3600 cal BCE. This is also supported by the roughly coeval terminal dates given for the preceding Lyalovo (Zaretskaya and Kostyleva, 2011) and Volga-Kama cultures (Lychagina, 2018), as well as the appearance of related neighbouring cultures, for example, in the Kama region (Nikitin, 2012; Lychagina, 2018), the southern forest steppe area (Korolev and Shalapinin, 2014), and north-western Russia and Finland (Nordqvist, 2018). Still, the dating of many of these cultural phases suffers from the same problems as of Volosovo.

A handful of contested datings place the end of Volosovo culture to the final centuries of the 3rd millennium cal BCE, or even later (Kostyleva and Utkin, 2010; Chernykh et al., 2011; Nikitin, 2012). On the other hand, the new AMS dates indicate that Volosovo activities at Sakhtysh II and IIa ceased before or towards the early 3rd millennium cal BCE; if this reflects the general decline of Volosovo culture must be still confirmed by more dates from Sakhtysh and elsewhere. In this context, the general cultural development must be accounted for. To what extent – if at all – the Volosovo people were present after the arrival of the Corded Ware culture-related Fatyanovo-Balanovo populations? Based on the current, albeit scant and inconclusive radiocarbon data this took place from ca. 2700 cal BCE onwards (Krenke et al., 2013).

volosovo-fatyanovo-balanovo
Corded Ware and Comb Ware hunter-gatherer-related populations in north-eastern Europe from ca. 2600 BC. See full map.

Comments

One of the interesting genetic papers in the near future will be the one that finally includes samples from Corded Ware groups in the forest zone (i.e. Fatyanovo-Balanovo and Abashevo), which will most likely confirm that they are the origin of the known genetic profile of Central and East Uralic-speaking peoples, seeing how West Uralic peoples show genetic continuity in the East Baltic area, coinciding with the Battle Axe culture.

Uralicists have come a long way from the 1990s, when the picture of Uralic before Balto-Slavic in the Baltic was already evident, and Uralians were identified with Comb Ware peoples. The linguistic data and relative chronology are still valid, despite the now outdated interpretations of absolute archaeological chronology, as happens with interpretations of Krahe or Villar about Old European.

As an example, here are some relevant excerpts from Languages in the Prehistoric Baltic Sea Region, by Kallio (2003):

NOTE. Kallio’s contribution appeared in the book Languages in Prehistoric Europe (2003), which I hold nostalgically close in my Indo-European library (now almost impossible to read fully). It is still one of my preferred books (from those made up of mostly unconnected chunks on European linguistic prehistory), because it contains Oettinger’s essential update of North-West Indo-European common vocabulary, which led us indirectly to our Modern Indo-European project from 2005 on.

In any case, the Uralic arrival in the region east of the Baltic Sea preceded the Indo-European one (…).

This theory that the ancestors of Finno-Saamic speakers arrived in the Baltic Sea region earlier than those of Balto-Slavic speakers is still rejected by some scholars (e.g. Napolskikh 1993: 41-44), who claim, for instance, that Finno-Saamic speakers would not have known salmons before they met Balts because the Finno-Saamic word for ‘salmon’ (i.e. *losi) is a borrowing from Baltic. Similarly, one could claim that English speakers would not have known salmons before they met Frenchmen because English salmon is a borrowing from French. In other words, Worter und Sachen are not necessarily borrowed hand in hand. Otherwise, it would not be so easy to explain how many Finnish names of body parts are borrowings from Baltic (e.g. hammas ‘tooth’, kaula ‘neck’, reisi ‘thigh’) and from Germanic (e.g. hartia ‘shoulder’, lantio ‘loin’, maha ‘stomach’).

A more probative argument is the fact that Balto-Slavic features in Finno-Saamic are mostly lexical ones (i.e. typical superstrate features), where Finno-Saamic features in Balto-Slavic are mostly non-lexical ones (i.e. typical substrate features). Note that there are more Balto-Slavic features in Finnic than in Saamic and more Finno-Saamic features in Baltic than in Slavic. This fact could be explained by presuming that Pre-Saamic was spoken north of the Corded Ware area and Pre-Slavic was spoken south of the Typical Pit-Comb Ware area, whereas Pre-Finnic and Pre-Baltic alone were spoken in the area, where both the Typical Pit-Comb Ware culture (ca. 4000-3600 BC) and the Corded Ware culture (ca. 3200-2300 BC) were situated. This area was most probably bilingual, until Finnic and Baltic won in the north and in the south, respectively.

As is well-known, the idea of Uralic substrate features in Balto-Slavic is not new (cf. e.g. Pokorny 1936/1968: 181-185). As recent studies (e.g. Bednarczuk 1997) have shown, their density is the most remarkable in the four Balto-Slavic languages spoken in the earlier Pit-Comb Ware area (i.e. Latvian, Lithuanian, Belorussian, Russian). On the other hand, occasional Uralisms in the other Balto-Slavic languages spoken west of the Vistula and south of the Pripyat may rather be considered adstrate features spread from the northeast.

comb-ware-uralic
Our beliefs from the 2000s. A hypothetic Uralic Comb Ware distribution before the arrival of a hypothetic North-West Indo-European-speaking Corded Ware. “Generalized distribution of the Pit-Comb Ware cultural complex (Mallory & Adams 1997: 430, Carpelan 1999: 257) and the most probable homelands of Saamic, Finnic, Mordvin, Mari, and Permic.”

The idea of Indo-European superstrate features in Finnic is not new either (cf. e.g. Posti 1953). As Jorma Koivulehto (1983) has recently shown, the earliest Indo-European loanword stratum in the westernmost Uralic branches alone can be considered Northwest Indo-European and connected with the Corded Ware culture (ca. 3200-2300 BC). Since this layer, there have been continuous contacts between Baltic and Finnic. According to Koivulehto (1990), the following stratum can be called Proto-Balt(o-Slav)ic and dated to the Late Neolithic period (ca. 2300-1500 BC). Note that this Proto-Balt(o-Slav)ic dating agrees with the established ones (cf. e.g. Shevelov 1964: 613-614, Kortlandt 1982: 181), when we remember the fact that archaeologists have also moved their datings back by centuries during the last decades.

Finally, there is also a Baltic loanword stratum which was not borrowed from the ancestral stage of Latvian, Lithuanian and/or Old Prussian but from some extinct Baltic language or dialect (Nieminen 1957). However, as these words still go back to the early Proto-Finnic stage, they can hardly be dated later than Bronze-Age ( ca. 1500-500 BC). Therefore, we may conclude that they were probably borrowed from a Baltic superstrate, which arrived in the Finnish Gulf area during the Corded Ware period and survived there until the Bronze Age, when it was no longer identical with other Baltic dialects. In any case, as later Baltic loanword strata concern southern Finnic languages alone, we may presume that this ‘North Baltic’superstrate had become extinct.

The traditional association of Uralic with Volosovo hunter-gatherers doesn’t make sense, since they neither miraculously survived for thousands of years nor mixed for hundreds of years with Corded Ware peoples, so we can now more confidently reject the recent assumption by Carpelan & Parpola that their language was adopted by incoming Fatyanovo, Balanovo and Abashevo groups, to develop into the known Uralic languages (more here). This includes one of the many models of the the Copenhagen group, who simplistically follow “Steppe ancestry” for Indo-Europeannes.

If one combines the known relative linguistic chronology with the North-West Indo-European hydrotoponymy layer, now more clearly identified as Old Europeans expanding with East Bell Beakers and derived Early Bronze Age groups, I think there is little space left for maneuvering out of the overwhelming evidence for a Uralic homeland in the forest-steppes, linked to the spread of late Sredni Stog/Corded Ware ancestry into north-eastern Europe and beyond the Urals.

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

Haplogroup R1a and CWC ancestry predominate in Fennic, Ugric, and Samoyedic groups

uralic-languages

Open access Genes reveal traces of common recent demographic history for most of the Uralic-speaking populations, by Tambets et al. Genome Biology (2018).

Interesting excerpts (emphasis mine):

Methods

A total of 286 samples of Uralic-speaking individuals, of those 121 genotyped in this study, were analysed in the context of 1514 Eurasian samples (including 14 samples published for the first time) based on whole genome single nucleotide polymorphisms (SNPs) (Additional file 1: Table S1). All these samples, together with the larger sample set of Uralic speakers, were characterized for mtDNA and chrY markers.

The question as which material cultures may have co-spread together with proto-Uralic and Uralic languages depends on the time estimates of the splits in the Uralic language tree. Deeper age estimates (6,000 BP) of the Uralic language tree suggest a connection between the spread of FU languages from the Volga River basin towards the Baltic Sea either with the expansion of the Neolithic culture of Combed Ware, e.g. [6, 7, 17, 26] or with the Neolithic Volosovo culture [7]. Younger age estimates support a link between the westward dispersion of Proto-Finno-Saamic and eastward dispersion of Proto-Samoyedic with a BA Sejma-Turbino (ST) cultural complex [14, 18, 27, 28] that mediated the diffusion of specific metal tools and weapons from the Altai Mountains over the Urals to Northern Europe or with the Netted Ware culture [23], which succeeded Volosovo culture in the west. It has been suggested that Proto-Uralic may have even served as the lingua franca of the merchants involved in the ST phenomenon [18]. All these scenarios imply that material culture of the Baltic Sea area in Europe was influenced by cultures spreading westward from the periphery of Europe and/or Siberia. Whether these dispersals involved the spread of both languages and people remains so far largely unknown.

The population structure of Uralic speakers

To contextualize the autosomal genetic diversity of Uralic speakers among other Eurasian populations (Additional file 1: Table S1), we first ran the principal component (PC) analysis (Fig. 2a, Additional file 3: Figure S1). The first two PCs (Fig. 2a, Additional file 3: Figure S1A) sketch the geography of the Eurasian populations along the East-West and North-South axes, respectively. The Uralic speakers, along with other populations speaking Slavic and Turkic languages, are scattered along the first PC axis in agreement with their geographic distribution (Figs. 1 and 2a) suggesting that geography is the main predictor of genetic affinity among the groups in the given area. Secondly, in support of this, we find that FST-distances between populations (Additional file 3: Figure S2) decay in correlation with geographical distance (Pearson’s r = 0.77, p < 0.0001). On the UPGMA tree based on these FST-distances (Fig. 2b), the Uralic speakers cluster into several different groups close to their geographic neighbours.

uralic-pca
Principal component analysis (PCA) and genetic distances of Uralic-speaking populations. a PCA (PC1 vs PC2) of the Uralic-speaking populations.

We next used ADMIXTURE [48], which presents the individuals as composed of inferred genetic components in proportions that maximize Hardy-Weinberg and linkage equilibrium in the overall sample (see the ‘Methods’ section for choice of presented K). Overall, and specifically at lower values of K, the genetic makeup of Uralic speakers resembles that of their geographic neighbours. The Saami and (a subset of) the Mansi serve as exceptions to that pattern being more similar to geographically more distant populations (Fig. 3a, Additional file 3: S3). However, starting from K = 9, ADMIXTURE identifies a genetic component (k9, magenta in Fig. 3a, Additional file 3: S3), which is predominantly, although not exclusively, found in Uralic speakers. This component is also well visible on K = 10, which has the best cross-validation index among all tests (Additional file 3: S3B). The spatial distribution of this component (Fig. 3b) shows a frequency peak among Ob-Ugric and Samoyed speakers as well as among neighbouring Kets (Fig. 3a). The proportion of k9 decreases rapidly from West Siberia towards east, south and west, constituting on average 40% of the genetic ancestry of FU speakers in Volga-Ural region (VUR) and 20% in their Turkic-speaking neighbours (Bashkirs, Tatars, Chuvashes; Fig. 3a). The proportion of this component among the Saami in Northern Scandinavia is again similar to that of the VUR FU speakers, which is exceptional in the geographic context. It is also notable that North Russians, sampled from near the White Sea, differ from other Russians by sporting higher proportions of k9 (10–15%), which is similar to the values we observe in their Finnic-speaking neighbours. Notably, Estonians and Hungarians, who are geographically the westernmost Uralic speakers, virtually lack the k9 cluster membership.

siberian-ancestry
Population structure of Uralic-speaking populations inferred from ADMIXTURE analysis on autosomal SNPs in Eurasian context. a Individual ancestry estimates for populations of interest for selected number of assumed ancestral populations (K3, K6, K9, K11). Ancestry components discussed in a main text (k2, k3, k5, k6, k9, k11) are indicated and have the same colours throughout. The names of the Uralic-speaking populations are indicated with blue (Finno-Ugric) or orange (Samoyedic). The full bar plot is presented in Additional file 3: Figure S3. b Frequency map of component k9

We also tested the different demographic histories of female and male lineages by comparing outgroup f3 results for autosomal and X chromosome (chrX) data for pairs of populations (Estonians, Udmurts or Khanty vs others) with high versus low probability to share their patrilineal ancestry in chrY hg N (see the ‘Methods’ section, Additional file 3: Figure S13). We found a minor but significant excess of autosomal affinity relative to chrX for pairs of populations that showed a higher than 10% chance of two randomly sampled males across the two groups sharing their chrY ancestry in hg N3-M178, compared to pairs of populations where such probability is lower than 5% (Additional file 3: Figure S13).

In sum, these results suggest that most of the Uralic speakers may indeed share some level of genetic continuity via k9, which, however, also extends to the geographically close Turkic speakers.

uralic-modern-europe

Identity-by-descent

We found that it is the admixture with the Siberians that makes the Western Uralic speakers different from the tested European populations (Additional file 3: Figure S4A-F, H, J, L). Differentiating between Estonians and Finns, the Siberians share more derived alleles with Finns, while the geographic neighbours of Estonians (and Finns) share more alleles with Estonians (Additional file 3: Figure S4M). Importantly, Estonians do not share more derived alleles with other Finnic, Saami, VUR FU or Ob-Ugric-speaking populations than Latvians (Additional file 3: Figure S4O). The difference between Estonians and Latvians is instead manifested through significantly higher levels of shared drift between Estonians and Siberians on the one hand and Latvians and their immediate geographic neighbours on the other hand. None of the Uralic speakers, including linguistically close Khanty and Mansi, show significantly closer affinities to the Hungarians than any non-FU population from NE Europe (Additional file 3: Figure S4R).

ibd-uralic-genetics
Share of ~ 1–2 cM identity-by-descent (IBD) segments within and between regional groups of Uralic speakers. For each Uralic-speaking population representing lines in this matrix, we performed permutation test to estimate if it shows higher IBD segment sharing with other population (listed in columns) as compared to their geographic control group. Empty rectangles indicate no excess IBD sharing, rectangles filled in blue indicate comparisons when statistically significant excess IBD sharing was detected between one Uralic-speaking population with another Uralic-speaking population (listed in columns), rectangles filled in green mark the comparisons when a Uralic-speaking population shows excess IBD sharing with a non-Uralic-speaking population. For each tested Uralic speaker (matrix rows) populations in the control group that were used to generate permuted samples are indicated using small circles. For example, the rectangle filled in blue for Vepsians and Komis (A) implies that the Uralic-speaking Vepsians share more IBD segments with the Uralic-speaking Komis than the geographic control group for Vepsians, i.e. populations indicated with small circles (Central and North Russians, Swedes, Latvians and Lithuanians). The rectangle filled in green for Vepsians and Dolgans shows that the Uralic-speaking Vepsians share more IBD segments with the non-Uralic-speaking Dolgans than the geographic control group

Time of Siberian admixture

The time depth of the Globetrotter (Fig. 5b) inferred admixture events is relatively recent—500–1900 AD (see also complementary ALDER results, in Additional file 13: Table S12 and Additional file 3: Figure S7)—and agrees broadly with the results reported in Busby et al. [55]. A more detailed examination of the ALDER dates, however, reveals an interesting pattern. The admixture events detected in the Baltic Sea region and VUR Uralic speakers are the oldest (800–900 AD or older) followed by those in VUR Turkic speakers (∼1200–1300 AD), while the admixture dates for most of the Siberian populations (>1500 AD) are the most recent (Additional file 3: Figure S7). The West Eurasian influx into West Siberia seen in modern genomes was thus very recent, while the East Eurasian influx into NE Europe seems to have taken place within the first millennium AD (Fig. 5b, Additional file 3: Figure S7).

Affinities of the Uralic speakers with ancient Eurasians

We next calculated outgroup f3-statistics [48] to estimate the extent of shared genetic drift between modern and ancient Eurasians (Additional file 14: Table S13, Additional file 3: Figures S8-S9). Consistent with previous reports [45, 50], we find that the NE European populations including the Uralic speakers share more drift with any European Mesolithic hunter-gatherer group than Central or Western Europeans (Additional file 3: Figure S9A-C). Contrasting the genetic contribution of western hunter-gatherers (WHG) and eastern hunter-gatherers (EHG), we find that VUR Uralic speakers and the Saami share more drift with EHG. Conversely, WHG shares more drift with the Finnic and West European populations (Additional file 3: Figure S9A). Interestingly, we see a similar pattern of excess of shared drift between VUR and EHG if we substitute WHG with the aDNA sample from the Yamnaya culture (Additional file 3: Figure S9D). As reported before [2, 45], the genetic contribution of European early farmers decreases along an axis from Southern Europe towards the Ural Mountains (Fig. 6, Additional file 3: Figure S9E-F).

yamna-cwc-qpgraph-admixture-uralic
Proportions of ancestral components in studied European and Siberian populations and the tested qpGraph model. a The qpGraph model fitting the data for the tested populations. Colour codes for the terminal nodes: pink—modern populations (‘Population X’ refers to test population) and yellow—ancient populations (aDNA samples and their pools). Nodes coloured other than pink or yellow are hypothetical intermediate populations. We putatively named nodes which we used as admixture sources using the main recipient among known populations. The colours of intermediate nodes on the qpGraph model match those on the admixture proportions panel. b Admixture proportions (%) of ancestral components. We calculated the admixture proportions summing up the relative shares of a set of intermediate populations to explain the full spectrum of admixture components in the test population. We further did the same for the intermediate node CWC’ and present the proportions of the mixing three components in the stacked column bar of CWC’. Colour codes for ancestral components are as follows: dark green—Western hunter gatherer (WHG’); light green—Eastern hunter gatherer (EHG’); grey—European early farmer (LBK’); dark blue—carriers of Corded Ware culture (CWC’); and dark grey—Siberian. CWC’ consists of three sub-components: blue—Caucasian hunter-gatherer in Yamnaya (CHGinY’); light blue—Eastern hunter-gatherer in Yamnaya (EHGinY’); and light grey—Neolithic Levant (NeolL’)

We then used the qpGraph software [48] to test alternative demographic scenarios by trying to fit the genetic diversity observed in a range of the extant Finno-Ugric populations through a model involving the four basic European ancestral components: WHG, EHG, early farmers (LBK), steppe people of Yamnaya/Corded Ware culture (CWC) and a Siberian component (Fig. 6, Additional file 3: Figure S10). We chose the modern Nganasans to serve as a proxy for the latter component because we see least evidence for Western Eurasian admixture (Additional file 3: Figure S3) among them. We also tested the Khantys for that proxy but the model did not fit (yielding f2-statistics, Z-score > 3). The only Uralic-speaking population that did not fit into the tested model with five ancestral components were Hungarians. The qpGraph estimates of the contributions from the Siberian component show that it is the main ancestry component in the West Siberian Uralic speakers and constitutes up to one third of the genomes of modern VUR and the Saami (Fig. 6). It drops, however, to less than 10% in most of NE Europe, to 5% in Estonians and close to zero in Latvians and Lithuanians.

Discussion

uralic-groups-haplogroup-r1a
Additional file 6: Table S5. Y chromosome haplogroup frequencies in Eurasia. Modified by me: in bold haplogroup N1c and R1a from Uralic-speaking populations, with those in red showing where R1a is the major haplogroup. Observe that all Uralic subgroups – Finno-Permic, Ugric, and Samoyedic – have some populations with a majority of R1a lineages.

One of the notable observations that stands out in the fineSTRUCTURE analysis is that neither Hungarians nor Estonians or Mordovians form genetic clusters with other Uralic speakers but instead do so with a broad spectrum of geographically adjacent samples. Despite the documented history of the migration of Magyars [63] and their linguistic affinity to Khantys and Mansis, who today live east of the Ural Mountains, there is nothing in the present-day gene pool of the sampled Hungarians that we could tie specifically to other Uralic speakers.

Perhaps even more surprisingly, we found that Estonians, who show close affinities in IBD analysis to neighbouring Finnic speakers and Saami, do not share an excess of IBD segments with the VUR or Siberian Uralic speakers. This is eIn this context, it is important to remind that the limited (5%, Fig. 6) East Eurasian impact in the autosomal gene pool of modern Estonians contrasts with the fact that more than 30% of Estonian (but not Hungarian) men carry chrY N3 that has an East Eurasian origin and is very frequent among NE European Uralic speakers [36]. However, the spread of chrY hg N3 is not language group specific as it shows similar frequencies in Baltic-speaking Latvians and Lithuanians, and in North Russians, who in all our analyses are very similar to Finnic-speakers. The latter, however, are believed to have either significantly admixed with their Uralic-speaking neighbours or have undergone a language shift from Uralic to Indo-European [38].ven more striking considering that the immediate neighbours—Finns, Vepsians and Karelians—do.

With some exceptions such as Estonians, Hungarians and Mordovians, both IBD sharing and Globetrotter results suggest that there are detectable inter-regional haplotype sharing ties between Uralic speakers from West Siberia and VUR, and between NE European Uralic speakers and VUR. In other words, there is a fragmented pattern of haplotype sharing between populations but no unifying signal of sharing that unite all the studied Uralic speakers.

Comments

The paper is obviously trying to find a “N1c/Siberian ancestry = Uralic” link, but it shows (as previous papers using ancient DNA) that this identification is impossible, because it is not possible to identify “N1c=Siberian ancestry”, “N1c=Uralic”, or “Siberian ancestry = Uralic”. In fact, the arrival of N subclades and Siberian ancestry are late, both events (probably multiple stepped events) are unrelated to each other, and represent east-west demic diffusion waves (as well as founder effects) that probably coincide in part with the Scythian and Turkic (or associated) expansions, i.e. too late for any model of Proto-Uralic or Proto-Finno-Ugric expansion.

On the other hand, it shows interesting data regarding ancestry of populations that show increased Siberian influence, such as those easternmost groups admixed with Yeniseian-like populations (Samoyedic), those showing strong founder effects (Finnic), or those isolated in the Circum-Artic region with neighbouring Siberian peoples in Kola (Saami). All in all, Hungarians, Estonians and Mordovians seem to show the original situation better than the other groups, which is also reflected in part in Y-DNA, conserved as a majority of R1a lineages precisely in these groups. Just another reminder that CWC-related ancestry is found in every single Uralic group, and that it represents the main ancestral component in all non-Samoyedic groups.

estonians-hungarians-mordvinian
Selection of the PCA, with the group of Estonians, Mordovians, and Hungarians selected.

The qpGraph shows the ancestor of Yamna (likely Khvalynsk) and Corded Ware stemming as different populations from a common (likely Neolithic) node – whose difference is based on the proportion of Anatolian-related ancestry – , that is, probably before the Indo-Hittite expansion; and ends with CWC groups forming the base for all Uralic peoples. Below is a detail of the qpGraph on the left, and my old guess (2017) on the right, for comparison:

yamna-corded-ware-qpgraph

#EDIT (22 sep 2018): I enjoyed re-reading it, and found this particular paragraph funny:

Despite the documented history of the migration of Magyars [63] and their linguistic affinity to Khantys and Mansis, who today live east of the Ural Mountains, there is nothing in the present-day gene pool of the sampled Hungarians that we could tie specifically to other Uralic speakers.

They are so obsessed with finding a link to Siberian ancestry and N1c, and so convinced of Kristiansen’s idea of CWC=Indo-European, that they forgot to examine their own data from a critical point of view, and see the clear link between all Uralic peoples with Corded Ware ancestry and R1a-Z645 subclades… Here is a reminder about Hungarians and R1a-Z282, and about the expansion of R1a-Z645 with Uralic peoples.

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