Bell Beakers and Mycenaeans from Yamnaya; Corded Ware from the forest steppe

eba-yamnaya-ancestry-hungary

I have recently written about the spread of Pre-Yamnaya or Yamnaya ancestry and Corded Ware-related ancestry throughout Eurasia, using exclusively analyses published by professional geneticists, and filling in the gaps and contradictory data with the most reasonable interpretations. I did so consciously, to avoid any suspicion that I was interspersing my own data or cherry picking results.

Now I’m finished recapitulating the known public data, and the only way forward is the assessment of these populations using the available datasets and free tools.

Understanding the complexities of qpAdm is fairly difficult without a proper genetic and statistical background, which I won’t pretend to have, so its tweaking to get strictly correct results would require an unending game of trial and error. I have sadly little time for this, even taking my tendency to procrastination into account… so I have used a simple model akin to those published before – in particular, the outgroup selection by Ning, Wang et al. (2019), who seem to be part of the only group interested in distinguishing Yamnaya-related from Corded Ware-related ancestry, probably the most relevant question discussed today in population genomics regarding the Proto-Indo-European and Proto-Uralic homelands.

eneolithic-steppe-best-fits
Supplementary Table 13. P values of rank=2 and admixture proportions in modelling Steppe ancestry populations as a three-way admixture of Eneolithic steppe Anatolian_Neolithic and WHG using 14 outgroups.
Left populations: Test, Eneolithic_steppe, Anatolian_Neolithic, WHG.
Right populations: Mbuti.DG, Ust_Ishim.DG, Kostenki14, MA1, Han.DG, Papuan.DG, Onge.DG, Villabruna, Vestonice16, ElMiron, Ethiopia_4500BP.SG, Karitiana.DG, Natufian, Iran_Ganj_Dareh_Neolithic.

I have used for all analyses below a merged dataset including the curated one of the Reich Lab, the latest on Central and South Asia by Narasimhan, Patterson et al. (2019), on Iberia by Olalde et al. (2019), and on the East Baltic by Saag et al. (2019), as well as datasets including samples from Wang et al. (2019) and Lamnidis et al. (2018). I used (and intend to use) the same merged dataset in all cases, despite its huge size, to avoid adding one more uncontrolled variable to the analyses, so that all results obtained can be compared.

I try to prepare in advance a bunch of relevant files with left pops and right pops for each model:

  1. It seems a priori more reasonable to use geographically and chronologically closer proxy populations (say, Trypillia or GAC for Steppe-related peoples) than hypothetic combinations of ancestral ones (viz. Anatolian farmer, WHG, and EHG).
  2. This also means using subgroups closer to the most likely source population, such as (Don-Volga interfluve) Yamnaya_Kalmykia rather than (Middle Volga) Yamnaya_Samara for the western expansion of late Repin/early Yamnaya, or the early Germany_Corded_Ware.SG or Czech_Corded Ware for the group closest to the Proto-Corded Ware population (see below), likely neighbouring the Upper Vistula region.
  3. I usually test two source populations for different targets, which seems like a much more efficient way of using computer resources, whenever I know what I want to test, since I need my PC back for its normal use; whenever I don’t know exactly what to test, I use three-way admixture models and look for subsets to try and improve the results.

I have probably left out some more complex models by individualizing the most relevant groups, but for the time being this would have to do. Also, no other formal stats have been used in any case, which is an evident shortcoming, ruling out an interpretation drawn directly and only from the results below.

Full qpAdm results for each batch of samples are presented in a Google Spreadsheet, with each tab (bottom of the page) showing a different combination of sources, usually in order of formally ‘best’ (first to the left) to ‘worst’ (last to the right) fits, although the order is difficult to select in highly heterogeneous target groups, as will be readily visible.

maykop-trypillia-intrusion-steppes
Disintegration, migration, and imports of the Azov–Black Sea region. First migration event (solid arrows): Gordineşti–Maikop expansion (groups: I – Bursuchensk; II – Zhyvotylivka; III – Vovchans’k; IV – Crimean; V – Lower Don; VI – pre-Kuban). Second migration event (hollow arrows): Repin expansion. After Rassamakin (1999), Demchenko (2016).

Corded Ware origins

The latest publications on the Yampil barrow complex have not improved much our understanding of the complexity of Corded Ware origins from an archaeological point of view, involving multiple cultural (hence likely population) influences. This bit is from Ivanova et al., Baltic-Pontic Studies (2015) 20:1, and most hypotheses of the paper remain unanswered (except maybe for the relevance of the Złota group):

In the light of the above outline therefore one should argue that the ‘architecture of barrows’ associated in the ‘Yampil landscape’ of the Middle Dniester Area with the Eneolithic (specifically, mainly with the TC), precedes the development of a similar phenomenon that can be observed from 2900/2800 BC in the Upper Dniester Area and drainage basin of the Upper Vistula, associated with the CWC [Goslar et al. 2015; Włodarczak 2006; 2007; 2008; Jarosz, Włodarczak 2007]. The most consuming research question therefore is whether ritual customs making use of Eneolithic (Tripolye) ‘barrow architecture’ could have penetrated northwards along the Dniester route, where GAC communities functioned. One could also ask what role the rituals played among the autochthons [Kośko 2000; Włodarczak 2008; 2014: 335; Ivanova, Toshchev 2015b].

This issue has already been discussed with a resulting tentative systemic taxonomy in the studies of Włodarczak, arguing for the Złota culture (ZC) in the Vistula region as an illustration of one of the (Małopolska) reception centres of civilization inspirations from the oldest Pontic ‘barrow culture’ circle associated with the Eneolithic and Early Bronze Age [Włodarczak 2008]. Notably, it is in the ZC that one can notice a set of cultural traits (catacomb grave construction, burial details, forms and decoration of vessels) analogous to those shared by the north-western Black Sea Coast groups of the forest-steppe Eneolithic (chiefly Zhyvotilovka-Volchansk) and the Late Tripolye circle (chiefly Usatovo-Gordinești-Horodiștea-Kasperovtsy).

gac-trypillia-usatovo-corded-ware
Globular Amphorae culture „exodus” to the Danube Delta: a – Globular Amphorae culture; b – GAC (1), Gorodsk (2), Vykhvatintsy (3) and Usatovo (4) groups of Trypillia culture; c – Coţofeni culture; d – northern border of the late phase of Baden culture;red arrows – direction of Globular Amphora culture expansion; blue arrow – direction of „reflux” of Globular Amphora culture (apud Włodarczak, 2008, with changes).

Taking into account that I6561 might be wrongly dated, we cannot include the Corded Ware-like sample of the end-5th millennium BC in the analysis of Corded Ware origins. That uncertainty in the chronology of the appearance of “Steppe ancestry” in Proto-Corded Ware peoples complicates the selection of any potential source population from the CHG cline.

Nevertheless, the lack of hg. R1a-M417 and sizeable Pre-Yamnaya-related ancestry in the sampled Pontic forest-steppe Eneolithic populations (represented exclusively by two samples from Dereivka ca. 3600-3400 BC) would leave open the interesting possibility that a similar ancestry got to the forest-steppe region between modern Poland and Ukraine during the known complex population movements of the Late Eneolithic.

It is known that Corded Ware-derived groups and Steppe Maykop show bad fits for Pre-Yamnaya/Yamnaya ancestry, and also that Steppe Maykop is a potential source of “Steppe-related ancestry” within the Eneolithic CHG mating network of the Pontic-Caspian steppes and forest-steppes. Testing Corded Ware for recent Trypillia and Maykop influences, proper of Late Trypillia and Late Maykop groups in the North Pontic area (such as Zhyvotylivka–Vovchans’k and Gordineşti) side by side with potential Pre-Yamnaya and Yamnaya sources makes thus sense:

Now, the main obvious difference between Khvalynsk-Yamnaya and Corded Ware is the long-lasting, pervasive Y-chromosome bottlenecks under R1b lineages in the former, compared to the haplogroup variability and late bottleneck under R1a-M417 in the latter, which speaks in favour – on top of everything else – of a different community of sub-Neolithic hunter-gatherers including hg. R1a-M417 hijacking the expansion of Steppe_Maykop-related ancestry around the Volhynian-Podolian Upland.

Akin to how Yamnaya patrilineal descendants hijacked regional EEF (±CWC) ancestry components mainly through exogamy, dragging them into the different expanding Bell Beaker groups (see below), but kept their Indo-European languages, these hunter-gatherers that admixed with peoples of “Steppe ancestry” were the most likely vector of expansion of Uralic languages in Eastern Europe.

corded-ware-from-trypillia-maykop
PCA of ancient Eurasian samples. Marked likely Proto-Corded Ware samples and potential origin of its PCA cluster based on qpAdm results. See full PCA and more related files.

Baltic Corded Ware

One of the most interesting aspects of the results above is the surprising heterogeneity of the different regional groups, which is also reflected in the Y-DNA variability of early Corded Ware samples.

Seeing how Baltic CWC groups, especially the early Latvia_LN sample, show particularly bad fits with the models above, it seems necessary to test how this population might have come to be. My first impression in 2017 was that they could represent early Corded Ware groups admixed with Yamnaya settlers through their interactions along the Dnieper-Dniester corridor.

However, I recently predicted that the most likely admixture leading to their ancestry and PCA cluster would involve a Corded Ware-like group and a group related to sub-Neolithic cultures of eastern Europe, whose best proxy to date are EHG-like Khvalynsk samples (i.e. excluding the outlier with Pre-Yamnaya ancestry, I0434):

corded-ware-pca-sub-neolithic-europe
Detail of the PCA of the Corded Ware expansion. See full PCA and more related files.

Late Corded Ware + Yamnaya vanguard

Relevant are also the mixtures of Corded Ware from Esperstedt, and particularly those of the sample I0104, which I have repeated many times in this blog I suspected to be influenced by vanguard Yamnaya settlers:

The infeasible models of CWC + Yamnaya_Kalmykia ± Hungary_Baden (see below for Bell Beakers) and the potential cluster formed with other samples from the Baltic suggest that it could represent a more complex set of mixtures with sub-Neolithic populations. On the other hand, its location in Germany, late date (ca. 2500 BC or later), and position in the PCA, together with the good fits obtained for Germany_Beaker as a source, suggest that the increase in Steppe-related ancestry + EEF makes it impossible for the model (as I set it) to directly include Yamnaya_Kalmykia, despite this excess Steppe-related ancestry actually coming from Yamnaya vanguard groups.

I think it is very likely that the future publication of EEF-admixed Yamnaya_Hungary samples (or maybe even Yamnaya vanguard samples) will improve the fits of this model.

These results confirm at least the need to distrust the common interpretation of mixtures including late Corded Ware samples from Esperstedt (giving rise to the “up to 75% Yamnaya ancestry of CWC” in the 2015 papers) as representative of the Corded Ware culture as a whole, and to keep always in mind that an admixture of European BA groups including Corded Ware Esperstedt as a source also includes East BBC-like ancestry, unless proven otherwise.

yamnaya-vanguard-corded-ware-chalcolithic-early
Yamnaya vanguard groups in Corded Ware territory before the expansion of Bell Beakers (ca. 2500 BC). See full map.

Bell Beaker expansion

A hotly (re)debated topic in the past 6 months or so, and for all the wrong reasons, is the origin of the Bell Beaker folk. Archaeology, linguistics, and different Y-chromosome bottlenecks clearly indicate that Bell Beakers were at the origin of the North-West Indo-European expansion in Europe, while the survival of Corded Ware-related groups in north-eastern Europe is clearly related to the expansion of Uralic languages.

NOTE. For the interesting case of Proto-Indo-Iranians expanding with Corded Ware-like ancestry, see more on the formation of Sintashta-Potapovka-Filatovka from East Uralic-speaking Abashevo and Pre-Proto-Indo-Iranian-speaking Poltavka herders. See also more on R1a in Indo-Iranians and on the social complexity of Sintashta.

Nevertheless, every single discarded theory out there seems to keep coming back to life from time to time, and a new wave of interest in “Bell Beaker from the Single Grave culture” somehow got revived in the process, too, because this obsession – unlike the “Bell Beakers from Iberia Chalcolithic” – is apparently acceptable in certain circles, for some reason.

We know that Iberian Beakers, British Beakers, or Sicilian EBA – representing the most likely closest source population of speakers of Proto-Galaico-Lusitanian, Pre-Celtic Indo-European, and Proto-Elymian, respectively – have already been successfully tested for a direct origin among Western European Beakers in Olalde et al. (2018), Olalde et al. (2019), and Fernandes et al. (2019).

This success in ascertaining a closer Beaker source is probably due to the physical isolation of the specific groups (related to Germany_Beaker, Netherlands_Beaker, and NE_Mediterranean_Beaker samples, respectively) after their migration into regions dominated by peoples without Steppe-related ancestry. Furthermore, Celtic-speaking populations expanding with Urnfield south of the Pyrenees also show a good fit with a source close to France_Beaker.

So I decided to test sampled Bell Beaker populations, to see if it could shed light to the most likely source population of individual Beaker groups and the direction of migration within Central Europe, i.e. roughly eastwards or westwards. As it was to be expected for closely related populations (see the relevant discussion here), an attempt to offer a simplistic analysis of direction based on formal stats does not make any sense, because most of the alternative hypotheses cannot be rejected:

Not only because of the similar values obtained, but because it is absurd to take p-values as a measure of anything, especially when most of these conflicting groups with slightly ‘better’ or ‘worse’ p-values represent multiple different mixtures of the type (Yamnaya + EEF) + (Corded Ware + EEF ± Yamnaya), impossible to distinguish without selecting proper, direct ancestral populations…

A further example of how explosive the Bell Beaker expansion was into different territories, and of their extensive local admixture, is shown by the unsuccessful attempt by Olalde et al. (2018) to obtain an origin of the EEF source for all Beaker groups (excluding Iberian Beakers):

bell-beaker-local-population-iberia
Investigating the genetic makeup of Beaker-complex-associated individuals. Testing different populations as a source for the Neolithic ancestry component in Beaker-complex-associated individuals. The table shows P values (* indicates values > 0.05) for the fit of the model: ‘Steppe_EBA + Neolithic/Copper Age’ source population.
burials-yamnaya-hungary
Map of attested Yamnaya pit-grave burials in the Hungarian plains; superimposed in shades of blue are common areas covered by floods before the extensive controls imposed in the 19th century; in orange, cumulative thickness of sand, unfavourable loamy sand layer. Marked are settlements/findings of Boleráz (ca. 3500 BC on), Baden (until ca. 2800 BC), Kostolac (precise dates unknown), and Yamna kurgans (from ca. 3100/3000 BC on).

Now, there is a simpler way to understand what kind of Steppe-related ancestry is proper of Bell Beakers. I tested two simple models for some Beaker groups: Yamnaya + Hungary Baden vs. Corded Ware + GAC Poland. After all, the Bell Beaker folk should prefer a source more closely related to either Yamnaya Hungary or Central European Corded Ware:

Interestingly, models including Yamnaya + Baden show good fits for the most important groups related to North-West Indo-Europeans, including Bell Beakers from Germany, the Netherlands, Italy, and Poland, representing the most likely closest source populations of speakers of Pre-Proto-Celtic, Pre-Proto-Germanic, Proto-Italo-Venetic, and Pre-Proto-Balto-Slavic, respectively.

The admixed Yamnaya samples from Hungary that will hopefully be published soon by the Jena Lab will most likely further improve these fits, especially in combination with intermediate Chalcolithic populations of the Middle and Upper Danube and its tributaries, to a point where there will be an absolute chronological and geographical genomic trail from the fully Yamnaya-like Yamnaya settlers from Hungary to all North-West Indo-European-speaking groups of the Early Bronze Age.

The only difference between groups will be the gradual admixture events of their source Beaker group with local populations on their expansion paths, including peoples of mainly EEF, CWC+EEF, or CWC+EEF+Yamnaya related ancestry. There is ample evidence beyond ancestry models to support this, in particular continued Y-DNA bottlenecks under typical Yamnaya paternal lineages, mainly represented by R1b-L51 subclades.

east-bell-beaker-group-expansion
Distribution of the Bell Beaker East Group, with its regional provinces, as of c. 2400 cal BC (after Heyd et al. 2004, modified). See full maps.

European Early Bronze Age

European EBA groups that might show conflicting results due to multiple admixture events with Corded Ware-related populations are the Únětice culture and the Nordic Late Neolithic.

The results for Únětice groups seem to be in line with what is expected of a Central European EBA population derived from Bell Beakers admixed with surrounding poulations of East Bell Beaker and/or late (Epi-)Corded Ware descent.

Potential models of mixture for Nordic Late Neolithic samples – despite the bad fits due to the lack of direct ancestral CWC and BBC groups from Denmark – seem to be impossible to justify as derived exclusively from Single Grave or (even less) from Battle Axe peoples, supporting immigration waves of Bell Beakers from the south and further admixture events with local groups through maritime domination.

PCA of ancient European samples. Marked are Bronze Age clusters. See full PCAs.

Balkans Bronze Age

The potential origin of the typical Corded Ware Steppe-related ancestry in the social upheaval and population movements of the Dnieper-Dniester forest-steppe corridor during the 4th millennium BC raises the question: how much do Balkan Bronze Age groups owe their ancestry to a population different than the spread of Pre-Yamnaya-like Suvorovo-Novodanilovka chieftains? Furthermore, which Bronze Age groups seem to be more likely derived exclusively from Pre-Yamnaya groups, and which are more likely to be derived from a mixture of Yamnaya and Pre-Yamnaya? Do the formal stats obtained correspond to the expected results for each group?

Since the expansion of hg. I2a-L699 (TMRCA ca. 5500 BC) need not be associated with Yamnaya, some of these values – together with the assessment of each individual archaeological culture – may question their origin in a Yamnaya-related expansion rather than in a Khvalynsk-related one.

NOTE. These are the last ones I was able to test yesterday, and I have not thought these models through, so feel free to propose other source and target groups. In particular, complex movements through the North Pontic area during the Late Eneolithic would suggest that there might have been different Steppe-ancestry-related vs. EEF-related interactions in the north-west and west Pontic area before and during the expansion of Yamnaya.

Mycenaeans

One of the key Indo-European populations that should be derived from Yamnaya to confirm the Steppe hypothesis, together with North-West Indo-Europeans, are Proto-Greeks, who will in turn improve our understanding of the preceding Palaeo-Balkan community. Unfortunately, we only have Mycenaean samples from the Aegean, with slight contributions of Steppe-related ancestry.

Still, analyses with potential source populations for this Steppe ancestry show that the Yamnaya outlier from Bulgaria is a good fit:

The comparison of all results makes it quite evident the why of the good fits from (Srubnaya-related) Bulgaria_MLBA I2163 or of Sintashta_MLBA relative to the only a priori reasonable Yamnaya and Catacomb sources: it is not about some hypothetical shared ancestor in Graeco-Aryan-speaking East Yamnaya– or even Catacomb-Poltavka-related groups, because all available Yamnaya-related peoples are almost indistinguishable from each other (at least with the sampling available today). These results reflect a sizeable contribution of similar EEF-related populations from around the Carpathians in both Steppe-related groups: Corded Ware and Yamnaya settlers from the Balkans.

mycenaeans-minyan-ware-greece-minoan
Cultural groups in and around the Balkans during the Early Bronze Age. See full maps.

qpAdm magic

In hobby ancestry magic, as in magic in general, it is not about getting dubious results out of thin air: misdirection is the key. A magician needs to draw the audience attention to ‘remarkable’ ancestry percentages coupled with ‘great’ (?) p-values that purportedly “prove” what the audience expects to see, distracting everyone from the true interesting aspects, like statistical design, the data used (and its shortcomings), other opposing models, a comparison of values, a proper interpretation…you name it.

I reckon – based on the examples above – that the following problems lie at the core of bad uses of qpAdm:

  1. In the formal aspect, the poor understanding of what p-values and other formal stats obtained actually mean, and – more importantly – what they don’t mean. The simplistic trend to accept results of a few analyses at face value is necessarily wrong, in so far as there is often no proper reasoning of what is being assessed and how, and there is never a previous opinion about what could be expected if the alternative hypotheses were true.
  2. In the interpretation aspect, the poor judgement of accompanying any results with simplistic, superficial, irrelevant, and often plainly wrong archaeological or linguistic data selected a posteriori; the inclusion of some racial or sociopolitical overtones in the mixture to set a propitious mood in the target audience; and a sort of ritualistic theatrics with the main theme of ‘winning’, that is best completed with ad hominems.

If you get rid of all this, the most reasonable interpretation of the output of a model proposed and tested should be similar to Nick Patterson’s words in his explanation of qpWave and qpAdm use:

Here we see that, at least in this analysis there are reasonable models with CordedWareNeolithic is a mix of either WHG or LBKNeolithic and YamnayaEBA. (…) The point of this note is not to give a serious phylogenetic analysis but the results here certainly support a major Steppe contribution to the Corded Ware population, which is entirely concordant with the archaeology [?].

Very far, as you can see, from the childish “Eureka! I proved the source!”-kind of thinking common among hobbyists.

The Mycenaean case is an illustrative example: if the Yamnaya outlier from Bulgaria were not available, and if one were not careful when designing and assessing those mixture models, the interpretation would range from erroneous (viz. a Graeco-Aryan substrate, as I initially thought) to impossible (say, inventing migration waves of Sintashta or Srubnaya peoples into Crete). The models presented above show that a contribution of Yamnaya to Mycenaeans couldn’t be rejected, and this alone should have been enough to accept Yamnaya as the most likely source population of “Steppe ancestry” in Proto-Greeks, pending intermediate samples from the Balkans. In other words, one could actually find that ‘the best’ p-values for source populations of Mycenaeans is a combination of modern Poles + Turks, despite the impracticality of such a model…

I haven’t been able to reproduce results which supposedly showed that Corded Ware is more likely to be derived from (Pre-)Yamnaya than other source population, or that Corded Ware is better suited as the ancestral population of Bell Beakers. The analyses above show values in line with what has been published in recent scientific papers, and what should be expected based on linguistics and archaeology. So I’ll go out on a limb here and say that it’s only through a careful selection of outgroups and samples tested, and of as few compared models as possible, that you could eventually get this kind of results and interpretation, if at all.

Whether that kind of special care for outgroups and samples is about (a) an acceptable fine-tuning of the analyses, (b) a simplistic selection dragged from the first papers published and applied indiscriminately to all models, or (c) cherry picking analyses until results fit the expected outcome, is a question that will become mostly irrelevant when future publications continue to support an origin of the expansion of ancient Indo-European languages in Khvalynsk- and Yamnaya-related migrations.

Feel free to suggest (reasonable) modifications to correct some of these models in the comments. Also, be sure to check out other values such as proportions, SD or SNPs of the different results that I might have not taken into account when assessing ‘good’ or ‘bad’ fits.

Related

Yamnaya ancestry: mapping the Proto-Indo-European expansions

steppe-ancestry-expansion-europe

The latest papers from Ning et al. Cell (2019) and Anthony JIES (2019) have offered some interesting new data, supporting once more what could be inferred since 2015, and what was evident in population genomics since 2017: that Proto-Indo-Europeans expanded under R1b bottlenecks, and that the so-called “Steppe ancestry” referred to two different components, one – Yamnaya or Steppe_EMBA ancestry – expanding with Proto-Indo-Europeans, and the other one – Corded Ware or Steppe_MLBA ancestry – expanding with Uralic speakers.

The following maps are based on formal stats published in the papers and supplementary materials from 2015 until today, mainly on Wang et al. (2018 & 2019), Mathieson et al. (2018) and Olalde et al. (2018), and others like Lazaridis et al. (2016), Lazaridis et al. (2017), Mittnik et al. (2018), Lamnidis et al. (2018), Fernandes et al. (2018), Jeong et al. (2019), Olalde et al. (2019), etc.

NOTE. As in the Corded Ware ancestry maps, the selected reports in this case are centered on the prototypical Yamnaya ancestry vs. other simplified components, so everything else refers to simplistic ancestral components widespread across populations that do not necessarily share any recent connection, much less a language. In fact, most of the time they clearly didn’t. They can be interpreted as “EHG that is not part of the Yamnaya component”, or “CHG that is not part of the Yamnaya component”. They can’t be read as “expanding EHG people/language” or “expanding CHG people/language”, at least no more than maps of “Steppe ancestry” can be read as “expanding Steppe people/language”. Also, remember that I have left the default behaviour for color classification, so that the highest value (i.e. 1, or white colour) could mean anything from 10% to 100% depending on the specific ancestry and period; that’s what the legend is for… But, fere libenter homines id quod volunt credunt.

Sections:

  1. Neolithic or the formation of Early Indo-European
  2. Eneolithic or the expansion of Middle Proto-Indo-European
  3. Chalcolithic / Early Bronze Age or the expansion of Late Proto-Indo-European
  4. European Early Bronze Age and MLBA or the expansion of Late PIE dialects

1. Neolithic

Anthony (2019) agrees with the most likely explanation of the CHG component found in Yamnaya, as derived from steppe hunter-fishers close to the lower Volga basin. The ultimate origin of this specific CHG-like component that eventually formed part of the Pre-Yamnaya ancestry is not clear, though:

The hunter-fisher camps that first appeared on the lower Volga around 6200 BC could represent the migration northward of un-admixed CHG hunter-fishers from the steppe parts of the southeastern Caucasus, a speculation that awaits confirmation from aDNA.

neolithic-chg-ancestry
Natural neighbor interpolation of CHG ancestry among Neolithic populations. See full map.

The typical EHG component that formed part eventually of Pre-Yamnaya ancestry came from the Middle Volga Basin, most likely close to the Samara region, as shown by the sampled Samara hunter-gatherer (ca. 5600-5500 BC):

After 5000 BC domesticated animals appeared in these same sites in the lower Volga, and in new ones, and in grave sacrifices at Khvalynsk and Ekaterinovka. CHG genes and domesticated animals flowed north up the Volga, and EHG genes flowed south into the North Caucasus steppes, and the two components became admixed.

neolithic-ehg-ancestry
Natural neighbor interpolation of EHG ancestry among Neolithic populations. See full map.

To the west, in the Dnieper-Dniester area, WHG became the dominant ancestry after the Mesolithic, at the expense of EHG, revealing a likely mating network reaching to the north into the Baltic:

Like the Mesolithic and Neolithic populations here, the Eneolithic populations of Dnieper-Donets II type seem to have limited their mating network to the rich, strategic region they occupied, centered on the Rapids. The absence of CHG shows that they did not mate frequently if at all with the people of the Volga steppes (…)

neolithic-whg-ancestry
Natural neighbor interpolation of WHG ancestry among Neolithic populations. See full map.

North-West Anatolia Neolithic ancestry, proper of expanding Early European farmers, is found up to border of the Dniester, as Anthony (2007) had predicted.

neolithic-anatolia-farmer-ancestry
Natural neighbor interpolation of Anatolia Neolithic ancestry among Neolithic populations. See full map.

2. Eneolithic

From Anthony (2019):

After approximately 4500 BC the Khvalynsk archaeological culture united the lower and middle Volga archaeological sites into one variable archaeological culture that kept domesticated sheep, goats, and cattle (and possibly horses). In my estimation, Khvalynsk might represent the oldest phase of PIE.

(…) this middle Volga mating network extended down to the North Caucasian steppes, where at cemeteries such as Progress-2 and Vonyuchka, dated 4300 BC, the same Khvalynsk-type ancestry appeared, an admixture of CHG and EHG with no Anatolian Farmer ancestry, with steppe-derived Y-chromosome haplogroup R1b. These three individuals in the North Caucasus steppes had higher proportions of CHG, overlapping Yamnaya. Without any doubt, a CHG population that was not admixed with Anatolian Farmers mated with EHG populations in the Volga steppes and in the North Caucasus steppes before 4500 BC. We can refer to this admixture as pre-Yamnaya, because it makes the best currently known genetic ancestor for EHG/CHG R1b Yamnaya genomes.

From Wang et al (2019):

Three individuals from the sites of Progress 2 and Vonyuchka 1 in the North Caucasus piedmont steppe (‘Eneolithic steppe’), which harbour EHG and CHG related ancestry, are genetically very similar to Eneolithic individuals from Khvalynsk II and the Samara region. This extends the cline of dilution of EHG ancestry via CHG-related ancestry to sites immediately north of the Caucasus foothills

eneolithic-pre-yamnaya-ancestry
Natural neighbor interpolation of Pre-Yamnaya ancestry among Neolithic populations. See full map. This map corresponds roughly to the map of Khvalynsk-Novodanilovka expansion, and in particular to the expansion of horse-head pommel-scepters (read more about Khvalynsk, and specifically about horse symbolism)

NOTE. Unpublished samples from Ekaterinovka have been previously reported as within the R1b-L23 tree. Interestingly, although the Varna outlier is a female, the Balkan outlier from Smyadovo shows two positive SNP calls for hg. R1b-M269. However, its poor coverage makes its most conservative haplogroup prediction R-M343.

The formation of this Pre-Yamnaya ancestry sets this Volga-Caucasus Khvalynsk community apart from the rest of the EHG-like population of eastern Europe.

eneolithic-ehg-ancestry
Natural neighbor interpolation of non-Pre-Yamnaya EHG ancestry among Eneolithic populations. See full map.

Anthony (2019) seems to rely on ADMIXTURE graphics when he writes that the late Sredni Stog sample from Alexandria shows “80% Khvalynsk-type steppe ancestry (CHG&EHG)”. While this seems the most logical conclusion of what might have happened after the Suvorovo-Novodanilovka expansion through the North Pontic steppes (see my post on “Steppe ancestry” step by step), formal stats have not confirmed that.

In fact, analyses published in Wang et al. (2019) rejected that Corded Ware groups are derived from this Pre-Yamnaya ancestry, a reality that had been already hinted in Narasimhan et al. (2018), when Steppe_EMBA showed a poor fit for expanding Srubna-Andronovo populations. Hence the need to consider the whole CHG component of the North Pontic area separately:

eneolithic-chg-ancestry
Natural neighbor interpolation of non-Pre-Yamnaya CHG ancestry among Eneolithic populations. See full map. You can read more about population movements in the late Sredni Stog and closer to the Proto-Corded Ware period.

NOTE. Fits for WHG + CHG + EHG in Neolithic and Eneolithic populations are taken in part from Mathieson et al. (2019) supplementary materials (download Excel here). Unfortunately, while data on the Ukraine_Eneolithic outlier from Alexandria abounds, I don’t have specific data on the so-called ‘outlier’ from Dereivka compared to the other two analyzed together, so these maps of CHG and EHG expansion are possibly showing a lesser distribution to the west than the real one ca. 4000-3500 BC.

eneolithic-whg-ancestry
Natural neighbor interpolation of WHG ancestry among Eneolithic populations. See full map.

Anatolia Neolithic ancestry clearly spread to the east into the north Pontic area through a Middle Eneolithic mating network, most likely opened after the Khvalynsk expansion:

eneolithic-anatolia-farmer-ancestry
Natural neighbor interpolation of Anatolia Neolithic ancestry among Eneolithic populations. See full map.
eneolithic-iran-chl-ancestry
Natural neighbor interpolation of Iran Chl. ancestry among Eneolithic populations. See full map.

Regarding Y-chromosome haplogroups, Anthony (2019) insists on the evident association of Khvalynsk, Yamnaya, and the spread of Pre-Yamnaya and Yamnaya ancestry with the expansion of elite R1b-L754 (and some I2a2) individuals:

eneolithic-early-y-dna
Y-DNA haplogroups in West Eurasia during the Early Eneolithic in the Pontic-Caspian steppes. See full map, and see culture, ADMIXTURE, Y-DNA, and mtDNA maps of the Early Eneolithic and Late Eneolithic.

3. Early Bronze Age

Data from Wang et al. (2019) show that Corded Ware-derived populations do not have good fits for Eneolithic_Steppe-like ancestry, no matter the model. In other words: Corded Ware populations show not only a higher contribution of Anatolia Neolithic ancestry (ca. 20-30% compared to the ca. 2-10% of Yamnaya); they show a different EHG + CHG combination compared to the Pre-Yamnaya one.

eneolithic-steppe-best-fits
Supplementary Table 13. P values of rank=2 and admixture proportions in modelling Steppe ancestry populations as a three-way admixture of Eneolithic steppe Anatolian_Neolithic and WHG using 14 outgroups.
Left populations: Test, Eneolithic_steppe, Anatolian_Neolithic, WHG.
Right populations: Mbuti.DG, Ust_Ishim.DG, Kostenki14, MA1, Han.DG, Papuan.DG, Onge.DG, Villabruna, Vestonice16, ElMiron, Ethiopia_4500BP.SG, Karitiana.DG, Natufian, Iran_Ganj_Dareh_Neolithic.

Yamnaya Kalmykia and Afanasievo show the closest fits to the Eneolithic population of the North Caucasian steppes, rejecting thus sizeable contributions from Anatolia Neolithic and/or WHG, as shown by the SD values. Both probably show then a Pre-Yamnaya ancestry closest to the late Repin population.

wang-eneolithic-steppe-caucasus-yamnaya
Modelling results for the Steppe and Caucasus cluster. Admixture proportions based on (temporally and geographically) distal and proximal models, showing additional AF ancestry in Steppe groups and additional gene flow from the south in some of the Steppe groups as well as the Caucasus groups. See tables above. Modified from Wang et al. (2019). Within a blue square, Yamnaya-related groups; within a cyan square, Corded Ware-related groups. Green background behind best p-values. In red circle, SD of AF/WHG ancestry contribution in Afanasevo and Yamnaya Kalmykia, with ranges that almost include 0%.

EBA maps include data from Wang et al. (2018) supplementary materials, specifically unpublished Yamnaya samples from Hungary that appeared in analysis of the preprint, but which were taken out of the definitive paper. Their location among Yamnaya settlers from Hungary is speculative, although most uncovered kurgans in Hungary are concentrated in the Tisza-Danube interfluve.

eba-yamnaya-ancestry
Natural neighbor interpolation of Pre-Yamnaya ancestry among Early Bronze Age populations. See full map. This map corresponds roughly with the known expansion of late Repin/Yamnaya settlers.

The Y-chromosome bottleneck of elite males from Proto-Indo-European clans under R1b-L754 and some I2a2 subclades, already visible in the Khvalynsk sampling, became even more noticeable in the subsequent expansion of late Repin/early Yamnaya elites under R1b-L23 and I2a-L699:

chalcolithic-early-y-dna
Y-DNA haplogroups in West Eurasia during the Yamnaya expansion. See full map and maps of cultures, ADMIXTURE, Y-DNA, and mtDNA of the Early Chalcolithic and Yamnaya Hungary.

Maps of CHG, EHG, Anatolia Neolithic, and probably WHG show the expansion of these components among Corded Ware-related groups in North Eurasia, apart from other cultures close to the Caucasus:

NOTE. For maps with actual formal stats of Corded Ware ancestry from the Early Bronze Age to the modern times, you can read the post Corded Ware ancestry in North Eurasia and the Uralic expansion.

eba-chg-ancestry
Natural neighbor interpolation of non-Pre-Yamnaya CHG ancestry among Early Bronze Age populations. See full map.
eba-ehg-ancestry
Natural neighbor interpolation of non-Pre-Yamnaya EHG ancestry among Early Bronze Age populations. See full map.
eba-whg-ancestry
Natural neighbor interpolation of WHG ancestry among Early Bronze Age populations. See full map.
eba-anatolia-farmer-ancestry
Natural neighbor interpolation of Anatolia Neolithic ancestry among Early Bronze Age populations. See full map.
eba-iran-chl-ancestry
Natural neighbor interpolation of Iran Chl. ancestry among Early Bronze Age populations. See full map.

4. Middle to Late Bronze Age

The following maps show the most likely distribution of Yamnaya ancestry during the Bell Beaker-, Balkan-, and Sintashta-Potapovka-related expansions.

4.1. Bell Beakers

The amount of Yamnaya ancestry is probably overestimated among populations where Bell Beakers replaced Corded Ware. A map of Yamnaya ancestry among Bell Beakers gets trickier for the following reasons:

  • Expanding Repin peoples of Pre-Yamnaya ancestry must have had admixture through exogamy with late Sredni Stog/Proto-Corded Ware peoples during their expansion into the North Pontic area, and Sredni Stog in turn had probably some Pre-Yamnaya admixture, too (although they don’t appear in the simplistic formal stats above). This is supported by the increase of Anatolia farmer ancestry in more western Yamna samples.
  • Later, Yamnaya admixed through exogamy with Corded Ware-like populations in Central Europe during their expansion. Even samples from the Middle to Upper Danube and around the Lower Rhine will probably show increasing contributions of Steppe_MLBA, at the same time as they show an increasing proportion of EEF-related ancestry.
  • To complicate things further, the late Corded Ware Espersted family (from ca. 2500 BC or later) shows, in turn, what seems like a recent admixture with Yamnaya vanguard groups, with the sample of highest Yamnaya ancestry being the paternal uncle of other individuals (all of hg. R1a-M417), suggesting that there might have been many similar Central European mating networks from the mid-3rd millennium BC on, of (mainly) Yamnaya-like R1b elites displaying a small proportion of CW-like ancestry admixing through exogamy with Corded Ware-like peoples who already had some Yamnaya ancestry.
mlba-yamnaya-ancestry
Natural neighbor interpolation of Yamnaya ancestry among Middle to Late Bronze Age populations (Esperstedt CWC site close to BK_DE, label is hidden by BK_DE_SAN). See full map. You can see how this map correlated with the map of Late Copper Age migrations and Yamanaya into Bell Beaker expansion.

NOTE. Terms like “exogamy”, “male-driven migration”, and “sex bias”, are not only based on the Y-chromosome bottlenecks visible in the different cultural expansions since the Palaeolithic. Despite the scarce sampling available in 2017 for analysis of “Steppe ancestry”-related populations, it appeared to show already a male sex bias in Goldberg et al. (2017), and it has been confirmed for Neolithic and Copper Age population movements in Mathieson et al. (2018) – see Supplementary Table 5. The analysis of male-biased expansion of “Steppe ancestry” in CWC Esperstedt and Bell Beaker Germany is, for the reasons stated above, not very useful to distinguish their mutual influence, though.

Based on data from Olalde et al. (2019), Bell Beakers from Germany are the closest sampled ones to expanding East Bell Beakers, and those close to the Rhine – i.e. French, Dutch, and British Beakers in particular – show a clear excess “Steppe ancestry” due to their exogamy with local Corded Ware groups:

Only one 2-way model fits the ancestry in Iberia_CA_Stp with P-value>0.05: Germany_Beaker + Iberia_CA. Finding a Bell Beaker-related group as a plausible source for the introduction of steppe ancestry into Iberia is consistent with the fact that some of the individuals in the Iberia_CA_Stp group were excavated in Bell Beaker associated contexts. Models with Iberia_CA and other Bell Beaker groups such as France_Beaker (P-value=7.31E-06), Netherlands_Beaker (P-value=1.03E-03) and England_Beaker (P-value=4.86E-02) failed, probably because they have slightly higher proportions of steppe ancestry than the true source population.

olalde-iberia-chalcolithic

The exogamy with Corded Ware-like groups in the Lower Rhine Basin seems at this point undeniable, as is the origin of Bell Beakers around the Middle-Upper Danube Basin from Yamnaya Hungary.

To avoid this excess “Steppe ancestry” showing up in the maps, since Bell Beakers from Germany pack the most Yamnaya ancestry among East Bell Beakers outside Hungary (ca. 51.1% “Steppe ancestry”), I equated this maximum with BK_Scotland_Ach (which shows ca. 61.1% “Steppe ancestry”, highest among western Beakers), and applied a simple rule of three for “Steppe ancestry” in Dutch and British Beakers.

NOTE. Formal stats for “Steppe ancestry” in Bell Beaker groups are available in Olalde et al. (2018) supplementary materials (PDF). I didn’t apply this adjustment to Bk_FR groups because of the R1b Bell Beaker sample from the Champagne/Alsace region reported by Samantha Brunel that will pack more Yamnaya ancestry than any other sampled Beaker to date, hence probably driving the Yamnaya ancestry up in French samples.

The most likely outcome in the following years, when Yamnaya and Corded Ware ancestry are investigated separately, is that Yamnaya ancestry will be much lower the farther away from the Middle and Lower Danube region, similar to the case in Iberia, so the map above probably overestimates this component in most Beakers to the north of the Danube. Even the late Hungarian Beaker samples, who pack the highest Yamnaya ancestry (up to 75%) among Beakers, represent likely a back-migration of Moravian Beakers, and will probably show a contribution of Corded Ware ancestry due to the exogamy with local Moravian groups.

Despite this decreasing admixture as Bell Beakers spread westward, the explosive expansion of Yamnaya R1b male lineages (in words of David Reich) and the radical replacement of local ones – whether derived from Corded Ware or Neolithic groups – shows the true extent of the North-West Indo-European expansion in Europe:

chalcolithic-late-y-dna
Y-DNA haplogroups in West Eurasia during the Bell Beaker expansion. See full map and see maps of cultures, ADMIXTURE, Y-DNA, and mtDNA of the Late Copper Age and of the Yamnaya-Bell Beaker transition.

4.2. Palaeo-Balkan

There is scarce data on Palaeo-Balkan movements yet, although it is known that:

  1. Yamnaya ancestry appears among Mycenaeans, with the Yamnaya Bulgaria sample being its best current ancestral fit;
  2. the emergence of steppe ancestry and R1b-M269 in the eastern Mediterranean was associated with Ancient Greeks;
  3. Thracians, Albanians, and Armenians also show R1b-M269 subclades and “Steppe ancestry”.

4.3. Sintashta-Potapovka-Filatovka

Interestingly, Potapovka is the only Corded Ware derived culture that shows good fits for Yamnaya ancestry, despite having replaced Poltavka in the region under the same Corded Ware-like (Abashevo) influence as Sintashta.

This proves that there was a period of admixture in the Pre-Proto-Indo-Iranian community between CWC-like Abashevo and Yamnaya-like Catacomb-Poltavka herders in the Sintashta-Potapovka-Filatovka community, probably more easily detectable in this group because of the specific temporal and geographic sampling available.

srubnaya-yamnaya-ehg-chg-ancestry
Supplementary Table 14. P values of rank=3 and admixture proportions in modelling Steppe ancestry populations as a four-way admixture of distal sources EHG, CHG, Anatolian_Neolithic and WHG using 14 outgroups.
Left populations: Steppe cluster, EHG, CHG, WHG, Anatolian_Neolithic
Right populations: Mbuti.DG, Ust_Ishim.DG, Kostenki14, MA1, Han.DG, Papuan.DG, Onge.DG, Villabruna, Vestonice16, ElMiron, Ethiopia_4500BP.SG, Karitiana.DG, Natufian, Iran_Ganj_Dareh_Neolithic.

Srubnaya ancestry shows a best fit with non-Pre-Yamnaya ancestry, i.e. with different CHG + EHG components – possibly because the more western Potapovka (ancestral to Proto-Srubnaya Pokrovka) also showed good fits for it. Srubnaya shows poor fits for Pre-Yamnaya ancestry probably because Corded Ware-like (Abashevo) genetic influence increased during its formation.

On the other hand, more eastern Corded Ware-derived groups like Sintashta and its more direct offshoot Andronovo show poor fits with this model, too, but their fits are still better than those including Pre-Yamnaya ancestry.

mlba-ehg-ancestry
Natural neighbor interpolation of non-Pre-Yamnaya EHG ancestry among Middle to Late Bronze Age populations. See full map.
mlba-chg-ancestry
Natural neighbor interpolation of non-Pre-Yamnaya CHG ancestry among Middle to Late Bronze Age populations. See full map.
mlba-anatolia-farmer-ancestry
Natural neighbor interpolation of Anatolia Neolithic ancestry among Middle to Late Bronze Age populations. See full map.
mlba-iran-chl-ancestry
Natural neighbor interpolation of Iran Chl. ancestry among Middle to Late Bronze Age populations. See full map.

NOTE For maps with actual formal stats of Corded Ware ancestry from the Early Bronze Age to the modern times, you should read the post Corded Ware ancestry in North Eurasia and the Uralic expansion instead.

The bottleneck of Proto-Indo-Iranians under R1a-Z93 was not yet complete by the time when the Sintashta-Potapovka-Filatovka community expanded with the Srubna-Andronovo horizon:

early-bronze-age-y-dna
Y-DNA haplogroups in West Eurasia during the European Early Bronze Age. See full map and see maps of cultures, ADMIXTURE, Y-DNA, and mtDNA of the Early Bronze Age.

4.4. Afanasevo

At the end of the Afanasevo culture, at least three samples show hg. Q1b (ca. 2900-2500 BC), which seemed to point to a resurgence of local lineages, despite continuity of the prototypical Pre-Yamnaya ancestry. On the other hand, Anthony (2019) makes this cryptic statement:

Yamnaya men were almost exclusively R1b, and pre-Yamnaya Eneolithic Volga-Caspian-Caucasus steppe men were principally R1b, with a significant Q1a minority.

Since the only available samples from the Khvalynsk community are R1b (x3), Q1a(x1), and R1a(x1), it seems strange that Anthony would talk about a “significant minority”, unless Q1a (potentially Q1b in the newer nomenclature) will pop up in some more individuals of those ca. 30 new to be published. Because he also mentions I2a2 as appearing in one elite burial, it seems Q1a (like R1a-M459) will not appear under elite kurgans, although it is still possible that hg. Q1a was involved in the expansion of Afanasevo to the east.

middle-bronze-age-y-dna
Y-DNA haplogroups in West Eurasia during the Middle Bronze Age. See full map and see maps of cultures, ADMIXTURE, Y-DNA, and mtDNA of the Middle Bronze Age and the Late Bronze Age.

Okunevo, which replaced Afanasevo in the Altai region, shows a majority of hg. Q1b, but also some R1b-M269 samples proper of Afanasevo, suggesting partial genetic continuity.

NOTE. Other sampled Siberian populations clearly show a variety of Q subclades that likely expanded during the Palaeolithic, such as Baikal EBA samples from Ust’Ida and Shamanka with a majority of Q1b, and hg. Q reported from Elunino, Sagsai, Khövsgöl, and also among peoples of the Srubna-Andronovo horizon (the Krasnoyarsk MLBA outlier), and in Karasuk.

From Damgaard et al. Science (2018):

(…) in contrast to the lack of identifiable admixture from Yamnaya and Afanasievo in the CentralSteppe_EMBA, there is an admixture signal of 10 to 20% Yamnaya and Afanasievo in the Okunevo_EMBA samples, consistent with evidence of western steppe influence. This signal is not seen on the X chromosome (qpAdm P value for admixture on X 0.33 compared to 0.02 for autosomes), suggesting a male-derived admixture, also consistent with the fact that 1 of 10 Okunevo_EMBA males carries a R1b1a2a2 Y chromosome related to those found in western pastoralists. In contrast, there is no evidence of western steppe admixture among the more eastern Baikal region region Bronze Age (~2200 to 1800 BCE) samples.

This Yamnaya ancestry has been also recently found to be the best fit for the Iron Age population of Shirenzigou in Xinjiang – where Tocharian languages were attested centuries later – despite the haplogroup diversity acquired during their evolution, likely through an intermediate Chemurchek culture (see a recent discussion on the elusive Proto-Tocharians).

Haplogroup diversity seems to be common in Iron Age populations all over Eurasia, most likely due to the spread of different types of sociopolitical structures where alliances played a more relevant role in the expansion of peoples. A well-known example of this is the spread of Akozino warrior-traders in the whole Baltic region under a partial N1a-VL29-bottleneck associated with the emerging chiefdom-based systems under the influence of expanding steppe nomads.

early-iron-age-y-dna
Y-DNA haplogroups in West Eurasia during the Early Iron Age. See full map and see maps of cultures, ADMIXTURE, Y-DNA, and mtDNA of the Early Iron Age and Late Iron Age.

Surprisingly, then, Proto-Tocharians from Shirenzigou pack up to 74% Yamnaya ancestry, in spite of the 2,000 years that separate them from the demise of the Afanasevo culture. They show more Yamnaya ancestry than any other population by that time, being thus a sort of Late PIE fossils not only in their archaic dialect, but also in their genetic profile:

shirenzigou-afanasievo-yamnaya-andronovo-srubna-ulchi-han

The recent intrusion of Corded Ware-like ancestry, as well as the variable admixture with Siberian and East Asian populations, both point to the known intense Old Iranian and Old/Middle Chinese contacts. The scarce Proto-Samoyedic and Proto-Turkic loans in Tocharian suggest a rather loose, probably more distant connection with East Uralic and Altaic peoples from the forest-steppe and steppe areas to the north (read more about external influences on Tocharian).

Interestingly, both R1b samples, MO12 and M15-2 – likely of Asian R1b-PH155 branch – show a best fit for Andronovo/Srubna + Hezhen/Ulchi ancestry, suggesting a likely connection with Iranians to the east of Xinjiang, who later expanded as the Wusun and Kangju. How they might have been related to Huns and Xiongnu individuals, who also show this haplogroup, is yet unknown, although Huns also show hg. R1a-Z93 (probably most R1a-Z2124) and Steppe_MLBA ancestry, earlier associated with expanding Iranian peoples of the Srubna-Andronovo horizon.

All in all, it seems that prehistoric movements explained through the lens of genetic research fit perfectly well the linguistic reconstruction of Proto-Indo-European and Proto-Uralic.

Related

Corded Ware ancestry in North Eurasia and the Uralic expansion

uralic-clines-nganasan

Now that it has become evident that Late Repin (i.e. Yamnaya/Afanasevo) ancestry was associated with the migration of R1b-L23-rich Late Proto-Indo-Europeans from the steppe in the second half of the the 4th millennium BC, there’s still the question of how R1a-rich Uralic speakers of Corded Ware ancestry expanded , and how they spread their languages throughout North Eurasia.

Modern North Eurasians

I have been collecting information from the supplementary data of the latest papers on modern and ancient North Eurasian peoples, including Jeong et al. (2019), Saag et al. (2019), Sikora et al. (2018), or Flegontov et al. (2019), and I have tried to add up their information on ancestral components and their modern and historical distributions.

Fortunately, the current obsession with simplifying ancestry components into three or four general, atemporal groups, and the common use of the same ones across labs, make it very simple to merge data and map them.

Corded Ware ancestry

There is no doubt about the prevalent ancestry among Uralic-speaking peoples. A map isn’t needed to realize that, because ancient and modern data – like those recently summarized in Jeong et al. (2019) – prove it. But maps sure help visualize their intricate relationship better:

natural-modern-srubnaya-ancestry
Natural neighbor interpolation of Srubnaya ancestry among modern populations. See full map.
kriging-modern-srubnaya-ancestry
Kriging interpolation of Srubnaya ancestry among modern populations. See full map

Interestingly, the regions with higher Corded Ware-related ancestry are in great part coincident with (pre)historical Finno-Ugric-speaking territories:

uralic-languages-modern
Modern distribution of Uralic languages, with ancient territory (in the Common Era) labelled and delimited by a red line. For more information on the ancient territory see here.

Edit (29/7/2019): Here is the full Steppe_MLBA ancestry map, including Steppe_MLBA (vs. Indus Periphery vs. Onge) in modern South Asian populations from Narasimhan et al. (2018), apart from the ‘Srubnaya component’ in North Eurasian populations. ‘Dummy’ variables (with 0% ancestry) have been included to the south and east of the map to avoid weird interpolations of Steppe_MLBA into Africa and East Asia.

modern-steppe-mlba-ancestry2
Natural neighbor interpolation of Steppe MLBA-like ancestry among modern populations. See full map.

Anatolia Neolithic ancestry

Also interesting are the patterns of non-CWC-related ancestry, in particular the apparent wedge created by expanding East Slavs, which seems to reflect the intrusion of central(-eastern) European ancestry into Finno-Permic territory.

NOTE. Read more on Balto-Slavic hydrotoponymy, on the cradle of Russians as a Finno-Permic hotspot, and about Pre-Slavic languages in North-West Russia.

natural-modern-lbk-en-ancestry
Natural neighbor interpolation of LBK EN ancestry among modern populations. See full map.
kriging-modern-lbk-en-ancestry
Kriging interpolation of LBK EN ancestry among modern populations. See full map

WHG ancestry

The cline(s) between WHG, EHG, ANE, Nganasan, and Baikal HG are also simplified when some of them excluded, in this case EHG, represented thus in part by WHG, and in part by more eastern ancestries (see below).

modern-whg-ancestry
Natural neighbor interpolation of WHG ancestry among modern populations. See full map.
kriging-modern-whg-ancestry
Kriging interpolation of WHG ancestry among modern populations. See full map.

Arctic, Tundra or Forest-steppe?

Data on Nganasan-related vs. ANE vs. Baikal HG/Ulchi-related ancestry is difficult to map properly, because both ancestry components are usually reported as mutually exclusive, when they are in fact clearly related in an ancestral cline formed by different ancient North Eurasian populations from Siberia.

When it comes to ascertaining the origin of the multiple CWC-related clines among Uralic-speaking peoples, the question is thus how to properly distinguish the proportions of WHG-, EHG-, Nganasan-, ANE or BaikalHG-related ancestral components in North Eurasia, i.e. how did each dialectal group admix with regional groups which formed part of these clines east and west of the Urals.

The truth is, one ought to test specific ancient samples for each “Siberian” ancestry found in the different Uralic dialectal groups, but the simplistic “Siberian” label somehow gets a pass in many papers (see a recent example).

Below qpAdm results with best fits for Ulchi ancestry, Afontova Gora 3 ancestry, and Nganasan ancestry, but some populations show good fits for both and with similar proportions, so selecting one necessarily simplifies the distribution of both.

Ulchi ancestry

modern-ulchi-ancestry
Natural neighbor interpolation of Ulchi ancestry among modern populations. See full map.
kriging-modern-ulchi-ancestry
Kriging interpolation of Ulchi ancestry among modern populations. See full map.

ANE ancestry

natural-modern-ane-ancestry
Natural neighbor interpolation of ANE ancestry among modern populations. See full map.
kriging-modern-ane-ancestry
Kriging interpolation of ANE ancestry among modern populations. See full map.

Nganasan ancestry

modern-nganasan-ancestry
Natural neighbor interpolation of Nganasan ancestry among modern populations. See full map.
kriging-modern-nganasan-ancestry
Kriging interpolation of Nganasan ancestry among modern populations. See full map.

Iran Chalcolithic

A simplistic Iran Chalcolithic-related ancestry is also seen in the Altaic cline(s) which (like Corded Ware ancestry) expanded from Central Asia into Europe – apart from its historical distribution south of the Caucasus:

modern-iran-chal-ancestry
Natural neighbor interpolation of Iran Neolithic ancestry among modern populations. See full map.
kriging-modern-iran-neolithic-ancestry
Kriging interpolation of Iran Chalcolithic ancestry among modern populations. See full map.

Other models

The first question I imagine some would like to know is: what about other models? Do they show the same results? Here is the simplistic combination of ancestry components published in Damgaard et al. (2018) for the same or similar populations:

NOTE. As you can see, their selection of EHG vs. WHG vs. Nganasan vs. Natufian vs. Clovis of is of little use, but corroborate the results from other papers, and show some interesting patterns in combination with those above.

EHG

damgaard-modern-ehg-ancestry
Natural neighbor interpolation of EHG ancestry among modern populations, data from Damgaard et al. (2018). See full map.
damgaard-kriging-ehg-ancestry
Kriging interpolation of EHG ancestry among modern populations. See full map.

Natufian ancestry

damgaard-modern-natufian-ancestry
Natural neighbor interpolation of Natufian ancestry among modern populations, data from Damgaard et al. (2018). See full map.
damgaard-kriging-natufian-ancestry
Kriging interpolation of Natufian ancestry among modern populations. See full map.

WHG ancestry

damgaard-modern-whg-ancestry
Natural neighbor interpolation of WHG ancestry among modern populations, data from Damgaard et al. (2018). See full map.
damgaard-kriging-whg-ancestry
Kriging interpolation of WHG ancestry among modern populations. See full map.

Baikal HG ancestry

damgaard-modern-baikalhg-ancestry
Natural neighbor interpolation of Baikal hunter-gatherer ancestry among modern populations, data from Damgaard et al. (2018). See full map.
damgaard-kriging-baikal-hg-ancestry
Kriging interpolation of Baikal HG ancestry among modern populations. See full map.

Ancient North Eurasians

Once the modern situation is clear, relevant questions are, for example, whether EHG-, WHG-, ANE, Nganasan-, and/or Baikal HG-related meta-populations expanded or became integrated into Uralic-speaking territories.

When did these admixture/migration events happen?

How did the ancient distribution or expansion of Palaeo-Arctic, Baikalic, and/or Altaic peoples affect the current distribution of the so-called “Siberian” ancestry, and of hg. N1a, in each specific population?

NOTE. A little excursus is necessary, because the calculated repetition of a hypothetic opposition “N1a vs. R1a” doesn’t make this dichotomy real:

  1. There was not a single ethnolinguistic community represented by hg. R1a after the initial expansion of Eastern Corded Ware groups, or by hg. N1a-L392 after its initial expansion in Siberia:
  2. Different subclades became incorporated in different ways into Bronze Age and Iron Age communities, most of which without an ethnolinguistic change. For example, N1a subclades became incorporated into North Eurasian populations of different languages, reaching Uralic- and Indo-European-speaking territories of north-eastern Europe during the late Iron Age, at a time when their ancestral origin or language in Siberia was impossible to ascertain. Just like the mix found among Proto-Germanic peoples (R1b, R1a, and I1)* or among Slavic peoples (I2a, E1b, R1a)*, the mix of many Uralic groups showing specific percentages of R1a, N1a, or Q subclades* reflect more or less recent admixture or acculturation events with little impact on their languages.

*other typically northern and eastern European haplogroups are also represented in early Germanic (N1a, I2, E1b, J, G2), Slavic (I1, G2, J) and Finno-Permic (I1, R1b, J) peoples.

ananino-culture-new
Map of archaeological cultures in north-eastern Europe ca. 8th-3rd centuries BC. [The Mid-Volga Akozino group not depicted] Shaded area represents the Ananino cultural-historical society. Fading purple arrows represent likely stepped movements of subclades of haplogroup N for centuries (e.g. Siberian → Ananino → Akozino → Fennoscandia [N-VL29]; Circum-Arctic → forest-steppe [N1, N2]; etc.). Blue arrows represent eventual expansions of Uralic peoples to the north. Modified image from Vasilyev (2002).

The problem with mapping the ancestry of the available sampling of ancient populations is that we lack proper temporal and regional transects. The maps that follow include cultures roughly divided into either “Bronze Age” or “Iron Age” groups, although the difference between samples may span up to 2,000 years.

NOTE. Rough estimates for more external groups (viz. Sweden Battle Axe/Gotland_A for the NW, Srubna from the North Pontic area for the SW, Arctic/Nganasan for the NE, and Baikal EBA/”Ulchi-like” for the SE) have been included to offer a wider interpolated area using data already known.

Bronze Age

Similar to modern populations, the selection of best fit “Siberian” ancestry between Baikal HG vs. Nganasan, both potentially ± ANE (AG3), is an oversimplification that needs to be addressed in future papers.

Corded Ware ancestry

bronze-age-corded-ware-ancestry
Natural neighbor interpolation of Srubnaya ancestry among Bronze Age populations. See full map.

Nganasan-like ancestry

bronze-age-nganasan-like-ancestry
Natural neighbor interpolation of Nganasan-like ancestry among Bronze Age populations. See full map.

Baikal HG ancestry

bronze-age-baikal-hg-ancestry
Natural neighbor interpolation of Baikal Hunter-Gatherer ancestry among Bronze Age populations. See full map.

Afontova Gora 3 ancestry

bronze-age-afontova-gora-ancestry
Natural neighbor interpolation of Afontova Gora 3 ancestry among Bronze Age populations. See full map.

Iron Age

Corded Ware ancestry

Interestingly, the moderate expansion of Corded Ware-related ancestry from the south during the Iron Age may be related to the expansion of hg. N1a-VL29 into the chiefdom-based system of north-eastern Europe, including Ananyino/Akozino and later expanding Akozino warrior-traders around the Baltic Sea.

NOTE. The samples from Levänluhta are centuries older than those from Estonia (and Ingria), and those from Chalmny Varre are modern ones, so this region has to be read as a south-west to north-east distribution from the Iron Age to modern times.

iron-age-corded-ware-ancestry
Natural neighbor interpolation of Srubnaya ancestry among Iron Age populations. See full map.

Baikal HG-like ancestry

The fact that this Baltic N1a-VL29 branch belongs in a group together with typically Avar N1a-B197 supports the Altaic origin of the parent group, which is possibly related to the expansion of Baikalic ancestry and Iron Age nomads:

iron-age-baikal-ancestry
Natural neighbor interpolation of Baikal HG ancestry among Iron Age populations. See full map.

Nganasan-like ancestry

The dilution of Nganasan-like ancestry in an Arctic region featuring “Siberian” ancestry and hg. N1a-L392 at least since the Bronze Age supports the integration of hg. N1a-Z1934, sister clade of Ugric N1a-Z1936, into populations west and east of the Urals with the expansion of Uralic languages to the north into the Tundra region (see here).

The integration of N1a-Z1934 lineages into Finnic-speaking peoples after their migration to the north and east, and the displacement or acculturation of Saami from their ancestral homeland, coinciding with known genetic bottlenecks among Finns, is yet another proof of this evolution:

iron-age-nganasan-ancestry
Natural neighbor interpolation of Nganasan ancestry among Iron Age populations. See full map.

WHG ancestry

Similarly, WHG ancestry doesn’t seem to be related to important population movements throughout the Bronze Age, which excludes the multiple North Eurasian populations that will be found along the clines formed by WHG, EHG, ANE, Nganasan, Baikal HG ancestry as forming part of the Uralic ethnogenesis, although they may be relevant to follow later regional movements of specific populations.

iron-age-whg-ancestry
Natural neighbor interpolation of WHG ancestry among Iron Age populations. See full map.

Conclusion

It seems natural that people used to look at maps of haplogroup distribution from the 2000s, coupled with modern language distributions, and would try to interpret them in a certain way, reaching thus the wrong conclusions whose consequences are especially visible today when ancient DNA keeps contradicting them.

In hindsight, though, assuming that Balto-Slavs expanded with Corded Ware and hg. R1a, or that Uralians expanded with “Siberian” ancestry and hg. N1a, was as absurd as looking at maps of ancestry and haplogroup distribution of ancient and modern Native Americans, trying to divide them into “Germanic” or “Iberian”…

The evolution of each specific region and cultural group of North Eurasia is far from being clear. However, the general trend speaks clearly in favour of an ancient, Bronze Age distribution of North Eurasian ancestry and haplogroups that have decreased, diluted, or become incorporated into expanding Uralians of Corded Ware ancestry, occasionally spreading with inter-regional expansions of local groups.

Given the relatively recent push of Altaic and Indo-European languages into ancestral Uralic-speaking territories, only the ancient Corded Ware expansion remains compatible with the spread of Uralic languages into their historical distribution.

Related

Villabruna cluster in Late Epigravettian Sicily supports South Italian corridor for R1b-V88

epipalaeolithic-whg-expansion

New preprint Late Upper Palaeolithic hunter-gatherers in the Central Mediterranean: new archaeological and genetic data from the Late Epigravettian burial Oriente C (Favignana, Sicily), by Catalano et al. bioRxiv (2019).

Interesting excerpts (emphasis mine):

Grotta d’Oriente is a small coastal cave located on the island of Favignana, the largest (~20 km2) of a group of small islands forming the Egadi Archipelago, ~5 km from the NW coast of Sicily.

The Oriente C funeral pit opens in the lower portion of layer 7, specifically sublayer 7D. Two radiocarbon dates on charcoal from the sublayers 7D (12149±65 uncal. BP) and 7E, 12132±80 uncal. BP are consistent with the associated Late Epigravettian lithic assemblages (Lo Vetro and Martini, 2012; Martini et al., 2012b) and refer the burial to a period between about 14200-13800 cal. BP, when Favignana was connected to the main island (Agnesi et al., 1993; Antonioli et al., 2002; Mannino et al. 2014).

sicily-grotta-oriente
A-B) Geographic location of Grotta d’Oriente.

The anatomical features of Oriente C are close to those of Late Upper Palaeolithic populations of the Mediterranean and show strong affinity with other Palaeolithic individuals of Sicily. As suggested by Henke (1989) and Fabbri (1995) the hunter-gatherer populations were morphologically rather uniform.

Genetic analysis

We confirmed the originally reported mitochondrial haplogroup assignment of U2’3’4’7’8’9. This haplogroup is present in both pre- and post-LGM populations, but is rare by the Mesolithic, when U5 dominates (Posth et al.2016).

Lipson et al. (2018) (their supplementary Figure S5.1) and Villalba-Mouco et al. (2019) (their Figure 2A) showed that European Late Palaeolithic and Mesolithic hunter-gatherers fall along two main axes of genetic variation. Multidimensional scaling (MDS) of f3-statistics shows that these axes form a “V” shape (Fig. 3). (…)

Focusing further on Oriente C, we find that it shares most drift with individuals from Northern Italy, Switzerland and Luxembourg, and less with individuals from Iberia, Scandinavia, and East and Southeast Europe (Fig. 4A-B). Shared drift decreases significantly with distance (Fig. 4C) and with time (Fig. 4D) although in a linear model of drift with distance and time as a covariate, only distance (p=1.3×10-6) and not time (p=0.11) is significant. Consistent with the overall E-W cline in hunter-gatherer ancestry, genetic distance to Oriente C increases more rapidly with longitude than latitude, although this may also be affected by geographic features. For example, Oriente C shares significantly more drift with the 8,000 year-old 1,400 km distant individual from Loschbour in Luxembourg (Lazaridis et al.,2014), than with the 9,000 year old individual from Vela Spila in Croatia (Mathieson et al.,2018) only 700 km away as shown by the D-statistic (Patterson et al.,2012) D (Mbuti, Oriente C, Vela Spila, Villabruna); Z=3.42. Oriente C’s heterozygosity was slightly lower than Villabruna (14% lower at 1240k transversion sites), but this difference is not significant (bootstrap P=0.12).

oriente-c-villabruna-f3-statistics
Multidimensional scaling of outgroup f3-statistics for Late 531 Upper Palaeolithic and Mesolithic hunter-gatherers.

Discussion and Conclusion

The robust record of radiocarbon dates proves that they reached Sicily not before 15-14 ka cal. BP, several millennia after the LGM peak. In our opinion, in fact, the hypothesis about an early colonization of Sicily by Aurignacians (Laplace, 1964; Chilardi et al., 1996) must be rejected, on the basis of a recent reinterpretation of the techno-typological features of the lithic industries from Riparo di Fontana Nuova (Martini et al., 2007; Lo Vetro and Martini, 2012; on this topic see also Di Maida et al., 2019).

These analyses have implications for understanding the origin and diffusion of the hunter-gatherers that inhabited Europe during the Late Upper Palaeolithic and Mesolithic. Our findings indicate that Oriente C shows a strong genetic relationship with Western European Late Upper Palaeolithic and Mesolithic hunter-gatherers, suggesting that the “Western hunter-gatherers” was a homogeneous population widely distributed in the Central Mediterranean, presumably as a consequence of continuous gene flow among different groups, or a range expansion following the LGM.

shared-drift-whg-villabruna-oriente-c
The same statistic as in A plotted with geographic position

The South Italian corridor

Once again, a hypothesis based on phylogeography – apart from scarce archaeological and palaeolinguistic data (“Semitic”-like topo-hydronymy and substrates in Europe) – seems to be confirmed step by step. Since the finding of the Villabruna individual of hg. R1b-L754 (likely R1b-V88, like south-eastern European lineages expanded with WHG ancestry), it was quite likely to find out that southern Europe was the origin of the expansion of R1b-V88 into Africa.

The most likely explanation for the presence of “archaic” R1b-V88 subclades among modern Sardinians was, therefore, that they represented a remnant from a Late Upper Palaeolithic/Early Mesolithic population that had not been replaced in subsequent migrations, and thus that the migration of these lineages into Northern Africa and the Green Sahara happened during a period when Italy was connected by a shallower Mediterranean (and more land connections) to Northern Africa.

late-epigravettian
Likely Late Epigravettian/Mesolithic expansion of R1b-V88 into Northern Africa. See full map.

Nevertheless, the arguments for a quite recent expansion of R1b-V88 through the Mediterranean and into Africa keep being repeated, probably based on ancestry from the few ancient (and many modern) populations that have been investigated to date, a simplistic approach prone to important errors that overarch whole migration models.

For example, in the recent paper by Marcus et al. (2019) the presence of these lineages among ancient Sardinians (from the late 4th millennium BC on) is interpreted as an expansion of R1b-V88 with the Cardial Neolithic based on their ancestry, disregarding the millennia-long gap between these samples and the presence of this haplogroup in Palaeolithic/Mesolithic Northern Iberia and Northern Italy, and the comparatively much earlier splits in the phylogenetic tree and dispersal among African populations.

Afroasiatic and Nostratic

I was asked recently if I really believed that we could reconstruct Proto-Nostratic and connect it with any ancestral population. My answer is simple: until the Chalcolithic – when the whole picture of Indo-Europeans, Uralians, Egyptians or Semites becomes quite clear – we have just very few (linguistic, archaeological, genetic) dots which we would like to connect, and we do so the best we can. The earlier the population and proto-language, the more difficult this task becomes.

NOTE. 1) I tentatively connected hg. R with Nostratic in a previous text – when it appeared that R1a expanded from around Lake Baikal, hence Eurasiatic; R1b from the south with AME-WHG ancestry, hence Afroasiatic; and R2 with Dravidian.

2) After that, I though it was more likely to be connected to AME ancestry and the Middle East, because of the apparent expansion of WHG from south-eastern Europe, and the potential association of Afroasiatic and (Elamo-?)Dravidian to Middle Eastern populations.

3) However, after finding more and more R1b samples expanding through northern Eurasia, spreading through the (then wider) steppe regions; and R1a essentially surviving among other groups in eastern Europe for thousands of years without being associated to significant migrations (like, say, hg. C after the Palaeolithic), it didn’t seem like this division was accurate, hence my most recent version.

But, in essence, it’s all about connecting the dots, and we have very few of them…

eurasiatic-phylum-ultraconserved-words
Phylogenetic tree from Pagel et al. (2013), partially in agreement with Kortlandt’s view on Eurasiatic. “Consensus phylogenetic tree of Eurasiatic superfamily (A) superimposed on Eurasia and (B) rooted tree with estimated dates of origin of families and of superfamily. (A) Unrooted consensus tree with branch lengths (solid lines) shown to scale and illustrating the correspondence between the tree and the contemporary north-south and east-west geographical positions of these language families. Abbreviations: P (proto) followed by initials of language family: PD, proto-Dravidian; PK, proto-Kartvelian; PU, proto-Uralic; PIE, proto–Indo-European; PA, proto-Altaic; PCK, proto–Chukchi-Kamchatkan; PIY, proto–Inuit-Yupik. The dotted line to PIY extends the inferred branch length into the area in which Inuit-Yupik languages are currently spoken: it is not a measure of divergence. The cross-hatched line to PK indicates that branch has been shortened (compare with B). The branch to proto-Dravidian ends in an area that Dravidian populations are thought to have occupied before the arrival of Indo-Europeans (see main text). (B) Consensus tree rooted using proto-Dravidian as the outgroup. The age at the root is 14.45 ± 1.75 kya (95% CI = 11.72–18.38 kya) or a slightly older 15.61 ± 2.29 kya (95% CI = 11.72–20.40 kya) if the tree is rooted with proto-Kartvelian. The age assumes midpoint rooting along the branch leading to proto-Dravidian (rooting closer to PD would produce an older root, and vice versa), and takes into account uncertainty around proto–Indo-European date of 8,700 ± 544 (SD) y following ref. 35 and the PCK date of 692 ± 67 (SD) y ago.”

In linguistics, I trust traditional linguists who tend to trust other more experimental linguists (like Hyllested or Kortlandt) who consider that – in their experience – an Indo-Uralic and a Eurasiatic phylum can be reconstructed. Similarly, linguists like Kortlandt are apparently (partially) supportive of attempts like that of Allan Bomhard with Nostratic – although almost everyone is critic of the Muscovite school‘s attachment to the Brugmannian reconstruction, stuck in pre-laryngeal Proto-Indo-Anatolian and similar archaisms.

I mostly use Nostratic as a way to give a simplistic ethnolinguistic label to the genetically related prehistoric peoples whose languages we will probably never know. I think it’s becoming clear that the strongest connection right now with the expansion of potential Eurasiatic dialects is offered by ANE-related populations (hence Y-chromosome bottlenecks under hg. R, Q, probably also N), however complicated the reconstruction of that hypothetic community (and its dialectalization) may be.

Therefore, the multiple expansions of lineages more or less closely associated to ANE-related peoples – like R1b-V88 in the case of Afrasian, or R2 in the case of Dravidians – are the easiest to link to the traditionally described Nostratic dialects and their highly hypothetic relationship.

green-sahara-neolithic
Reconstruction of North African vegetation during past green Sahara periods. Estimated and reconstructed MAP for the Holocene GSP (6–10 kyr BP) projected onto a cross-section along the eastern Sahara (left panel) and map view of reconstructed MAP, vegetation and physiographic elements [7,8,11,45] (right panel). Image from Larrasoaña et al. (2013).

What should be clear to anyone is that the attempt of many modern Afroasiatic speakers to connect their language to their own (or their own community’s main) haplogroups, frequently E and/or J, is flawed for many reasons; it was simplistic in the 2000s, but it is absurd after the advent of ancient DNA investigation and more recent investigation on SNP mutation rates. R1b-V88 should have been on the table of discussions about the expansion of Afroasiatic communities through the Green Sahara long ago, whether one supports a Nostratic phylum or not.

The fact that the role of R1b bottlenecks and expansions in the spread of Afroasiatic is usually not even discussed despite their likely connection with the most recent population expansions through the Green Sahara fitting a reasonable time frame for Proto-Afroasiatic reconstruction, a reasonable geographical homeland, and a compatible dialectal division – unlike many other proposed (E or J) subclades – reveals (once again) a lot about the reasons behind amateur interest in genetics.

Just like seeing the fixation in (and immobility of) recent writings about the role of I1, I2, or (more recently) R1a in the Proto-Indo-European expansion, R1b with Vasconic, or N1c with Proto-Uralic.

NOTE. That evident interest notwithstanding, it is undeniable that we have a much better understanding of the expansions of R1b subclades than other haplogroups, probably due in great part to the easier recovery of ancient DNA from Eurasia (and Europe in particular), for many different – sociopolitical, geographical, technological – reasons. It is quite possible that a more thorough temporal transect of ancient DNA from the Middle East and Africa might radically change our understanding of population movements, especially those related to the Afroasiatic expansion. I am referring in this post to interpretations based on the data we currently have, despite that potential R1b-based bias.

Related

Sea Peoples behind Philistines were Aegeans, including R1b-M269 lineages

New open access paper Ancient DNA sheds light on the genetic origins of early Iron Age Philistines, by Feldman et al. Science Advances (2019) 5(7):eaax0061.

Interesting excerpts (modified for clarity, emphasis mine):

Here, we report genome-wide data from human remains excavated at the ancient seaport of Ashkelon, forming a genetic time series encompassing the Bronze to Iron Age transition. We find that all three Ashkelon populations derive most of their ancestry from the local Levantine gene pool. The early Iron Age population was distinct in its high genetic affinity to European-derived populations and in the high variation of that affinity, suggesting that a gene flow from a European-related gene pool entered Ashkelon either at the end of the Bronze Age or at the beginning of the Iron Age. Of the available contemporaneous populations, we model the southern European gene pool as the best proxy for this incoming gene flow. Last, we observe that the excess European affinity of the early Iron Age individuals does not persist in the later Iron Age population, suggesting that it had a limited genetic impact on the long-term population structure of the people in Ashkelon.

philistines-pca
Ancient genomes (marked with color-filled symbols) projected onto the principal components inferred from present-day west Eurasians (gray circles). The newly reported Ashkelon populations are annotated in the upper corner.

Genetic discontinuity between the Bronze Age and the early Iron Age people of Ashkelon

In comparison to ASH_LBA, the four ASH_IA1 individuals from the following Iron Age I period are, on average, shifted along PC1 toward the European cline and are more spread out along PC1, overlapping with ASH_LBA on one extreme and with the Greek Late Bronze Age “S_Greece_LBA” on the other. Similarly, genetic clustering assigns ASH_IA1 with an average of 14% contribution from a cluster maximized in the Mesolithic European hunter-gatherers labeled “WHG” (shown in blue in Fig. 2B) (15, 22, 26). This component is inferred only in small proportions in earlier Bronze Age Levantine populations (2 to 9%).

In agreement with the PCA and ADMIXTURE results, only European hunter-gatherers (including WHG) and populations sharing a history of genetic admixture with European hunter-gatherers (e.g., as European Neolithic and post-Neolithic populations) produced significantly positive f4-statistics (Z ≥ 3), suggesting that, compared to ASH_LBA, ASH_IA1 has additional European-related ancestry.

We find that the PC1 coordinates positively correlate with the proportion of WHG ancestry modeled in the Ashkelon individuals, suggesting that WHG reasonably tag a European-related ancestral component within the ASH_IA1 individuals.

philistines-admixture
We plot the ancestral proportions of the Ashkelon individuals inferred by qpAdm using Iran_ChL, Levant_ChL, and WHG as sources ±1 SEs. P values are annotated under each model. In cases when the three-way model failed (χ2P < 0.05), we plot the fitting two-way model. The WHG ancestry is necessary only in ASH_IA1.

The best supported one (χ2P = 0.675) infers that ASH_IA1 derives around 43% of ancestry from the Greek Bronze Age “Crete_Odigitria_BA” (43.1 ± 19.2%) and the rest from the ASH_LBA population.

(…) only the models including “Sardinian,” “Crete_Odigitria_BA,” or “Iberia_BA” as the candidate population provided a good fit (χ2P = 0.715, 49.3 ± 8.5%; χ2P = 0.972, 38.0 ± 22.0%; and χ2P = 0.964, 25.8 ± 9.3%, respectively). We note that, because of geographical and temporal sampling gaps, populations that potentially contributed the “European-related” admixture in ASH_IA1 could be missing from the dataset.

The transient impact of the “European-related” gene flow on the Ashkelon gene pool

The ASH_IA2 individuals are intermediate along PC1 between the ASH_LBA ones and the earlier Bronze Age Levantines (Jordan_EBA/Lebanon_MBA) in the west Eurasian PCA (Fig. 2A). Notably, despite being chronologically closer to ASH_IA1, the ASH_IA2 individuals position closer, on average, to the earlier Bronze Age individuals.

philistines-y-dna
See more information on Y-DNA SNP calls, including ASH067 as R1b-M269 (xL151).

The transient excess of European-related genetic affinity in ASH_IA1 can be explained by two scenarios. The early Iron Age European-related genetic component could have been diluted by either the local Ashkelon population to the undetectable level at the time of the later Iron Age individuals or by a gene flow from a population outside of Ashkelon introduced during the final stages of the early Iron Age or the beginning of the later Iron Age.

By modeling ASH_IA2 as a mixture of ASH_IA1 and earlier Bronze Age Levantines/Late Period Egyptian, we infer a range of 7 to 38% of contribution from ASH_IA1, although no contribution cannot be rejected because of the limited resolution to differentiate between Bronze Age and early Iron Age ancestries in this model.

Hg. R1b-M269 and the Aegean

I already predicted this relationship of Philistines and Aegeans (Greeks in particular) months ago, based on linguistics, archaeology, and phylogeography, although it was (and still is) yet unclear if these paternal lineages might have come from other nearby populations which might be descended from Common Anatolians instead, given the known intense contacts between Helladic and West Anatolian groups.

luwian-civilization-sea-peoples
The alternative view: The Sea Peoples can be traced back to the Aegean, so they could also have consisted of Luwian petty kingdoms, who had formed an alliance and attacked Hatti from the south.

The deduction process for the Greek connection was quite simple:

Palaeo-Balkan populations

We know that R1b-Z2103 expanded with Yamna, including West Yamna settlers: they appear in Vučedol, which means they formed part of the earliest expansion waves of Yamna settlers into the Carpathian Basin, and they also appear scattered among Bell Beakers (apart from dominating East Yamna and Afanasevo), which suggests that they were possibly one of the most successful lineages during the late Repin/early Yamna expansion.

The “Steppe ancestry” associated with I2a-L699 samples among Balkan BA peoples may have also been associated with recent Bronze Age expansions, and this haplogroup’s presence among modern Balkan peoples may also suggest that it expanded with Palaeo-Balkan languages. Nevertheless, we don’t know which specific lineages and “Steppe ancestry” they represent, sadly.

These samples may well be related to remnants of previous Balkan populations like Cernavodă or Ezero, because there has been no peer-reviewed attempt at distinguishing Khvalynsk-/Novodanilovka- from Sredni Stog- from Yamnaya-related populations (see here), and some groups that are associated with this ancestry, like Corded Ware, are known to be culturally distinct from Yamna.

In any case, Proto-Greeks from the southern Balkans (say, Sitagroi IV and related groups) are probably going to show, based on Palaeo-Balkan substrate and Pre-Greek substrate and on the available Mycenaean samples, a process of decreasing proportion of R1b-Z2103 lineages relative to local ones, and a relatively similar cline of Yamna:EEF ancestry from northern to southern areas, at least in the periods closest to the Yamna expansion.

NOTE. The finding of “archaic” R1b-L389 (R1b-V1636) and R1a-M198 subclades among modern Greeks and the likely Neolithic origin of these paternal lineages around the Caucasus suggest that their presence in Greece may be from any of the more recent migrations that have happened between Anatolia and the Balkans, especially during the Common Era, rather than Indo-Anatolian migrations; probably very very recently.

-chalcolithic-late-balkans
Bronze Age cultures in the Balkans and the Aegean. See full map including ancient samples with Y-DNA, mtDNA, and ADMIXTURE.

Minoans and haplogroup J

In the Aegean, it is already evident that the population changed language partly through cultural diffusion, probably through elite domination of Proto-Greek speakers. Whether that happened before the invasion into the Greek Peninsula or after it is unclear, as we discussed recently, because we only have one reported Y-chromosome haplogroup among Mycenaeans, and it is J (probably continuing earlier lineages).

Now we have more samples from the so-called Emporion 2 cluster in Olalde et al. (2019), which shows Mycenaean-like eastern Mediterranean ancestry and 3 (out of 3) samples of haplogroup J, which – given the origin of the colony in Phocea – may be interpreted as the prevalence of West Anatolian-like ancestry and lineages in the eastern part of the Aegean (and possibly thus south Peloponnese), in line with the modern situation.

NOTE. It does not seem likely that those R or R1b-L23 samples from the Emporion 1 cluster are R1b-Z2103, based on their West European-like ancestry, although they still may be, because – as we know – ancestry (unlike haplogroup) changes too easily to interpret it as an ancestral ethnolinguistic marker.

anatolia-greek-aegean
PCA of ancient samples related to the Aegean, with Minoans, Mycenaeans (including the Emporion 2 cluster in the background) Anatolia N-Ch.-BA and Levantine BA-LBA populations, including Tel Shadud samples. See more PCAs of ancient Eurasian populations.

Greeks and haplogroup R1b-M269

Therefore, while the presence of R1b-Z2103 among ancient Balkan peoples connected to the Yamna expansion is clear, one might ask if R1b-Z2103 really spread up to the Peloponnese by the time of the Mycenaean Civilization. That has only one indirect answer, and it’s most likely yes.

We already had some R1b-Z2103 among Thracians and around the Armenoid homeland, which offers another clue at the migration of these lineages from the Balkans. The distribution of different “archaic” R1b-Z2103 subclades among modern Balkan populations and around the Aegean offered more support to this conclusion.

But now we have two interesting ancient populations that bear witness to the likely intrusion of R1b-M269 with Proto-Greeks:

An Ancient Greek of hg. R1b

A single ancient sample supports the increase in R1b-Z2103 among Greeks during the “Dorian” invasions that triggered the Dark Ages and the phenomenon of the Aegean Sea Peoples. It comes from a Greek lab study, showing R1b1b (i.e. R1b-P297 in the old nomenclature) as the only Y-chromosome haplogroup obtained from the sampling of the Gulf of Amurakia ca. 470-30 BC, i.e. before the Roman foundation of Nikopolis, hence from people likely from Anaktorion in Ancient Acarnania, of Corinthian origin.

ancient-greeks-y-dna-mtdna

Even with the few data available – and with the caution necessary for this kind of studies from non-established labs, which may be subject to many different kinds of errors – one could argue that the western Greek areas, which received different waves of migrants from the north and shows a higher distribution of R1b-Z2103 in modern times, was probably more heavily admixed with R1b-Z2103 than southern and eastern areas, which were always dominated by Greek-speaking populations more heavily admixed with locals.

The Dorian invasion and the Greek Dark Ages may thus account for a renewed influx of R1b-Z2103 lineages accompanying the dialects that would eventually help form the Hellenic Koiné. In a sense, it is only natural that demographically stronger populations around the Bronze Age Aegean would suffer a limited (male) population replacement with the succeeding invasions, starting with a higher genetic impact in the north-west and diminishing as they progressed to the south and the east, coupled with stepped admixture events with local populations.

This would be therefore the late equivalent of what happened at the end of the 3rd millennium BC, with Mycenaeans and their genetic continuity with Minoans.

pre-greek-ssos
Distribution of Pre-Greek place-names ending in -ssos/-ssa or -sos/-sa. See original images and more on the south/east cline distribution of Pre-Greek place-names here.

Sea peoples of hg. R1b-M269

Thanks to Wang et al. (2018) supplementary materials we knew that one of the two Levantine LBA II samples from Tel Shadud (final 13th–early 11th c. BC) published in van den Brink (2017) was of hg. R1b-M269 – in fact, the one interpreted as a Canaanite official residing at this site and emulating selected funerary aspects of Egyptian mortuary culture.

Both analyzed samples, this elite individual and a commoner of hg. J buried nearby, were genetically similar and indistinguishable from local populations, though:

Principal Components Analysis of L112 and L126 was carried out within the framework described in Lazaridis et al. (2016). This analysis showed that the two individuals cluster genetically, with similar estimated proportions of ancestry from diverse West Eurasian ancestral sources. These results are consistent with the hypothesis that they derive from the same population, or alternatively that they derive from two quite closely related populations.

We know that ancestry changes easily within a few generations, so there was not much information to go on, except for the fact that – being R1b-M269 – this individual could trace his paternal ancestor at some point to Proto-Indo-Europeans.

One might think that, because many haplogroups in this spreadsheet were wrong, this is also wrong; nevertheless, many haplogroups are correctly identified by Yleaf, and finding R1b-M269 in the Levant after the expansion of Sea Peoples could not be that surprising, because they were most likely related to populations of the Aegean Sea. Any other related hg. R1b (R1b-M73, R1b-V88, even R1b-V1636) wouldn’t fit as well as R1b-M269.

sea-peoples-egypt-rameses-iii

However, the early expansion of Proto-Indo-Aryans into the Middle East, as well as the later expansion of Armenians from the Balkans through Anatolia and of West Iranians from the east may have all potentially been related to this sample. But still, the previous linguistic and archaeological theories concerning the Philistines and the expansion of Sea Peoples in the Levant made this sample a likely (originally) Greek “Dorian” lineage, rather than the other (increasingly speculative) alternatives.

In any case, it was obvious to anyone – that is, to anyone with a minimum knowledge of how population genomics works – that just the two samples from van den Brink (2017) couldn’t be used to get to any conclusions about the ancestral origin of these individuals (or their differences) beyond Levantine peoples, because their ancestry was essentially (i.e. statistically) the same as the other few available ancient samples from nearby regions and similar periods.

If anything, the PCA suggested an origin of the R1b sample closer to Aegean populations relative to the J individual (see PCA above), and this should have been supported also by amateur models, without any possible confirmation (as with the ASH_IA2 cluster in this paper). However, if you have followed online discussions of Tel Shadud R1b-M269 sample since it was mentioned first on Eupedia months ago – including another wave of misguided speculation based on the ancestry of both individuals triggered by a discussion on this blog -, you have once more proof of how misleading ancestry analyses can be in the wrong hands.

NOTE. This is the Nth proof (and that only in 2019) of how it’s best to just avoid amateur analyses and interpretations altogether, as I did in the recent publication of the books. All those who didn’t take into account whatever was commented about the ancestry of these samples haven’t lost a single bit of relevant information on Levantine peoples, and have had more time for useful reads, compared to those dedicated to endless void speculation, once again gone awfully wrong, as does everything related to cocky ancient DNA crackpottery 😉

bronze-age-late-aegean
Late Bronze Age population movements in the Eastern Mediterranean and the Middle East. See full map including ancient DNA samples with Y-DNA, mtDNA, and ADMIXTURE.

Admittedly, though, even accepting the evident Mediterranean origin of this lineage, one could have argued that this sample may have been of R1b-L151 subclade, if one were inclined to support the theory that Italic peoples were behind Sea Peoples expanding east – and consequently that the ancestors of Etruscans had migrated eastward into the Aegean (e.g. into Lemnos), so that it could be asserted that Tyrsenian might have been a remnant language of an ancient population of northern Italy.

Philistines

Fortunately, some of the samples recovered in Feldman et al. (2019) that could be analyzed (those of the cluster ASH_IA1) offer a very specific time frame where European ancestry appeared (ca. 1250 BC) before it subsequently became fully diluted (as seen in cluster ASH_IA2) among the prevalent Levantine ancestry of the area.

Also fortunately, this precise cluster shows another R1b-M269 sample, likely R1b-Z2103 (because it is probably xL151), and this sample together with others from the same cluster prove that the ancestry related to the original southern European incomers was:

  1. Recent, related thus to LBA population movements, as expected; and
  2. More closely related to coeval Aegeans, including Mycenaeans with Steppe-related ancestry.

NOTE. I say “fortunately” because, as you can imagine if you have dealt with amateurish discussions long enough, without this cluster with evident Aegean ancestry and the R1b-M269 (Z2103) sample precisely associated to it, some would enter again in endless comment loops created by ancestry magicians, showing how Aegean peoples were not behind Sea Peoples, or not behind Philistines, or not behind the R1b-M269 among Philistines, depending on their specific agendas.

aegean-sea-peoples
Map of the Sea People invasions in the Aegean Sea and Eastern Mediterranean at the end of the Late Bronze Age (blue arrows).. Some of the major cities impacted by the raids are denoted with historical dates. Inland invasions are represented by purple arrows. From Kaniewski et al. (2011). Some of the major cities impacted by the raids are denoted with historical dates. Inland invasions are represented by purple arrows.

The results of the paper don’t solve the question of the exact origin of all Sea Peoples (not even that of Philistines), but it is quite clear that most of those forming this seafaring confederation must have come from sites around the Aegean Sea. This supports thus the traditional origin attributed to them, including a hint at the likely expansion of Eastern Mediterranean ancestry and lineages into the Italian Peninsula precisely from the Aegean, as some oral communications have already disclosed.

As an indirect conclusion from the findings in this paper, then, we can now more confidently support that Tyrsenian speakers most likely expanded into the Appenines and the Alps originally from a Tyrsenian-speaking LBA population from Lemnos, due to the social unrest in the whole Aegean region, and might have become heavily admixed with local Italic peoples quite quickly, as it happened with Philistines, resulting in yet another case of language expansion through (the simplistically called) elite domination.

Conclusion

Even more interesting than these specific findings, this paper confirms yet another hypothesis based on phylogeography, and proves once again two important starting points for ancient DNA interpretation that I have discussed extensively in this blog:

  • The rare R1b-M269 Y-chromosome lineage of Tel Shadud offered ipso facto the most relevant clue about the ancestral geographical origin of this Canaanite elite male’s paternal family, most likely from the north-west based on ancient phylogeography, which indirectly – in combination with linguistics and archaeology – supported the ancestral ethnolinguistic identification of Philistines with the Aegean and thus with (a population closest to) Ancient Greeks.
  • Ancestry analyses are often fully unreliable when assessing population movements, especially when few samples from incomplete temporal-geographical transects are assessed in isolation, because – unlike paternal (and maternal) haplogroups – ancestry might change fully within a few generations, depending on the particular anthropological setting. Their investigation is thus bound by many limitations – of design, statistical, and anthropological (i.e. archaeological and linguistic) – which are quite often not taken into account.

These cornerstones of ancient DNA interpretation have been already demonstrated to be valid not only for Levantine populations, as in this case, but also for Balkan peoples, for Bell Beakers, for steppe populations (like Khvalynsk, Sredni Stog, Yamna, Corded Ware), for Basques, for Balto-Slavs, for Ugrians and Samoyeds, and for many other prehistoric peoples.

I rest my case.

Related

Genetic continuity among Uralic-speaking cultures in north-eastern Europe

east-europe-bronze-age

The recent study of Estonian Late Bronze Age/Iron Age samples has shown, as expected, large genetic continuity of Corded Ware populations in the East Baltic area, where West Uralic is known to have been spoken since at least the Early Bronze Age.

The most interesting news was that, unexpectedly for many, the impact of “Siberian ancestry” (whatever that actually means) was small, slow, and gradual, with slight increases found up to the Middle Ages, compatible with multiple contact events in north-eastern Europe. Haplogroup N became prevalent among Finnic populations only through late bottlenecks, as research of modern populations have long suggested, and as ancient DNA research hinted since at least 2015.

I risked to correlate the arrival of chiefs from the south-west with the infiltration of N1c-VL29 subclades during the transition to the Iron Age, coupled with that minimal “Siberian” ancestry (see e.g. here and here). Now we know that the penetration of this non-CW ancestry started, as predicted, in the Iron Age; that it was highly variable in the few samples where it appeared, with ca. 1-4%, while most Iron Age individuals show 0%; and that it was not especially linked to individuals of N1c-Vl29 lineages.

It is also basically confirmed, based on the (ancient and Modern Swedish) N1c-L550 subclades found among Iron Age Estonians, that N1c-VL29 lineages and the so-called “Siberian” ancestry will be found simultaneously around the Baltic coastal areas, and that different lineages must have suffered later founder effects among Finns, which suggests that these alliances through exogamy brought exactly as much language change in Sweden, Lithuania, or Poland, as they did in the East Baltic region…

On the other hand, the paper has also shown a potential movement of Corded Ware-derived peoples, if the change from LBA to IA samples is meaningful; in fact, even more Corded Ware-like than Baltic and Estonian BA populations. The exact origin of that movement is difficult to pinpoint, and it may not be related to the arrival of Akozino warrior-traders from the south-east, since theirs seems to be a minor impact proper of elites in a chiefdom system around the Baltic.

fortified-settlements-lba-ia
Distribution of fortified settlements (filled circles) and other hilltop sites (empty circles) of the Late Bronze Age and Pre-Roman Iron Ages in the East Baltic region. Tentative area of most intensive contacts between Baltic and Balto-Finnic communities marked with a dashed line. Image modified from (Lang 2016).

Also suggesting a potential movement is the ‘southern’ shift observed in the West and East Baltic areas, likely showing the arrival of Proto-East Baltic speakers (such as the Trzciniec outlier), as we have already discussed in this blog. The unexpected increase in Corded Ware-like ancestry in the Eastern Baltic, coupled with the expected large continuity of hg. R1a-Z283 in the homeland of Balto-Finnic expansions, gives even more support to the known complex system of exogamy along the Baltic coasts, and offers another potential reason for the rise of Baltic-speaking territories in the West Baltic: elite domination.

It is nevertheless important to understand that, even among the most “genetic continuous” regions like Estonia, not a single population in Europe is heir of some ancestral, immutable people. Not in terms of haplogroups, and not in terms of admixture. Balto-Finnic speakers, however continuous they might seem (e.g. in Southern Estonians) aren’t an exception.

After all, this blog was (re)born to fight the currently prevalent sheer stupidity surrounding the simplistic “R1a/steppe ancestry=Indo-European” association, so I wouldn’t like to see it replaced with some other stupid continuity or purity ideas within 10 to 20 years…

Late Uralic stems from East Corded Ware groups

With the currently available tools – linguistics, archaeology, and now genetics -, I don’t think there is any argument to date to question the direct connection of the Late Proto-Uralic expansion with all Eastern Corded Ware groups (i.e. Battle Axe, Fatyanovo-Balanovo, and Abashevo), and thus at least with the unifying A-horizon of Corded Ware and the bottlenecks under R1a-Z645.

NOTE. The only out-group among Corded Ware cultures is the Single Grave culture. It appears to be an early Corded Ware offshoot, reflected in their non-unitary cultural traits (distinct from later unifying waves), in their varied patrilineal clans, and in the short-lasting cultural effect in northern Europe before their complete demise under pressure of expanding Yamna/Bell Beaker peoples from the Danube. The culture’s minimal (if any) effects on succeeding peoples might be seen mostly in the (mainly phonetic) Uralic substrate found in Balto-Slavic – although this may also stem from a more eastern influence, close to the Baltic – and in the contacts of Celtic with Uralic. The huge time depth between this early hypothetic Uralic layer in northern Europe and the emergence of peoples inhabiting these territories in recorded history have no doubt been erroneously interpreted as a lack of Uralic presence in the area.

1) That connection was evident in the Yamna – CWC differences in archaeology, and especially later, with at least Fatyanovo-Balanovo and Abashevo representing the obvious replacement of the Volosovo culture before further expansions of CWC-related groups west and east of the Urals.

The mythical millennia-long continuity of Volosovo hunter-gatherers, including centuries among Corded Ware peoples, as expected lately by the Copenhagen group (and anyone who doesn’t want to question the 1960s association of Indo-European with CWC) must be rejected today in population genomics, as the recent studies of ancient and modern populations show, and as ancient DNA from the region will confirm.

2) In linguistics, the survival of Volosovo as The Uralic-speaking culture was also hardly believable. From Kallio (2015):

While we can say at least something about Uralic substrates in Northeastern Europe, non-Uralic substrates cannot at all easily be identified, because of multiple language shifts, viz. first from non-Uralic to Uralic and then from Uralic to Russian. Yet the Soviet Uralicist Boris Serebrennikov (1956, 1959) argued that there are some non-Uralic substrate toponyms in the Volga-Oka region, but his idea was never taken seriously in the west (cf. Sauvageot 1958), and it pretty soon also sank into oblivion in Russia, even though it can still occasionally pop up there in non-onomastic circles (cf. Napolskikh 1995: 18–19). However, not all the hypotheses on non-Uralic substrates in Northeastern Europe should be rejected (see e.g. Helimski 2001b).

bronze-age-early-languages-east-europe
Tentative map of the distribution of known languages in Eastern Europe during the Early Bronze Age. See full map.

Helimski (2001) argues for a non-Uralic topo-hydronomy in Northern Russia, whose population may have kept their languages up to the Common Era despite the Corded Ware expansion, which is in line with the survival of some non-Indo-European languages everywhere in Europe after the expansion of Yamna and its offshoots:

It should be borne in mind that these [Uralic] hydronyms reached us mainly through Northern Russian and, accordingly, with a tendency to phonetic-morphological adaptation and unification (for river names it is “natural” to be, like the word ‘river’ itself, feminine and to end in -a). Taking into account this circumstance, it may turn out to be non-useless for etymological identification of at least some of the hydronyms on the Finno-Ugric basis.

On the other hand, I wouldn’t exclude the possibility that some parts of this large geographical area were never (completely) Finno-Ugric. The population that created the most important part of the hydronymy of the Russian North could be finally pushed aside or assimilated only at the end of the 1st – beginning of the 2nd millennium AD, during the Russian colonization, retaining the memory of the White-Eyed Chude in its own memory.

NOTE. For more on this non-IE substrate in (especially West) Uralic, see e.g. Zhivlov (2015),

The same non-Uralic substrate is most likely behind most of the shared traits by Mordvinic and Balto-Finnic (see below).

3) In genetics, I don’t think the picture could get any clearer. I don’t know what “Steppe ancestry = Indo-European” proponents expected from 2019, if they expected anything at all (I haven’t seen any coherent model, proposal, or prediction for a long time now), but I doubt the recent results are compatible with any of their implied expectations.

corded-ware-pca-sub-neolithic-europe
Detail of the PCA of the Corded Ware expansion. See full PCA and more related files.

Notice, from the PCA above, how this Baltic Late Neolithic group shows actually a shift from Sredni Stog (see PCA with Sredni Stog) towards typical Khvalynsk-Urals-related ancestry, i.e. populations from eastern European forested regions, derived from hunter-gatherer pottery groups, as I have proposed for a very long time, since the first time a Baltic LN “outlier” appeared. It’s amazing how some amateurs can find 0.1% of any Siberian outlier’s ancestry among Uralians 4,000 years later, but fail to see the direct connection here. The esoteric uses of qpAdm, I guess…

Especially noticeable is the extra WHG-like ancestry and corresponding shift, seen especially marked in late Polish CWC samples, but also in Baltic CWC and especially in one Sweden Battle Axe sample, all of them shifting apparently closer to Pitted Ware and SHG. While that may have been interpreted as an in situ admixture in Scandinavia before, the late Polish CWC samples show likely a resurgence of local populations, so we can assume that both shifts (to SHG- and EHG-like populations) of available CWC samples around the Baltic are clearly part of the WHG:EHG continuum that will be found in the eastern European sub-Neolithic cultures, from Narva to Volosovo.

This WHG-related ancestry is clearly predominant in groups with which Battle Axe peoples admixed, based on the shift towards Pitted Ware, which – I can only guess based on modern Volga Finns – is different from the shift we will see in Netted Ware, more towards the Khvalynsk-Urals cluster. This is in line with the expansion of Battle Axe eastward through coastal areas (West to East Baltic and Finland into Sweden), while Fatyanovo peoples probably emerged from a slightly different route, but also a northern one, if one is to follow archaological similarities and their chronology.

bronze-age-europe-baltic
Detail of the PCA of European Bronze Age populations. See full PCA and more related files.

During the Iron Age, the only peoples that probably shifted strongly (based on modern populations) are West Baltic ones, getting closer to the available Late Trzciniec samples, and even closer to the Trzciniec outlier, i.e. away from the earlier Eastern Corded Ware cluster, and towards Central European groups like Czech EBA or Poland EBA, both of them clearly derived from Bell Beakers, but also admixed with (and thus shifted toward) CW-like populations.

If one looks carefully at the previous PCA on Bronze Age populations, and the next one on Iron Age clusters, it is evident that adding the Swedish LN outlier to East Baltic BA (both strongly related to Battle Axe populations) essentially gives us the continuity of East Baltic BA into the Iron Age. This cluster is continued also in two outliers from Sigtuna, a Viking town close to the Gulf of Finland, known to be an important trading site, 1,500 years later. Not much of a change around the Gulf of Finland, then:

iron-age-eastern-europe
Detail of the PCA of East and North European Iron Age populations. See full PCA and more related files.

Based on the two simplistic Uralic clines one might see described (among the many that certainly existed, from Corded Ware to different Eurasian populations), and just like BOO was for some months fashionable as “Samic”, some may be tempted to say that certain Sintashta or Srubna outliers close to the Urals mark the True Uralic™ peoples. Because, of course they do. Ghost haplogroup N and stuff. And Corded Ware never ever Uralic. Because Gimbutas, and my IE R1a grandfather.

NOTE. Funny thing here: there might be Corded Ware, Iranian, Slavic, Germanic, etc… outliers or out-groups, and they might form the widest genetic clusters ever seen, but they are all of one language, because archaeology and linguistics; however, one “outlier” (also, put your own definition of “outlier” here, let’s say 1% of whatever, and strontium isotope potentially from 100 km away) ca. 600 BC in the Baltic who (surprise!) happens to show hg. N, and he signals the first incoming True Uralic™ speaker from wherever… It won’t be the first or the last time some people resort to “the complexity of Uralic-speaking peoples” in ancestry, just to look for “hg. N = Uralic” like crazy. You only need common sense to understand that this is not how this works. Amateur genomics can’t get more embarrassing than the current “let’s look for ‘Siberian ancestry’ in every individual of haplogroup N” trend. Or maybe it can, and it will, but I can’t see it yet.

If one were to insist on looking for ‘foreign’ contributions among Iron Age Estonians, though, I think one should also check out first archaeology, and then the PC3 (or, more graphically, a 3D plot), to understand what might be happening with the many Uralic clines derived from Corded Ware, before starting to play around with bioinformatic tools to discover a teeny tiny 1% admixture of the wrong population, and rushing to build far-fetched narratives. Apparently, one of the different clines formed roughly between southern (steppe – forest-steppe) and northern (tundra-taiga) populations in Uralians is also seen in some Iron Age Estonian individuals – especially in some late samples from Ingria…This is not my main interest, so I will leave this here for others to keep wasting their time chasing the white whale of the 0.5% of True Uralic™ ancestry in ancient Baltic samples of hg. N.

pca-3d-estonians-iron-age-boo-samic
Still images of the 3D plot of Eurasian samples. Typical PC1 vs. PC2 visualization to the left, and shift of the view to PC3 on the right image. See full PCA and more related files.

An exclusive Volga-Kama homeland for Disintegrating Uralic?

Since I don’t believe in macro-regions of largely continuous ethnolinguistic communities, as I have often said about Slavic (naively associated with prehistoric tribes of Eastern Europe) or Germanic (absurdly considered to be represented by Battle Axe), it is difficult for me to believe that Battle Axe-derived cultures remained of the same Finno-Samic dialects since the Corded Ware expansion…unless we live in Westeros, where everything happens “for thousands of years”.

I have to admit, then, that the now prevalent identification among Uralicists has become quite attractive:

  • Fatyanovo-Balanovo as Finno-Permic:
    • Fatyanovo/Netted Ware with West Uralic (also called Finno-Mordvinic).
    • Balanovo/Chirkovo-Kazan with Central Uralic (Mari-Permic).
  • Abashevo, into the Andronovo-like Horizon through the Seima-Turbino phenomenon, with East Uralic (also Ugro-Samoyedic).

Exactly like the identification of Yamna Hungary – Bell Beaker transition as the North-West Indo-European homeland, it gives us simplicity and small and late ethnolinguistic communities, away from the traditionally overused big and early language territories.

This late homeland would be supported, among others, by:

  • The presence of Indo-Iranian loanwords in Finno-Permic and Ugric (probably also in Samoyedic, either lost, or – much more likely – underresearched), compatible with the immediate contact between Abashevo – Sintashta-Potapovka-Filatovka and Fatyanovo-Balanovo.
  • The supposed expansion of Netted Ware from Fatyanovo to the north-west, which may be explained as the split and expansion of Balto-Finnic and Samic ca. 1900 BC.
  • A longer-lasting Finno-Permic (West+Central Uralic) community contrasting with the early separation of East Uralic.
  • The compatibility of this late expansion with the late expansion of Pre-Germanic from Denmark with the Dagger Period, and of Balto-Slavic with Trzciniec, which puts all three dialects reaching the Baltic Sea in the EBA.

NOTE. I meant to update the linguistic text to include the most recently favoured phylogenetic tree of Uralic languages after Häkkinen (2007, 2009, 2014), which has very quickly become the new normal among Uralicists, but I don’t think I will have enough time to review the necessary papers for that. I am rushing to publish a printed edition, so the text will wind up being a mixture of “traditional” (meaning, basically, pre-2010s) description of Uralic dialects but using modern divisions; say, “West Uralic” instead of “Finno-Samic”. By the way, I am still amazed that none of my reader-haters (or any online user discussing Uralic migrations, for that matter) have come up with the questions that the new division pose, and it supports my suspicion about the complete lack of interest in linguistics of most (a)DNA fans, except for the occasional use of old and free PDFs Googled to support new narratives invented expressly for some qpAdm results…

textile-ceramics-europe-bronze-age
Textile ceramic styles and influence of Bronze Age cultures divided in clusters.

Problems with this Parpola-Carpelan’s (2012-2018) interpretation include:

  • The differentiation between Fennoscandian Textile Ceramics vs. Netted Ware, which is not warranted in archaeology. The assumption that Netted Ware expanded to the Baltic Sea (as Kallio does, following the traditional view) is thus weak, and it was probably a question of cultural contacts coupled with short-distance population movements/exchange in both directions (from the Baltic to the Volga and vice versa). In fact, the culture division relies on some fairly common and technically simple ornamentation patterns, widespread all over northern Europe, even before the Corded Ware expansion, and it is very difficult to separate certain neighboring Textile Ceramics from Netted Ware groups in southern Finland (i.e. Sarsa-Tomitsa groups).
  • The strict and radical direction described for the Netted Ware by Carpelan, as an eastward and northward expansion, within a very short time frame (ca. 1900-1800 BC), based on few radiocarbon dates, which seems to me like a very risky assumption. We know how this kind of descriptions of direction of culture expansion based on radiocarbon dates has turned out in much more complex “packages”, like the Bell Beaker culture… In fact, the earliest dates for Textile Ware are from the East Baltic, earlier than those of Netted Ware.
  • The assumption that Balto-Finnic traits shared with Mordvinic are a) late and b) meaningful for dialectalization of two closely related dialects, when it is clear that both dialects separated quite early. Phonologically Finnic is more conservative, morphologically less so, and the shared traits include a handful of non-Uralic substrate words which can’t be traced to a single common source, hence they were adopted when both languages had already separated… All in all, Finnic – Mordvinic correspondances are not even close to Italo-Celtic ones, which is clearly fully incompatible with a proposal of a Finnic separation from Mordvinic coinciding with the LBA-IA transition.

Especially problematic for Parpola’s model is the lack of genetic impact in Bronze Age or Iron Age Estonians, not reaching a significant level under any possible statistical threshold – which I am sure was quite disappointing for some of my readers -, but is in line with major archaeological continuity of groups the from region, only disturbed in cultural (and Y-chromosome) terms by the expansion of Akozino warrior-traders all over the Baltic Sea. Any proposed population movement will be very difficult to support in genetics, given the Corded Ware-derived populations that we will see in both regions, and the continued Baltic-Volga contacts since the Corded Ware expansion.

Problems with an interpretation of such a small impact in population genomics includes the similarly weak impacts and haplogroup infiltrations that can be seen among populations basically everywhere in Eurasia, during any given period, and much greater genetic impacts that are supposed to be (or that were certainly) followed by ethnolinguistic continuity.

akozino-malar-axes-fennoscandia
Distribution of the Akozino-Mälar axes according to Sergej V. Kuz’minykh (1996: 8, Abb. 2).

The Battle Axe question

From Kallio (2015), about choosing a tentative homeland for Proto-Uralic:

(…) linguistically uniform Proto-Uralic would have been spoken in the Volga-Oka region until the mid-third millennium BC when the Proto-Uralic-speaking area would have expanded to the Volga-Kama region as well. By the end of the same millennium, this expansion would have led to the earliest dialectal splits within Uralic into Finno-Mordvin, Mari-Permic, and Ugro-Samoyed. The splitting up of these three soon followed during the early second millennium BC when the Uralic-speaking area finally stretched from the Baltic Sea in the west to the Altai mountains in the east. Indeed, no matter where Proto-Uralic was spoken, the branching into the nine well-attested subgroups (viz. Finnic, Saami, Mordvin, Mari, Permic, Hungarian, Mansi, Khanty, and Samoyed) must have taken less than a millennium, because their shared phonological and morphosyntactic isoglosses are rather limited (see Salminen 2002). The traditional view that all this branching would have taken several millennia violates everything linguistic typology teaches us about the rate of language change.

The basic problem of this identification of Fatyanovo-Balanovo as West-Central Uralic and Abashevo as East Uralic is the nature of the Battle Axe culture, including the Bronze Age East Baltic and Gulf of Finland area. Even if it is accepted that Fatyanovo-Balanovo represented all Western groups, Battle Axe must have represented West Uralic-like dialects.

The ethnolinguistic identification of Battle Axe depends ultimately on the nature of contacts of Fatyanovo/Netted Ware with Battle Axe/Textile Ceramics. If both groups were close and interacted profusely, as it seems, it doesn’t seem granted that we will be able to distinguish a close Para-West Uralic dialect of Scandinavia from the actual expanding Balto-Finnic and Samic dialects, if they were actually linked to the Netted Ware expansion. Also from Kallio (2015):

No doubt the most convincing substrate theory has recently been put forward by the Saami Uralicist Ante Aikio (2004), who has not only rehabilitated but also improved the old idea of a non-Uralic substrate in Saami. His study shows that there were still non-Uralic languages spoken in Northern Fennoscandia as recently as the first millennium AD. Most of all, they were not only genetically non-Uralic but also typologically non-Uralic-looking, bearing a closer resemblance to the so-called Palaeo-European substrates (for which see e.g. Schrijver 2001; Vennemann 2003).

In comparison, the case of Finnic is much more difficult. The fact that Proto-Uralic was not spoken in the East Baltic region means that this area must have originally been non-Uralic-speaking, but so far the evidence for a non-Uralic substrate in Finnic has consisted of appellatives and proper names with no etymology (cf. Ariste 1971; Saarikivi 2004a). Contrary to the proposed substrate words in Saami, those in Finnic show no structural non-Uralisms, as if they had indeed been borrowed from some genetically related or at least typologically similar languages, as I suggested above. Also none of them is more recent than the Middle Proto-Finnic stage, which makes them at least two millennia old. All this agrees with archaeological evidence discussed earlier that the Uralicization of the East Baltic region occurred during the Bronze Age (ca. 1900–500 BC).

The discussion of the paper continues with an unsuccessful attempt to find a hypothetical ancient Indo-European substrate that Kallio believes must be associated with the expansion of Corded Ware, in line with the traditional belief. For example, the often mentioned – almost folk etymology-like, unsurprisingly popular among amateurs – ‘Neva’ as derived from IE “young” is logically rejected…Unlike Parpola, Kallio’s view seems to be confident that Netted Ware (as Textile Ware) expanded into the East Baltic, on both sides of the Gulf of Finland, already during the Bronze Age.

As it has become apparent in population genomics, none of them was right, and Textile Ceramics will essentially show – like Netted Ware – a large genetic continuity of Corded Ware peoples in the whole north-eastern European forest zone – despite small regional population movements, obviously -, which necessarily implies that the whole Corded Ware culture – and not only Fatyanovo-Balanovo and Abashevo – were Uralic-speaking territories.

The similarities in terms of culture and Y-DNA bottlenecks between Battle Axe and Fatyanovo-Balanovo also imply that the linguistic differences between these groups were probably not many, and became strongly divided only after their territorial division. Continued contacts between Battle Axe- and Fatyanovo-derived groups can explain the proposed contacts (Finnic with Samic, Finnic with Mordvinic) after their linguistic-but-not-physical separation.

east-european-fatyanovocwc
East European movement directions (arrows) of the representatives of the Central European Corded Ware Culture (according to I.I. Artemenko).

Battle Axe spoke “Para-Balto-Finnic”?

The Balto-Finnic-speaking nature of Battle Axe is thus supported by:

  • The lack of non-Uralic substrates in Balto-Finnic territory (Kallio 2015).
  • The early separation of Samic and Finnic from Mordvinic, and the virtual identity of Proto-West-Uralic and Proto-Uralic, which suggests that Proto-Uralic spread fast (Parpola 2012).
  • The scarce non-Uralic topo-hydronymy in the East Baltic and around the Gulf of Finland (Saarikivi 2004), comparable to that on the Upper Volga region.
  • The strong influence of a Balto-Finnic-like substrate on Pre-Germanic (or, in Kallio’s opinion, the same Scandinavian substrate influencing both Germanic and Balto-Finnic at the same time), and the continued influence of Balto-Finnic on Proto-Baltic and Proto-Slavic.
  • The continued influence of Corded Ware-derived groups in central-east Sweden in Finland and the East Baltic in terms of agricultural innovations appearing in the LBA, compatible with Schrijver’s proposal of intermediate Germanic-shifted Balto-Finnic groups and Balto-Finnic groups influenced by their pronunciation.
  • The intense Palaeo-Germanic and late Balto-Slavic / early Proto-Baltic superstrate on Balto-Finnic, which place all three dialects around the Baltic Sea since the Early Bronze Age.
  • The easy replacement of a hypothetic Para-Balto-Finnic dialect by incoming Proto-Balto-Finnic-speaking peoples (say, with textile ceramics), without much linguistic impact.

In fact, the continuous contacts of the East Baltic with the Volga, and especially the close interaction with Akozino warrior-traders just before the Tarand-grave period, could be the actual origin of the recent (if any) Finnic-Mordvinic connections that need to be traced back to the LBA-IA (maybe here the number ‘ten’), since most of them can be related to a Pit-Comb Ware culture substrate and earlier contacts through the forest zone, which Samic (due to its early split and presence to the north of the Gulf of Finland during the BA) does not share. In fact, some of them can be traced back to Balto-Finnic first

These are the most often mentioned, in order of descending relevance for a shared ancient community:

  • Noun paradigms and the form and function of individual cases.
  • The geminate *mm (foreign to Proto-Uralic before the development of Fennic under Germanic influence) and other non-Uralic consonant clusters.
  • The change of numeral *luka ‘ten’ with (non-Uralic) *kümmen.
  • The presence of loanwords of non-Uralic origin, related to farming and trees, potentially Palaeo-European in nature.

It’s not only a question of quantity. Are these shared Mordvinic – Balto-Finnic traits really more relevant than, say, those between Italo-Celtic, which are supposed to have formed a community for a very short period at the end of the 3rd millennium around the Alps? Are these traits even sufficient to propose a common early Mordvinic-Finnic group within West Uralic, rather than loose Mordvinic – Balto-Finnic contacts, i.e. contacts between East Baltic (Textile Ceramics) and Volga-Kama (Netted Ware)?

Based on the alternative (Kallio’s) view of continued contacts between Textile Ceramics groups, even without knowing anything about linguistics, you can guess that Parpola is spinning very thin when assuming that these changes suggest that Balto-Finnic may have expanded with Akozino warrior-traders, separating thus ca. 800 BC from Mordvinic…

Genetic findings now clearly help dismiss any meaningful population impact in the LBA-IA transition, although any linguist can obviously argue for linguistic change in spite of major genetic continuity. But then we are stuck in the pre-ancient DNA era, so what’s ancient DNA for.

netted-ware-textile-ceramics
Middle Bronze Age cultures of Eastern Europe.

Genetic continuity = language continuity?

In the end, it’s very difficult to say how much language continuity there is around Estonia since the arrival of Corded Ware peoples. Looking at Modern Estonians, they have been clearly influenced by recent contacts with Baltic- and Germanic-speaking peoples clustering to the south-west in the PCA. They seem to have also received contacts from north(-east)ern peoples, likely from Finland, evidenced by their shifts toward the modern Estonian cluster during and after the Middle Ages, with a slight increase in Siberian ancestry and N1c subclades associated with Lovozero Ware. How much language change did these contacts bring? Maybe an expansion of Gulf of Finland Finnic (Northern Estonian) over Inland Finnic (Southern Estonian) and Gulf of Riga Finnic (Livonian)? Difficult to know, exactly, but, in the traditional view of Balto-Finnic dialectal distribution among Uralicists like Kallio, possibly no change at all.

So, if the obvious changes in the Estonia_MA cluster relative to Estonia_IA cluster and Estonia_Modern relative to Estonia_MA do not represent radical language change…Why would Estonia_IA represent a change relative to Estonia_BA, when it is statistically basically the same? Or Estonia_BA relative to CWC_Baltic? Because of the infiltration of haplogroup N1c around the whole Baltic? Because of the occasional 1% “Siberian” ancestry in some non-locals of varied haplogroups across the whole Baltic area?

In spite of all this, the amount of special pleading we are seeing among openly Nordicist amateurs when discussing the Uralic homeland relative to the Indo-European question in genetics has become a matter of plain willful ignorance. Like the living corpses of the Anatolian homeland, the Armenian homeland, the OIT proponents, or the nativist Basque R1b association, the personal involvement in the revival of “R1a=Indo-European” and “N=Uralic” trends is just painful to watch.

[Next post in this line, if I manage to make time for it: “Genetic (dis)continuity in Central Europe“. Let’s see if early Balts and early Slavs, as well as Germanic peoples, show a cluster closer to Danubian EBA (viz. Maros), Hungary-Balkans BA, and Urnfield-related samples than their predecessors in their areas, i.e. away from East Corded Ware groups… If you want, you can enjoy for the moment the new PCAs I could get done and the tentative map of languages in the Early Bronze Age, that will probably give you the right idea about early Indo-European and Uralic population movements]

bronze-age-early-indo-european
European Early Bronze Age: tentative language map based on linguistics, archaeology, and genetics. See full map.

Related

Złota a GAC-CWC transitional group…but not the origin of Corded Ware peoples

koszyce-gac-zlota-cwc

Open access Unraveling ancestry, kinship, and violence in a Late Neolithic mass grave, by Schroeder et al. PNAS (2019).

Interesting excerpts of the paper and supplementary materials, about the Złota group variant of Globular Amphora (emphasis mine):

A special case is the so-called Złota group, which emerged around 2,900 BCE in the northern part of the Małopolska Upland and existed until 2,600-2,500 BCE. Originally defined as a separate archaeological “culture” (15), this group is mainly defined by the rather local introduction of a distinct form of burial in the area mentioned. Distinct Złota settlements have not yet been identified. Nonetheless, because of the character of its burial practices and material culture, which both retain many elements of the GAC and yet point forward to the Corded Ware tradition, and because of its geographical location, the Złota group has attracted significant archaeological attention (15, 16).

The Złota group buried their dead in a new, distinct type of funerary structure; so-called niche graves (also called catacomb graves). These structures featured an entrance shaft or pit and, below that, a more or less extensive niche, sometimes connected to the entrance area by a narrow corridor. Local limestone was used to seal off the entrance shaft and to pave the floor of the niche, on which the dead were usually placed along with grave goods. This specific and relatively sophisticated form of burial probably reflects contacts between the northern Małopolska Upland and the steppe and forest-steppe communities further to the east, who also buried their dead in a form of catacomb graves. Individual cases of the use of ochre and of deformation of skulls in Złota burials provide further indications of such a connection (15). At the same time, the Złota niche grave practice also retains central elements of the GAC funerary tradition, such as the frequent practice of multiple burials in one grave, often entailing redeposition and violation of the anatomical order of corpses, and thus differs from the catacomb grave customs found on the steppes which are strongly dominated by single graves. Nonetheless, at Złota group cemeteries single burial graves appear, and even in multiple burial graves the identity of each individual is increasingly emphasized, e.g. by careful deposition of the body and through the personal nature of grave goods (16).

globular-amphorae-corded-ware-zlota-amphorae
Correspondence analysis of amphorae from the Złota-graveyards reveals that there is no typological break between Globular Amphorae and Corded Ware Amphorae, including ‘Strichbündelamphorae’ (after Furholt 2008)

Just like its burial practices, the material culture and grave goods of the Złota group combine elements of the GAC, such as amber ornaments and central parts of the ceramic inventory, with elements also found in the Corded Ware tradition, such as copper ornaments, stone shaft-hole axes, bone and shell ornaments, and other stylistic features of the ceramic inventory. In particular, Złota group ceramic styles have been seen as a clear transitional phenomenon between classical GAC styles and the subsequent Corded Ware ceramics, probably playing a key role in the development of the typical cord decoration patterns that came to define the latter (17).

As briefly summarized above, the Złota group displays a distinct funerary tradition and combination of material culture traits, which give the clear impression of a cultural “transitional situation”. While the group also appears to have had long-distance contacts directed elsewhere (e.g. to Baden communities to the south), it is the combination of Globular Amphora traits, on the one hand, and traits found among late Yamnaya or Catacomb Grave groups to the east as well as the closely related Corded Ware groups that emerged around 2,800 BCE, on the other hand, that is such a striking feature of the Złota group and which makes it interesting when attempting to understand cultural and demographic dynamics in Central and Eastern Europe during the early 3rd millennium BCE.

catacomb-grave-ksiaznice
Catacomb grave no. 2a/06 from Książnice, Złota culture (acc. to Wilk 2013). Image from Włodarczak (2017)

Książnice (site 2, grave 3ZC), Świętokrzyskie province. This burial, a so-called niche grave of the Złota type (with a vertical entrance shaft and perpendicularly situated niche), was excavated in 2006 and contained the remains of 8 individuals, osteologically identified as three adult females and five children, positioned on limestone pavement in the niche part of the grave. Radiocarbon dating of the human remains indicates that the grave dates to 2900-2630 BCE, 95.4% probability (Dataset S1). The grave had an oval entrance shaft with a diameter of 60 cm and depth of 130 cm; the depth of the niche reached to 170 cm (both measured from the modern surface), and it also contained a few animal bones, a few flint artefacts and four ceramic vessels typical of the Złota group. Książnice is located in the western part of the Małopolska Upland, which only has a few Złota group sites but a stronger presence of other, contemporary groups (including variants of the Baden culture).

Wilczyce (site 90, grave 10), Świętokrzyskie province. A rescue excavation in 2001 uncovered a niche grave of the Złota type, which had a round entrance shaft measuring 90 cm in diameter. The grave was some 60-65 cm deep below the modern surface and the bottom of the niche was paved with thin limestone plates, on which remains of three individuals had been placed; two adults, one female and one male, and one child. Four ceramic vessels of Złota group type were deposited in the niche along with the bodies. Wilczyce is located in the Sandomierz Upland, an area with substantial presence of both the Globular Amphora culture and Złota group, as well as the Corded Ware culture from 2800 BCE.

zlota-gac-cwc
Genetic affinities of the Koszyce individuals and other GAC groups (here including Złota) analyzed in this study. (A) Principal component analysis of previously published and newly sequenced ancient individuals. Ancient genomes were projected onto modern reference populations, shown in gray. (B) Ancestry proportions based on supervised ADMIXTURE analysis (K = 3), specifying Western hunter-gatherers, Anatolian Neolithic farmers, and early Bronze Age steppe populations as ancestral source populations. LP, Late Paleolithic; M, Mesolithic; EN, Early Neolithic; MN, Middle Neolithic; LN, Late Neolithic; EBA, Early Bronze Age; PWC, Pitted Ware culture; TRB, Trichterbecherkultur/Funnelbeaker culture; LBK, Linearbandkeramik/Linear Pottery culture; GAC, Globular Amphora culture; Złota, Złota culture. Image modified to outline in red GAC and Złota groups.

To further investigate the ancestry of the Globular Amphora individuals, we performed a supervised ADMIXTURE (6) analysis, specifying typical western European hunter-gatherers (Loschbour), early Neolithic Anatolian farmers (Barcın), and early Bronze Age steppe populations (Yamnaya) as ancestral source populations (Fig. 2B). The results indicate that the Globular Amphora/Złota group individuals harbor ca. 30% western hunter-gatherer and 70% Neolithic farmer ancestry, but lack steppe ancestry. To formally test different admixture models and estimate mixture proportions, we then used qpAdm (7) and find that the Polish Globular Amphora/Złota group individuals can be modeled as a mix of western European hunter-gatherer (17%) and Anatolian Neolithic farmer (83%) ancestry (SI Appendix, Table S2), mirroring the results of previous studies.

zlota-steppe-ancestry-cwc
Table S2. qpADM results. The ancestry of most Globular Amphora/Złota group individuals
can be modelled as a two-way mixture of Mesolithic western hunter-gatherers (WHG), and early Anatolian Neolithic farmers (Barcın). The five individuals from Książnice (Złota group) show evidence for additional gene flow, most likely from an eastern source.

The lack of a direct genetic connection of Corded Ware peoples with the Złota group despite their common “steppe-like traits” – shared with Yamna – reveals, once more, how the few “Yamna-like” traits of Corded Ware do not support a direct connection with Indo-Europeans, and are the result of the expansion of the so-called steppe package all over Europe, and particularly among cultures closely related to the Khvalynsk expansion, and later under the influence of expanding Yamna peoples.

The results from Książnice may support that early Corded Ware peoples were in close contact with GAC peoples in Lesser Poland during the complex period of GAC-Trypillia-CWC interactions, and especially close to the Złota group at the beginning of the 3rd millennium BC. Nevertheless, patrilineal clans of Złota apparently correspond to Globular Amphorae populations, with the only male sample available yet being within haplogroup I2a-L801, prevalent in GAC.

NOTE. The ADMIXTURE of Złota samples in common with GAC samples (and in contrast with the shared Sredni Stog – Corded Ware “steppe ancestry”) makes the possibility of R1a-M417 popping up in the Złota group from now on highly unlikely. If it happened, that would complicate further the available picture of unusually diverse patrilineal clans found among Uralic speakers expanding with early Corded Ware groups, in contrast with the strict patrilineal and patrilocal culture of Indo-Europeans as found in Repin, Yamna and Bell Beakers.

Once again the traditional links between groups hypothesized by archaeologists – like Gimbutas and Kristiansen in this case – are wrong, as is the still fashionable trend in descriptive archaeology, of supporting 1) wide cultural relationships in spite of clear-cut inter-cultural differences (and intra-cultural uniformity kept over long distances by genetically-related groups), 2) peaceful interactions among groups based on few common traits, and 3) regional population continuities despite cultural change. These generalized ideas made some propose a steppe language shared between Pontic-Caspian groups, most of which have been proven to be radically different in culture and genetics.

gimbutas-kurgan-indo-european
The background shading indicates the tree migratory waves proposed by Marija Gimbutas, and personally checked by her in 1995. Image from Tassi et al. (2017).

Furthermore, paternal lines show once again marked bottlenecks in expanding Neolithic cultures, supporting their relevance to follow the ethnolinguistic identity of different cultural groups. The steppe- or EHG-related ancestry (if it is in fact from early Corded Ware peoples) in Książnice was thus probably, as in the case of Trypillia, in the form of exogamy with females of neighbouring groups:

The presence of unrelated females and related males in the grave is interesting because it suggests that the community at Koszyce was organized along patrilineal lines of descent, adding to the mounting evidence that this was the dominant form of social organization among Late Neolithic communities in Central Europe. Usually, patrilineal forms of social organization go hand in hand with female exogamy (i.e., the practice of women marrying outside their social group). Indeed, several studies (11, 12) have shown that patrilocal residence patterns and female exogamy prevailed in several parts of Central Europe during the Late Neolithic. (…) the high diversity of mtDNA lineages, combined with the presence of only a single Y chromosome lineage, is certainly consistent with a patrilocal residence system.

funnelbeaker-trypillia-corded-ware
Map of territorial ranges of Funnel Beaker Culture (and its settlement concentrations in Lesser Poland), local Tripolyan groups and Corded Ware Culture settlements (■) at the turn of the 4th/3rd millennia BC.

Since ancient and modern Uralians show predominantly Corded Ware ancestry, and Proto-Uralic must have been in close contact with Proto-Indo-European for a very long time – given the different layers of influence that can be distinguished between them -, it follows as logical consequence that the North Pontic forest-steppes (immediately to the west of the PIE homeland in the Don-Volga-Ural steppes) is the most likely candidate for the expansion of Proto-Uralic, accompanying the spread of Sredni Stog ancestry and a bottleneck under R1a-M417 lineages.

The early TMRCAs in the 4th millennium BC for R1a-M417 and R1a-Z645 support this interpretation, like the R1a-M417 sample found in Sredni Stog. On the other hand, the resurgence of typical GAC-like ancestry in late Corded Ware groups, with GAC lineages showing late TMRCAs in the 3rd millennium BC, proves the disintegration of Corded Ware all over Europe (except in Textile Ceramics- and Abashevo-related groups) as the culture lost its cohesion and different local patrilineal clans used the opportunity to seize power – similar to how eventually I2a-L621 infiltrated eastern (Finno-Ugrian) groups.

Related

Uralic speakers formed clines of Corded Ware ancestry with WHG:ANE populations

steppe-forest-tundra-biomes-uralic

The preprint by Jeong et al. (2018) has been published: The genetic history of admixture across inner Eurasia Nature Ecol. Evol. (2019).

Interesting excerpts, referring mainly to Uralic peoples (emphasis mine):

A model-based clustering analysis using ADMIXTURE shows a similar pattern (Fig. 2b and Supplementary Fig. 3). Overall, the proportions of ancestry components associated with Eastern or Western Eurasians are well correlated with longitude in inner Eurasians (Fig. 3). Notable outliers include known historical migrants such as Kalmyks, Nogais and Dungans. The Uralic- and Yeniseian-speaking populations, as well as Russians from multiple locations, derive most of their Eastern Eurasian ancestry from a component most enriched in Nganasans, while Turkic/Mongolic speakers have this component together with another component most enriched in populations from the Russian Far East, such as Ulchi and Nivkh (Supplementary Fig. 3). Turkic/Mongolic speakers comprising the bottom-most cline have a distinct Western Eurasian ancestry profile: they have a high proportion of a component most enriched in Mesolithic Caucasus hunter-gatherers and Neolithic Iranians and frequently harbour another component enriched in present-day South Asians (Supplementary Fig. 4). Based on the PCA and ADMIXTURE results, we heuristically assigned inner Eurasians to three clines: the ‘forest-tundra’ cline includes Russians and all Uralic and Yeniseian speakers; the ‘steppe-forest’ cline includes Turkic- and Mongolic-speaking populations from the Volga and Altai–Sayan regions and Southern Siberia; and the ‘southern steppe’ cline includes the rest of the populations.

eurasian-clines-uralic-altaic
The first two PCs summarizing the genetic structure within 2,077 Eurasian individuals. The two PCs generally mirror geography. PC1 separates western and eastern Eurasian populations, with many inner Eurasians in the middle. PC2 separates eastern Eurasians along the northsouth cline and also separates Europeans from West Asians. Ancient individuals (color-filled shapes), including two Botai individuals, are projected onto PCs calculated from present-day individuals.

For the forest-tundra populations, the Nganasan + Srubnaya model is adequate only for the two Volga region populations, Udmurts and Besermyans (Fig. 5 and Supplementary Table 8).

For the other populations west of the Urals, six from the northeastern corner of Europe are modelled with additional Mesolithic Western European hunter-gatherer (WHG) contribution (8.2–11.4%; Supplementary Table 8), while the rest need both WHG and early Neolithic European farmers (LBK_EN; Supplementary Table 2). Nganasan-related ancestry substantially contributes to their gene pools and cannot be removed from the model without a significant decrease in the model fit (4.1–29.0% contribution; χ2 P ≤ 1.68 × 10−5; Supplementary Table 8).

west-urals-finno-ugrians-qpadm
Supplementary Table 8. QpAdm-based admixture modeling of the forest-tundra cline populations. For the 13 populations west of the Urals, we present a four-way admixture model, Nganasan+Srubnaya+WHG+LBK_EN, or its minimal adequate subset. Modified from the article, to include colors for cultures, and underlined best models for Corded Ware ancestry among Uralians.

NOTE. It doesn’t seem like Hungarians can be easily modelled with Nganasan ancestry, though…

For the 4 populations east of the Urals (Enets, Selkups, Kets and Mansi), for which the above models are not adequate, Nganasan + Srubnaya + AG3 provides a good fit (χ2 P ≥ 0.018; Fig. 5 and Supplementary Table 8). Using early Bronze Age populations from the Baikal Lake region (‘Baikal_EBA’; Supplementary Table 2) as a reference instead of Nganasan, the two-way model of Baikal_EBA + Srubnaya provides a reasonable fit (χ2 P ≥ 0.016; Supplementary Table 8) and the three-way model of Baikal_EBA + Srubnaya + AG3 is adequate but with negative AG3 contribution for Enets and Mansi (χ2 P ≥ 0.460; Supplementary Table 8).

east-urals-ugric-samoyedic-qpadm
Supplementary Table 8. QpAdm-based admixture modeling of the forest-tundra cline populations. For the four populations east of the Urals, we present three admixture models: Baikal_EBA+Srubnaya, Baikal_EBA+Srubnaya+AG3 and Nganasan+Srubnaya+AG3. For each model, we present qpAdm p-value, admixture coefficient estimates and associated 5 cM jackknife standard errors (estimate ± SE). Modified from the article, to include colors for cultures, and underlined best models for Corded Ware ancestry among Uralians.

Bronze/Iron Age populations from Southern Siberia also show a similar ancestry composition with high ANE affinity (Supplementary Table 9). The additional ANE contribution beyond the Nganasan + Srubnaya model suggests a legacy from ANE-ancestry-rich clines before the Late Bronze Age.

bronze-age-iron-age-karasuk-mezhovska-tagar-qpadm
Supplementary Table 9. QpAdm-based admixture modeling of Bronze and Iron Age populations of southern Siberia. For ancieint individuals associated with Karasuk and Tagar cultures, Nganasan+Srubnaya model is insufficient. For all five groups, adding AG3 as the third ancestry or substituting Nganasan with Baikal_EBA with higher ANE affinity provides an adequate model. For each model, we present qpAdm p-value, admixture coefficient estimates and associated 5 cM jackknife standard errors (estimate ± SE). Models with p-value ≥ 0.05 are highlighted in bold face. Modified from the article, to include colors for cultures, and underlined best models for Corded Ware ancestry among Uralians.

Lara M. Cassidy comments the results of the study in A steppe in the right direction (you can read it here):

Even among the earliest available inner Eurasian genomes, east–west connectivity is evident. These, too, form a longitudinal cline, characterized by the easterly increase of a distinct ancestry, labelled Ancient North Eurasian (ANE), lowest in western European hunter-gatherers (WHG) and highest in Palaeolithic Siberians from the Baikal region. Flow-through from this ANE cline is seen in steppe populations until at least the Bronze Age, including the world’s earliest known horse herders — the Botai. However, this is eroded over time by migration from west and east, following agricultural adoption on the continental peripheries (Fig. 1b,c).

Strikingly, Jeong et al. model the modern upper steppe cline as a simple two-way mixture between western Late Bronze Age herders and Northeast Asians (Fig. 1c), with no detectable residue from the older ANE cline. They propose modern steppe peoples were established mainly through migrations post-dating the Bronze Age, a sequence for which has been recently outlined using ancient genomes. In contrast, they confirm a substantial ANE legacy in modern Siberians of the northernmost cline, a pattern mirrored in excesses of WHG ancestry west of the Urals (Fig. 1b). This marks the inhospitable biome as a reservoir for older lineages, an indication that longstanding barriers to latitudinal movement may indeed be at work, reducing the penetrance of gene flows further south along the steppe.

eurasian-clines-uralic-turkic-mongol-altaic
The genomic formation of inner Eurasians. b–d, Depiction of the three main clines of ancestry identified among Inner Eurasians. Sources of admixture for each cline are represented using proxy ancient populations, both sampled and hypothesised, based on the study’s modelling results. The major eastern and western ancestries used to model each cline are shown in bold; the peripheral admixtures that gave rise to these are also shown. Additional contributions to subsections of each cline are marked with dashed lines. b, The northernmost cline, illustrating the legacy of WHG and ANE-related populations. c,d, The upper (c) and lower (d) steppe clines are shown, both of which have substantial eastern contributions related to modern Tungusic speakers. The authors propose these populations are themselves the result of an admixture between groups related to the Nganasan, whose ancestors potentially occupied a wider range, and hunter-gatherers (HGs) from the Amur River Basin. While the upper steppe cline in c can be described as a mixture between this eastern ancestry and western steppe herders, the current model for the southern steppe cline as shown in d is not adequate and is likely confounded by interactions with diverse bordering ancestries. Credit: Ecoregions 2017, Resolve https://ecoregions2017.appspot.com/

Given the findings as reported in the paper, I think it should be much easier to describe different subclines in the “northernmost cline” than in the much more recent “Turkic/Mongolic cline”, which is nevertheless subdivided in this paper in two clines. As an example, there are at least two obvious clines with “Nganasan-related meta-populations” among Uralians, which converge in a common Steppe MLBA (i.e. Corded Ware) ancestry – one with Palaeo-Laplandic peoples, and another one with different Palaeo-Siberian populations:

siberian-clines-uralic-altaic
PCA of ancient and modern Eurasian samples. Ancient Palaeo-Laplandic, Palaeosiberian, and Altai clines drawn, with modern populations labelled. See a version with higher resolution.

The inclusion of certain Eurasian groups (or lack thereof) in the PCA doesn’t help to distinguish these subclines visually, and I guess the tiny “Naganasan-related” ancestral components found in some western populations (e.g. the famous ~5% among Estonians) probably don’t lend themselves easily to further subdivisions. Notice, nevertheless, the different components of the Eastern Eurasian source populations among Finno-Ugrians:

uralic-admixture-qpadm
Characterization of the Western and Eastern Eurasian source ancestries in inner Eurasian populations. [Modified from the paper, includes only Uralic populations]. a, Admixture f3 values are compared for different Eastern Eurasian (Mixe, Nganasan and Ulchi; green) and Western Eurasian references (Srubnaya and Chalcolithic Iranians (Iran_ChL); red). For each target group, darker shades mark more negative f3 values. b, Weights of donor populations in two sources characterizing the main admixture signal (date 1 and PC1) in the GLOBETROTTER analysis. We merged 167 donor populations into 12 groups (top right). Target populations were split into five groups (from top to bottom): Aleuts; the forest-tundra cline populations; the steppe-forest cline populations; the southern steppe cline populations; and ‘others’.

Also remarkable is the lack of comparison of Uralic populations with other neighbouring ones, since the described Uralic-like ancestry of Russians was already known, and is most likely due to the recent acculturation of Uralic-speaking peoples in the cradle of Russians, right before their eastward expansions.

west-eurasian-east-eurasian-ancestry
Supplementary Fig. 4. ADMIXTURE results qualitatively support PCA-based grouping of inner Eurasians into three clines. (A) Most southern steppe cline populations derive a higher proportion of their total Western Eurasian ancestry from a source related to Caucasus, Iran and South Asian populations. (B) Turkic- and Mongolic-speaking populations tend to derive their Eastern Eurasian ancestry more from the Devil’s Gate related one than from Nganasan-related one, while the opposite is true for Uralic- and Yeiseian-speakers. To estimate overall western Eurasian ancestry proportion, we sum up four components in our ADMIXTURE results (K=14), which are the dominant components in Neolithic Anatolians (“Anatolia_N”), Mesolithic western European hunter-gatherers (“WHG”), early Holocene Caucasus hunter-gatherers (“CHG”) and Mala from southern India, respectively. The “West / South Asian ancestry” is a fraction of it, calculated by summing up the last two components. To estimate overall Eastern Eurasian ancestry proportion, we sum up six components, most prevalent in Surui, Chipewyan, Itelmen, Nganasan, Atayal and early Neolithic Russian Far East individuals (“Devil’s Gate”). Eurasians into three clines. (A) Most southern steppe cline populations derive a higher proportion of their total Western Eurasian ancestry from a source related to Caucasus, Iran and South Asian populations. (B) Turkic- and Mongolic-speaking populations tend to derive their Eastern Eurasian ancestry more from the Devil’s Gate related one than from Nganasan-related one, while the opposite is true for Uralic- and Yeiseian-speakers. To estimate overall western Eurasian ancestry proportion, we sum up four components in our ADMIXTURE results (K=14), which are the dominant components in Neolithic Anatolians (“Anatolia_N”), Mesolithic western European hunter-gatherers (“WHG”), early Holocene Caucasus hunter-gatherers (“CHG”) and Mala from southern India, respectively. The “West / South Asian ancestry” is a fraction of it, calculated by summing up the last two components. To estimate overall Eastern Eurasian ancestry proportion, we sum up six components, most prevalent in Surui, Chipewyan, Itelmen, Nganasan, Atayal and early Neolithic Russian Far East individuals (“Devil’s Gate”).

A comparison of Estonians and Finns with Balts, Scandinavians, and Eastern Europeans would have been more informative for the division of the different so-called “Nganasan-like meta-populations”, and to ascertain which one of these ancestral peoples along the ancient WHG:ANE cline could actually be connected (if at all) to the Cis-Urals.

Because, after all, based on linguistics and archaeology, geneticists are not supposed to be looking for populations from the North Asian Arctic region, for “Siberian ancestry”, or for haplogroup N1c – despite previous works by their peers – , but for the Bronze Age Volga-Kama region…

Related