The complex origin of Samoyedic-speaking populations

uralic-turkic

Open access Siberian genetic diversity reveals complex origins of the Samoyedic-speaking populations, by Karafet et al. Am J Hum Biol (2018) e23194.

Interesting excerpts (emphasis mine):

Siberian groups

Consistent with their origin, Mongolic-speaking Buryats demonstrate genetic similarity with Mongols, and Turkic-speaking Altai-Kizhi and Teleuts are drawn close to CAS groups. The Tungusic-speaking Evenks collected in central and eastern Siberia cluster together and overlap with Yukagirs. Dolgans are widely scattered in the plot, justifying their recent origin from one Evenk clan, Yakuts, and Russian peasants in the 18th century (Popov, 1964). Uralic-speaking populations comprise a very wide cluster with Komi drawn to Europe, and Khants showing a closer affinity with Selkups, Tundra and Forest Nentsi. Yenisey-speaking Kets are intermingled with Selkups. Interestingly, Samoyedic-speaking Nganasans from the Taymyr Peninsula form a separate tight cluster closer to Evenks, Yukagirs, and Koryaks.

pca-siberian-uralic
Principal component analysis (PCA) using the “drop one in” technique for 27 present-day (N = 424) and 6 ancient populations (N = 20). PCA was performed on 281 093 SNPs from the intersection of our data with publicly available ancient Siberian samples

ADMIXTURE and the “Siberian component”

Among Siberians, the Komi are primarily Europeans, while Nganasans, Evenks, Yukagirs, and Koryaks are nearly 100% East Asians. At K = 4 finer scale subcontinental structure can be distinguished with the emergence of a “Siberian” component. This component is highly pronounced in the Nganasans. Outside Siberia, this component is present in Germany and in CAS at low frequency. Within ancient cultures, this component has the highest frequency in three BA Karasuk samples. It is also found in Mal’ta, ENE Afanasievo and BA Andronovo, but not in Ust’-Ishim and BA Okunevo. At K = 5, the “Siberian” component is roughly subdivided into two components with different geographic distributions. The “Nganasan” component is frequent in nearly all Siberian populations, except the Komi, Kets and Selkups. The newly derived “Selkup-Ket” component is found at high frequencies in western Siberian populations. It is observed in BA Karasuk and in Mal’ta. At K = 6, the western Siberian “Nentsi-Khant” ancestry component was developed in Forest and Tundra Nentsi, Khants. This component is also present at low levels in EUR, CAS, Tibet, and southern Siberia.

Identity-by-descent

The Dolgans share more segments with the Nganasans than within themselves (54.13 vs 41.72, Mann-Whitney test, P = .000000000001562546). The result is not surprising as the demographic data showed that the Nganasans were subjected to intense assimilation by the Dolgans in the second half of the 20th century (Goltsova, Osipova, Zhadanov, & Villems, 2005). Tundra Nentsi share more IBD with Forest Nentsi than within themselves (83.96 vs 50.3, P = .000055) possibly due to the common origin and long-term gene flow. The Ket and Selkup populations allocate significantly more IBD blocks between populations than with individuals from their own population (121.2 cM vs 85.9 cM for Kets, P = .000008, and 121.2 cM vs 114.9 cM for Selkups, P = .043).

admixture-siberian
ADMIXTURE plot. Clustering of 444 individuals from 27 present-day and 6 ancient populations (281 093 SNPs) assuming K6 to K7 clusters. Individuals are shown as vertical bars colored in ratio to their estimated ancestry within each cluster

Haplogroup N in Siberia

Although Siberia exhibits 42 haplogroups, the vast majority of Siberian Y-chromosomes belong only to 4 of the 18 major clades (N = 46.2%; C = 20.9%; Q = 14.4%; and R = 15.2%). The Y-chromosome haplogroup N is widely spread across Siberia and Eastern Europe (Ilumae et al., 2016; Karafet et al., 2002; Wong et al., 2016) and reaches its maximum frequency among Siberian populations such as Nganasans (94.1%) and Yakuts (91.9%). Within Siberia, two sister subclades N-P43 and N-L708 show different geographic distributions. N-P43 and derived haplogroups N-P63 and N- P362 (phylogenetically identical to N-B478* and N-B170, respectively) (Ilumae et al., 2016) are extremely rare in other major geographic regions. Likely originating in western Siberia, they are limited almost entirely to northwest Siberia, the Volga- Uralic regions, and the Taymyr Peninsula (ie, do not extend to eastern Siberia). Conversely, clade N-L708 is frequent in all Siberian populations except the Kets and Selkups, reaching its highest frequency in the Yakuts (91.9%).

Surprisingly, not a single sign of the proposed reindeer pastoralist horde led by Nganasans into north-eastern Europe. This is strange because “Siberian” migrants hypothetically imposed their language over Indo-Europeans quite recently, apparently after the Iron Age

Interesting comparisons among Siberian groups, though.

Related

Minimal gene flow from western pastoralists in the Bronze Age eastern steppes

jeong-steppes-mongolia

Open access paper Bronze Age population dynamics and the rise of dairy pastoralism on the eastern Eurasian steppe, by Jeong et al. PNAS (2018).

Interesting excerpts (emphasis mine):

To understand the population history and context of dairy pastoralism in the eastern Eurasian steppe, we applied genomic and proteomic analyses to individuals buried in Late Bronze Age (LBA) burial mounds associated with the Deer Stone-Khirigsuur Complex (DSKC) in northern Mongolia. To date, DSKC sites contain the clearest and most direct evidence for animal pastoralism in the Eastern steppe before ca. 1200 BCE.

Most LBA Khövsgöls are projected on top of modern Tuvinians or Altaians, who reside in neighboring regions. In comparison with other ancient individuals, they are also close to but slightly displaced from temporally earlier Neolithic and Early Bronze Age (EBA) populations from the Shamanka II cemetry (Shamanka_EN and Shamanka_EBA, respectively) from the Lake Baikal region. However, when Native Americans are added to PC calculation, we observe that LBA Khövsgöls are displaced from modern neighbors toward Native Americans along PC2, occupying a space not overlapping with any contemporary population. Such an upward shift on PC2 is also observed in the ancient Baikal populations from the Neolithic to EBA and in the Bronze Age individuals from the Altai associated with Okunevo and Karasuk cultures.

pca-eurasians-karasuk-khovsgol
Image modified from the article. Karasuk cluster in green, closely related to sample ARS026 in red. Principal Component Analysis (PCA) of selected 2,077 contemporary Eurasians belonging to 149 groups. Contemporary individuals are plotted using three-letter abbreviations for operational group IDs. Group IDs color coded by geographic region. Ancient Khövsgöl individuals and other selected ancient groups are represented on the plot by filled shapes. Ancient individuals are projected onto the PC space using the “lsqproject: YES” option in the smartpca program to minimize the impact of high genotype missing rate.

(…) two individuals fall on the PC space markedly separated from the others: ARS017 is placed close to ancient and modern northeast Asians, such as early Neolithic individuals from the Devil’s Gate archaeological site (22) and present-day Nivhs from the Russian far east, while ARS026 falls midway between the main cluster and western Eurasians.

Upper Paleolithic Siberians from nearby Afontova Gora and Mal’ta archaeological sites (AG3 and MA-1, respectively) (25, 26) have the highest extra affinity with the main cluster compared with other groups, including the eastern outlier ARS017, the early Neolithic Shamanka_EN, and present-day Nganasans and Tuvinians (Z > 6.7 SE for AG3). Main cluster Khövsgöl individuals mostly belong to Siberian mitochondrial (A, B, C, D, and G) and Y (all Q1a but one N1c1a) haplogroups.

mongolia-botai-ehg-ane-cline
The genetic affinity of the Khövsgöl clusters measured by outgroup-f3 and -f4 statistics. (A) The top 20 populations sharing the highest amount of >genetic drift with the Khövsgöl main cluster measured by f3(Mbuti; Khövsgöl, X). (B) The top 15 populations with the most extra affinity with each of the three Khövsgöl clusters in contrast to Tuvinian (for the main cluster) or to the main cluster (for the two outliers), measured by f4(Mbuti, X; Tuvinian/Khövsgöl, Khövsgöl/ARS017/ARS026). Ancient and contemporary groups are marked by squares and circles, respectively. Darker shades represent a larger f4 statistic.

Previous studies show a close genetic relationship between WSH populations and ANE ancestry, as Yamnaya and Afanasievo are modeled as a roughly equal mixture of early Holocene Iranian/ Caucasus ancestry (IRC) and Mesolithic Eastern European hunter-gatherers, the latter of which derive a large fraction of their ancestry from ANE. It is therefore important to pinpoint the source of ANE-related ancestry in the Khövsgöl gene pool: that is, whether it derives from a pre-Bronze Age ANE population (such as the one represented by AG3) or from a Bronze Age WSH population that has both ANE and IRC ancestry.

The amount of WSH contribution remains small (e.g., 6.4 ± 1.0% from Sintashta). Assuming that the early Neolithic populations of the Khövsgöl region resembled those of the nearby Baikal region, we conclude that the Khövsgöl main cluster obtained ∼11% of their ancestry from an ANE source during the Neolithic period and a much smaller contribution of WSH ancestry (4–7%) beginning in the early Bronze Age.

khovsgol-shamanka-sintashta
Admixture modeling of Altai populations and the Khövsgöl main cluster using qpAdm. For the archaeological populations, (A) Shamanka_EBA and (B and C) Khövsgöl, each colored block represents the proportion of ancestry derived from a corresponding ancestry source in the legend. Error bars show 1 SE. (A) Shamanka_EBA is modeled as a mixture of Shamanka_EN and AG3. The Khövsgöl main cluster is modeled as (B) a two-way admixture of Shamanka_EBA+Sintashta and (C) a three-way admixture Shamanka_EN+AG3+Sintashta.

Apparently, then, the first individual with substantial WSH ancestry in the Khövsgöl population (ARS026, of haplogroup R1a-Z2123), directly dated to 1130–900 BC, is consistent with the first appearance of admixed forest-steppe-related populations like Karasuk (ca. 1200-800 BC) in the Altai. Interestingly, haplogroup N1a1a-M178 pops up (with mtDNA U5a2d1) among the earlier Khövsgöl samples.

I will repeat what I wrote recently here: Samoyedic arrived in the Altai with Karasuk and hg R1a-Z645 + Steppe_MLBA-like ancestry, admixed with Altai populations, clustering thus within an Ancient Altai cline. Only later did N1a1a subclades infiltrate Samoyedic (and Ugric) populations, bringing them closer to their modern Palaeo-Siberian cline. The shared mtDNA may support an ancestral EHG-“Siberian” cline, or else a more recent Afanasevo-related origin.

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. Read more.

Also interesting, Q1a2 subclades and ANE ancestry making its appearance everywhere among ancestral Eurasian peoples, as Chetan recently pointed out.

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.

           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.

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.

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
  • The traditional multilingualism of Siberian populations

    uralic-languages

    New paper (behind paywall) A case-study in historical sociolinguistics beyond Europe: Reconstructing patterns of multilingualism in a linguistic community in Siberia, by Khanina and Meyerhoff, Journal of Historical Sociolinguistics (2018) 4(2).

    The Nganasans have been eastern neighbours of the Enets for at least several centuries, or even longer, as indicated in Figures 2 and 3.10 They often dwelled on the same grounds and had common households with the Enets. Nganasans and Enets could intermarry (Dolgikh 1962a), while the Nganasans did not marry representatives of any other ethnic groups. As a result, it was not unusual for Enets and Nganasans to live in the same tent and/or to have common relatives. Such close contact must clearly have favoured acquisition of Nganasan by Enets children and of Enets by Nganasan children from an early age.

    The Nenets have been close neighbours of all the Enets groups more recently (Figures 2 and 3). In the seventeenth century, there were only warlike contacts between the Nenets and the Enets, while in the eighteenth century the Nenets started to live on the traditional Enets lands, on the western bank of the Yenisey river, with more peaceful interactions reported. (…) Since then the same situation of intermarriages and common households has been attested for these western Enets neighbours as with the Nganasans (Dolgikh 1962a), and this has also created conditions favouring early acquisition of both languages by children.

    enets-nganasan-early
    The Enets and neighbouring peoples in the middle of the seventeenth century; map by Yuri Koryakov (http://lingvarium.org), adapted from Dolgikh (1960).

    As for the Evenkis and the Selkups, the Enets had regular contact with these peoples (Figures 2 and 3), though they were not their close neighbours: in fact, geographically, the Selkups were not neighbours at all by the end of the nineteenth century. The Evenkis had always been direct south-eastern neighbours (…) Contacts with Selkups could be trade based, or they could simply be occasional encounters on adjacent lands. (…) [With Evenkis] some sporadic contacts were similar in nature to those with the Selkups, however many other contacts were war-like. Traditionally, the Enets considered the Evenkis to have a martial spirit, and the Evenkis were known as being accustomed to stealing Enets women. A number of stories in Dolgikh (1961) concern Evenkis stealing Enets women and Enets men going to Evenki lands to find and return them. It is clear, therefore, that if Evenki or Selkup were acquired by the Enets, this happened later in life, and this acquisition required particular conditions for it, i. e. it was not readily acquired through regular or harmonious contact (as with Nganasan).

    In a pattern similar to the situation with Nganasan, in the second half of the twentieth century most Enets elders could speak Nenets (Vasil’jev 1963; Eugen Helimski p.c., the lead author’s fieldwork experience).

    enets-nganasan-late
    The Enets and neighbouring indigenous peoples: end of the nineteenth century – beginning of the twentieth century; map by Yuri Koryakov (http://lingvarium.org), adapted from
    Bruk (1961).

    At the start of the period studied, in the 1850s, the Enets linguistic community could be characterized as multilingual in the following five languages: Enets, Nganasan, Nenets, Evenki, and Russian (Figure 4). The number of Enets individuals who were able to converse in each of the other four languages differed and generally was a property of the individuals who had regular social contact with speakers of the other four languages. (…) Note that in all cases of interethnic communication there could well be a lack of perfect proficiency in a language for which the multilingualism is ascribed to the Enets community or Enets individuals: as Braunmüller and Ferraresi (2003: 3) put it: “Nobody would ever have expected to know other languages ‘perfectly’ (whatever that may mean in detail). This expectation seems to be a quite modern idea when discussing issues of bilingualism or multilingualism in general”.

    The complex interactions of Siberian populations during the 17th-19th centuries offer a reasonably good picture of the life in the centuries before these accounts, when Samoyedic peoples migrated northwards, and Palaeo-Siberian and Tungusic populations were gradually assimilated into their Uralic culture and language, through intermarriage and close contacts among naturally nomadic populations.

    You can read more about the origin of Nganasans – and other modern Samoyedic-speaking peoples – as Palaeo-Siberian populations (hence probably speaking Palaeo-Siberian languages more or less related to each other) who adopted Samoyedic languages in Wikipedia, which offers a summary of Boris Dolgikh’s On the Origin of the Nganasans (1962). Dolgikh is one of the main sources of information for these Siberian groups, as is reflected in this paper, too.

    samoyedic
    Map of distribution of Samoyedic languages (red) in the XVII century (approximate; hatching) and in the end of XX century (continuous background). Notice late expansion to north and west into the typical territory where Nomadic peoples roamed. Modified from Wikipedia, with the Tuva region labelled (see a recent genetic study on the Tuva region, one of the most likely to be originally Samoyedic-speaking).

    Why some geneticists are using Nganasans – in fact the latest Palaeo-Siberians to learn Samoyedic, already during historic times – as a model for the expansion of Uralic? I have never understood that. Among the many cases of circular reasoning based on modern populations that have been created since the start of population genomics, the use of Nganasans as a model of ‘true Uralians’ is probably the most clearly frontally opposed to what was well known in anthropology before geneticists started this new field.

    If Kallio is right, most “eastern homeland” proposals are due to the interest of Russian nationalism, which is sadly quite likely to be influencing genetic research, too. It’s like letting Hindu nationalists influence publications on steppe-related migrations. As David Reich puts it in his book:

    The tensest twenty-four hours of my scientific career came in October 2008, when my collaborator Nick Patterson and I traveled to Hyderabad to discuss these initial results with Singh and Thangaraj.

    Our meeting on October 28 was challenging. Singh and Thangaraj seemed to be threatening to nix the whole project. Prior to the meeting, we had shown them a summary of our findings, which were that Indians today descend from a mixture of two highly divergent ancestral populations, one being “West Eurasians.” Singh and Thangaraj objected to this formulation because, they argued, it implied that West Eurasian people migrated en masse into India. They correctly pointed out that our data provided no direct evidence for this conclusion. They even reasoned that there could have been a migration in the other direction, of Indians to the Near East and Europe. (…) They also implied that the suggestion of a migration from West Eurasia would be politically explosive. They did not explicitly say this, but it had obvious overtones of the idea that migration from outside India had a transformative effect on the subcontinent.

    If you add the nation-building myths in Eastern Europe (like the Russian Euro-Asian movements) to the now prevalent Indo-European—CWC idea, and a Siberian ancestry peaking in the Arctic, with little demographic or political relevance of modern Uralic-speaking peoples, you have clearly an explosive sociopolitical mix (based on a mythical Pan-Eurasian Indo-Slavonic) in the making…

    euro-asian-empire-dugin
    Russia as the Euro-Asian Empire. Source: A. Dugin (1999), p. 415. From Eberhardt (2018).

    Related

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

    uralic-languages

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

    Interesting excerpts (emphasis mine):

    Methods

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

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

    The population structure of Uralic speakers

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

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

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

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

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

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

    uralic-modern-europe

    Identity-by-descent

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

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

    Time of Siberian admixture

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

    Affinities of the Uralic speakers with ancient Eurasians

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

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

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

    Discussion

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

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

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

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

    Comments

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

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

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

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

    yamna-corded-ware-qpgraph

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

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

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

    Related

    On the origin and spread of haplogroup R1a-Z645 from eastern Europe

    indo-european-uralic-migrations-corded-ware

    In my recent post about the origin and expansion of haplogroup R1b-L51, Chetan made an interesting comment on the origin and expansion of R1a-Z645. Since this haplogroup is also relevant for European history and dialectal North-West Indo-European and Indo-Iranian expansion, I feel compelled to do a similar post, although the picture right now is more blurry than that of R1b-L51.

    I find it interesting that many geneticists would question the simplistic approach to the Out of Africa model as it is often enunciated, but they would at the same time consider the current simplistic model of Yamna expansion essentially right; a model – if anyone is lost here – based on proportions of the so-called Yamnaya™ ancestral component, as found in a small number of samples, from four or five Eneolithic–Chalcolithic cultures spanning more than a thousand years.

    The “75% Yamnaya ancestry of Corded Ware”, which has been given so much publicity since 2015, made geneticists propose a “Yamna → Corded Ware → Únětice / Bell Beaker” migration model, in order of decreasing Yamnaya proportions. Y-DNA and solid archaeological models suggested that this model was wrong, and recent findings have proven it was. In fact, the CWC sample closest to Yamna was a late outlier of Esperstedt in Central Europe, whose ancestry is most likely directly related to Yamna settlers from Hungary.

    These wrong interpretations have been now substituted by data from two new early samples from the Baltic, which cluster closely to Yamna, and which – based on the Y-DNA and PCA cluster formed by all Corded Ware samples – are likely the product of female exogamy with Yamna peoples from the neighbouring North Pontic region (as we are seeing, e.g. in the recent Nikitin et al. 2018).

    NOTE. There is also another paper from Nikitin et al. (2017), with more ancient mtDNA, “Subdivisions of haplogroups U and C encompass mitochondrial DNA lineages of Eneolithic-Early Bronze Age Kurgan populations of western North Pontic steppe”. Link to paper (behind paywall). Most interesting data is summarized in the following table:

    yamna-corded-ware-mtdna

    Even after the publication of Olalde et al. (2018) and Wang et al. (2018) – where expanding Yamna settlers and Bell Beakers are clearly seen highly admixed within a few generations, and are found spread across a wide Eurasian cline (sharing one common invariable trait, the paternally inherited haplogroup, as supported by David Reich) – fine-scale studies of population structure and social dynamics is still not a thing for many, even though it receives more and more advocates among geneticists (e.g. Lazaridis, or Veeramah).

    NOTE. I have tried to explain, more than once, that the nature and origin of the so-called “Yamnaya ancestry” (then “steppe ancestry”, and now subdivided further as Steppe_EMBA and Steppe_MLBA) is not known with precision before Yamna samples of ca. 3000 BC, and especially that it is not necessarily a marker of Indo-European speakers. Why some people are adamant that steppe ancestry and thus R1a must be Indo-European is mostly related to a combination of grandaddy’s haplogroup, the own modern ethnolinguistic attribution, and an aversion to sharing grandpa with other peoples and cultures.

    In the meantime, we are seeing the “Yamnaya proportion” question often reversed: “how do we make Corded Ware stem from Yamna, now that we believed it?”. This is a funny circular reasoning, akin to the one used by proponents of the Franco-Cantabrian origin of R1b, when they look now at EEF proportions in Iberian R1b-L23 samples. It seems too comic to be true.

    R1a and steppe ancestry

    The most likely origin of haplogroup R1a-Z645 is to be found in eastern Europe. Samples published in the last year support this region as a sort of cradle of R1a expansions:

    • I1819, Y-DNA R1a1-M459, mtDNA U5b2, Ukraine Mesolithic ca. 8825-8561 calBCE, from Vasilievka.
    • I5876, Y-DNA R1a, mtDNA U5a2a, Ukraine Mesolithic 7040-6703 calBCE, from Dereivka.
    • I0061, hg R1a1-M459 (xR1a1a-M17), mtDNA C1, ca. 6773-6000 calBCE (with variable dates), from Yuzhnyy Oleni Ostrov in Karelia.
    • Samples LOK_1980.006 and LOK_1981.024.01, of hg MR1a1a-M17, mtDNA F, Baikalic cultures, dated ca. 5500-5000 BC.
    • Sample I0433, hg R1a1-M459(xM198), mtDNA U5a1i, from Samara Eneolithic, ca. 5200-4000 BCE
    • Samples A3, A8, A9, of hg R1a1-M459, mtDNA H, from sub-Neolithic cultures (Comb Ware and Zhizhitskaya) at Serteyea, although dates (ca. 5th-3rd millennium BC) need possibly a revision (from Chekunova 2014).

    NOTE. The fact that Europe is better sampled than North Asia, coupled with the finding of R1a-M17 in Baikalic cultures, poses some problems as to the precise origin of this haplogroup and its subclades. While the first (Palaeolithic or Mesolithic) expansion was almost certainly from Northern Eurasia to the west – due to the Mal’ta sample – , it is still unknown if the different subclades of R1a in Europe are the result of local developments, or rather different east—west migrations through North Eurasia.

    Y-Full average estimates pointed to R1a-M417 formation ca. 6500 BC, TMRCA ca. 3500 BC, and R1a-Z645 formation ca. 3300 BC, TMRCA ca. 2900 BC, so the most likely explanation was that R1a-Z645 and its subclades – similar to R1b-L23 subclades, but slightly later) expanded quickly with the expansion of Corded Ware groups.

    The presence of steppe ancestry in Ukraine Eneolithic sample I6561, of haplogroup R1a-M417, from Alexandria, dated ca. 4045-3974 calBCE, pointed to the forest steppe area and late Sredni Stog as the most likely territory from where the haplogroup related to the Corded Ware culture expanded.

    However, the more recent Y-SNP call showing R1a-Z93 (L657) subclade rendered Y-Full’s (at least formation) estimates too young, so we have to rethink the actual origin of both subclades, R1a-Z93 (formation ca. 2900 BC, TMRCA ca 2700 BC), and R1a-Z283 (formation ca. 2900 BC, TMRCA ca. 2800 BC).

    Contrary to what we thought before this, then, it is possible that the expansion of Khvalynsk-Novodanilovka chieftains through the steppes, around the mid-5th millennium BC, had something to do with the expansion of R1a-Z645 to the north, in the forest steppe.

    We could think that the finding of Z93 in Alexandria after the expansion of Khvalynsk-Novodanilovka chiefs would make it more likely that R1a-Z645 will be found in the North Pontic area. However, given that Lower Mikhailovka and Kvitjana seem to follow a steppe-related cultural tradition, different to forest steppe cultures (like Dereivka and Alexandria), and that forest steppe cultures show connections to neighbouring northern and western forest regions, the rest of the expanding R1a-Z645 community may not be related directly to the steppe at all.

    Adding a hypothetical split and expansion of Z645 subclades to the mid-/late-5th millennium could place the expansion of this haplogroup to the north and west, pushed by expanding Middle PIE-speaking steppe peoples from the east:

    distribution-horse-scepters
    Schematic depiction of the spread of horse-head scepters in the Middle Eneolithic, representing expanding Khvalynsk-Novodanilovka chieftains. See a full version with notes here.

    The Złota culture

    I have already written about the Podolia-Volhynia region: about the North Pontic steppe cultures in contact with this area, and about the chaotic period of migrations when Corded Ware seem to have first emerged there among multi-directional and multi-ethnic migrants.

    This is what Włodarczak (2017) says about the emergence of Corded Ware with ‘steppe features’ after the previous expansion of such features in Central Europe with Globular Amphorae peoples. He refers here to the Złota culture (appearing ca. 2900-2800 BC) in Lesser Poland, believed to be the (or a) transitional stage between GAC and Corded Ware, before the emergence of the full-fledged “Corded Ware package”.

    So far, to the north of the Carpathian Mountains, including Polish lands, no graves indicating their relationship with communities of the steppe zone have been found. On the contrary, the funeral rites always display a local, central European nature. However, individual elements typical of steppe communities do appear, such as the “frog-like” arrangement of the body (Fig. 20), or items associated with Pit Grave milieux (cf. Klochko, Kośko 2009; Włodarczak 2014). A spectacular example of the latter is the pointed-base vessel of Pit Grave culture found at the cemetery in Święte, site 11 near Jarosław (Kośko et al. 2012). These finds constitute a confirmation of the importance of the relationships between communities of Pit Grave culture and Corded Ware culture. They are chronologically diverse, although most of them are dated to 2600-2400 BC – that is, to the “classic” period of Corded Ware culture.

    funnelbeaker-trypillia-corded-ware
    Map of territorial ranges of Funnel Beaker Culture (and its settlement concentrations in Lesser Poland), local Trypillian groups and early Corded Ware Culture settlements (◼) at the turn of the 4th/3rd millennia BC.

    However, when discussing the relationships with the steppe communities, Polish lands deserve particular attention since part of the groups inhabiting it belonged to the eastern province of Corded Ware culture (cf. Häusler 2014), which neighboured Pit Grave culture both from the east and south. In addition, there was a tradition of varied relationships with the north Pontic zone, which began to intensify from the second half of the 4th millennium BC (Kośko, Szmyt, 2009; Kośko, Klochko, 2009). These connections are especially readable in Małopolska and Kujawy (Kośko 2014; Włodarczak 2014). The emergence of the community of Globular Amphora culture in the north Pontic zone at the end of the 4th and the beginnings of the 3rd millennium BC (Szmyt 1999) became a harbinger of a cultural closening between the worlds of central Europe and the steppe.

    The second important factor taking place at that time was the expansion of the people of Pit Grave culture in a westerly direction, along the Danube thoroughfare. As a result of this, also to the south of the Carpathian Mountains, e.g., along the upper Tisza River, a new “kurgan” cultural system was formed. As one outcome, the areas of central Europe, above all Małopolska, found themselves in the vicinity of areas inhabited by communities characterized by new principles of social organization and a new funeral rite. Around 2800 BC these changes became evident in different regions of Poland, with the most numerous examples being documented in south-eastern Poland and Kujawy. The nature of the funeral rite and the features of the material culture perceptible at that time do not have straight forward analogies in the world of north Pontic communities. In this respect, the “A-horizon” is a phenomenon of local, central European origin. The events preceding the emergence of the said horizon (that is, the expansion of the people of Pit Grave culture into the area north of the arc of the Carpathians) are nowadays completely unidentifiable and remain merely an interesting theoretical matter (cf. e.g., Kośko 2000). Therefore, analysis of the archaeological sources cannot confirm the first archaeogenetic analysis suggesting a bond between the communities of the Pit Grave culture and Corded Ware culture (e.g., Haak et al. 2015).

    Artefacts of the “A-horizon”, i.e., shaft-hole axes, amphorae (Fig. 21), beakers, and pots with a plastic wavy strip (Fig. 7) are found in different funerary and settlement contexts, sometimes jointly with finds having characteristics of various cultures (e.g., in graves of Złota culture, or at settlements of Rzucewo culture). Hence, they primarily represent a chronological phase (c. 2800-2600 BC), one obviously related to the expansion of a new ideology.

    Eastern CWC expansion

    Before continuing tracing the Corded Ware culture’s main features, it is worth it to trace first their movement forward in time, as Corded Ware settlers, from Poland to the east.

    Circum-Baltic CWC

    According to Klochko and Kośko (1998):

    The colonizing Neolithic waves are continued by the Circum-Baltic Corded Ware culture, closely related to the traditions of the Single Grave culture and traditions of the Northern European Lowlands. After ca. 2900 BC, certain cultural systems with ‘corded’ traits –genetically related to the catchment area of the south-western Baltic – appear in the drainages of the Nemen, Dvina, Upper Dnieper, and even the Volga. These communities are considered the vector of Neolithisation in the Forest Zone.

    east-european-fatyanovocwc
    East European movement directions (arrows) of the representatives of the Central European Corded Ware Culture. Modified from I.I. Artemenko.

    The picture in the Baltic (Pamariu / Rzucewo) and Finland (Battle Axe) is thus more or less clearly connected with early dates ca. 2900-2800 BC:

    There is a clear interaction sphere between the eastern Gulf of Finland area – reaching from Estonia to the areas of present-day Finland and the Karelian Isthmus in Russia –, evidenced e.g. by the sharp-butted axes, derived from the Estonian Karlova axe.

    Interesting in this regard is the expansion of the Corded Ware culture in Finland, into a far greater territory than previously thought, that is poorly represented in most maps depicting the extent of the culture in Europe. Here is summary of CWC findings in Finland, using images from Nordqvist and Häkäla (2014):

    finland-cwc
    Corded Ware culture remains in Finland, excluding the so-called ‘imitations’. [Notice in the top left image the often depicted border of the culture]. Combination of maps from Nordqvist & Häkälä (2014)

    Middle Dnieper and Fatyanovo

    The earliest Middle Dnieper remains are related to CWC graves between the Upper Vistula and the Bug, containing pottery with Middle Dnieper traits, dated probably ca. 2700 BC, which links it with the expansion of the A-horizon. In fact, during the period ca. 2800-2400 BC, the area of Lesser Poland (with its numerous kurgans and catacomb burials) is considered the western fringe of an area spreading to the east, to the middle Dniester and middle Dnieper river basins, i.e. regions bordering the steppe oecumene. This ‘eastern connection’ of funeral ritual, raw materials, and stylistic traits of artefacts is also identified in some graves of the Polish Lowlands (Włodarczak 2017).

    gac-cwc-baltic-dnieper
    Cultural situation in Eastern Europe in approximately the middle of the III mill. BC. Key: 1 – areas settled by Globular Amphora culture populations; 2 – areas penetrated by Globular Amphora culture populations; 3 – border between central and eastern group; 4 – Pamariu/Rzucewo culture area; 5 – zone of Pamariu/Rzucewo culture influences; 6 – directions of Comb Pottery culture influence; 7 – Zhizhitskaya culture; 8 – eastern border of “pure” Corded Ware site; 9 – North Belarussian culture; 10 – Middle Dnieper culture; 11 – Fatyanovo culture; 12 – Yamnaya culture; 13 – eastern border of Dniester group; 14 – Kemi-Oba culture and influences; 15 – Foltesti culture; 16 – syncretic sites with evidence of Globular Amphora culture traits (1 – Nida; 2 – Butinge; 3 – Palanga; 4 – Juodkrante; 5 – Azyarnoye; 6 – Mali Rogi; 7 – Prorva; 8 – Strumen/Losha; 9 – Syabrovichi; 10 – Luchin-Zavale; 11 – Lunevo (?); 12 – Belynets; 13 – Losiatyn; 14 – Corpaci; 15 – Ocnita; 16-17 – Camenca; 18 – Marculesti; 19 – Orhei; 20 – Efimovka; 21 – Tatarbunary; 22 – Novoselitsa; 23 – Primorskoye; 24 – Sanzhiyka; 25 – Akkermen; 26 – Maydanetskoye; 27 – Grigorevka; 28 – Kholmskoye; 29 – Purcari; 30 – Roscani; 31 – Semenovka; 32 – Grishevka; 33 – Durna Skela; 34 – Iskovshchina; 35 – Primorskoye); 17 – borders of ecological zones. From Szmyt (2010)

    The Fatyanovo (or Fatyanovo-Balanovo) culture was the easternmost group of the Corded Ware culture, and occupied the centre of the Russian Plain, from Lake Ilmen and the Upper Dnieper drainage to the Wiatka River and the middle course of the Volga. From the few available dates, the oldest ones from the plains of the Moskva river, and from the late Volosovo culture containing also Fatyanovo materials, and in combination they show a date of ca. 2700 BC for its appearance in the region. The Volosovo culture of foragers eventually disappeared when the Fatyanovo culture expanded into the Upper and Middle Volga basin.

    The origin of the culture is complicated, because it involves at its earliest stage different Corded Ware influences in neighbouring sites, at least on the Moskva river plains (Krenke et al. 2013): some materials (possibly earlier) show Circum-Baltic and Polish features; other sites show a connection to western materials, in turn a bridge to the Middle Dnieper culture. This suggests that groups belonging to different groups of the corded ware tradition penetrated the Moscow region.

    The split of subclades Z93 – Z283

    If we take into account that the split between R1-Z93 and R1a-Z283 must have happened during the 5th millennium BC, we have R1a-Z93 likely around the middle Dnieper area (as supported by the Alexandria sample), and R1a-Z283 possibly to the north(-west), so that it could have expanded easily into Central Europe, and – through the northern, Baltic region – to the east.

    Where exactly lies the division is unclear, but for the moment all reported Circum-Baltic samples with Z645 subclades seem to belong to Z282, while R1a samples from Sintashta/Potapovka (including the Poltavka outlier) point to Abashevo being dominated by R1a-Z93 subclades.

    We have to assume, then, that an original east-west split betwen R1a-Z283 and R1a-Z93 turned, in the eastern migrations, into a north-south split between Z282 and Z93, where Finland and Battle Axe in general is going to show Z282, and Middle Dnieper – Abashevo Z93 subclades.

    copper-age-early-cwc
    Early Copper Age migrations ca. 3100-2600 BC.

    I can think of two reasons why this is important:

    1. Depending on how Proto-Corded Ware peoples expanded, we may be talking about one community overcoming the other and imposing its language. Because either
      • clans of both Z93 and Z283 were quite close and kept intense cultural contacts around Dnieper-Dniester area; or
      • if the split is as early as the 5th millennium BC, and both communities separated then without contact, we are probably going to see a difference in the language spoken by both of them.
    2. In any case, the main north-south division of eastern Corded Ware groups is pointing to an important linguistic division within the Uralic-speaking communities, specifically between a Pre-Finno-Ugric and a Pre-Samoyedic one, and potentially between Pre-Finno-Permic and Pre-Ugric.

    These may seem irrelevant questions – especially for people interested only in Indo-European migrations. However, for those interested in the history of Eurasian peoples and languages as a whole, they are relevant: even those who support an ‘eastern’ origin of Proto-Uralic, like Häkkinen, or Parpola (who are, by the way, in the minority, because most Uralicists would point to eastern Europe well before the Yamna expansion), place the Finno-Ugric expansion with the Netted Ware culture as the latest possible Finno-Ugric immigrants in Fennoscandia.

    The Netted Ware culture

    The image below shows the approximate expansion of Corded Ware peoples of Battle Axe traditions in Finland, as well as neighbouring Fennoscandian territories, from ca. 2800 BC until the end of the 3rd millennium. A controversial 2nd (late) wave of the so-called Estonian Corded Ware is popular in texts about this region, but has not been substantiated, and it seems to be a regional development, rather than the product of migrations.

    finland-corded-ware
    Left: Corded Ware remains in Finland from ca. 2800 BC, according to Nordqvist & Häkälä (2014), combined in a single image. Right: Distribution of the Corded Ware culture within Finland. Mapped (black dots) are finds of typical stone battle axes, used as a proxy (data from [8]). The red isolines indicate average permanent snow cover period from 1981 to 2010 (data from [9]). A recent study estimates the snow cover period ca 4500 years ago would have been 40–50 days less than today [10]. Overlying coloration refers to the lactose persistance (LP) allele gradient in modern northeastern Europe (see the electronic supplementary material, appendix B: Material and methods and table 1, for details); lozenge dots specify the dataset mean points for the triangulation. From Cramp et al. (2014).

    As we have seen, Fatyanovo represents the most likely cultural border zone between Circum-Baltic peoples reaching from the Russian Battle Axe to the south, and Middle Dnieper peoples reaching from Abashevo to the north. In that sense, it also represents the most likely border culture between north-western (mainly R1a-Z282) and south-eastern (R1a-Z93) subclades.

    With worsening climatic conditions (cooler seasons) at the end of the 3rd millennium, less settlements are apparent in the archaeological record in Finland. After ca. 2000 BC, two CWC-related cultures remain: in the coast, the Kiukainen culture, derived from the original Circum-Baltic Corded Ware settlers, reverts to a subsistence economy which includes hunting and fishing, and keeps mainly settlements (from the best territories) along the coast. In the inland, Netted Ware immigrants eventually appear from the south.

    cultures-western-finland
    Image modified from Cramp et al. (2014) “The timeline shows the archaeological cultures
    discussed here alongside actual sherds sampled and typical vessel forms (after [26–28]) (latter not shown to scale). Distribution maps show the geographical range of (f) Typical Comb Ware, (g) Corded Ware, (h) Kiukainen Ware and (i) Bronze Age cultures in the region (after [10,20,29]).”

    The Netted Ware culture emerged in the Upper Volga–Oka region, derived from the Abashevo culture and its interaction with the Seima-Turbino network, and spread ca. 1900-1800 BC to the north into Finland, spreading into eastern regions previously occupied by cultures producing asbestos and organic-tempered wares (Parpola 2018).

    NOTE. Those ‘contaminated’ by the Copenhagen fantasy map series may think that Volosovo hunter-gatherers somehow survived the expansion of Fatyanovo-Balanovo and Abashevo, hidden for hundreds of years in the forest, and then reappeared and expanded the Netted Ware culture. Well, they didn’t. At least not in archaeological terms, and certainly not with the genetic data we have.

    If we combine all this information, and we think about these peoples in terms of Pre-Finno-Permic and Pre-Ugric languages developing side by side, we get a really interesting picture (see here for Proto-Fennic estimates):

    • The Battle Axe around the Baltic Sea – including the Gulf of Finland and Scandinavia – would be the area of expansion of Pre-Finno-Permic peoples, of R1a-Z283 subclades, which became later concentrated mainly on coastal regions;
    • the southern areas may correspond to Pre-Ugric peoples, which expanded later to the north with Netted Ware (see image below) – their precise subclades may be dependent on what will be found in Fatyanovo;
    • and Pre-Samoyedic peoples (of R1a-Z93 subclades) would have become isolated somewhere in the Cis- or (more likely) Trans-Urals region after 2000 BC, possibly from the interaction of the latest Balanovo stages and the Seima-Turbino phenomenon.
    netted-ware-parpola
    Distribution of the Netted Ware according to Carpelan (2002: 198). A: Emergence of the Netted Ware on the Upper Volga c. 1900 calBC. B: Spread of Netted Ware by c. 1800 calBC. C: Early Iron Age spread of Netted Ware. (After Carpelan 2002: 198 > Parpola 2012a: 151.)

    These communities in contact would have allowed for:

    • the known Indo-Iranian loanwords in Finno-Ugric to spread through a continuum of early dialects formed by Abashevo – Fatyanovo – Battle Axe groups;
    • the Finno-Saamic substrate of Germanic to be associated with Battle Axe groups in Scandinavia;
    • the important Palaeo-Germanic loanwords in Finno-Saamic spreading with long-term contacts (from Pre-Germanic to the Proto-Germanic, and later North Germanic period) through the Baltic Sea, between Scandinavia and the Gulf of Finland;
    • and Tocharian contacts with Samoyedic (although limited, and in part controversial), which point to its early expansion to the east of the Ural Mountains.

    On the other hand, if one is inclined to believe that R1a and steppe ancestry do represent Indo-European speakers… which language was spoken from the Gulf of Finland well into the north, the inland, and Karelia, and in Northern Russia, by Corded Ware peoples and their cultural heirs (like Kiukainen or Netted Ware) for almost three thousand years?

    Because we know that no other peoples of different haplogroups dominated over eastern Fennoscandia until the Iron Age, and N1c and Siberian ancestry expanded separately, and probably due to late bottlenecks, especially with Fennic peoples expanding recently to the north at the expense of the Saami population.

    After the expansion of Bell Beaker peoples, the geographic distribution of late Corded Ware groups in the second half of the 3rd millennium, just before their demise – and before the expansion of Netted Ware to the north – , can be depicted thus as follows:

    bronze_age_early_cut
    Early Bronze Age Europe.

    Territories in cyan must then represent, for some people who believe in an archaic Indo-Slavonic of sorts, the famous Fennoscandian Balto-Slavic to the north (before they were displaced by incoming Finno-Saamic peoples of hg N1c during the Iron Age and up to the Middle Ages); and the also famous Tundra-Forest Indo-Iranian in the Upper Volga area, a great environment for the development of the two-wheeled chariot…

    But let’s leave the discussion on imaginary IE dialects for another post, and continue with the real question at hand.

    A steppe funerary connection?

    Back to Złota as a transitional culture, we have already seen how the corded ware vessels characteristic of the Classic CWC are related to Globular Amphora tradition, and show no break with this culture. It is usually believed that the funerary rites were adopted from steppe influence, too. That is probably right; but it does not mean that it came from Yamna or other coeval (or previous) steppe culture; at least not directly.

    NOTE. A similar problem is seen when we read that Mierzanowice or Trzciniec show “Corded Ware” traits from a neighbouring CWC group, when CWC groups disappeared long before these cultures emerged. For cultural groups that are separated centuries from each other, an assertion as to their relationship needs specifics in terms of dates and material connection, or it is plainly wrong.

    These are the funerary ritual features from Złota (later specialized in Corded Ware), as described by Włodarczak (2017):

    • Single burial graves; along with the habit of interring the deceased in multiple burial graves, but emphasizing their individual character by careful deposition of the body and personal nature of the grave goods.
    • Grave goods with materials and stylistiscs belonging to an older system (e.g. amber products); and others correlated to the ‘new world’ of the CWC, such as flint products made of the raw materials tipical of Lesser Poland’s CWC, copper ornaments, stone shaft-hole axes, bone and shell ornaments, and characteristic forms of vessels like beakers and amphoras.
    • Military goods, which would become prevalent in later periods, are present in a moderate number, compatible with their lesser importance.
    • There are also cases of the characteristic catacomb (“niche”) graves – with an entrance pit, a more extensive niche, and a narrow corridor leading to a vault – , as well as some individual cases of application of ochre and deformation of skulls.
    catacomb-grave-ksiaznice
    Catacomb grave no. 2a/06 from Książnice, Złota culture (acc. to Wilk 2013). Image from Włodarczak (2017)

    It seems that the Złota funerary tradition was also “transitional”, like corded ware vessels, into the classical Corded Ware ideology. But “transitional” from what exactly? Yamna? Probably not.

    The Lublin-Volhynia culture

    One needs not look for a too distant culture to find similarities. Włodarczak (2017) points to CWC in south-eastern Poland and Kuyavia showing, by the time of the Yamna expansion, a funeral rite and features of the material culture without straightforward analogies in the world of north Pontic communities, and thus suggests that the “A-horizon” is a local phenomenon of central European origin.

    This assertion is interesting, in so far as most Corded Ware samples investigated to date seem to come precisely from an East-Central territory near the Ukraine forest steppe, with a cluster already established by the end of the 5th millennium:

    PCA-caucasus-lola-ane-chg
    Image modified from Wang et al. (2018). Samples projected in PCA of 84 modern-day West Eurasian populations (open symbols). Previously known clusters have been marked and referenced. An EHG and a Caucasus ‘clouds’ have been drawn, leaving Pontic-Caspian steppe and derived groups between them.See the original file here.

    The following text is from Stanisław Wilk (2018), about the Lublin-Volhynian (and related) cemeteries at Wyciąże and Książnice:

    lublin-volhynia-culture
    A reach of the Wyciąże-Złotniki group and Lublin-Volhynian culture in the south-eastern Poland and western Ukraine: 1. Area of the Wyciąże-Złotniki group; 2. Area of the Lublin-Volhynian culture. A. Cemetery of the Lublin-Volhynian culture at site 2 in Książnice; B. Cemetery of the Wyciąże-Złotniki group at site 5 in Kraków Nowa Huta-Wyciąże (drawing by S. Wilk based on Zakościelna 2006 and Nowak 2014, on a background downloaded from https://maps-for-free.com/).

    Regardless of the differences between the two necropolises (such as the number of burials, the area which has been explored, the orientation and layout of burials), it seems that they have several key elements in common:

    • concentration of graves in separate cemeteries;
    • differentiation of burials with regard to sex (the principle of the ‘left ̶ right’ side, different burial goods for males and females);
    • stratification of graves with regard to the richness of their inventories (this mainly applied to copper artefacts);
    • occurrence of indicators of the richest male burials (a copper dagger in Wyciąże, a copper battle axe, a small axe and a chisel in Książnice);
    • allocation of a separate area for elite burials (the eastern burial area in Książnice, and the southeastern and north-central part of the necropolis in Wyciąże), as well as one for egalitarian burials (the western area in Książnice, and the south-central and western part of the cemetery in Wyciąże).
    lublin-volhynian-eneolithic-cemetery
    Plan of the Lublin-Volhynian culture cemetery at site 2 in Książnice: 1. female graves; 2. man graves; 3. copper traces; 4. cenothap; 5. cremation grave; 6. partial grave; 7. estimated area of the L-VC cemetery; 8. estimated area of an elite and poor burial fields; 9. area of burials containing copper artefacts (drawing by S. Wilk).

    The above-mentioned characteristics prove that the patterns of social and religious behaviours from areas lying beyond the Carpathian Mountains exerted a strong influence on the two societies living in Lesser Poland.

    Anna Zakościelna, while describing the similarities between the burial ritual of the late Polgár groups and cultures from areas on the Tisza river and the Lublin-Volhynia culture, claimed that:

    a characteristic feature of the burial ritual of both cultures was practicing various group norms, which required different treatment of the deceased depending on their sex, age and social rank. As in the Lublin-Volhynia culture, the opposition ‘male – female’ can the most clearly be observed ̶ particularly, in the consistent positioning of males on the right, and females, on the left side. And, there is much indication that this ritual norm divided the deceased from early childhood (Sofaer Derevensky 1997: 877, Tab. 1; Lichter 2001: 276- 280, 322-323) (Zakościelna 2010: 227-228).

    It seems that these observations can also be extended to the Wyciąże-Złotniki group.

    Another question is whether the evidence of the influences of the copper civilization observed in both cemeteries emerged as a result of the literal copying of patterns from the south, or whether the latter were only a source of inspiration for local solutions.

    Looking at this problem form the perspective of the details of burial ritual, between the Carpathian Basin and Lesser Poland, we can observe clear differences, among others, in the size of cemeteries and orientation of burials. While, in the Carpathian Basin there were large necropolises, consisting of several dozen burials located in rows, with the dominant orientation along the SE-NW and E-W axis (Lichter 2001: Abb. 123, 143; Kadrow 2008: 87); in Lesser Poland there were small cemeteries of several to a dozen or so burials, mostly oriented along the S-N axis (in the Lublin-Volhynia culture; Zakościelna 2010: 66), as well as S-E and NE-SW (in the Wyciąże-Złotniki group; Kaczanowska 2009: 77). Similarly, there are differences in the details of the burial goods. North of the Carpathians, there is a much smaller frequency of copper artefacts, particularly in the group of prestigious, heavy items (battle axes, axes and daggers), as well as a complete lack of objects made of gold. Want is more, the pottery found in the graves has a distinct local character, only supplemented by imitating or imports from areas beyond the Carpathians (Zakościelna 2006: 85; Nowak 2014: 273; a different opinion Kozłowski 2006: 57). Therefore, the suggestion made by Nowak seems right ̶ namely, that these influences were not caused by migrations of groups of the population living on the Tisza river to Lesser Poland, but were rather due to processes of selective cultural transmission (Nowak 2014: 273).

    Therefore, the sharing of a similar funerary rite (as happened later between Lublin-Volhynia and Złota), although it shows a strong cultural connection with autochthonous cultures, is obviously not the same as sharing ancestors; and even if it were so, they would not need to be paternal ancestors. But it shows that important Corded Ware cultural traits are local developments, and it disconnects thus still more supposed CWC ‘steppe traits’ from steppe cultures, and connects them with the first steppe-related cultural wave that reached central Europe in the 5th millennium BC.

    Prehistoric Pontic—Caspian links

    How would a Lublin-Volhynia culture be related to the North Pontic area ca. 4500-3000 BC? We can enjoy the map series of Baltic—Pontic migrations by Viktor Klochko (2009), and make a wild guess:

    baltic-pontic-routes
    Pontic—Baltic routes of migrations during the Eneolithic. Top left: Linear Pottery expansion. Top right: Funnel Beaker expansion. Bottom left: late Trypillia expansion. Bottom right: GAC expansion.

    And then read the account of Sławomir Kadrow, in Exchange of People, Ideas and Things between Cucuteni-Trypillian Complex and Areas of South-Eastern Poland (2016):

    In the second half of the 5th millennium BC (horizon 1), communities of the Tripolye culture, phases BI-BII, had contacts with the population of the late (IIa) phase of the Malice culture. The areas settled by both cultural complexes were located at a great distance from each other. The communities of the Tripolye culture adopted selected features of Malice ceramic production (fig 2). This seems to have resulted from marital exchange: on a moderate scale, Tripolye men sought out their wives in the area of the Malice culture and, according to patrilocal marriage customs, the women then moved to the Tripolye settlements, sporadically transferring ready-made ceramic products, so-called imports, to the Tripolye culture. Thus, the wives were responsible for the considerably more numerous imitations of the Malice ceramics and the long-lasting, though selective, traditions of Malice pottery passed down in their new environment. The patrilocal marriage customs involving the Malice women and the Tripolye men (never the other way round), and the fact that pottery was women’s domain, led to the unidirectional transfer of vessels, technology and norms of ceramic production from the Malice culture to the Tripolye culture.

    The turn of the 5th and the 4th millennia and the early 4th millennium BC (horizon 2) witnessed the deepening interaction between the populations of the youngest (IIb) phase of the Malice culture and the classic (II) phase of the Lublin-Volhynia culture on the one hand and the communities of phase BII of the Tripolye culture on the other. The Danube and the Tripolye settlement complexes came into contact on the upper Dniester and between the Styr and the Horyn rivers in Volhynia. This helped to continue the previous forms of marital exchange, which resulted in further popularisation of the ceramics and the traditions of ceramic production typical of the Danube cultures, i.e. the Malice and the Lublin-Volhynia cultures, and also the Polgár culture, in the areas settled by the Tripolye cultural complex.

    As the civilizational norms of the Eneolithic (Copper) Age became widespread in that period, the forms of interaction described above acquired new elements. The deepening internal diversification of the early Eneolithic communities of the Lublin-Volhynia culture led to a growing demand for prestige objects, which was met with import or imitation of copper artefacts, mainly those from the Carpathian Basin, and with flint tools produced from long blades. That type of flint production depended largely on new technologies derived from the Tripolye culture, as proven by such borrowings as troughlike retouch or the very idea and technology for the production of long flint blades in the Lublin-Volhynia culture. It seems that the influx of Tripolye settlers into flintbearing areas in Volhynia and on the upper Dniester, adjacent to the settlement centres of the late phase of the Malice culture and the Lublin-Volhynia culture, created sufficient conditions for the expanding influence of the Tripolye flint working on the communities of the Eneolithic Lublin-Volhynia culture.

    In the mid-4th millennium BC (horizon 3), those forms of interaction between the Danube communities (the late phase of the Lublin-Volhynian culture) and the Tripolye communities (phase CI)were continued. Elements of the Danube pottery still grew in popularity in the Tripolye population, while selected features of the Tripolye flint working were adopted by the Lublin-Volhynia culture.

    In that period, the population of the Funnel Beaker culture of the pre-classic and early classic phases (the beginnings of Gródek 1 and Bronocice III), until then absent from those areas, quite quickly drove out and replaced the Danube population in western Volhynia and the upper Dniester basin. This caused significant changes in the forms and intensity of the intercultural interaction, which became fully apparent already in the 2nd half of the 4th millennium BC.

    In the following period (horizon 4), the population of the classic phase of the Funnel Beaker culture (Gródek 1, Bronocice III) settled more and more intensively the upper Dniester basin, up to the Hnyla Lypa river, and western Volhynia, up to the Styr river. East of those rivers, the Funnel Beaker settlers created considerable areas where they mixed with settlers from early phase CII of the Tripolye culture. Their coexistence, lasting there for many generations, resulted in deepening the interactions between members of both cultural complexes and in developing entirely new forms of relationships.

    (…)

    The intensifying interaction between the communities of the Funnel Beaker culture and the Tripolye culture, early phase CII, in the 2nd half of the 4th millennium BC (horizon 4) was an introduction to, and perhaps a condition for, even more frequent contacts in the next period, the first centuries of the 3rd millennium BC (horizon 5). In that case, the interaction was mainly triggered by multidirectional migrations of larger human groups, involving a significant part of the population of all cultures from the areas discussed here. The Tripolye communities of younger phase CII settled Volhynia, its eastern areas in particular, from the south and the south-east, while groups representing the younger phases of the Funnel Beaker culture (Gródek 2), often with Baden features (Bronocice IV and V), moved increasingly into the western part of that region. The Yamna communities expanded along the lower and central Danube to the west, whereas the populations of the late phase of the Baden culture took the opposite direction, reaching as far as Kiev in the northeast, and contributed to the cultural character of the Sofievka group.

    The communities of the Globular Amphora culture migrated from the north-west, from eastern Poland, towards the Danube Delta and as far as the Dnieper in the east, while the multicultural population from the areas around the mouth of the Danube moved in the opposite direction, carrying with them cultural elements from Thrace, or even from Anatolia. Some of them returned to the starting point (to south-eastern Poland), bringing with them a new form of pottery, so-called Thuringian amphora, borrowed from the late Trypillian Usatovo group. This resulted in origins of the Złota culture, a cultural phenomenon that gave beginnings to the oldest Corded Ware culture. Inventories of both cultures contained the already mentioned Thuringian amphorae.

    lublin-volhynia-alexandria
    Graves and cemeteries with gender differentiated burial rites in Europe; A — Hamangia and Varna cultures; B — Tiszapolgar and Bodrogkeresztur cultures; C — Lublin-Volhynia culture; D — Brześć Kujawski culture. Added star symbol with approximate location of the Alexandria site. Modified image from Sławomir Kadrow (2016)

    Here is a more recent assessment (2017) of the latest radiocarbon analyses of the available settlements of cultures in the area, published by Marek Novak (announced in a previous post), which gives the following data on Wyciąże-Złotniki, Lublin-Volhynia, and Wyciąże/Niedźwiedź:

    This scheme unambiguously suggests both the overlapping and contiguous nature of cultural development in western Lesser Poland within the Middle Neolithic. The basic elements of this development are: 1) the Wyciąże-Złotniki group and the Lublin-Volhynian culture, until c. 3650–3550 cal BC; 2) the Funnel Beaker culture proper, which appeared c. 3750–3700c al BC, and existed until c. 3300–3250 cal BC, perhaps accompanied by the Wyciąże/Niedźwiedź materials from c. 3650–3550 cal BC; and 3) the Baden culture and the Funnel Beaker/Baden assemblages from 3100 and 3300–3100 cal BC, respectively, until 2850–2750 and 2850 cal BC, with – possibly – later Funnel Beaker culture and Wyciąże/ Niedźwiedź materials, existing until c. 3100 cal BC.

    The final scheme shows that the Lublin-Volhynian culture could have coincided with the Wyciąże-Złotniki group. In view of the territorial relationship between them, relations from the point of view of material culture, primarily in the field of pottery, become particularly interesting. It is relatively easy to see clear similarities between these units. However, the most evident similarities apply only to some categories of ceramics, including, for example, vessels with Scheibenhenkel handles. What is more, in the period between the late 38th and early 36th centuries BC, the early Funnel Beaker and possibly early Baden influences are superimposed on this Lublin-Volhynian/Wyciąże-Złotniki ‘mix’.

    [About Corded Ware: The] development of this unit in central Europe, including western Lesser Poland, [] usually point to c. 2800 cal BC (Włodarczak 2006a). (…) the calibration curve makes it possible to alternatively refer several dates earlier than c. 3100 to c. 2850–2800 cal BC.

    Conclusion

    There is no direct archaeological link of Lublin-Volhynia-related groups with Corded Ware, beyond the fact that they shared homeland and Central European (‘steppe-related’) traits, as found in the Złota culture. But there is no direct link of Yamna with Corded Ware, either, whether in terms of culture or population.

    So, given the evident link of R1a-Z93 and steppe ancestry with the forest steppe ca. 4000 BC, the surrounding North Pontic areas in contact along the Dniester, Dnieper, Bug, and Prut are the best candidates for the appearance of R1a-Z283: steppe cultures to the south and south-west; sub-Neolithic (Comb Ware) groups to the north in the forest zone; and Eneolithic groups to the west and north-west.

    Seeing how ‘ancestral components’ and PCA cluster can change within a few generations, the question of the spread of R1a-Z645 subclades is still not settled by a single sample in Alexandria. However, based on the explosive expansions we are seeing from small territories, it would not be surprising to find R1a-Z93 and R1a-Z283 side by side in the same small area within the forest steppe.

    NOTE. An archaeological link may not mean anything relevant in genetics, especially – as in this case – when no clear migration event has been traced to date. We have seen exactly that with Kristiansen’s proposal of a long-term genetic admixture of Yamna with Trypillia and GAC to form Corded Ware, which didn’t happen. The cultural and ideological connection of CWC peoples with Lublin-Volhynian tradition may be similar to the already known connection with GAC, and not mean anything in genetic finds; at least in terms of Y-DNA haplogroup.

    We believed in the 2000s that Corded Ware represented the expansion of Late Proto-Indo-European, because the modern map of haplogroup R1a showed a distribution similar to how we thought the European and Indo-Iranian languages could have expanded. This has been proven wrong, and that’s what ancient DNA is for; not to confirm the own ideas or models, or to support modern ideologies.

    It is impossible to know if R1a-Z645 comes from the steppe, forest steppe, or forest zone, until more samples are published. I don’t think there will be any big surprise, no matter where it is eventually found. By now, adding linguistic reconstruction to archaeological traits, and to the genetic data from Yamna and Corded Ware settlers, the only clear pattern is that patrilineal clans expanded, during the Final Eneolithic / Chalcolithic:

    • Late Proto-Indo-European with Yamna and R1b-L23 subclades, given the known genomic data from Khvalynsk, Yamna, Afanasevo, Bell Beaker, Catacomb, and Poltavka—Sintashta/Potapovka.
    • Uralic with Corded Ware and R1a-Z645 subclades, given the known genomic data from Fennoscandia and the Forest Zone.

    Everything else is just wishful thinking at this moment.

    Related

    Y-DNA haplogroups of Tuvinian tribes show little effect of the Mongol expansion

    uralic-turkic

    Open access Estimating the impact of the Mongol expansion upon the gene pool of Tuvans, by Balanovskaya et al., Vavilov Journal of genetics and breeding (2018), 22(5):611-619.

    Abstract (emphasis mine):

    With a view to trace the Mongol expansion in Tuvinian gene pool we studied two largest Tuvinian clans – those in which, according to data of humanities, one could expect the highest Central Asian ancestry, connected with the Mongol expansion. Thus, the results of Central Asian ancestry in these two clans component may be used as upper limit of the Mongol influence upon the Tuvinian gene pool in a whole. According to the data of 59 Y-chromosomal SNP markers, the haplogroup spectra in these Tuvinian tribal groups (Mongush, N = 64, and Oorzhak, N = 27) were similar. On average, two-thirds of their gene pools (63 %) are composed by North Eurasian haplogroups (N*, N1a2, N3a, Q) connected with autochtonous populations of modern area of Tuvans. The Central Asian haplogroups (C2, O2) composed less then fifth part (17 %) of gene pools of the clans studied. The opposite ratio was revealed in Mongols: there were 10 % North Eurasian haplogroups and 75 % Central Asian haplogroups in their gene pool. All the results derived – “genetic portraits”, the matrix of genetic distances, the dendrogram and the multidimensional scaling plot, which mirror the genetic connections between Tuvinian clans and populations of South Siberia and East Asia, demonstrated the prominent similarity of the Tuvinian gene pools with populations from and Khakassia and Altai. It could be therefore assumed that Tuvinian clans Mongush and Oorzhak originated from autochtonous people (supposedly, from the local Samoyed and Kets substrata). The minor component of Central Asian haplogroups in the gene pool of these clans allowed to suppose that Mongol expansion did not have a significant influence upon the Tuvinan gene pool at a whole.

    tuvan-clans-y-dna

    Interesting excerpts:

    Haplogroup C2 peaks in Central Asia (Wells et al., 2001; Zerial et al., 2003), though its variants are abundant in other peoples of Siberia and Far East. For instance, in one of Buryat clans, namely Ekhirids, hg C2 frequency is 88 % (Y-base); in Kazakhs from different regions of Kazakhstan, total occurrence of hg C2 variants averages between 17 and 81 % (Abilev et al., 2012; Zhabagin et al., 2013, 2014, 2017), in populations of the Amur River (such as Nanais, Negidals, Nivkhs, Ulchs) – between 40 and 65 %, in Evenks – up to 68 % (Y-base), in Kyrgyz people of Pamir-Alay – up to 22 %, correspondingly; of all Turkic peoples of Altai, relatively high hg C2 frequency (16 %) is detected only in Telengits (Balanovskaya et al., 2014; Balaganskaya et al., 2011a, 2016). In Tuvinian clans under the study, hg C2 frequency is rather low – 19 % in Mongush and 11 % in Oorzhak, while in Mongols it makes up almost two thirds of the entire gene pool an comprises different genetic lines (subhaplogroups).

    tuvinian-y-chromosome
    Y-chromosomal haplogroup spectra in gene pools of Tuvinian Oorzhak and Mongush clans and of the neighboring populations of South Siberia and Central Asia.

    Haplogroup N is abundant all over North Eurasia from Scandinavia to Far East (Rootsi et al., 2007). The study on whole Y-chromosome sequencing conducted with participation of our group (Ilumäe et al., 2016) subdivided this haplogroup into several branches with their regional distribution. In gene pools of the Tuvans involved, hg N was represented by two sub-clades, namely N1a2 and N3a.

    Sub-clade N1a2 peaks in populations of West Siberia (in Nganasans, frequency is 92 %) and South Siberia (in Khakas 34 %, in Tofalars 25 %) (Y-base). In Tuvans, N1a2 occurrence is nearly 16 % in Mongush and 15 % in Oorzhak clans, respectively, while in Mongols, the frequency is three times less (5 %). Hg N1a2 is supposed to display the impact of the Samoyedic component to the gene pool of Tuvinian clans (Kharkov et al., 2013).

    Sub-clade N3a is major in the Oorzhak clan comprising almost half of the gene pool (45 %); it is represented by two sub-clades, namely N3a* and N3a5. The same sub-branches are specific to the Mongush clan as well, though with lower frequencies: N3a* – 9 % and N3a5 – 14 % (see Table). In Khori-Buryats from the Transbaikal region, a high frequency is observed – 82 % (Kharkov et al., 2014), while in Mongols, N3a5 occurs rather rarely (6 %). Hg N3a* was detected in populations of South Siberia only, and was widely spread in Khakas-Sagays and Shors (up to 40 %) (Ilumäe et al., 2016) (Y-base).

    samoyedic
    Map of distribution of Samoyedic languages (red) in the XVII century (approximate; hatching) and in the end of XX century (continuous background). Modified from Wikipedia, with the Tuva region labelled.

    Within the pan-Eurasian haplogroup R1a1a, two large genetic lines (sub-haplogroups) are identified: “European” (marker M458) and “Asian” (marker Z93) the latter almost never occurring in Europe (Balanovsky, 2015) but abundant in South Siberia and northern Hindustan. In the Altai-Sayan region, high frequencies of the “Asian” branch are spread in many peoples – Shors, Tubalars, Altai-Kizhi people, Telengits, Sagays, Kyzyl Khakas, Koibals, Teleuts (Y-base) (Kharkov et al., 2009). Hg R1a1a comprises perceptible parts of gene pools of Tuvinian clans (19 % in Mongush, and 15 % in Oorzhak), though its occurrence in Mongols is much lower (6 %). Those results also count in favor of the hypothesis of autochtonous component dominance even in the gene pools of clans potentially most influenced by Mongolian ancestry. If we add R1a1a variants to the “North Eurasian” haplogroups, the “not-Central Asian” component will compose average four fifth of the entire gene pools for Tuvinian clans (in Mongush 77 %, and in Oorzhak 81 %), being only 16 % in Mongols. Such data are definitely contrary to the hypothesis of a crucial influence of the Mongol expansion upon the development of Tuvinian gene pool.

    I found interesting the high proportion of R1a-Z93 subclades among Sagays in Khakhasia, which stem from a local Samoyed substratum, as described by the paper…

    Featured Image: Map of Uralic and Altaic languages, from Wikipedia.

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