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

pskov-novgorod-russia

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

Interesting excerpts:

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

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

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

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

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

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

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

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

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

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

A really hard choice between equally plausible models.

Related

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

Neolithic and Bronze Age Anatolia, Urals, Fennoscandia, Italy, and Hungary (ISBA 8, 20th Sep)

jena-isba8

I will post information on ISBA 8 sesions today as I see them on Twitter (see programme in PDF, and sessions from yesterday).

Official abstracts are listed first (emphasis mine), then reports and images and/or link to tweets. Here is the list for quick access:

Russian colonization in Yakutia

Exploring the genomic impact of colonization in north-eastern Siberia, by Seguin-Orlando et al.

Yakutia is the coldest region in the northern hemisphere, with winter record temperatures below minus 70°C. The ability of Yakut people to adapt both culturally and biologically to extremely cold temperatures has been key to their subsistence. They are believed to descend from an ancestral population, which left its original homeland in the Lake Baykal area following the Mongol expansion between the 13th and 15th centuries AD. They originally developed a semi-nomadic lifestyle, based on horse and cattle breeding, providing transportation, primary clothing material, meat, and milk. The early colonization by Russians in the first half of the 17th century AD, and their further expansion, have massively impacted indigenous populations. It led not only to massive epidemiological outbreaks, but also to an important dietary shift increasingly relying on carbohydrate-rich resources, and a profound lifestyle transition with the gradual conversion from Shamanism to Christianity and the establishment of new marriage customs. Leveraging an exceptional archaeological collection of more than a hundred of bodies excavated by MAFSO (Mission Archéologique Française en Sibérie Orientale) over the last 15 years and naturally kept frozen by the extreme cold temperatures of Yakutia, we have started to characterize the (epi)genome of indigenous individuals who lived from the 16th to the 20th century AD. Current data include the genome sequence of approximately 50 individuals that lived prior to and after Russian contact, at a coverage from 2 to 40 fold. Combined with data from archaeology and physical anthropology, as well as microbial DNA preserved in the specimens, our unique dataset is aimed at assessing the biological consequences of the social and biological changes undergone by the Yakut people following their neolithisation by Russian colons.

NOTE: For another interesting study on Yakutian tribes, see Relationships between clans and genetic kin explain cultural similarities over vast distances.

Ancient DNA from a Medieval trading centre in Northern Finland

Using ancient DNA to identify the ancestry of individuals from a Medieval trading centre in Northern Finland, by Simoes et al.

Analyzing genomic information from archaeological human remains has proved to be a powerful approach to understand human history. For the archaeological site of Ii Hamina, ancient DNA can be used to infer the ancestries of individuals buried there. Situated approximately 30 km from Oulu, in Northern Finland, Ii Hamina was an important trade place since Medieval times. The historical context indicates that the site could have been a melting pot for different cultures and people of diversified genetic backgrounds. Archaeological and osteological evidence from different individuals suggest a rich diversity. For example, stable isotope analyses indicate that freshwater and marine fish was the dominant protein source for this population. However, one individual proved to be an outlier, with a diet containing relatively more terrestrial meat or vegetables. The variety of artefacts that was found associated with several human remains also points to potential differences in religious beliefs or social status. In this study, we aimed to investigate if such variation could be attributed to different genetic ancestries. Ten of the individuals buried in Ii Hamina’s churchyard, dating to between the 15th and 17th century AD, were screened for presence of authentic ancient DNA. We retrieved genome-wide data for six of the individuals and performed downstream analysis. Data authenticity was confirmed by DNA damage patterns and low estimates of mitochondrial contamination. The relatively recent age of these human remains allows for a direct comparison to modern populations. A combination of population genetics methods was undertaken to characterize their genetic structure, and identify potential familiar relationships. We found a high diversity of mitochondrial lineages at the site. In spite of the putatively distant origin of some of the artifacts, most individuals shared a higher affinity to the present-day Finnish or Late Settlement Finnish populations. Interestingly, different methods consistently suggested that the individual with outlier isotopic values had a different genetic origin, being more closely related to reindeer herding Saami. Here we show how data from different sources, such as stable isotopes, can be intersected with ancient DNA in order to get a more comprehensive understanding of the human past.

A closer look at the bottom left corner of the poster (the left columns are probably the new samples):

finland-medieval-admixture

Plant resources processed in HG pottery from the Upper Volga

Multiple criteria for the detection of plant resources processed in hunter-gatherer pottery vessels from the Upper Volga, Russia, by Bondetti et al.

In Northern Eurasia, the Neolithic is marked by the adoption of pottery by hunter-gatherer communities. The degree to which this is related to wider social and lifestyle changes is subject to ongoing debate and the focus of a new research programme. The use and function of early pottery by pre-agricultural societies during the 7th-5th millennia BC is of central interest to this debate. Organic residue analysis provides important information about pottery use. This approach relies on the identification and isotopic characteristics of lipid biomarkers, absorbed into the pores of the ceramic or charred deposits adhering to pottery vessel surfaces, using a combined methodology, namely GC-MS, GC-c-IRMS and EA-IRMS. However, while animal products (e.g., marine, freshwater, ruminant, porcine) have the benefit of being lipid-rich and well-characterised at the molecular and isotopic level, the identification of plant resources still suffers from a lack of specific criteria for identification. In huntergatherer contexts this problem is exacerbated by the wide range of wild, foraged plant resources that may have been potentially exploited. Here we evaluate approaches for the characterisation of terrestrial plant food in pottery through the study of pottery assemblages from Zamostje 2 and Sakhtysh 2a, two hunter-gatherer settlements located in the Upper Volga region of Russia.

GC-MS analysis of the lipids, extracted from the ceramics and charred residues by acidified methanol, suggests that pottery use was primarily oriented towards terrestrial and aquatic animal products. However, while many of the Early Neolithic vessels contain lipids distinctive of freshwater resources, triterpenoids are also present in high abundance suggesting mixing with plant products. When considering the isotopic criteria, we suggest that plants were a major commodity processed in pottery at this time. This is supported by the microscopic identification of Viburnum (Viburnum Opulus L.) berries in the charred deposits on several vessels from Zamostje.

The study of Upper Volga pottery demonstrated the importance of using a multidisciplinary approach to determine the presence of plant resources in vessels. Furthermore, this informs the selection of samples, often subject to freshwater reservoir effects, for 14C dating.

Studies on hunter-gatherer pottery – appearing in eastern Europe before Middle Eastern Neolithic pottery – may be important to understand the arrival of R1a-M17 lineages to the region before ca. 7000 BC. Or not, right now it is not very clear what happened with R1b-P297 and R1a-M17, and with WHG—EHG—ANE ancestry

Bronze Age population dynamics and the rise of dairy pastoralism on the eastern Eurasian steppe

Bronze Age population dynamics and the rise of dairy pastoralism on the eastern Eurasian steppe, by Warinner et al.

Recent paleogenomic studies have shown that migrations of Western steppe herders (WSH), beginning in the Eneolithic (ca. 3300-2700 BCE), profoundly transformed the genes and cultures of Europe and Central Asia. Compared to Europe, the eastern extent of this WSH expansion is not well defined. Here we present genomic and proteomic data from 22 directly dated Bronze Age khirigsuur burials from Khövsgöl, Mongolia (ca. 1380-975 BCE). Only one individual showed evidence of WSH ancestry, despite the presence of WSH populations in the nearby Altai-Sayan region for more than a millennium. At the same time, LCMS/ MS analysis of dental calculus provides direct protein evidence of milk consumption from Western domesticated livestock in 7 of 9 individuals. Our results show that dairy pastoralism was adopted by Bronze Age Mongolians despite minimal genetic exchange with Western steppe herders.

Detail of the images:

mongol-bronze-age-pca

mongol-bronze-age-f4-ancestry

North Asian mitogenomes hint at the arrival of pastoralists from West to East ca. 2800-1000 BC

north-asia-mitogenomes

Open access Investigating Holocene human population history in North Asia using ancient mitogenomes, by Kılınç et al., Scientific Reports (2018) 8: 8969.

Abstract (emphasis mine):

Archaeogenomic studies have largely elucidated human population history in West Eurasia during the Stone Age. However, despite being a broad geographical region of significant cultural and linguistic diversity, little is known about the population history in North Asia. We present complete mitochondrial genome sequences together with stable isotope data for 41 serially sampled ancient individuals from North Asia, dated between c.13,790 BP and c.1,380 BP extending from the Palaeolithic to the Iron Age. Analyses of mitochondrial DNA sequences and haplogroup data of these individuals revealed the highest genetic affinity to present-day North Asian populations of the same geographical region suggesting a possible long-term maternal genetic continuity in the region. We observed a decrease in genetic diversity over time and a reduction of maternal effective population size (Ne) approximately seven thousand years before present. Coalescent simulations were consistent with genetic continuity between present day individuals and individuals dating to 7,000 BP, 4,800 BP or 3,000 BP. Meanwhile, genetic differences observed between 7,000 BP and 3,000 BP as well as between 4,800 BP and 3,000 BP were inconsistent with genetic drift alone, suggesting gene flow into the region from distant gene pools or structure within the population. These results indicate that despite some level of continuity between ancient groups and present-day populations, the region exhibits a complex demographic history during the Holocene.

north-asians-mtdna-haplogroup-frequency
Relationship between ancient North Asians and other populations based on haplogroup frequencies. Ancient North Asians as a single group (SIB, n = 41) and as divided into three different regional groups including Cis-Baikal (CISB, n = 23), Trans-Baikal (TRAB, n = 7) and Yakutia (YAK, n = 9) or as divided into three temporal groups including Early (7,000 BP, n = 11), Middle (4800 BP, n = 16) and Late (3000 BP, n = 11). Two individuals from Krasnoyarsk and Blagoveshensk are not included in regional groups due to their distinct geographical locations. (a) Barplot showing haplogroup frequencies on a dataset of 1,780 individuals. PCA plot based on haplogroup frequencies calculated using (b) 291 individuals with full mitochondrial sequences. Ancient North Asians are included as a single population. (c) 1,780 individuals. Ancient North Asians are included as three different regional groups in the analysis. See also Supplementary Tables S1, S4–S12 and Fig. S3a and b in Supplementary Information.

Interesting excerpts:

Although highly dependent on sample size and thus prone to generalization, haplotype sharing analysis between three spatial groups and other modern and ancient populations (Supplementary Table S15) revealed that the TRAB group shared most lineages with ancient Kazakh Altai (KA) and modern Nganasan (NGN)39,40,41,42. The CISB group shared most lineages with Tubalar39,42, KA43 and Early Bronze Age groups of Russia (BO)12, which might reflect the Siberian roots of BO, consistent with MDS based on Fst (Fig. 3b). The YAK group shared most lineages with the CISB, BO and Tubalar groups. These results showed that despite being from different sides of the Lake Baikal, the CISB and YAK groups shared most lineages with the Tubalar and also both of them were to a certain degree affiliated to the BO of the Cis-Baikal region, thus, reflecting a shared common ancestry. Furthermore, the CISB and YAK groups share lineages supporting the hypothesis of a lasting continuity in this large geographical territory. However, the TRAB group may have different legacy with affinities to ancient Kazakh Altai and modern Nganasan groups (that, actually, may have relocated from the Trans-Baikal region in times post-dating our sample).

north-asian-mtdna-plot
Relationship between ancient North Asians and other ancient and present-day populations based on Slatkin’s linearized pairwise FST. MDS plot based on Slatkin’s linearized pairwise FST calculated using (a) full mitochondrial DNA sequences. (b) HVRI sequences. See also Fig. S3c and d in Supplementary Information, Supplementary Tables S13–S15.

Two findings, however, were intriguing. One was the discovery of only weak support for a single regional population in comparisons between Early vs. Late as well as Middle vs. Late groups in the region. This may be explained by population structure, as the Late group comprised geographically very distant individuals, such as individuals from Krasnoyarsk Krai and Amur Oblast, not represented in the other diachronic groups (Table S9). Another explanation for rejecting the null hypothesis of continuity between the Middle and Late (4,800–3,000 BP) groups might be due to an interruption and the arrival of pastoralists at the beginning of the Iron Age between 3,670 to 2,760  BP as suggested by the archaeological record32. Thus, the introduction of the new lifeways, technologies and material culture expressions might also here be associated to an increased mobility into the area.

The second point was the estimated reduction in maternal effective population size and haplotype diversity around 7,000 BP. Intriguingly, climate modelling and radiocarbon dating studies53 suggest that climatic change and a collapse of the riverine ecosystems might have affected the human populations in Cis Baikal between 7,000–6,000 BP in line with our results. This finding was further supported by archaeological studies pointing to a possible hiatus38,54,55.

Although our results provide a first glimpse into population structure and diversity in North Asia during the Holocene which link to trend in the archaeological record, complete genome sequences will provide a higher resolution of more complex demographic events in the region.

Yet another hint at the west-east (and not east-west) population movement in Eurasia after the Corded Ware and Yamna expansions, without any significant change in the other direction until the Iron Age (as we know from Fennoscandian samples), which leaves still less space to propose incoming Uralic-speaking groups from Asia…

Related:

Genomic history of Northern Eurasians includes East-West and North-South gradients

russia-uralic-ibd

Open Access article on modern populations (including ancient samples), Between Lake Baikal and the Baltic Sea: genomic history of the gateway to Europe, by Triska et al., BMC Genetics 18(Suppl 1):110, 2017.

Abstract:

Background
The history of human populations occupying the plains and mountain ridges separating Europe from Asia has been eventful, as these natural obstacles were crossed westward by multiple waves of Turkic and Uralic-speaking migrants as well as eastward by Europeans. Unfortunately, the material records of history of this region are not dense enough to reconstruct details of population history. These considerations stimulate growing interest to obtain a genetic picture of the demographic history of migrations and admixture in Northern Eurasia.

Results
We genotyped and analyzed 1076 individuals from 30 populations with geographical coverage spanning from Baltic Sea to Baikal Lake. Our dense sampling allowed us to describe in detail the population structure, provide insight into genomic history of numerous European and Asian populations, and significantly increase quantity of genetic data available for modern populations in region of North Eurasia. Our study doubles the amount of genome-wide profiles available for this region.

We detected unusually high amount of shared identical-by-descent (IBD) genomic segments between several Siberian populations, such as Khanty and Ket, providing evidence of genetic relatedness across vast geographic distances and between speakers of different language families. Additionally, we observed excessive IBD sharing between Khanty and Bashkir, a group of Turkic speakers from Southern Urals region. While adding some weight to the “Finno-Ugric” origin of Bashkir, our studies highlighted that the Bashkir genepool lacks the main “core”, being a multi-layered amalgamation of Turkic, Ugric, Finnish and Indo-European contributions, which points at intricacy of genetic interface between Turkic and Uralic populations. Comparison of the genetic structure of Siberian ethnicities and the geography of the region they inhabit point at existence of the “Great Siberian Vortex” directing genetic exchanges in populations across the Siberian part of Asia.

EEHG-CHG-Neolithic_Farmer-ANE
f3 values to estimate (a) Eastern European Hunter-Gatherer, b Neolithic Farmer, c Caucasus hunter-gatherer, and d) Mal’ta (Ancient North Eurasian) ancestry in modern humans

Slavic speakers of Eastern Europe are, in general, very similar in their genetic composition. Ukrainians, Belarusians and Russians have almost identical proportions of Caucasus and Northern European components and have virtually no Asian influence. We capitalized on wide geographic span of our sampling to address intriguing question about the place of origin of Russian Starovers, an enigmatic Eastern Orthodox Old Believers religious group relocated to Siberia in seventeenth century. A comparative reAdmix analysis, complemented by IBD sharing, placed their roots in the region of the Northern European Plain, occupied by North Russians and Finno-Ugric Komi and Karelian people. Russians from Novosibirsk and Russian Starover exhibit ancestral proportions close to that of European Eastern Slavs, however, they also include between five to 10 % of Central Siberian ancestry, not present at this level in their European counterparts.

russian-uralic-turkicadmixture
Admixture proportions in studied populations, K = 6. Populations from the Extended dataset. Abbreviated population codes: NSK – Russians from Novosibirsk; STV -Starover Russians; ARK: Bashkirs from Arkhangelskiy district; BRZ – Bashkirs from Burzyansky district

Conclusions
Our project has patched the hole in the genetic map of Eurasia: we demonstrated complexity of genetic structure of Northern Eurasians, existence of East-West and North-South genetic gradients, and assessed different inputs of ancient populations into modern populations.

Featured image, from the article: “Departures from the expected IBD. Shown populations exceed the expected IBD sharing by more than two standard deviations.”

Related:

Königsberg (AKA Kaliningrad) under international law: Russian, German, Polish, Lithuanian, or simply Prussian?

The progress of the ‘star wars’ (AKA missile shield) affair, which Russia seemed willing to aggravate by talking about plans to station missiles in Kaliningrad, without any concerns whatsoever for the welfare of Kaliningraders and Europeans, should make the European Union reexamine its current policy under the Kaliningrad Strategy, of collaborating with Russia by facilitating the transit of goods and persons and helping its socio-economic development.

Instead of just hearing what Russians have to claim before the international community, the EU should ask the international community by which right keeps the Russian Federation hold on Königsberg territory, and should demand from Russia a date for devolution, no matter how hard Russian media propaganda tries to avoid the question:

Although disputes over the status of Russia’s westernmost exclave of Kaliningrad have practically ceased, this should be regarded as a signal that all the parties concerned are aware of the serious repercussions that instability in that region could cause.

Geopolitical Stability has been by far the most repeated pro-Russian argument since the 90’s, also in official European Union forums (see Freedom to Kaliningrad thread); it is easily summed up into a “let’s maintain the statu quo to avoid destabilizing the region”. The murmuring of those plans to use Kaliningrad as missile base made by Russian military officials to the press, to escalate tensions in the missile shield affair, has shown how the Russian Federation respects the will of Europeans for stability in the region. Not to talk about Russia’s lack of respect for the lives of thousands of European citizens in this winter’s gas disputes, or its lack of respect for Estonian democratic decisions, or its support for the authoritarian Belarusian regime of Lukashenko

Other great arguments made by pro-Russians include “Nazi Germany”, “World War II” and “Mother Russia”, and are easily read elsewhere in Russian media and blogs when the Kaliningrad question is mentioned. Nevertheless, most Kaliningraders – whether ethnic Russians or not – show often an open mind about the return options. And even official Russian media like Russia Today recognize still in 2009 (only in English texts for outsiders) the Lithuanian claims to the territory and its return; East German rights are still taboo in Russian ‘free’ media, while Polish claims are probably too weak to be worth mentioning:

The region became an administrative unit of Russia [sic] in 1946 after the Potsdam conference and the partition of Germany. Although it solidified as an administrative entity, with the collapse of the Soviet Union, the issue of reassimilating the Kaliningrad region into its historic entity of Lithuania arose.

According to a thorough study on the question (The Kaliningrad Challenge, 2003) Russia has been always concerned about the risk of separatism in Kaliningrad, which might be greater than expected if the European Report The EU and Kaliningrad (2002) is correct in assessing that Kaliningrad’s level of development is closer to Lithuania and Latvia than previously thought. In that sense, ethnic Russian Kaliningraders see Kaliningrad in the future as another Baltic Republic, either still somehow federated to Russia with great autonomy or fully independent. Moreover,

There are opinion polls – now more frequently held within blogs and forums – which show that Kaliningraders occasionally imagine their future not so much as a fourth Baltic Republic, but as part of a return to Germany

As it has been already argued on the situation of Königsberg/Kaliningrad region and the Northern Territories/Southern Kuril Islands under international law:

In a similar way, the Soviets also refused to discuss the final peace settlement in Europe after the Second World War. It is important to emphasize that neither the United States nor Britain agreed at Potsdam or anywhere else to the transfer of East Prussia or part of the Königsberg Region to the Soviet Union. Thus, although the Kaliningrad Region is currently administered by Russia, it is not a legal part of Russia.

Stalin was seeking a deal on East Prussia at the Tehran conference in 1943, drawing a line in red pencil on the map “to illustrate the fact that, if part of eastern Prussia, including the ports of Könisberg and Tilsit, were given to the Soviet Union, he would be prepared to accept the Curzon line […] as the frontier between the Soviet Union and Poland.”

This line goes roughly along the current border between the Kaliningrad Region and Poland, but Stalin’s red line on the map went virtually through the cities of Königsberg and Insterburg (see the Map). Charles E. Bolen, the interpreter for the American delegation, says in his memoirs that during their discussion, Stalin and Churchill virtually agreed on the future borders of Poland, but the official American record of the conversation says that “although nothing was stated, it was apparent that the British were going to take this suggestion back to London to the Poles.”

On February 11, 1945, at the Crimea (Yalta) Conference, the Big Three agreed on the Curzon Line as the boundary between Poland and the USSR. However, the archival material clearly shows that there had not been any legally binding agreement made between the allies about the transfer of the Königsberg Region to the Soviet Union at any of the Second World War conferences. This is why Stalin attempted to secure his gains at the Potsdam conference in Berlin, which took place from July 17 to August 2, 1945.

After the end of the Second World War, the Kaliningrad question began by Stalin’s personal will of revenge against Germany:

Königsberg was neither appended outright to the Soviet Union nor was it to be considered part of the Soviet zone of occupation, which had been outlined earlier in the agreement.

[The Soviet Union] acted decisively to completely eradicate the German presence in Königsberg and replace it with Soviet presence. This began even before the end of hostilities with the Reich:

Königsberg was destroyed in the last weeks of the war when there was no real reason to assault it. When the soldiers of the Byelorussian front were dying in its streets in the first week of April, 1945, the rest of the Red Army was already besieging Berlin. Seven centuries of history went up in smoke in one week of shelling and bombing. By then, the decision to annihilate East Prussia and grant Königsberg to the Soviet Union had already been taken, so the reason for its destruction remains a mystery. Did Stalin take the decision in a fit of war revenge? Did he think that the setting of an ancient bourgeois city would hamper the development of the new Soviet city he wanted to build in its place? Or did he fear that, unless turned into a pile of ruins, Königsberg might not be conceded to him by the Allies after all? Pictures and models in the bunker-cum-museum where the capitulation of the city was signed are revealing. Most of the destruction was done after-wards, when the victors took to the task of building a new city on the ruins of the old…

While the destruction of the city’s infrastructure was underway, an equally brutal purge of its population through gang rapes and indiscriminate crimes was carried out:

The demography of that part of Lithuania Minor which is under direct Soviet administration, the “Oblast,” has changed in the most radical way in all its history. The original population of the area — German as well as Lithuanian — has disappeared completely. Many had fled before the Soviet armed forces invaded the area in 1945; those who remained — several hundred thousand — either perished from hunger or disease or were deported to Siberia; the others were expelled to Germany in 1949. They all — about 1,200,000 before World War II — were replaced by about 600,000 settlers from the northern and central parts of Russia. The administration and economy of the “Oblast” has been reorganized to conform with Soviet models and practices. It has been fortified to serve the strategic aims of the Soviet Union.

Modern Claims in Europe

After the fall of the Soviet Union, there were 4 main alternatives for the future of Kaliningrad, following Raymond A. Smith’s article The Status of The Kaliningrad Oblast Under International Law (1992), which argues in favour of the Lithuanian claim, but which also addresses some historical and political questions:

From the historical [point of view] sovereignty over the territory of the Kaliningrad Oblast passed over the course of centuries from the the indigenous Old Prussian population, to the Teutonic Order, to the Kingdom of Poland, to the Kingdom of Prussia (later the German Empire) and finally, perhaps, to the USSR/RSFSR. It is not surprising, then, to find that each of these entities (with the exception, of course, of the Teutonic Order) has a conceivable claim to this territory. This section examines the legal basis, or lack thereof, of the actual or potential claim of each entity, as well as the potential claim of the indigenous population.

  • The German Claim: Some Germans challenge the validity of both the Final Settlement and the original “dismemberment” of the German Reich.
    Their arguments are complex but can be reduced in essence to two claims:

    1. the Allies had no power to allow German territory to be annexed by other countries
    2. the West Germany and even the modern Federal Republic of Germany are not coextensive with the German Reich and are therefore not competent to speak for it in its entirety

    The first proposition is supported by numerous charges: that the guarantees of self-determination in the Atlantic Charter, the UN Charter, and the Vienna Convention on the Law of Treaties were ignored; that the Ancient Roman principle of ex injuria non oritur jus prohibits punishing Germany by unilateral confiscation of its territory; that the powers of the Allies as occupiers were strictly curtailed by the Hague Laws of War of 1907; that use of German lands as “compensation” to Poland for lands lost to the Soviet Union has no basis in international law; and many others.

  • The Russian Claim: As the historical overview recounted, the working premise of the Potsdam Conference was that the Soviets would receive the Oblast at the final peace conference. The Allies specifically committed themselves to supporting the Soviet claim in the Final Settlement, but when that settlement was finally signed in 1990, specific title was not transferred. Why the Final Settlement did not include a specific statement of transfer is unclear. The seemingly most probable reason is that the transfer of Kaliningrad to the Soviet Union is considered a fait accompli and that the legal niceties of including a specific mention of transfer were outweighed by potential political embarassment such a mention might have caused the Kohl government. Such a position assumes that the tranfer has already taken place, an assertion which rests on shaky ground.

    Similarly, the Act of Military Surrender specifically indicates that the occupation itself did not effect the annexation of Germany. Thus, although Germany surrendered unconditionally, none of its territories were automatically annexed to any other state. Such annexation would have to be made explicit in a legally binding document. Only “administration” was established by the Potsdam Agreement, however, and “administration” is definitely not the same as “annexation” under international law.

    Rather than present arguments based on international law, Stalin advanced the law of revenge. ‘The Russians had suffered so much and lost so much blood, they were anxious to have some small satisfaction to [sic] tens of millions of their inhabitants who had suffered in the war,” Stalin said at Potsdam.

    In the absence of ethnic and historical claims to shore up their questionable legal claim, then, the only argument which the Soviet Union can depend upon is the principle of prescriptive claim. This principle transfers title to land when a country has held it for a long period of time without protest by the land’s original owners or by the international community at large. No specific time frame is suggested for the acquisition of prescriptive claim. Grotius suggested 100 years, a figure which the Permanent Court of International Justice endorsed in 1933. The International Court of Justice, on the other hand, said that fifty years had been long enough for a boundary between Venezuela and British Guiana to have legal effect.

  • The Polish Claim: Poland has no ethnic claim to the Oblast. Although the southern half of East Prussia was occupied mainly by Polish Masurians, they had almost no presence in the northern part.

    Poland’s historic claim is only marginally stronger. For some two centuries, Prussia was a fief of the Polish King, but during that period the area remained firmly under German control. In any case, title was decisively transferred by the Treaty of Wehlau in 1657. During World War II many Poles operated under the belief that all of East Prussia would become theirs, but they were never legally promised the territory in its entirety.

  • Lithuanian Claim: The claim of the Lithuanian state could rely upon both ethnic and historical grounds.
    1. The Lithuanians may argue that
      the first peoples to hold sovereignty over the region were ethnic Lithuanians and closely related Old Prussians, and
    2. the pre-1945 population outside the cities of the Oblast was largely of Lithuanian origin. If the status of the Oblast were to be altered in the future, then, the Lithuanian state could have a strong argument for assimilating this remainder of Lithuania Minor.

    The idea of unifying the Oblast with the rest of Lithuania has strong historical precedents. Lithuanian assemblies met in Chicago and New York in 1914, The Hague in 1916 and Berne in 1917 to demand an independent Lithuania including all of Lithuania Minor. An assembly in Vilnius in 1917 restated the problem to define the new Lithuania within its “ethnographic borders,” a concept endorsed by a later assembly in Voronezh the same year.
    Finally, on November 30, 1918, the National Council of Prussian Lithuania issued the Declaration of Tilsit:

    Taking into account that everything that exists has a right to continue existing and that we, Lithuanians who live here in Prussian Lithuania, are the majority of the population of this land, we demand, on the basis of Wilson’s right of national self-determination, that Lithuania Minor be joined to Lithuania Major

    The clearest catch here is that any annexation of the Oblast by Lithuania might hinge upon the democratic decision of an indigenous Lithuanian majority to authorize such an annexation. And, as we have seen, virtually none of the indigenous Lithuanian population remains in the Oblast, having fled or been killed or exiled after World War II. This raises the final claim to be discussed — that of the indigenous population.

  • The Claim of the Native Population: The right to national self-determination is one of the main cornerstones of the contemporary international legal order. Eight of Wilson’s Fourteen Points refer to such concerns. The Atlantic Charter’s third and fourth principles call for self-determination in matters of both boundaries and choice of government. The Charter of the United Nations calls for colonial powers to foster self-determination in “non-self governing territories”. That right might be interpreted as concerning:
    1. The Oblast’s postwar ethnic Russian settlers – as opposed to central Soviet or Russian authorities.
    2. the traditional population which was decimated or expelled en masse after World War II, which is defended on the grounds that forcible deportations of native populations is clearly in violation of international law – native Königsbergers expelled after World War II, then, have a right under international law to choose to return to their native land.

    On that question, there is the precedent of United Nations action regarding the settlement of Gibraltar:

    As in the case of the Oblast, the key issue was whether the rightful native population of the Rock should be considered to be the contemporary residents or an earlier population who had been compelled to depart in 1704. The British argued that over the centuries since 1704 a permanent and authentic population had been developed on the Rock, which now had the right to determine their own fate. The Spanish countered that the post-1704 population were “pseudo-Gibraltarians” and that the rightful rulers of Gibraltar Rock were the descendants of Spaniards who had resettled, for the most part, in the nearby city of San Roque.

    Under pressure from the United Nations to end its colonial occupation of Gibraltar and in an attempt to settle the status of the Rock once and for all, the British government conducted a plebiscite in 1967. The choices were stark — full political affiliation with either Great Britain or with Spain — and the result was unequivocal: 12,138 to 44 in favor of Great Britain. Nonetheless, the U.N. General Assembly once again condemned British occupation of Gibraltar, this time in the strongest language yet. It, in essence, declared the plebiscite null, accused the British of resisting decolonization, and called once again for immediate negotiations between Great Britain and Spain for a transfer of sovereignty.

    Whatever the merits of the Gibraltar case, the precedent for the Oblast is clear. If the rights of native populations can stretch back to 1704, then surely the postwar expellees from the Oblast would have an unambiguous right to return to their homeland and choose its political fate — be that choice in-dependence or association with another state. The current population of the Oblast would presumably have no say in the territory’s political future.

    The key difference between Gibraltar and the Oblast is that in the former case, there actually is a population in San Roque able and willing to resettle the Rock. No analagous “population-in-exile” exists in the case of the Oblast. Rather, much of the population of Königsberg was killed or died in exile. Those who were deported to Germany (and their descendants) in all likelihood now enjoy a standard of living which is, at least quantitatively, many times better than any which would be possible in the backward conditions of the Oblast. Further, most — although far from all — Germans seem to have accepted the loss of the prewar lands; the idea of reclaiming part of East Prussia would not necessarily resonate with much of the population. It seems extremely unlikely, then, that more than a handful of such native German Königsbergers would wish to uproot and resettle in the Oblast.

Even with German and Lithuanian strong claims about the Soviet colony of Königsberg opposing the legality of Stalin’s annexation, Russia did in the 90’s what it was used to in such cases when the Soviet Union was still a Great Power: they took the easy way, and annexed the territory to Russia, expecting the international community to accept it. Which is nice, because the EU as a Great Power will therefore be entitled to follow the same principle in the future…

In my personal opinion, the European Union faces today 3 alternatives, given Russia’s will to retain Stalin’s European exclave no matter how illegal or illegitimate it is from an international point of view:

  1. Support modern Kaliningraders in their demands of greater autonomy within the Russian Federation – and maybe a future separation from it -, which is the fairest position under modern international law, which demands non-belligerant positions (against Russia in this case) and respect for human rights – Russian settlers and their families. This is certainly the option of most Kaliningraders of Russian ethnicity, as well as most EU-politicians.
  2. Support Germany’s or Lithuania’s claims (or both), seeking to integrate Kaliningrad within the European Union, maybe as a sort of a Baltic territory co-administered by both Germany and Lithuania, financing the return of (families of) expellees to Königsberg, and the return of (willing) families of Russian settlers to Russia. This is the option preferred by many Germans and (I guess) most Lithuanians.
  3. Support the creation of a modern Baltic Prussian State (Prusa), which could help unite the Pro-Baltic (and Pro-European) attitude of Russian Kaliningraders, the will of native peoples and their families to return to East Prussia, as well as claims of EU member states to integrate Königsberg in Europe, by embracing Old Prussian history of the territory and its peoples. Modern organizations supporting the revival of the Old Prussian language would probably support its revitalization in Königsberg include the future Research Institute of Prussology and the Prussian language organization in Poland.

The third is my preferred option, not because I am some kind of language revival freak (what I possibly am, given that I also support Old Prussian language revival), but because what many (want to) regard simply as ethnic German and ethnic Lithuanian inhabitants of East Prussia in 1945 were in fact descendants of Old Prussians who had lost their language in favour of either German or Lithuanian languages, depending on the territories they dwelled when they ceased to speak Prussian. Given that historical, cultural and linguistic background of the Königsberg (or East Prussian) territory, the European Union should take action supporting the return of those expelled peoples and their families to their ancient territory, which they were forced to leave half a century ago.

There is therefore no need to support the adscription of East Prussia to modern countries or peoples, be it Russia, Germany, Poland or Lithuania. And the only alternative to modern peoples, cultures and states is to support a linguistic and cultural revival of a Prussian people and language that should have never disappeared.