Baltic Finns in the Bronze Age, of hg. R1a-Z283 and Corded Ware ancestry

estonian-bronze-age-dna

Open access The Arrival of Siberian Ancestry Connecting the Eastern Baltic to Uralic Speakers further East, by Saag et al. Current Biology (2019).

Interesting excerpts:

In this study, we present new genomic data from Estonian Late Bronze Age stone-cist graves (1200–400 BC) (EstBA) and Pre-Roman Iron Age tarand cemeteries (800/500 BC–50 AD) (EstIA). The cultural background of stone-cist graves indicates strong connections both to the west and the east [20, 21]. The Iron Age (IA) tarands have been proposed to mirror “houses of the dead” found among Uralic peoples of the Volga-Kama region [22].

(…) The 33 individuals included 15 from EstBA, 6 from EstIA, 5 from Pre-Roman to Roman Iron Age Ingria (500 BC–450 AD) (IngIA), and 7 from Middle Age Estonia (1200–1600 AD) (EstMA) and yielded endogenous DNA ∼4%–88%, average genomic coverages ∼0.017–0.734×, and contamination estimates <4% (Table S1). We analyzed the data in the context of modern and other ancient individuals, including from Neolithic Estonia [13].

estonian-y-dna-bronze-iron-age
Archaeological Information, Genetic Sex, mtDNA and Y Chromosome Haplogroups, and Average Coverage of the Individuals of This Study. Modified from the paper to mark distinct Y-DNA haplogroups in the LBA and IA.

We identified chrY hgs for 30 male individuals (Tables 1 and S2; STAR Methods). All 16 successfully haplogrouped EstBA males belonged to hg R1a, showing no change from the CWC period, when this was also the only chrY lineage detected in the Eastern Baltic [11, 13, 30, 31]. Three EstIA and two IngIA individuals also belonged to hg R1a, but three EstIA males belonged to hg N3a, the earliest so far observed in the Eastern Baltic. Three EstMA individuals belonged to hg N3a, two to hg R1a, and one to hg J2b. ChrY lineages found in the Baltic Sea region before the CWC belong to hgs I, R1b, R1a5, and Q [10, 11, 12, 13, 17, 32]. Thus, it appears that these lineages were substantially replaced in the Eastern Baltic by hg R1a [10, 11, 12, 13], most likely through steppe migrations from the east [30, 31]. (…) Our results enable us to conclude that, although the expansion time for R1a1 and N3a3′5 in Eastern Europe is similar [25], hg N3a likely reached Estonia or at least became comparably frequent to modern Estonia [1] only during the BA-IA transition.

A clear shift toward West Eurasian hunter-gatherers is visible between European LN and BA (including Baltic CWC) and EstBA individuals, the latter clustering together with Latvian and Lithuanian BA individuals [11]. EstIA, IngIA, and EstMA individuals project between BA individuals and modern Estonians, partially overlapping with both.

(…) EstBA individuals are clearly distinguishable from Estonian CWC individuals as the former have more of the blue component most frequent in WHGs and less of the brown and yellow components maximized in Caucasus hunter-gatherers and modern Khanty, respectively. The individuals of EstBA, EstIA, IngIA, EstMA, and modern Estonia are quite similar to each other on average, indicating that the relatively high proportion of WHG ancestry in modern Eastern Baltic populations compared to other present-day Europeans [15] traces back to the BA.

estonian-pca-published
Detail of the PCA, modified from the paper to label populations. Estonian Bronze Age and Iron Age samples cluster close to Early Corded Ware from the Baltic.. Principal-component analysis results of modern West Eurasians with ancient individuals projected onto the first two components (PC1 and PC2). BA, Bronze Age; EF, early farmers; HG, hunter-gatherers; IA, Iron Age; IMA, Iron/Middle Ages; LN, Late Neolithic; LNBA, Late Neolithic/Bronze Age; MA, Middle Ages

When comparing Estonian CWC and EstBA using autosomal outgroup f3 and Patterson’s D statistics (Table S3), the latter is more similar to other Baltic BA populations, to Baltic IA and Middle Age (MA) populations, and also to populations similar to WHGs and Scandinavian hunter-gatherers (SHGs), but not to Estonian CCC (Figures 2A and S2A; Data S1). The increase in WHG or SHG ancestry could be connected to western influences seen in material culture [20, 21] and facilitated by a decline in local population after the CCC-CWC period [20]. A slight trend of bigger similarity of Estonian CWC to forest or steppe zone populations and of EstBA to European early farmer populations can also be seen.

(…) When comparing to modern populations, Estonian CWC is slightly more similar to Caucasus individuals but EstBA to Baltic populations and Finnic speakers (Figure 2B; Data S1). Outgroup f3 and D statistics do not reveal apparent differences when comparing EstBA to EstIA, EstIA to IngIA, and EstIA to EstMA (Data S1).

estonian-ba-ia-ancestry
qpAdm results. Error bars indicate one SE. Central MN, Central European Middle Neolithic; EstBA, Estonian Bronze Age; EstIA, Estonian Iron Age; IngIA, Ingrian Iron Age; EstMA, Estonian Middle Ages; WHG, western hunter-gatherers.

These results highlight how uniparental and autosomal data can lead to different demographic inferences—the genetic change between CWC and BA not seen in uniparental lineages is clear in autosomal data and the appearance of chrY hg N in the IA is not matched by a clear shift in autosomal profiles.

EstBA individuals have no Nganasan-related ancestry and EstIA, IngIA, and EstMA individuals on average have 2% or 4% (Figure 3; Data S1). The differentiation remains when using BA or IA Fennoscandian populations [26] instead of Nganasans (Data S1). Notably, the proportion of Nganasan-related ancestry varies between 0% and 12% among sampled EstIA, IngIA, and EstMA individuals (Data S1), which may suggest its relatively recent admixture into the target population. Moreover, two individuals from Kunda (0LS10 and V10) have the highest proportions of Nganasan ancestry among EstIA (6% and 8%), one of them has chrY hg N3a, and isotopic analysis suggests neither individual being born in Kunda [34].

About these two males from Tarand-graves, ‘foreign’ to Kunda:

0LS10: Male from tarand III (burial 9; TÜ 1325: L777), age 17–25 years [34]. He had a fragment of a sheep/goat bone and ceramics as grave goods. This burial has two radiocarbon dates: 2430 ± 35 BP (Poz-10801; 760–400 cal BC) and 2530 ± 41 BP (UBA-26114; 800–530 cal BC) [34]. According to the isotopic analysis, the person was not born in the vicinity of Kunda; his place of birth is still unknown (but south-western Finland and Sweden are excluded) [34]. Sampled tooth r P1.

V10: Male from tarand XI (burial 24; TÜ 1325: L1925), age 25–35 years [34], date 2484 ± 40 BP (UBA-26115; 790–430 cal BC) [34]. He had a few potsherds near the skull. Likewise, this person was not locally born [34]. Sampled tooth l P1.

estonia-bronze-iron-age-steppe-siberian
Autosomal Analyses’ Results for Gyvakarai1 as the closest available Corded Ware source for Balto-Finnic populations.

The paper shows thus:

  • Major continuity of ancestry from Corded Ware to modern Estonians, with only slight changes in different periods. In fact, one of the best fits for the Late Bronze Age ancestry is Gyvakarai1, one of the Corded Ware “outliers” described as “closer to Yamna”, which I already said may be closer to Sredni Stog/EHG populations instead. Another interesting take is that the change from Bronze Age to Iron Age corresponds to an increase in Baltic Corded Ware-related ancestry, rather than being driven by Siberian ancestry.
  • pca-mittnik-gyvakarai
    File modified by me from Mittnik et al. (2018) to include the approximate position of the most common ancestral components, and an identification of potential outliers. Zoomed-in version of the European Late Neolithic and Bronze Age samples. “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). From Mittnik et al. (2018).
  • A Volosovo-related migration of hg. N1c with Netted Ware into the area seems to be discarded, based on the full replacement of paternal lines and continuity of R1a-Z283. It is only during the Tarand-grave period when a system of chiefdoms (spread from Ananyino/Akozino) brings haplogroup N1c to the Gulf of Finland. During the Iron Age, the proportion of paternal lineages is still clearly in favour of R1a (50% in the coast, 100% in Ostrobothnia), which indicates a gradual replacement led by elites, likely because of the incorporation of Akozino warrior-traders spreading all over the Baltic, bringing the described shared Mordvinic traits in Fennic.
  • finno-ugric-haplogroup-n
    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. Fading purple arrows represent likely stepped movements of subclades of haplogroup N for centuries (e.g. Siberian → Ananino → Akozino → Fennoscandia [N-VL29]; Circum-Arctic → forest-steppe [N1, N2]; etc.). Blue arrows represent eventual expansions of Uralic peoples to the north. Modified image from Vasilyev (2002).
  • The arrival of Akozino warrior-traders (bringing N1c and R1a lineages) was probably linked to this minimal “Nganasan-like” ancestry of some samples in the transition to the Iron Age. This arrival is supported by samples 0LS10 (the earliest hg. N1c) and V10 (of hg. R1a), both dated to ca. 800-400 BC, with V10 showing the highest “Nganasan-like” ancestry with 4.8%, both of them neighbouring samples showing 0%. This variable admixture among local and foreign paternal lineages might support the described social system of family alliances with intermarriages. In fact, a medieval sample, 0LS03_1 (hg. R1a) also shows a recent “Nganasan-like” ancestry, which probably points to the integration of different Arctic-related ancestry components among Modern Estonians, in this case related to Finnish expansions and thus integration of Levänluhta-related ancestry, as per the supplementary data.
  • NOTE. Such minimal proportions of “Nganasan-like” ancestry evidence the process of admixture of Volga Finns in Akozino territory through their close interactions with Permians of Ananyino, who in turn acquired this Palaeo-Arctic admixture most likely during the expansion of the linguistic community to hunter-gatherer territories, to the north of the Cis-Urals. This process of stepped infiltration and expansion without language change is not dissimilar to the one seen among Indo-Iranians and Balto-Slavs of hg. R1b, or Vasconic speakers of hg. I2a, although in the case of Baltic Finns of hg. R1a the process of infiltration and expansion of hg. N1c is much less dramatic, with no radical replacement anywhere before the huge bottlenecks observable in Finns.

  • The expansion of haplogroup N1c among Finnic populations, as we are going to see in samples from the Middle Ages such as Luistari, is the consequence of late founder effects after huge bottlenecks expected based on the analysis of modern populations. The expansion of N1c-VL29 is different in origin from that of N1c-Z1936 among Samic (later integrated into Finnish populations), most likely from the east and originally associated with Lovozero Ware.
haplogroup_n3a3
Frequency-Distribution Maps of Individual Subclade N3a3 / N1a1a1a1a1a-CTS2929/VL29, probably initially with Akozino warrior-traders. Map from Ilumäe et al. (2016).

In spite of all this, the conclusion of the paper is (surprise!) that Siberian ancestry and hg. N heralded the arrival of Finnic to the Gulf of Finland in the Iron Age… However, this conclusion is supposedly* supported, not by their previous papers, but by a recent phylogenetic study by Honkola et al. (2013), which doesn’t actually argue for such a late ‘arrival’: it argues for the split of Balto-Finnic around 1500 BC.

NOTE. I say ‘supposedly’ because Kristiina Tambets, for example, has been following the link of Uralic with haplogroup N since the 2000s, so this is not some conclusion they just happened to misread from some random paper they Googled. In those initial assessments, she argued that the “ancient homeland” of the Tat C mutation suggested that Finno-Ugrians were in Fennoscandia before Indo-Europeans. Apparently, since haplogroup N appears later and from the east, it is now more important to follow this haplogroup than what is established in archaeology and linguistics.

Even in the referred paper, this split is considered an in situ development, since the phylogenetic study takes the information – among others – 1) from Parpola and Carpelan, who consider Netted Ware, a culture derived from Fatyanovo/Abashevo and Volosovo, as the culprit of the Finno-Ugric expansion; and 2) from Kallio (2006), who clearly states that Proto-Balto-Finnic (like Proto-Finno-Samic) was spoken around the Gulf of Finland during the Bronze Age. Both of them set the terminus ante quem of the language presence in the Baltic ca. 1900 BC.

Anyways, as a consequence of geneticists keeping these untenable pre-ancient DNA haplogroup-based arguments today, I expect to see this “Finnic” language expansion also described for the Western Baltic, Scandinavia or northern Europe, when this same proportion of hg. N1c and “Nganasan” ancestry is observed in Iron Age samples around the Baltic Sea. The nativist trends that this domination of “Finns” all over Northern Europe 2,500 years ago will create will be even more fun to read than the current ones…

EDIT (10 May 2019) How I see the reaction of many to ancient DNA, in keeping their old theories:

Related

Uralic speakers formed clines of Corded Ware ancestry with WHG:ANE populations

steppe-forest-tundra-biomes-uralic

The preprint by Jeong et al. (2018) has been published: The genetic history of admixture across inner Eurasia Nature Ecol. Evol. (2019).

Interesting excerpts, referring mainly to Uralic peoples (emphasis mine):

A model-based clustering analysis using ADMIXTURE shows a similar pattern (Fig. 2b and Supplementary Fig. 3). Overall, the proportions of ancestry components associated with Eastern or Western Eurasians are well correlated with longitude in inner Eurasians (Fig. 3). Notable outliers include known historical migrants such as Kalmyks, Nogais and Dungans. The Uralic- and Yeniseian-speaking populations, as well as Russians from multiple locations, derive most of their Eastern Eurasian ancestry from a component most enriched in Nganasans, while Turkic/Mongolic speakers have this component together with another component most enriched in populations from the Russian Far East, such as Ulchi and Nivkh (Supplementary Fig. 3). Turkic/Mongolic speakers comprising the bottom-most cline have a distinct Western Eurasian ancestry profile: they have a high proportion of a component most enriched in Mesolithic Caucasus hunter-gatherers and Neolithic Iranians and frequently harbour another component enriched in present-day South Asians (Supplementary Fig. 4). Based on the PCA and ADMIXTURE results, we heuristically assigned inner Eurasians to three clines: the ‘forest-tundra’ cline includes Russians and all Uralic and Yeniseian speakers; the ‘steppe-forest’ cline includes Turkic- and Mongolic-speaking populations from the Volga and Altai–Sayan regions and Southern Siberia; and the ‘southern steppe’ cline includes the rest of the populations.

eurasian-clines-uralic-altaic
The first two PCs summarizing the genetic structure within 2,077 Eurasian individuals. The two PCs generally mirror geography. PC1 separates western and eastern Eurasian populations, with many inner Eurasians in the middle. PC2 separates eastern Eurasians along the northsouth cline and also separates Europeans from West Asians. Ancient individuals (color-filled shapes), including two Botai individuals, are projected onto PCs calculated from present-day individuals.

For the forest-tundra populations, the Nganasan + Srubnaya model is adequate only for the two Volga region populations, Udmurts and Besermyans (Fig. 5 and Supplementary Table 8).

For the other populations west of the Urals, six from the northeastern corner of Europe are modelled with additional Mesolithic Western European hunter-gatherer (WHG) contribution (8.2–11.4%; Supplementary Table 8), while the rest need both WHG and early Neolithic European farmers (LBK_EN; Supplementary Table 2). Nganasan-related ancestry substantially contributes to their gene pools and cannot be removed from the model without a significant decrease in the model fit (4.1–29.0% contribution; χ2 P ≤ 1.68 × 10−5; Supplementary Table 8).

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Supplementary Table 8. QpAdm-based admixture modeling of the forest-tundra cline populations. For the 13 populations west of the Urals, we present a four-way admixture model, Nganasan+Srubnaya+WHG+LBK_EN, or its minimal adequate subset. Modified from the article, to include colors for cultures, and underlined best models for Corded Ware ancestry among Uralians.

NOTE. It doesn’t seem like Hungarians can be easily modelled with Nganasan ancestry, though…

For the 4 populations east of the Urals (Enets, Selkups, Kets and Mansi), for which the above models are not adequate, Nganasan + Srubnaya + AG3 provides a good fit (χ2 P ≥ 0.018; Fig. 5 and Supplementary Table 8). Using early Bronze Age populations from the Baikal Lake region (‘Baikal_EBA’; Supplementary Table 2) as a reference instead of Nganasan, the two-way model of Baikal_EBA + Srubnaya provides a reasonable fit (χ2 P ≥ 0.016; Supplementary Table 8) and the three-way model of Baikal_EBA + Srubnaya + AG3 is adequate but with negative AG3 contribution for Enets and Mansi (χ2 P ≥ 0.460; Supplementary Table 8).

east-urals-ugric-samoyedic-qpadm
Supplementary Table 8. QpAdm-based admixture modeling of the forest-tundra cline populations. For the four populations east of the Urals, we present three admixture models: Baikal_EBA+Srubnaya, Baikal_EBA+Srubnaya+AG3 and Nganasan+Srubnaya+AG3. For each model, we present qpAdm p-value, admixture coefficient estimates and associated 5 cM jackknife standard errors (estimate ± SE). Modified from the article, to include colors for cultures, and underlined best models for Corded Ware ancestry among Uralians.

Bronze/Iron Age populations from Southern Siberia also show a similar ancestry composition with high ANE affinity (Supplementary Table 9). The additional ANE contribution beyond the Nganasan + Srubnaya model suggests a legacy from ANE-ancestry-rich clines before the Late Bronze Age.

bronze-age-iron-age-karasuk-mezhovska-tagar-qpadm
Supplementary Table 9. QpAdm-based admixture modeling of Bronze and Iron Age populations of southern Siberia. For ancieint individuals associated with Karasuk and Tagar cultures, Nganasan+Srubnaya model is insufficient. For all five groups, adding AG3 as the third ancestry or substituting Nganasan with Baikal_EBA with higher ANE affinity provides an adequate model. For each model, we present qpAdm p-value, admixture coefficient estimates and associated 5 cM jackknife standard errors (estimate ± SE). Models with p-value ≥ 0.05 are highlighted in bold face. Modified from the article, to include colors for cultures, and underlined best models for Corded Ware ancestry among Uralians.

Lara M. Cassidy comments the results of the study in A steppe in the right direction (you can read it here):

Even among the earliest available inner Eurasian genomes, east–west connectivity is evident. These, too, form a longitudinal cline, characterized by the easterly increase of a distinct ancestry, labelled Ancient North Eurasian (ANE), lowest in western European hunter-gatherers (WHG) and highest in Palaeolithic Siberians from the Baikal region. Flow-through from this ANE cline is seen in steppe populations until at least the Bronze Age, including the world’s earliest known horse herders — the Botai. However, this is eroded over time by migration from west and east, following agricultural adoption on the continental peripheries (Fig. 1b,c).

Strikingly, Jeong et al. model the modern upper steppe cline as a simple two-way mixture between western Late Bronze Age herders and Northeast Asians (Fig. 1c), with no detectable residue from the older ANE cline. They propose modern steppe peoples were established mainly through migrations post-dating the Bronze Age, a sequence for which has been recently outlined using ancient genomes. In contrast, they confirm a substantial ANE legacy in modern Siberians of the northernmost cline, a pattern mirrored in excesses of WHG ancestry west of the Urals (Fig. 1b). This marks the inhospitable biome as a reservoir for older lineages, an indication that longstanding barriers to latitudinal movement may indeed be at work, reducing the penetrance of gene flows further south along the steppe.

eurasian-clines-uralic-turkic-mongol-altaic
The genomic formation of inner Eurasians. b–d, Depiction of the three main clines of ancestry identified among Inner Eurasians. Sources of admixture for each cline are represented using proxy ancient populations, both sampled and hypothesised, based on the study’s modelling results. The major eastern and western ancestries used to model each cline are shown in bold; the peripheral admixtures that gave rise to these are also shown. Additional contributions to subsections of each cline are marked with dashed lines. b, The northernmost cline, illustrating the legacy of WHG and ANE-related populations. c,d, The upper (c) and lower (d) steppe clines are shown, both of which have substantial eastern contributions related to modern Tungusic speakers. The authors propose these populations are themselves the result of an admixture between groups related to the Nganasan, whose ancestors potentially occupied a wider range, and hunter-gatherers (HGs) from the Amur River Basin. While the upper steppe cline in c can be described as a mixture between this eastern ancestry and western steppe herders, the current model for the southern steppe cline as shown in d is not adequate and is likely confounded by interactions with diverse bordering ancestries. Credit: Ecoregions 2017, Resolve https://ecoregions2017.appspot.com/

Given the findings as reported in the paper, I think it should be much easier to describe different subclines in the “northernmost cline” than in the much more recent “Turkic/Mongolic cline”, which is nevertheless subdivided in this paper in two clines. As an example, there are at least two obvious clines with “Nganasan-related meta-populations” among Uralians, which converge in a common Steppe MLBA (i.e. Corded Ware) ancestry – one with Palaeo-Laplandic peoples, and another one with different Palaeo-Siberian populations:

siberian-clines-uralic-altaic
PCA of ancient and modern Eurasian samples. Ancient Palaeo-Laplandic, Palaeosiberian, and Altai clines drawn, with modern populations labelled. See a version with higher resolution.

The inclusion of certain Eurasian groups (or lack thereof) in the PCA doesn’t help to distinguish these subclines visually, and I guess the tiny “Naganasan-related” ancestral components found in some western populations (e.g. the famous ~5% among Estonians) probably don’t lend themselves easily to further subdivisions. Notice, nevertheless, the different components of the Eastern Eurasian source populations among Finno-Ugrians:

uralic-admixture-qpadm
Characterization of the Western and Eastern Eurasian source ancestries in inner Eurasian populations. [Modified from the paper, includes only Uralic populations]. a, Admixture f3 values are compared for different Eastern Eurasian (Mixe, Nganasan and Ulchi; green) and Western Eurasian references (Srubnaya and Chalcolithic Iranians (Iran_ChL); red). For each target group, darker shades mark more negative f3 values. b, Weights of donor populations in two sources characterizing the main admixture signal (date 1 and PC1) in the GLOBETROTTER analysis. We merged 167 donor populations into 12 groups (top right). Target populations were split into five groups (from top to bottom): Aleuts; the forest-tundra cline populations; the steppe-forest cline populations; the southern steppe cline populations; and ‘others’.

Also remarkable is the lack of comparison of Uralic populations with other neighbouring ones, since the described Uralic-like ancestry of Russians was already known, and is most likely due to the recent acculturation of Uralic-speaking peoples in the cradle of Russians, right before their eastward expansions.

west-eurasian-east-eurasian-ancestry
Supplementary Fig. 4. ADMIXTURE results qualitatively support PCA-based grouping of inner Eurasians into three clines. (A) Most southern steppe cline populations derive a higher proportion of their total Western Eurasian ancestry from a source related to Caucasus, Iran and South Asian populations. (B) Turkic- and Mongolic-speaking populations tend to derive their Eastern Eurasian ancestry more from the Devil’s Gate related one than from Nganasan-related one, while the opposite is true for Uralic- and Yeiseian-speakers. To estimate overall western Eurasian ancestry proportion, we sum up four components in our ADMIXTURE results (K=14), which are the dominant components in Neolithic Anatolians (“Anatolia_N”), Mesolithic western European hunter-gatherers (“WHG”), early Holocene Caucasus hunter-gatherers (“CHG”) and Mala from southern India, respectively. The “West / South Asian ancestry” is a fraction of it, calculated by summing up the last two components. To estimate overall Eastern Eurasian ancestry proportion, we sum up six components, most prevalent in Surui, Chipewyan, Itelmen, Nganasan, Atayal and early Neolithic Russian Far East individuals (“Devil’s Gate”). Eurasians into three clines. (A) Most southern steppe cline populations derive a higher proportion of their total Western Eurasian ancestry from a source related to Caucasus, Iran and South Asian populations. (B) Turkic- and Mongolic-speaking populations tend to derive their Eastern Eurasian ancestry more from the Devil’s Gate related one than from Nganasan-related one, while the opposite is true for Uralic- and Yeiseian-speakers. To estimate overall western Eurasian ancestry proportion, we sum up four components in our ADMIXTURE results (K=14), which are the dominant components in Neolithic Anatolians (“Anatolia_N”), Mesolithic western European hunter-gatherers (“WHG”), early Holocene Caucasus hunter-gatherers (“CHG”) and Mala from southern India, respectively. The “West / South Asian ancestry” is a fraction of it, calculated by summing up the last two components. To estimate overall Eastern Eurasian ancestry proportion, we sum up six components, most prevalent in Surui, Chipewyan, Itelmen, Nganasan, Atayal and early Neolithic Russian Far East individuals (“Devil’s Gate”).

A comparison of Estonians and Finns with Balts, Scandinavians, and Eastern Europeans would have been more informative for the division of the different so-called “Nganasan-like meta-populations”, and to ascertain which one of these ancestral peoples along the ancient WHG:ANE cline could actually be connected (if at all) to the Cis-Urals.

Because, after all, based on linguistics and archaeology, geneticists are not supposed to be looking for populations from the North Asian Arctic region, for “Siberian ancestry”, or for haplogroup N1c – despite previous works by their peers – , but for the Bronze Age Volga-Kama region…

Related

Pre-Germanic and Pre-Balto-Finnic shared vocabulary from Pitted Ware seal hunters

corded-ware-pitted-ware

I said I would write a post about topo-hydronymy in Europe and Iberia based on the most recent research, but it seems we can still enjoy some more discussions about the famous Vasconic Beakers, by people longing for days of yore. I don’t want to spoil that fun with actual linguistic data (which I already summarized) so let’s review in the meantime one of the main Uralic-Indo-European interaction zones: Scandinavia.

Seal hunting

One of the many eye-catching interpretations – and one of the few interesting ones – that could be found in the relatively recent article Talking Neolithic: Linguistic and Archaeological Perspectives on How Indo-European Was Implemented in Southern Scandinavia, by Iversen & Kroonen AJA (2017) was this:

The borrowing of lexical items from hunter-gatherers into Germanic refers to the potential adoption of Proto-Germanic *selhaz “seal” (Old Norse selr, Old English seolh, Old High German selah) as well as Early Proto-Balto-Finnic *šülkeš “seal” (Finnish hylje, Estonian hüljes) from the marine-oriented Sub-Neolithic Pitted Ware culture.

kroonen-iversen
Modified from Kristiansen et al. (2017), with red circle around the hypothesized interaction of Germanic with hunter-gatherers. “Schematic representation of how different Indo-European branches have absorbed words (circles) from a lost Neolithic language or language group (dark fill) in the reconstructed European linguistic setting of the third millennium BC, possibly involving one or more hunter gatherer languages (light fill) (after Kroonen & Iversen 2017)”.

This is what Kroonen thought about this word in his Etymological Dictionary of Proto-Germanic (2006):

Gmc. *selha– m. ‘seal’ – ON selr m. ‘id.’, Far. selur m. ‘id.’, OSw. siæl m. ‘id.’, Sw. själ c. ‘id.’, OE seolh m. ‘id.’, E seal, OS selah m. ‘id.’, EDu. seel, seel-hont m. ‘id.’, Du. zee-hond c. ‘id.’, OHG selah m. ‘id.’, MHG sele m. ‘id.’ (GM).

A Germanic word with no certain IE etymology. The link with Lith. selė́ti ‘to crawl’ (Torp 1909: 436) is erroneous, as this verb corresponds to PGm. *stelan- (q.v.). The *h may nevertheless correspond to the PIE animal suffix *-ko-, for which see *elha{n)- ‘elk’ and *baruga- ‘boar’.

Focusing on this substrate etymon, coupled with archaeology and ancient DNA, in the recent SAA 84th Annual Meeting (Abstracts in PDF):

Kroonen, Guus (Leiden University) and Rune Iversen

[196] The Linguistic Legacy of the Pitted Ware Culture

The Scandinavian hunter-, fisher- and gatherer-based Pitted Ware culture is chronologically situated in the Neolithic. However, it challenges our traditional view on cultural and social evolution by representing a return to an otherwise abandoned hunter-gatherer lifestyle. In general, the Pitted Ware culture must be seen as an offshoot of the “Sub-Neolithic” societies inhabiting wide parts of northern and northeastern Europe in the fourth and third millennium B.C.E.

Isotopic and aDNA studies have shown that people of the east Swedish Pitted Ware culture, both dietarily and genetically were distinct from the early farmers in this region, the Funnel Beaker culture. Isotopic data shows a marked predominance of seal in the diet, which has given the Pitted Ware people the nickname “Inuit of the Baltic”.

As regards language, it is to be expected that people practicing a Pitted Ware lifestyle spoke a non-Indo-European language. In fact, there is some linguistic evidence that can support this claim. It is conceivable that both the Germanic and Finnish word for “seal” were ultimately borrowed from a language spoken in a Pitted Ware context. Once more, the linguistic evidence turns out to offer important information complementary to that of archaeology and archaeo-genetics.

prehistoric-seal-hunters
Stone Age Seal Hunters, by Måns Sjöberg.

Apparently, the idea of non-IE substrate languages in contact with Germanic in Scandinavia is fashionable for the Copenhagen group, probably due to their particular interpretation of the recent genetic papers, hence the multiple Germanic-Fennic connections to be reviewed through this new prism. While the ulterior motive of this proposal may be to try and connect yet again Germanic with CWC Denmark, I would argue that the effect is actually the opposite.

An early borrowing via Uralic

The word has always been considered a more likely loan from one language to the other, and – because of the quite popular idea of Uralic native to Fennoscandia – it was often seen as a likely borrowing of Germanic from Balto-Finnic. In any possible case, the borrowing in either direction must be quite early, for obvious reasons:

  • If the borrowing had been via late Palaeo-Germanic, the ending in *-xa– would have been reflected in Balto-Finnic, hence an early Palaeo-Germanic to Pre-Balto-Finnic stage would be necessary.
  • If the borrowing had been via late Balto-Finnic, the initial sibilant would be already aspirated, being adopted as *-x– in Palaeo-Germanic, while the ending in *-k– would have remained as such if it was adopted after Grimm’s law ceased to be active.
  • Similarly, a borrowing from a common, non-Indo-European & non-Uralic source would require that it happened during the early stages of both proto-languages to have undergone their respective phonetic changes, and both borrowings chronologically close to each other, to assume a similar vocalism and consonantism of the ultimate source.
wiik-indo-european-uralic-substrate
The idea of seal-hunting Uralic substrate of Pitted Ware is not new. Image modified from The Uralic and Finno-Ugric Phonetic Substratum, by Kalevi Wiik, Linguistica Uralica (1997).

Furthermore, regarding the most likely way of expansion of this loanword, due to the different vowels and sibilants present in Uralic but not in Indo-European:

  • A direct loan from Pre-Germanic **selkos – which shows a regular thematic declension – to Pre-Balto-Finnic *šülkeš doesn’t seem to be a reasonable assumption.
  • NOTE. A Germanic borrowing from alternative Gmc. genitive *silxis could only work in a Pre-Germanic to Pre-Balto-Finnic model, hence only if the Gmc. form can be reconstructed for an earlier stage. Even then, for the same reason stated above, the opposite could be more reasonably argued, i.e. that this form is the original one adopted in Germanic: Pre-PBF *šülkeš > Pre-Gmc. *silkis, reinterpreted as an -o- stem in its declension.

  • If we reconstruct an older Pre-Finno-Samic (i.e. with Finno-Permic-like vocalism) **šëlkëš, a borrowing into Pre-Germanic **selkos would work. Even though no Saami derivative exists to confirm such a possibility, this would be supported by the known common evolution of Finno-Samic dialects in close contact with Pre-Germanic.
  • Admittedly, even accepting the existence of a Finno-Samic stem, a potential substrate word could not be discarded. In fact, while **šëlkë- could perfectly be a Uralic root, the ending in *-š can’t be easily interpreted. Therefore, a third, non-Indo-European & non-Uralic source is a plausible explanation.

NOTE. Arguably, Proto-Finno-Samic could have adopted Gmc. *kh or *x exceptionally as PFS *k. However, early Palaeo-Germanic borrowings in Finno-Samic show a consistent regular consonant change as described above. For more on this, see Finno-Samic borrowings.

This likely Uralic first nature of the loanword is important for the discussion below.

Pitted Ware culture

pitted-ware-pyheensilta-ware-culture
Middle Neolithic A period. Distribution of Pyheensilta Ware, Funnel Beaker Culture in Sweden, and Pitted Ware Culture in northern Europe during the Middle Neolithic A period, c. 3300–2800 cal BC. Find locations with numbers demarcate sites where cereal grains have been found and later AMS radiocarbon dated. Figure was created by SV using QGIS 3.4. (https://www.qgis.org/) and Natural Earth data (https://www.naturalearthdata.com/). Image from Vanhanen et al. (2019).

About the Pitted Ware culture, this is what the recent paper by Vanhanen et al. (2019), from the University of Finland (including Volker Heyd) had to say:

The origins of the PWC are controversial. In one likely scenario, Comb Ceramic and Mesolithic hunter-gatherers first interacted with FBC during the last centuries of the EN and became specialized maritime hunter-gatherers. The PWC pushed south and westwards during the Middle Neolithic (MN), c. 3300–2300 BC, along the northern Baltic shoreline and adjacent islands, eventually reaching as far west as Denmark and southern Norway. Around 2800 BC, after the FBC ceased to exist, the Corded Ware Culture (CWC) migrated into the PWC area. The end date for the PWC and CWC is approximately 2300 BC, when the material culture was replaced by the Late Neolithic (LN) culture<. Spanning nearly a millennium virtually unchanged, the PWC maintained a coherent society and a successful economic model. PWC people lived in marine-oriented settlements, commonly dwelled in huts and produced relatively large amounts of ceramic vessels. This speaks to the partly sedentary nature of their habitation, at least for their base camps. These specialist hunter-gatherers obtained the great majority of their subsistence from maritime sources, such as seal, fish, and sea birds. Considering the amount of bones, sealing was of paramount importance, causing these peoples to be labelled ‘hard-core sealers’ or even the ‘Inuit of the Baltic’.

The Middle Neolithic Pitted Ware culture is dated ca. 3500–2300 BC, so we would be seeing here Pre-Germanic and Pre-Balto-Finnic peoples arriving near the Pitted Ware culture. That would leave us with one of both languages expanding with Corded Ware peoples, and the other with Bell Beakers. Since Battle Axe-derived cultures around the Gulf of Finland are associated with Balto-Finnic groups, and Bell Beakers arriving ca. 2400 started the Dagger Period, commonly associated with the Pre-Germanic community, I think the connection of each group with their language is self-evident.

pitted-ware-cored-ware-culture
Middle Neolithic B period. Distribution of Corded Ware Culture and Pitted Ware Culture in northern Europe during the Middle Neolithic B period, c. 2800–2300 cal BC. Find locations with numbers demarcate sites where cereal grains have been found and later AMS radiocarbon dated. Figure was created by SV using QGIS 3.4. (https://www.qgis.org/) and Natural Earth data (https://www.naturalearthdata.com/). Modified from Vanhanen et al. (2019).

NOTE. You can read some interesting information about prehistoric and recent seal hunting in the Baltic in the blog post “Själen” – Seal Hunting in the Northern Baltic Sea.

Germanic-Fennic phonetic evolution

The common Germanic – Balto-Finnic phonetic evolution, especially Verner’s law in Palaeo-Germanic and qualitative gradation in Proto-Balto-Finnic, has been variably interpreted as:

  • Uralic in Scandinavia influenced by Germanic (Verner’s law source of the gradation), by Koivulehto and Vennemann (1996).
  • Germanic over a Uralic substratum in Scandinavia, by Wiik (1997).
  • Both Germanic and Balto-Finnic influenced by a third language, an “extinct non-Uralic source” spoken in Fennoscandia before the arrival of Uralic and Indo-European, by Kallio (2001); maybe the same substrate proposed to have influenced the accent shift in Germanic similar to Uralic.
  • Balto-Finnic speakers adopting Pre-Germanic in Scandinavia, in contact with Balto-Finnic speakers retaining their language, by Schrijver in Language Contact and the Origins of the Germanic Languages (2014)– although first suggested by him in the 1990s.

NOTE. There are other (some much older) proposals of a Uralic substrate in Scandinavia, but I think those above summarize the most common positions tenable today.

If you add all linguistic, archaeological, and now genetic connections, it is really strange to keep arguing for so many surprisingly fitting common substrates and/or contact languages for both. Especially because the Pre-Germanic community – if originally from southern Scandinavia and not further south (see e.g. Kortlandt’s theory) – was marked by the Dagger Period, as accepted by most archaeologists (including Kristiansen), and we know that Bell Beakers – who triggered the Dagger period – might have arrived a little late to the Pitted Ware disintegration in most seal-hunting areas of southern Scandinavia.

bell-beaker-density
Density analysis based (Bell Beaker per km2) on the distribution of Bell Beaker per region (ca. 2700-2200 BC). Combination of different levels of b-spline interpolation. Exaltation of the values through square root usage. Modified from Michael Bilger (2018).

In other words, how many common substrate languages can we propose for Germanic (and Balto-Finnic)? Just from Kroonen we have already the Semitic-like TRB, and the seal-hunting Pitted Ware culture. Apparently, the culprit of the common phonetic evolution must be some (other?) culture that both Pre-Germanic and Pre-Balto-Finnic assimilated (or with which both were in contact) in Fennoscandia.

NOTE. I believe no data supports the attribution of those Germanic borrowings to the TRB culture, especially if one assumes they belong to an Afroasiatic branch, as did Kroonen. His initial assumption about an expansion of R1b-M269 associated with the Neolithic from Anatolia, and thus with Afroasiatic, must today be rejected. Much more likely is the incorporation of most of these loanwords during the expansion of North-West Indo-Europeans from Yamna Hungary.

How many “common” substrates from different regions and cultures is too much? Arguably, it’s not a question of quantity (because the overall probability remains the same), but a question of quality of arguments.

In my opinion, both a) the marked seal-hunting subsistence economy of the Pitted Ware culture and b) the difficult reconstruction of a fitting ‘natural’ PIE or PU stem warrant this proposal of a third source, just like the European agricultural substrate of North-West Indo-European and Palaeo-Balkan languages, as well as the Asian agricultural substrate of Indo-Iranian are the most logical interpretation of words not found in other IE dialects. The only problem in this case is the lack of other Scandinavian substrate words to compare its typology against.

scandinavia-neolithic-flint-daggers
Close contacts in Fennoscandia. The distribution of Scandinavian flint daggers (A) in the east and south Baltic region and possible trends of “down the line” trade (B). Good size and quality flint zone in the south-west Baltic region is hatched (C). According to: Wojciechowski 1976; Olausson 1983, fig. 1; Madsen 1993, 126; Libera 2001; Kriiska & Tvauri 2002, 86. Image modified from Piličiauskas (2010).

Common Scandinavian substratum

The theory of a Pitted Ware borrowing is therefore quite convincing from a cultural point of view, at the same time as it fits the linguistic data. However, one reason why I dislike the interpretation of a dual origin is that our knowledge of Uralic languages is fairly limited, whereas that of Indo-European branches and hence Proto-Indo-European is huge. To put it otherwise: if a common word appears in both, and it is most likely (culturally and linguistically) not Indo-European, it certainly means that it was borrowed in Germanic. What are the a priori chances of it coming directly from a third substrate language for both dialects, instead of coming directly from Pre-Balto-Finnic?

From Schrijver (2014):

What did happen, apparently, is that Finnic speakers had enough access to the way in which Germanic speakers pronounced Balto-Finnic in order to model their own pronunciation of Balto-Finnic on it. In other words, Balto-Finns conversed with bilingual speakers of Germanic and Balto-Finnic whose pronunciation of both was essentially Germanic. But access to the Germanic language itself was not sufficient to allow Balto-Finns to become bilingual themselves, either because social segregation prevented this or because contact with Germanic was severed before widespread bilingualism set in. This limited access to Germanic would allow us to understand why Balto-Finnic did not go the way of the vernacular languages that came in contact with Latin in the Roman Empire, where access to Latin was open to almost everybody and massive language shift in favour of Latin ensued.

NOTE. For a more detailed discussion, you can read the whole chapter dedicated to this question. I summarized it in Pre-Germanic born out of a Proto-Finnic substrate in Scandinavia.

On the other hand, about the ad hoc interpretation by Kallio (2001) of hypothetic third languages strongly influencing in the same way both the Palaeo-Germanic- and Balto-Finnic-speaking communities, Schrijver (2014) comments:

The idea that perhaps both languages moved towards a lost third language, whose speakers may have been assimilated to both Balto-Finnic and Germanic, provides a fuller explanation but suffers from the drawback that it shifts the full burden of the explanation to a mysterious ‘language X’ that is called upon only in order to explain the developments in Proto-Germanic and Balto-Finnic. That comes dangerously close to circular reasoning.

early-bronze-age-nordic-dagger-period
Early Bronze Age cultures of Northern Europe (roughly ca. 2200-1750). Dagger period representing the expansion of BBC-derived groups from southern Scandinavia.

NOTE. The proposal of some kind of “SHG/EHG-based Fennoscandian substrate” seems funny to me, for two reasons: firstly, there is usually no talk about which culture spread that common language, how it survived, how it was in contact with both groups and until when, etc. (see below for possibilities); secondly, apparently the evident survival of West European EEF communities driven by at least two cultural groups – El Argar and the poorly known groups from the Atlantic façade north of the Pyrenees – is, for the same people proposing this simplistic SHG/EHG idea, somehow not fitting for the prehistory of Proto-Iberian and Proto-Aquitanian, respectively…

The same argument that one could use against the direct borrowing of both dialects from Pitted Ware, but much more strongly, can be thus wielded against a common, centuries-long phonetic evolution of both Balto-Finnic and Germanic caused by close interactions with (and/or substrate influence of) some third language. Which unitary culture and when exactly could that have happened around the Baltic Sea?

  • Was it Pitted Ware the mysterious substrate language? Seems rather unlikely, due to the early demise of the Pitted Ware culture in contrast to the long-lasting common influence seen in both dialects.
  • Was it Pitted Ware in southern Scandinavia, but Comb Ware in the Gulf of Finland? Is there a direct genetic connection between both cultures? And how likely is a common phonology of an ancestral Comb Ware-like substrate language surviving separately in Finland and Sweden? Even accepting these assumptions, we would be stuck again in the Indo-European Beakers vs. Uralic Battle Axe model.
  • Was it a succession of cultures, from some Scandinavian culture that was replaced by some incoming ethnolinguistic group, then influencing the other? This non-IE, non-Uralic substrate would then need to be proposed, given the chronological and archaeological constraints, as an effect of Pitted Ware over Pre-Finno-Baltic spoken by Battle Axe peoples in Scandinavia, then replaced by Pre-Germanic peoples arriving later with Bell Beakers. A reverse direction and later chronology (say, Germanic replaced by Balto-Finnic from Netted Ware arriving from the Volga) wouldn’t work as well.
  • Was it Asbestos Ware as a late Comb Ware group influencing both? How likely is such a continued influence in Southern Scandinavia and the Gulf of Finland? Even if we accepted this influence that miraculously didn’t affect Samic (most likely located between the Balto-Finnic-speaking Gulf of Finland and northern Fennoscandian Asbestos Ware groups), it would necessarily mean that Germanic and Balto-Finnic were spoken neighbouring exactly the same Asbestos Ware groups in Scandinavia. That is, essentially, that the BBC-derived Dagger Period represented Pre-Germanic, while Battle Axe-derived groups around the Gulf of Finland were Balto-Finnic.

Mixing linguistics with archaeology (now complemented with genetics) also risks circular reasoning. But, how else can someone propose a third substrate language for a phonetic change, necessarily represented by Fennoscandian groups potentially separated by thousands of years? In this age of population genomics we can’t simply talk about theoretical models anymore: we must refer to Fennoscandian cultures and populations in a very specific time frame, as Kronen & Iversen do in their proposal. Not only is such a third unknown language usually a weak explanation for a common development of two unrelated languages; in this case it finds no support whatsoever.

Seals and the Arctic

Another interesting aspect about this Fennic-Germanic comparandum is its relevance to the Uralic homeland problem.

uralic-languages-modern
Current distribution of Uralic languages. Nenets and Saami are among the best positioned to retain the ‘original’ Uralic seal-hunting vocabulary.

Since the publication of Mittnik et al. (2018), Lamnidis et al. (2018), and Sikora et al. (2018), the new normal is apparently to consider Corded Ware Finland as Germanic-speaking, the Gulf of Finland as Balto-Slavic-speaking, while the Kola peninsula and whichever Palaeo-Arctic peoples preceded Nganasans and Nenets as ancient Uralians. Uh-huh, OK.

But, if prehistoric Arctic peoples practiced specialized seal-hunting economies, and Uralians were one among such populations – supposedly one widespread from the Barents Sea to the Lapteve Sea…how come no common Uralic word for ‘seal’ exists? In other words, why would these True™ Uralic peoples expanding from the Arctic need to borrow a word for ‘seal’ from neighbouring populations in every single seal-hunting region they are attested?

grey-seal-distribution
Historical distribution of grey seals, an important part of the diet around the Baltic Sea. Image modified from Wikimedia to include Skagerrak and Kattegat regions.

About Saami, which some have recklessly proposed to be derived from Bronze Age N1c-L392 samples from the Kola Peninsula (against the good judgment of the authors of the paper), this is what we know from their word for ‘seal’, from Grünthal (2004):

Ter Saami vīrre ‘seal; wolf’ displays two meanings that refer to clearly different animals. Neither of them is borrowed from the source language because the word descends from Russian zver’ ‘animal’ (T.I.Itkonen 1958: 756). Another word, Skolt Saami näúdd ‘seal, wolf’, has been similarly used in the two meanings. The evidence of North Saami návdi ‘wolf; creature, fur animal; beast’ (Sammallahti 1989: 305; Lagercrantz (1939: 518) presents the alternative meanings in the opposite order; E. Itkonen (1969: 148) lists the meanings ‘wildes Tier; Raubtier (bes. Wolf); Pelztier’) suggesting that ‘wolf’ is the primary sense and ‘seal’ is a metaphorical extension of it. More precisely, it is an example of a mythic metaphor (cf. Siikala 1992). According to the old folk belief, seal was a wolf and the Skolt Saamis preferred not to eat its meat (T.I.Itkonen 1958: 906). Before that the metonymic meaning ‘wolf’ rose from the less specified meanings, and originally návdi is a Scandinavian or Finnic loan word in Saamic, cf. Old Norse naut ‘vieh, rind’, Icelandic and Norwegian naut, Swedish nöt < Germanic *nauta ‘property’ (Hellquist 1980: 721, T.I.Itkonen 1958: 275, Lagercrantz 1939: 518, de Vries 1961: 406; E. Itkonen (1969: 148) considers Finnic, cf. Finnish nauta ‘bovine’ (< Germanic) as a possible alternative source for the Saamic word).

NOTE. Possibly comparable, for the mythic metaphor proper of Scandinavian folk belief, are Germanic derivatives built as ‘seal-hound’ and/or ‘sea-hound’.

sea-distribution-arctic
Seals formed a great part of the diet for Palaeo-Arctic populations. Boundaries of regions used to predict sea ice, superimposed over the distributions of the five ringed seal subspecies. Image modified from Kelly et al. (2010).

About Nenets (quite close to the Naganasans of pure “Siberian ancestry”), here is what Edward Vajda, an expert in Palaeo-Siberian languages, has to say:

Nenets techniques for hunting the animals of the Arctic Ocean seem to have been borrowed from the first Arctic aborigines. Thus, the Nenets word for seal is nyak, the Eskimo word is nesak. Also, the Nenets word for a one-piece Arctic clothing is lu; the Korak word on the Kamchatka peninsula for clothing is l’ku. All of these groups may have borrowed the words from some original circumpolar aborigines. More probably, the first settlers of Arctic Europe were cousins of the present-day Eskimo, Chukchi and other residents of the far northeast region of Asia. Nenets folklore also speaks of the aborigines living in ice dugouts (igloos).

On the other hand, Proto-Uralic shows a Chalcolithic steppe-like culture, with common words for metal and metalworking, for agriculture, and for domesticated animals, most likely including cattle. They were close to Indo-Europeans since at least before the Tocharian split, and probably earlier than that (even if one does not accept the Indo-Uralic phylum). And there were clearly strong contacts of Finno-Ugric with Indo-Iranian, and especially of Finno-Samic with Germanic.

uralic-cline
Uralic clines from Corded Ware groups to the east. A clear reason for the lack of common seal-hunting vocabulary. Modified from Tambets et al. (2018). Principal component analysis (PCA) and genetic distances of Uralic-speaking populations. a PCA (PC1 vs PC2) of the Uralic-speaking populations. You can see another PCA including ancient samples.

Some among my readers may now be thinking about these totally believable proposals of prehistoric cultures around Lake Baikal representing the True™ Uralic homeland; because haplogroup N1c, and because some 0.5% more “Devil’s Gate Cave ancestry” in Estonians than in Lithuanians; despite the fact that 1) the so-called “Siberian ancestry” formed an ancestral cline with EHG in North Eurasia, that 2) N1c-L392 lineages seem to appear among many Asian peoples of different languages, and that 3) recent prehistoric N1c-L392 lines expanded clearly with Micro-Altaic languages.

Like, who would have hunted seals in Lake Baikal, right? The problem is, seals represented one of their main game, essential for their subsistence economy. From Novokonova et al. (2015):

One of the key reasons for the density of human settlement in the Baikal region compared to adjacent areas of Siberia is that the lake and its nearby rivers offer an abundance of aquatic food resources, including several endemic species, with perhaps the most well known being the Baikal seal. This freshwater seal is only found in Lake Baikal and portions of its tributaries. It shares lifecycle and behavioral patterns with other small northern ice-adapted seals, and is genetically and morphologically most closely related to the ringed seal (Pusa hispida). The nerpa can grow up to 1.8 m long and weigh as much as 130 kg, with the males tending to be slightly larger than the females.

Zooarchaeological analyses of the 16,000 Baikal seal remains from this well-dated site clearly show that sealing began here at least 9000 calendar years ago. The use of these animals at Sagan-Zaba appears to have peaked in the Middle Holocene, when foragers used the site as a spring hunting and processing location for yearling and juvenile seals taken on the lake ice. After 4800 years ago, seal use declined at the site, while the relative importance of ungulate hunting and fishing increased. Pastoralists began occupying Sagan-Zaba at some point during the Late Holocene, and these groups too utilized the lake’s seals. Domesticated animals are increasingly common after about 2000 years ago, a pattern seen elsewhere in the region, but spring and some summer hunting of seals was still occurring. This use of seals by prehistoric herders mirrors patterns of seal use among the region’s historic and modern groups.

Bronze Age movements in Fennoscandia

Regarding the shrinkage and expansion of different farming economic strategies in Scandinavia since the Neolithic, with potential relevance for population movements and thus ethnolinguistic change – either from Balto-Finnic peoples migrating back from eastern Sweden, or Germanic peoples moving to eastern Finland – from Vanhanen et al. (2019):

Cultivated plants at CWC sites in Finland were not discovered in the current investigation (Supplementary Results) or earlier studies. In Finland, the keeping of domestic animals is indicated by the evidence of dairy lipids and mineralized goat hairs. Charred remains and impressions of cultivated plants have been discovered at CWC sites in Estonia and east-central Sweden (Fig. 3: 12). In the eastern Baltic region, the earliest bones of domestic animals and a shift in subsistence occurred with the CWC. Whether CWC produced the cereals and other agricultural products found at PWC sites is difficult to estimate because only small amounts of plant remains have ever been discovered at CWC sites. The CWC seemingly reached east-central Sweden from regions further to the east, where there is evidence of animal husbandry, but only very few signs of plant cultivation.

For the Late Neolithic (LN), cereal grains have been found north of Mälaren and along the Norrland coast. In mainland Finland, the first cereal grains occur during the LN or Bronze Age, c. 1900–1250 cal BC. The earliest bones of sheep/goat from mainland Finland are earlier, dating back to 2200–1950 cal BC. Finds of Scandinavian bronze artefacts indicate an influx from east-central Sweden, which might well be a source area for these agricultural innovations. A similar development is found in the eastern Baltic region, where the earliest directly radiocarbon-dated cereals originate from the Bronze Age, 1392–1123 cal BC (2 sigma). Thus, agriculture was evident during the Bronze Age in the eastern Baltic, but at least animal keeping and probably crop cultivation were present earlier during the CWC phase.

It has been known for a while already that the only options left for the expansion of Finno-Saami into Fennoscandia are either Battle Axe (continued in Textile Ceramics) or Netted Ware (as proposed e.g. by Parpola), based, among other data, on language contacts, language estimates, cultural evolution, and population genomics. Data like this one on seal-hunting vocabulary also support the most likely option, which entails the identification of Corded Ware as the vector of expansion of Uralic languages.

NOTE. Also interesting in this regard is the lack of Slavic words for ‘seal’ – borrowed, in Russian from Samic, and in other Slavic dialects from Russian, Latin, or other languages -, and the coinage of a new term in East Baltic. Rather odd for an “autochthonous” Proto-Baltic (supposedly in contact with Pitted Ware, Germanic, and Balto-Finnic, then), and for a Proto-Slavic stemming from the Baltic. Quite appropriate, though, for a Proto-East Baltic arriving in the Baltic with Trzciniec and for a Proto-Slavic community evolving further south.

So, what new episode in this renewed 2000s R1b/R1a/N1c soap opera is it going to be, when eastern Fennoscandia shows Corded Ware-derived peoples of “steppe ancestry” (and mainly R1a-Z645 lineages) continue during the Bronze Age? Will the resurge and/or infiltration of I2 – maybe even N1c – lineages among Corded Ware-derived cultures of north-eastern Europe support or challenge this model, and why? Make your bet below.

Related

The cradle of Russians, an obvious Finno-Volgaic genetic hotspot

pskov-novgorod-russia

First look of an accepted manuscript (behind paywall), Genome-wide sequence analyses of ethnic populations across Russia, by Zhernakova et al. Genomics (2019).

Interesting excerpts:

There remain ongoing discussions about the origins of the ethnic Russian population. The ancestors of ethnic Russians were among the Slavic tribes that separated from the early Indo-European Group, which included ancestors of modern Slavic, Germanic and Baltic speakers, who appeared in the northeastern part of Europe ca. 1,500 years ago. Slavs were found in the central part of Eastern Europe, where they came in direct contact with (and likely assimilation of) the populations speaking Uralic (Volga-Finnish and Baltic- Finnish), and also Baltic languages [11–13]. In the following centuries, Slavs interacted with the Iranian-Persian, Turkic and Scandinavian peoples, all of which in succession may have contributed to the current pattern of genome diversity across the different parts of Russia. At the end of the Middle Ages and in the early modern period, there occurred a division of the East Slavic unity into Russians, Ukrainians and Belarusians. It was the Russians who drove the colonization movement to the East, although other Slavic, Turkic and Finnish peoples took part in this movement, as the eastward migrations brought them to the Ural Mountains and further into Siberia, the Far East, and Alaska. During that interval, the Russians encountered the Finns, Ugrians, and Samoyeds speakers in the Urals, but also the Turkic, Mongolian and Tungus speakers of Siberia. Finally, in the great expanse between the Altai Mountains on the border with Mongolia, and the Bering Strait, they encountered paleo-Asiatic groups that may be genetically closest to the ancestors of the Native Americans. Today’s complex patchwork of human diversity in Russia has continued to be augmented by modern migrations from the Caucasus, and from Central Asia, as modern economic migrations take shape.

pskov-novgorod-pca-eurasia-yakut
Sample relatedness based on genotype data. Eurasia: Principal Component plot of 574 modern Russian genomes. Colors reflect geographical regions of collection; shapes reflect the sample source. Red circles show the location of Genome Russia samples.

In the current study, we annotated whole genome sequences of individuals currently living on the territory of Russia and identifying themselves as ethnic Russian or as members of a named ethnic minority (Fig. 1). We analyzed genetic variation in three modern populations of Russia (ethnic Russians from Pskov and Novgorod regions and ethnic Yakut from the Sakha Republic), and compared them to the recently released genome sequences collected from 52 indigenous Russian populations. The incidence of function-altering mutations was explored by identifying known variants and novel variants and their allele frequencies relative to variation in adjacent European, East Asian and South Asian populations. Genomic variation was further used to estimate genetic distance and relationships, historic gene flow and barriers to gene flow, the extent of population admixture, historic population contractions, and linkage disequilibrium patterns. Lastly, we present demographic models estimating historic founder events within Russia, and a preliminary HapMap of ethnic Russians from the European part of Russia and Yakuts from eastern Siberia.

pskov-novgorod-pca-finno-permic
Sample relatedness based on genotype data. Western Russia and neighboring countries: Principal Component plot of 574 modern Russian genomes. Colors reflect geographical regions of collection; shapes reflect the sample source. Red circles show the location of Genome Russia samples.

The collection of identified SNPs was used to inspect quantitative distinctions among 264 individuals from across Eurasia (Fig. 1) using Principal Component Analysis (PCA) (Fig. 2). The first and the second eigenvectors of the PCA plot are associated with longitude and latitude, respectively, of the sample locations and accurately separate Eurasian populations according to geographic origin. East European samples cluster near Pskov and Novgorod samples, which fall between northern Russians, Finno-Ugric peoples (Karelian, Finns, Veps etc.), and other Northeastern European peoples (Swedes, Central Russians, Estonian, Latvians, Lithuanians, and Ukrainians) (Fig. 2b). Yakut individuals map into the Siberian sample cluster as expected (Fig. 2a). To obtain an extended view of population relationships, we performed a maximum likelihood-based estimation of ancestry and population structure using ADMIXTURE [46](Fig. 2c). The Novgorod and Pskov populations show similar profiles with their Northeastern European ancestors while the Yakut ethnic group showed mixed ancestry similar to the Buryat and Mongolian groups.

pskov-novgorod-yakut-admixture
Population structure across samples in 178 populations from five major geographic regions (k=5). Samples are pooled across three different studies that covered the territory of Russian Federation (Mallick et al. 2016 [36], Pagani et al. 2016 [37], this study). The optimal k-value was selected by value of cross validation error. Russian samples from all studies (highlighted in bold dark blue) show a slight gradient from Eastern European (Ukrainian, Belorussian, Polish) to North European (Estonian Karelian, Finnish) structures, reflecting population history of northward expansion. Yakut samples from different studies (highlighted in bold red) also show a slight gradient from Mongolian to Siberian people (Evens), as expected from their original admixture and northward expansions. The samples originated from this study are highlighted, and plotted in separated boxes below.

Possible admixture sources of the Genome Russia populations were addressed more formally by calculating F3 statistics, which is an allele frequency-based measure, allowing to test if a target population can be modeled as a mixture of two source populations [48]. Results showed that Yakut individuals are best modeled as an admixture of Evens or Evenks with various European populations (Supplemental Table S4). Pskov and Novgorod showed admixture of European with Siberian or Finno-Ugric populations, with Lithuanian and Latvian populations being the dominant European sources for Pskov samples.

direction-expansion-russians
The heatmaps of gene flow barriers show for each point at the geographical map the interpolated differences in allele frequencies (AF) between the estimated AF at the point with AFs in the vicinity of this point. The direction of the maximal difference in allele frequencies is coded by colors and arrows.

So, Russians expanding in the Middle Ages as acculturaded Finno-Volgaic peoples.

Or maybe the true Germano-Slavonic™-speaking area was in north-eastern Europe, until the recent arrival of Finno-Permians with the totally believable Nganasan-Saami horde, whereas Yamna -> Bell Beaker represented Vasconic-Caucasian expanding all over Europe in the Bronze Age. Because steppe ancestry in Fennoscandia and Modern Basques in Iberia.

A really hard choice between equally plausible models.

Related

Magyar tribes brought R1a-Z645, I2a-L621, and N1a-L392(xB197) lineages to the Carpathian Basin

hungarian-conquerors-turks

The Nightmare Week of “N1c=Uralic” proponents continues, now with preprint Y-chromosome haplogroups from Hun, Avar and conquering Hungarian period nomadic people of the Carpathian Basin, by Neparaczki et al. bioRxiv (2019).

Abstract:

Hun, Avar and conquering Hungarian nomadic groups arrived into the Carpathian Basin from the Eurasian Steppes and significantly influenced its political and ethnical landscape. In order to shed light on the genetic affinity of above groups we have determined Y chromosomal haplogroups and autosomal loci, from 49 individuals, supposed to represent military leaders. Haplogroups from the Hun-age are consistent with Xiongnu ancestry of European Huns. Most of the Avar-age individuals carry east Eurasian Y haplogroups typical for modern north-eastern Siberian and Buryat populations and their autosomal loci indicate mostly unmixed Asian characteristics. In contrast the conquering Hungarians seem to be a recently assembled population incorporating pure European, Asian and admixed components. Their heterogeneous paternal and maternal lineages indicate similar phylogeographic origin of males and females, derived from Central-Inner Asian and European Pontic Steppe sources. Composition of conquering Hungarian paternal lineages is very similar to that of Baskhirs, supporting historical sources that report identity of the two groups.

Interesting excerpts (emphasis mine):

All N-Hg-s identified in the Avars and Conquerors belonged to N1a1a-M178. We have tested 7 subclades of M178; N1a1a2-B187, N1a1a1a2-B211, N1a1a1a1a3-B197, N1a1a1a1a4-M2118, N1a1a1a1a1a-VL29, N1a1a1a1a2-Z1936 and the N1a1a1a1a2a1c1-L1034 subbranch of Z1936. The European subclades VL29 and Z1936 could be excluded in most cases, while the rest of the subclades are prevalent in Siberia 23 from where this Hg dispersed in a counter-clockwise migratory route to Europe (…). All the 5 other Avar samples belonged to N1a1a1a1a3-B197, which is most prevalent in Chukchi, Buryats, Eskimos, Koryaks and appears among Tuvans and Mongols with lower frequency.

haplogroup-n-pca
First two components of PCA from Hg N1a subbranch distribution in 51 populations including Avars and Conquerors. Colors indicate geographic regions. Three letter codes are given in Supplementary Table S5.

By contrast two Conquerors belonged to N1a1a1a1a4-M2118, the Y lineage of nearly all Yakut males, being also frequent in Evenks, Evens and occurring with lower frequency among Khantys, Mansis and Kazakhs.

Three Conqueror samples belonged to Hg N1a1a1a1a2-Z1936 , the Finno-Permic N1a branch, being most frequent among northeastern European Saami, Finns, Karelians, as well as Komis, Volga Tatars and Bashkirs of the Volga-Ural region.Nevertheless this Hg is also present with lower frequency among Karanogays, Siberian Nenets, Khantys, Mansis, Dolgans, Nganasans, and Siberian Tatars.

The west Eurasian R1a1a1b1a2b-CTS1211 subclade of R1a is most frequent in Eastern Europe especially among Slavic people. This Hg was detected just in the Conqueror group (K2/18, K2/41 and K1/10). Though CTS1211 was not covered in K2/36 but it may also belong to this sub-branch of Z283.

Hg I2a1a2b-L621 was present in 5 Conqueror samples, and a 6th sample form Magyarhomorog (MH/9) most likely also belongs here, as MH/9 is a likely kin of MH/16 (see below). This Hg of European origin is most prominent in the Balkans and Eastern Europe, especially among Slavic speaking groups. It might have been a major lineage of the Cucuteni-Trypillian culture and it was present in the Baden culture of the Chalcolithic Carpathian Basin.

hungarian-conquerors-y-dna
Image modified from the paper, with drawn red square around lineages of likely Ugric origin, and squares around R1a-Z93, R1a-Z283, N1a-Z1936, and N1a-M2004 samples. Y-Hg-s determined from 46 males grouped according to sample age, cemetery and Hg. Hg designations are given according to ISOGG Tree 2019. Grey shading designate distinguished individuals with rich grave goods, color shadings denote geographic origin of Hg-s according to Fig. 1. For samples K3/1 and K3/3 the innermost Hg defining marker U106* was not covered, but had been determined previously.

We identified potential relatives within Conqueror cemeteries but not between them. The uniform paternal lineages of the small Karos3 (19 graves) and Magyarhomorog (17 graves) cemeteries approve patrilinear organization of these communities. The identical I2a1a2b Hg-s of Magyarhomorog individuals appears to be frequent among high-ranking Conquerors, as the most distinguished graves in the Karos2 and 3 cemeteries also belong to this lineage. The Karos2 and Karos3 leaders were brothers with identical mitogenomes 11 and Y-chromosomal STR profiles (Fóthi unpublished). The Sárrétudvari commoner cemetery seems distinct from the others, containing other sorts of European Hg-s. Available Y-chromosomal and mtDNA data from this cemetery suggest that common people of the 10th century rather represented resident population than newcomers. The great diversity of Y Hg-s, mtDNA Hg-s, phenotypes and predicted biogeographic classifications of the Conquerors indicate that they were relatively recently associated from very diverse populations.

Surprising about the Hungarian conquerors – although in line with the historical accounts – is the varied patrilineal origin of clans, including Q1a, G2a2b, I1, E1b1b, R1b, J1, or J2 – some of which (depending on specific lineages) may have appeared earlier in the Carpathian Basin or south-eastern Europe.

However, out of the 27 conqueror elite samples, 17 are of haplogroups most likely related to Ugric populations beyond the Urals: R1a-Z645, I2-L621, and two specific N1a-L392 lineages (see below). In fact, there are three high-ranking conqueror elites of hg. I2-L621 (one of them termed a “leader”, brother to an unpublished leader of Karos3, and all of them possibly family), one of hg. R1a-Z280, one of hg. R1a-Z93 (which should be added to the Árpáds), and one of hg. N1a-Z1936, which gives a good idea of the ruling class among the elite Ugric settlers.

NOTE. The Q1a sample is also likely to be found in the mixed population of the West Siberian forest-steppes, since it was found in Mesolithic-Neolithic samples from eastern Europe to Lake Baikal, and in Bronze Age Siberian groups, although admittedly it may have formed part of an Avar Transtisza group, or even earlier Hunnic or Scythian groups along the steppes. Without precise subclades it’s impossible to know.

arrival-of-hungarians-arpad
The seven chieftains of the Hungarians, detail of Arrival of the Hungarians, from Árpád Feszty’s and his assistants’ vast (1800 m2) cyclorama, painted to celebrate the 1000th anniversary of the Magyar conquest of Hungary, now displayed at the Ópusztaszer National Heritage Park in Hungary. Image from Wikipedia.

I2a-L621

I2a-L621 (xS17250) or I2a1b2 in the old nomenclature, is found in 6 early conquerors (including one leader), on a par with R1a and N samples. This haplogroup is found widely distributed in ancient samples, due to its early split (formed ca. 9200 BC, TMRCA ca. 4500 BC) and expansion, probably with Neolithic populations. I can’t seem to find samples of this early haplogroup from the Carpathian Basin, as mentioned in the text, although it wouldn’t be strange, because it appears also in Neolithic Iberia, and in modern populations from western Europe.

Nevertheless, I2a-L621 samples seem to be concentrated mainly in Mesolithic-Neolithic cultures of Fennoscandia, and appeared also in Sikora et al. (2017) in a sample of the High Middle Ages from Sunghir (ca. AD 1100-1200), probably from the Vladimir-Suzdalian Rus’, in a region where clearly tribes of Volga Finns were being assimilated at the time. The reported SNP call by Genetiker is A16681 (see Yfull), deep within I2a-CTS10228. It is possibly also behind a modern Saami from Chalmny Varre (ca. AD 1800) of hg. I2a in Lamnidis et al. (2018).

Lacking precise subclades from Hungarian conquerors this is pure speculation, but modern samples may also point to I2a-CTS10228 (formed ca. 3100 BC, TMRCA ca. 1800 BC) as a Finno-Ugric lineage in common with R1a, which must have expanded to the Urals and beyond with eastern Corded Ware groups or (more likely) succeeding cultures. This is in line with the association of certain I2a lineages with modern Uralic peoples or populations from their historical regions in eastern Europe, and linked thus to the most likely homeland of Uralians in the eastern European forests:

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, and also of I lineages. Data from Tambets et al. (2018).

R1a-Z645

Regarding the important question of the ethnic makeup of Ugric populations stemming from the Urals, the most interesting (and expected) data is the presence of R1a-Z645 lineages among high-ranking conquerors, in particular four R1a-Z280 subclades proper of Finno-Ugrians.

This proves that, in line with the old split and expansion of R1a-CTS1211 (formed ca. 2600 BC, TMRCA ca. 2400 BC), and its finding in Bronze Age Fennoscandian samples, only some late R1a-Z280 (xZ92) lineages (see Z280 on YFull) may show a clear identification with early acculturated Uralic speakers, with the main early acculturated Balto-Slavic R1a haplogroup remaining R1a-M458.

I recently hypothesized this late connection of Slavs with very specific R1a-Z280 (xZ92) lineages based on analyses of modern populations (like Slovenians), because the connection of ancient Finno-Ugrians with modern Z92 samples was already evident:

(…) subclades of hg. R1a1a1b1a2-Z280 (xR1a1a1b1a2a-Z92) seem to have also been involved in early Slavic expansions, like R1a1a1b1a2b3a-CTS3402 (formed ca. 2200 BC, TMRCA ca. 2200 BC), found among modern West, South, and East Slavic populations and in Fennoscandia, prevalent e.g. among modern Slovenians which points to a northern origin of its expansion (Maisano Delser et al. 2018).

This finding also supports the expected shared R1a-Z280 lineages among ancient Finno-Ugric populations, as predicted from the study of modern Permic and Ugric peoples in Dudás et al. (2019).

r1a-z282-z280-z2125-distribution
Modified image, from Underhill et al. (2015). Spatial frequency distributions of Z282 (green) and Z93 (blue) affiliated haplogroups. Notice the distribution of R1a-Z280 (xZ92), i.e. R1a-M558, compared to the ancient Finno-Ugric distribution.

Furthermore, while we don’t have precise R1a-Z93 lineages to compare with the new Hunnic sample reported, we already know that some archaic R1a-Z2124 subclades stem from the forest-steppe areas of the Cis- and Trans-Urals, and the two newly reported R1a-Z93 Hungarian conqueror elites, like those of the Árpád dynasty, probably belong to them.

There is an obvious lack of continuity in specific paternal lineages among the Hunnic, the Avar, and the Conqueror periods, which makes any simplistic identification of all R1a-Z93 lineages as stemming from Avars, Huns, or the Iron Age Pontic-Caspian steppes clearly flawed. Comparing R1a-Z93 in Hungarian Conquerors with Huns is like comparing them with samples of the Srubna or earlier periods… Similarly, comparing the Hunnic R1b-U106 or the early Avar I1 to later Hungarian samples is not warranted without precise subclades, because they most likely correspond to different Germanic populations: Goths among Huns, then Longobards, then likely peoples descended from Franks and Irish Monks (the latter with R1b-P312).

N1a-L392

Second behind R1a subclades are, as expected, N1a-L392 (N1c in the old nomenclature).

Avars are dominated by a specific N1a-L392 subclade, N1a-B197, as we recently discovered in Csáky et al. (2019).

Hungarian conquerors show three N1a-Z1936 subclades, which is known to stem from the northern Ural region, including the Arctic (likely Palaeo-Laplandic peoples) and cross-stamped cultures of the northern Eurasian forests.

haplogroup_n3a4
Frequency-Distribution Maps of Individual Subclade N3a4 / N1a1a1a1a2-Z1936, probably with the Samic (first) and Fennic (later) expansions into Paleo-Lakelandic and Palaeo-Laplandic territories.

On the other hand, the two N1a-M2118 lineages are more clearly associated with Palaeo-Siberian populations east of the Urals, but became incorporated into the Ugric stock in the Trans-Urals region probably in the same way as N1a-Z1936, by infiltration from (and acculturation of) hunter-gatherers of forest and taiga cultures.

NOTE. You can read more about the infiltration of N1a lineages in the recent post Corded Ware—Uralic (IV): Hg R1a and N in Finno-Ugric and Samoyedic expansions, and in the specific sections for each Uralic group in A Clash of Chiefs.

haplogroup-n1a-M2118
Frequency-Distribution Maps of Individual Sub-clades of hg N3a2, by Ilumäe et al. (2016).

Conclusion

The picture offered by the paper on Hungarian Conquerors, while in line with historical accounts of multi-ethnic tribes incorporating regional lineages, shows nevertheless patrilineal clans clearly associated with Uralic peoples, in a distribution which could have been easily inferred from ancient Trans-Uralian forest-steppe cultures and modern samples (even regarding I2a-L621).

In spite of this, there is a great deal of discussion in the paper about specific N1a subclades in Hungarian conquerors, while the presence of R1a-Z280 (among early Magyar elites!) is interpreted, as always, as recently acculturated Slavs. This is sadly coupled with the simplistic identification of I2a-L621 as of local origin around the Carpathians.

The introduction of the paper to the history of Hungarians is also weird, for example giving credibility to the mythic accounts of the Árpád dynasty’s origin in Attila, which is in line, I guess, with what the authors intended to support all along, i.e. the association of Magyars with Turks from the Eurasian steppes, which they are apparently willing to achieve by relating them to haplogroup R1a-Z93

The conclusion is thus written to appease modern nation-building myths more than anything else, like many other papers before it:

It is generally accepted that the Hungarian language was brought to the Carpathian Basin by the Conquerors. Uralic speaking populations are characterized by a high frequency of Y-Hg N, which have often been interpreted as a genetic signal of shared ancestry. Indeed, recently a distinct shared ancestry component of likely Siberian origin was identified at the genomic level in these populations, modern Hungarians being a puzzling exception36. The Conqueror elite had a significant proportion of N Hgs, 7% of them carrying N1a1a1a1a4-M2118 and 10% N1a1a1a1a2-Z1936, both of which are present in Ugric speaking Khantys and Mansis. At the same time none of the examined Conquerors belonged to the L1034 subclade of Z1936, while all of the Khanty Z1936 lineages reported in 37 proved to be L1034 which has not been tested in the 23 study. Population genetic data rather position the Conqueror elite among Turkic groups, Bashkirs and Volga Tatars, in agreement with contemporary historical accounts which denominated the Conquerors as “Turks”. This does not exclude the possibility that the Hungarian language could also have been present in the obviously very heterogeneous, probably multiethnic Conqueror tribal alliance.

So, back to square one, and new circular reasoning: If ancient populations from north-eastern Europe believed to represent ancient Finno-Ugrians are of R1a-Z645 lineages, it’s because they were not Finno-Ugric speakers. If ancient and modern populations known to be of Finno-Ugric language show clear connections with R1a-Z645, it’s because they are “multi-ethnic”.

The only stable basis for discussion in genetic papers, apparently, is the own making of geneticists, with their traditional 2000s “R1a=Indo-European” and “N1c=Uralic”, coupled with national beliefs. It does not matter how many predictions based on that have been proven wrong, or how many predictions based on the Corded Ware = Uralic expansion have been proven right.

Related

Scytho-Siberians of Aldy-Bel and Sagly, of haplogroup R1a-Z93, Q1b-L54, and N

iron-age-sakas-aldy-bel-scythians

Recently, a paper described Eastern Scythian groups as “Uralic-Altaic” just because of the appearance of haplogroup N in two Pazyryk samples.

This simplistic identification is contested by the varied haplogroups found in early Altaic groups, by the early link of Cimmerians with the expansion of hg. N and Q, by the link of N1c-L392 in north-eastern Europe with Palaeo-Laplandic, and now (paradoxically) by the clear link between early Mongolic expansion and N1c-L392 subclades.

A new paper (behind paywall) offers insight into the prevalent presence of R1a-Z93 among eastern Scytho-Siberian groups (most likely including Samoyedic speakers in the forest-steppes), and a new hint to the westward expansion of haplogroups Q and N (probably coupled with the so-called “Siberian ancestry”) from the east with different groups of Iron Age steppe nomads:

Genetic kinship and admixture in Iron Age Scytho-Siberians, by Mary et al. Human Genetics (2019).

Interesting excerpts (emphasis mine):

From an archeological and historical point of view, the term “Scythians” refers to Iron Age nomadic or seminomadic populations characterized by the presence of three types of artifacts in male burials: typical weapons, specific horse harnesses and items decorated in the so-called “Animal Style”. This complex of goods has been termed the “Scythian triad” and was considered to be characteristic of nomadic groups belonging to the “Scythian World” (Yablonsky 2001). This “Scythian World” includes both the Classic (or European) Scythians from the North Pontic region (7th–3th century BC) and the Southern Siberian (or Asian) populations of the Scythian period (also called Scytho-Siberians). These include, among others, the Sakas from Kazakhstan, the Tagar population from the Minusinsk Basin (Republic of Khakassia), the Aldy-Bel population from Tuva (Russian Federation) and the Pazyryk and Sagly cultures from the Altai Mountains.

mtdna-scytho-siberians
Proportions of Scythian mtDNA haplogroups. Western (blue) and eastern (pink) Eurasian lineages are equally distributed in the Arzhan Scytho-Siberian sample. The U5a2a1 haplogroup shared between the two Scythian groups studied is in bold

In this work, we first aim to address the question of the familial and social organization of Scytho-Siberian groups by studying the genetic relationship of 29 individuals from the Aldy-Bel and Sagly cultures using autosomal STRs. (…) were obtained from 5 archeological sites located in the valley of the Eerbek river in Tuva Republic, Russia (Fig. 1). All the mounds of this archeological site were excavated but DNA samples were not collected from all of them. 14C dates mainly fall within the Hallstatt radiocarbon calibration plateau (ca. 800–400 cal BC) where the chronological resolution is poor. Only one date falls on an earlier segment of calibration curve: Le 9817–2650 ± 25 BP, i.e. 843–792 cal BC with a probability of 94.3% (using the OxCal v4.3.2 program). This sample (Bai-Dag 8, Kurgan 1, grave 10) is not from one of the graves studied but was used to date the kurgan as a whole.

Y-chromosome haplogroups were first assigned using the ISOGG 2018 nomenclature. In order to improve the precision of haplogroup definition, we also analyzed a set of Y-chromosome SNP (Supplementary Table 2). Nine samples belonged to the R1a-M513 haplogroup (defined by marker M513) and two of these nine samples were characterized as belonging to the R1a1a1b2-Z93 haplogroup or one of its subclades. Six samples belonged to the Q1b1a-L54 haplogroup and five of these six samples belonged to the Q1b1a3-L330 subclade. One sample belonged to the N-M231 haplogroup.

haplogroups-scythian-siberians

The distribution of these haplogroups in the population must be confronted with the prevalence of kinship among the samples. Although five individuals belonged to haplogroup Q1b1a3-L330, three of them (ARZ-T18, ARZ-T19 and ARZ-T20) were paternally related (Fig. 2). It must, therefore, be considered that haplogroup Q1b1a3-L330 is present in three independent instances (given that the remaining two instances exhibit no close familial relationship with other samples or one another). All five were buried on the Eki-Ottug 1 archaeological site (although in two different kurgans).

In the same way, although two groups, of two and three individuals, shared haplotypes belonging to the R1a-M513 haplogroup, these groups likely include a father/son pair (ARZ-T2 and ARZ-T12). Therefore, among nine R1a-M513 men, we found six independent haplotypes, one being present in two independent instances. All R1a-M513 haplotypes, however, including those attributed to the R1a1a1b2-Z93 subclade, only differed by one-step mutations, across 5 loci at most. All R1a-M513 individuals were buried on the same site, Eki-Ottug 2, in a single Kurgan.

y-haplogroups-r1a-n-q1b

Haplogroup R1a-M173 was previously reported for 6 Scytho-Siberian individuals from the Tagar culture (Keyser et al. 2009) and one Altaian Scytho-Siberian from the Sebÿstei site (Ricaut et al. 2004a), whereas haplogroup R1a1a1b2-Z93 (or R1a1a1b-S224) was described for one Scythian from Samara (Mathieson et al. 2015) and two Scytho-Siberians from Berel and the Tuva Republic (Unterländer et al. 2017). On the contrary, North Pontic Scythians were found to belong to the R1b1a1a2 haplogroup (Krzewińska et al. 2018), showing a distinction between the two groups of Scythians. (…) The absence of R1b lineages in the Scytho-Siberian individuals tested so far and their presence in the North Pontic Scythians suggest that these 2 groups had a completely different paternal lineage makeup with nearly no gene flow from male carriers between them.

The seven other male individuals studied in this work were found to carry Eastern Eurasian Y haplogroups Q1b1a and one of its subclades (n = 6) and N (n = 1). Haplogroup Q1b1a-L54 was previously described in four males from the Bronze Age in the Altai Mountains (Hollard et al. 2014, 2018) and was clearly associated with Siberian populations (Regueiro et al. 2013).

The N-M231 haplogroup emerged from haplogroup K in Southern Asia around 21,000 years BCE, maybe in Southern China (Shi et al. 2013; Ilumäe et al. 2016). Previous studies attested to its presence in samples from Neolithic and Bronze Age in China (Li et al. 2011; Cui et al. 2013). Waves of northwestern expansion of this haplogroup are described as beginning during the Paleolithic period (Derenko et al. 2006; Shi et al. 2013) but traces of this expansion in archeological samples were reported only in two Scytho-Siberian males from the Altai (Pilipenko et al. 2015).

The sample of haplogroup N comes from the Aldy-Bel culture (ARZ-T15), from the Eerbek site, but has no radiocarbon date. All Q1b-L330 samples come from the Sagly culture, and three are paternally related. The other Q1b-L54 sample is from other tombs in one kurgan at Aldy Bel.

It seems that – exactly as expected – different waves of steppe nomads brought different lineages at a time (the Iron Age) when many regions incorporated different eastern lineages without necessarily changing language. Just like the expansion of N among Ugrians and Samoyeds, and N1c among Finno-Permic peoples, and like many other lineages expanding with federation-like groups in eastern, central, and western Europe

Related

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).

y-str-haplogroup-n-mongolian-ugrians
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.

haplogroup_n1
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.

bashkir-mari-southern-mansi
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.

y-dna-hungarians-ugric-mansi
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.

N1c-Tat:

(…) 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).

n1c-n-tat-uralic-ugric
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.

r1a-z280-ugrians
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.

r1a-z93-ugrians
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)

Comments

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.

r1a-z282-z280-z2125-distribution
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.

Conclusion

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.

finno-ugric-haplogroup-n
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. Fading purple arrows represent likely stepped movements of subclades of haplogroup N for centuries (e.g. Siberian → Ananino → Akozino → Fennoscandia [N-VL29]; Circum-Arctic → forest-steppe [N1, N2]; etc.). Blue arrows represent eventual expansions of Uralic peoples to the north. Modified image from Vasilyev (2002).

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.

Related

The Pazyryk culture spoke a “Uralic-Altaic” language… because haplogroup N

Matrilineal and patrilineal genetic continuity of two iron age individuals from a Pazyryk culture burial, by Tikhonov, Gurkan, Peler, & Dyakonov, Int J Hum Genet (2019).

Relevant excerpts (emphasis mine):

Of particular interest to the current study are the archaeogenetic investigations associated with the exemplary mound 1 from the Ak-Alakha-1 site on the Ukok Plateau in the Altai Republic (Polosmak 1994a; Pilipenko et al. 2015). This typical Pazyryk “frozen grave” was dated around 2268±39 years before present (Bln-4977) (Gersdorff and Parzinger 2000). Initial anthropological findings suggested an undisturbed dual inhumation comprising “a middle-aged European- type man” and “a young European-type woman”, both of whom presumably had a high social status among the Pazyryk elite (Polosmak 1994a). In contrast, recent archaeogenetic investigations revealed somewhat contradicting results since analyses at both the amelogenin gene and Y-chromosome short tandem repeat (Y-STR) loci clearly established that both Scythians were actually males and had paternal and maternal lineages that are typically associated with eastern Eurasians (Pilipenko et al. 2015). Through the use of mitochondrial, autosomal and Y-chromosomal DNA typing systems, it was possible to not only investigate the potential relationships between the two ancient Scythians but also to gather initial phylogenetic and phylogeographic information on their paternal and maternal lineages (Pilipenko et al. 2015).

Based on the Y-STR data available, the two Ak-Alakha-1 Scythians had an in silico haplogroup assignment of N, which first appeared in southeastern Asia and then expanded in southern Siberia (Rootsi et al. 2007; Pilipenko et al. 2015).

Current study aims to investigate the geographical distributions of the ancient and contemporary matches and close genetic variants of the maternal and paternal lineages observed in the two Scythians from the exemplary Ak-Alakha-1 kurgan.

tikhonov-scythian-modern
Geographic distribution of the exact matches with the Scythian (PZ1) Y-STR (17-loci) and mtDNA (HVR1) haplotypes detailed in Tables 1a and 1b. Boundaries of the Altai Republic within the Russian Federation are shown with dashed lines, along with an approximate position of the Ak-Alakha-1 burial site, which is denoted with an ‘x’ on the map. Countries shaded in gray refer to those that have full 17-loci Y-STR and/or mtDNA HVR1 match(es) with the PZ1 haplotypes. Inset in the top and bottom left corners are the Altai and Uzbekistan maps, respectively, both scaled-up to allow better representation of the samples derived from these countries. There were no other exact matches from around parts of the globe that are not shown on the map, except for a single contemporary mtDNA haplotype from US, which presumably belonged to an ‘East Asian’ individual. Inset in the top right corner provides a scale for the number of haplotypes observed, but only up to three samples, which is valid for the entire map as well as the inset maps, irrespective of the differences in the scales of the actual map and inset maps themselves. For sample pools larger than three, the same linear scale provided on the inset in the top right corner still applies; please refer to Tables 1a and b for actual sample pool sizes. Samples are depicted on the entire map and the insets maps with circles and diamonds for the Y-STR and mtDNA haplotypes, respectively. Black and white coloring for samples depict whether the haplotype(s) are contemporary or ancient, respectively. Location of the PZ1 mtDNA and Y-STR haplotypes are shown on top of each other.

In response to aggressive Xiongnu expansion into the Altai region around the 2nd century BCE, some members of the Pazyryk culture may have started moving up North, and eventually reached the Vilyuy River at the beginning of 1st century CE. Notably, there is clear population continuity between the Uralic people such as Khants, Mansis and Nganasans, Paleo-Siberian people such as Yukaghirs and Chuvantsi, and the Pazyryk people even when considering just the two mtDNA and Y-STR haplotypes from the Ak-Alakha-1 mound 1 kurgan (Tables 1a, b, Table 2, Fig. 1). These concepts are also in agreement with the famous Yakut ethnographer Ksenofontov, who suggested that technologies associated with ferrous metallurgy were brought to the Vilyuy Valley at around 1st century CE by the first (proto)Turkic-speaking pioneers (Ksenofontov 1992). Yakut ethnogenesis per se possibly involved two major stages, the first being the proto-Turkic epoch through the arrival of Scytho-Siberian culture originating from Southern Siberia, such as that associated with the Pazyryk culture and the second being the proper Turkic epoch.

Nomadic peoples from the Central Asian steppes are East Iranian speakers whenever they are of haplogroup R1a, but “Uralic-Altaic” speakers whenever they are of haplogroup N. True story.

So they followed a haplogroup ca. 37,000 years old, in a sample dated some 2,300 years ago, whose precise subclade and ancient history is (yet) unknown, compared it to present-day populations, and the result is that they spoke “Uralic-Altaic” because haplogroup N and continuity. Sound familiar? Yep, it’s the kind of reasoning you might be reading right now about Iberian Bell Beakers, about Bell Beakers, or even about Yamna and their relationship to a Vasconic-Caucasian language, based on haplogroup R1b in modern Basques. Another true story.

Anyway, based on the multi-ethnic federations created during this time, and on the ancestral components visible in the different groups (see a post on Karasuk by Chad Rohlfsen), the Pazyryk culture’s language is unknown, and it could be, as a matter of fact (apart from the obvious East Iranian connection):

We also know that haplogroup N and Siberian ancestry expanded into cultures of Northern Eurasia precisely with the creation of the new social paradigm of chiefdoms and alliances, roughly at the same time as Scythians expanded, with the first sample of haplogroup N in Hungary appearing with Cimmerians.

finno-ugric-haplogroup-n
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. Fading purple arrows represent likely stepped movements of subclades of haplogroup N for centuries (e.g. Siberian → Ananino → Akozino → Fennoscandia [N-VL29]; Circum-Arctic → forest-steppe [N1, N2]; etc.). Blue arrows represent eventual expansions of Uralic peoples to the north. Modified image from Vasilyev (2002).

While the study of modern populations is interesting, the problem I have with the paper is the reasoning of “language of ancient haplogroups based on modern populations”, and especially with the concept of “Uralic-Altaic”, and the highly hypothetic “Proto-Turkic” nomadic steppe pastoralists before “Hunnic Turkic” (which is itself questionable), before the “real Turkic” layer (being the authors apparently Turkic themselves), and the supposed “continuity” of Eastern Uralic and Turkic groups in Asia since the Out of Africa migration. The combination of all of this in the same text is just disturbing.

If you look at it from the bright side, at least these samples were not of haplogroup R1a-Z280, or we would be talking about great Slavonic Scythians showing continuity from Russia with love, as the paper threatened to do in its introduction…

If you are enjoying the comeback of this retro 2000s comedy in 2019 (based on the classic nativist “R1a=IE”, “R1b=Basque”, and “N=Uralic” combo) it’s because you – like me – are putting yourself in this guy’s shoes every time a new episode of funny self-destruction appears:

chosen-poorly

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