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 Kossinian 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.

Related:

Indo-European demic diffusion model, 3rd edition

pca-yamna-corded-ware

I have just uploaded the working draft of the third version of the Indo-European demic diffusion model. Unlike the previous two versions, which were published as essays (fully developed papers), this new version adds more information on human admixture, and probably needs important corrections before a definitive edition can be published.

The third version is available right now on ResearchGate and Academia.edu. I will post the PDF at Academia Prisca, as soon as possible:

pca-map-yamna-corded-ware-bell-beaker
Map overlaid by PCA including Yamna, Corded Ware, Bell Beaker, and other samples

Feel free to comment on the paper here, or (preferably) in our forum.

A working version (needing some corrections) divided by sections, illustrated with up-to-date, high resolution maps, can be found (as always) at the official collaborative Wiki website indo-european.info.

Analysis of R1b-DF27 haplogroups in modern populations adds new information that contrasts with ‘steppe admixture’ results

R1b-DF27-iberia

New open access article published in Scientific Reports, Analysis of the R1b-DF27 haplogroup shows that a large fraction of Iberian Y-chromosome lineages originated recently in situ, by Solé-Morata et al. (2017).

Abstract

Haplogroup R1b-M269 comprises most Western European Y chromosomes; of its main branches, R1b-DF27 is by far the least known, and it appears to be highly prevalent only in Iberia. We have genotyped 1072 R1b-DF27 chromosomes for six additional SNPs and 17 Y-STRs in population samples from Spain, Portugal and France in order to further characterize this lineage and, in particular, to ascertain the time and place where it originated, as well as its subsequent dynamics. We found that R1b-DF27 is present in frequencies ~40% in Iberian populations and up to 70% in Basques, but it drops quickly to 6–20% in France. Overall, the age of R1b-DF27 is estimated at ~4,200 years ago, at the transition between the Neolithic and the Bronze Age, when the Y chromosome landscape of W Europe was thoroughly remodeled. In spite of its high frequency in Basques, Y-STR internal diversity of R1b-DF27 is lower there, and results in more recent age estimates; NE Iberia is the most likely place of origin of DF27. Subhaplogroup frequencies within R1b-DF27 are geographically structured, and show domains that are reminiscent of the pre-Roman Celtic/Iberian division, or of the medieval Christian kingdoms.

Some people like to say that Y-DNA haplogroup analysis, or phylogeography in general, is of no use anymore (especially modern phylogeography), and they are content to see how ‘steppe admixture’ was (or even is) distributed in Europe to draw conclusions about ancient languages and their expansion. With each new paper, we are seeing the advantages of analysing ancient and modern haplogroups in ascertaining population movements.

Quite recently there was a suggestion based on steppe admixture that Basque-speaking Iberians resisted the invasion from the steppe. Observing the results of this article (dates of expansion and demographic data) we see a clear expansion of Y-DNA haplogroups precisely by the time of Bell Beaker expansion from the east. Y-DNA haplogroups of ancient samples from Portugal point exactly to the same conclusion.

The situation of R1b-DF27 in Basques, as I have pointed out elsewhere, is probably then similar to the genetic drift of Finns, mainly of N1c lineages, speaking today a Uralic language that expaned with Corded Ware and R1a subclades.

The recent article on Mycenaean and Minoan genetics also showed that, when it comes to Europe, most of the demographic patterns we see in admixture are reminiscent of the previous situation, only rarely can we see a clear change in admixture (which would mean an important, sudden replacement of the previous population).

Equating the so-called steppe admixture with Indo-European languages is wrong. Period.

The following are excerpts from the article (emphasis is mine):

Dates and expansions

The average STR variance of DF27 and each subhaplogroup is presented in Suppl. Table 2. As expected, internal diversity was higher in the deeper, older branches of the phylogeny. If the same diversity was divided by population, the most salient finding is that native Basques (Table 2) have a lower diversity than other populations, which contrasts with the fact that DF27 is notably more frequent in Basques than elsewhere in Iberia (Suppl. Table 1). Diversity can also be measured as pairwise differences distributions (Fig. 5). The distribution of mean pairwise differences within Z195 sits practically on top of that of DF27; L176.2 and Z220 have similar distributions, as M167 and Z278 have as well; finally, M153 shows the lowest pairwise distribution values. This pattern is likely to reflect the respective ages of the haplogroups, which we have estimated by a modified, weighted version of the ρ statistic (see Methods).

Z195 seems to have appeared almost simultaneously within DF27, since its estimated age is actually older (4570 ± 140 ya). Of the two branches stemming from Z195, L176.2 seems to be slightly younger than Z220 (2960 ± 230 ya vs. 3320 ± 200 ya), although the confidence intervals slightly overlap. M167 is clearly younger, at 2600 ± 250 ya, a similar age to that of Z278 (2740 ± 270 ya). Finally, M153 is estimated to have appeared just 1930 ± 470 ya.

Haplogroup ages can also be estimated within each population, although they should be interpreted with caution (see Discussion). For the whole of DF27, (Table 3), the highest estimate was in Aragon (4530 ± 700 ya), and the lowest in France (3430 ± 520 ya); it was 3930 ± 310 ya in Basques. Z195 was apparently oldest in Catalonia (4580 ± 240 ya), and with France (3450 ± 269 ya) and the Basques (3260 ± 198 ya) having lower estimates. On the contrary, in the Z220 branch, the oldest estimates appear in North-Central Spain (3720 ± 313 ya for Z220, 3420 ± 349 ya for Z278). The Basques always produce lower estimates, even for M153, which is almost absent elsewhere.

R1b-DF27-tree
Simplified phylogenetic tree of the R1b-M269 haplogroup. SNPs in italics were not analyzed in this manuscript.

Demography

The median value for Tstart has been estimated at 103 generations (Table 4), with a 95% highest probability density (HPD) range of 50–287 generations; effective population size increased from 131 (95% HPD: 100–370) to 72,811 (95% HPD: 52,522–95,334). Considering patrilineal generation times of 30–35 years, our results indicate that R1b-DF27 started its expansion ~3,000–3,500 ya, shortly after its TMRCA.

As a reference, we applied the same analysis to the whole of R1b-S116, as well as to other common haplogroups such as G2a, I2, and J2a. Interestingly, all four haplogroups showed clear evidence of an expansion (p > 0.99 in all cases), all of them starting at the same time, ~50 generations ago (Table 4), and with similar estimated initial and final populations. Thus, these four haplogroups point to a common population expansion, even though I2 (TMRCA, weighted ρ, 7,800 ya) and J2a (TMRCA, 5,500 ya) are older than R1b-DF27. It is worth noting that the expansion of these haplogroups happened after the TMRCA of R1b-DF27.

R1b-DF27-PCA
Principal component analysis of STR haplotypes. (a) Colored by subhaplogroup, (b) colored by population. Larger squares represent subhaplogroup or population centroids.

Sum up and discussion

We have characterized the geographical distribution and phylogenetic structure of haplogroup R1b-DF27 in W. Europe, particularly in Iberia, where it reaches its highest frequencies (40–70%). The age of this haplogroup appears clear: with independent samples (our samples vs. the 1000 genome project dataset) and independent methods (variation in 15 STRs vs. whole Y-chromosome sequences), the age of R1b-DF27 is firmly grounded around 4000–4500 ya, which coincides with the population upheaval in W. Europe at the transition between the Neolithic and the Bronze Age. Before this period, R1b-M269 was rare in the ancient DNA record, and during it the current frequencies were rapidly reached. It is also one of the haplogroups (along with its daughter clades, R1b-U106 and R1b-S116) with a sequence structure that shows signs of a population explosion or burst. STR diversity in our dataset is much more compatible with population growth than with stationarity, as shown by the ABC results, but, contrary to other haplogroups such as the whole of R1b-S116, G2a, I2 or J2a, the start of this growth is closer to the TMRCA of the haplogroup. Although the median time for the start of the expansion is older in R1b-DF27 than in other haplogroups, and could suggest the action of a different demographic process, all HPD intervals broadly overlap, and thus, a common demographic history may have affected the whole of the Y chromosome diversity in Iberia. The HPD intervals encompass a broad timeframe, and could reflect the post-Neolithic population expansions from the Bronze Age to the Roman Empire.

While when R1b-DF27 appeared seems clear, where it originated may be more difficult to pinpoint. If we extrapolated directly from haplogroup frequencies, then R1b-DF27 would have originated in the Basque Country; however, for R1b-DF27 and most of its subhaplogroups, internal diversity measures and age estimates are lower in Basques than in any other population. Then, the high frequencies of R1b-DF27 among Basques could be better explained by drift rather than by a local origin (except for the case of M153; see below), which could also have decreased the internal diversity of R1b-DF27 among Basques. An origin of R1b-DF27 outside the Iberian Peninsula could also be contemplated, and could mirror the external origin of R1b-M269, even if it reaches there its highest frequencies. However, the search for an external origin would be limited to France and Great Britain; R1b-DF27 seems to be rare or absent elsewhere: Y-STR data are available only for France, and point to a lower diversity and more recent ages than in Iberia (Table 3). Unlike in Basques, drift in a traditionally closed population seems an unlikely explanation for this pattern, and therefore, it does not seem probable that R1b-DF27 originated in France. Then, a local origin in Iberia seems the most plausible hypothesis. Within Iberia, Aragon shows the highest diversity and age estimates for R1b-DF27, Z195, and the L176.2 branch, although, given the small sample size, any conclusion should be taken cautiously. On the contrary, Z220 and Z278 are estimated to be older in North Central Spain (N Castile, Cantabria and Asturias). Finally, M153 is almost restricted to the Basque Country: it is rarely present at frequencies >1% elsewhere in Spain (although see the cases of Alacant, Andalusia and Madrid, Suppl. Table 1), and it was found at higher frequencies (10–17%) in several Basque regions; a local origin seems plausible, but, given the scarcity of M153 chromosomes outside of the Basque Country, the diversity and age values cannot be compared.

Within its range, R1b-DF27 shows same geographical differentiation: Western Iberia (particularly, Asturias and Portugal), with low frequencies of R1b-Z195 derived chromosomes and relatively high values of R1b-DF27* (xZ195); North Central Spain is characterized by relatively high frequencies of the Z220 branch compared to the L176.2 branch; the latter is more abundant in Eastern Iberia. Taken together, these observations seem to match the East-West patterning that has occurred at least twice in the history of Iberia: i) in pre-Roman times, with Celtic-speaking peoples occupying the center and west of the Iberian Peninsula, while the non-Indoeuropean eponymous Iberians settled the Mediterranean coast and hinterland; and ii) in the Middle Ages, when Christian kingdoms in the North expanded gradually southwards and occupied territories held by Muslim fiefs.

DF27-iberia-france
Contour maps of the derived allele frequencies of the SNPs analyzed in this manuscript. Population abbreviations as in Table 1. Maps were drawn with SURFER v. 12 (Golden Software, Golden CO, USA).

I wouldn’t trust the absence of R1b-DF27 outside France as a proof that its origin must be in Western Europe – especially since we have ancient DNA, and that assertion might prove quite wrong – but aside from that the article seems solid in its analysis of modern populations.

Related:

Text and figures from the article, licensed under a Creative Commons Attribution 4.0 International License. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

Genetic origins of Minoans and Mycenaeans and their continuity into modern Greeks

mycenaean-minoan

A new article has appeared in Nature, Genetic origins of the Minoans and Mycenaeans, by Lazaridis et al. (2017), referenced by Science.

Abstract:

The origins of the Bronze Age Minoan and Mycenaean cultures have puzzled archaeologists for more than a century. We have assembled genome-wide data from 19 ancient individuals, including Minoans from Crete, Mycenaeans from mainland Greece, and their eastern neighbours from southwestern Anatolia. Here we show that Minoans and Mycenaeans were genetically similar, having at least three-quarters of their ancestry from the first Neolithic farmers of western Anatolia and the Aegean, and most of the remainder from ancient populations related to those of the Caucasus3 and Iran. However, the Mycenaeans differed from Minoans in deriving additional ancestry from an ultimate source related to the hunter–gatherers of eastern Europe and Siberia, introduced via a proximal source related to the inhabitants of either the Eurasian steppe or Armenia. Modern Greeks resemble the Mycenaeans, but with some additional dilution of the Early Neolithic ancestry. Our results support the idea of continuity but not isolation in the history of populations of the Aegean, before and after the time of its earliest civilizations.

Samples are scarce, and there is only one Y-DNA haplogroup of Mycenaeans, J2a1 (in Galatas Apatheia, ca. 1700-1200), which shows continuity of haplogroups from Minoan samples, so it does not clarify the potential demic diffusion of Proto-Greeks marked by R1b subclades.

Regarding admixture analyses, it is explicitly or implicitly (according to the press release) stated that:

  • There is continuity between Mycenaeans and living people, so that the major components of the Greeks’ ancestry was in place already in the Bronze Age, after the migration of the earliest farmers from Anatolia.
  • Anatolians may have been the source of “eastern” Caucasian ancestry in Mycenaeans, and maybe of early Indo-European languages (i.e. earlier than Proto-Greek) in the region.
  • The “northern” steppe population (speaking a Late Indo-European dialect, then) had arrived only in mainland Greece, with a 13-18% admixture, by the time studied.
  • Samples before the Final Neolithic (ca. 4100 BC) do not possess either type of ancestry, suggesting that the admixture detected occurred during the fourth to second millennium BC.
  • Admixture from Levantine or African influence (i.e. Egyptian or Phoenician colonists) cannot be supported with admixture.

All in all, there is some new interesting information, and among them the possibility of obtaining ancient DNA from arid regions, which is promising for future developments in the field.

EDIT (20/8/2017): The article received widespread media attention, and two blog posts were linked to by the main author in his Twitter account: Who are you calling Mycenaean?, and On genetics and the Aegean Bronze Age. Apart from the obviously wrong reductio ad Hitlerum that pops up in any discussion on Indo-Europeans or genetics (even I do it regarding fans of admixture analysis), I don’t know why these created so much fuss (and hate) among geneticists. There seems to be a war brewing between Archaeology and Genetics.

Razib Khan writes The Revolution Which Came To Archaeology Without Archaeologists?, and I guess this is how many people feel in the field, but if they had studied some minimal archaeology of the samples they are studying they would know that their conclusions would come as no surprise, in any case. They can solve old archaeological questions, and they can help create new hypothesis. That’s it. Regarding the study Mr. Khan believes did come as a surprise to archaeologists, that on Bell Beakers, I would like to remind him of the predictions Volker Heyd did about genetics already in 2007, based only on Archaeology.

Related:

Featured map: samples studied, from the article.

Something is very wrong with models based on the so-called ‘steppe admixture’ – and archaeologists are catching up

steppe-admixture

Russian archaeologist Leo Klejn has published an article Discussion: Are the Origins of Indo-European Languages Explained by the Migration of the Yamnaya Culture to the West?, which includes the criticism received from Wolfgang Haak, Iosif Lazaridis, Nick Patterson, and David Reich (mainly on the genetic aspect), and from Kristian Kristiansen, Karl-Göran Sjögren, Morten Allentoft, Martin Sikora, and Eske Willerslev (mainly on the archaeological aspect).

I will not post details of Klejn’s model of North-South Proto-Indo-European expansion – which is explained in the article, and relies on the north-south cline of ‘steppe admixture’ in the modern European population -, since it is based on marginal anthropological methods and theories, including glottochronological dates, and archaeological theories from the Russian school (mainly Zalyzniak), which are obviously not mainstream in the field of Indo-European Studies, and (paradoxically) on the modern distribution of ‘steppe admixture’…

The most interesting aspects of the article are the reactions to the criticism, some of which can be used from the point of view of the Indo-European demic diffusion model, too. It is sad, however, that they didn’t choose to answer earlier to Heyd’s criticism (or to Heyd’s model, which is essentially also that of Mallory and Anthony), instead of just waiting for proponents of the least interesting models to react…

The answer by Haak et al.:

Klejn mischaracterizes our paper as claiming that practitioners of the Corded Ware culture spoke a language ancestral to all European Indo-European languages, including Greek and Celtic. This is incorrect: we never claim that the ancestor of Greek is the language spoken by people of the Corded Ware culture. In fact, we explicitly state that the expansion of steppe ancestry might account for only a subset of Indo-European languages in Europe. Klejn asserts that ‘a source in the north’ is a better candidate for the new ancestry manifested in the Corded Ware than the Yamnaya. While it is indeed the case that the present-day people with the greatest affinity to the Corded Ware are distributed in north-eastern Europe, a major part of the new ancestry of the Corded Ware derives from a population most closely related to Armenians (Haak et al., 2015) and hunter-gatherers from the Caucasus (Jones et al., 2015). This ancestry has not been detected in any European huntergatherers analysed to date (Lazaridis et al., 2014; Skoglund et al., 2014; Haak et al., 2015; Fu et al., 2016), but made up some fifty per cent of the ancestry of the Yamnaya. The fact that the Corded Ware traced some of its ancestry to the southern Caucasus makes a source in the north less parsimonious.

In our study, we did not speculate about the date of Proto-Indo-European and the locations of its speakers, as these questions are unresolved by our data, although we do think the genetic data impose constraints on what occurred. We are enthusiastic about the potential of genetics to contribute to a resolution of this longstanding issue, but this is likely to require DNA from multiple, as yet unsampled, ancient populations.

Klejn response to that:

Allegedly, I had accused the authors of tracing all Indo-European languages back to Yamnaya, whereas they did not trace all of them but only a portion! Well, I shall not reproach the authors for their ambiguous language: it remains the case that (beginning with the title of the first article) their qualifications are lost and their readers have understood them as presenting the solution to the whole question of the origins of Indo-European languages.

(…) they had in view not the Proto-Indo-European before the separation of the Hittites, but the language that was left after the separation. Yet, this was still the language ancestral to all the remaining Indo-European languages, and the followers of Sturtevan and Kluckhorst call only this language Proto-Indo-European (while they call the initial one Indo-Hittite). The majority of linguists (specialists in Indo-European languages) is now inclined to this view. True, the breakup of this younger language is several hundred years more recent (nearly a thousand years later according to some glottochronologies) than the separation of Anatolian languages, but it is still around a thousand years earlier than the birth of cultures derived from Yamnaya.
More than that, I analysed in my criticism both possibilities — the case for all Indo-European languages spreading from Yamnaya and the case for only some of them spreading from Yamnaya. In the latter case, it is argued that only the languages of the steppes, the Aryan (Indo- Iranian) are descended from Yamnaya, not the languages of northern Europe. Together with many scholars, I am in agreement with the last possibility. But, then, what sense can the proposed migration of the Yamnaya culture to the Baltic region have? It would bring the Indo-Iranian proto-language to that region! Yet, there are no traces of this language on the coasts of the Baltic!

My main concern is that, to my mind, one should not directly apply conclusions from genetics to events in the development of language because there is no direct and inevitable dependence between events in the life of languages, culture, and physical structure (both anthropological and genetic). They can coincide, but often they all follow divergent paths. In each case the supposed coincidence should be proved separately.

The authors’ third objection concerns the increase of the genetic similarity of European population with that of the Yamnaya culture. This increases in the north of Europe and is weak in the south, in the places adjacent to the Yamnaya area, i.e. in Hungary. This gradient is clearly expressed in the modern population, but was present already in the Bronze Age, and hence cannot be explained by shifts that occurred in the Early Iron Age and in medieval times. However, the supposed migration of the Yamnaya culture to the west and north should imply a gradient in just the opposite direction!

Regarding the arguments of Kristiansen and colleagues:

[They argue that] in two early burials of the Corded Ware culture (one in Germany, the other in Poland) some single attributes of Yamnaya origin have been found.

(…) if this is the full extent of Yamnaya infiltration into central Europe—two burials (one for each country) from several thousands (and from several hundreds of early burials)—then it hardly amounts to large-scale migration.

Quite recently we have witnessed the success of a group of geneticists from Stanford University and elsewhere (Poznik et al., 2016). They succeeded in revealing varieties of Y-chromosome connected with demographic expansions in the Bronze Age. Such expansion can give rise to migration. Among the variants connected with this expansion is R1b, and this haplogroup is typical for the Yamnaya culture. But what bad luck! This haplogroup connected with expansion is indicated by the clade L11, while the Yamnaya burials are associated with a different clade, Z2103, that is not marked by expansion. It is now time to think about how else the remarkable results reached by both teams of experienced and bright geneticists may be interpreted.

Regarding the work of Heyd,

(…) with regard to the barrow burials of the third millennium BC in the basin of the Danube, although they have been assigned to the Yamnaya culture, I would consider them as also belonging to
another, separate culture, perhaps a mixed culture: its burial custom is typical of the Yamnaya, but its pottery is absolutely not Yamnaya, but local Balkan with imports of distinctive corded beakers (Schnurbecher). I would not be surprised if
Y-chromosome haplogroups of this population were somewhat similar to those of the Yamnaya, while mitochondrial groups were indigenous. As yet, geneticists deal with great blocks of populations and prefer to match them to very large and generalized cultural blocks, while archaeology now analyses more concrete and smaller cultures, each of which had its own fate.

Iosif Lazaridis shares more thoughts on the discussion in his Twitter account:

As we mentioned in Haak, Lazaridis et al. (2015), the Yamnaya are the best proximate source for the new ancestry that first appears with the Corded Ware in central Europe, as it has the right mix of both ANE (related to Native Americans, MA1, and EHG), but also Armenian/Caucasus/Iran-like southern component of ancestry. The Yamnaya is a westward expansive culture that bears exactly the two new ancestral components (EHG + Caucasus/Iran/Armenian-like).
As for the Y-chromosome, it was already noted in Haak, Lazaridis et al. (2015) that the Yamnaya from Samara had Y-chromosomes which belonged to R-M269 but did not belong to the clade common in Western Europe (p. 46 of supplement). Also, not a single R1a in Yamnaya unlike Corded Ware (R1a-dominated). But Yamnaya samples = elite burials from eastern part of the Yamnaya range. Both R1a/R1b found in Eneolithic Samara and EHG, so in conclusion Yamnaya expansion still the best proximate source for the post-3,000 BCE population change in central Europe. And since 2015 steppe expansion detected elsewhere (Cassidy et al. 16, Martiniano et al. 17, Mittnik et al. 17, Mathieson et al. 17, Lazaridis et al. 2016 (South Asia) and …?…

I love the smell of new wording in the morning… viz. Yamnaya best proximate source for Corded Ware, Corded Ware might account for only a subset of Indo-European languages, Corded Ware representing Aryan languages (probably Klejn misinterprets what the authors mean, i.e. some kind of Indo-Slavonic or Germano-Balto-Slavic group)…

We shall expect more and more ambiguous rewording and more adjustments of previous conclusions as new papers and new criticisms appear.

Related:

Featured image from the article: Distribution of the ‘Yamnaya’ genetic component in the populations of Europe (data taken from Haak et al., 2015). The intensity of the colour corresponds to the contribution of this component in various modern populations

Another hint at the role of Corded Ware peoples in spreading Uralic languages into north-eastern Europe, found in mtDNA analysis of the Finnish population

corded-ware-migration-yamna

Open article at Scientific Reports (Nature): Identification and analysis of mtDNA genomes attributed to Finns reveal long-stagnant demographic trends obscured in the total diversity, by Översti et al. (2017).

Of special interest is its depiction of Finland’s past as including the expansion of Corded Ware population of mtDNA U5b1b2 (and probably Y-DNA R1a-M417 subclades), most likely Uralic speakers of the Forest Zone, to the north of the Yamna culture (where Late Proto-Indo-European was spoken).

A later expansion of other subclades – particularly Y-DNA N1c -, was probably associated with the later western expansion of the Eurasian Seima-Turbino phenomenon, and its current prevalence in Finnish Y-DNA haplogroups might have been the consequence of the population decline ca. 1500 BC, and later Iron Age population bottleneck (with the population peak ca. 500 AD) described in the article.

That would more naturally explain the ‘cultural diffusion’ of Finnic languages into invading eastern N1c lineages, a diffusion which would have been in fact a long-term, quite gradual replacement of previously prevalent Y-DNA R1a subclades in the region, as supported by the prevalent “steppe” component in genome-wide ancestry of Finns.

Therefore, there were probably no sudden, strong population (and thus cultural) changes associated with the arrival of N1c lineages, like the ones seen with R1a (Corded Ware / Uralic) and R1b (Yamna / Proto-Indo-European) expansions in Europe.

How the Saami fit into this scheme is not yet obvious, though.

Abstract:

In Europe, modern mitochondrial diversity is relatively homogeneous and suggests an ubiquitous rapid population growth since the Neolithic revolution. Similar patterns also have been observed in mitochondrial control region data in Finland, which contrasts with the distinctive autosomal and Y-chromosomal diversity among Finns. A different picture emerges from the 843 whole mitochondrial genomes from modern Finns analyzed here. Up to one third of the subhaplogroups can be considered as Finn-characteristic, i.e. rather common in Finland but virtually absent or rare elsewhere in Europe. Bayesian phylogenetic analyses suggest that most of these attributed Finnish lineages date back to around 3,000–5,000 years, coinciding with the arrival of Corded Ware culture and agriculture into Finland. Bayesian estimation of past effective population sizes reveals two differing demographic histories: 1) the ‘local’ Finnish mtDNA haplotypes yielding small and dwindling size estimates for most of the past; and 2) the ‘immigrant’ haplotypes showing growth typical of most European populations. The results based on the local diversity are more in line with that known about Finns from other studies, e.g., Y-chromosome analyses and archaeology findings. The mitochondrial gene pool thus may contain signals of local population history that cannot be readily deduced from the total diversity.

From its results:

In general, there appears to be two loose and largely overlapping clusters among the Finn-characteristic haplogroups: the first between 1,000–2,000 ybp and the second around 3,300–5,500 ybp. The age of the older cluster coincides temporally with the arrival of the Corded-Ware culture and, notably, the spread of agriculture in Finland. The arrival and spread of agriculture, temporally corresponding with the age estimates for most of the haplogroups characteristic of Finns, might be a sign of population size increase enabled by the new mode of subsistence, resulting in reduced drift and accumulation of genetic diversity in the population.

(…)

Another insight in the past population sizes in Finland is based on radiocarbon-dated archaeological findings in different time periods. These analyses suggest two prehistoric population peaks in Finland, the Stone Age peak (c. 5,500 ybp) and the Metal Age peak (~1,500 ybp). Both of these peaks were followed by a population decline, which appears to have reached its ebb around 3,500 ybp. These developments are not distinguishable in the BSPs. However, these ages correspond well to the two haplogroup age clusters described above. The presumably less severe Iron Age population bottleneck seen in the archaeological data, 1,500–1,300 ybp, temporally coincides with the population size reduction visible for the Finn-characteristic subhaplogroups.

Related:

Discovered via Eurogenes.