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).

west-urals-finno-ugrians-qpadm
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

Corded Ware—Uralic (IV): Hg R1a and N in Finno-Ugric and Samoyedic expansions

haplogroup-uralians

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

Let me begin this final post on the Corded Ware—Uralic connection with an assertion that should be obvious to everyone involved in ethnolinguistic identification of prehistoric populations but, for one reason or another, is usually forgotten. In the words of David Reich, in Who We Are and How We Got Here (2018):

Human history is full of dead ends, and we should not expect the people who lived in any one place in the past to be the direct ancestors of those who live there today.

Haplogroup N

Another recurrent argument – apart from “Siberian ancestry” – for the location of the Uralic homeland is “haplogroup N”. This is as serious as saying “haplogroup R1” to refer to Indo-European migrations, but let’s explore this possibility anyway:

Ancient haplogroups

We have now a better idea of how many ancient migrations (previously hypothesized to be associated with westward Uralic migrations) look like in genetic terms. From Damgaard et al. (Science 2018):

These serial changes in the Baikal populations are reflected in Y-chromosome lineages (Fig. SA; figs. S24 to S27, and tables S13 and SI4). MAI carries the R haplogroup, whereas the majority of Baikal_EN males belong to N lineages, which were widely distributed across Northern Eurasia (29), and the Baikal_LNBA males all carry Q haplogroups, as do most of the Okunevo_EMBA as well as some present-day Central Asians and Siberians.

The only N1c1 sample comes from Ust’Ida Late Neolithic, 180km to the north of Lake Baikal, which – together with the Bronze Age sample from the Kola peninsula, and the medieval sample from Ust’Ida – gives a good idea of the overall expansion of N subclades and Siberian ancestry among the Circum-Arctic peoples of Eurasia, speakers of Palaeo-Siberian languages.

eurasian-n-subclades
Geographical location of ancient samples belonging to major clade N of the Y-chromosome.

Modern haplogroups

What we should expect from Uralic peoples expanding with haplogroup N – seeing how Yamna expands with R1b-L23, and Corded Ware expands with R1a-Z645 – is to find a common subclade spreading with Uralic populations. Let’s see if it works like that for any N-X subclade, in data from Ilumäe et al. (2016):

haplogroup_n1
Geographic-Distribution Map of hg N3 / N1c / N1a.

Within the Eurasian circum-Arctic spread zone, N3 and N2a reveal a well-structured spread pattern where individual sub-clades show very different distributions:

N1a1-M46 (or N-TAT), formed ca. 13900 BC, TMRCA 9800 BC

   N1a1a2-B187, formed ca. 9800 BC, TMRCA 1050 AD:

The sub-clade N3b-B187 is specific to southern Siberia and Mongolia, whereas N3a-L708 is spread widely in other regions of northern Eurasia.

     N1a1a1a-L708, formed ca. 6800 BC, TMRCA 5400 BC.

       N1a1a1a2-B211/Y9022, formed ca. 5400 BC, TMRCA 1900 BC:

The deepest clade within N3a is N3a1-B211, mostly present in the Volga-Uralic region and western Siberian Khanty and Mansi populations.

         N1a1a1a1a-L392/L1026), formed ca. 4400 BC, TMRCA 2800 BC:

The neighbor clade, N3a3’6-CTS6967, spreads from eastern Siberia to the eastern part of Fennoscandia and the Baltic States

haplogroup_n3a3
Frequency-Distribution Maps of Individual Subclade N3a3 / N1a1a1a1a1a-CTS2929/VL29, probably initially with Akozino warrior-traders.

           N1a1a1a1a1a-CTS2929/VL29, formed ca. 2100 BC, TMRCA 1600 BC:

In Europe, the clade N3a3-VL29 encompasses over a third of the present-day male Estonians, Latvians, and Lithuanians but is also present among Saami, Karelians, and Finns (Table S2 and Figure 3). Among the Slavic-speaking Belarusians, Ukrainians, and Russians, about three-fourths of their hg N3 Y chromosomes belong to hg N3a3.

In the post on Finno-Permic expansions, I depicted what seems to me the most likely way of infiltration of N1c-L392 lineages with Akozino warrior-traders into the western Finno-Ugric populations, with an origin around the Barents sea.

This includes the potential spread of (a minority of) N1c-B211 subclades due to contacts with Anonino on both sides of the Urals, through a northern route of forest and forest-steppe regions (equivalent to the distribution of Cherkaskul compared to Andronovo), given the spread of certain subclades in Ugric populations.

NOTE. An alternative possibility is the association of certain B211 subclades with a southern route of expansion with Pre-Scythian and Scythian populations, under whose influence the Ananino culture emerged -which would imply a very quick infiltration of certain groups of haplogroup N everywhere among Finno-Ugrics on both sides of the Urals – , and also the expansion of some subclades with Turkic-speaking peoples, who apparently expanded with alliances of different peoples. Both (Scythian and Turkic) populations expanded from East Asia, where haplogroup N (including N1c) was present since the Neolithic. I find this a worse model of expansion for upper clades, but – given the YFull estimates and the presence of this haplogroup among Turkic peoples – it is a possibility for many subclades.

           N1a1a1a1a2-Z1936, formed ca. 2800 BC, TMRCA 2400 BC:

The only notable exception from the pattern are Russians from northern regions of European Russia, where, in turn, about two-thirds of the hg N3 Y chromosomes belong to the hg N3a4-Z1936—the second west Eurasian clade. Thus, according to the frequency distribution of this clade, these Northern Russians fit better among other non-Slavic populations from northeastern Europe. N3a4 tends to increase in frequency toward the northeastern European regions but is also somewhat unexpectedly a dominant hg N3 lineage among most Turcic-speaking Volga Tatars and South-Ural Bashkirs.

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.

The expansion of N1a-Z1936 in Fennoscandia is most likely associated with the expansion of Saami into asbestos ware-related territory (like the Lovozero culture) during the Late Iron Age – and mixture with its population – , and with the later Fennic expansion to the east and north, replacing their language, as well as with Arctic and forest populations assimilated during Permic, Ugric, and Samoyedic expansions to the north.

           N1a1a1a1a4-M2019 (previously N3a2), formed ca. 4400 BC, TMRCA 1700 BC:

Sub-hg N3a2-M2118 is one of the two main bifurcating branches in the nested cladistic structure of N3a2’6-M2110. It is predominantly found in populations inhabiting present-day Yakutia (Republic of Sakha) in central Siberia and at lower frequencies in the Khanty and Mansi populations, which exhibit a distinct Y-STR pattern (Table S7) potentially intrinsic to an additional clade inside the sub-hg N3a2

The second widespread sub-clade of hg N is N2a. (…):

   N1a2b-P43 (B523/FGC10846/Y3184), formed ca. 6800 BC, TMRCA ca. 2700 BC:

The absolute majority of N2a individuals belong to the second sub-clade, N2a1-B523, which diversified about 4.7 kya (95% CI = 4.0–5.5 kya). Its distribution covers the western and southern parts of Siberia, the Taimyr Peninsula, and the Volga-Uralic region with frequencies ranging from from 10% to 30% and does not extend to eastern Siberia (…)

haplogroup_n2
Geographic-Distribution Map of hg N2a1 / N1a2b-P43

The “European” branch suggested earlier from Y-STR patterns turned out to consist of two clades

     N1a2b2a-Y3185/FGC10847, formed ca. 2200 BC, TMRCA 800 BC:

N2a1-L1419, spread mainly in the northern part of that region.

     N1a2b2b1-B528/Y24382, formed ca. 900 BC, TMRCA ca. 900 BC:

N2a1-B528, spread in the southern Volga-Uralic region.

Haplogroup R1a

We also have a good idea of the distribution of haplogroup R1a-Z645 in ancient samples. Its subclades were associated with the Corded Ware expansion, and some of them fit quite well the early expansion of Finno-Permic, Ugric, and Samoyedic peoples to the east.

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.

This is how the modern distribution of R1a among Uralians looks like, from the latest report in Tambets et al. (2018):

  • Among Fennic populations, Estonians and Karelians (ca. 1.1 million) have not suffered the greatest bottleneck of Finns (ca. 6-7 million), and show thus a greater proportion of R1a-Z280 than N1c subclades, which points to the original situation of Fennic peoples before their expansion. To trust Finnish Y-DNA to derive conclusions about the Uralic populations is as useful as relying on the Basque Y-DNA for the language spread by R1b-P312
  • Among Volga-Finnic populations, Mordovians (the closest to the original Uralic cluster, see above) show a majority of R1a lineages (27%).
  • Hungarians (ca. 13-15 million) represent the majority of Ugric (and Finno-Ugric) peoples. They are mainly R1a-Z280, also R1a-Z2123, have little N1c, and lack Siberian ancestry, and represent thus the most likely original situation of Ugric peoples in 4th century AD (read more on Avars and Hungarians).
  • Among Samoyedic peoples, the Selkup, the southernmost ones and latest to expand – that is, those not heavily admixed with Siberian populations – , also have a majority of R1a-Z2123 lineages (see also here for the original Samoyedic haplogroups to the south).

To understand the relevance of Hungarians for Ugric peoples, as well as Estonians, Karelians, and Mordovians (and northern Russians, Finno-Ugric peoples recently Russified) for Finno-Permic peoples, as opposed to the Circum-Arctic and East Siberian populations, one has to put demographics in perspective. Even a modern map can show the relevance of certain territories in the past:

population-density
Population density (people per km2) map of the world in 1994. From Wikipedia.

Summary of ancestry + haplogroups

Fennic and Samic populations seem to be clearly influenced by Palaeo-Laplandic peoples, whereas Volga-Finnic and especially Permic populations may have received gene flow from both, but essentially Palaeo-Siberian influence from the north and east.

The fact that modern Mansis and Khantys offer the highest variation in N1a subclades, and some of the highest “Siberian ancestry” among non-Nganasans, should have raised a red flag long ago. The fact that Hungarians – supposedly stemming from a source population similar to Mansis – do not offer the same amount of N subclades or Siberian ancestry (not even close), and offer instead more R1a, in common with Estonians (among Finno-Samic peoples) and Mordvins (among Volga-Finnic peoples) should have raised a still bigger red flag. The fact that Nganasans – the model for Siberian ancestry – show completely different N1a2b-P43 lineages should have been a huge genetic red line (on top of the anthropological one) to regard them as the Uralian-type population.

We know now that ethnolinguistic groups have usually expanded with massive (usually male-biased) migrations, and that neighbouring locals often ‘resurge’ later without changing the language. That is seen in Europe after the spread of Bell Beakers, with the increase of previous ancestry and lineages in Scandinavia during the formation of the Nordic ethnolinguistic community; in Central-West Europe, with the resurgence of Neolithic ancestry (and lineages) during the Bronze Age over steppe ancestry; and in Central-East Europe (with Unetice or East European Bronze Age groups like Mierzanowice, Trzciniec, or Lusatian) showing an increase in steppe ancestry (and resurge of R1a subclades); none of them represented a radical ethnolinguistic change.

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).

It is not hard to model the stepped arrival, infiltration, and/or resurge of N subclades and “Siberian ancestries”, as well as their gradual expansion in certain regions, associated with certain migrations first – such as the expansions to the Circum-Arctic region, and later the Scythian- and Turkic-related movements – , as well as limited regional developments, like the known bottleneck in Finns, or the clear late expansion of Ugric and Samoyedic languages to the north among nomadic Palaeo-Siberians due to traditions of exogamy and multilingualism. This fits quite well with the different arrival of N (N1c and xN1c) lineages to the different Uralic-speaking groups, and to the stepped appearance of “Siberian ancestry” in the different regions.

The aternative

It is evident that a lot of people were too attached to the idea of Palaeolithic R1b lineages ‘native’ to western Europe speaking Basque languages; of R1a lineages speaking Indo-European and spreading with Yamna; and N lineages ‘native’ to north-eastern Europe and speaking Uralic, and this is causing widespread weeping and gnashing of teeth (instead of the joy of discovering where one’s true patrilineal ancestors come from, and what language they spoke in each given period, which is the supposed objective of genetic genealogy…)

Since an Indo-Germanic branch (as revived now by some in the Copenhaguen group to fit Kristiansen’s theory of the 1980s with recent genetic data) does not make any sense in linguistics, the finding of R1a in Yamna would not have led where some think it would have, because North-West Indo-European would still be the main Late PIE branch in Europe. Don’t take my word for it; take James P. Mallory’s (2013).

mallory-adams-tree
The levels of Indo-European reconstruction, from Mallory & Adams (2006).

If an (unlikely) Indo-Slavonic group were posited, though, such a group would still be bound (with Indo-Iranian) to the steppes with East Yamna/Poltavka (admixing with Abashevo migrants, but retaining its language), developing Sintashta/Potapovka → Srubna/Andronovo, and R1a lineages would have equally undergone the known bottlenecks of the steppes where they replaced R1b-Z2103 – which this eastern group shares with Balkan languages, a haplogroup that links therefore together the Graeco-Aryan group.

As far as I know – and there might be many other similar pet theories out there – there have been proposals of “modern Balto-Slavic-like” populations (in an obvious circular reasoning based on modern populations) in some Scythian clusters of the Iron Age.

NOTE. I will not enter into “Balto-Slavic-like R1a” of the Late Bronze Age or earlier because no one can seriously believe at this point of development of Population Genetics that autosomal similarity predating 1,500+ years the appearance of Slavs equates to their (ethnolinguistic) ancestral population, without a clear intermediate cultural and genetic trail – something we lack today in the Slavic case even for the late Roman period…

finno-saamic-palaeo-germanic-substratum
The Finnic and Saamic separation looks shallower than it actually is. Invisible convergence can be ‘triangulated’ with the help of Germanic layers of mutual loanwords (Häkkinen 2012).

We also know of R1a-Z280 lineages in Srubna, probably expanding to the west. With that in mind, and knowing that Palaeo-Germanic was in close contact with Finno-Samic while both were already separated but still in contact, and that Palaeo-Germanic was also in contact and closely related to a ‘Temematic’ distinct from Balto-Slavic (and also that early Proto-Baltic and Proto-Slavic from the Roman Iron Age and later were in contact with western Uralic) this will be the linguistic map of the Iron Age if R1a is considered to expand Indo-European from some kind of “patron-client” relationship with west Yamna:

palaeo-germanic-italo-celtic
Eastern European language map during the Late Bronze Age / Iron Age, if R1a spread Indo-European languages and Eastern Yamna spoke Indo-Slavonic. Palaeo-Germanic (i.e. Pre- to Proto-Germanic) needs to be in contact with both the Samic Lovozero population and the Fennic west Circum-Arctic one. Italic and Celtic in contact with Pre-Germanic. Germanic in contact with Temematic. Balto-Slavic in contact with Iranian, and near Fennic to allow for later loanwords. For Germanic and Temematic, see Kortlandt (2018).

You might think I have some personal or political reason against this kind of proposals. I haven’t. We have been proposing Indo-European to be the language of the European Union for more than 10 years, so to support R1b-Italo-Celtic in the whole Western Europe, R1a-Germanic in Central and Eastern Europe, and R1a-Indo-Slavonic in the steppes (as the Danish group seems to be doing) has nothing inherently bad (or good) for me. If anything, it gives more reason to support the revival of North-West Indo-European in Europe.

My problem with this proposal is that it is obviously beholden to the notion of the uninterrupted cultural, historic and ethnic continuity in certain territories. This bias is common in historiography (von Falkenhausen 1993), but it extends even more easily into the lesser known prehistory of any territory, and now more than ever some people feel the need to corrupt (pre)history based on their own haplogroups (or the majority haplogroups of their modern countries). However, more than on philosophical grounds, my rejection is based on facts: this picture is not what the combination of linguistic, archaeological, and genetic data shows. Period.

Nevertheless, if Yamna + Corded Ware represented the “big and early expansion” of Germanic and Italo-Celtic peoples proper of the dream Nazi’s Lebensraum and Fascist’s spazio vitale proposals; Uralians were Siberian hunter-gatherers that controlled the whole eastern and northern Russia, and miraculously managed to push (ethnolinguistically) Neolithic agropastoralists to the west during and after the Iron Age, with gradual (and often minimal) genetic impact; and Balto-Slavic peoples were represented by horse riders from Pokrovka/Srubna, hiding then somewhere around the forest-steppe until after the Scythian expansion, and then spreading their language (without much genetic impact) during the early Middle Ages…so be it.

See also

Related

Corded Ware—Uralic (III): “Siberian ancestry” and Ugric-Samoyedic expansions

siberian-ancestry-tambets

This is the third of four posts on the Corded Ware—Uralic identification. See

An Eastern Uralic group?

Even though proposals of an Eastern Uralic (or Ugro-Samoyedic) group are in the minority – and those who support it tend to search for an origin of Uralic in Central Asia – , there is nothing wrong in supporting this from the point of view of a western homeland, because the eastward migration of both Proto-Ugric and Pre-Samoyedic peoples may have been coupled with each other at an early stage. It’s like Indo-Slavonic: it just doesn’t fit the linguistic data as well as the alternative, i.e. the expansion of Samoyedic first, different from a Finno-Ugric trunk. But, in case you are wondering about this possibility, here is Häkkinen’s (2012) phonological argument:

ugro-samoyedic-uralic

The case of Samoyedic is quite similar to that of Hungarian, although the earliest Palaeo-Siberian contact languages have been lost. There were contacts at least with Tocharian (Kallio 2004), Yukaghir (Rédei 1999) and Turkic (Janhunen 1998). Samoyedic also:

a) has moved far from the related languages and has been exposed to strong foreign influence

b) shares a small number of common words with other branches (from Sammallahti 1988: only 123 ‘Uralic’ words, versus 390 ‘Uralic’ + ‘Finno-Ugric’ words found in other branches than Samoyedic = 31,5 %)

c) derives phonologically from the East Uralic dialect.

The phonological level is taxonomically more reliable, since it lacks the distortion caused by invisible convergence and false divergence at the lexical level. Thus we can conclude that the traditional taxonomic model, according to which Samoyedic was the first branch to split off from the Proto-Uralic unity, is just as incorrect as the view that Hungarian was the first branch to split off.

Seima-Turbino

Late Uralic can be traced back to metallurgical cultures thanks to terms like PU *wäśka ‘copper/bronze’ (borrowed from Proto-Samoyedic *wesä into Tocharian); PU *äsa and *olna/*olni, ‘lead’ or ‘tin’, found in *äsa-wäśka ‘tin-bronze’; and e.g. *weŋći ‘knife’, borrowed into Indo-Iranian (through the stage of vocalization of nasals), appearing later as Proto-Indo-Aryan *wāćī ‘knife, awl, axe’.

It is known that the southern regions of the Abashevo culture developed Proto-Indo-Iranian-speaking Sintashta-Petrovka and Pokrovka (Early Srubna). To the north, however, Abashevo kept its Uralic nature, with continuous contacts allowing for the spread of lexicon – mainly into Finno-Ugric – , and phonetic influence – mainly Uralisms into Proto-Indo-Iranian phonology (read more here).

The northern part of Abashevo (just like the south) was mainly a metallurgical society, with Abashevo metal prospectors found also side by side with Sintashta pioneers in the Zeravshan Valley, near BMAC, in search of metal ores. About the Seima-Turbino phenomenon, from Parpola (2013):

From the Urals to the east, the chain of cultures associated with this network consisted principally of the following: the Abashevo culture (extending from the Upper Don to the Mid- and South Trans-Urals, including the important cemeteries of Sejma and Turbino), the Sintashta culture (in the southeast Urals), the Petrovka culture (in the Tobol-Ishim steppe), the Taskovo-Loginovo cultures (on the Mid- and Lower Tobol and the Mid-Irtysh), the Samus’ culture (on the Upper Ob, with the important cemetery of Rostovka), the Krotovo culture (from the forest steppe of the Mid-Irtysh to the Baraba steppe on the Upper Ob, with the important cemetery of Sopka 2), the Elunino culture (on the Upper Ob just west of the Altai mountains) and the Okunevo culture (on the Mid-Yenissei, in the Minusinsk plain, Khakassia and northern Tuva). The Okunevo culture belongs wholly to the Early Bronze Age (c. 2250–1900 BCE), but most of the other cultures apparently to its latter part, being currently dated to the pre-Andronovo horizon of c. 2100–1800 BCE (cf. Parzinger 2006: 244–312 and 336; Koryakova & Epimakhov 2007: 104–105).

post-eneolithic-steppe-asia
Schematic map of the Middle Bronze Age cultures (steppe and foreststeppe
zone)

The majority of the Sejma-Turbino objects are of the better quality tin-bronze, and while tin is absent in the Urals, the Altai and Sayan mountains are an important source of both copper and tin. Tin is also available in southern Central Asia. Chernykh & Kuz’minykh have accordingly suggested an eastern origin for the Sejma-Turbino network, backing this hypothesis also by the depiction on the Sejma-Turbino knives of mountain sheep and horses characteristic of that area. However, Christian Carpelan has emphasized that the local Afanas’evo and Okunevo metallurgy of the Sayan-Altai area was initially rather primitive, and could not possibly have achieved the advanced and difficult technology of casting socketed spearheads as one piece around a blank. Carpelan points out that the first spearheads of this type appear in the Middle Bronze Age Caucasia c. 2000 BCE, diffusing early on to the Mid-Volga-Kama-southern Urals area, where “it was the experienced Abashevo craftsmen who were able to take up the new techniques and develop and distribute new types of spearheads” (Carpelan & Parpola 2001: 106, cf. 99–106, 110). The animal argument is countered by reference to a dagger from Sejma on the Oka river depicting an elk’s head, with earlier north European prototypes (Carpelan & Parpola 2001: 106–109). Also the metal analysis speaks for the Abashevo origin of the Sejma-Turbino network. Out of 353 artefacts analyzed, 47% were of tin-bronze, 36% of arsenical bronze, and 8.5% of pure copper. Both the arsenical bronze and pure copper are very clearly associated with the Abashevo metallurgy.

seima-turbino-phenomenon-parpola
Find spots of artefacts distributed by the Sejma-Turbino intercultural trader network, and the areas of the most important participating cultures: Abashevo, Sintashta, Petrovka. Based on Chernykh 2007: 77.

The Abashevo metal production was based on the Volga-Kama-Belaya area sandstone ores of pure copper and on the more easterly Urals deposits of arsenical copper (Figure 9). The Abashevo people, expanding from the Don and Mid-Volga to the Urals, first reached the westerly sandstone deposits of pure copper in the Volga and Kama basins, and started developing their metallurgy in this area, before moving on to the eastern side of the Urals to produce harder weapons and tools of arsenical copper. Eventually they moved even further south, to the area richest in copper in the whole Urals region, founding there the very strong and innovative Sintashta culture.

Regarding the most likely expansion of Eastern Uralic peoples:

Nataliya L’vovna Chlenova (1929–2009; cf. Korenyako & Ku’zminykh 2011) published in 1981 a detailed study of the Cherkaskul’ pottery. In her carefully prepared maps of 1981 and 1984 (Figure 10), she plotted Cherkaskul’ monuments not only in Bashkiria and the Trans-Urals, but also in thick concentrations on the Upper Irtysh, Upper Ob and Upper Yenissei, close to the Altai and Sayan mountains, precisely where the best experts suppose the homeland of Proto-Samoyed to be.

cherkaskul-andronovo
Distribution of Srubnaya (Timber Grave, early and late), Andronovo (Alakul’ and Fëdorovo variants) and Cherkaskul’ monuments. After Parpola 1994: 146, fig. 8.15, based on the work of N. L. Chlenova (1984: map facing page 100).

Ugric

The Cherkaskul’ culture was transformed into the genetically related Mezhovka culture (c. 1500–1000 BCE), which occupied approximately the same area from the Mid-Kama and Belaya rivers to the Tobol river in western Siberia (cf. Parzinger 2006: 444–448; Koryakova & Epimakhov 2007: 170–175). The Mezhovka culture was in close contact with the neighbouring and probably Proto-Iranian speaking Alekseevka alias Sargary culture (c. 1500–900 BCE) of northern Kazakhstan (Figure 4 no. 8) that had a Fëdorovo and Cherkaskul’ substratum and a roller pottery superstratum (cf. Parzinger 2006: 443–448; Koryakova & Epimakhov 2007: 161–170). Both the Cherkaskul’ and the Mezhovka cultures are thought to have been Proto-Ugric linguistically, on the basis of the agreement of their area with that of Mansi and Khanty speakers, who moreover in their Fëdorovo-like ornamentation have preserved evidence of continuity in material culture (cf. Chlenova 1984; Koryakova & Epimakhov 2007: 159, 175).

mezhovska-sargary-irmen
Cultures of the Final Bronze Age of the Urals and western Siberia (steppe
and forest-steppe zone).

The Mezhovka culture was succeeded by the genetically related Gamayun culture (c. 1000–700 BCE) (cf. Parzinger 2006: 446; 542–545).

From the Gamayun culture descend Trans-Urals cultures in close contact with Finno-Permic populations of the Cis-Ural region:

  • [Proto-Mansi] Itkul’ culture (c. 700–200 BCE) distributed along the eastern slope of the Ural Mountains (cf. Parzinger 2006: 552–556). Known from its walled forts, it constituted the principal Trans-Uralian centre of metallurgy in the Iron Age, and was in contact with both the Anan’ino and Akhmylovo cultures (the metallurgical centres of the Mid-Volga and Kama-Belaya region) and the neighbouring Gorokhovo culture.
    • [Proto-Hungarian] via the Vorob’evo Group (c. 700–550 BCE) (cf. Parzinger 2006: 546–549), to the Gorokhovo culture (c. 550–400 BCE) of the Trans-Uralian forest steppe (cf. Parzinger 2006: 549–552). For various reasons the local Gorokhovo people started mobile pastoral herding and became part of the multicomponent pastoralist Sargat culture (c. 500 BCE to 300 CE), which in a broader sense comprized all cultural groups between the Tobol and Irtysh rivers, succeeding here the Sargary culture. The Sargat intercommunity was dominated by steppe nomads belonging to the Iranian-speaking Saka confederation, who in the summer migrated northwards to the forest steppe
  • [Proto-Khanty] Late Bronze Age and Early Iron Age cultures related to the Gamayunskoe and Itkul’ cultures that extended up to the Ob: the Nosilovo, Baitovo, Late Irmen’, and Krasnoozero cultures (c. 900–500 BCE). Some were in contact with the Akhmylovo on the Mid-Volga.
sargat-gorokhovo-bolscherechye
Cultural groups of the Iron Age in the forest-steppe zone of western
Siberia. (

Samoyedic

Parpola (2012) connects the expansion of Samoyedic with the Cherkaskul variant of Andronovo. As we know, Andronovo was genetically diverse, which speaks in favour of different groups developing similar material cultures in Central Asia.

Juha Janhunen, author of the etymological dictionary of the Samoyed languages (1977), places the homeland of Proto-Samoyedic in the Minusinsk basin on the Upper Yenissei (cf. Janhunen 2009: 72). Mainly on the basis of Bulghar Turkic loanwords, Janhunen (2007: 224; 2009: 63) dates Proto-Samoyedic to the last centuries BCE. Janhunen thinks that the language of the Tagar culture (c. 800–100 BCE) ought to have been Proto-Samoyedic (cf. Janhunen 1983: 117– 118; 2009: 72; Parzinger 2001: 80 and 2006: 619–631 dates the Tagar culture c. 1000–200 BCE; Svyatko et al. 2009: 256, based on human bone samples, c. 900 BCE to 50 CE). The Tagar culture largely continues the traditions of the Karasuk culture (c. 1400–900 BCE), (…)

chicha-irmen-tagar-baraba-forest-siberian
Map showing the location of Chicha-1.

For the most recent expansions of Samoyedic languages to the north, into Palaeo-Siberian populations, read more about the traditional multilingualism of Siberian populations.

Genetics

Siberian ancestry

The use of a map of “Siberian ancestry” peaking in the arctic to show a supposedly late Uralic population movement (starting in the Iron Age!) seems to be the latest trend in population genomics:

siberian-ancestry-map
Frequency map of the so-called ‘Siberian’ component. From Tambets et al. (2018) (see below for ADMIXTURE in specific populations).

I guess that would make this map of Neolithic farmer ancestry represent an expansion of Indo-European from the south, because Anatolia, Greece, Italy, southern France, and Iberia – where this ancestry peaks in modern populations – are among the oldest territories where Indo-European languages were recorded:

reich-farmer-ancestry
Modern genome-wide data shows that the primary gradient of farmer ancestry in Europe does not flow southeast-to-northwest but instead in an almost perpendicular direction, a result of a major migration of pastoralists from the east that displaced much of the ancestry of the first farmers.

Probably not the right interpretation of this kind of simplistic data about modern populations, though…

The most striking thing about the “Siberian ancestry” white whale is that nobody really knows what it is; just like we did not know what “Yamnaya ancestry” was, until the most recent data is making the picture clearer. Its nature is changing with each new paper, and it can be summed up by “some ancestry we want to find that is common to Uralic-speaking peoples, and should not be CWC-related”. Tambets et al. (2018) explain quite well how they “found it”:

Overall, and specifically at lower values of K, the genetic makeup of Uralic speakers resembles that of their geographic neighbours. The Saami and (a subset of) the Mansi serve as exceptions to that pattern being more similar to geographically more distant populations (Fig. 3a, Additional file 3: S3). However, starting from K = 9, ADMIXTURE identifies a genetic component (k9, magenta in Fig. 3a, Additional file 3: S3), which is predominantly, although not exclusively, found in Uralic speakers. This component is also well visible on K = 10, which has the best cross-validation index among all tests (Additional file 3: S3B). The spatial distribution of this component (Fig. 3b) shows a frequency peak among Ob-Ugric and Samoyed speakers as well as among neighbouring Kets (Fig. 3a). The proportion of k9 decreases rapidly from West Siberia towards east, south and west, constituting on average 40% of the genetic ancestry of FU speakers in Volga-Ural region (VUR) and 20% in their Turkic-speaking neighbours (Bashkirs, Tatars, Chuvashes; Fig. 3a).

siberian-ancestry-modern
Population structure of Uralic-speaking populations inferred from ADMIXTURE analysis on autosomal SNPs in Eurasian context. Individual ancestry estimates for populations of interest for selected number of assumed ancestral populations (K3, K6, K9, K11). Ancestry components discussed in a main text (k2, k3, k5, k6, k9, k11) are indicated and have the same colours throughout. The names of the Uralic-speaking populations are indicated with blue (Finno-Ugric) or orange (Samoyedic). Image from Tambets et al. (2018).

However, this ‘something’ that some people occasionally find in some Uralic populations is also common to other modern and ancient groups, and not so common in some other Uralic peoples. Simply put:

siberian-ancestry-modern-populations
Image modified from Lamnidis et al. (2018). Red line representing maximum “Siberian admixture” in Eastern European hunter-gatherers. In blue, Uralic-speaking groups. “Plot of ADMIXTURE (K=3) results containing West Eurasian populations and the Nganasan. Ancient individuals from this study are represented by thicker bars.”

I already said this in the recent publication of Siberian samples, where a renamed and radiocarbon dated Finnish_IA clearly shows that Late Iron Age Saami (ca. 400 AD) had little “Siberian ancestry”, if any at all, representing the most likely Fennic (and Samic) ancestral components before their expansion into central and northern Finland, where they admixed with circum-polar peoples of asbestos ware cultures.

I will say that again and again, any time they report the so-called “Siberian ancestry” in Uralic samples, no matter how it is defined each time: it does not seem to be that special something people are looking for, but rather (at least in a great part) a quite old ancestral component forming an evident cline with EHG, whose best proximate source are Baikal_EN (and/or Devil’s Gate) at this moment, and thus also East European hunter-gatherers for Western Uralic peoples:

dzudzuana-baikal-en-admixture
Image modified from Lazaridis et al. (2018). In red: samples with Baikal_EN ancestry in speculative estimates. In pink: samples with Baikal_EN ancestry in conservative estimates (probably marking a recent arrival of Baikal_En ancestry, see here). Modeling present-day and ancient West-Eurasians. Mixture proportions computed with qpAdm (Supplementary Information section 4). The proportion of ‘Mbuti’ ancestry represents the total of ‘Deep’ ancestry from lineages that split prior to the split of Ust’Ishim, Tianyuan, and West Eurasians and can include both ‘Basal Eurasian’ and other (e.g., Sub-Saharan African) ancestry. (Left) ‘Conservative’ estimates. Each population 367 cannot be modeled with fewer admixture events than shown. (Right) ‘Speculative’ estimates. The highest number of sources (≤5) with admixture estimates within [0,1] are shown for each population. Some of the admixture proportions are not significantly different from 0 (Supplementary Information section 4).

So either Samara_HG, Karelia_HG, and many other groups from eastern Europe all spoke Uralic according to this ADMIXTURE graphic (and the formation of steppe ancestry in the Volga-Ural region brought the Proto-Indo-European language to the steppes through the CHG/ANE expansion), or a great part of this “Siberian ancestry” found in modern Uralic-speaking populations is not what some people would like to think it is…

Modern populations

PCA clines can be looked for to represent expansions of ancient populations. Most recently, Flegontov et al. (2018) are attempting to do this with Asian populations:

For some Turkic groups in the Urals and the Altai regions and in the Volga basin, a different admixture model fits the data: the same West Eurasian source + Uralic- or Yeniseian-speaking Siberians. Thus, we have revealed an admixture cline between Scythians and the Iranian farmer genetic cluster, and two further clines connecting the former cline to distinct ancestry sources in Siberia. Interestingly, few Wusun-period individuals harbor substantial Uralic/Yeniseian-related Siberian ancestry, in contrast to preceding Scythians and later Turkic groups characterized by the Tungusic/Mongolic-related ancestry. It remains to be elucidated whether this genetic influx reflects contacts with the Xiongnu confederacy. We are currently assembling a collection of samples across the Eurasian steppe for a detailed genetic investigation of the Hunnic confederacies.

jeong-population-clines
Three distinct East/West Eurasian clines across the continent with some interesting linguistic correlates, as earlier reported by Jeong et al. (2018). Alexander M. Kim.

There are potential errors with this approach:

The main one is practical – does a modern cline represent an ancestral language? The answer is: sometimes. It depends on the anthropological context that we have, and especially on the precision of the PCA:

clines-himalayan
Genetic structure of the Himalayan region populations from analyses using unlinked SNPs. (A) PCA of the Himalayan and HGDP-CEPH populations. Each dot represents a sample, coded by region as indicated. The Himalayan region samples lie between the HGDP-CEPH East Asian and South Asian samples on the right-hand side of the plot. From Arciero et al. (2018).

The ‘Europe’, ‘Middle East’, etc. clines of the above PCA do not represent one language, but many. For starters, the PCA includes too many (and modern) populations, its precision is useless for ethnolinguistic groups. Which is the right level? Again, it depends.

The other error is one of detail of the clines drawn (which, in turn, depends on the precision of the PCA). For example, we can draw two paralell lines (or even one line, as in Flegontov et al. above) in one PCA graphic, but we still don’t have the direction of expansion. How do we know if this supposed “Uralic-speaking cline” goes from one region to the other? For that level of detail, we should examine closely modern Uralic-speaking peoples and Circum-Arctic populations:

uralic-cline
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

The real ancient Uralic cluster (drawn above in blue) is thus probably from a North-East European source (probably formed by Battle Axe / Fatyanovo-Balanovo / Abashevo) to the east into Siberian populations, and to the north into Laplandic populations (see below also on Mezhovska ancestry for the drawn ‘European cline’, which some may a priori wrongly assume to be quite late).

The fact that the three formed clines point to an admixture of CWC-related populations from North-Eastern Europe, and that variation is greater at the Palaeo-Laplandic and Palaeo-Siberian extremities compared to the CWC-related one, also supports this as the correct interpretation.

However, judging by the two main clines formed, one could be alternatively inclined to interpret that Palaeo-Laplandic and Palaeo-Siberian populations formed a huge ancestral “Uralic” ghost cluster in Siberia (spanning from the Palaeo-Laplandic to the Palaeo-Siberian one), and from there expanded Finno-Samic on one hand, and “Volga-Ugro-Samoyed” on the other. That poses different problems: an obvious linguistic and archaeological one – which I assume a lot of people do not really care about – , and a not-so-obvious genetic one (see below for ancient samples and for the expansion of haplogroup N).

To understand the simplest solution better, one can just have a look at the PCA from Bell Beaker samples in Olalde et al. (2018), which (as Reich has already explained many times) expanded directly from Yamna R1b-L23 lineages:

olalde_pca_clines
Image modified from Olalde et al. (2018). PCA of 999 Eurasian individuals. Marked is the Espersted Outlier with the approximate position of Yamna Hungary, probably the source of its admixture. Different Bell Beaker clines have been drawn, to represent approximate source of expansions from Central European sources into the different regions.

Unlike this PCA with ancient samples, where Bell Beaker clines could be a rough approximation to the real sources for each population, and where a cluster spanning all three depicted Early Bronze Age clusters could give a rough proximate source of European Bell Beakers in Hungary (and where one can even distinguish the Y-DNA bottlenecks in the L23 trunk created by each cline) the PCA of modern Uralic populations is probably not suitable for a good estimate of the ancient situation, which may be found shifted up or down of the drawn “Uralic” cluster along East European groups.

After all, we already know that the Siberian cline shows probably as much an ancient admixture event – from the original Uralic expansion to the east with Corded Ware ancestry – as another more recent one – a westward migration of Siberian ancestry (or even more than one). While we know with more or less exactitude what happened with the Palaeo-Laplandic admixture by expanding Proto-Finno-Samic populations (see here), the Proto-Ugric and Pre-Samoyedic populations formed probably more than one cline during the different ancient migrations through central Asia.

Ancient populations

Apparently, the Corded Ware expansion to the east was not marked by a huge change in ancestry. While the final version of Narasimhan et al. (2018) may show a little more detail about other forest-steppe Seima-Turbino/Andronovo-related migrations (and thus also Eastern Uralic peoples), we have already had enough information for quite some time to get a good idea.

mezhovska-pca
Principal component analysis. PCA of ancient individuals (according colours see legend) projected on modern West Eurasians (grey). Iron Age Scythians are shown in black; CHG, Caucasus hunter-gatherer; LNBA, late Neolithic/Bronze Age; MN, middle Neolithic; EHG, eastern European huntergatherer; LBK_EN, early Neolithic Linearbandkeramik; HG, hunter-gatherer; EBA, early Bronze Age; IA, Iron Age; LBA, late Bronze Age; WHG, western hunter-gatherer.dataset (grey). Iron Age Scythians are shown in black; CHG, Caucasus hunter-gatherer; LNBA, late Neolithic/Bronze Age; MN, middle Neolithic; EHG, eastern European hunter-gatherer; LBK_EN, early Neolithic Linearbandkeramik; HG, hunter-gatherer; EBA, early Bronze Age; IA, Iron Age; LBA, late Bronze Age; WHG, western hunter-gatherer.

Mezhovska‘s position is similar to the later Pre-Scythian and Scythian populations. There are some interesting details: apart from haplogroup R1a-Z280 (CTS1211+), there is one R1b-M269 (PF6494+), probably Z2103, and an outlier (out of three) in a similar position to the recently described central/southern Scythian clusters.

NOTE. The finding of R1b-M269 in the forest-steppe is probably either 1) from an Afanasevo-Okunevo origin, or 2) from an admixture with neighbouring Andronovo-related populations, such as Sargary. A third, maybe less likely option is that this haplogroup admixed with Abashevo directly (as it happened in Sintashta, Potapovka, or Pokrovka) and formed part of early Uralic migrations. In any case, since Mezhovska is a Bronze Age society from the Urals region, its association with R1b-Z2103 – like the association of R1b-Z2103 in Scythian clusters – cannot be attributed to “Thracian peoples”, a link which is (as I already said) too simplistic.

The drawn “European cline” of Hungarians (see above), leading from ‘west-like’ Mansi to Hungarian populations – and hosting also Finnic and Estonian samples – , cannot therefore be attributed simply to late “Slavic/Balkan-like” admixture.

Karasuk – located further to the east – is basically also Corded Ware peoples showing clearly a recent admixture with local ANE / Baikal_EN-like populations. In terms of haplogroups it shows haplogroup Q, R1a-Z2124, and R1a-Z2123, later found among early Hungarians, and present also in ancient Samoyedic populations now acculturated.

The most interesting aspect of both Mezhovska and Karasuk is that they seem to diverge from a point close to Ukraine_Eneolithic, which is the supposed ancestral source of Corded Ware peoples (read more about the formation of “steppe ancestry”). This means that Eastern Uralians derive from a source closer to Middle Dnieper/Abashevo populations, rather than Battle Axe (shifted to Latvian Neolithic), which is more likely the source prevalent in Finno-Permic peoples.

Their initial admixture with (Palaeo-)Siberian populations is thus seen already starting by this time in Mezhovska and especially in Karasuk, but this process (compared to modern populations) is incomplete:

f4-test-karasuk-mezhovska
Visualization of f-statistics results. f4(Test, LBK; Han, Mbuti) values are plotted on x axis and f4(Test, LBK; EHG, Mbuti) values on y axis, positive deviations from zero show deviations from a clade between Test and LBK. A red dashed line is drawn between Yamnaya from Samara and Ami. Iron Age populations that can be modelled as mixtures of Yamnaya and East Eurasians (like the Ami) are arrayed around this line and appear to be distinct from the main North/South European cline (blue) on the left of the x axis.
karasuk-mezhovska-admixture
ADMIXTURE results for ancient populations. Red arrows point to the Iron Age Scythian individuals studied. LBK_EN: Early Neolithic Linearbandkeramik; EHG: Eastern European hunter-gatherer; Motala_HG: hunter-gatherer from Motala (Sweden); WHG: western hunter-gatherer; CHG: Caucasus hunter-gatherer; IA: Iron Age; EBA: Early Bronze Age; LBA: Late Bronze Age.

We know now that Samic peoples expanded during the Late Iron Age into Palaeo-Laplandic populations, admixing with them and creating this modern cline. Finns expanded later to the north (in one of their known genetic bottlenecks), admixing with (and displacing) the Saami in Finland, especially replacing their male lines.

So how did Ugric and Samoyedic peoples admix with Palaeo-Siberian populations further, to obtain their modern cline? The answer is, logically, with East Asian migrations related to forest-steppe populations of Central Asia after the Mezhovska and Karasuk periods, i.e. during the Iron Age and later. Other groups from the forest-steppe in Central Asia show similar East Asian (“Siberian”) admixture. We know this from Narasimhan et al. (2018):

(…) we observe samples from multiple sites dated to 1700-1500 BCE (Maitan, Kairan, Oy_Dzhaylau and Zevakinsikiy) that derive up to ~25% of their ancestry from a source related to present-day East Asians and the remainder from Steppe_MLBA. A similar ancestry profile became widespread in the region by the Late Bronze Age, as documented by our time transect from Zevakinsikiy and samples from many sites dating to 1500-1000 BCE, and was ubiquitous by the Scytho-Sarmatian period in the Iron Age.

We already have some information about these later migrations:

siberian-genetic-component-chronology
Very important observation with implication of population turnover is that pre-Turkic Inner Eurasian populations’ Siberian ancestry appears predominantly “Uralic-Yeniseian” in contrast to later dominance of “Tungusic-Mongolic” sort (which does sporadically occur earlier). Alexander M. Kim

The Ugric-speaking Sargat culture in Western Siberia shows the expected mixture of haplogroups (ca. 500 BC – 500 AD), with 5 samples of hg N and 2 of hg R1a1, in Pilipenko et al. (2017). Although radiocarbon dates and subclades are lacking, N lineages probably spread late, because of the late and gradual admixture of Siberian cultures into the Sargat melting pot.

The Samoyedic-speaking Tagar culture also shows signs of a genetic turnover in Pilipenko et al. (2018):

The observed reduction in the genetic distance between the Middle Tagar population and other Scythian like populations of Southern Siberia(Fig 5; S4 Table), in our opinion, is primarily associated with an increase in the role of East Eurasian mtDNA lineages in the gene pool (up to nearly half of the gene pool) and a substantial increase in the joint frequency of haplogroups C and D (from 8.7% in the Early Tagar series to 37.5% in the Middle Tagar series). These features are characteristic of many ancient and modern populations of Southern Siberia and adjacent regions of Central Asia, including the Pazyryk population of the Altai Mountains.

Before the Iron Age, the Karasuk and Mezhovska population were probably already somehow ‘to the north’ within the ancient Steppe-Altai cline (see image below9 created by expanding Seima-Turbino- and Andronovo-related populations. During the Iron Age, further Siberian contributions with Iranian expansions must have placed Uralians of the Central Asian forest-steppe areas much closer to today’s Palaeo-Siberian cline.

However, the modern genetic picture was probably fully developed only in historic times, when Samoyedic and Ugric languages expanded to the north, only in part admixing further with Palaeo-Siberian-speaking nomads from the Circum-Arctic region (see here for a recent history of Samoyedic Enets), which justifies their more recent radical ‘northern shift’.

east-uralic-clines
Modified image from Jeong et al. (2018), supplementary materials. 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 north-south 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.

This late acquisition of the language by Palaeo-Siberian nomads (without much population replacement) also justifies the wide PCA clusters of very small Siberian populations. See for example in the PCA from Tambets et al. (2018):

uralic-ugric-samoyedic-modern-clines
Approximate Ugric and Samoyedic clines (exluding apparent outliers). 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

For their relationship with modern Mansi, we have information on Hungarian conqueror populations from Neparáczki et al. (2018):

Moreover, Y, B and N1a1a1a1a Hg-s have not been detected in Finno-Ugric populations [80–84], implying that the east Eurasian component of the Conquerors and Finno-Ugric people are probably not directly related. The same inference can be drawn from phylogenetic data, as only two Mansi samples appeared in our phylogenetic trees on the side branches (S1 Fig, Networks; 1, 4) suggesting that ancestors of the Mansis separated from Asian ancestors of the Conquerors a long time ago. This inference is also supported by genomic Admixture analysis of Siberian and Northeastern European populations [85], which revealed that Mansis received their eastern Siberian genetic component approximately 5–7 thousand years ago from ancestors of modern Even and Evenki people. Most likely the same explanation applies to the Y-chromosome N-Tat marker which originated from China [86,87] and its subclades are now widespread between various language groups of North Asia and Eastern Europe [88].

The genetic picture of Hungarians (their formed cline with Mansi and their haplogroups) may be quite useful for the true admixture found originally in Mansi peoples at the beginning of the Iron Age. By now it is clear even from modern populations that Steppe_MLBA ancestry accompanied the Uralic expansion to the east (roughly approximated in the graphic with Afanasievo_EBA + Bichon_LP EasternHG_M):

siberian-population-expansions
Admixture modelling using qpAdm. Maps showing locations and ancestry proportions of ancient (left) and modern (right) groups. From Sikora et al. (2018).

Continue reading the final post of the series: Corded Ware—Uralic (IV): Haplogroups R1a and N in Finno-Ugric and Samoyedic.

See also

Related

  • The traditional multilingualism of Siberian populations
  • Iron Age bottleneck of the Proto-Fennic population in Estonia
  • Y-DNA haplogroups of Tuvinian tribes show little effect of the Mongol expansion
  • Corded Ware—Uralic (I): Differences and similarities with Yamna
  • Haplogroup R1a and CWC ancestry predominate in Fennic, Ugric, and Samoyedic groups
  • The Iron Age expansion of Southern Siberian groups and ancestry with Scythians
  • Evolution of Steppe, Neolithic, and Siberian ancestry in Eurasia (ISBA 8, 19th Sep)
  • Mitogenomes from Avar nomadic elite show Inner Asian origin
  • On the origin and spread of haplogroup R1a-Z645 from eastern Europe
  • Oldest N1c1a1a-L392 samples and Siberian ancestry in Bronze Age Fennoscandia
  • Consequences of Damgaard et al. 2018 (III): Proto-Finno-Ugric & Proto-Indo-Iranian in the North Caspian region
  • The concept of “Outlier” in Human Ancestry (III): Late Neolithic samples from the Baltic region and origins of the Corded Ware culture
  • Genetic prehistory of the Baltic Sea region and Y-DNA: Corded Ware and R1a-Z645, Bronze Age and N1c
  • More evidence on the recent arrival of haplogroup N and gradual replacement of R1a lineages in North-Eastern Europe
  • Another hint at the role of Corded Ware peoples in spreading Uralic languages into north-eastern Europe, found in mtDNA analysis of the Finnish population
  • New Ukraine Eneolithic sample from late Sredni Stog, near homeland of the Corded Ware culture