



Interesting excerpts (emphasis mine):

The gap between Russian and Western scholarship










Distribution as a criterion in the dating of loanwords








Evidence for Andronovo Aryan and Indo-Aryan loanwords?





List of Indo-European and Indo-Iranian Etymologies





Ambiguous early loans (can be either from PII or PI)


List of uncertain Indo-Iranian etymologies




Early loans into differentiated branches

Proto-West Uralic

Only in Finnic:



Only in Saami:



Only in Finnic and Saami:








Only in Mordvin, Finnic and/or Saami and Mari

*juma ‘good’


Only in Mari:








Only in Khanty and Mansi (regular cases):






Only in Samoyed:





Pan-Uralic loans







Discussion about the distribution and its impact on Uralic taxonomy






Impacts on dating and the location of the contact zones









Genetic continuity among Uralic-speaking cultures in north-eastern Europe











Late Uralic stems from East Corded Ware groups


























An exclusive Volga-Kama homeland for Disintegrating Uralic?



  • Fatyanovo-Balanovo as Finno-Permic:


This late homeland would be supported, among others, by:









The Battle Axe question











Battle Axe spoke “Para-Balto-Finnic”?





  • Noun paradigms and the form and function of individual cases.




Middle Bronze Age cultures of Eastern Europe.

Genetic continuity = language continuity?







R1a-Z280 and R1a-Z93 shared by ancient Finno-Ugric populations; N1c-Tat expanded with Micro-Altaic

Two important papers have appeared regarding the supposed link of Uralians with haplogroup N.

Avars of haplogroup N1c-Tat

Preprint Genetic insights into the social organisation of the Avar period elite in the 7th century AD Carpathian Basin, by Csáky et al. bioRxiv (2019).

Interesting excerpts (emphasis mine):

After 568 AD the Avars settled in the Carpathian Basin and founded the Avar Qaganate that was an important power in Central Europe until the 9th century. Part of the Avar society was probably of Asian origin, however the localisation of their homeland is hampered by the scarcity of historical and archaeological data.

Here, we study mitogenome and Y chromosomal STR variability of twenty-six individuals, a number of them representing a well-characterised elite group buried at the centre of the Carpathian Basin more than a century after the Avar conquest.

The Y-STR analyses of 17 males give evidence on a surprisingly homogeneous Y chromosomal composition. Y chromosomal STR profiles of 14 males could be assigned to haplogroup N-Tat (also N1a1-M46). N-Tat haplotype I was found in four males from Kunpeszér with identical alleles on at least nine loci. The full Y-STR haplotype I, reconstructed from AC17 with 17 detected STRs, is rare in our days. Only nine matches were found among haplotypes in YHRD database, such as samples from the Ural Region, Northern Europe (Estonia, Finland), and Western Alaska (Yupiks). We performed Median Joining (MJ) network analysis using N-Tat haplotypes with ten shared STR loci (Fig. 3, Table S9). All modern N-Tat samples included in the network had derived allele of L708 as well. Haplotype I (Cluster 1 in Fig. 3) is shared by eight populations on the MJ network among the 24 identical haplotypes. Cluster 1 represents the founding lineage, as it is described in Siberian populations, because this haplotype is shared by the most populations and it is more diverse than Cluster 2.

Nine males share N-Tat haplotype II (on a minimum of eight detected alleles), all of them buried in the Danube-Tisza Interfluve. We found 30 direct matches of this N-Tat haplotype II in the YHRD database, using the complete 17 STR Y-filer profile of AC1, AC12, AC14, AC15, AC19 samples. Most hits came from Mongolia (seven Buryats and one Khalkh) and from Russia (six Yakuts), but identical haplotypes also occur in China (five in Xinjiang and four in Inner Mongolia provinces). On the MJ network, this haplotype II is represented by Cluster 2 and is composed of 45 samples (including 32 Buryats) from six populations (Fig. 3).

Median Joining network of 162 N-Tat Y-STR haplotypes Allelic information of ten Y-STR loci were used for the network. Only those Avar samples were included, which had results for these ten Y-STR loci. The founder haplotype I (Cluster 1) is shared by eight populations including three Mongolian, three Székely, three northern Mansi, two southern Mansi, two Hungarian, eight Khanty, one Finn and two Avar (AC17, AC26) chromosomes. Haplotype II (Cluster 2) includes 45 haplotypes from six populations studied: 32 Buryats, two Mongolians, one Székely, one Uzbek, one Uzbek Madjar, two northern Mansi and six Avars (AC1, AC12, AC14, AC15, AC19 and KSZ 37). Haplotype III (indicated by a red arrow) is AC8. Information on the modern reference samples is seen in Table S9.

A third N-Tat lineage (type III) was represented only once in the Avar dataset (AC8), and has no direct modern parallels from the YHRD database. This haplotype on the MJ network (see red arrow in Fig. 3) seems to be a descendent from other haplotype cluster that is shared by three populations (two Buryat from Mongolia, three Khanty and one Northern Mansi samples). This haplotype cluster also differs one molecular step (locus DYS393) from haplotype II. We classified the Avar samples to downstream subgroup N-F4205 within the N-Tat haplogroup, based on the results of ours and Ilumäe et al.18 and constructed a second network (Fig. S4). The N-F4205 network results support the assumption that the N-Tat Avar samples belong to N-F4205 subgroup (see SI chapter 1d for more details).

Based on our calculation, the age of accumulated STR variance (TMRCA) within N-Tat lineage for all samples is 7.0 kya (95% CI: 4.9 – 9.2 kya), considering the core haplotype (Cluster 1) to be the founding lineage. Y haplogroup N-Tat was not detected by large scale Eurasian ancient DNA studies but it occurs in late Bronze Age Inner Mongolia and late medieval Yakuts, among them N-Tat has still the highest frequency.

Two males (AC4 and AC7) from the Transtisza group belong to two different haplotypes of Y-haplogroup Q1. Both Q1a-F1096 and Q1b-M346 haplotypes have neither direct nor one step neighbour matches in the worldwide YHRD database. A network of the Q1b-M346 haplotype shows that this male had a probable Altaian or South Siberian paternal genetic origin.

EDIT (5 APR 2019): The paper offers an interesting late sample before the arrival of Hungarian conquerors, although we don’t know which precise lineage the sample belongs to:

One sample in our dataset (HC9) comes from this population, and both his mtDNA (T1a1b) and Y chromosome (R1a) support Eastern European connections. (…) Furthermore, we excluded sample HC9 from population-genetic statistical analyses because it belongs to a later period (end of 7th – early 9th centuries)

Apparently, then, results are consistent with what was already known from studies of modern populations:

According to Ilumäe et al. study, the frequency peak of N-F4205 (N3a5-F4205) chromosomes is close to the Transbaikal region of Southern Siberia and Mongolia, and we conclude that most Avar N-Tat chromosomes probably originated from a common source population of people living in this area, completely in line with the results of Ilumäe et al.

Geographic-Distribution Map of hg N3 from Ilumäe et al.

Finno-Ugrians share haplogroup R1a-Z280

Another paper, behind paywall, Genetic history of Bashkirian Mari and Southern Mansi ethnic groups in the Ural region, by Dudás et al. Molecular Genetics and Genomics (2019).

Interesting excerpts (emphasis mine):

Y‑chromosome diversity

The most frequent haplogroups of the Bashkirian Maris were N1b-P43 (42%), R1a-Z280 (16%), R1a-Z93 (16%), N1c-Tat (13%), and J2-M172 (7%). Furthermore, subgroup R1b-M343 accounted for 4% and I2a-P37 covered 2% of the lineages. None of the Mari N1c Y chromosomes belonged to the N1c subgroups investigated (L1034, VL29, Z1936).

In the case of the Southern Mansi males, the most frequent haplogroups were N1b-P43 (33%), N1c-L1034 (28%) and R1a-Z280 (19%). The frequencies of the remaining haplogroups were as follows: R1a-M458 (6%), I1-L22 (3%), I2a-P37 (3%), and R1b-P312 (3%). The haplotype and haplogroup diversities of the Bashkirian Mari group were 0.9929 and 0.7657, whereas these values for the Southern Mansi were 0.9984 and 0.7873, respectively. The results show that, in both populations, haplotypes are much more diverse than haplogroups.

Haplogroup frequencies of the Bashkirian Mari and the Southern Mansi ethnic groups in Ural region

Genetic structure

(..) the studied Bashkirian Mari and Southern Mansi population groups formed a compact cluster along with two Khanty, Northern Mansi, Mari, and Estonian populations based on close Fst-genetic distances (< 0.05), with nonsignificant p values (p > 0.05) except for the Estonian population. All of these populations belong to the Finno-Ugric language family. Interestingly, the other Mansi population studied by Pimenoff et al. (2008) (pop # 38) was located a great distance from the Southern Mansi group (0.268). In addition, the Bashkir population (pop # 6) did not show a close genetic affinity to the Bashkirian Mari group (0.194), even though it is the host population. However, the Russian population from the Eastern European region of Russia (pop # 49) showed a genetic distance of 0.055 with the Southern Mansi group. All Hungarian speaking populations (pops 13, 22, 23, 24, 50, and 51) showed close genetic affinities to each other and to the neighbouring populations, but not to the two studied populations.

Multidimensional scaling (MDS) plot constructed on Fstgenetic distances of Y haplogroup frequencies of 63 populations compared. The haplogroup frequency data used for population comparison together with references are seen in Online Resource 2 (ESM_2). Pairwise Fst-genetic distances and p values between 63 populations were calculated as shown in Online Resource 3 (ESM_3) Fig. 4 Multidimensional scaling (MDS) plot constructed on Rstgenetic distances of 10 STR-based Y haplotype frequencies of 21 populations compared. Image modified to include labels of modern populations.

Phylogenetic analysis

Median-joining networks were constructed for:

N-P43 (earlier N1b):

(…) TMRCA estimates for this haplogroup were made for all P43 samples (n = 157) 8.7 kya (95% CI 6.7–10.8 kya), for the N-P43 Asian.


(…) 75% of Buryats belonged to Haplotype 2, indicating that the Buryats studied by us is a young and isolated population (Bíró et al. 2015). Bashkirian Mari samples derive from Haplotype 2 via Haplotype 3 (see dark purple circles on the top of Fig. 6a). Haplotype 3 contained six males (2 Buryat, 1 Northern Mansi, and 3 Khanty samples from Pimenoff et al. 2008). The biggest Bashkirian Mari haplotype node (3 Mari samples) was positioned three mutational steps away from Haplotype 1 and the remaining Mari samples can be derived from this haplotype. Southern Mansi haplotypes were scattered within the network except for two, which formed a smaller haplotype node with two Northern Mansi and two Khanty samples from Pimenoff et al. (2008).

Median-Joining Networks (MJ) of 153 N-Tat (a) and 26 N-L1034 (b) haplotypes constructed. The circle sizes are proportional to the haplotype frequencies. The smallest area is equivalent to one individual. For N-Tat network, we used data from Southern Mansi (n = 11), Bashkirian Mari (n = 6) samples with Hungarian (n = 12), Hungarian speaking Székely (n = 6), Northern Mansi (n = 14), Mongolian (n = 16), Buryat (n = 44), Finnish (n = 13), Uzbek Madjar (n = 2), Uzbek (n = 3), Khanty (n = 4) populations studied earlier by us (Fehér et al. 2015; Bíró et al. 2015) and Khanty (n = 18) and Mansi (n = 4) studied by Pimenoff et al. (2008)

R1a-Z280 haplotypes, shared by Maris, Mansis, and Hungarians, hence ancient Finno-Ugrians:

The founder R1a-Z280 haplotype was shared by four samples from four populations (1 Bashkirian Mari; 1 Southern Mansi; 1 Hungarian speaking Székely; and 1 Hungarian), as presented in Fig. 7 (Haplotype 1). Haplotype 2 included five males (3 Bashkirian Mari and 2 Hungarian), as it can be seen in Fig. 7. Haplotype 4 included two shared haplotypes (1 Bashkirian Mari and one Hungarian speaking Csángó). The remaining two Bashkirian Mari haplotypes differ from the founder haplotype (Haplotype 1) by two mutational steps via Hungarian or Hungarian and Bashkirian Mari shared haplotypes. Beside Haplotype 1, the remaining Southern Mansi haplotypes were shared with Hungarians (Haplotype 5 or turquoise blue and red-coloured circles above Haplotype 7) or with Hungarians and Hungarian speaking Székely group (Haplotypes 3, 5, and 6). Haplotype 7 included ten Hungarian speakers (Hungarian, Székely, and Csángó). One Hungarian and one Uzbek Khwarezm shared haplotype can be found in Fig. 7 as well (red and white-coloured circle). All the other haplotypes were scattered in the network. The age of accumulated STR variation within R1a-Z280 lineage for 93 samples is estimated to be 9.4 kya (95% CI 6.5–12.4 kya) considering Haplotype 1 (Fig. 7) to be the founder.

Median-Joining Networks (MJ) of 93 R1a-Z280 haplotypes constructed. The circle sizes are proportional to the haplotype frequencies. The smallest area is equivalent to one individual. We used haplotype data from Bashkirian Mari (n = 7), Southern Mansi (n = 7), Hungarian (n = 52), Hungarian speaking Székely (n = 11), Hungarian speaking Csángó (n = 10), Uzbek Ferghana (n = 2), Uzbek Tashkent (n = 1), Uzbek Khwarezm (n = 1) and Northern Mansi (n = 2) populations

R1a-Z93 as isolated lineages among Permic and Ugric populations:

Figure 8 depicts an MJ network of R1a-Z93* samples using 106 haplotypes from the 14 populations (Fig. 8). All of the Bashkirian Mari samples (7 haplotypes) formed a very isolated branch and differed from the one Hungarian haplotype (Fig. 8, see Haplotype 1) by seven mutational steps as well from two Uzbek Tashkent samples (see Haplotype 3). Another Hungarian sample shared two haplotypes of Uzbek Khwarezm samples in Haplotype 4. This haplotype can be derived from Haplotype 3 (Uzbek Tashkent). Haplotype 2 included one Hungarian and one Khakassian male. The remaining three Hungarian haplotypes are outliers in the network and are not shared by any sample. The other population samples included in the network either form independent clusters such as Altaians, Khakassians, Khanties, and Uzbek Madjars or were scattered in the network. The age of accumulated STR variation (TMRCA) within R1a-Z93* lineage for 106 samples is estimated as 11.6 kya (95% CI 9.3–14.0 kya) considering an Armenian haplotype (Fig. 8, “A”) to be the founder and the median haplotype.

Median-Joining Networks (MJ) of 106 R1a-Z93 haplotypes constructed. The circle sizes are proportional to the haplotype frequencies. The smallest area is equivalent to one individual. We used the next haplotype data: 7 Bashkirian Mari, 6 Khanty, 4 Uzbek Madjar, 5 Uzbek Ferghana, 9 Uzbek Tashkent, 7 Uzbek Khwarezm, 2 Mongolian, 2 Buryat, 6 Hungarian samples tested by us for this study or published earlier (Bíró et al. 2015) and populations (3 Armenian; 3 Afghan Tajik;
16 Altaian; 24 Khakassian; 12 Kyrgyz) from Underhill et al. (2015)


The results of modern populations for N (especially N1c) subclades show really wide clusters and ancient TMRCA, consistent with their known ancient and wide distribution in northern and eastern Eurasian groups, and thus with infiltration of different lineages with eastern nomads (and northern Arctic populations) coupled with later bottlenecks, as well as acculturation of groups.

EDIT (2 APR): Interesting is the specific subclade to which ancient Mongolic-speaking Avars belong (information from Yfull) N1c-F4205 (TMRCA ca. 500 BC), subclade of N1c-Y6058 (formed ca. 2800 BC, TMRCA ca. 2800 BC). This branch also gives the “European” branch N1c-CTS10760 (formed ca. 2800 BC, TMRCA ca. 2100 BC), and is subclade of a branch of N1c-L392 (formed ca. 4400 BC, TMRCA ca. 2800 BC). A northern expansion of N1c-L392 is probably represented by its branch N1c-Z1936 (formed ca. 2800, TMRCA ca. 2100 BC), the most likely candidate to appear in the Kola Peninsula in the Bronze Age as the Palaeo-Laplandic population (see here). Read more about potential routes of expansion of haplogroup N.

On the other hand, R1a-Z280 lineages form a tight cluster connecting Permic with Ugric groups, with R1a-Z93 showing early isolation (probably) between Cis-Urals and Trans-Urals regions. While both Corded Ware lineages in Finno-Ugrians are most likely related to the Abashevo expansion through Seima-Turbino and the Andronovo-like Horizon (and potentially later Eurasian expansions), a plausible hypothesis would be that Finno-Ugrians are related to an expansion of R1a-Z283 haplogroups (we already knew about the Finno-Permic connection), while the ancient connection between Permians and Hungarians with R1a-Z93 would correspond to this haplogroup’s potentially tighter link with an early Samoyedic split.

I don’t think that an explosive expansion of eastern Corded Ware groups of R1a-Z645 lineages will show a clear-cut division of haplogroups among Eastern Uralic groups, though, and culturally I doubt we will have such a clear image, either (similar to how the explosive expansion of Bell Beakers cannot be easily divided by regional/language group into R1b-L151 subclades before the known bottlenecks). Relevant in this regard are the known Z93 samples from the Árpád dynasty.

Nevertheless, this data may represent a slightly more recent wave of R1a-Z280 lineages linked to the expansion of Ugric into the Trans-Uralian region, after their split from Finno-Permic, still in close contact with Indo-Iranians in Poltavka and Sintashta-Potapovka, evident from the early and late Indo-Iranian borrowings, during a common period when Samoyedic had already separated.

Such a “Z283 over Z93” layer in the Trans-Urals (and Cis-Urals?) forest-steppes would be similar to the apparent replacement of Z284 by Z282 in the Eastern Baltic during the Bronze Age (possibly with the second or Estonian Battle Axe wave or, much more likely during later population movements). Such an early R1a-Z93 split could potentially be supported also by the separation into bottlenecks under “Northern” (R1a-Z283) Finno-Ugric-speaking Abashevo-related groups and “Southern” (R1a-Z93) acculturated Indo-Iranian-speaking Abashevo migrants developing Sintashta-Potapovka admixing with Poltavka R1b-Z2103 herders.

Modified image, from Underhill et al. (2015). Spatial frequency distributions of Z282 (green) and Z93 (blue) affiliated haplogroups.. Notice the potential Finno-Ugric-associated distribution of Z282 (especially R1a-M558, a Z280 subclade), the expansion of R1a-Z2123 subclades with Central Asian forest-steppe groups.


Let’s review some of the most common myths about Hungarians (and Finno-Ugrians in general) repeated ad nauseam, side by side with my assertions:

❌ N (especially N1c-Tat) in ancient and modern samples represent the True Uralic™ N1c peoples including Magyar tribes? Nope.

✅ Ancient N (especially N1c-Tat) lineages among Uralic populations expanded relatively recently, and differently in different regions (including eastern steppe nomads and northern arctic populations) not associated with a particular language or language group? Yep (read the series on Corded Ware = Uralic expansion).

❌ Modern Hungarian R1a-Z280 lineages represent the majority of the native population, poor Slavic ‘peasants’ from the Carpathian Basin, forcibly acculturated by a minority of bad bad Hungarian hordes? Nope.

✅ Modern Hungarian R1a-Z280 subclades represent Ugric lineages in common with ancient R1a-Z645 Finno-Ugric populations from north-eastern Europe and the Trans-Urals? Yep (see Avars and Ugrians).

❌ Modern Hungarian R1a-Z93 lineages represent acculturated Iranian/Turkic peoples from the steppes? Not likely.

✅ Modern Hungarian R1a-Z93 lineages represent a remnant of the expansion of Corded Ware to the east, potentially more clearly associated with Samoyedic? Much more likely.


Sooo, the theory of a “diluted” Y-DNA in Modern Hungarians from originally fully N-dominated conquerors subjugating native R1a-Z280 Slavs from the Carpathian Basin is not backed up by genetic studies? The ethnic Iranian-Turkic R1a-Z93 federation in the steppes that ended up speaking Magyar is not real?? Who would’ve thunk.

Another true story whose rejection in genetics could not be predicted, like, not at all.

Totally unexpected, too, the drift of “R1a=IE” fans with the newest genetic findings towards a Molgen-like “Yamna/R1b = Vasconic-Caucasian”, “N1c = Uralic-Altaic”, and “R1a = the origin of the white world in Mother Russia”. So much for the supposed interest in “Steppe ancestry” and fancy statistics.


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


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):


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.

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.

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.



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).

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).

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.



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.


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.


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:


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


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.