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:

The Indo-European demic diffusion model, and the “R1b – Indo-European” association

yamna_bell_beaker_cut

Beginning with the new year, I wanted to commit myself to some predictions, as I did last year, even though they constantly change with new data.

I recently read Proto-Indo-European homelands – ancient genetic clues at last?, by Edward Pegler, which is a good summary of the current state of the art in the Indo-European question for many geneticists – and thus a great example of how well Genetics can influence Indo-European studies, and how badly it can be used to interpret actual cultural events – although more time is necessary for some to realize it. Notice for example the distribution of ‘Yamnaya’ in 3000 BC, all the way to Latvia (based on the initial findings of Mathieson et al. 2017), and the map of 2000 BC with ‘Corded Ware’, both suggesting communities linked by admixture and unrelated to actual cultures.

Some people – especially those interested in keeping a simplistic picture of Europe, either divided into admixture groups or simplistic R1b-Vasconic / R1a-Indo-European / N1c-Uralic (or any combination thereof) – want (others) to believe that I am linking ‘Indo-Europeans’ with haplogroup R1b. That is simply not true. In fact, my model dismisses such simplistic identifications of the reconstructible proto-languages with any modern peoples, admixtures, or haplogroups.

vasconic-uralic
Simplistic Vasconic/R1b-Uralic/N1c distribution, and intruding Indo-European/R1a, according to Wiik.

The beauty of the model lies, therefore, precisely in that if you take any modern group speaking Indo-European languages, none can trace back their combination of language, admixture, and/or haplogroup to a common Indo-European-speaking people. All our ancestral lines have no doubt changed language families (and indeed cultures), they have admixed, and our European regions’ paternal lines have changed, so that any dreams of ‘purity’ or linguistic/cultural/regional continuity become absurd.

That conclusion, which should be obvious to all, has been denied for a long time in blogs and forums alike, and is behind the effort of many of those involved in amateur genetics.

Main linguistic aim

The main consequence of the model, as the title of the paper suggests, is that reconstructible Indo-European proto-languages expanded with people, i.e. with actual communities, which is what we can assert with the help of Genomics. From a personal (or ethnic, or political) point of view genomics is useless, but from an anthropological (and thus linguistic) point of view, genomics can be a very useful tool to decide between alternative models of language diffusion, which has given lots of headaches to those of us involved in Indo-European studies.

The demic diffusion theory for the three main stages of the proto-language expansion was originally, therefore, a dismissal of impossible-to-prove cultural diffusion models for the proto-language – e.g. the adoption of Late Proto-Indo-European by Corded Ware groups due to a patron-client relationship (as proposed by Anthony), or a long-lasting connection between cultures (as proposed by Kristiansen, and favoured by “constellation analogy” proponents like Clackson, who negated the existence of common proto-languages). It also means the acceptance of the easiest anthropological model for language change: migration and – consequently – replacement.

By the time of the famous 2015 papers, I had been dealing for some time with the idea that the shared features between Indo-Iranian and Balto-Slavic may have been due to a common substrate, and must have therefore had some reflection in genomic finds. The data on these papers, and the addition of a weak connection between Pre-Germanic and Balto-Slavic communities, together with their clearest genetic link – R1a-M417 subclades (especially European Z283) – made it still easier to propose a Corded Ware substrate, partially common to the three.

Allentoft Corded Ware
Allentoft et al. “Arrows indicate migrations — those from the Corded Ware reflect the evidence that people of this archaeological culture (or their relatives) were responsible for the spreading of Indo-European languages. All coloured boundaries are approximate.”

Before the famous 2015 papers (and even after them, if we followed their interpretation), we were left to wonder why the supposed vector of expansion of Indo-European languages, Corded Ware migrants – represented by R1a-Z645 subclades, and supposedly continued unchanged into modern populations in its ‘original’ ancestral territories, Balto-Slavic and Indo-Iranian – , were precisely the (phonetically) most divergent Indo-European languages – relative to the parent Late Indo-European proto-language.

My paper implied therefore the dismissal of an unlikely Indo-Slavonic group, as proposed by Kortlandt, and of a still less factible Germano-Slavonic, or Germano-Indo-Slavonic (?) group, as loosely implied by some in the past, and maybe supported in certain archaeological models (viz. Kristiansen or partially Anthony), and presently by some geneticists since their simplistic 2015 papers on “massive migrations from the steppe“, and amateur genetic fans with infinite pet theories, indeed.

A common Corded Ware substrate to Balto-Slavic and Indo-Iranian, and common also partially between Balto-Slavic and Germanic (as supported by Kortlandt, too, albeit with different linguistic connotations), would explain their common features. The Corded Ware culture (and Uralic, tentatively proposed by me as the group’s main language family) is a strong potential connection between them, further supported by phylogeography, too.

Other consequences

Interpretations in my paper help thus dismiss the simplistic Yamna -> Corded Ware -> Bell Beaker migration model implied with phylogeography in the 2000s, and revived again by geneticists and Kristiansen’s workgroup based on the famous 2015 papers, whereby – due to the “Yamnaya ancestral component” – the Yamna culture would have been composed of communities of R1a-M417 and R1b-M269 lineages which remained against all odds ‘related but separated’ for more than two thousand years, sharing a common unitary language (why? and how?), and which expanded from Yamna (mainly R1b-L23) into Corded Ware (mainly R1a-M417) and then into Bell Beaker (mainly R1b-L51), in imaginary migration waves whose traces Archaeology has not found, or Anthropology described, before.

While phylogeography (especially the distribution of ancient samples of certain R1b and R1a subclades) was the main genetic aspect I used in combination with Archaeology and Anthropology to challenge the reliability of the “Yamnaya ancestral component” in assessing migrations – and thus Kristiansen’s now-popular-again modified Kurgan model – , my main aim was to prove a recent expansion of Late Proto-Indo-European from the steppe, and a still more recent expansion of a common group of speakers of North-West Indo-European, the language ancestral to Italo-Celtic, Germanic, and probably Balto-Slavic (or ‘Temematic’, the NWIE substrate of Balto-Slavic, according to some linguists).

My arguments serve for this purpose, and modern distributions of haplogroups or admixture are fully irrelevant: I am ready to change my view at any time, regarding the role of any haplogroup, or ancestral component, archaeological data, or anthropological migration model, to the extent that it supports the soundest linguistic model.

proto-indo-european-stages
Stages of Proto-Indo-European evolution. IU: Indo-Uralic; PU: Proto-Uralic; PAn: Pre-Anatolian; PToch: Pre-Tocharian; Fin-Ugr: Finno-Ugric. The period between Balkan IE and Proto-Greek could be divided in two periods: an older one, called Proto-Greek (close to the time when NWIE was spoken), probably including Macedonian, and spoken somewhere in the Balkans; and a more recent one, called Mello-Greek, coinciding with the classically reconstructed Proto-Greek, already spoken in the Greek peninsula (West 2007). Similarly, the period between Northern Indo-European and North-West Indo-European could be divided, after the split of Pre-Tocharian, into a North-West Indo-European proper, during the expansion of Yamna to the west, and an Old European period, coinciding with the formation and expansion of the East Bell Beaker group.

Gimbutas’ old theory of sudden and recent expansion served well to support a real community of Proto-Indo-European speakers, as did later the Yamna -> Corded Ware -> Bell Beaker theory that circulated in the 2000s based on modern phylogeography, and as did later partially Anthony’s updated steppe theory (2007). On the other hand, Kristiansen’s long-lasting connections among north-west Pontic steppe cultures and Globular Amphorae and Trypillian cultures, did not fit well with a close community expanding rapidly – although recent genetic data on Trypillia and Globular Amphorae might be compelling him to improve his migration theory.

So, if data turns out to be not as I expect now, I will reflect that in future versions of the paper. I have no problem saying I am wrong. I have been wrong many times before, and something I am certain is that I am wrong now in many details, and I am going to be in the future.

If, for example, R1b-L23(xZ2105) is demonstrated to come from Hungary and not the steppe (as supported by Balanovsky) or R1a-M417 samples are proved to have expanded with West Yamna settlers (as recently proposed by Anthony, see below the Balto-Slavic question), I would support the same model from a linguistic point of view, but modified to reflect these facts. Or if a direct migration link is found in Archaeology from Yamna to Corded Ware, and from Corded Ware to Bell Beaker (as proposed in the 2015 papers), I will revise that too (again, see the image below). Or, if – as Lazaridis et al. (2017) paper on Minoans and Mycenaeans suggested – the Anatolian hypothesis (that is, one of the multiple ones proposed) turns out to be somehow right, I will support it.

calcolithic-expansion
My map of Late Proto-Indo-European expansion (A Grammar of Modern Indo-European, 2006), following Gimbutas and Mallory.

Haplogroups are the least important aspect of the whole model, they are just another data that has to be taken into account for a throrough explanation of migrations. It has become essential today because of the apparent lack of vision on the part of geneticists, who failed to use them to adjust their findings of admixture with findings of haplogroup expansions, favouring thus a marginal theory of long-lasting steppe expansion instead of the mainstream anthropological models.

Since many of these alternative scenarios seem less and less likely with each new paper, it is probably more efficient to talk about which developments are most likely to challenge my model.

Main points

My main predictions – based mostly on language guesstimates, archaeological cultures, and anthropological models of migration -, even with the scarce genomic data we had, have been proven right until know with new samples from Mathieson et al. (2017) and Olalde et al. (2017), among other papers of this past year. These were my original assumptions:

(1) A Middle Proto-Indo-European expansion defined by the appearance of steppe ancestry + reduction in haplogroup diversity and expansion of (mainly) R1b-M269 and R1b-L23 lineages;

(2) A Late Proto-Indo-European expansion defined by steppe ancestry + reduction in haplogroup diversity and expansion of (mainly) R1b-L23 subclades; and

(3) A North-West Indo-European expansion defined by steppe ancestry + reduction in haplogroup diversity and expansion of (mainly) R1b-L51 subclades.

The expansion of Corded Ware peoples, associated with steppe ancestry + reduction in haplogroup diversity and expansion of (mainly) R1a-Z645 subclades, represents thus a different migration, which is compatible with the different nature of the Corded Ware culture, unrelated to Yamna and without migration waves from one to the other (although there were certainly contacts in neighbouring regions).

As you can see, neither of the 3+1 expansion models imply that no other haplogroup can be found in the culture or regions involved (others have in fact been found, and still the models remain valid): these migrations imply a reduction of haplogroup diversity, and the expansion of certain subclades as is common in population expansions throughout history. While we all accept this general idea, some people have difficulties accepting just those cases not compatible with their dreams of autochthonous continuity.

Nevertheless, there are still voids in genetic investigation.

Controversial aspects

In my humble opinion, these are potential conflict periods and the most likely areas of change for the future of the theory:

1. When and how did R1b-M269 lineages become “chiefs” in the steppe?

Based on scarce data from Khvalynsk, it seems that during the Neolithic there were many haplogroups in the North Pontic and North Caspian steppes. A reduction to R1b-M269 subclades must have happened either just before or (as I support) during (the migrations that caused) the Suvorovo-Novodanilovka expansion among Sredni Stog, probably coinciding also with the expansion (or one of the expansions) of CHG ancestry (and thus the appearance of ‘Steppe component’ in the steppe). My theory was based initially on Anthony’s account and TMRCA of haplogroups of modern populations (both ca. 4200-4000 BC), but recent samples of the Balkans (R1b-M269 and steppe ancestry) seem to trace the population expansion some centuries back.

If my assessment is correct, then modern populations of haplogroup R1b-M269* and R1b-L23* in the Balkans probably reflect that ancient expansion, and samples related to Proto-Anatolian cultures in the Balkans will most likely be of R1b-M269 subclades and R1b-L23*. After admixture in the Balkans, posterior migrations of Anatolian languages into Anatolia might be associated with a different admixture component and haplogroups, we don’t have enough data yet.

If the haplogroup reduction and expansion in Khvalynsk happened later than the Suvorovo-Novodanilovka expansion, then we might find the expansion of Pre- or Proto-Anatolian associated with many different haplogroups, such as R1b (xM269), R1a, I, J, or G2, and more or less associated with steppe ancestry in the Balkans.

Another reason for finding such variety of haplogroups in ancient samples from the Balkans would be that this Khvalynsk group of “chiefs” traversed – and mixed with – the Sredni Stog population. Nevertheless, if we suppose homogeneity in haplogroups in Khvalynsk during the expansion, a high proportion of different haplogroups explained by admixture with the local population of Sredni Stog would challenge the whole “chief domination” explanation by Anthony, and we would have to return to the “different culture” theory by Rassamakin and potentially an older migration from Khvalynsk. In any case, both researchers show clear links of the Suvorovo-Novodanilovka phenomenon to Khvalynsk, and a differentiation with the surrounding Sredni Stog culture.

A less likely model would support the identification of the whole Eneolithic Pontic-Caspian steppe as a loose Indo-Hittite-speaking community, which would be in my opinion too big a territory and too loose a cultural bond to justify such a long-lasting close linguistic connection. This will probably be the refuge of certain people looking desperately for R1a-IE connections. However, the nature of the western steppe will remain distinct from Late Proto-Indo-European, which must have developed in the Yamna culture, so autochthonous continuity is not on the table anymore, in any case…

suvorovo-novodanilovka-region
Coexistence of the Varna-Gumelniţa culture and the Suvorovo phase of the sceptre-bearer communities. 1 — Fălciu; 2 — Fundeni-Lungoţi; 3 — Novoselskaja; 4 — Suvorovo; 5 — Casimcea; 6 — Kjulevča; 7 — Reka Devnja; 8 — Drama; 9 — Gonova mogila; 10 — Reževo; 11 — geographically separate Decea variant of the sceptre bearer group (after Govedarica, Manzura 2011: Abb. 5, adapted).

2. How did R1a-M417 (and especially R1a-Z645) haplogroups came to dominate over the Corded Ware cultures?

If I am right (again, based on TMRCA of modern populations), then it is precisely at the time of the potential expansion of Proto-Corded Ware from the Dnieper-Dniester forest, forest-steppe, and steppe regions, ca 3300-3000. Furholt’s recent radiocarbon analysis and suggestions of a Lesser Poland origin of the third or A-horizon, on which disparate archaeologists such as Anthony or Klejn rely now, seem to suggest also that Corded Ware was a cultural complex rather than a compact culture reflecting a migration of peoples – similar thus to the Bell Beaker complex.

This cultural complex interpretation of Corded Ware contrasts with the quite homogeneous late samples we have, suggesting clear migration waves in northern Europe, at least at some point in time, so Genomics will be a great tool to ascertain when and from where approximately did Corded Ware peoples expand. Right now, it seems that Eneolithic Ukraine populations are the closest to its origin, so the traditional interpretation of its regional origin by Kristiansen or Anthony remains valid.

3. How was Indo-Iranian adopted by Corded Ware invaders?

This is rather an anthropological question. We need reasonable models of founder effect/cultural diffusion necessary for that to happen – similar to the ones necessary to explain the arrival of N1c subclades into north-east Europe, or the arrival of R1b subclades in Basque/Iberian-speaking regions in south-west Europe. My description of potential events in the eastern steppe – based partially on Anthony – is merely a short sketch. Genomic data is unlikely to offer more than it does today (replacement of haplogroups, and gradually of some steppe component, by late Corded Ware groups in the steppe), but let’s see what new samples can contribute.

As for what some Indians – and other people willing to confront them – are looking for, regarding R1a-M417 and/or Indo-European origins in India, I don’t see the point, we already know a) that the origin of the expansion is in the steppe and b) that Hindu nationalist biggots will not accept results from research that oppose their views. I don’t expect huge surprises there, just more fruitless discussions (fomented by those who live from trolling or conspiracies)…

4. Yamna settlers from Hungary

Anthony’s new theory – and the nature of Balto-Slavic – hinges on the presence of R1a-M417 subclades (associated with later Corded Ware samples) in Yamna settlers of Hungary, potentially originally from the North Pontic area, where the oldest sample has been found.

My ‘modified’ version of Anthony’s new model (the only I deem just remotely factible) includes the expansion of a Proto-Corded Ware from Lesser Poland, but (given the overwhelming R1b found in East Bell Beaker), with R1a-M417 being associated with the region. How to explain this language change with objective data? Well, we have Bell Beaker expanding to these areas at a later time, so we would need to find R1b-L23 settlers in Lesser Poland, and then a resurge of R1a-M417 haplogroup. If not, resorting yet again to cultural diffusion Yamna “patrons” to Corded Ware “clients” of Lesser Poland would bring us to square one, now with the ‘steppe ancestry’ controversy included…

Since some Eastern Europeans are (for no obvious reason whatsoever) putting their hopes on that IE-R1a-CWC association, let’s hope some samples of R1a-M417 in Yamna or Hungary give them a break, so that they can begin accepting something closer to mainstream anthropological models. We could then work from there a Yamna-> Bell Beaker / North-West Indo-European association truce, and from there keep accepting that no single haplogroup from Yamna settlers is linked with modern languages, cultures or ethnic groups.

yamna-region
localization of Central-European funerary monuments with elements of the Pit Grave culture (after Bátora 2006);

5. How and when was Balto-Slavic associated with haplogroup R1a?

If we accept the Southern or Graeco-Aryan nature of Balto-Slavic with influence from an absorbed North-West Indo-European dialect, “Temematic” (as Kortlandt does), then Indo-Slavonic adopted in the steppe from Potapovka by Sintashta and Poltavka populations divided ca. 2000 BC into Indo-Iranian (migrating to the east with Andronovo), and Balto-Slavic (migrating westward with the Srubna culture). History from there is not straightforward, and it should follow Srubna, Thraco-Cimmerian, or other late expansions from cultures of the steppe.

On the other hand, if it is a Northern dialect related closely to Germanic and Italo-Celtic (in a North-West Indo-European group), then its origin has to be found in the initial expansion of East Bell Beakers, and its development into either the Únětice culture (of Balkan and thus potentially “Southern IE” influence), or the Mierzanowice-Nitra culture (of Corded Ware and thus potentially Uralic influence), or maybe from both, given the intermediate substrate found in Germanic and Balto-Slavic.

It is my opinion that the association of Balto-Slavic with haplogroup R1a is quite early after the East Bell Beaker expansion, probably initially with the subclade typically associated with West Slavic, R1a-M458. I have not much data to support this (apart from the most common linguistic model), just modern haplogroup distribution maps and common TMRCA, and highly hypothetical archaeological-anthropological models. Genetics will hopefully bring more data.

Let’s see also what information on ancient haplogroups we can obtain from the Tollense valley (already showing a close cluster with modern West Slavic populations) and steppe regions.

6. How did Germanic, Celtic, and Italic expand?

Germanic is probably the most interesting one. Following the expansion of R1b-L51 subclades (especially R1b-U106) and steppe ancestry (a confounding factor, with the previous expansion of R1a-Z284 subclades) in Scandinavia is going to be fascinating. Anthropological models already point to a linguistic and archaeological expansion of Pre-Germanic with Bell Beaker peoples.

The expansion of Celtic seems to be associated with chiefdoms, untraceable today in terms of haplogroups, and it seems thus different from previous expansions. New studies might tell how that happened, if it was actually in successive ways, as proposed, or maybe we don’t have enough data yet to reach conclusions.

We don’t know either how Italic expanded into the Italian Peninsula, or whether Latin expanded with peoples from Italy, if at all, or it was mostly a cultural diffusion event, as it seems.

Regarding Etruscan, while I think it is a controversy initiated based on fantastic accounts, and ignited with few finds of Middle Eastern ancestry (that seem logical from the point of view of regional contacts), it will be important for Italian linguists and archaeologists, also to accept the most likely scenario.

As for Palaeo-Hispanic languages, while steppe ancestry is found quite reduced in R1b-L51 subclades (after so many different expansions and admixture events since the departure from the steppe), their distribution from the Chalcolithic onwards and the resurgence of native haplogroups may serve to ascertain which Pre-Roman tribes were associated with the oldest regions where these subclades dominated. For that aim, a closer look at the developments in Aquitania and other pre-Roman Vasconic- and Iberian-speaking regions may shed some light on how founder effects might develop to leave the native language intact (in a case similar to the adoption of Indo-Iranian by post-Corded Ware Sinthastha and Potapovka in the eastern Pontic-Caspian steppe).

NOTE: Although mostly unrelated, linguistic questions may also be somehow altered with a change of migration models. For example, our current Corded Ware Substrate Hypothesis – strongly contested by Kortlandt and others – implies that Uralic was potentially the language spoken by Eneolithic Ukraine / Proto-Corded Ware peoples, therefore early Uralic languages were spoken by Corded Ware peoples, as a substrate for Germanic and Balto-Slavic, and Balto-Slavic and Indo-Iranian. If an Indo-Hittite branch different from Late PIE is accepted for Eneolithic Ukraine (thus suggesting a millennia-long cultural-historical community in the steppe), then the model still stands (e.g. Ger. and BSl. *-mos/-mus, as stated by Kortlandt, would correspond to the oldest morphological IE layer). As you can read in the different versions of our model, the different possibilities for the common substrate are stated, and the most likely one selected. But the most likely a priori option sometimes turns out to be wrong…

NOTE 2: You can comment whatever you want here, but I opened a specific thread in our forum if you want serious comments on the model to stuck and be further discussed.

Featured images: from the book Interactions, changes and meanings. Essays in honour of Igor Manzura on the occasion of his 60th birthday. Țerna S., Govedarica B. (eds.). 2016. Kishinev: Stratum Plus.

See also:

New Ukraine Eneolithic sample from late Sredni Stog, near homeland of the Corded Ware culture

eneolithic-europe

Just one day after publishing the draft of the Indo-European demic diffusion model, 3rd version, Mathieson et al. (2017) have updated some information in a new version of their article, including a new interesting sample from late Sredni Stog. It gives support to what I predicted, regarding the potential origin of the third Corded Ware horizon.

After my first version, findings in Olalde et al. (2017) and Mathieson et al. (2017) supported some of my predictions. Now after my third, their new data also supports another prediction. Because the model is based on solid linguistic and archaeological models. Here is an excerpt from the Indo-European demic diffusion model, 3rd ed. (pp. 55-56):

At the end of the Trypillian culture, herding/hunting trends intensified, and the agricultural system collapsed, with people moving to the steppe zone, as confirmed by the presence of numerous graves to the south (Rassamakin 1999). At the same time, the Trypillian world absorbed a foreign tradition related to materials of settlement sites of the Dnieper steppes – such as the late Sredni Stog culture –, like cord impressions and burial rites similar to the later Corded Ware culture, marking also the transformation of decors and changes in their interpretation (Palaguta 2007).

The similarity in burial rituals between Yamna and Corded Ware made Gimbutas define a common “Kurgan people”, whose relationship has also been long supported by Kristiansen (Kristiansen 1989; Kristiansen et al. 2017). An equivalence of both burial rites has been, however, rejected (Häusler 1963, 1978, 1983), and it is generally agreed that the Yamna culture did not expand to the north of the Tisza River.

The importance of horse exploitation in Deriivka, in the forest-steppe zone of the north Pontic region along the Dnieper region, during the Middle Eneolithic period (probably ca. 3700-3530 BC), suggests that horses played a significant role in the life of this Sredni Stog community (Anthony and Brown 2003). In its late period (ca. 4000-3500 BC), this culture had adopted corded ware pottery, and stone battle-axes.

However, this [sic] western steppe peoples were mainly hunters (Rassamakin 1999), and the ‘herding skill’ essential for wild horse domestication seems absent (Kuzmina 2003). All this has been confirmed with zooarchaeological evidence and new molecular and stable isotope results, suggesting an absence of horse domestication in territories of the late Sredni Stog culture in the north Pontic steppe (Mileto et al. 2017), before the advent of migrants from the Indo-European-speaking Repin culture.

The new sample described in Mathieson et al. (2017), dated ca. 4200 BC (but within a wide range, 5000-3500 BC) is from a site classified as of late Sredni Stog (although potentially from Post-Mariupol / Kvitjana), a culture of hunters who probably did not breed domesticated horses (even after the period of conquest and dominance of Suvorovo-Novodanilovka chiefs, from Indo-Hittite-speaking early Khvalynsk, who had domesticated horses), and – more importantly – is of R1a-M417 lineage, shows high so-called “Yamna component” in ADMIXTURE, and clusters among Corded Ware samples in PCA approximately a thousand years before this culture’s expansion. Information from the supplementary material:

An Eneolithic cemetery of the Sredny Stog II culture was excavated by D. Telegin in 1955-1957 near the village of Alexandria, Kupyansk district, Kharkov region on the left bank of the river Oskol. A total of 33 individuals were recovered. Based on craniometric analysis (I.Potekhina 1999) it was suggested that the Eneolithic inhabitants of Alexandria were not homogeneous and resulted from admixture of local Neolithic hunter-gatherers and early farmers, possibly Trypillian groups. We report genetic data from one individual: I6561

pca-mathieson_ukraine_eneolithic
PCA, Admixture of Ukraine Eneolithic and other samples from Mathieson et al. (2017)

Another individual from Eneolithic Ukraine (of R1b1 xM269 lineage) clusters quite closely with Neolithic samples from the Baltic, which points to the strong connection between both – southern and northern – regions of east-central Europe before the period of great Chalcolithic expansions, and the potential origin of the spread of R1b (xM269) lineages with the Corded Ware culture.

The so-called ‘Yamna component’ – an infamous name which, as expected, is turning out to be very wrong – has been found quite elevated in this sample, previous and completely unrelated to the Yamna culture and its expansion, and similar to the (later) Corded Ware samples. In fact, we are seeing that Corded Ware samples are actually clustering closely with east-central Europe (excluding the CWC outlier), and not with Yamna and other Indo-European-speaking steppe cultures.

pca-mathieson2-ukraine-eneolithic
‘Yamna component’ (in yellow) in the North-West Pontic Steppe and the Balkans, including Eneolithic Ukraine samples

It will be fun to see the mess that certain researchers have made (and will still make in the near future) of their findings coupled with the concept of “Yamna component”, when trying to describe the “proxy ancestral populations” of European Copper Age and Bronze Age cultures… Difficult times ahead for many, after the collapse of the simplistic Yamna -> Corded Ware -> Bell Beaker genetic model laid out since Haak et al. (2015) and Allentoft et al. (2015).

[EDIT 27 September 2017] Not directly related, but here is today’s interesting discussion on Twitter surrounding the ancestral populations of the “Yamnaya component”, for illustration of the discussions to come when this ancestry is divided into different, more precise, older (Neolithic) steppe components, and these in turn shown to contribute to different European and Asian Chalcolithic and Bronze Age cultures:

Given the variance found in the three samples from Eneolithic Ukraine (comparable to the variance found in east Bell Beaker samples), we may now be getting closer to the precise territory and culture where the Corded Ware culture might have formed, which cannot be much further from the Dnieper-Dniester region before the Yamna expansion to the west ca. 3300 BC, judging from the elevated steppe component.

It seems, because of the proximity of both cultures and the similar dates of their migrations, that the westward expansion of the Yamna culture may have indeed provided an important push (among some strong ‘pull’ forces) for peoples of the expansion of the Corded Ware culture.

eneolithic_Caucasus
Diachronic map of Eneolithic migrations in the Caucasus ca. 4000-3100 BC

It keeps being demonstrated that archaeologists like Anthony, Heyd, Mallory, etc. were right where others tried to interpret admixture based on few samples and their own imagination, without any knowledge (or interest) whatsoever about Archaeology or Linguistics.

So Genetics reinforces the solidest models of Archaeology and Linguistics? Professional academics being mostly right in their careful research, and amateur geneticists playing with software being wrong? Who would have thought… More and more papers help thus shut up naysayers who state (again and again) that new algorithms are here to revolutionise these academic fields.

As Heyd predicted more than 10 years ago, and as many pointed out in terms of linguistic influence (like Mallory or Prescott) the transformation of Yamna settlers into the east Bell Beaker culture, and this culture‘s spread into western and northern Europe, must be noticed in genetic investigation.

The expansion of peoples is known to be associated with the spread of a certain admixture component + the expansion and reduction in variability of a haplogroup (i.e. few male lineages are usually more successful during the expansion): Neolithic farmers from the Middle East expanding with haplogroup G2a; Natufian component (Levant hunter-gatherers or later, Neolithic farmers) and haplogroup E southward into Africa; CHG component expansion with haplogroup J; WHG expansion into east Europe with haplogroup R1b; etc.

There were (at least) two main expansion processes involving Proto-Indo-European: one causing the branching off of the language ancestral to Anatolian, and another during the spread of Late Indo-European dialects. Based on this, and on known archaeological models, I have predicted since the first version of the demic diffusion model:

  • Based on haplogroups found until then in Yamna (R1b-M269), Corded Ware (R1a-M417, especially Z645), and Bell Beaker (R1b-L151):
    • that mainly R1b-L23 (especially L51) lineages and more steppe admixture would be found in east Bell Beaker – confirmed some two months after my publication by Olalde et al. (2017);
    • and that mainly R1a-M417 (especially Z645) subclades will be found in Corded Ware samples.
  • Based on the finding of “Yamna component” in the Corded Ware culture: that this admixture must have come from somewhere else. I pointed out to eastern Europe, including the forest and forest-steppe zone especially in the natural continuum of the Dniester-Dnieper region. Especially after Mathieson et al. (2017), in my second and third versions of the model, I have more specifically suggested a southern origin in the region, nearer to where the CHG ancestry must have come from (the Caucasus and cultures formed in contact with it), according to mainstream archaeological data, i.e. cultures of the North Pontic steppe / steppe-forest. But of course, until more samples are available, more CHG ancestry in other cultures of the Forest Zone cannot be discarded.

For the vast majority of academics, more samples (regionally proportioned) are needed only from early Corded Ware, as we have from Bell Beaker: if they are (as expected) mostly R1a-M417, then everything is clear, and it will finally mean the end for the tiring, now almost ‘traditional’ association R1a – Proto-Indo-European. Some more samples from the potential homeland of the third Corded Ware horizon, most likely Ukraine (Podolia and Volynia regions), nearer to the time of the Corded Ware expansion, would also be great, to locate the actual ancestral population of Corded Ware migrants – recognisable by the main presence of haplogroup R1a-Z645 (formed ca. 3500 BC), and elevated “Yamna component” before the arrival of the Yamna culture…

If, however, early Corded Ware samples of R1b-L23 subclades are found in certain quantity, especially old samples from east-central Europe (excluding Yamna migrants along the Prut), the tricky question of Late Indo-European cultural diffusion will remain: Did Corded Ware peoples adopt a Late Indo-European language from clans of R1b-L23 lineages? That is what Kristiansen and Anthony have been betting for, a cultural diffusion, caused by:

  • A long-lasting contact, according to Kristiansen (1989,…,2017). He defends that Sredni Stog adopted the language – but obviously not the same culture – from the east, but that it is a genetic and cultural mix from Globular Amphora, Trypillia, and steppe cultures. This has been Kristiansen’s model for almost 30 years, and it follows Marija Gimbutas’ outdated theory of the “Kurgan people”.
  • A rapid change according to Anthony (2007). He associates the adoption of Pre-Germanic with the domination of Yamna chiefs over Usatovo people, and the adoption of Balto-Slavic by the people from (Corded Ware) Middle Dnieper group because of the technical superiority of neighbouring Yamna herders.

Linguistics, with the growing support of a North-West Indo-European group, points clearly to a European expansion of a community speaking the ancestral language of Italo-Celtic, Germanic, and probably Balto-Slavic. Archaeology, too, showed migration from Yamna only to south-eastern Europe (correcting Gimbutas’ Kurgan model) and later with east Bell Beaker mainly into central, western, and northern Europe.

Even Kristiansen admits that only after the arrival of Bell Beaker in Scandinavia was a linguistic community (i.e. Germanic) formed – although he places the center of gravity in Úněticean influence, and (yet again) a cultural diffusion event into the Danish Dagger period.

Because of more and more data contrasting with old theories, some have elected to develop weak, indemonstrable links, to keep supporting e.g. Gimbutas’ concept of “Kurgan people” in Archaeology, and a sudden, early expansion of all PIE dialects at once in Linguistics. It seems that, after so much fuss about the (misleading) ‘Yamna component’ concept – and so many far-fetched assumptions by amateur geneticists -, the Corded Ware connection will once again hinge on weak, indemonstrable cultural diffusion theories, be it ‘Kurgan peoples’ (including now, of course, Eneolithic cultures of Ukraine) or any culture from eastern Europe that will reveal some close samples to Corded Ware migrants, in terms of PCA, ADMIXTURE, or haplogroup.

So once we find mainly R1a-Z645 in more Corded Ware samples (and this haplogroup and more “Yamna component” in non-Yamna cultures of Eneolithic Ukraine, and potentially Poland or Belarus) we all may finally expect a peaceful acceptance of reality, at least in Genetics? Nope. No siree. Nein. Not then, not ever.

Why? Because some people want their paternal lineage to have lived in their historical region, and spoken their historical language, since time immemorial. It won’t matter if Archaeology, Linguistics, Genetics, etc. don’t support their claims: if they need to use some aspects of admixture, or haplogroups (or a combination of them) from carefully selected samples instead of looking at the whole picture; if they have to support that Indo-Europeans came from a culture different than Yamna, in- or outside of the steppe or forest-steppe, be it the Balkans, Anatolia, Armenia, or the Moon; if their proto-language should then come directly from Indo-Hittite, or from a Germano-Slavonic, or Indo-Slavonic, or Indo-Germanic group, or whatever invented dialectal branch necessary to fit their model, or if they have to support the ‘constellation analogy’ of Clackson, or thousands of years of development for each branch; etc. They will support whatever is necessary.

And this adaptation, obviously, has no end. It’s stupid, I know. But that’s how we are, how we think. We have seen that these sad trends continue no matter what, for decades, and not only regarding Indo-European. Some common examples include:

  • Indo-Aryan-speaking Indians defending an autochthonous origin of R1a and Indo-European; as well as the ‘opposite’ autochtonous continuity theory of Dravidian-speaking Indians (based on ASI ancestry, haplogroup R2, mtDNA haplogroup M, or whatever is at hand).
  • Western Europeans defending an autochthonous origin of the R1b haplogroup, with a Palaeolithic or Mesolithic origin, including the language, viz. the recent Indo-European from the Atlantic façade theories (in the Celtic from the West series, by Koch and Cunliffe); the now fading Palaeolithic Continuity Theory; and many other forgotten Eurocentric proposals; as well as the more recent informal hints of a central European/Balkan homeland based on the Villabruna cluster and south-eastern Mesolithic finds, which is at risk of being related to a Balkan origin of Proto-Indo-European
    • There is also the ‘opposite’ theory of the autochthonous origin of the Basques, including Proto-Iberians and potentially other peoples like Paleo-Sardinians, based on the previously popular Vasconic-Uralic hypothesis (and an ancient Europe divided into R1b and N1c1 haplogroups), which is still widely believed in certain regions.
  • Finno-Ugric speakers of N1c1 lineage defending an autochtonous origin of the lineage and language in eastern Europe.
  • Nordic speakers supporting the autochthonous nature of Germanic and haplogroup I1 to Scandinavia.
  • Armenian speakers delighted to see a proposal of Indo-European homeland in the Armenian highlands, be it supported by glottalic consonants, CHG ancestrty, R1b (xM269) or J lineages…
  • Greek speakers now willing to support continuity of haplogroup J as a ‘native’ Greek lineage, of people speaking Proto-Greek (and in earlier times PIE), because of two Minoan, and one Mycenaean samples found in Lazaridis et al. (2017).
  • Even Turks linking Yamna with the expansion of Turkic languages. That one is fun to read, almost like a parody for the rest – substituting “Indo-European” for “Turkic”.
  • For years, a lot of people – me included (at least since 2005) – believed, because of modern maps of R1a distribution, that R1a and Corded Ware are the vector of Indo-European languages. For those of us who don’t have any personal or national tie with this haplogroup, this notion has been easy to change with new data. For others, it obviously isn’t, and it won’t be.
R1a_distribution_Eurasia
Modern R1a distribution in Eurasia (Wikipedia, 2008). The typical, simplistic map we relied on in the 2000s to derive wrong conclusions based on Genetics, conclusions which are sadly still alive and kicking…

For all these people, a sample, result, or conclusion from any paper, just dubiously in favour, means everything, but a thousand against mean nothing, or can be reinterpreted to support their fantasies.

The Kossinnian autochthonous continuity” crap permeates this relatively new subfield of Human Evolutionary Genetics, as it permeated Indo-European studies (first Linguistics, then Archaeology) in its infancy. It seems to be a generalised human trend, no doubt related to some absurd inferiority complex, mixed with historical romanticism, a certain degree of chauvinism, and (falling in the eternal Godwin’s Law of our field) some outdated, childish notion of ‘supremacy’ linked with the expansion of the own language and people.

Such simplistic and popular models are also lucrative, judging by the boom in demand for DNA analysis, which companies embellish with modern fortune tellers (or fortune tellers themselves sell for a price), promising to ascertain your ‘ancestry proportions’ using automated algorithms, so that you don’t have to get lost in complex genetic data and prehistoric accounts, which can’t help you define your “ethnicity”…

Some just don’t want to realize that the spread of prehistoric languages (like Late Indo-European dialects) was a complex, non-uniform, stepped process, devoid of modern romantic concepts, which in genetic terms necessarily included later founder effects and cultural diffusions, so that no one can trace their haplogroup, lineage, family, region, or country to any single culture, language, or ethnic group. The same, by the way, can be said of peoples and countries in historic times.

As I said before, we shall expect supporters of the Kurgan model (and thus the expansion of R1a-Z645 with Yamna) to wait for just one sample of R1a-M417 in Yamna and/or Bell Beaker (which will eventually be found), and just one sample of R1b-M269 in Corded Ware (which will also eventually be found), to blow the horn of victory in this naïve competition against time, general knowledge, and (essentially) themselves.

A sad consequence of how we are is that, because of the obvious influence of these stupid modern ethnolinguistic agendas, because we are not all rowing in the same direction, genetic results and conclusions are still perceived as far-fetched and labile, and thus most archaeologists and linguists prefer not to include genetic results in their investigation. And those who dare to do so, are badly counselled by those who go with the tide, so that their papers become almost instantly outdated.

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