Demographic history and genetic adaptation in the Himalayan region

Open access Demographic history and genetic adaptation in the Himalayan region inferred from genome-wide SNP genotypes of 49 populations, by Arciero et al. Mol. Biol. Evol (2018), accepted manuscript (msy094).

Abstract (emphasis mine):

We genotyped 738 individuals belonging to 49 populations from Nepal, Bhutan, North India or Tibet at over 500,000 SNPs, and analysed the genotypes in the context of available worldwide population data in order to investigate the demographic history of the region and the genetic adaptations to the harsh environment. The Himalayan populations resembled other South and East Asians, but in addition displayed their own specific ancestral component and showed strong population structure and genetic drift. We also found evidence for multiple admixture events involving Himalayan populations and South/East Asians between 200 and 2,000 years ago. In comparisons with available ancient genomes, the Himalayans, like other East and South Asian populations, showed similar genetic affinity to Eurasian hunter-gatherers (a 24,000-year-old Upper Palaeolithic Siberian), and the related Bronze Age Yamnaya. The high-altitude Himalayan populations all shared a specific ancestral component, suggesting that genetic adaptation to life at high altitude originated only once in this region and subsequently spread. Combining four approaches to identifying specific positively-selected loci, we confirmed that the strongest signals of high-altitude adaptation were located near the Endothelial PAS domain-containing protein 1 (EPAS1) and Egl-9 Family Hypoxia Inducible Factor 1 (EGLN1) loci, and discovered eight additional robust signals of high-altitude adaptation, five of which have strong biological functional links to such adaptation. In conclusion, the demographic history of Himalayan populations is complex, with strong local differentiation, reflecting both genetic and cultural factors; these populations also display evidence of multiple genetic adaptations to high-altitude environments.

himalayan-map
Population samples analysed in this study. A. Map of South and East Asia, highlighting the four regions examined, and the colour assigned to each. B. Samples from the Tibetan Plateau. C.Samples from Nepal. D. Samples from Bhutan and India. The circle areas are proportional to the sample sizes. The three letter population codes in B-D are defined in supplementary table S1.

Relevant excerpts:

Genetic affinity to ancestral populations

We explored the genetic affinity between the Himalayan populations and five ancient genomes using f3-outgroup statistics. Himalayans show greater affinity to Eurasian hunter-gatherers (MA-1, a 24,000- year-old Upper Palaeolithic Siberian), and the related Bronze Age Yamnaya, than to European farmers (5,500-4,800 years ago; Fig. 5A) or to European hunter-gatherers (La Braña, 7,000 years ago; Fig. 5B), like other South and East Asian populations. We further explored the affinity of Himalayan populations by comparing them with the 45,000-year-old Upper Palaeolithic hunter-gatherer (Ust’-Ishim) and each of MA-1, La Braña, or Yamnaya. Himalayan individuals cluster together with other East Asian populations and show equal distance from Ust’-Ishim and the other ancient genomes, probably because Ust’-Ishim belongs to a much earlier period of time (supplementary fig. S15). We also explored genetic affinity between modern Himalayan populations and five ancient Himalayans (3,150 1,250 years old) from Nepal. The ancient individuals cluster together with modern Himalayan populations in a worldwide PCA (supplementary fig. S16), and the f3-outgroup statistics show modern high-altitude populations have the closest affinity with these ancient Himalayans, suggesting that these ancient individuals could represent a proxy for the first populations residing in the region (supplementary fig. S17 and supplementary table S4). Finally, we explored the genetic affinity of Himalayan samples with the archaic genomes of Denisovans and Neanderthals (Skoglund and Jakobsson 2011), and found that they show a similar sharing pattern with Denisovans and Neanderthals to the other South and East Asian populations. Individuals belonging to four Nepalese, one Cambodian, and three Chinese populations show the highest Denisovan sharing (after populations from Australia and Papua New Guinea) but these values are not significantly greater than other South and East Asian populations (supplementary figs. S18 and S19).

himalayan-pca
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. B. PCA of the Himalayan populations alone. Each dot represents a sample, coded by country or region as indicated. Most samples lie on an arc between Bhutanese and Nepalese samples; Toto (India) are seen as extreme outlier in the bottom left corner, while Dhimal (Nepal) and Bodo (India) also form outliers.

NOTE. The variance explained in the PCA graphics seems to be too high. This happened recently also with the Damgaard et al. (2018) papers (see here the comment by Iosif Lazaridis).

Similarities and differences between high-altitude Himalayan

The most striking example is provided by the Toto from North India, an isolated tribal group with the lowest genetic diversity of the Himalayan populations examined here, indicated by the smallest long-term Ne (supplementary fig. S5), and a reported census size of 321 in 1951 (Mitra 1951), although their numbers have subsequently increased. Despite this extreme substructure, shared common ancestry among the high-altitude populations (Fig. 2C and Fig. 3) can be detected, and the Nepalese in general are distinguished from the Bhutanese and Tibetans (Fig. 2C) and they also cluster separately (Fig. 3). In a worldwide context, they share an ancestral component with South Asians (supplementary fig. S2). On the other hand, the Tibetans do not show detectable population substructure, probably due to a much more recent split in comparison with the other populations (Fig. 2C and supplementary fig. S6). The genetic similarity between the high-altitude populations, including Tibetans, Sherpa and Bhutanese, is also supported by their clustering together on the phylogenetic tree, the PCA generated from the co-ancestry matrix generated by fineSTRUCTURE (supplementary fig. S10 and S11), the lack of statistical significance for most of the D-statistics tests (Yoruba, Han; high-altitude Himalayan 1, high-altitude Himalayan 2), and the absence of correlation between the increased genetic affinity to lowland East Asians and the spatial location of the Himalayan populations (supplementary figs. S12 and S13). Together, these results suggest the presence of a single ancestral population carrying advantageous variants for high-altitude adaptation that separated from lowland East Asians, and then spread and diverged into different populations across the Himalayan region. (…)

Recent admixture events

himalayan-admixture
Genetic structure of the Himalayan region populations from analyses using unlinked SNPs. C. ADMIXTURE (K values of 2 to 6, as indicated) analysis of the Himalayan samples. Note that most increases in the value of K result in single population being distinguished. Population codes in C are defined in supplementary table S1.

Himalayan populations show signatures of recent admixture events, mainly with South and East Asian populations as well as within the Himalayan region itself. Newar and Lhasa show the oldest signature of admixture, dated to between 2,000 and 1,000 years ago. Majhi and Dhimal display signatures of admixture within the last 1,000 years. Chetri and Bodo show the most recent admixture events, between 500 and 200 years ago (Fig. 4, supplementary tables S3). The comparison between the genetic tree and the linguistic association of each Himalayan population highlights the agreement between genetic and linguistic sub-divisions, in particular in the Bhutanese and Tibetan populations. Nepalese populations show more variability, with genetic sub-clusters of populations belonging to different linguistic affiliations (Fig. 3B). Modern high-altitude Himalayans show genetic affinity with ancient genomes from the same region (supplementary fig. S17), providing additional support for the idea of an ancient high-altitude population that spread across the Himalayan region and subsequently diverged into several of the present-day populations. Furthermore, Himalayan populations show a similar pattern of allele sharing with Denisovans as other South-East Asian populations (supplementary fig. S18 and S19). Overall, geographical isolation, genetic drift, admixture with neighbouring populations and linguistic subdivision played important roles in shaping the genetic variability we see in the Himalayan region today.

Related:

Copenhagen group: Germanic and Balto-Slavic from Bell Beaker; Indo-Anatolian homeland in the Caucasus

Article of general knowledge in Der Spiegel, Invasion from the Steppe, with comments from Willerslev and Kristiansen, appeared roughly at the same time as the Damgaard et al. Nature (2018) and Science (2018) papers were published.

NOTE. You can read the article (in German) from Kristiansen’s Academia.edu account.

Excerpts translated from German (emphasis mine):

On the Y-DNA data

Particularly striking is the genetic signature from the steppe on the Y chromosome. From this the researchers conclude that the majority of migrants were males. Kristian Kristiansen, chief archaeologist in the Willerslev team, also has an idea of ​​how this could be explained: “Maybe it’s a rite of initiation, as it was spread among the steppe peoples,” he says.

The younger sons of the Yamnaya herders, who were excluded from the succession, had to seek their fortune on their own. As part of a solemn ritual, they threw themselves to wolves’ skins and then swarmed in warlike gangs to buy their own herds by cattle-stealing.

(…)

An ally that they seem to have brought from their homeland may also have contributed to the genetic success of the steppe people: Yersinia pestis, the plague bacterium. Its genes were found by researchers from the Max Planck Institute in Jena – and apparently it emerged exactly at the same time as the Yamnaya thrust began.

About the Hittites

(…) And yet now, where Asia and Europe meet geographically, there is no trace of the Yamnaya genes. The wander-loving people from the Pontic-Caspian steppe apparently found neither the way across the Balkans nor through the Caucasus mountains.

Now the researchers are puzzled: How can it be that a language goes on a walk, without the accompanying speakers coming along? Is it possible that the Indo-European seeped into Anatolia, much like the English language spread today without the need for Englishmen?

Archaeologist Kristiansen does not believe it. The researchers would find it hard to reconsider their theories, he says: “Especially the first chapter of the story has to be rewritten.”

He suspects that there was a predecessor of the Yamnaya culture, in which a kind of Proto-Proto-Indo-European was spoken. And he also has a suspicion, where this people could have drifted around: The Caucasus, says Kristiansen, was their homeland. But that remains unproven: “There’s another hole left,” he admits.

invasion-yamnaya-steppe
Spread of Indo-European languages

About the Botai

The study of [the Botai] genome revealed that it was genetically radically different from the members of the Yamnaya culture. The Botai, it seems, consistently avoided any contact with their neighbors – even though they must have crossed the territory of the Botai on their migratory waves.

Willerslev assumes that the art of keeping horses from the Yamnaya steppe nomads was adopted from these peoples, and then they developed it further. At some point, the Botai could then have itself become doomed by its groundbreaking innovation: While the descendants of the Yamnaya spread over half of Eurasia, the Botai disappeared without leaving a trace.

Even more interesting than the few words that set the Copenhagen group’s views for future papers (such as the expected Maykop samples with EHG ancestry) is the artistic sketch of the Indo-European migrations, probably advised by the group.

A simple map does not mean that all members of the Danish workgroup have changed their view completely, but I would say it is a great improvement over the previous “arrows of migration” (see here), and it is especially important that they show a more realistic picture of ancient migrations to general readers.

NOTE. Especially absurd is the identification of the ‘Celtic’ expansion with the first Bell Beakers in the British Isles (that idea is hold by few, such as Koch and Cunliffe in their “Celtic from the West” series). Also inexact, but not so worrying, are the identification of ‘Germanic’ in Germany/Únětice, or the spread of ‘Baltic’ and ‘Slavic’ directly to East Europe (i.e. I guess Mierzanowice/Nitra -> Trzciniec), which is probably driven by the need to assert a close connection with early Iranians and thus with their satemization trends.

Also, as we know now thanks to Narasimhan et al. (2018), there is no need to support that convoluted west arrow (representing CWC) from West Yamna to Central Europe, and then to East Yamna, since the Proto-Indo-Iranian community – represented by the Steppe MLBA cloud that later expanded Indo-Aryan and Iranian languages – has a more direct connection with the in situ admixture of Poltavka/Abashevo within the Volga-Ural region.

I think we can keep this from the article:

Their results, as well as those of the competition labs at Harvard University and Jena’s Max Planck Institute for the History of Humanity, leave no doubt: Yes, the legendary herdsmen in the Pontic-Caspian steppe really existed. They belonged to the so-called Yamnaya culture, and they spread, as linguists had predicted, in massive migrations towards Central Europe and India – a later triumph for linguists.

This can be added to a recent comment by de Barros Damgaard:

The project has been an extremely enriching and exciting process. We were able to direct many very different academic fields towards a single coherent approach. By asking the right questions, and keeping limitations of the data in mind, contextualizing, nuancing, and keeping dialogues open between scholars of radically different backgrounds and approaches, we have carved out a path for a new field of research. We have already seen too many papers come out in which models produced by geneticists working on their own have been accepted without vital input from other fields, and, at the other extreme, seen archaeologists opposing new studies built on archaeogenetic data, due to a lack of transparency between the fields.

Data on ancient DNA is astonishing for its ability to provide a fine-grained image of early human mobility, but it does stand on the shoulders of decades of work by scholars in other fields, from the time of excavation of human skeletons to interpreting the cultural, linguistic origins of the samples. This is how cold statistics are turned into history.

Related:

The Caucasus a genetic and cultural barrier; Yamna dominated by R1b-M269; Yamna settlers in Hungary cluster with Yamna

caucasus-europe

Open access The genetic prehistory of the Greater Caucasus, by Wang et al. bioRxiv (2018).

The Caucasus Mountains as a prehistoric barrier

I think the essential message we can extract from the paper is that the Caucasus was a long-lasting cultural and genetic barrier, although (obviously) it was not insurmontable.

Our results show that at the time of the eponymous grave mound of Maykop, the North Caucasus piedmont region was genetically connected to the south. Even without direct ancient DNA data from northern Mesopotamia, the new genetic evidence suggests an increased assimilation of Chalcolithic individuals from Iran, Anatolia and Armenia and those of the Eneolithic Caucasus during 6000-4000 calBCE23, and thus likely also intensified cultural connections. Within this sphere of interaction, it is possible that cultural influences and continuous subtle gene flow from the south formed the basis of Maykop.

caucasus-mountains-eneolithic
The zoomed map shows the location of sites in the Caucasus. The size of the circle reflects number of individuals that produced genome-wide data. The dashed line illustrates a hypothetical geographic border between genetically distinct Steppe and Caucasus clusters.

Also, unlike more recent times, the North Caucasian piedmont and foothill of the Caucasus region was more strongly connected to Northern Iran than to the steppe, at least until the Bronze Age.

(…) our data shows that the northern flanks were consistently linked to the Near East and had received multiple streams of gene flow from the south, as seen e.g. during the Maykop, Kura-Araxes and late phase of the North Caucasus culture.

Northern Caucasus dominated by R1b, southern Caucasus by J and G2

caucasus-y-dna
Comparison of Y-chromosome (A) 1123 and mitochondrial (B) haplogroup distribution in the Steppe and Caucasus cluster.

The first samples from the Eneolithic (one ca. 4300 BC?, the other ca. 4100 BC) are R1b1, without further subclades, so it is difficult to say if they were V88. On the PCA, they seem to be an important piece of the early Khvalynsk -> early Yamna transition period, since they cluster closer to (or even among) subsequent Yamna samples.

From 3000 BC onwards, all samples from the Northern Caucasus group of Yamna are R1b-M269, which right now is probably no surprise for anyone.

The Catacomb culture is dominated by R1b-Z2103, which agrees with what we saw in the unclassified Ukraine Eneolithic sample. However, the new samples (clustering close to Yamna, but with slightly ‘to the south’ of it) don’t seem to cluster closely to that first sample, so that one may still remain a real ‘outlier’, showing incoming influence (through exogamy) from the north.

If anyone was still wondering, no R1a in any of the samples, either. This, and the homogeneous R1b-Z2103 community in Catacomb (a culture in an intermediate region between Late Yamna to the West, and Poltavka to the East), together with Poltavka dominated by R1b-Z2103, too, should put an end to the idea that Steppe MLBA (Sintashta-Petrovka/Potapovka) somehow formed in the North Pontic steppe and appeared directly in the Volga-Ural region. A Uralic/Indo-Iranian community it is, then.

The admixed population from the Caucasus probably points to an isolated region of diverse peoples and languages even in this period, which justifies the strong differences among the historic language families attested in the Caucasus.

So, not much space for Anatolian migrating with those expected Maykop samples with EHG ancestry, unless exogamy is proposed as a source of language change.

PCA-caucasus
ADMIXTURE and PCA results, and chronological order of ancient Caucasus individuals. Samples from Hungary are surrounded by red circles (see below for ADMIXTURE data) (a) ADMIXTURE results (k=12) of the newly genotyped individuals (fillbred symbols with black outlines) sorted by genetic clusters (Steppe and Caucasus) and in chronological order (coloured bars indicate the relative archaeological dates, (b) white circles the mean calibrated radiocarbon date and the errors bars the 2-sigma range. (d) shows these projected onto a PCA of 84 modern-day West Eurasian populations (open symbols).

Yamna Hungary, and the previous Yamna “outliers”

Those western “Yamna outliers”, as I expected, were part of some late Khvalynsk/early Yamna groups that cluster “to the south” of eastern Yamna samples:

Another important observation is that all later individuals in the steppe region, starting with Yamnaya, deviate from the EHG-CHG admixture cline towards European populations in the West. This documents that these individuals had received Anatolian farmer-related ancestry, as documented by quantitative tests and recently also shown for two Yamnaya individuals from Ukraine (Ozera) and one from Bulgaria24. For the North Caucasus region, this genetic contribution could have occurred through immediate contact with groups in the Caucasus or further south. An alternative source, explaining the increase in WHG-related ancestry, would be contact with contemporaneous Chalcolithic/EBA farming groups at the western periphery of the Yamnaya culture distribution area, such as Globular Amphora and Tripolye (Cucuteni–Trypillia) individuals from Ukraine, which also have been shown to carry Anatolian Neolithic farmer-derived ancestry24.

On the other hand, it is interesting that – although no information is released about these samples – Yamna Bulgaria is now a clear outlier, among very “Yamnaya”-like Yamna settlers from Hungary, most likely from the Carpathian basin, and new Yamna LCA/EBA samples, possibly from Late Yamna (see them also marked in the PCA above):

yamnaya-hungary-admixture
Modified image, with red rectangles surrounding (unexplained) Hungarian samples (c) ADMIXTURE results of relevant prehistoric individuals mentioned in the text (filled symbols)

The important admixture of Yamna settlers with native populations, seen in expanding East Bell Beakers of R1b-L23 lineages from ca. 2500 BC on, must have therefore happened at the same time as the adoption of the proto-Bell Beaker package, i.e. precisely during the Carpathian Basin / Lower Danube settlements, and not in West Yamna.

yamnaya-hungary-lca-eba
Modified image, with red rectangles surrounding (unexplained) Yamna samples Modelling results for the Steppe and Caucasus cluster. Admixture proportions based on (temporally and geographically) distal and proximal models, showing additional Anatolian farmer-related ancestry in Steppe groups as well as additional gene flow from the south in some of the Steppe groups as well as the Caucasus groups

So, it can’t get clearer that Late Neolithic Baltic and Corded Ware migrants, sharing R1a-Z645 lineages and a different admixture, related to Eneolithic North Pontic groups such as Sredni Stog (see above ADMIXTURE graphics of CWC and Eneolithic Ukraine samples), did not come from West Yamna migrants, either.

So much for the R1a/R1b Yamna community that expanded Late PIE into Corded Ware.

NOTE. Andrew Gelman has coined a term for a curious phenomenon (taken from an anonymous commenter): “Eureka bias”, which refers not only to how researchers stick to previously reported incorrect results or interpretations, but also to how badly they react to criticism, even if they understand that it is well-founded. Directly applicable to the research groups that launched the Yamna-CWC idea (and the people who followed them) based on the fallacious “Yamnaya ancestry” concept, and who are still rooting for some version of it, from now on with exogamy, patron-client relationships, Eneolithic Indo-Slavonic, and whatnot. Unless, that is, Anthony’s latest model is right, and Yamna Hungary is suddenly full of R1a-Z645 samples…

Images used are from the article. They are available under a CC-BY-NC-ND 4.0 International license. (Yes, I know, I modified them. To mark special newly reported samples from Yamna Hungary and Yamna LCA/EBA. I expect this to count as fair use).

Related:

Sintashta-Petrovka and Potapovka cultures, and the cause of the Steppe EMBA – MLBA differences

Interesting recent papers on Sintashta and related Volga-Ural MLBA communities, with relevant excerpts (emphasis mine):

Social Organization of the Sintashta-Petrovka Groups of the Late Bronze Age and a Cause for Origin of Social Elites (Based on Materials of the Settlement of Kamenny Ambar), by Chechushkov et al. Stratum Plus (2018) Nº2.

Abstract (official, in English):

The formation of social complexity often unfolded in non-unilineal ways in those regions of the world where the surplus product remained low enough to support institutionalized power and state bureaucracy. The Bronze Age of Northern Eurasia is a vivid example where social complexity arose based on herding economy, while population density remained low enough not to form territorially separate competing groups. Studying of such societies sheds light on how and under what conditions the social elite emerged. The undertaken analysis suggests that the formation, development, and decline of social complexity in the Bronze Age steppe societies were directly related to the intensification of subsistence practices and colonization of new territories. At the same time, some members of the society took upon themselves the role of community life’s managers, and, in return, received privileged statuses. The environment and the economy changing, the need for such functions disappeared. As a result, the Bronze Age social elites dissolved in the mass and lost their privileged statuses.


Open access Sintashta as a transcultural phenomenon, by N.B. Vinogradov, Архив Поволжская археология №1 (23) 2018.

Abstract (official, in English)

The paper features a substantiation of the understanding of Sintashta-type monuments dating back to the boundary of the Middle and Late Bronze Ages in the Southern Trans- Urals as a transcultural phenomenon, the establishment and operation of which has been associated with the clans of miners, smelters and smiths of the Southern Ural cultures of the studied period. In the author’s opinion, the variety of ceramic complexes from Sintashta burial mounds suggest a reconsideration of several cultural traditions and the peculiar nature of family and marriage relations practiced by the clans of Sintashta-type communities.

Interesting excerpts, from the conclusion (translated from Russian):

1. In contrast to the pastoral cattle-breeding cultures of the Alakul cultural-historical community, the Sintashta clan-communities, in the author’s opinion, were a more specific transcultural phenomenon with an original model of life organization, uniting clans of miners, metallurgists, blacksmiths and casters, sometimes from several neighboring archaeological cultures (my italics – N.V.), in particular, the Abashevo culture of the South Urals, some “proto-Srubna” culture of the Southern Urals and quasi-Eneolithic cultures of the Southern Urals and of Northern Kazakhstan.

2. The Sintashta phenomenon as a community of clans of miners – metallurgists – smiths functioned relatively independently, outside or under conditions of partial jurisdiction (?) Of the elites of the above mentioned cultures.

3. At the historical level, the facts presented by the author concerning both funeral rites and ceramics can be understood as a reflection of the characteristics, first of all, of the family-marriage relations system within the specialized communities. And it is not by chance that the ceramics of Sintashta cemeteries carry in themselves often reinterpreted (especially in the case of ornamentation) traces of several cultural traditions. The variety of ceramic complexes of Sintashta monuments and the rethinking, reworking of marker elements of ornament on vessels testify, in the author’s view, about the distinctiveness, the specifics of family relations in Sintashta communities.


Paleoanthropological Data as a Source of Reconstruction of the Process of Social Formation and Social Stratifi cation (based on the Sintashta and Potapovo sites of the Bronze Age), by Kitov et al. Stratum Plus (2018) Nº 2.

Abstract (official, in English):

The paper is devoted to the analysis of craniological materials from the cemeteries of the Bronze Age of the Volga-Ural region (Sintashta and Potapovo assemblages). The characteristic feature of the physical appearance of this population is the combination of different morphological variations with a dominant and the presence of the Uraloid components. At the same time, a group of individuals with a specific, different from other individuals, skull structure is distinguished: maturized, broad-faced men with a set of striking features in the face. Analysis of the funerary rites of these individuals indicates their high social status in the Sintashta-Potapovo society. The addition of such an anthropological complex occurred in the Eneolithic on the territory of modern Kazakhstan as a result of contacts of steppe sharply profiled Europeoid populations and groups of Uraloid origin. This led to the formation of a population, originally of metisic origin, conventionally called “steppe Kazakhstan”, which took part in the process of morphogenesis, and, indirectly, the cultural genesis of Sintashta and Potapovo communities.

While this paper reports mainly athropometric data, the team forms part of the Samara Valley project – including Khokhlov.

Here are interesting excerpts from the general conclusions (translated from Russian):

Summing up, it can be noted that the distinguishing feature of the carriers of the Sintashta and Potapovka traditions is the sharp heterogeneity of the anthropological features, the cause of which were active ethno- and culturogenetic processes in the Volga-Ural region at the turn of the 3rd/2nd millennium BC. One of the active components of these processes was probably a population group with specific craniological data, distinct from the rest of the craniocomplexes. These included mature, broad-leaning male individuals with a set of vivid signs in the structure of the face, such as unfolded and flattened cheekbones, and a strong nose protrusion.

The peculiarities of the burial rite speak about their high social position in the society: burials were made in large central burial pits, accompanied by abundant sacrificial remains in the form of skulls and limbs of horses, large and small cattle, rich funeral complements including bronze tools and weapons, artifacts of metal production, attributes of the chariot complex. It should be noted that such a craniological type is present in every mound of the Sintashta-Potapovka circle of monuments, and is found on the wide territory of the steppes and forest-steppes of the Volga region, the Southern Urals, and the Trans-Urals. The addition of the similar anthropological complex occurred in the Eneolithic due to the contacts, on the one hand, of steppe sharply profiled Europoid populations that extended to the east and, on the other hand, encountered groups of uraloid origin, which led to the formation of a population, originally of metisic origin, which can be conditionally called “steppe Kazakhstan”.

Related:

Consequences of Damgaard et al. 2018 (III): Proto-Finno-Ugric & Proto-Indo-Iranian in the North Caspian region

copper-age-early_yamna-corded-ware

The Indo-Iranian – Finno-Ugric connection

On the linguistic aspect, this is what the Copenhagen group had to say (in the linguistic supplement) based on Kuz’mina (2001):

(…) a northern connection is suggested by contacts between the Indo-Iranian and the Finno-Ugric languages. Speakers of the Finno-Ugric family, whose antecedent is commonly sought in the vicinity of the Ural Mountains, followed an east-to-west trajectory through the forest zone north and directly adjacent to the steppes, producing languages across to the Baltic Sea. In the languages that split off along this trajectory, loanwords from various stages in the development of the Indo-Iranian languages can be distinguished: 1) Pre-Proto-Indo-Iranian (Proto-Finno-Ugric *kekrä (cycle), *kesträ (spindle), and *-teksä (ten) are borrowed from early preforms of Sanskrit cakrá- (wheel, cycle), cattra- (spindle), and daśa- (10); Koivulehto 2001), 2) Proto-Indo-Iranian (Proto-Finno-Ugric *śata (one hundred) is borrowed from a form close to Sanskrit śatám (one hundred), 3) Pre-Proto-Indo-Aryan (Proto-Finno-Ugric *ora (awl), *reśmä (rope), and *ant- (young grass) are borrowed from preforms of Sanskrit ā́rā- (awl), raśmí- (rein), and ándhas- (grass); Koivulehto 2001: 250; Lubotsky 2001: 308), and 4) loanwords from later stages of Iranian (Koivulehto 2001; Korenchy 1972). The period of prehistoric language contact with Finno-Ugric thus covers the entire evolution of Pre-Proto-Indo-Iranian into Proto-Indo-Iranian, as well as the dissolution of the latter into Proto-Indo- Aryan and Proto-Iranian. As such, it situates the prehistoric location of the Indo-Iranian branch around the southern Urals (Kuz’mina 2001).

NOTE. While I agree with the evident ancestral nature of the *kekrä borrowing, I will repeat it here again: I don’t believe that the distinction of late Proto-Indo-Iranian from ‘Pre-Proto-Indo-Aryan’ loans is warranted; not for words reconstructed from recent Finno-Ugric languages.

copper-age-late-urals
The time and place for Finno-Ugric and Indo-Iranian contacts. Late Copper Age migrations in Asia ca. 2800-2300 BC.

In this period of a Pre-Proto-Indo-Iranian community, which is to be associated with East Yamna/Poltavka, ca. 3000-2400 BC – as accepted in the supplement from de Barros Damgaard et al. (Nature 2018) – , both Poltavka and Abashevo/Balanovo herders were expanding ca. 2800-2600 BC to the east (and Abashevo already admixing into Poltavka territory), near the southern Urals.

There is no other, clearer, later connection between Finno-Ugric and Proto-Indo-Iranian speakers. Even the arrival of the Seima-Turbino phenomenon (after ca. 2000 BC), if it brought migrants to North-East Europe, would not fit the linguistic, archaeological, or genetic data. It is by now quite clear that Seima-Turbino does not fit with incoming N1c1 lineages and/or Siberian ancestry, either, for those looking for these as potential signs of incoming Uralic speakers.

While the Copenhagen group did not have access to data from Sintashta ca. 2100 BC onwards – now available in Narasimhan et al. (2018) – when submitting the papers, we already know that there was a clear long period of slow progressive admixture in the North Caspian region. It can be seen in the genetic contribution of Yamna to incoming Abashevo groups, and in the R1b-L23 samples still appearing in Sintashta until ca. 1800 BC (as I predicted could happen).

Since the first sample signalling incoming Abashevo migrants is found in the Poltavka outlier dated ca. 2700 BC (of R1a-Z93 lineage), this represents a rather unique, several centuries long process of admixture in the North Caspian region, different from the massive Afanasevo or Bell Beaker migrations in Asia and Europe, whereby a great part of the native male population was suddenly replaced.

This offers further support for language continuity despite genetic replacement in the development of East Yamna/Poltavka (part of the Steppe EMBA cline, formed by Yamna and Afanasevo) mixing with Abashevo migrants (probably identical to Corded Ware samples) to form Potapovka, Sintashta, and later Srubna, and Andronovo communities (all forming, with Corded Ware groups, a wide Eurasian Steppe MLBA cloud). See the available data from Narasimhan et al. (2018).

yamna-late-proto-indo-european
Image modified from Narasimhan et al. (2018), including the most likely proto-language identification of different groups. Original description “Modeling results including Admixture events, with clines or 2-way mixtures shown in rectangles, and clouds or 3-way mixtures shown in ellipses”. See the original full image here.

The continuous interactions and migrations left thus eventually two communities in the southern Urals genetically similar, but ethnolinguistically diverse:

  • To the north, Abashevo-Balanovo – but potentially also Fatyanovo, and related North-East European late Corded Ware groups – borrowed necessary words from Indo-Iranian neighbours, while maintaining their Finno-Ugric language and culture.
  • To the south, immigrants (or their descendants) of Abashevo origin expanding among Pre-Proto-Indo-Iranian-speaking North Caspian communities assimilated the surrounding culture and language, giving it their own accent (i.e. ‘satemizing’ it) and turning it into Proto-Indo-Iranian (see e.g. Parpola’s account).

Anthropologically, this ‘long-term founder effect’ that appears as genetic replacement is probably explained by the faster life history in MLBA North Caspian populations, likely due to a combination of changing environmental and social circumstances.

NOTE. The prevalent explanation before the latest studies on the Sintashta society were social strife and isolation of small groups, an argument I used in my demic diffusion model. Other, similar cases of proven linguistic continuity despite genetic replacement are seen in Iberian Bronze Age after the expansion of R1b-L23 lineages (with Vasconic, Iberian, and Tartessian surviving at least until proto-historic times), and in Remote Oceania.

bronze_age_early_Asia-andronovo
Diachronic map of migrations in Asia ca. 2250-1750 BC

Implications for Late PIE migrations

I am happy to see that people are resorting now to dialectal classifications and Y-DNA to explain the findings in Old Hittites, Tocharians (and related migrations), and Indo-Iranians. It is especially interesting to see precisely this Danish group downplay the relevance of ancestry and favor complex anthropological models when assessing migrations and ethnolinguistic identification.

So let’s talk about the growing elephant in the room.

It seems we all accept now Tocharian’s more archaic Late PIE nature, which is supported by waves of late Khvalynsk migrants starting probably ca. 3300 BC, as seen in different samples to the east in Central Asia, and to the south in Iran. Almost all of them share R1b-L23 lineages.

NOTE. Whereas their early LPIE dialects have not survived to historic times, the rather speculative hypotheses of Euphratic and Gutian languages may be of interest.

We also know of the coetaneous migrants that settled to the west of the Don River (in the territory of the previous late Sredni Stog culture), to form the western South-Bug / Lower Don groups, which, together with the Volga-Ural / North Caucasian groups formed the early Yamna culture, that dominated from ca. 3300 BC over the Pontic-Caspian steppe.

It is only logical that the other attested languages belonging to the common Late PIE trunk must come from these groups, which must have stuck together for quite some time – after the recently proven late Khvalynsk migrations – , to allow for the spread of isoglosses (not found in Tocharian) among them.

This is agreed, even by the Copenhagen group, who expressly state that Yamna is to be identified with the rest of Late PIE languages after the Tocharian-related migrations.

copper-age-early_yamna-corded-ware
Early Yamna community and its migrations ca. 3000 BC onwards.

The period of an early Yamna community constrained to the Pontic-Caspian steppe (ca. 3300-3000 BC) is followed by renewed waves of Late Proto-Indo-European migrations, during which areal contacts and innovations (even between unrelated LPIE branches) can still be reconstructed.

These later migrations can be precisely described as follows (after the latest studies):

  • Yamna migrants, of mixed R1b-L51 and R1b-Z2103 lineages, settle ca. 3000-2600 BC along the lower Danube, in the Balkans and the Carpathian basin, giving rise later to groups of:
  • In the Pontic-Caspian steppe, early Yamna groups evolve into (from west to east) Late Yamna, Catacomb, and Poltavka groups, ca. 2800-2300 BC, all still dominated by R1b-L23 lineages (see discussion on the Catacomb sample), with:
    • Poltavka peoples admixing with Abashevo migrants to form admixed Potapovka and Sintashta-Petrovka groups, showing still after ca. 1800 BC a mixed society of R1a-Z93 and R1b-Z2103 lineages (see Narasimhan et al. 2018);
      • Expanding early Proto-Iranian and Proto-Indo-Aryan groups in Srubna (to the west) and Andronovo (to the east), during the first half of the 2nd millennium BC, dominate over the Bronze Age steppe and Central Asia with expanding R1a-Z93 lineages.

Conclusion

chalcolithic_late_Europe_Bell_Beaker
Diachronic map of Late Copper Age migrations including Classical Bell Beaker (east group) expansion from central Europe ca. 2600-2250 BC

1) East Bell Beakers clearly dominated culturally and genetically over almost all of Europe, ca. 2500-2000 BC, including previous Corded Ware territory, representing thus the most recent massive migration of steppe peoples in Europe, and being the only pan-European culture derived from Late Proto-Indo-European-speaking Yamna. They must therefore be identified with North-West Indo-European speakers, as proposed by Mallory (2013), and not just Italo-Celtic (as supported recently by the Danish school, based on Gimbutas’ outdated model):

1.A) For Germanic, we already have proof that an appropriate, unitary Scandinavian society, ripe for the development of a common Pre-Germanic language (that expanded much later, during the Iron Age, as Proto-Germanic) could have developed only after the arrival of Bell Beakers (see Prescott 2017). The association of proto-historic Germanic tribes mainly with the expansion of R1b-U106 lineages bears witness to that.

NOTE. Even without taking into account the likely L51 samples from Khvalynsk, it is by now quite clear that R1b-L51 lineages were already admixed in Yamna settlers from the Carpathian Basin, and any subclade of U106, L21, DF27, or U152 can thus be found everywhere in Europe associated with any of those North-West Indo-European migrations. What we are seing later, as in the East Bell Beaker migrants arriving in the British Isles (L21), Iberia (DF27), or the Netherlands/Scandinavia (U106), is the further reduction in variability coupled with the expansion of a few sucessful families (and their lineages), as we know it usually happens during migrations.

1.B) For Balto-Slavic, it seems they were not part of the eastern Corded Ware peoples: the Copenhagen group denies an Indo-Slavonic group in the Nature paper, referring instead to a dominion of early Iranians in the steppes, following their traces to proto-historic and historic Iranian-speaking peoples. And we knew already that Bell Beakers dominated over Central-East Europe, before the resurge of R1a-Z645 lineages in the region, which is compatible with the North-West Indo-European nature of their language undergoing a satemization process similar (but not equal to) to the Indo-Iranian one (see the full discussion on Balto-Slavic here).

NOTE. The few ancestral traits common to Germanic and Balto-Slavic are today considered a common substrate language to both, and not due to close contacts (and still less a common branch, as was proposed in the 1st half of the 20th c.). You can read e.g. Kortlandt’s Baltic, Slavic, Germanic (2017), or our Corded Ware substrate hypothesis (2017). In both theories, the referenced substrate is likely a non-Indo-European language, and in both cases it is related to the Corded Ware culture, which represents their most common immediate ancestral population before the spread of Bell Beakers.

2) The late Corded Ware groups of Finland and Estonia, as well as Fatyanovo and Abashevo (and succeeding groups of Eastern Europe) may now be more clearly associated with Proto-Finno-Ugric dialects, and thus probably Corded Ware groups in general with Uralic languages, whose western branches have not survived to this day, with their culture and language being replaced quite early by expanding Bell Beakers.

NOTE. While the demise of Central and Central-East European CWC groups is evident, continuous contacts among Battle Axe culture groups in Scandinavia and the Gulf of Finland through the Baltic Sea – and the strong Bronze Age Palaeo-Germanic influence on Finnic languages (stronger than earlier Indo-Iranian borrowings) may point to the continuity of Proto-Finnic in Northern Scandinavia, which may force a reinterpretation of the prehistoric location of Proto-Finnic-speaking groups.

Those supporting a Corded Ware expansion of Germanic or Balto-Slavic with R1a subclades, now rejecting the expansion of Proto-Indo-European from an Anatolian homeland (following the spread of Neolithic farmer ancestry), and negating the close Proto-Indo-Iranian – Uralic contacts, are willfully ignoring linguistic, archaeological, and genetic data whenever it does not fit with their previous theories.

Good times ahead to chase false syllogisms and contradictions everywhere.

Related:

Consequences of Damgaard et al. 2018 (II): The late Khvalynsk migration waves with R1b-L23 lineages

chalcolithic_early-asia

This post should probably read “Consequences of Narasimhan et al. (2018),” too, since there seems to be enough data and materials published by the Copenhagen group in Nature and Science to make a proper interpretation of the data that will appear in their corrected tables.

The finding of late Khvalynsk/early Yamna migrations, identified with early LPIE migrants almost exclusively of R1b-L23 subclades is probably one of the most interesting findings in the recent papers regarding the Indo-European question.

Although there are still few samples to derive fully-fledged theories, they begin to depict a clearer idea of waves that shaped the expansion of Late Proto-Indo-European migrants in Eurasia during the 4th millennium BC, i.e. well before the expansion of North-West Indo-European, Palaeo-Balkan, and Indo-Iranian languages.

Late Khvalynsk expansions and archaic Late PIE

Like Anatolian, Tocharian has been described as having a more archaic nature than the rest of Late PIE. However, Pre-Tocharian belongs to the Late PIE trunk, clearly distinguishable phonetically and morphologically from Anatolian.

It is especially remarkable that – even though it expanded into Asia – it has more in common with North-West Indo-European, hence its classification (together with NWIE) as part of a Northern group, unrelated to Graeco-Aryan.

The linguistic supplement by Kroonen et al. accepts that peoples from the Afanasevo culture (ca. 3000-2500 BC) are the most likely ancestors of Tocharians.

NOTE. For those equating the Tarim Mummies (of R1a-Z93 lineages) with Tocharians, you have this assertion from the linguistic supplement, which I support:

An intermediate stage has been sought in the oldest so-called Tarim Mummies, which date to ca. 1800 BCE (Mallory and Mair 2000; Wáng 1999). However, also the language(s) spoken by the people(s) who buried the Tarim Mummies remain unknown, and any connection between them and the Afanasievo culture on the one hand or the historical speakers of Tocharian on the other has yet to be demonstrated (cf. also Mallory 2015; Peyrot 2017).

New samples of late Khvalynsk origin

These are are the recent samples that could, with more or less certainty, correspond to migration waves from late Khvalynsk (or early Yamna), from oldest to most recent:

  • The Namazga III samples from the Late Eneolithic period (in Turkmenistan), dated ca. 3360-3000 BC (one of haplogroup J), potentially showing the first wave of EHG-related steppe ancestry into South Asia. Not related to Indo-Iranian migrations.

NOTE. A proper evaluation with further samples from Narasimhan et al. (2018) is necessary, though, before we can assert a late Khvalynsk origin of this ancestry.

  • Afanasevo samples, dated ca. 3081-2450 BC, with all samples dated before ca. 2700 BC uniformly of R1b-Z2103 subclades, sharing a common genetic cluster with Yamna, showing together the most likely genomic picture of late Khvalynsk peoples.

NOTE 1. Anthony (2007) put this expansion from Repin ca. 3300-3000 BC, while his most recent review (2015) of his own work put its completion ca. 3000-2800. While the migration into Afanasevo may have lasted some time, the wave of migrants (based on the most recent radiocarbon dates) must be set at least before ca. 3100 BC from Khvalynsk.

NOTE 2. I proposed that we could find R1b-L51 in Afanasevo, presupposing the development of R1b-L51 and R1b-Z2103 lineages with separating clans, and thus with dialectal divisions. While finding this is still possible within Khvalynsk regions, it seems we will have a division of these lineages already ca. 4250-4000 BC, which would require a closer follow-up of the different inner late Khvalynsk groups and their samples. For the moment, we don’t have a clear connection through lineages between North-West Indo-European groups and Tocharian.

tocharian-early-copper-age
Early Copper Age migrations in Asia ca. 3300-2800, according to Anthony (2015).
  • Subsequent and similar migration waves are probably to be suggested from the new sample of Karagash, beyond the Urals (attributed to the Yamna culture, hence maintaining cultural contacts after the migration waves), of R1b-Z2103 subclade, ca. 3018-2887 BC, potentially connected then to the event that caused the expansion of Yamna migrants westward into the Carpathians at the same time. Not related to Indo-Iranian migrations.
  • The isolated Darra-e Kur sample, without cultural adscription, ca. 2655 BC, of R1b-L151 lineage. Not related to Indo-Iranian migrations.
  • The Hajji Firuz samples: I4243 dated ca. 2326 BC, of haplogroup I1b, with a clear inflow of steppe ancestry; and I2327 (probably to be dated to the late 3rd millennium BC or after that), of R1b-Z2103 lineage. Not related to Indo-Iranian migrations.

NOTE. A new radiocarbon dating of I2327 is expected, to correct the currently available date of 5900-5000 BC. Since it clusters nearer to Chalcolithic samples from the site than I4243 (from the same archaeological site), it is possible that both are part of similar groups receiving admixture around this period, or maybe I2327 is from a later period, coinciding with the Iron Age sample F38 from Iran (Broushaki et al. 2016), with which it closely clusters. Also, the finding of EHG-related ancestry in Maykop samples dated ca. 3700-3000 BC (maybe with R1b-L23 subclades) offers another potential source of migrants for this Iranian group.

NOTE. Samples from Narasimhan et al. (2018) still need to be published in corrected tables, which may change the actual subclades shown here.

These late Khvalynsk / early Yamna migration waves into Asia are quite early compared to the Indo-Iranian migrations, whose ancestors can only be first identified with Volga-Ural groups of Yamna/Poltavka (ca. 3000-2400 BC), with its fully formed language expanding only with MLBA waves ca. 2300-1200 BC, after mixing with incoming Abashevo migrants.

While the authors apparently forget to reference the previous linguistic theories whereby Tocharian is more archaic than the rest of Late PIE dialects, they refer to the ca. 1,000-year gap between Pre-Tocharian and Proto-Indo-Iranian migrations, and thus their obvious difference:

The fact that Tocharian is so different from the Indo-Iranian languages can only be explained by assuming an extensive period of linguistic separation.

Potential linguistic substrates in the Middle East

A few words about relevant substrate language proposals.

Euphratic language

What Gordon Whittaker proposes is a North-West Indo-European-related substratum in Sumerian language and texts ca. 3500 BC, which may explain some non-Sumerian, non-Semitic word forms. It is just one of many theories concerning this substratum.

eneolithic_steppe
Diachronic map of Eneolithic migrations ca. 4000-3100 BC

This is a summary of his findings from his latest writing on the subject (a chapter of a book on Indo-European phonetics, from the series Copenhagen Studies in Indo-European):

In Sumerian and Akkadian vocabulary, the cuneiform writing system, and the names of deities and places in Southern Mesopotamia a body of lexical material has been preserved that strongly suggests influence emanating from a superstrate of Indo-European origin. his Indo-European language, which has been given the name Euphratic, is, at present, attested only indirectly through the filters of Sumerian and Akkadian. The attestations consist of words and names recorded from the mid-4th millennium BC (Late Uruk period) onwards in texts and lexical lists. In addition, basic signs that originally had a recognizable pictorial structure in proto-cuneiform preserve (at least from the early 3rd millennium on) a number of phonetic values with no known motivation in Sumerian lexemes related semantically to the items depicted. This suggests that such values are relics from the original logographic values for the items depicted and, thus, that they were inherited from a language intimately associated with the development of writing in Mesopotamia. Since specialists working on proto-cuneiform, most notably Robert K. Englund of the Cuneiform Digital Library Initiative, see little or no evidence for the presence of Sumerian in the corpus of archaic tablets, the proposed Indo-European language provides a potential solution to this problem. It has been argued that this language, Euphratic, had a profound influence on Sumerian, not unlike that exerted by Sumerian and Akkadian on each other, and that the writing system was the primary vehicle of this influence. he phonological sketch drawn up here is an attempt to chart the salient characteristics of this influence, by comparing reconstructed Indo-European lexemes with similarly patterned ones in Sumerian (and, to a lesser extent, in Akkadian).

His original model, based on phonetic values in basic proto-cuneiform signs, is quite imaginative and a very interesting read, if you have the time. His Academia.edu account hosts most of his papers on the subject.

We could speculate about the potential expansion of this substrate language with the commercial contacts between Uruk and Maykop (as I did), now probably more strongly supported because of the EHG found in Maykop samples.

NOTE. We could also put it in relation with the Anatolian language of Mari, but this would require a new reassessment of its North-West Indo-European nature.

Nevertheless, this theory is far from being mainstream, anywhere. At least today.

NOTE. The proposal remains still hypothetic, because of the flaws in the Indo-European parallels – similar to Koch’s proposal of Indo-European in Tartessian inscriptions. A comprehensive critic approach to the theory is found in Sylvie Vanséveren’s A “new” ancient Indo-European language? On assumed linguistic contacts between Sumerian and Indo-European “Euphratic”, in JIES (2008) 36:3&4.

Gutian language

References to Gutian are popping up related to the Hajji Firuz samples of the mid-3rd millennium.

The hypothesis was put forward by Henning (1978) in purely archaeological terms.

This is the relevant excerpt from the book:

(…) Comparativists have asserted that, in spite of its late appearance, Tokharian is a relatively archaic form of Indo-European.3 This claim implies that the speakers of this group separated from their Indo-European brethren at a comparatively early date. They should accordingly have set out on their migrations rather early, and should have appeared within the Babylonian sphere of influence also rather early. Earlier, at any rate, than the Indo-Iranians, who spoke a highly developed (therefore probably later) form of Indo-European. Moreover, as some of the Indo-Iranians after their division into Iranians and Indo-Aryans4 appeared in Mesopotamia about 1500 B.C., we should expect the Proto-Tokharians about 2000 B.C. or even earlier.

If, armed with these assumptions as our working hypothesis, we look through the pages of history, we find one nation – one nation only – that perfectly fulfills all three conditions, which, therefore, entitles us to recognize it as the “Proto-Tokharians”. Tis name was Guti; the intial is also spelled with q (a voiceless back velar or pharyngeal), but the spelling with g is the original one. The closing -i is part of the name, for the Akkadian case-endings are added to it, nom. Gutium etc. Guti (or Gutium, as some scholars prefer) was valid for the nation, considered as an entity, but also for the territory it occupied.
(…).

The text goes on to follow the invasion of Babylonia by the Guti, and further eastward expansions supposedly connected with these, to form the attested Tocharians.

The referenced text by Thorkild Jakobsen offers the interesting linguistic data:

Among the Gutian rulers is one Elulumesh, whose name is evidently Akkadian Elulum slightly “Gutianized” by the Gutian case(?) ending -eš.40 This Gutian ruler Elulum is obviously the same man whom we find participating in the scramble for power after the death of Shar-kali-sharrii; his name appears there in Sumerian form without mimation as Elulu.

The Gutian dynasty, from ca. 22nd c. BC appears as follows:

gutian-rulers

I don’t think we could derive a potential relation to any specific Indo-European branch from this simple suffix repeated in Gutian rulers, though.

The hypothesis of the Tocharian-like nature of the Guti (apart from the obvious error of considering them as the ancestors of Tocharians) remains not contrasted in new works since. It was cited e.g. by Gamkrelidze and Ivanov (1995) to advance their Armenian homeland, and by Mallory and Adams in their Encyclopedia (1997).

It lies therefore in the obscurity of undeveloped archaeological-linguistic hypotheses, and its connection with the attested R1b-Z2103 samples from Iran is not (yet) warranted.

Related:

No large-scale steppe migration into Anatolia; early Yamna migrations and MLBA brought LPIE dialects in Asia

eurasian-hittite-samples

Another, simultaneous paper with the Eurasian samples from Nature, The first horse herders and the impact of early Bronze Age steppe expansions into Asia, by de Barros Damgaard et al., Science (2018).

A lot of interesting data, I will try to analyse its main implications, if only superficially, in sections.

Anatolian samples

Anatolia_EBA from Ovaören, and Anatolia_MLBA (this including Assyrian and Old Hittite samples), all from Kalehöyük, show almost no change in Y-DNA lineages (three samples J2a, one G2a), and therefore an origin of these people in common with CHG and Iranian Neolithic populations is likely. No EHG ancestry is found. And PCA cluster is just somehow closer to Europe, but not to EHG populations.

NOTE. Hittite is attested only in the late first half of the 2nd millennium, although the authors cite (in the linguistic supplement) potential evidence from the palatial archives of the ancient city of Ebla in Syria to argue that Indo-European languages may have been already spoken in the region in the late 3rd millennium BCE.

Regarding the Assyrian samples (one J2a) from Ovaören:

Layer V of GT-137 was the richest in terms of architectural finds and dates to the Early Bronze Age II. In this layer, 2 different structures and a well were uncovered. The well was filled with stones, pottery, and human skeletons (Figs. S2 and S3). In total, skeletons belonging to 22 individuals, including adults, young adults, and children, must belong to the disturbed Early Bronze Age II graves adjacent to the well (103). Pottery and stones found below the skeletons demonstrate that the water well was consciously filled and closed. The fill consists of dumped stones, sherds and skeletons, and the closing stones demonstrate that the water well was consciously filled and cancelled.

Regarding the site most likely associated with the emergence of Old Hittite (two samples J2a1, one G2a2b1), this is what we know:

The Middle Bronze Age at Kaman-Kalehöyük represented by stratum IIIc yields material remains (seals and ceramics) contemporary with the international trade system managed by expatriate Assyrian merchants evidenced at the nearby site of Kültepe/Kanesh. It is therefore also referred to as belonging to the “Assyrian Colony Period” (98). The stratum has revealed three burned architectural units, and it has been suggested that the seemingly site-wide conflagration might be connected to a destruction event linked with the emergence of the Old Hittite state (99). (…) Omura (100) suggests that the rooms could belong to a public building, and that it might even be a small trade center based on the types of artifacts recovered. Omura (100) has concluded that the evidence from the first complex indicates a battle between 2 groups took place at the site. It is possible that a group died inside the buildings, mostly perishing in the fire, while another group died in the courtyard.

NOTE. For more on the Old Hittite period, you can read this for example.

Regarding PCA:

The PCA (Fig. 2B) indicates that all the Anatolian genome sequences from the Early Bronze Age ( -2200 BCE) and Late Bronze Age (-1600 BCE) cluster with a previously sequenced Copper Age ( -3900- 3700 BCE) individual from Northwestern Anatolia and lie between Anatolian Neolithic (Anatolia_ N) samples and CHG samples but not between Anatolia_N and EHG samples.

(…) we are not able to reject a two-population qpAdm model in which these groups derive -60% of their ancestry from Anatolian farmers and -40% from CHG-related ancestry (p-value = 0.5). This signal is not driven by Neolithic Iranian ancestry.

pca-ancient-modern-eurasians
Principal Component Analysis estimated with ancient and modern Eurasians.

NOTE. Anatolian Iron Age samples, from the Hellenistic period, which was obviously greatly influenced by different, later Indo-European migrations, does show a change in PCA.

Regarding CHG ancestry:

Ancient DNA findings suggest extensive population contact between the Caucasus and the steppe during the Copper Age (-5000-3000 BCE) (1, 2, 42). Particularly, the first identified presence of Caucasian genomic ancestry in steppe populations is through the Khvalynsk burials (2, 47) and that of steppe ancestry in the Caucasus is through Armenian Copper Age individuals (42). These admixture processes likely gave rise to the ancestry that later became typical of the Yamnaya pastoralists (7), whose IE language may have evolved under the influence of a Caucasian language, possibly ‘from the Maykop culture (50, 55). This scenario is consistent with both the “Copper Age steppe” (4) and the “Caucasian” models for the origin of the Proto-Anatolian language (56).

The CHG specific ancestry and the absence of EHG-related ancestry in Bronze Age Anatolia would be in accordance with intense cultural interactions between populations in the Caucasus and Anatolia observed during the late 5th millennium BCE that seem to come to an end in the first half of the 4th millennium BCE with the village-based egalitarian Kura-Araxes society (59, 60), thus preceding the emergence and dispersal of Proto-Anatolian.

Our results indicate that the early spread of IE languages into Anatolia was not associated with any large-scale steppe-related migration, as previously suggested (61). Additionally, and in agreement with the later historical record of the region (62), we find no correlation between genetic ancestry and exclusive ethnic or political identities among the populations of Bronze Age Central Anatolia, as has previously been hypothesized ( 63).

The Anatolian question

There is no steppe ancestry or R1b-M269 lineages near early historic Hittites. Yet.

Nevertheless, we already know about potentially similar cases:

So there seems to be thus no theoretical problem in accepting:

  • That neither steppe ancestry nor R1b-M269 subclades, already diminished in Bulgaria in the mid-5th millennium, did reach Anatolia, but only those Common Anatolian-speaking Aegean groups over whose ancestors Proto-Anatolians (marked by incoming EHG ancestry) would have previously dominated in the Balkans.
  • That steppe ancestry and R1b-M269 subclades did in fact arrive in the Aegean, but EHG was further diluted among the CHG-related population by the time of the historic Anatolian-speaking peoples in central Anatolia. Or, the most likely option, that their trace have not been yet found. Probably the western Luwian peoples, near Troy, were genetically closer to Common Anatolians.

Both of these scenarios are interesting, in that they show potential links between Pre-Greek peoples of Hellas (related to Anatolians) and the Pelasgian substrate of early Greek dialects, since they show a similar recent CHG-related wave from the East.

What we can assert right now is that Proto-Anatolian must have separated quite early for this kind of data to show up. This should mean an end to the Late PIE origin of Anatolian, if there was some lost soul from the mid-20th century still rooting for this.

As I said in my review of Lazaridis’ latest preprint, we will have to wait for the appropriate potential routes of expansion of Proto-Anatolian to be investigated. As he answered, the lack of EHG poses a problem for steppe expansion into Anatolia, but there is still no better alternative model proposed.

Eurasian-CHG-ancestry
Model-based clustering analysis of present-day and ancient individuals assuming K = 6 ancestral components. The main ancestry components at K = 6 correlate well with CHG (turquoise), a major component of Iran_N, Namazga_CA and South Asian dines; EHG (pale blue), a component of the steppe dine and present in South Asia; East Asia (yellow ochre), the other component of the steppe d ine also in Tibeto-Burman South Asian populations; South Indian (pink), a core component of South Asian populations; Anatolian_N (purple), an important component of Anatolian Bronze Age and Steppe_MLBA; Onge (dark pink) forms its own component.

This is what the authors have to say:

Our findings are thus consistent with historical models of cultural hybridity and “Middle Ground” in a multi-cultural and multi-lingual but genetically homogeneous Bronze Age Anatolia (68, 69). Current linguistic estimations converge on dating the Proto-Anatolian split from residual PIE to the late 5th or early 4th millennia BCE (58, 70) and place the breakup of Anatolian IE inside Turkey prior to the mid-3rd millennium (53, 71,72).

We cannot at this point reject a scenario in which the introduction of the Anatolian IE languages into Anatolia was coupled with the CHG-derived admixture prior to 3700 BCE, but note that this is contrary to the standard view that PIE arose in the steppe north of the Caucasus (4) and that CHG ancestry is also associated with several non-IE-speaking groups, historical and current. Indeed, our data are also consistent with the first speakers of Anatolian IE coming to the region by way of commercial contacts and small-scale movement during the Bronze Age. Among comparative linguists, a Balkan route for the introduction of Anatolian IE is generally considered more likely than a passage through the Caucasus, due, for example, to greater Anatolian IE presence and language diversity in the west (73). Further discussion of these options is given in the archaeological and linguistic supplementary discussions (48, 49).

If you are asking yourselves why the Danish school (of Allentoft, Kristiansen, and Kroonen, co-authors of this paper) was not so fast to explain the findings the same way the proposed their infamous Indo-European – steppe ancestry association (i.e. ancestry = language, ergo CHG = PIE in this case), and resorted to mainstream anthropological models instead to explain the incongruence, I can think of two main reasons:

The possibility of having an early PIE around the Caucasus, potentially closely related not only to Uralic to the north, but also to Caucasian languages, Sumerian, Afroasiatic, Elamo-Dravidian, etc. could be a good reason for those excited with these few samples to begin dealing with macro-language proposals, such as Eurasiatic and Nostratic. If demonstrated to be true, a Northern Iranian origin of Middle PIE would also help relieve a little bit the pressure that some are feeling about the potentially male-driven Indo-European continuity (even if not “autochthonous”) associated with the expansion of R1b-L23 subclades.

On the other hand, I am a firm supporter of solid anthropological models of migration, and of “late and small” language expansions, usually accompanied by demic diffusion, which has been demonstrated to be linked with haplogroup expansion and reduction in variability.

Therefore, for the moment, even if it is weak – as weak as it always was (but still stronger than Gimbutas’ Maykop route) – the Balkan route seems like the best fit for all the data combined.

In fact, we already have steppe ancestry moving into the Lower Danube and Bulgaria in the mid-5th millennium. Let’s not forget that.

Yamna expansion to the East

Interesting data from an early East Yamna offshoot at Karagash, ca. 3018-2887 BC, of R1b-Z2106 lineage, which shows some ancestry, lineage, and cultural continuity in Sholpan, ca. 2620-2468 BC, in Kazakhstan.

This sample might be part of another descendant group from the migration waves that reached Afanasevo, and can thus be related to other early Asian R1b-L23 samples found in Narasimhan et al. (2018).

On the formation of Yamna and its CHG contribution, from the supplementary material:

  1. An admixture event, where Yamnaya is formed from a CHG population related to KK1 [=Kotias, dated ca. 7800 BC] and an ANE population related to Sidelkino and Botai. We inferred 54% of the Yamnaya ancestry to come from CHG and the remaining 46% to come from ANE.
  2. A split event, where the CHG component of Yamnaya splits from KK1. The model inferred this time at 27 kya (though we note the larger models in Sections S2.12.4 and S2.12.5 inferred a more recent split time [see below graphic]).
  3. A split event, where the ANE component of Yamnaya splits from Sidelkino. This was inferred at about about 11 kya.
  4. A split event, where the ANE component of Yamnaya splits from Botai. We inferred this to occur 17 kya. Note that this is above the Sidelkino split time, so our model infers Yamnaya to be more closely related to the EHG Sidelkino, as expected.
  5. An ancestral split event between the CHG and ANE ancestral populations. This was inferred to occur around 40 kya.
yamnaya-chg-ancestry
A 10-leaf model based on combining the models in Fig. S16 and Fig. S19 and re-estimating the model parameters.

On the expansion of domestication

CHG is not found in Botai, no gene flow from Yamna is found in its samples, and they are more related to East Asians, while Yamna is related to West Eurasians:

The lack of evidence of admixture between Botai horse herders and western steppe pastoralists is consistent with these latter migrating through the central steppe but not settling until they reached the Altai to the east (4). More significantly, this lack of admixture suggests that horses were domesticated by hunter-gatherers not previously familiar with farming, as were the cases for dogs (38) and reindeer (39). Domestication of the horse thus may best parallel that of the reindeer, a food animal that can be milked and ridden, which has been proposed to be domesticated by hunters via the “prey path” (40); indeed anthropologists note similarities in cosmological beliefs between hunters and reindeer herders (41). In contrast, most animal domestications were achieved by settled agriculturalists (5).

NOTE. I am not sure, but they seem to hint that there were separate events of horse domestication and horse-riding technique by the Botai and Yamna populations due to their lack of genetic contribution from the latter to the former. I guess they did not take into account farming spreading to the steppe without genetic contribution beyond the Dnieper… In fact, the superiority in horse-riding shown by the expanding Yamna peoples – as they state – should also serve to suggest from where the original technique expanded.

Indo-Iranian migrations

On the expansion of Yamna, and the different expansion of Steppe MLBA (with Indo-Iranian speakers) into Asia, further supporting Narasimhan et al. (2018), they have this to say:

However, direct influence of Yamnaya or related cultures of that period is not visible in the archaeological record, except perhaps for a single burial mound in Sarazm in present-day Tajikistan of contested age (44, 45). Additionally, linguistic reconstruction of proto-culture coupled with the archaeological chronology evidences a Late (-2300-1200 BCE) rather than Early Bronze Age (-3000-2500 BCE) arrival of the Indo-Iranian languages into South Asia (16, 45, 46). Thus, debate persists as to how and when Western Eurasian genetic signatures and IE languages reached South Asia.

Samples from the Namazga region (current Turkmenistan) from the Iron Age show an obvious influence from steppe MLBA (ca. 2300-1200 BC), and not steppe EBA (i.e. Yamna), population, in contrast with samples from the Chalcolithic (ca. 3300 BC), which don’t show this influence. This helps distinguish prior contacts with Iran Neolithic from the actual steppe population that expanded Indo-Iranian into Asia.

Very interesting therefore the Namazga CA sample (ca. 855 BC), of R1a-Z93 subclade, showing the sign of immigrant Indo-Aryans in the region. For more on this we will need an evaluation in common with the corrected data from Narasimhan et al. (2018), and all, including de Barros (Nature 2018), in combination with statistical methods to ascertain differences between early Indo-Aryans and Iranians.

namazga-expansion-south-asia
A summary of the four qpAdm models fitted for South Asian populations. For each modern South Asian population. we fit different models with qpAdm to explain their ancestry composition using ancient groups and present the f irst model that we could not reject in the following priority order: 1. Namazga_CA + Onge, 2. Namazga_CA + Onge + Late Bronze Age Steppe, 3. Namazga_CA + Onge + Xiongnu_lA (East Asian proxy). and 4. Turkmenistan_lA + Xiongnu_lA. Xiongnu_lA were used here to represent East Asian ancestry. We observe that while South Asian Dravidian speakers can be modeled as a mixture of Onge and Namazga_CA. an additional source related to Late Bronze Age steppe groups is required for IE speakers. In Tibeto-Burman and Austro-Asiatic speakers. an East Asian rather than a Steppe_MLBA source is required.

Siberian peoples and N1c lineages

We have already seen how the paper on Eurasian steppe samples tries to assign Uralic to Neolithic peoples east of the Urals. The association with Okunevo is unlikely, since most are of haplogroup Q1a2, but they seem to suggest (combining both papers) that they accompanied N lineages from Siberian hunter-gatherers (present e.g. in Botai or Shamanka II, during the Early Neolithic), and formed part of (or suffered from) different demic diffusion waves:

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.

NOTE. Also interesting to see no R1a in Baikal hunter-gatherers after ca. 3500 BC, and a prevalence of N lineages as supported in a previous paper on the Kitoi culture, which some had questioned in the past.

In fact, the only N1c1 sample comes from Ust’Ida Late Neolithic, 180km to the north of Lake Baikal, apparently before the expansion of Q1a2a lineages during the EBA period. While this sample may be related to those expanded later in Finno-Ugric territory (although it may only be related to those expanded much later with Yakuts), other samples are not clearly from those found widely distributed among North-East Europeans only after the Iron Age, or – as in the case of Shamanka II (N1c2), they are clearly not of the same haplogroup.

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

Wrap-up

It is great to see the paper and the supplementary material deal with Y-DNA haplogroups and their relevance for migrations with such detail. Especially because this paper comes from the same Copenhagen-based research group that originally associated ancestry with language, creating thus today’s mess based on steppe ancestry.

Regarding Y-DNA data, once again almost 100% of samples from late Khvalynsk/Yamna and derived cultures (like Afanasevo and Bell Beaker) are R1b-L23, no single R1a-M417 lineage found, and few expected by now, if any, within Late Proto-Indo-European territory.

While they claim to take Y-DNA into account to assess migrations – as they do for example with Asian cultures – , their previous model of a Yamna “R1a-R1b community” remains oddly unchanged, and they even insist on it in the supplementary materials, as they do in their parallel Nature paper.

They have also expressly mitigated the use of ancestral components to assess populations, citing the ancestral and modern association of CHG ancestry with different ethnolinguistic groups in the Middle East, to dismiss any rushed conclusions on the origin of Anatolian, and consequently of Middle PIE. And they did so evidently because it did not fit the anthropological data that is mainstream today (supporting a Balkan route), which is the right thing to do.

However, they have apparently not stopped to reconsider the links of CWC and steppe ancestry to ancestral and modern Uralic peoples – although they expressly mention the strong connection with modern Karelians in the supplementary material.

Also, after Narasimhan et al. (2018), there is a clear genetic continuity with East Yamna (in ancestry as in R1b-L23 subclades), so their interpretations about Indo-Iranian in this paper and especially de Barros (Nature 2018) – regarding the Abashevo -> Sintashta/Srunba/Andronovo connection – come, again, too late.

Related:

Eurasian steppe dominated by Iranian peoples, Indo-Iranian expanded from East Yamna

yamna-indo-iranian-expansion

The expected study of Eurasian samples is out (behind paywall): 137 ancient human genomes from across the Eurasian steppes, by de Barros Damgaard et al. Nature (2018).

Dicussion (emphasis mine):

Our findings fit well with current insights from the historical linguistics of this region (Supplementary Information section 2). The steppes were probably largely Iranian-speaking in the first and second millennia bc. This is supported by the split of the Indo-Iranian linguistic branch into Iranian and Indian33, the distribution of the Iranian languages, and the preservation of Old Iranian loanwords in Tocharian34. The wide distribution of the Turkic languages from Northwest China, Mongolia and Siberia in the east to Turkey and Bulgaria in the west implies large-scale migrations out of the homeland in Mongolia since about 2,000 years ago35. The diversification within the Turkic languages suggests that several waves of migration occurred36 and, on the basis of the effect of local languages, gradual assimilation to local populations had previously been assumed37. The East Asian migration starting with the Xiongnu accords well with the hypothesis that early Turkic was the major language of Xiongnu groups38. Further migrations of East Asians westwards find a good linguistic correlate in the influence of Mongolian on Turkic and Iranian in the last millennium39. As such, the genomic history of the Eurasian steppes is the story of a gradual transition from Bronze Age pastoralists of West Eurasian ancestry towards mounted warriors of increased East Asian ancestry—a process that continued well into historical times.

This paper will need a careful reading – better in combination with Narasimhan et al. (2018), when their tables are corrected – , to assess the actual ‘Iranian’ nature of the peoples studied. Their wide and long-term dominion over the steppe could also potentially explain some early samples from Hajji Firuz with steppe ancestry.
fku

eurasian-steppe-samples
Principal component analyses. The principal components 1 and 2 were plotted for the ancient data analysed with the present-day data (no projection bias) using 502 individuals at 242,406 autosomal SNP positions. Dimension 1 explains 3% of the variance and represents a gradient stretching from Europe to East Asia. Dimension 2 explains 0.6% of the variance, and is a gradient mainly represented by ancient DNA starting from a ‘basal-rich’ cluster of Natufian hunter-gatherers and ending with EHGs. BA, Bronze Age; EMBA, Early-to-Middle Bronze Age; SHG, Scandinavian hunter-gatherers.

For the moment, at first sight, it seems that, in terms of Y-DNA lineages:

  • R1b-Z93 (especially Z2124 subclades) dominate the steppes in the studied periods.
  • R1b-P312 is found in Hallstatt ca. 810 BC, which is compatible with its role in the Celtic expansion.
  • R1b-U106 is found in a West Germanic chieftain in Poprad (Slovakia) ca. 400 AD, during the Migration Period, hence supporting once again the expansion of Germanic tribes especially with R1b-U106 lineages.
  • A new sample of N1c-L392 (L1025) lineage dated ca. 400 AD, now from Lithuania, points again to a quite late expansion of this lineage to the region, believed to have hosted Uralic speakers for more than 2,000 years before this.
  • A sample of haplogroup R1a-Z282 (Z92) dated ca. 1300 AD in the Golden Horde is probably not quite revealing, not even for the East Slavic expansion.
  • Also, interestingly, some R1b(xM269) lineages seem to be associated with Turkic expansions from the eastern steppe dated around 500 AD, which probably points to a wide Eurasian distribution of early R1b subclades in the Mesolithic.

NOTE. I have referenced not just the reported subclades from the paper, but also (and mainly) further Y-SNP calls studied by Open Genomes. See the spreadsheet here.

Interesting also to read in the supplementary materials the following, by Michaël Peyrot (emphasis mine):

1. Early Indo-Europeans on the steppe: Tocharians and Indo-Iranians

The Indo-European language family is spread over Eurasia and comprises such branches and languages as Greek, Latin, Germanic, Celtic, Sanskrit etc. The branches relevant for the Eurasian steppe are Indo-Aryan (= Indian) and Iranian, which together form the Indo-Iranian branch, and the extinct Tocharian branch. All Indo-European languages derive from a postulated protolanguage termed Proto-Indo-European. This language must have been spoken ca 4500–3500 BCE in the steppe of Eastern Europe21. The Tocharian languages were spoken in the Tarim Basin in present-day Northwest China, as shown by manuscripts from ca 500–1000 CE. The Indo-Aryan branch consists of Sanskrit and several languages of the Indian subcontinent, including Hindi. The Iranian branch is spread today from Kurdish in the west, through a.o. Persian and Pashto, to minority languages in western China, but was in the 2nd and 1st millennia BCE widespread also on the Eurasian steppe. Since despite their location Tocharian and Indo-Iranian show no closer relationship within Indo-European, the early Tocharians may have moved east before the Indo-Iranians. They are probably to be identified with the Afanasievo Culture of South Siberia (ca 2900 – 2500 BCE) and have possibly entered the Tarim Basin ca 2000 BCE103.

The Indo-Iranian branch is an extension of the Indo-European Yamnaya Culture (ca 3000–2400 BCE) towards the east. The rise of the Indo-Iranian language, of which no direct records exist, must be connected with the Abashevo / Sintashta Culture (ca 2100 – 1800 BCE) in the southern Urals and the subsequent rise and spread of Andronovo-related Culture (1700 – 1500 BCE). The most important linguistic evidence of the Indo-Iranian phase is formed by borrowings into Finno-Ugric languages104–106. Kuz’mina (2001) identifies the Finno-Ugrians with the Andronoid cultures in the pre-taiga zone east of the Urals107. Since some of the oldest words borrowed into Finno-Ugric are only found in Indo-Aryan, Indo-Aryan and Iranian apparently had already begun to diverge by the time of these contacts, and when both groups moved east, the Iranians followed the Indo-Aryans108. Being pushed by the expanding Iranians, the Indo-Aryans then moved south, one group surfacing in equestrian terminology of the Anatolian Mitanni kingdom, and the main group entering the Indian subcontinent from the northwest.

steppe-migrations-pastoralists
Summary map. Depictions of the five main migratory events associated with the genomic history of the steppe pastoralists from 3000 bc to the present. a, Depiction of Early Bronze Age migrations related to the expansion of Yamnaya and Afanasievo culture. b, Depiction of Late Bronze Age migrations related to the Sintashta and Andronovo horizons. c, Depiction of Iron Age migrations and sources of admixture. d, Depiction of Hun-period migrations and sources of admixture. e, Depiction of Medieval migrations across the steppes.

2. Andronovo Culture: Early Steppe Iranian

Initially, the Andronovo Culture may have encompassed speakers of Iranian as well as Indo-Aryan, but its large expansion over the Eurasian steppe is most probably to be interpreted as the spread of Iranians. Unfortunately, there is no direct linguistic evidence to prove to what extent the steppe was indeed Iranian speaking in the 2nd millennium BCE. An important piece of indirect evidence is formed by an archaic stratum of Iranian loanwords in Tocharian34,109. Since Tocharian was spoken beyond the eastern end of the steppe, this suggests that speakers of Iranian spread at least that far. In the west of the Tarim Basin the Iranian languages Khotanese and Tumshuqese were spoken. However, the Tocharian B word etswe ‘mule’, borrowed from Iranian *atswa- ‘horse’, cannot derive from these languages, since Khotanese has aśśa- ‘horse’ with śś instead of tsw. The archaic Iranian stratum in Tocharian is therefore rather to be connected with the presence of Andronovo people to the north and possibly to the east of the Tarim Basin from the middle of the 2nd millennium BCE onwards110.

Since Kristiansen and Allentoft sign the paper (and Peyrot is a colleague of Kroonen), it seems that they needed to expressly respond to the growing criticism about their recent Indo-European – Corded Ware Theory. That’s nice.

They are obviously trying to reject the Corded Ware – Uralic links that are on the rise lately among Uralicists, now that Comb Ware is not a suitable candidate for the expansion of the language family.

IECWT-proponents are apparently not prepared to let it go quietly, and instead of challenging the traditional Neolithic Uralic homeland in Eastern Europe with a recent paper on the subject, they selected an older one which partially fit, from Kuz’mina (2001), now shifting the Uralic homeland to the east of the Urals (when Kuz’mina asserts it was south of the Urals).

Different authors comment later in this same paper about East Uralic languages spreading quite late, so even their text is not consistent among collaborating authors.

Also interesting is the need to resort to the questionable argument of early Indo-Aryan loans – which may have evidently been Indo-Iranian instead, since there is no way to prove a difference between both stages in early Uralic borrowings from ca. 4,500-3,500 years ago…

EDIT (10/5/2018) The linguistic supplement of the Science paper deals with different Proto-Indo-Iranian stages in Uralic loans, so on the linguistic side at least this influence is clear to all involved.

A rejection of such proposals of a late, eastern homeland can be found in many recent writings of Finnic scholars; see e.g. my references to Parpola (2017), Kallio (2017), or Nordqvist (2018).

NOTE. I don’t mind repeating it again: Uralic is one possibility (the most likely one) for the substrate language that Corded Ware migrants spread, but it could have been e.g. another Middle PIE dialect, similar to Proto-Anatolian (after the expansion of Suvorovo-Novodanilovka chiefs). I expressly stated this in the Corded Ware substrate hypothesis, since the first edition. What was clear since 2015, and should be clear to anyone now, is that Corded Ware did not spread Late PIE languages to Europe, and that some east CWC groups only spread languages to Asia after admixing with East Yamna. If they did not spread Uralic, then it was a language or group of languages phonetically similar, which has not survived to this day.

Their description of Yamna migrations is already outdated after Olalde et al. & Mathieson et al. (2018), and Narasimhan et al. (2018), so they will need to update their model (yet again) for future papers. As I said before, Anthony seems to be one step behind the current genetic data, and the IECWT seems to be one step behind Anthony in their interpretations.

At least we won’t have the Yamna -> Corded Ware -> BBC nonsense anymore, and they expressly stated that LPIE is to be associated with Yamna, and in particular the “Indo-Iranian branch is an extension of the Indo-European Yamnaya Culture (ca 3000–2400 BCE) to the East” (which will evidently show an East Yamna / Poltavka society of R1b-L23 subclades), so that earlier Eneolithic cultures have to be excluded, and Balto-Slavic identification with East Europe is also out of the way.

Related: