The genetic and cultural barrier of the Pontic-Caspian steppe – forest-steppe ecotone

steppe-forest-steppe-biomes

We know that the Caucasus Mountains formed a persistent prehistoric barrier to cultural and population movements. Nevertheless, an even more persistent frontier to population movements in Europe, especially since the Neolithic, is the Pontic-Caspian steppe – forest-steppe ecotone.

Like the Caucasus, this barrier could certainly be crossed, and peoples and cultures could permeate in both directions, but there have been no massive migrations through it. The main connection between both regions (steppe vs. forest-steppe/forest zone) was probably through its eastern part, through the Samara region in the Middle Volga.

The chances of population expansions crossing this natural barrier anywhere else seem quite limited, with a much less porous crossing region in the west, through the Dnieper-Dniester corridor.

A Persistent ecological and cultural frontier

It is very difficult to think about any culture that transgressed this persistent ecological and cultural frontier: many prehistoric and historical steppe pastoralists did appear eventually in the neighbouring forest-steppe areas during their expansions (e.g. Yamna, Scythians, or Turks), as did forest groups who permeated to the south (e.g. Comb Ware, GAC, or Abashevo), but their respective hold in foreign biomes was mostly temporary, because their cultures had to adapt to the new ecological environment. Most if not all groups originally from a different ecological niche eventually disappeared, subjected to renewed demographic pressure from neighbouring steppe or forest populations…

The Samara region in the Middle Volga may be pointed out as the true prehistoric link between forests and steppes (see David Anthony’s remarks), something reflected in its nature as a prehistoric sink in genetics. This strong forest – forest-steppe – steppe connection was seen in the Eurasian technocomplex, during the expansion of hunter-gatherer pottery, in the expansion of Abashevo peoples to the steppes (in one of the most striking cases of population admixture in the area), with Scythians (visible in the intense contacts with Ananyino), and with Turks (Volga Turks).

steppe-forest-steppe-europe
Simplified map of the distribution of steppes and forest-steppes (Pontic and Pannonian) and xeric grasslands in Eastern Central Europe (with adjoining East European ranges) with their regionalisation as used in the review (Northern—Pannonic—Pontic). Modified from Kajtoch et al. (2016).

Before the emergence of pastoralism, the cultural contacts of the Pontic region (i.e. forest-steppes) with the Baltic were intense. In fact, the connection of the north Pontic area with the Baltic through the Dnieper-Dniester corridor and the Podolian-Volhynian region is essential to understand the spread of peoples of post-Maglemosian and post-Swiderian cultures (to the south), hunter-gatherer pottery (to the north), TRB (to the south), Late Trypillian groups (north), GAC (south), or Comb Ware (south) (see here for Eneolithic movements), and finally steppe ancestry and R1a-Z645 with Corded Ware (north). After the complex interaction of TRB, Trypillia, GAC, and CWC during the expansion of late Repin, this traditional long-range connection is lost and only emerges sporadically, such as with the expansion of East Germanic tribes.

A barrier to steppe migrations into northern Europe

One may think that this barrier was more permeable, then, in the past. However, the frontier is between steppe and forest-steppe ecological niches, and this barrier evolved during prehistory due to climate changes. The problem is, before the drought that began ca. 4000 BC and increased until the Yamna expansion, the steppe territory in the north Pontic region was much smaller, merely a strip of coastal land, compared to its greater size ca. 3300 BC and later.

This – apart from the cultural and technological changes associated with nomadic pastoralism – justifies the traditional connection of the north Pontic forest-steppes to the north, broken precisely after the expansion of Khvalynsk, as the north Pontic area became gradually a steppe region. The strips of north Pontic and Azov steppes and Crimea seem to have had stronger connections to the Northern Caucasus and Northern Caspian steppes than with the neighbouring forest-steppe areas during the Upper Palaeolithic, Mesolithic, and Neolithic.

NOTE. We still don’t know the genetic nature of Mikhailovka or Ezero, steppe-related groups possibly derived from Novodanilovka and Suvorovo close to the Black Sea (which possibly include groups from the Pannonian plains), and how they compare to neighbouring typically forest-steppe cultures of the so-called late Sredni Stog groups, like Dereivka or partly Kvityana.

steppe-forest-steppe-migration-routes
Typical migration routes through European steppes and forest-steppes. Red line represents the persistent cultural and genetic barrier, with the latest evolution in steppe region represented by the shift from dashed line to the north. Arrows show the most common population movements. Modified from Kajtoch et al. (2016).

Despite the Pontic-Caspian steppes and forest-steppes neighbouring each other for ca. 2,000 km, peoples from forested and steppe areas had an obvious advantage in their own regions, most likely due to the specialization of their subsistence economy. While this is visible already in Palaeolithic and Mesolithic hunter-gatherers, the arrival of the Neolithic package in the Pontic-Caspian region incremented the difference between groups, by spreading specialized animal domestication. The appearance of nomadic pastoralism adapted to the steppe, eventually including the use of horses and carts, made the cultural barrier based on the economic know-how even stronger.

Even though groups could still adapt and permeate a different territory (from steppe to forest-steppe/forest and vice-versa), this required an important cultural change, to the extent that it is eventually complicated to distinguish these groups from neighbouring ones (like north-west Pontic Mesolithic or Neolithic groups and their interaction with the steppes, Trypillia-Usatovo, Scythians-Thracians, etc.). In fact, this steppe – forest-steppe barrier is also seen to the east of the Urals, with the distinct expansion of Andronovo and Seima-Turbino/Andronovo-like horizons, which seem to represent completely different ethnolinguistic groups.

As a result of this cultural and genetic barrier, like that formed by the Northern Caucasus:

1) No steppe pastoralist culture (which after the emergence of Khvalynsk means almost invariably horse-riding, chariot-using nomadic herders who could easily pasture their cows in the huge grasslands without direct access to water) has ever been successful in spreading to the north or north-west into northern Europe, until the Mongols. No forest culture has ever been successful in expanding to the steppes, either (except for the infiltration of Abashevo into Sintashta-Potapovka).

2) Corded Ware was not an exception: like hunter-gatherer pottery before it (and like previous population movements of TRB, late Trypillia, GAC, Comb Ware or Lublin-Volhynia settlers) their movements between the north Pontic area and central Europe happened through forest-steppe ecological niches due to their adaptation to them. There is no reason to support a direct connection of CWC with true steppe cultures.

3) The so-called “Steppe ancestry” permeated the steppe – forest-steppe ecotone for hundreds of years during the 5th and early 4th millennium BC, due to the complex interaction of different groups, and probably to the aridization trend that expanded steppe (and probably forest-steppe) to the north. Language, culture, and paternal lineages did not cross that frontier, though.

EDIT (4 FEB 2019): Wang et al. is out in Nature Communications. They deleted the Yamna Hungary samples and related analyses, but it’s interesting to see where exactly they think the trajectory of admixture of Yamna with European MN cultures fits best. This path could also be inferred long ago from the steppe connections shown by the Yamna Hungary -> Bell Beaker evolution and by early Balkan samples:

wang-yamna-connection
Prehistoric individuals projected onto a PCA of 84 modern-day West Eurasian populations (open symbols). Dashed arrows indicate trajectories of admixture: EHG—CHG (petrol), Yamnaya—Central European MN (pink), Steppe—Caucasus (green), and Iran Neolithic—Anatolian Neolithic (brown). Modified from the original, a red circle has been added to the Yamna-Central European MN admixture.

Related

On the origin of haplogroup R1b-L51 in late Repin / early Yamna settlers

steppe-eneolithic-migrations

A recent comment on the hypothetical Central European origin of PIE helped me remember that, when news appeared that R1b-L51 had been found in Khvalynsk ca. 4250-4000 BC, I began to think about alternative scenarios for the expansion of this haplogroup, with one of them including Central Europe.

Because, if YFull‘s (and Iain McDonald‘s) estimation of the split of R1b-L23 in L51 and Z2103 (ca. 4100 BC, TMRCA ca. 3700 BC) was wrong, by as much as the R1a-Z645 estimates proved wrong, and both subclades were older than expected, then maybe R1b-L51 was not part of the Yamna expansion, but rather part of an earlier expansion with Suvorovo-Novodanilovka into central Europe.

That is, R1b-L51 and R1b-Z2103 would have expanded wih Khvalynsk-Novodanilovka migrants, and they would have either disappeared among local populations, or settled and expanded with successful lineages in certain regions. I think this may give rise to two potential models.

A hidden group in the European east-central steppes?

Here is what Heyd (2011), for example, has to say about the effect of the Khvalynsk-Novodanilovka expansion in the 4th millennium BC, with the first Kurgan wave that shuttered the social, economic, and cultural foundations of south-eastern Europe (before the expansion of west Yamna migrants in the region):

indo-european-anatolian-uralic-migrations
Proto-Anatolian migrations with Khvalynsk-Novodanilovka expansion, including ADMIXTURE data from Wang et al. (2018).

As the Boleraz and Baden tumuli cases in Serbia and Hungary demonstrate, there are earlier, 4th millennium cal. B.C. round tumuli in the Carpathian basin. There are also earlier north-Pontic steppe populations who infiltrated similar environments west of the Black Sea prior to the rise of the Yamnaya culture. This situation can be traced back to the 2nd half of the 5th millennium cal. B.C. to a group of distinct burials, zoomorphic maceheads, long flint blades, triangular flint points, etc., summarized under the term Suvurovo-Novodanilovka (Govedarica 2004; Rassamakin 2004; Anthony 2007; Heyd forthcoming 2011). They also erected round personalized tumuli, though smaller in size and height, above inhumations of single individuals. Suvorovo and Casimcea are the key examples in the lower Danube region of Romania. In northeast Bulgaria, the primary grave of Polska Kosovo (ochre-stained supine extended body position: information communicated by S. Alexandrov) can also be seen as such, as should the Targovishte-“Gonova mogila” primary grave 1 in the Thracian plain with a burial arranged in a supine position with flexed legs, southeast-northwest orientated, and strewed with ochre (Kanchev 1991 , p. 56- 57; Ivanova Gaydarska 2007). In addition to the many copper and shell beads, the 17.4cm long obsidian blade is exceptional, which links this grave to the Csongrád-“Kettoshalom” grave in the south Hungarian plain (Ecsedy 1979). It also yielded an obsidian blade ( 13.2cm long) and copper, shell and limestone beads.

suvorovo-novodanilovka-expansion-europe
The Southeast European distribution of graves of the Suvorovo-Novodanilovka group and such unequipped ones mentioned in the text which can be attributed by burial custom and stratigraphic position in the barrow, plus zoomorphic and abstract animal head sceptres as well as specific maceheads with knobs as from Decea Maresului (mid-5th millennium until around 4000 BC). Heyd (2016).

However, no traces of a tumulus have been recorded above the Kettoshalom tomb. Conventionally, it is dated to the Bodrogkeresztur-period in east Hungary, shortly after 4000 cal. B.C., which would correspond very well with the suggested Cernavodă I (or its less known cultural equivalent in the Thracian plain) attribution for the “Gonova mogila” grave, a cultural background to which the Csongrád grave should have also belonged. Bodrogkeresztur and Cernavodă I periods are not the only examples of 4th millennium cal. B.C. tumuli and burials displaying this steppe connection. Indeed we can find this early steppe impact throughout the 4th millennium cal. B.C. These include adscriptions to the Horodiștea II (Corlateni-Dealul Stadole, grave I: Burtanescu l 998, p. 37; Holbocai, grave 34: Coma 1998, p. 16); to Gordinești-Cernavodă 11 (Liești-Movila Arbănașu, grave 22: Brudiu 2000); to Gorodsk-Usatovo (Corlăteni Dealul Cetăţii, grave I: Comșa 1998, p. 17- 18, in Romania; Durankulak, grave 982: Vajsov 2002, in Bulgaria); and to Cernavodă III(Golyama Detelina, tum. 4: Leshtakov, Borisov 1995), and early (end of 4th millennium cal. B.C.) Ezero in Ovchartsi, primary grave (Kalchev 1994, p. 134-138) and Golyama Detelina, tum. 2 (Kanchev 1991) in Bulgaria. Also the Boleráz and Baden tumuli of Banjevac-Tolisavac and Mokrin in the south Carpathian basin account for this, since one should perhaps take into account primary grave 12 of the Sárrédtudavari-Orhalom tumulus in the Hungarian Alfold: a left-sided crouched juvenile ( 15- 17 y) individual in an oval, NW-SE orientated grave pit 14C dated to 3350-3100 cal. B.C. at 2 sigma (Dani, Ncpper 2006). Neither the burial custom (no ochre strewing or depositing a lump of ochre has been recorded), nor date account for its ascription to the Yamnaya!

All of these tumuli and burials demonstrate, though, that there is already a constant but perhaps low-level 4th millennium cal. B.C. steppe interaction, linking the regions of the north of the Black Sea with those of the west, and reaching deep into the Carpathian basin. This has to be acknowledged. even if these populations remain small, bounded to their steppe habitat with an economy adapted to this special environment, and are not always visible in the record. Indirect hints may help in seeing them, such as the frequent occurrence of horse bones, regarded as deriving from domesticated horses, in Hungarian Baden settlements (Bokonyi 1978; Benecke 1998), and in those of the south German Cham Culture (Matuschik 1999, p. 80-82) and the east German Bernburg Culture (Becker 1999; Benecke 1999). These occur, however, always in low numbers, perhaps not enough to maintain and regenerate a herd. Does this point us towards otherwise archaeologically hidden horsebreeders in the Carpathian basin, before the Yamnaya? In any case, I hope to make one case clear: these are by no means Yamnaya burials in the strict definition! Attribution to the Yamnaya in its strict definition applies.

pit-graves-central-europe
Distribution of Pit-Grave burials west of the Black Sea likely dating to the 2nd half of the 4th millennium BC (triangles: side-crouched burials; filled circles: supine extended burials; open circles: suspected). In Alin Frînculeasa, Bianca Preda, Volker Heyd, Pit-Graves, Yamnaya and Kurgans along the Lower Danube.

Also, about the expansion of Yamna settlers along the steppes:

However, it should have been made clear by the distribution map of the Western Yamnaya that they were confining themselves solely to their own, well-known, steppe habitat and therefore not occupying, or pushing away and expelling, the locally settled farming societies. Also, living solely in the steppes requires another lifestyle, and quite different economic and social bases, most likely very different to the established farming societies. Although surely regarded as incoming strangers, they may therefore not have been seen as direct competitors. This argument can be further enforced when remembering that the lowlands and the steppes in the southeast of Europe had already been populated throughout the 4th millennium cal. B.C., as demonstrated above, by societies with a similar north-Pontic steppe origin and tradition, albeit in lower numbers. It is only for these groups that the Yamnaya may have become a threat, but their common origin and perhaps a similar economic/ social background with comparable lifestyles would surely have assisted to allow rapid assimilation. More important, though, is that farming societies in this region may therefore have been accustomed to dealing and interacting with different people and ethnic strangers for a long time. (…)

When assessing farming and steppe societies’ interaction from a general point of view, attitudes can diverge in three main directions:

  1. the violent one; with raids, fights, struggles, warfare, suppression and finally the superiority and exploitation of the one over the other;
  2. the peaceful one; with a continuous exchange of gifts, goods, work, information and genes in a balanced reciprocal system, leading eventually to the merging of the two societies and creation of a new identity;
  3. the neutral one; with the two societies ignoring each other for a long time.

What we see from trying to understand the record of the Yamnaya, based on their tumuli and burials, and the local and neighbouring contemporary societies, based on their settlements, hoards, and graves, is likely a mixture of all three scenarios, with the balance perhaps more towards exchange in a highly dynamic system with alterations over time. However, violence and raids cannot be ruled out; they would be difficult to see in the archaeological record; or only indirectly, such as the building of hill forts, particularly the defence-like chain of Vucedol hillforts along the south shore of the Danube on the Serbian/Croatian border zone (Tasic 1995a), and the retreat of people into them (Falkenstein 1998, p. 261-262), with other interpretations also possible. And finally, we are dealing here with very different local and neighbouring societies, as well as with more distant contemporary ones, looking, in reality, rather like a chequer board of societies and archaeological cultures (see Parzinger 1993 for the overview). These display different regional backgrounds and traditions leading to different social and settlement organizations, different economic bases and material cultures in the wide areas between Prut and Maritza rivers, and Black Sea and Tisza river. They surely found their individual way of responding to the incoming and settling Yamnaya people.

yamna-tumuli-west-carpathians
Yamnaya tumuli signalling the expansion of West Yamna from ca. 3100 BC (especially after ca. 2950 BC). Heyd (2011).

The best data we have about this potential non-Yamna origin of R1b-L51 – and thus in favour of its admixture in the Carpathian basin – lies in:

  1. The majority of R1a-Z2103 subclades found to date among Yamna samples.
  2. The presence of R1b-Z2103 in the Catacomb culture – in the Northern Caucasus and in Ukraine.
  3. The limited presence of (ancient and modern) R1b-L51 in eastern Europe and India, whose isolated finds are commonly (and simplistically) attributed to ‘late migrations’.
  4. The presence of R1b-L51 (xZ2103) in cultures related to the ‘Yamna package’, but supposedly not to Yamna settlers. So for example I7043, of haplogroup R1b-L151(xU106,xP312), ca. 2500-2200 BC from Szigetszentmiklós-Üdülősor, probably from the Bell Beaker (Csepel group), but maybe from the early Nagýrev culture.
  5. The expansion of its subclades apparently only from a single region, around the Carpathian basin, in contrast to R1b-Z2103.
  6. The already ‘diluted’ steppe admixture found in the earliest samples with respect to Yamna, which points to the appearance after the Yamna admixture with the local population.
  7. Ukrainian archaeologists (in contrast to their Russian colleagues) point to the relevance of North Pontic cultures like Kvitjana and Lower Mikhailovka in the development of Early Yamna in the west, and some eastern European researchers also believe in this similarity.
  8. If R1b-Z2103 and R1b-L51 had expanded with Suvorovo-Novodanilovka migrants to the west, and had admixed later as Hungary_LCA-LBA-like peoples with Yamna migrants during the long-term contacts with other ‘kurganized cultures’ ca. 2900-2500 BC in the Great Hungarian Plains, it could explain some peculiar linguistic traits of North-West Indo-European, and also why R1b-Z2103 appears in cultures associated with this earlier ‘steppe influence’ (i.e. not directly related to Yamna) such as Vučedol (with a R1b-Z2103 sample, see below). That could also explain the presence of R1b-L151(xP312, xU106) in similar Balkan cultures, possibly not directly related to Yamna.
PCA-r1b-l51
Image modified from Wang et al. (2018). PCA of ancient and modern samples. Red circle in dashed line around Varna, Greece Neolithic, and (approximate position of) Smyadovo outliers, part of Khvalynsk-Novodanilovka settlers.

A hidden group among north or west Pontic Eneolithic steppe cultures?

The expansion of Khvalynsk as Novodanilovka into the North Pontic area happened through the south across the steppe, near the coast, with the forest-steppe region working as a clear natural border for this culture of likely horse-riding chieftains, whose economy was probably based on some rudimentary form of mobile pastoralism.

Although archaeologists are divided as to the origin of each individual Middle Eneolithic group near the Black Sea after the end of the Khvalynsk-Novodanilovka period, it seems more or less clear that steppe cultures like Cernavodă, Lower Mikhailovka, or Kvitjana are closer (or “more archaic”) in their steppe features, which connects them to Volga–Ural and Northern Caucasus cultures, like Northern Caucasus, Repin or Khvalynsk.

On the other hand, forest-steppe cultures like Dereivka (including Alexandria) show innovative traits and contacts with para- or sub-Neolithic cultures to the north, like Comb-Pit Ware groups, apart from corded decoration influenced by Trypillian groups to the west, especially in their later (‘Proto-Corded Ware‘) stage after ca. 3500 BC.

If Ukrainian researchers like Rassamakin are right, Early Yamna expanded not only from Repin settlers, but also from local steppe cultures adopting Repin traits to develop an Early Yamna culture, similar to how eastern (Volga–Ural groups) seem to have synchronously adopted Early Yamna without massive affluence of Repin settlements.

Furthermore, local traits develop in southern groups, like anthropomorphic stelae (shared with Kemi-Oba, direct heir of Lower Mikhailovka), and rich burials featuring wagons. These traits are seen in west Yamna settlers.

north-pontic-kvityana-dereivka-repin
Modified from Rassamakin (1999), adding red color to Repin expansion. The system of the latest Eneolithic Pointic cultures and the sites of the Zhivotilovo-Volchanskoe type: 1) Volchanskoe; 2) Zhivotilovka; 3) Vishnevatoe; 4) Koisug.

Problems of this model include:

  1. On the North Pontic area – in contrast to the Volga–Ural region – , there was a clear “colonization” wave of Repin settlers, also supported by Ukrainian researchers, based on the number of new settlements and burials, and on the progressive retreat of Dereivka, Kvitjana, as well as (more recent) Maykop- and Trypillia-related groups from the North Pontic area ca. 3350/3300 BC. It seems unlikely that these expansionist, semi-nomadic, cattle-breeding, patrilineally-related steppe clans that were driving all native populations out of their territories suddenly decided, at some point during their spread into the North Pontic area ca. 3300-3100 BC, to join forces with some foreign male lineages from the area, and then continue their expansion to the west…
  2. Similar to the fate of R1b-P297 subclades in the Baltic after the expansion of Corded Ware migrants, previous haplogropus of the North Pontic region – such as R1a, R1b-V88, and I2 subclades basically disappeared from the ancient DNA record after the expansion of Khvalynsk-Novodanilovka, and then after the expansion of Yamna, as is clear from Yamna, Afanasevo, and Bell Beaker samples obtained to date. This, in combination with what we know about Y-chromosome bottlenecks in post-Neolithic expansions, leaves little space to think that a big enough territorial group with a majority of “native” haplogroups could survive later expansions (be it R1b-L51 or R1a-Z645).
  3. Supporting an expansion of the same male (and partly female) population, the Yamna admixture from east to west is quite homogeneous, with the only difference found in (non-significant) EEF-like proportion which becomes elevated in distant areas [apart from significant ‘southern’ contribution to certain outlier samples]. Based on the also homogeneous Y-DNA picture, the heterogeneity must come, in general, from the female exogamy practiced by expanding groups.
  4. There is a short period, spanning some centuries (approximately 3300-2700 BC), in which the North Pontic area – especially the forest-steppe territories to the west of the Dnieper, i.e. the Upper Dniester, Boh, and Prut-Siret areas – are a chaos of incoming and emigrating, expanding and shrinking groups of different cultures, such as late Trypillian groups, Maykop-related traits, TRB, GAC, (Proto-)Corded Ware, and Early Yamna settlements. No natural geographic frontier can be delimited between these groups, which probably interacted in different ways. Nevertheless, based on their cultural traits, admixture, and especially on their Y-DNA, it seems that they never incorporated foreign male lineages, beyond those they probably had during their initial expansion trends.
  5. The further expansionist waves of Early Yamna seen ca. 3100 BC, from the Danube Delta to the west, give an overall image of continuously expanding patrilineal clans of R1b-M269 subclades since the Khvalynsk-Novodanilovka migration, in different periodic steps, mostly from eastern Pontic-Caspian nuclei, usually overriding all encountered cultures and (especially male) populations, rather than showing long-term collaboration and interaction. Such interaction is seen only in exceptional cases, e.g. the long-term admixture between Abashevo and Poltavka, as seen in Proto-Indo-Iranian peoples and their language.
PCA-Ukraine-r1b-l51
Image modified from Wang et al. (2018). PCA of ancient and modern samples. Arrows depicting Khvalynsk -> Yamna drift (blue), and hypothetic approximate Ukraine Eneolithic -> Yamna drift accompanying R1b-L51 (red).

Consequences

We are living right now an exemplary ego-, (ethno-)nationalism-, and/or supremacy-deflating moment, for some individuals of eastern and northern European descent who believed that R1a or ‘steppe ancestry proportions’ meant something special. The same can be said about those who had interiorized some social or ethnolinguistic meaning for the origin of R1b in western Europe, N1c in north-eastern Europe, as well as Greeks, Iranians, Armenians, or Mediterranean peoples in general of ‘Near Eastern’ ancestry or haplogroups, or peoples of Near Eastern origin and/or language.

These people had linked their haplogroups or ancestry with some fantasy continuity of ‘their’ ancestral populations to ‘their’ territories or languages (or both), and all are being proven wrong.

Apart from teaching such people a lesson about what simplistic views are useful for – whether it is based on ABO or RH group, white skin, blond hair, blue eyes, lactase persistence, or on the own ancestry or Y-DNA haplogroup -, it teaches the rest of us what can happen in the near future among western Europeans. Because, until recently, most western Europeans were comfortably settled thinking that our ancestors were some remnant population from an older, Palaeolithic or Mesolithic population, who acquired Indo-European languages by way of cultural diffusion in different periods, including only minor migrations.

Judging by what we can see now among some individuals of Northern and Eastern European descent, the only thing that can worsen the air of superiority among western Europeans is when they realize (within a few years, when all these stupid battles to control the narrative fade) that not only are they the cultural ‘heirs’ of the Graeco-Roman tradition that began with the Roman Empire, but that most of them are the direct patrilineal descendants of Khvalynsk, Yamna, Bell Beaker, and European Bronze Age peoples, and thus direct descendants of Middle PIE, Late PIE, and NWIE speakers.

steppe-chalcolithic-migrations
Steppe-related migrations ca. 3100-2600 BC with tentative linguistic identification.

The finding of R1b-L51 and R1b-Z2103 among expanding Suvorovo-Novodanilovka chieftains, with pockets of R1b-L51 remaining in steppe-like societies of the Balkans and the Carpathian Basin, would have beautifully complemented what we know about the East Yamna admixture with R1a-Z93 subclades (Uralic speakers) ca. 2600-2100 BC to form Proto-Indo-Iranian, and about the regional admixtures seen in the Balkans, e.g. in Proto-Greeks, with the prevalent J subclades of the region.

It would have meant an end to any modern culture or nation identifying themselves with the ‘true’ Late PIE and Yamna heirs, because these would be exclusively associated with the expansion of R1b-Z2103 subclades with late Repin, and later as the full-fledged Late PIE with Yamna settlers to south-east and central Europe, and to the southern Urals. The language would have had then obviously undergone different language changes in all these territories through long-lasting admixture with other populations. In that sense, it would have ended with the ideas of supremacy in western Europe before they even begin.

The most likely future

However limited the evidence, it seems that R1b-L51 expanded with Yamna, though, based on the estimates for the haplogroups involved, and on marginal hints at the variability of L23 subclades within Yamna and neighbouring populations. If R1b-L51 expanded with West Repin / Early Yamna settlers, this is why they have not yet been found among Yamna samples:

steppe-eneolithic-migrations
Simplified map of Repin expansions from ca. 3500/3400 BC.
  • The subclade division of Yamna settlers needs not be 50:50 for L51:Z2103, either in time or in space. I think this is the simplistic view underlying many thoughts on this matter. Many different expanding patrilineal clans of L23 subclades may have been more or less successful in different areas, and non-Z2103 may have been on the minority, or more isolated relative to Z2103-clans among expanding peoples on the steppe, especially on the east. In fact, we usually talk in terms of “Z2103 vs. L51” as if
    1. these two were the only L23 subclades; and
    2. both had split and succeeded (expanding) synchronously;

    that is, as if there had not been multiple subclades of both haplogroups, and as if there had not been different expansion waves for hundreds of years stemming from different evolving nuclei, involving each time only limited (successful) clans. Many different subclades of haplogroups L23 (xZ2103, xL51), Z2103, and L51 must have been unsuccessful during the ca. 1,500 years of late Khvalynsk and late Repin-Early Yamna expansions in which they must have participated (for approximately 60-75 generations, based on a mean 20-25 years).

  • If we want to imagine a pocket of ‘hidden’ L51 for some region of the North Pontic or Carpathian region, the same can be imagined – and much more likely – for any unsampled territory of expanding late Repin/Early Yamna settlers from the Lower Don – Lower Volga region (probably already a mixed society of L51 and Z2103 subclades since their beginning, as the early Repin culture, ca. 3800 BC), with L51 clans being probably successful to the west.
  • The Repin culture expanded only in small, mobile settlements from the Lower Don – Lower Volga to the north, east, and south, starting ca. 3500/3400 BC, in the waves that eventually gave a rather early distant offshoot in the Altai region, i.e. Afanasevo. Starting ca. 3300 BC in the archaeological record, the majority of R1b-Z2103 subclades found to date in Afanasevo also supports either
    • a mixed Repin society, with Z2103-clans predominating among eastern settlers; or
    • a Repin society marked by haplogroup L51, and thus a cultural diffusion of late Repin/Early Yamna traits among neighbouring (Khvalynsk, Samara, etc.) groups of essentially the same (early Khvalynsk-Novodanilovka) genetic stock in the Volga–Ural region.

    Both options could justify a majority of Z2103 in the Lower Volga–Ural region, with the latter being supported by the scattered archaeological remains of late Repin in the region before the synchronous emergence of Early Yamna findings in the whole Pontic-Caspian steppe.

  • Most Z2103 from Yamna samples to date are from around 3100 BC (in average) onward, and from the right bank of the Lower Don to the east, particularly from the Lower Volga–Ural area (especially the Samara region), which – based on the center of expansion of late Repin settlers – may be depicting an artificially high Z2103-distribution of the whole Yamna community.
repin-expansion-khvalynsk-cultures
Repin expansion into the Volga–Ural region from ca. 3500/3400 BC. Map made by me based on maps and data from Morgunova (2014, 2016). Lopatino is marked with number 64.
  • Yamna sample I0443, R1b-L23 (Y410+, L51-), ca. 3300-2700 BCE from Lopatino II, points to an intermediate subclade between L23 and L51, near one of the supposed late Repin sites (based on kurgan burials with late Repin cultural traits) in the Samara region.
  • Other Balkan cultures potentially unrelated to the Yamna expansion also show Z2103 (and not only L51) subclades, like I3499 (ca. 2884-2666 calBC), of the Vučedol culture, from Beli Manastir-Popova zemlja, which points to the infiltration of Yamna peoples in other cultures. In any case, the appearance of R1b-L23 subclades in the region happens only after the Yamna expansion ca. 3100 BC, probably through intrusions into different neighbouring regions, if these Balkan cultures are not directly derived from Yamna settlements (which is probably the case of the Csepel Bell Beaker or early Nagýrev sample, see above).
  • The diversity of haplogroups found in or around the Carpathian Basin in Late Chalcolithic / Early Bronze Age samples, including L151(xP312, xU106), P312, U106, Z2103, makes it the most likely sink of Yamna settlers, who spread thus with expanding family clans of different R1b-L23 subclades.
  • Even though some Yamna vanguard groups are known to have expanded up to Saxony-Anhalt before ca. 2700 BC, haplogroup Z2103 seems to be restricted to more eastern regions, which suggests that R1b-L51 was already successful among expanding West Yamna clans in Hungary, which gave rise only later to expanding East Bell Beakers (overwhelmingly of L151 subclades). The source of R1b-L51 and L151 expansion over Z2103 must lie therefore in the West Yamna period, and not in the Bell Beaker expansion.
indo-european-uralic-migrations-yamna-gac
Yamna migrants ca. 3300-2600. Most likely site of admixture with GAC circled in red.
  • The R1b-Z2103 found in Poltavka, Catacomb, and to the south point to a late migration displacing the western R1b-L51, only after the late Repin expansion. This is also seen in the steppe ancestry and R1b-Z2103 south of the Caucasus, in Hajji Firuz, which points to this route as a potential source of the supposed “Earliest Proto-Indo-Iranian” (the mariannu term) of the Near East. A similar replacement event happened some centuries later with expanding R1a-Z93 subclades from the east wiping out haplogroup R1b-Z2103 from the Pontic-Caspian steppe.
  • Many ancient samples from Khvalynsk, Northern Caucasus, Yamna, or later ones are reported simply as R1b-M269 or L23, without a clear subclade, so the simplistic ‘Yamna–Z2103’ picture is not real: if one takes into account that Z2103 might have been successful quite early in the eastern region, it is more likely to obtain a successful Y-SNP call of a Z2103 subclade in the Volga–Ural region than a xZ2103 one.
  • There are some modern samples of R1b-L51 in eastern Europe and Asia, whose common simplistic attribution to “late expansions” is usually not substantiated; and also ancient R1b-L51 samples might be confirmed soon for Asia.
  • ‘Western’ features described by archaeologists for West Yamna settlers, associated with Kemi Oba and southern Yamna groups in the North Pontic area – like rich burials with anthropomorphic stelae and wagons – are actually absent in burials from settlers beyond Bulgaria, which does not support their affiliation with these local steppe groups of the Black Sea. Also, a mix with local traditions is seen accross all Early Yamna groups of the Pontic-Caspian steppe, and still genetics and common cultural traits point to their homogeneization under the same patrilineal clans expanding continuously for centuries. The maintenance of local traditions (as evidenced by East Bell Beakers in Iberia related to Iberian Proto-Beakers) is often not a useful argument in genetics, especially when the female population is not replaced.
yamna-settlers-hungary
Yamna settlers in the Great Pannonian Plain, showing only kurgans of Hungary ca. 2950-2500 BC. Yamna Hungary was one of the biggest West Yamna provinces. From Hórvath et al. (2013).

Conclusion

This is what we know, using linguistics, archaeology, and genetics:

  • Middle Proto-Indo-European expanded with Khvalynsk-Novodanilovka after ca. 4800 BC, with the first Suvorovo settlements dated ca. 4600 BC.
  • Archaic Late Proto-Indo-European expanded with late Repin (or Volga–Ural settlers related to Khvalynsk, influenced by the Repin expansion) into Afanasevo ca. 3500/3400 BC.
  • Late Proto-Indo-European expanded with Early Yamna settlers to the west into central Europe and the Balkans ca. 3100 BC; and also to the east (as Pre-Proto-Indo-Iranian) into the southern Urals ca. 2600 BC.
  • North-West Indo-European expanded with Yamna Hungary -> East Bell Beakers, from ca. 2500 BC.
  • Proto-Indo-Iranian expanded with Sintashta, Potapovka, and later Andronovo and Srubna from ca. 2100 BC.

It seems that the subclades from Khvalynsk ca. 4250-4000 BC were wrongly reported – like those of Narasimhan et al. (2018). However, even if they are real and YFull estimates have to be revised, and even if the split had happened before the expansion of Suvorovo-Novodanilovka, the most likely origin of R1b-L51 among Bell Beakers will still be the expansion of late Repin / Early Yamna settlers, and that is what ancient DNA samples will most likely show, whatever the social or political consequences.

The only relevance of the finding of R1b-L51 in one place or another – especially if it is found to be a remnant of a Middle PIE expansion coupled with centuries of admixture and interaction in the Carpathian Basin – is the potential influence of an archaic PIE (or non-IE) layer on the development of North-West Indo-European in Yamna Hungary -> East Bell Beaker. That is, more or less like the Uralic influence related to the appearance of R1a-Z93 among Proto-Indo-Iranians, of R1a-Z284 among Pre-Germanic peoples, and of R1a-Z282 among Balto-Slavic peoples.

I think there is little that ancient DNA samples from West Yamna could add to what we know in general terms of archaeology or linguistics at this point regarding Late PIE migrations, beyond many interesting details. I am sure that those who have not attributed some random 6,000-year-old paternal ancestor any magical (ethnic or nationalist) meaning are just having fun, enjoying more and more the precise data we have now on European prehistoric populations.

As for those who believe in magical consequences of genetic studies, I don’t think there is anything for them to this quest beyond the artificially created grand-daddy issues. And, funnily enough, those who played (and play) the ‘neutrality’ card to feel superior in front of others – the “I only care about the truth”-type of lie, while secretly longing for grandpa’s ethnolinguistic continuity – are suffering the hardest fall.

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:

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: