Climatic conditions in the Cis-Ural Steppe region and the Repin culture

cis-ural-region

New paper (behind paywall) Climate and Vegetation Changes over the Past 7000 Years in the Cis-Ural Steppe, by Khokhlova, Morgunova, Khokhlov, and Gol’eva, Eurasian Soil Sc. (2018) 51: 506.

Abstract (emphasis mine):

A multilayered archaeological site Turganik Settlement in the valley of the Tok River in the Cis- Ural steppe (Orenburg oblast) was examined with the use of paleopedological and microbiomorph methods. Ancient people inhabited this area in the Latest Neolithic (Eneolithic) (5th millennium BC) and Early Bronze (4th millennium BC) ages. It was found that cultural layers dating back to the Atlantic period of the Holocene had been formed under conditions of a predominance of grassy–forb vegetation with a small portion of tree species and dry climate; the ancient settlement was not affected by floods and was suitable for permanent living. It is probable that soils of the chestnut type with salinization and solonetzic features were developed in that time. The final stages of the accumulation of cultural layers were marked by strong shortterm floods, whose sediments partly masked the features of the previous long arid epoch. The highest degree of aridity was at the end of the Atlantic period. In the Subboreal and Subatlantic periods, soils of the meadowchernozemic type were formed; the spore–pollen spectra of these periods are characterized by a higher portion of tree species and by the presence of phytoliths of meadow grasses. The climatic conditions were generally colder and more humid, though some short-term aridization stages could take place. Some of these stages are recorded in the thickness of the studied sediments.

Interesting excerpts:

Paleosols buried under archaeological monuments of different periods represent a valuable archieve of information about the paleoenvironment. Most of the works in the field of archaeological pedology deal with earthen burial sites and kurgans [2, 5, 7, 9, 11]. Paleosols buried under the kurgans present us the paleoenvironmental records of a relatively short time before they were buried under the kurgan bodies. The study of kurgans created in different times makes it possible to characterize paleosol sequences storing information on longer periods of time in the second half of the Holocene. However, groups of kurgans that were consecutively created during the entire time span of kurgan construction, beginning from about 6000 yrs BC to the Early Medieval epoch, are few in number [8, 18, 22]. Even for such groups of kurgans, there are considerable time intervals that cannot be characterized because kurgans were not constructed during them. Hence, it is impossible to study the soils buried during these intervals. To reconstruct paleoenvironmental conditions for them, certain interpolation is required [29].

At the same time, there are archaeological sites — ancient settlements — the material of which gradually accumulated during very long time, e.g., beginning from the Middle Holocene to the present. Though such objects may also contain “missing layers” for certain periods, when the processes of denudation, erosion, or deflation predominated over sediment accumulation, they represent an almost continuous record on information about the paleoenvironmental conditions from the beginning of their functioning [25]. The most valuable among such sites are those that retain information on those periods of the Holocene that cannot be characterized on the basis of available data on the paleosols buried under the kurgans. In particular, this is the Atlantic period (7500–5000 yrs ago), because the construction of kurgans began only in the second half of this period (within the Volga–Ural interfluve [14]), and such ancient kurgans are rarely found. We studied such an archaeological site in the valley of the Tok River in the Cis-Ural steppe zone in Orenburg oblast. This site is known as the ancient Turganik settlement.

We studied different layers of the Turganik settlement with the use of a set of methods in order reconstruct the paleoenvironmental conditions (climate and vegetation) for the entire period of the accumulation of these sediments.

Thus, the Atlantic period of the Holocene in the Cis-Uralian steppe was characterized by dry climatic conditions with the driest stage during the Early Bronze Age (the Early Yamnaya culture of the middle of the fourth millennium BC). The Subboreal and Subatlantic periods were generally colder and wetter, though they also included short-term aridization phases, some of which were recorded in the sediment thickness.

This site is at the core of the interaction of Samara, Khvalynsk, and Repin cultures during the Eneolithic.

You can read more about it and the nature of Repin described by Morgunova (in favour of Gimbutas’ model), as combining traditions from Eneolithic steppe cultures from Khvalynsk to Sredni Stog, e.g. in Pottery from the Volga area in the Samara and South Urals region from Eneolithic to Early Bronze Age (2015).

repin-khvalynsk
Eneolithic settlements (1–5, 7, 10–16, 20, 22–43, 48, 50), burial grounds (6, 8–9, 17–19, 21, 47, 49) and kurgans (44–46) of the steppe Ural-Volga region: 1 Ivanovka; 2 Turganik; 3 Kuzminki; 4 Mullino; 5 Davlekanovo; 6 Sjezheye (burial ground); 7 Vilovatoe; 8 Ivanovka; 9 Krivoluchye; 10–13 LebjazhinkaI-III-IV-V; 14 Gundorovka; 15–16 Bol. Rakovka I-II; 17–18 Khvalunsk I-II; 19 Lipoviy Ovrag; 20 Alekseevka; 21 Khlopkovskiy; 22 Kuznetsovo I; 23 Ozinki II; 24 Altata; 25 Monakhov I; 26 Oroshaemoe; 27 Rezvoe; 28 Varpholomeevka; 29 Vetelki; 30 Pshenichnoe; 31 Kumuska; 32 Inyasovo; 33 Shapkino VI; 34 Russkoe Truevo I; 35 Tsaritsa I-II; 36 Kamenka I; 37 Kurpezhe-Molla; 38 Istay; 39 Isekiy; 40 Koshalak; 41 Kara-Khuduk; 42 Kair-Shak VI; 43 Kombakte; 44 Berezhnovka I-II; 45 Rovnoe; 46 Politotdelskoe; 47 burial near s. Pushkino; 48 Elshanka; 49 Novoorsk; 50 Khutor Repin.

The migration model of Anthony (2007, 2015), who collaborated with this group, is a more precise description of how peoples from the east of the Don River (mainly Khvalynsk/Repin cultures) migrated to develop a greater Yamna community, with Repin-type material culture expanding east of the Urals (into Afanasevo) and west of the Don River (into previous Sredni Stog/Kvitjana territory) – which I followed for the Indo-European demic diffusion model (we have recent samples of other potential Khvalynsk/Repin-related migrations).

NOTE. I usually refer to this Khvalynsk/Repin migration in genomics as of ‘Khvalynsk migrants’, for simplicity purposes, given that the few samples we have are from Khvalynsk, and that cultural regions east of the Don are difficult to differentiate precisely. However, it remains to be seen if – as I proposed – there are genetic differences between Repin and Khvalynsk groups, especially regarding R1b-L23 subclades – I proposed mainly Z2103 for Khvalynsk, L51 for Repin, a difference which has not been confirmed for the moment in Afanasevo, probably of Pre-Tocharian dialect, an archaic Northern dialect of Late PIE.

Anthony’s model of Khvalynsk/Repin as Yamna forefathers is probably, as we are seeing in Yamna samples, the right interpretation of peoples behind pots, compared to Gimbutas’ general idea of expanding kurgans of the 1970s.

On the other hand, the alternative Russian school version of Yamna developing from a heterogeneous community of Khvalynsk-Sredni Stog-Lower Danube cultures is probably by now to be fully dismissed, in archaeology (as Morgunova says) as in genetics.

Related:

Steppe and Caucasus Eneolithic: the new keystones of the EHG-CHG-ANE ancestry in steppe groups

indo-uralic-ehg-chg-ane-ancestry

Some interesting excerpts from Wang et al. (2018):

An interesting observation is that steppe zone individuals directly north of the Caucasus (Eneolithic Samara and Eneolithic steppe) had initially not received any gene flow from Anatolian farmers. Instead, the ancestry profile in Eneolithic steppe individuals shows an even mixture of EHG and CHG ancestry, which argues for an effective cultural and genetic border between the contemporaneous Eneolithic populations in the North Caucasus, notably Steppe and Caucasus. Due to the temporal limitations of our dataset, we currently cannot determine whether this ancestry is stemming from an existing natural genetic gradient running from EHG far to the north to CHG/Iran in the south or whether this is the result of farmers with Iranian farmer/ CHG-related ancestry reaching the steppe zone independent of and prior to a stream of Anatolian farmer-like ancestry, where they mixed with local hunter-gatherers that carried only EHG ancestry.

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

Concerning the influences from the south, our oldest dates from the immediate Maykop predecessors Darkveti-Meshoko (Eneolithic Caucasus) indicate that the Caucasus genetic profile was present north of the range ~6500 BP, 4500 calBCE. This is in accordance with the Neolithization of the Caucasus, which had started in the flood plains of the great rivers in the South Caucasus in the 6th millennium BCE from where it spread to the West and Northwest Caucasus during the 5th millennium BCE9, 49. It remains unclear whether the local CHG ancestry profile (represented by Late Upper Palaeolithic/Mesolithic individuals from Kotias Klde and Satsurblia in today’s Georgia) was also present in the North Caucasus region before the Neolithic. However, if we take the Caucasus hunter-gatherer individuals from Georgia as a local baseline and the oldest Eneolithic Caucasus individuals from our transect as a proxy for the local Late Neolithic ancestry, we notice a substantial increase in Anatolian farmer-related ancestry. This in all likelihood is linked to the process of Neolithization, which also brought this type of ancestry to Europe. As a consequence, it is possible that Neolithic groups could have reached the northern flanks of the Caucasus earlier50 (Supplementary Information 1) and in contact with local hunter gatherers facilitated the exploration of the steppe environment for pastoralist economies. Hence, additional sampling from older individuals is needed to fill this temporal and spatial gap.

The newest paper of the Reich/Jena group has brought samples (probably) much nearer to the actual CHG and ANE contribution seen in Eneolithic steppe peoples than the previously available Kotias Klde, Satsurblia, Afontova Gora 3, or Mal’ta.

It is impossible to say without direct access to the samples, but it is very likely that we will soon be able to break down different gross contributions from groups similar to these Steppe/Caucasus Neolithic ancestral groups into the diverse Eneolithic cultures of the Pontic-Caspian steppe, and thus trace more precisely each of these cultures to their genetic (and thus ethnolinguistic) heirs.

qpgraph-eneolithic-steppe
Admixture Graph modelling of the population history of the Caucasus region. We started with a skeleton tree without admixture including Mbuti, Loschbour and MA1. We grafted onto this EHG, CHG, Globular_Amphora, Eneolithic_steppe, Maykop, and Yamnaya_Caucasus, adding them consecutively to all possible edges in the tree and retaining only graph solutions that provided no differences of |Z|>3 between fitted and estimated statistics. The worst match is |Z|=2.824 for this graph. We note that the maximum discrepancy is f4(MA1, Maykop; EHG, Eneolithic_steppe) = -3.369 if we do not add the 4% EHG ancestry to Maykop. Drifts along edges are multiplied by 1000 and dashed lines represent admixture.”

Some more representative samples from Eneolithic steppe, steppe-forest and forest zone cultures of Eastern Europe will probably help with the fine-scale structure of different Chalcolithic groups, especially the homeland of early Corded Ware groups.

These new samples seem another good reason (like the Botai and R1b-M73) to rethink the role of (what I assumed were) different westward Mesolithic Eurasian waves of expansion influencing the formation of an Indo-Uralic and Indo-European community in Eastern Europe, and return to the simpler idea of local contributions from North Caucasus and steppe peoples absorbed by expanding EHG-like groups.

Related:

East Bell Beakers, an in situ admixture of Yamna settlers and GAC-like groups in Hungary

indo-european-yamnaya-corded-ware

I wanted to repeat what I said last week in two different posts (see on the new Caucasus and Yamna Hungary samples, and on local groups in contact with Yamna settlers).

We already knew that expanding East Bell Beakers had received influence from a population similar to the available Globular Amphorae culture samples.

  1. Without Yamna settlers, but with Yamna Ukraine and East Bell Beaker samples, including an admixed Yamna Bulgaria sample (from Olalde & Mathieson 2017, and then with their Nature 2018 papers), the most likely interpretation was that Yamna settlers had received GAC ancestry probably during their migration through the Balkans, before turning into East Bell Beakers. However, some comments still supported that it was Corded Ware migrants the ones behind the formation of East Bell Beakers. I couldn’t understand it.
  2. Now we have (with Wang et al. 2018) Yamna settlers (identical to other Yamna groups and Afanasevo migrants) and GAC-like peoples coexisting with them in Hungary, with a Late Chalcolithic Yamna sample from Hungary showing a greater contribution from GAC. However, I still read discussions on Yamna settlers receiving GAC admixture from Corded Ware in Eastern Europe, from GAC in the Dnieper-Dniester area, in Budzhak/Usatovo, etc. I can’t understand this, either.
  3. I will post here the data we have, with the simplest maps and images showing the simplest possible model. No more long paragraphs.

    NOTE. All this data does not mean that this model is certain, especially because we don’t have direct access to the samples. But it is the simplest and most likely one. Sometimes 2+2=4. Even if it turns out later to be false.

    EDIT (30 MAY 2018): In fact, as I commented in the first post about these samples, there is a Yamna LCA/EBA sample probably from Late Yamna (in the North Pontic steppe, west of the Catacomb culture), which shows GAC-like contribution. However, this admixture is lesser than that of Hungary LCA/EBA1 sample, and both Yamna groups (Hungary and steppe) were probably already more sedentary, which also supports different contributions from nearby local GAC-like groups to each region, rather than maintained long-range internal genetic contributions from a single source near the steppe…

    indo-european-uralic-migrations-yamna-gac
    Yamna migrants ca. 3300-2600. Most likely site of admixture with GAC circled in red.
    yamna_bell_beaker
    Yamna – Bell Beaker migration according to Heyd (2007, 2012). Most likely site of admixture with GAC is marked by the evolution of Blue to Red color.
    PCA-yamna-hungary
    PCA results. Samples from Yamna Hungary are surrounded by red circles, GAC-like Hungarian groups surrounded by light brown (see below for ADMIXTURE data) Notice the most likely Yamna Hungary sample with GAC admixture clustering closely to CWC Esperstedt outlier, and thus to some East Bell Beaker samples. (d) shows these projected onto a PCA of 84 modern-day West Eurasian populations (open symbols).
    gac-like-hungary-yamnaya
    Modified image, with red rectangles surrounding (unreleased) Hungarian samples from Yamna and GAC-like groups. (c) ADMIXTURE results of relevant prehistoric individuals mentioned in the text (filled symbols)
    yamnaya-hungary-lca-eba
    Modified image, with red rectangles surrounding (unreleased) Yamna samples Notice greater GAC contribution to late Yamna Hungary sample. 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
    yamnaya-hungary-globular-amphora
    Modified table from Wang et al. (2018) Supplementary materials (in bold, Yamna and related samples; in red, newly reported samples). Notice greater GAC contribution to late Yamna Hungary sample. “Supplementary Table 18. P values of rank=1 and admixture coefficients of modelling the Steppe ancestry populations as a two-way admixture of the Eneolithic_steppe and Globular_Amphora using 14 outgroups. Left populations: Steppe cluster, Eneolithic_steppe, Globular Amphora Right populations: Mbuti.DG, Ust_Ishim.DG, Kostenki14, MA1, Han.DG, Papuan.DG, Onge.DG, Villabruna, Vestonice16, ElMiron, Ethiopia_4500BP.SG, Karitiana.DG, Natufian, Iran_Ganj_Dareh_Neolithic.”

    The CWC outlier from Esperstedt

    I already said that my initial interpretation of the Esperstedt outlier, dated ca. 2430 BC, as due to a late contribution directly from the steppe (i.e. from long-range contacts between late Corded Ware groups from Europe and late groups from the steppe) was probably wrong, seeing how (in Olalde et al. 2017) early East Bell Beaker samples from Hungary and Central Europe clustered closely to this individual.

    Now we see that fully ‘Yamnaya-like’ Yamna settlers lived in Hungary probably for two or three centuries ca. 2900-2600 BC, and the absorption of known (or unknown) Yamna vanguard groups found up to Saxony-Anhalt before 2600 BC would be enough to justify the genomic findings of this individual.

    An outlier it is, then. But probably from admixture with nearby Yamna-like people.

    olalde_pca
    Image modified by me, from Olalde et al. (2017). PCA of 999 Eurasian individuals. Marked is the Espersted Outlier.

    Related:

The R1b-L23/Late PIE expansions, and the ‘R1a – Indo-European’ association

indo-european-yamnaya-corded-ware

I wrote a series of posts at the end of 2017 / beginning of 2018, to answer the wrong assumptions I could read in forums and blogs since 2015.

I decided not to publish them then, seeing how many successive papers were confirming my Indo-European demic diffusion model in a (surprisingly) clear-cut way.

Nevertheless, because I keep reading the same comments no matter what gets published, even in mid-2018 – the latest ones in our Facebook page (“was haplogroup X Indo-European?”), and in this very blog (“I see it very difficult to link Bell Beaker with Balto-Slavic, when now Balto-Slavic people are strikingly R1a-dominated”); and because I see even more misunderstandings and personal attacks, I have decided to publish them.

This way I will be able to explain my “R1b-L23/Proto-Indo-Europeans” theory with simplistic maps (however badly I hate such maps when I find them on Google searches), and I will also have a page to redirect those who don’t want to dismiss the “R1a – Indo-European association”, instead of answering comments about this question each time they pop up…

Here you have the links to the posts – and also on the menu above (there is a lot of rambling, because they are from a period of less clear data on Yamna and Corded Ware; today I would have never written such long discussions, they are mostly unnecessary):

  1. Haplogroup is not language, but R1b-L23 expansion was associated with Proto-Indo-Europeans
  2. The history of the simplistic ‘haplogroup R1a — Indo-European’ association
  3. Tips for dialogue with those supporting the R1a/Indo-European association

Related:

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

invasion-from-the-steppe-yamnaya

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:

Immigration and transhumance in the Early Bronze Age Carpathian Basin

Interesting excerpts about local Hungarian groups that had close contacts with Yamna settlers in the Carpathian Basin, from the paper Immigration and transhumance in the Early Bronze Age Carpathian Basin: the occupants of a kurgan, by Gerling, Bánffy, Dani, Köhler, Kulcsár, Pike, Szeverényi & Heyd, Antiquity (2012) 86(334):1097-1111.

The most interesting of the local people is the occupant of grave 12, which is the earliest grave in the kurgan and the main statistical range of its radiocarbon date clearly predates the arrival of the western Yamnaya groups c. 3000 BC. This is also confirmed by the burial rite, which is not typical for the Yamnaya (Dani 2011: 29–33; Heyd in press), although some heterogeneity may apply in Yamnaya communities too. The migrant group, graves nos. 4, 7, 9 and 11, all occupy late stratigraphic positions in the mound, and have radiocarbon dates in the second quarter of the third millennium BC. It is also noteworthy that they are all adult or mature men. The contextual data, their physical distribution over the space of the whole kurgan, and the variety of burial practices, indicate several generations of burials. The cultural attributes of this group are summarised in Figure 5. Overall, their closest match lies in the Livezile group from the eastern and southern Apuseni Mountains, which is also the likely place of origin of the buried persons.

yamna-settlements-hungary
Cultural geography of the Carpathian Basin in the first half of the third millennium BC (in black: archaeological cultures and groups dating roughly to the first quarter; in red: those dating to the second quarter). Indicated also are regions and sites mentioned in the text.

The key question is, what cultural process could be responsible for attracting these men from their homeland to the Great Hungarian Plain, over several generations? Their sex and age uniformity indicate they are a social sub-set within a larger group, implying that only a portion of their society was on the move. Exogamy can probably be excluded, since one would expect more women than men to move in prehistoric times; not to mention the distance of more than 200km between the places of potential origin and burial.

One hypothesis would see these men involved in the exchange of goods, with long-term relations between the mountain and steppe communities. Normally living in, or next to, the Apuseni, these men would journey for weeks into the plain, returning to the same places and people over many decades. Ethnographic examples of such travels to exchange objects and ideas, and perhaps people, are numerous (e.g. Helms 1988). However, the child’s (grave 7a) local isotopic signature would remain unexplained, and one has to wonder for how many generations an exchange continues for four men to die near the Őrhalom.

A second hypothesis is essentially an economic model of transhumance, with livestock passing the winter and spring in the milder regions of the Great Hungarian Plain, and returning to higher pastures in the warmer months (Arnold & Greenfield 2006). Such systems can endure for centuries, provided the social relations underpinning them are stable. This has the advantage of accounting for relatively long periods of time spent away from home, as herdsmen guarded their animals, and perhaps some women and their children came too, which would account for the child’s presence, and the pottery relations of the Livezile group. Furthermore, regular visits to a region would increase the likelihood of Livezile transhumant herders becoming integrated locally. The second quarter of the third millennium BC was a period when Yamnaya ideology, and thus its internal coherence, might have already diminished. This would likely have resulted in a weakened grip by Yamnaya people on pastures and territory, consequently allowing Livezile herders, and potentially others, to step in and take over locally, perhaps first on a seasonal basis and then permanently.

On West Yamna settlers in Hungary

yamnaya-hungary-globular-amphora
Modified table from Wang et al. (2018) Supplementary materials (in bold, Yamna and related samples; in red, newly reported samples). “Supplementary Table 18. P values of rank=1 and admixture coefficients of modelling the Steppe ancestry populations as a two-way admixture of the Eneolithic_steppe and Globular_Amphora using 14 outgroups. Left populations: Steppe cluster, Eneolithic_steppe, Globular Amphora Right populations: Mbuti.DG, Ust_Ishim.DG, Kostenki14, MA1, Han.DG, Papuan.DG, Onge.DG, Villabruna, Vestonice16, ElMiron, Ethiopia_4500BP.SG, Karitiana.DG, Natufian, Iran_Ganj_Dareh_Neolithic.”

By disclosing very interesting information on (yet unpublished) Yamna samples from Hungary, the latest preprint from the Reich Lab has rendered irrelevant – in a rather surprising turn of events – (what I expected would be) future discussions on West Yamna settlers potentially sharing a similar ancestry with Baltic Late Neolithic / Corded Ware settlers (see here for more details).

Interesting excerpts regarding the tight cluster formed by all Yamna samples:

Individuals from the North Caucasian steppe associated with the Yamnaya cultural formation (5300-4400 BP, 3300-2400 calBCE) appear genetically almost identical to previously reported Yamnaya individuals from Kalmykia20 immediately to the north, the middle Volga region19, 27, Ukraine and Hungary, and to other Bronze Age individuals from the Eurasian steppes who share the characteristic ‘steppe ancestry’ profile as a mixture of EHG and CHG/Iranian ancestry23, 28. These individuals form a tight cluster in PCA space (Figure 2) and can be shown formally to be a mixture by significantly negative admixture f3-statistics of the form f3(EHG, CHG; target) (Supplementary Fig. 3).

Using qpAdm with Globular Amphora as a proximate surrogate population (assuming that a related group was the source of the Anatolian farmer-related ancestry), we estimated the contribution of Anatolian farmer-related ancestry into Yamnaya and other steppe groups. We find that Yamnaya individuals from the Volga region (Yamnaya Samara) have 13.2±2.7% and Yamnaya individuals in Hungary 17.1±4.1% Anatolian farmer-related ancestry (Fig.4; Supplementary Table 18)– statistically indistinguishable proportions.

yamna_bell_beaker
Yamna – Bell Beaker migration according to Heyd (2007, 2012)

Before this paper, we had the solidest anthropological models backed by Y-DNA against conflicting data from certain statistical tools applied to a few samples (which some used to contradict what was mainstream in Academia).

NOTE. I have discussed this extensively in this blog, and more than once. See for example my posts on R1a speaking IE (July 2017), on the Eneolithic Ukraine sample (September 2017), or on the “Yamnaya ancestral component” (November 2017).

Today, we have everything – including statistical tools – showing a genetically homogeneous, Late PIE-speaking late Khvalynsk/Yamna community expanding into its known branches, confirming what was described using traditional anthropological disciplines:

  • Late Khvalynsk expanding into Afanasevo ca. 3300-3000 BC with an archaic Late PIE dialect, which was attested much later as Tocharian;
  • East Yamna/Poltavka admixing with Uralic-speaking Abashevo migrants probably ca. 2600-2100 BC to form Proto-Indo-Iranian-speaking Sintashta-Petrovka and Potapovka;
  • and now also Yamna settlers: those in Hungary admixing (probably ca. 2800-2500 BC) with the local population to form North-West Indo-European-speaking East Bell Beakers; those from the Balkans forming other IE-speaking Balkan cultures, including the peoples that admixed in Greece, as seen in Mycenaeans.

If Volker Heyd is right with this and other papers – and he has been right until now in his predictions regarding Yamna, Bell Beaker, and Corded Ware cultures – , the change in ancestry will probably begin to be noticed in Yamna samples from Hungary and the Lower Danube during the second quarter of the 3rd millennium, a period defined by the addition of a more fashionable western Proto-Bell Beaker package to the fading traditional Yamna cultural package.

EDIT (19 MAY 2018): I corrected some sentences and added interesting information.

Related:

The unique elite Khvalynsk male from a Yekaterinovskiy Cape burial

Recent paper (behind paywall) The Unique Burial of the Ekaterinovsky Cape Early Eneolithic Cemetery in the Middle Volga Region, by Korolev et al. Stratum Plus (2018) Nº2.

Abstract (official, in English):

This is the first time we published the results of a comprehensive study of burial 45 of the eneolithic cemetery called Ekaterinovsky Cape. The burial contains the skeleton of a young man with traumatic injuries of the skull, leg and hand bones of other individuals, skeleton of a young specimen of a domestic goat (Capra hircus) that was abundantly sprinkled with red ocher. Grave goods include three stone scepters of different types, a large item made of horn in the shape of a bird’s head, a stone adze, knife-like plates of quartzite, beads from the flaps of the shells (Unio), marmot cutters, decoration made from a beaver’s tooth. The uniqueness of the burial is determined by the combination of the composition of the grave goods and traces of ritual practices. To conclude, we suggest the buried man could belong to the elite of the society that left this burial ground.

NOTE. About my terminology, Russian has a lenited pronunciation of E in this case, hence the “Ye-” transliteration of the name of the town (and the site) in Google as Yekaterinovka. The “more etymological” transliteration is with “E”, as they use here, although Russians paradoxically use phonetic transliterations of foreign terms. I prefer the lenited transliteration to distinguish the Russian site from other Ekaterinovkas, though.

ekaterinovsky
Schematic view from burial 45. Male of 20-25 years, ca. 4400-4200 BC.

Interesting excerpt (translated from Russian):

Perhaps, we should correlate three very closely related damages [on the skull] with certain rituals, with which scepters could be associated. Each scepter could be a symbolic expression of a part of society, a type of activity, reaching a certain age and social status. This assumption does not seem incredible in combination with other extant, no less impressive, details of the funeral rite. Of great interest is the ornithomorphic rod of the horn. The location of the wand in the head and right half of the breast emphasizes its special significance in ritual practice and in funeral rites. Direct analogies to this product in other burial places of the cemetery are absent, and outside it authors are not known.

NOTE. Although the paper is in Russian and is behind paywall, it is really cheap, and can be easily translated with Google Translate if you can’t read Russian, so – unlike usual papers from the big publishing companies – you could support the journal by paying for it. You can read more about this burial at Pikabu, too. Photos and text in that post are not the same as in the paper, though, so it seems that the author of the text got the information either directly or from another source.

On the genetic data

Here is what I could gather about the report I shared of R1b-L51 lineages in Samara:

1) Yes, the comment at MolGen.org contains a more or less accurate summary of the oral communication actually given. And no, no more interesting data – from a genetic point of view – was presented.

2) What A.A. Khokhlov reported was preliminary genetic information from some samples, and an outside lab shared this information with him.

NOTE. It is well-known that David Anthony, also part of the Samara Valley project, provided the Reich Lab with Khvalynsk and Yamna samples from the region, so it would not be a surprise that these had been in fact assessed by the Reich Lab, too. This is my assumption, though, and I may be wrong.

3) What the report conveys is that “all samples investigated” belonged to R1b-P312 and R1b-U106, so I understand there are in principle more than two samples, whatever Google Translate says.

4) As R. Rocca said in Anthrogenica, the reported R1b1a1a2a1a1c2b2b1a2 (U106 subclade) is exactly the same one reported in Narasimhan et al. (2018) for the sample from the Iron Age site Loebanr 1 (Swat proto-historic graves) ca. 950 BC.

NOTE. That would be another hint at the origin of the preliminary data, together with the timing of the report (January), probably coinciding with the final assessment of samples which appeared in Narasimhan et al. (2018). That would explain the similar weird Y-SNP calls from software yHaplo (as reported by Narasimhan in Twitter). This is all again conjecture, though.

R1b-P312 is not reported in Narasimhan et al. (2018) for any sample (that would be “R1b1a1a2a1b”, following the standard used in their tables). Because the V88 sample in Khvalynsk, as well as other previously known V88 samples, are correctly reported as within the V88 branch, we may be talking about anything in the R1b tree from L754 (xV88) on. Most likely at or beyond the subclade of the Zvejnieki sample of hg R1b1a1 (classified as of R1b1a1a2a1), i.e. from P297 on.

NOTE. Since R1b-Z2103 samples are correctly reported, it is unlikely that the reported samples are from this branch, either.

ekaterinovka-burial-45
Graphic reconstruction from the elite male of grave 45, by R. M. Galeev.

It is possible, then, that we will have haplogroup R1b-M269 or L23 instead of L51, after all, and there would be then no major corrections to be made, either to the estimated dates from McDonald or Yfull (with their current differences), or to my predictions for early and late Khvalynsk, Repin, and Yamna

NOTE. In fact, the appearance of R1b-M269* and/or L23* linked to expanding Khvalynsk could be the perfect end to the resurging theories on Armenian or Western European origin of this haplogroup.

5) The full official genetic data is expected within a year (precise date unknown), so unless someone knows of a related draft in the making (which could publish them earlier), I would keep my expectations low for an official confirmation of the precise subclade any time soon.

NOTE. The best likely proxy for the reported data, if the above assumptions on Y-SNP calls and the software used are correct, is therefore to check out – whenever the corrected tables are published – the samples in Narasimhan et al. (2018) now classified as of R1b1a1a2a1(-) subclades. Or to experiment with the software and available BAM files to see which ones give this result…

6) I don’t know if Khokhlov’s book on Samaran archaeology will contain a reference to the samples, but I doubt it could contribute much more to the genetic data.

The meaning of Yekaterinovka

Of course, the Yekaterinovskiy Cape burials are just a tiny sampling of the dozens of settlements known from Khvalynsk, and the known ones represent just a tiny part of the hundreds that the culture probably had while it developed for more than a thousand years. In that sense, you may say that it is statistically not significant.

Nevertheless, as Anthony’s team recently said when explaining the relevance of their findings at Radzorskoe, the potential implications of any discovery at any of the few studied sites are very important. In this case, by confirming that late Khvalynsk became dominated early by R1b-M269, as was later Yamna, and as were early Yamna offshoots like Afanasevo and Bell Beaker.

I really don’t have anything more to add, whether in comments or per email. That’s as much information and speculation as you can get from me (or from them, I guess). If you want more, you can write to the team members yourselves.

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.

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.

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