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

European hydrotoponymy (V): Etruscans and Rhaetians after Italic peoples

italy-mediterranean-bronze-age

There is overwhelming evidence that the oldest hydrotoponymic layer in Italy (and especially Etruria) is of Old European nature, which means that non-Indo-European-speaking (or, at least, non-Old-European-speaking) Etruscans came later to the Apennine Peninsula.

Furthermore, there is direct and indirect linguistic, archaeological, and palaeogenomic data supporting that the intrusive Tursānoi came from the Aegean during the Late Bronze Age, possibly through the Adriatic, and that their languages spread to Etruria and probably also to the eastern Alps.

Hydrotoponymic layer

The following are translated excerpts (emphasis mine) from Lenguas, genes y culturas en la Prehistoria de Europa y Asia suroccidental, by Villar et al. Universidad de Salamanca (2007):

villar-vascos
Lenguas, genes y culturas en la Prehistoria de Europa y Asia suroccidental (2007). Buy the ebook online (or the printed version, if available).

‘(Indo-)Mediterranean’ substrate?

The name Indo-Mediterranean substrate was spread in Italy by the work of V. Pisani. Other Italian scholars continued this idea, such as W. Belardi, L. Heilmann, D. Silvestri, etc. In their hands, the nuclear area of ​​the Indo-Mediterranean substratum was established as follows: “il mondo culturale indomediterráneo trova i suoi più importanti centri di gravitazione (e, soltanto secondariamente, di espansione) nel Mediterràneo Orientale (Creta, Cipro, Asia Minore), nella ‘regione dei due fiumi’ (area di espansione subarea) e nella valle dell’Indo (civiltà de Harappa e Mohenjo Daro)”. From there they could have spread to other areas, such as the western Mediterranean. Even at one point there was talk of “a Mediterranean oasis in the Baltic”, whose main basis was the existence of numerous lexical elements, real or supposedly pre-Indo-European in the Baltic languages.

One of the paradoxes of the theory of the Mediterranean substrate is that the lexical or toponymic components that are attributed to it can rarely be explained etymologically from the surviving languages ​​of said supposed substrate; sometimes they are not even very compatible with what we know of the non-Indo-European languages ​​of the corresponding area. For example, neither Basque nor Iberian have an ancestral and autochthonous phoneme /p/, while that phoneme is frequent in substrate words (cf. among the few mentioned above *pal- and *lap-). In fact, for these three languages ​​other alternative origins have been imagined, so that they would not be representatives of the local substrate: Basque (North Africa, the Caucasus), Iberian (North Africa), Etruscan (Asia Minor). Thus, under such hypotheses the non-Indo-European languages ​​attested in Italy and the Iberian Peninsula would not be autochthonous, but as immigrant as the Indo-European languages.

akwa-hydronyms
Akʷa hydronyms. The majority of old serial elements are found in Italy, with 9, where they don’t appear as second element. Different to the southern areas, they are found in especially frequent compounds in the acha-Namen in Germany, and hyper-represented (as usual) in Lithuania, which shows strictly 8 ancient names.

Italy and Iberia

Let’s review data on Italy:

I. Serial tponyms and hydronyms of Italy:

  1. ub-: Caecubus, Egubium, Litubium, Marrubium, Olobia, Rutuba, Tardoba, Tardubius, Verubius, etc.
  2. uc-: Aluca, Arucia, Arugus, Ausucum, Ausugum, Motuca, Uccia.
  3. ur-: Orinos, Stura, Stura, Astura, Tibur, Caburrum, Calorem.
  4. urc-: Coturga, Orgus, Urcia, Urcinia, Urgo.
  5. bai-: Baebiani.
  6. tuc-: Tucianus (pagus).
  7. murc-: Murcia, Murgantia, Murgantia.
  8. *war: Varduli, Barduli.

ub-hydronyms

II. Non-serial toponyms and hydronyms of Italy: Aesis, Aisis, Ana, Ania, Anios, Arsia, Astura, Ausa, Ausonia, Ausculum, Bardinisca vallis, Barduli, Basentius, Basta, Boron, Cabienses (Cabia), Caburrum, Cales, Cales, Casta Ballenis, Ceresium, Cerili, Corsica, Cortona, Curicum, Ispelum, Ispila, Isporos, Istonium, Istria, lacus, Latis, Latium, Laurentum, Laurentes, Luca, Lucania, Lucera, Maleventum, mare, Marrucini, Minio, Minius, Oscela, Osci, Ossa, Ostia, Paestum, Pisaurum, Pisaurus, Sabini, Sagis, Savo, Sila, Silarus, Silis, Soletum, etc.

italy-iberia-hydronymy-toponymy

Not few of the coincident place names between the southern Iberian and Italic material are rigorous cognates. We understand by such the names that not only coincide in the root or in the serial element, but in the whole root set plus suffixes, or – if it is a compound – in the two sets of roots plus suffixes. In addition to the ones that we are going to present below, there are others that we did not mention because the Iberian correlate was not found within the southern group, but in other geographical areas, as is the case, for example, with the Italian Mantua and the Spanish Mantua (Carpetania).

As can be seen, the parallels between the southern Iberian toponymic area and the Italic one are so wide and strict that the mere calculation of probabilities makes any attempt to attribute them to the mere chance of random homophony irrational. And the improbability of chance increases as coincidences are added in new places in Europe. What will not prevent, for sure, that some would resort to it as an explanation, in particular those who are reluctant to abandon the conception of the prehistory of the European continent that underlies their usual approaches, which suffer an irreparable strike when they are confronted with these data.

The second aspect, the compatibility of this material with Indo-European etymology, offers another significant correlation: the “southern” series that are also found in the Ibero-Pyrenean region and in Italy (and the rest of western Europe) are compatible with Indo-European etymologies; (…)

I will spare the reader of all proposed Indo-European etymologies, most of which are fairly evident. Those interested should buy one of the books, or both.

or-hydronyms

Etruria

(…) in the whole of Italy there is a considerable collection of toponyms and hydronyms of “Southern Iberian” type, whose joint inventory we have contributed to above. From them we find in Etruria Ause, Veturris / Bituriza, Castola, Hasta, Cortona, Luca, Minio, Osa / Ossa, Pissai, Pistoria. The Hispanic and Italian correlates of those names are:

iberian-etruscan-indo-european

However, the inventory of ancient names and hydronyms of Etruria compatible without discussion with well-known Indo-European etymologies is much wider: Albina, Alma, Alsium, Arnine, Arnos, Arnus, Aventia, Marta, Pallia, Umbro, Vetulonium, Volsinii. Furthermore, the majority of Etrurian hydronyms have non-Latin Indo-European etymology: Albina, Alma, Arnine, Arnos, Arnus, Auser, Aventia, Marta, Minio, Osa, Ossa, Pallia, Umbro. And very few of the others (Clusinus, Cremera, Lingeus, Trasumenus, Vesidia) could claim an Etruscan etymology, if only one could do so.

In summary, the territory occupied by Etruscans presents a hydro-toponymic situation very similar to that of the rest of Italy and Western Europe: it exhibits a very deep toponymic stratum of Indo-European character to which most hydronyms attested in antiquity belong. As we know the history of Etruria from the end of the 1st millennium BC, and we know that no other Indo-European peoples mediated between the Etruscans and the Romanization of the territory, we must conclude that this ancient toponymy was there before the Etruscans arrived or emerged in that place. And, when the Etruscans settled there, they did not have the opportunity to put names of their language to the rivers in general, because they had already received them from a previous people and the Etruscans limited themselves to learning them, adapting them to their language, and transmitting them in turn to the Romans. When the latter Romanized Etruria, they limited themselves to incorporating those names and adapting them to Latin.

maro-maranto

Etruscans

The ‘foreign’ Tyrsenians

Here is a recapitulation of the main reasons why Etruscans were recently intrusive to Italy, as they appeared in The Origin of the Etruscans, by Beekes (2003):

NOTE. You can read another version of the text in PDF, as the main paper from Biblioteca Orientalis LIX(3-4) 2002.

  1. The tradition as given by Herodotus and Dionysius of Halikarnassos.
  2. The story that the Etruscans were Pelasgians.
  3. The use of the term ‘Tyrsēnoi’ for both Etruscans and a people in north-western Asia Minor. Above we argued that the eastern Tyrsēnoi are the remnant of a population. This means that the Tyrsēnoi/Etruscans came from this area.
  4. The Lemnos inscription.
  5. To the testimony of Lemnos must now be added that Herodotus says that the people of Plakiê and Skylakê spoke the same language as the Etruscans.
  6. etruscan-homeland

  7. The kumdanlı inscription. (…) lake Egridir (of which the old name is unknown, unless it was just Limnai). This is just over the border of classical Lydia. The inscription dates from the second century ad and is given by Ramsay (i883); the same inscription is cited by Sundwall (i9i3, 22i). It mentions three people as Tyrsēnoi(67, 68, i02). Though very late, the inscription is of great interest, as it is the only time that we have inscriptional evidence for Tyrsēnoi in Asia Minor. (And nobody will argue that these were Etruscans from Italy.) (…)
  8. The suffix -ānos. The suffix -ānos in the name Tyrsēnoi (with ē from ā) points to the north-west of Asia Minor. It has long since been recognized that this suffix for ethnic names is at home in north-west Asia Minor; some think that it is of non-Greek origin; cf. Αβυδηνός , Ολυμπιηνός, Περγαμηνός, Σαρδηνός; (see Chantraine i933, 206; Schwyzer 490 (6); De Simone i993, 88ff.). This proves that the name Tyrsēnoi originated in the north-west of Asia Minor. (…)
  9. Loanwords. As to the language, Steinbauer (i999, 367) observes that Etruscan shows most connections (loanwords) with Lydian (…)
  10. Tarchon. The definite proof of the oriental origin of the Etruscans is that a ‘hero’ of great significance is Tarchon (Briquel i99i). He is clearly the Stormgod Tarhun(t)-, the highest god of the Luwians and Hittites.
  11. Nanas. This identification is strongly confirmed by the story that the Etruscans were Pelasgians who came from Greece under Nanas (Nanos), mentioned by Hellanikos. This name was long ago recognized as an Anatolian ‘Lallname’.
  12. The triumphus complex. In his study of the Roman triumphus Versnel has shown that (i970, 293): ‘the Etruscans brought the New Year festival with them from Asia Minor, together with the god who formed the centre of it, a god whom the Greeks called Dionysos, the Etruscans Tinia (or by an Italic name Voltumna), a figure of the ‘dying and rising’ type, who was invoked by the cry *thriambe and who on New Year’s Day was represented by the king.’ And on p. 300: ‘The Etruscans brought the New Year festival with them from Asia Minor and gave Rome two ceremonies: the ludi Romani as the festival of the New Year, the triumph as the festival of the victory. … Only along this way is it possible to explain the data: i. the Dionysiac call to epiphany triumpe, introduced via Etruria; 2. the identification of the Roman victorious general and of the magistrate leading the games with the god Iuppiter; 3. the typological and historic relation between the ludi Romani and the triumph.’
  13. The double axe. On a smaller issue Versnel concludes (p. 299): ‘When this bipennis [‘double axe’], property of ‘Zeus Bakchos’, carried as symbol of sacred power by Lydian kings, is encountered again as the symbol of the royal authority of the Etruscan kings, particularly of the supreme king of the federation of cities, this may be considered an important indication of the Asia Minor origin of the entire underlying ideology, and of the ceremony of investiture in which the bipennis played a part.’ These conclusions are of primary importance, as they concern a deeprooted complex of religious views that cannot have been taken over from elsewhere.
  14. The Kabeiroi. One might also recall the Latin word camillus, which means a young boy of noble birth who assists with ritual actions. (…) Probably more evidence can be found in the field of religion, such as the much discussed hepatoscopy. It seems quite probable to me that the lituus, the crosier used by the Roman priests, is Anatolian (see e.g. Wainwright i959, 2i0; cf. Haas i99i, Abb. 75, the Stormgod standing on an animal with his lituus over his shoulder).
  15. The Etruscan way of life. There was in antiquity much criticism on Etruscan customs, concerning cruelty, sexual behaviour, and the behaviour of women. (…) Dionysius concluded from the fact that they were so strange that they had always lived in Italy, whereas it is of course much more natural to explain it by assuming that they were strangers.
  16. No withdrawal area. We have seen above that Tuscany is not a ‘withdrawal area’, where an ancient people may hold out when the country is invaded. On the contrary, it is a desirable area which the Indo-European peoples, had they come later, would certainly have occupied. (But it went the other way: the Etruscans came long after the Indo-Europeans and settled there/conquered the country.)
  17. sea-peoples-expansion-tyrsenians
    The Sea Peoples in the Eastern Mediterranean c. 1200 BC. Map by Ian Mladjov.
  18. Archaeology. Many scholars would like to see archaeological evidence, but I think that it is quite possible that we shall never find any.
  19. The 1200 crisis. In 1200 the whole Mediterranean was in commotion; the Mycenaean and Hittite worlds, between which the TyrseOEnoi lived, disappeared. So the movement of the Etruscans fits very well in the general picture. That this was the setting of the migration of the Etruscans has been assumed by many earlier scholars.
  20. The ten saecula. As to the time, it has been argued that the Etruscans thought that their world would last ten saecula (Briquel i999, 58; Pfiffig i975, i59ff.). The way of counting provides several problems, however (…) If we accept it, we arrive at 968 bc. Now we do not know from when one started counting. This might have been a decisive victory over the Umbrians, or a kind of unification of the Etruscans, or the founding of an important city. It could well be that this was some 200 years after the arrival of the Etruscans, which would take us to 1168 bc. (…)
  21. The famine. Herodotus states that the reason for the departure of the Tyrsēnoi was a long famine. This has been identified as the famine about i200. (…)
  22. The sea-peoples. (…) The phenomenon as a whole stands, it seems; the problem is the details: which peoples took part in which movements? In our case, as the Lukka are mentioned (which were very probably the Lycians), the Tyrsēnoi may have been involved as well. So the question is whether the T(w)r(w)š, mentioned by Merneptah, were the Tyrsēnoi. We have no confirmation, but it seems quite possible.
  23. The journey. We know from the abundant finds of ceramics in the i3th century that the Mycenaeans knew the sea-route to Italy. (…)
  24. The Umbrians. Pliny (3, ii2) states that the Etruscans conquered 300 cities from the Umbrians (Trecenta eorum oppida Tusci debellasse reperiuntur.). This clearly refers to the ‘Landnahme’. This statement is confirmed by the river Umbro (mod. Ombrone), which flows in its full length in Etruscan territory. The river will have given its name to the people, or vice versa. Anyhow, the river will have flowed in Umbrian territory; so the Etruscans must have pushed the Umbrians out.
  25. The name Sergestus, of a prominent friend of Aeneas, seems identical with Lydian Srkastu- and Phrygian Surkastos (…) it is excluded that (Virgil) got it from Lydia or Phrygia, or Asa Minor in general. So he must have got it at home, from a source that was acqainted with Etruscan traditions. This means that the name was known to the Etruscans (or those who studied their traditions). Above I proposed that it lives on in Etr. Sekst-alu-.

You can read the full text (and its appendices) for further evidences adduced by Beekes, who considers the matter mostly settled.

Local Italic peoples

Another main reason for the intrusion of Tyrsenians among local groups is the ancient connection between Italic languages, which most likely formed an ancient Apennine dialect continuum:

  • the core Italic group with Latino-Faliscan and Palaeo-Sabellic – probably also including an Ausonian-Siculian branch – separated ca. 1500-1000 BC;
  • NOTE. Sicel is believed to have arrived in Sicily with Ausonian-Siculian speakers either around the 13th c. or in the middle of the 11th c. BC (or in both waves), from their ancient settlements in the mainland, driving prior inhabitants (Elymians) to the east of the island, which sets another clear terminus ante quem for the expansion of Italic languages in southern Italy.

  • and the possibly more distantly related North Picene and Venetic, connecting all roughly to an early to mid-2nd millennium BC language.

This continuum was probably broken (with language replacement and displacement events) with the 12th c. BC turmoil and the emergence of new social hierarchies. The adoption of older place and river names, as well as the lack of long-lasting influence on neighbouring languages, suggests that the predominance of the Etruscan language in its proto-historic territory was probably gradual and quite recent.

NOTE. For more on guesstimates, relative chronological expansions and potential archaeological identifications, see e.g. “Ausgliederung und Aufgliederung der italischen Sprachen”, by Helmut Rix In: Languages in Prehistoric Europe (2003). Or, basically, any recent (linguistic) text on the distribution and attribution of ancient Apennine languages to the Ital(o-Venet)ic group.

Italic-venetic-etruscan-languages-map
Languages of pre-Roman Italy and nearby islands. Italo-Venetic languages surrounded with shadowed red border. I1, South Picene; I2, Umbrian; I3, Sabine; I4, Faliscan; I5, Latin; I6, Volscian and Hernican; I7, Central Italic (Marsian, Aequian, Paeligni, Marrucinian, Vestinian); I8, Oscan, Sidicini, Pre-Samnite; I9, Sicel; IE1, Venetic; IE2, North Picene; IE3, Ligurian; IE4, Elymian; IE5, Messapian; C1, Lepontic; C2, Gaulish; G1-G2-G3, Greek dialects (G1: Ionic, G2: Aeolic, G3: Doric); P1, Punic; N1, Rhaetian; N2, Etruscan; N3, Nuragic. Image modified from Davius Sanctex.

Archaeology

The main criticism against this ethnolinguistic model of foreign Tyrsenians comes, surprisingly, from the lack of archaeological data to support this arrival. Or, rather, fitting anthropological interpretations of a culture of Asia Minor with similar hierarchical societies (?). From Review of R. S. P. Beekes, The Origin of the Etruscans, by Mahoney, Etruscan Studies (2008):

A crucial part of Beekes’ argument, however, is that there is a significant cultural break in Etruria around 1200, at the beginning of the Early Bronze Age or Proto-Villanovan period (p. 34, citing Briquel and Torelli). The introduction of cremation can be dated to around this period, and there is also evidence for a new hierarchical social organization (convenient summary in Barker and Rasmussen, p. 53-60). Beekes simply says that there is a change, and changes of this sort can come about when new people move in to an existing society, so therefore this change is consistent with his theory. That is correct as far as it goes, but what is missing is any consideration of how and why people coming in from Asia Minor would cause the particular changes that take place in Etruria. Can we argue that the society of the pre-migration Tyrsenians was hierarchical in the same way as those of the various Indo-European-speaking peoples in the region? Beekes simply says “what we still would like to have is material objects, or art traditions etc., from Etruria agreeing with their homeland” (p. 34). What we would really like to have is evidence for the organization of society in this alleged homeland.

Weird as this criticism is, here it is yet another example of the social change brought about under Eastern Mediterranean influences during the Final Bronze Age, from a recent paper (behind paywall) Mobile elites at Frattesina: flows of people in a Late Bronze Age ‘port of trade’ in northern Italy, by Cavazzuti et al. Antiquity (2019):

Introduction

The collapse of the Terramare system c. 1150 BC was followed by a sudden and substantial depopulation of the central part of the Po Plain (Cardarelli 2009). At the beginning of the Final Bronze Age, the southern part of the Po Valley was almost abandoned. In contrast, in the northern part of the Po Valley, some villages survived (…) Concurrently, a new territorial system arose, pivoting around the socio-economic pole of Frattesina (Calzavara Capuis et al. 1984; Bietti Sestieri et al. 2015; Cupitò et al. 2015). Therefore, within the area of the wider Terramare ‘culture’, local responses to the crisis led to different outcomes, some of which were relatively successful and others catastrophic. Economic factors—both in terms of internal carrying capacity and degree of openness to external relations—probably played a key role in determining different responses to the tensions.

The communities of the Terramare, especially in the southern area, were probably not flexible enough to adapt their political structure and modes of production to the needs of a rapidly changing world. Moreover, the domino effect from the overall geo-political instability of the twelfth century BC, in a highly interconnected system such as the Mediterranean, was undoubtedly another factor (Cardarelli 2009). The lack of evidence in the southern Terramare area for connections with the Aegean and the Levant suggests a more ‘closed’ system located on the edge of the ‘globalised’ world of the Late Bronze Age. In contrast, there is well-documented evidence from the largest terramare on the northern side of the Po River for possible incipient institutionalised, well-connected elites—particularly at Fondo Paviani, which has yielded locally produced pottery in Levantine and Late Helladic IIIC Aegean-Mycenaean styles (Bettelli et al. 2015).

The display of austere equality that dominated the Middle and Late Bronze Age ‘urnfields’ (Salzani 2005; Cardarelli 2014) strongly limited funerary expressions of social differentiation. Internal inequalities nonetheless existed between different co-resident extended families and lineages comprising tens of individuals at most (e.g. at Casinalbo; Cardarelli et al. 2014: 722–28), and, above all, between large centres, such as the terramara at Fondo Paviani and dependent satellite settlements (Balista et al. 2005; Cupitò et al. 2015). It seems reasonable therefore to hypothesise that groups based at nodal sites in the system attracted more prestige goods from exotic places, along with individuals from distant areas, while small villages attracted people mainly from within a local radius (Cavazzuti et al. 2019a). Within this dynamic cultural context, the Final Bronze Age funerary evidence from Frattesina documents a more elaborate display of power and wealth concentrated in the hands of elites. At Le Narde (Frattesina’s cemetery), this privileged segment of society, probably with its own entourage, is clearly represented by a small number of burials with several indicators of prestige.

bow-fibula-italy-aegean
Distribution of the violin-bow fibulae with two temple knots in the different source categories. Map by Sabine Pabst (2018).

Results

(…) the individual in burial Narde1-168 may have achieved the status of a ‘warrior-chief’, as symbolised by the presence of an Allerona-type sword (Bianco Peroni 1970). This was ritually broken and deposited in pieces inside the grave, along with a bronze pin, a pair of tweezers and other ornaments (Figure 8). (…) yielded a strontium isotope ratio (0.70983) that is incompatible with the local 0–20km baseline (Table 3), but fits within the 20–50km range. By contrast, the value obtained from the femoral cortical bone (0.70924) is consistent with the local range of Frattesina. This means that this individual moved to the site after early childhood—possibly during youth or early adulthood—and he probably spent the last years of his life there, at the apex of the community.

Marshall Sahlins (1981), in his famous article ‘The stranger-king: or Dumézil among the Fijians’, compares the dynamics of power in the Fiji Islands to the Indo-European tradition, arguing that human societies tend to locate power as originating from the outside (Sahlins 1981, 2008; see also Ling & Rowlands 2015). Sahlins focuses on origin myths across ancient polities in the Indo-European language area, which systematically feature a dichotomy between what the Romans called gravitas and celeritas. The former refers to the conservative, peaceful and productive character of an established native community, while celeritas represents the disruptive, transformative violence personified in the stranger king, who “erupts upon a pastoral scene of peaceful husbandry and political equality (or at least limited authority)” (Sahlins 1981: 112).

grave-goods-frattesina-warrior-chief-allerona-sword
The grave goods and cremated bones of burial Narde1-168 (after Salzani 1989). Urn height is 0.26m, sword length is 0.46m.

The individual buried in grave Narde1-168 at Frattesina was probably neither a true ‘king’, nor a true ‘stranger’. Despite its uniqueness, his grave resembles those of the rest of the community and is included within a large collective—or at least not evidently exclusive—burial mound. ‘Warrior-chief’ perhaps would be a more appropriate definition for this individual. Moreover, his place of origin was not so distant as to define him as a ‘stranger’. Nonetheless, Sahlins’s archetype of the ‘stranger-king’ evokes the power of alterity; burial Narde1-168 perfectly embodies celeritas, which breaks with the gravitas of the former Terramare tradition and guided whatever survived the collapse towards a new social model. Since the discovery of Frattesina and its cemeteries, Italian scholars have debated the mechanisms underlying the origin and economic success of the settlement, and the degree of foreign (i.e. Cypriot and Levantine) involvement in this process as suggested by archaeological finds (Cupitò et al. 2015). The new isotopic data presented here demonstrate that even though some individuals may have come from the Levant—where the available 87Sr/86Sr baseline ranges from 0.7079–0.7086 (Sheridan & Gregoricka 2015; Gregoricka & Guise Sheridan 2016)—or were from other exotic places, they nonetheless represent a minority of the population and, in any case, not the upper elite. The latter appear quite mobile, although probably from within the broader hinterland radius.

Adriatic or western route?

One of the interesting questions, and probably non-trivial for the correct interpretation of ancestry in future ancient DNA samples, is from where exactly did Tyrsenians come from, and more importantly where exactly did the arrive, and when. I have the impression that a Tyrrhenian Sea route is more commonly depicted (as in my maps) due to the historical predominance of Etruscans in the west, but that archaeologists usually consider the Adriatic – and thus a spread from the Po River Valley and/or Pannonia – a more likely route for Tyrsenian speakers, and probably rightly so.

NOTE. The tentative (and highly speculative) classification of fragmentary Rhaetian as more archaic than Etruscan relative to Lemnian may give further support to this route.

Failing a precise time transect from a population geographically close to the origin of their expansion in central or northern Italy, we are bound to see the same misinterpretations of the data we have seen in the case of Sea Peoples of hg. R1b behind Philistines. Nevertheless, here are some interesting predictions of population movements by Pabst (2013) based particularly on the Stätzling-/Allerona-sword from Narde in Veneto, which have been confirmed for the moment with isotope analyses, showing that some peoples of Frattesina had previously lived in the eastern Mediterranean, and that local elites had a much closer origin:

staetzling-swords
Distribution of the various blade profiles of the Stätzling (l) and Casale (H) type of leaf blade sword: 1 White symbols: blade with rapier-like ribs. – 2 black symbols: flat rhombic blade profile.- 3 Large gray symbols: a blade with a narrow midrib and longitudinal grooves.- Small gray symbols: lenticular or indefinite blade profile. (Map S. Pabst).

An Ingot fragment from the hoard of Hočko Pohorje in Styria, Slovenia indicates that possibly also Pannonia was involved in the 12th century BC (or during stage Ha A1) in the East and Central Mediterranean copper trade. According to the chemical composition or the high iron content, it is particularly close to individual finds from Sardinia, Italy and Anatolia.

The people behind the Stätzling swords could have been the potentates of this supraregional trade in the Adriatic and Ionian seas. This is also to be expected from the presence of late Mycenaean populations on the upper Adriatic. This is indicated – in addition to individual Mycenaean ceramics imports – especially in the Aegean Stätzling sword from the fly cave of Škocjan in the hinterland of Trieste, in this exchange network of the 12th century BC. However, not only people from the late Mycenaean cultural area were involved in the process. For native elites are suspected behind the mostly locally manufactured Stätzling swords in Pannonia and Italy, according to the analysis of the grave find 227 of Narde; perhaps local organizers of the trade, as allies of the Mycenaean chiefs.

Palaeogenomics

Palaeogenomics might help shed light upon the complex matter of the Tyrsenian emergence in Europe. Even though Rhaetian is a fragmentary language, it seems that it is related to Etruscan, and neither are remnant languages from the Bronze Age, but rather intrusive languages to Italy and Central Europe.

It is more than likely, then, that ancient DNA will show an increase in Aegean ancestry during the Late/Final Bronze Age in central and/or northern Italy, even if this change is found rapidly diluted within generations, as happened with the Aegean ancestry among Philistines, who – in spite of this dilution – also left their prolonged linguistic mark in the Levant.

This is the summary I made of an online report from oral communication A 12,000-year Genetic History of Rome and the Italian Peninsula, by Hannah Moots, the 6th February 2019, with 134 samples from Lazio and surrounding areas:

bronze-iron-age-romans-etruscans-osco-umbrians-map
Bronze Age – Iron Age evolution of Italy Top Left: Early Bronze Age cultures. Bottom left: PCA of groups from the Bronze Age; marked in red are previous Italy Bell Beakers. Top Right: Early Iron Age cultures. Bottom right: PCA of groups from the Iron Age – Middle Ages; marked in red are the approximate location of described ancient Italian clusters, one including Etruscans, Osco-Umbrians, Picentes, etc., and the wider cluster of Romans (dates unknown). See full maps and PCAs.

While Bronze Age samples of west-central Italy show a clear homogenisation of the genetic pool, with a shift in the PCA towards central Europe (away from the previous CHG/Iran Neolithic influence), and thus close to the modern Sardinian cluster, the few investigated Iron Age samples from the Republican period (ca. 700–20 BC) show a widespread genetic cluster encompassing the modern Italian ones, overlapping North Italian (ca. 60%) or South Italian/Sicilian (ca. 40%) clusters. The arrival or increase of EHG-, Levant Neolithic-, or CHG/IN-related ancestry in samples from this period suggest influence from previous population movements during the LBA from the north or through the Mediterranean, respectively. The Imperial Period shows influence from CHG/IN-related ancestry, but only sporadically Levant Neolithic.

NOTE. For more on the referred northern and southern Italian clusters, see Population structure of modern-day Italians reveals patterns of ancient and archaic ancestries in Southern Europe, by Raveane et al. bioRxiv (2018).

italian-north-south-clusters
Principal component analysis projecting 63 ancient individuals onto the components inferred from modern individuals. A) Principal component analysis projecting 63 ancient individuals onto the components inferred from 3,282 modern individuals assigned, through a CP/fS analysis, to European West Asian and Caucasian clusters.

The alternative view

Kristiansen is among those who offer an alternative view in the archaeological question, supporting the opposite direction of population movements: of Terramare migrants in Greece, a theory which is not to be lightly dismissed, in the complex setting of population movements across the Mediterranean during the Final Bronze Age.

As a weak linguistic support for such a movement, one can find the hypothesis of Eteo-Cretans as Osco-Umbrian speakers, based on de Ligt’s speculative interpretation of the Praisos inscription (Talanta 2008-2009).

It seems that, even if these views are also correct, the overwhelming evidence is for a foreign origin of Tyrsenians:

  • lack of Tyrsenian hydrotoponymic layers in Italy or central Europe;
  • guesstimates and “split” distribution of Italo-Venetic languages;
  • foreign culture and influences of Etruscans;
  • recent predominance and influence of Etruscan language and culture;
  • East Mediterranean peoples in the LBA Po Valley (isotope analyses);
  • genetics of Sea Peoples from the Aegean.

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

“Steppe ancestry” step by step: Khvalynsk, Sredni Stog, Repin, Yamna, Corded Ware

dzudzuana_pca-large

Wang et al. (2018) is obviously a game changer in many aspects. I have already written about the upcoming Yamna Hungary samples, about the new Steppe_Eneolithic and Caucasus Eneolithic keystones, and about the upcoming Greece Neolithic samples with steppe ancestry.

An interesting aspect of the paper, hidden among so many relevant details, is a clearer picture of how the so-called Yamnaya or steppe ancestry evolved from Samara hunter-gatherers to Yamna nomadic pastoralists, and how this ancestry appeared among Proto-Corded Ware populations.

anatolia-neolithic-steppe-eneolithic
Image modified from Wang et al. (2018). Marked are in orange: equivalent Steppe_Maykop ADMIXTURE; in red, approximate limit of Anatolia_Neolithic ancestry found in Yamna populations; in blue, Corded Ware-related groups. “Modelling results for the Steppe and Caucasus cluster. Admixture proportions based on (temporally and geographically) distal and proximal models, showing additional Anatolian farmer-related ancestry in Steppe groups as well as additional gene flow from the south in some of the Steppe groups as well as the Caucasus groups.”

Please note: arrows of “ancestry movement” in the following PCAs do not necessarily represent physical population movements, or even ethnolinguistic change. To avoid misinterpretations, I have depicted arrows with Y-DNA haplogroup migrations to represent the most likely true ethnolinguistic movements. Admixture graphics shown are from Wang et al. (2018), and also (the K12) from Mathieson et al. (2018).

1. Samara to Early Khvalynsk

The so-called steppe ancestry was born during the Khvalynsk expansion through the steppes, probably through exogamy of expanding elite clans (eventually all R1b-M269 lineages) originally of Samara_HG ancestry. The nearest group to the ANE-like ghost population with which Samara hunter-gatherers admixed is represented by the Steppe_Eneolithic / Steppe_Maykop cluster (from the Northern Caucasus Piedmont).

Steppe_Eneolithic samples, of R1b1 lineages, are probably expanded Khvalynsk peoples, showing thus a proximate ancestry of an Early Eneolithic ghost population of the Northern Caucasus. Steppe_Maykop samples represent a later replacement of this Steppe_Eneolithic population – and/or a similar population with further contribution of ANE-like ancestry – in the area some 1,000 years later.

PCA-caucasus-steppe-samara

This is what Steppe_Maykop looks like, different from Steppe_Eneolithic:

steppe-maykop-admixture

NOTE. This admixture shows how different Steppe_Maykop is from Steppe_Eneolithic, but in the different supervised ADMIXTURE graphics below Maykop_Eneolithic is roughly equivalent to Eneolithic_Steppe (see orange arrow in ADMIXTURE graphic above). This is useful for a simplified analysis, but actual differences between Khvalynsk, Sredni Stog, Afanasevo, Yamna and Corded Ware are probably underestimated in the analyses below, and will become clearer in the future when more ancestral hunter-gatherer populations are added to the analysis.

2. Early Khvalynsk expansion

We have direct data of Khvalynsk-Novodanilovka-like populations thanks to Khvalynsk and Steppe_Eneolithic samples (although I’ve used the latter above to represent the ghost Caucasus population with which Samara_HG admixed).

We also have indirect data. First, there is the PCA with outliers:

PCA-khvalynsk-steppe

Second, we have data from north Pontic Ukraine_Eneolithic samples (see next section).

Third, there is the continuity of late Repin / Afanasevo with Steppe_Eneolithic (see below).

3. Proto-Corded Ware expansion

It is unclear if R1a-M459 subclades were continuously in the steppe and resurged after the Khvalynsk expansion, or (the most likely option) they came from the forested region of the Upper Dnieper area, possibly from previous expansions there with hunter-gatherer pottery.

Supporting the latter is the millennia-long continuity of R1b-V88 and I2a2 subclades in the north Pontic Mesolithic, Neolithic, and Early Eneolithic Sredni Stog culture, until ca. 4500 BC (and even later, during the second half).

Only at the end of the Early Eneolithic with the disappearance of Novodanilovka (and beginning of the steppe ‘hiatus’ of Rassamakin) is R1a to be found in Ukraine again (after disappearing from the record some 2,000 years earlier), related to complex population movements in the north Pontic area.

NOTE. In the PCA, a tentative position of Novodanilovka closer to Anatolia_Neolithic / Dzudzuana ancestry is selected, based on the apparent cline formed by Ukraine_Eneolithic samples, and on the position and ancestry of Sredni Stog, Yamna, and Corded Ware later. A good alternative would be to place Novodanilovka still closer to the Balkan outliers (i.e. Suvorovo), and a source closer to EHG as the ancestry driven by the migration of R1a-M417.

PCA-sredni-stog-steppe

The first sample with steppe ancestry appears only after 4250 BC in the forest-steppe, centuries after the samples with steppe ancestry from the Northern Caucasus and the Balkans, which points to exogamy of expanding R1a-M417 lineages with the remnants of the Novodanilovka population.

steppe-ancestry-admixture-sredni-stog

4. Repin / Early Yamna expansion

We don’t have direct data on early Repin settlers. But we do have a very close representative: Afanasevo, a population we know comes directly from the Repin/late Khvalynsk expansion ca. 3500/3300 BC (just before the emergence of Early Yamna), and which shows fully Steppe_Eneolithic-like ancestry.

afanasevo-admixture

Compared to this eastern Repin expansion that gave Afanasevo, the late Repin expansion to the west ca. 3300 BC that gave rise to the Yamna culture was one of colonization, evidenced by the admixture with north Pontic (Sredni Stog-like) populations, no doubt through exogamy:

PCA-repin-yamna

This admixture is also found (in lesser proportion) in east Yamna groups, which supports the high mobility and exogamy practices among western and eastern Yamna clans, not only with locals:

yamnaya-admixture

5. Corded Ware

Corded Ware represents a quite homogeneous expansion of a late Sredni Stog population, compatible with the traditional location of Proto-Corded Ware peoples in the steppe-forest/forest zone of the Dnieper-Dniester region.

PCA-latvia-ln-steppe

We don’t have a comparison with Ukraine_Eneolithic or Corded Ware samples in Wang et al. (2018), but we do have proximate sources for Abashevo, when compared to the Poltavka population (with which it admixed in the Volga-Ural steppes): Sintashta, Potapovka, Srubna (with further Abashevo contribution), and Andronovo:

sintashta-poltavka-andronovo-admixture

The two CWC outliers from the Baltic show what I thought was an admixture with Yamna. However, given the previous mixture of Eneolithic_Steppe in north Pontic steppe-forest populations, this elevated “steppe ancestry” found in Baltic_LN (similar to west Yamna) seems rather an admixture of Baltic sub-Neolithic peoples with a north Pontic Eneolithic_Steppe-like population. Late Repin settlers also admixed with a similar population during its colonization of the north Pontic area, hence the Baltic_LN – west Yamna similarities.

NOTE. A direct admixture with west Yamna populations through exogamy by the ancestors of this Baltic population cannot be ruled out yet (without direct access to more samples), though, because of the contacts of Corded Ware with west Yamna settlers in the forest-steppe regions.

steppe-ancestry-admixture-latvia

A similar case is found in the Yamna outlier from Mednikarovo south of the Danube. It would be absurd to think that Yamna from the Balkans comes from Corded Ware (or vice versa), just because the former is closer in the PCA to the latter than other Yamna samples. The same error is also found e.g. in the Corded Ware → Bell Beaker theory, because of their proximity in the PCA and their shared “steppe ancestry”. All those theories have been proven already wrong.

NOTE. A similar fallacy is found in potential Sintashta→Mycenaean connections, where we should distinguish statistically that result from an East/West Yamna + Balkans_BA admixture. In fact, genetic links of Mycenaeans with west Yamna settlers prove this (there are some related analyses in Anthrogenica, but the site is down at this moment). To try to relate these two populations (separated more than 1,000 years before Sintashta) is like comparing ancient populations to modern ones, without the intermediate samples to trace the real anthropological trail of what is found…Pure numbers and wishful thinking.

Conclusion

Yamna and Corded Ware show a similar “steppe ancestry” due to convergence. I have said so many times (see e.g. here). This was clear long ago, just by looking at the Y-chromosome bottlenecks that differentiate them – and Tomenable noticed this difference in ADMIXTURE from the supplementary materials in Mathieson et al. (2017), well before Wang et al. (2018).

This different stock stems from (1) completely different ancestral populations + (2) different, long-lasting Y-chromosome bottlenecks. Their similarities come from the two neighbouring cultures admixing with similar populations.

If all this does not mean anything, and each lab was going to support some pre-selected archaeological theories from the 1960s or the 1980s, coupled with outdated linguistic models no matter what – Anthony’s model + Ringe’s glottochronological tree of the early 2000s in the Reich Lab; and worse, Kristiansen’s CWC-IE + Germano-Slavonic models of the 1940s in the Copenhagen group – , I have to repeat my question again:

What’s (so much published) ancient DNA useful for, exactly?

See also

Related

Trypillia and Greece Neolithic outliers: the smoking gun of Proto-Anatolian migrations?

neolithic-migrations-khvalynsk-novodanilovka-anatolian

(Continued from the post Corded Ware culture origins: The Final Frontier).

Looking at the PCA of Wang et al. (2018), I realized that Sredni Stog / Corded Ware peoples seem to lie somewhere between:

  • the eastern steppe (i.e. Khvalynsk-Yamna); and
  • Lower Danube and Balkan cultures affected by Anatolian- and steppe-related (i.e. Khvalynsk-Novodanilovka) migrations.

This multiethnic interaction of the western steppe fits therefore the complex archaeological description of events in the North Pontic, Lower Danube, and Dnieper-Dniester regions. Here are some interesting samples related to those long-lasting contacts:

1. I3719 (mtDNA H1, Y-DNA I2a2a) Ukraine Neolithic sample from Dereivka ca. 4949–4799 BC, described in Mathieson et al. (2018) as of “entirely northwestern-Anatolian-Neolithic-related ancestry”.

2. ANI163 from Varna I ca. 4711–4550 BC (mtDNA H7a1), and I2181 from Balkans Chalcolithic (Smyadovo, in Bulgaria) ca. 4500 BC (mtDNA HV15, Y-DNA R) show the first steppe ancestry in regions known to be affected by the expansion of Suvorovo chiefs.

3. The Yamna Bulgaria outlier (Y-DNA I2a2a1b1b), 3012-2900 calBCE, shows apparently an admixture with cultures of that region (but 1,500 years later).

PCA-trypillia-greece-neolithic-outlier-anatolian
Image modified from Wang et al. (2018). Samples projected in PCA of 84 modern-day West Eurasian populations (open symbols). Previously known clusters have been marked and referenced. Marked and labelled are the Balkan samples referenced in this text An EHG and a Caucasus ‘clouds’ have been drawn, leaving Pontic-Caspian steppe and derived groups between them. See the original file here.

Trypillia and Corded Ware

4. There is one ‘Trypillia outlier’ among five samples from the Verteba cave in Wang et al. (2018): I1927 (Y-DNA G2a2b2a1a1b1a1a1, mtDNA H1b), ca. 3619-2936 BC, a sample published previously in Nikitin et al. (2017) and Mathieson et al. (2017). We were very quick to dismiss Trypillia (three samples of haplogroup G2a, one sample E) and GAC as a source of Corded Ware admixture, but archaeology clearly shows important population movements at the end of the fourth millennium between late Trypillia groups, GAC, and post-Sredni Stog populations, and genetics is showing that in both cultures, too.

I am not a fan of the ‘lack of samples’ argument, but (similar to Old Hittite samples related to all Anatolian speakers) one site is not enough to describe a culture that spanned millennia and many different early and late groups. One among five Trypillian samples (from a single site), showing a late date (ca. 3228 BC) compared to the other samples (ca. 3700 BC), and quite close to the only three Ukraine Eneolithic samples we have may mean much more than what we may a priori think, i.e. some simplistic unidirectional punctual ‘intrusion’ of steppe ancestry, and instead hint at the known close contacts of late Trypillian groups and North Pontic cultures, including also the Caucasus.

NOTE. The big difference in PCA among GAC-like Hungary LCA – EBA samples (see above two star symbols close to Ukraine Neolithic outlier in the PCA, in contrast with the other three at the bottom) may also be significant, although we don’t have any data about their culture, sites, or the relationship between them.

trypillia-verteba-cave
Location of Verteba Cave in relation to different stages and neighbouring groups of the Cucuteni-Trypillia culture. Image from the paper A Subterranean Sanctuary of the Cucuteni-Trypillia Culture in Western Ukraine, by Kadrow and Pokutta (2016).

Greece Neolithic outlier: Proto-Anatolians?

5. Especially interesting is I6423, one of the Greece Neolithic samples referred to in Wang et al. (2018), which is obviously an outlier among the three used in the paper. It does not seem to correspond to any of the ancient DNA samples published to date; it is not in Hofmanova et al. (2016), in Lazaridis et al. (2017), or in Mathieson et al. (2018).

Since the Neolithic in Greece could mean any period from ca. 6500 BC to ca. 3200 BC, I guess we are talking here about some migration related to the expansion of Khvalynsk-Novodanilovka chieftains after ca. 4500 BC, because it appears on the PCA precisely on the same spot as Varna and Smyadovo outliers, and its ADMIXTURE shows similar components

admixture-ukraine-eneolithic-greece-neolithic
Image modified from Wang et al. (2018). “ADMIXTURE results of relevant prehistoric individuals mentioned in the text (filled symbols)”. ‘Outlier’ samples referred to in this post have been marked in red. See the original file here.

So, this may be the smoking gun of Proto-Anatolian (or maybe early Common Anatolian) expansion with steppe migrants up to the border of Western Anatolia, and we may be able to get rid of those unfounded doubts about Anatolian origins once and for all…

NOTE. Also interesting seems another Greece Neolithic sample, I6420, in ADMIXTURE, although its position in the PCA (near Minoans and Mycenaeans) does not necessarily point to potential steppe influence, but rather to the extra ‘eastern (Caucasus/Iran-related) ancestry’ contribution found in Minoans and in Mycenaeans (and Anatolia Chalcolithic) compared to previous samples of the region. The third Greece Nelithic sample, I5427 (mtDNA K1a24), from Diros, Alepotrypa Cave, is dated 6005-5879 BC (mean 5892 BC), and appeared first in Mathieson et al. (2018).

greece-neolithic
Modified from Wang et al. (2018) (Greece_Neolithic in red). Supplementary Table 6. P values of rank=1 in modelling the two-way admixture in the Caucasus cluster. 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. Source 2 populations in bold print are used as examples in modelling the Caucasus cluster groups (See Supplementary Table 7).
greece-neolithic-caucasus
Modified from Wang et al. (2018) (Greece_Neolithic in red). Supplementary Table 10. P values of rank=2 and ancestry proportions in modelling a three-way admixture in the Caucasus cluster testing additional contribution from Iran_ChL. Here, we used an extended set of outgroup populations populations to constrain standard errors: 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, EHG, WHG, Levant_N.

If this Greece Neolithic sample is not related to Yamna migrations – and its use for statistical analysis of Caucasus samples from Wang et al. (2018) suggests that it is not – , it may have important consequences:

If it is located near the Western Anatolian coast – especially near Troy – there won’t be much to add about the potential site of entry of Common Anatolian languages into Anatolia… I have read some comments about how ‘impossible’ it was for steppe migrants and their language to ‘invade’ the more advanced cultures of Anatolia from the west, but it seems as ‘impossible’ as it was for Barbarians to invade the Roman Empire and impose their language as elites in certain regions. (And yes, we have at least one important weak political period among Middle Eastern cultures in the early 3rd millennium BC, similar to the period of the Fall of the Western Roman Empire).

indo-european-anatolian-uralic-migrations
Most likely Proto-Anatolian expansion in the North Pontic and Balkan area with Khvalynsk-Novodanilovka chieftains, including ADMIXTURE data from Wang et al. (2018).

If it is located somewhere more ‘central’ in the Greek Peninsula, then it could also be used to support the Anatolian nature of the controversial Pre-Greek (‘Pelasgian’) substrate. While we know that Greek (at least since Mycenaean) shows a strong Pre-Greek cultural and linguistic heritage (also reflected in its genetic continuity), the nature of that language is usually believed to be non-Indo-European, and Anatolian contacts are rather few and coincident with the Mycenaean period. I don’t think this sample can tell much about the Pre-Greek language, though, because – if it is really Neolithic, and comparing it with later Minoan and Mycenaean samples – it seems a clear outlier.

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

If it is, however, related to later Yamna migrations after ca. 3000 BC (and, like the ‘Ukraine Eneolithic’ sample that is likely from Catacomb, it is classified as Neolithic just because it cannot be attributed to precise Helladic periods), then we may be in front of the first obvious Yamna migrants in Greece. If that is the case (which I doubt), the sample wouldn’t be so informative for PIE dialectal expansions, because by now it is evident that we will find steppe ancestry and R1b-Z2103 subclades accompanying Yamna migrants in the southern Balkans, and probably well into Mycenaean Greece.

NOTE. Whatever the case, I am sure that for those fond of absurd autochthonous continuity theories, as well as for anti-steppe conspirationists, this sample will be just another good way of arguing for anything, ranging from a rejection of the Middle PIE – Late PIE division, to a support for some mythic ancient autochtonous Proto-Graeco-Anatolian group, or maybe some ancient Graeco–Indo-Slavonic split, or whatever new dialectal stage one can invent to support the own genealogical fantasies…

So, if it actually is a Neolithic sample, let’s hope that it shows a clear R1b-M269 (xL23 or early L23) subclade distinct from those (likely Z2103) expanded later with Late PIE-speaking Yamna (and probably to be found among Mycenaeans), so that there can be no more place for ethnic fantasies.

EDIT (28 JUL 2018): Added information on Greece Neolithic and Trypillia samples

Related

Luwians: the missing link with the Aegean Bronze Age, including Troy and the Sea Peoples

luwians-sea-peoples

A very interesting monograph on the Luwian Civilization, and its potential connection with Wilusa (Troy) from the end of the third millennium and throughout the Bronze Age: The Luwian Civilization – The Missing Link in the Aegean Bronze Age, by Eberhard Zangger (also available in German: Die luwische Kultur – Das fehlende Element in der Ägäischen Bronzezeit).

Abstract:

Aegean prehistory suffered from a bias when the field was conceived 100 years ago and subsequent research has never questioned the fundamental paradigms of the discipline. As a consequence, only one third of the Aegean coasts have thus far been attributed to ancient civilizations. This leaves tremendous opportunities for current and future generations of archaeologists – on the somewhat neglected eastern side of the Aegean. Practically all contemporary sources indicate that Late Bronze Age petty states used to form military alliances. The Assuwa league, mentioned in Hittite texts from around 1400 BCE, is a good example, but so are the mercenary forces mustered by Muwatalli and the various accounts of united tribes from the Aegean (aka “Sea Peoples”) and even – in later recollections of past events – Homer’s catalogues of ships and Trojan contingents. “The Luwian Civilization” argues that such a coalition of the petty states in western Asia Minor may have succeeded in bringing down the Hittite hegemony over central Asia Minor.

Excerpt (from the Introduction):

Possibly due to its vast extent and complicated topography, for thousands of years the majority of western Asia Minor was politically fragmented into many petty kingdoms and principalities. This certainly weakened the region in its economic and political significance, but it also delayed the recognition of a more or less consistent Luwian culture.

From a linguistics point of view, however, the Luwian culture is relatively well known. From about 2000 BCE Luwian personal names and loanwords appear in Assyrian documents retrieved from the trading town Kültepe (also Kaniš or Neša). Assyrian merchants who lived in Asia Minor at the time described the indigenous population as nuwa’um, corresponding to “Luwians.” At about the same time, early Hittite settlements arose a little further north at the upper Kızılırmak River. In documents from the Hittite capital Hattuša written in Akkadian cuneiform, western Asia Minor is originally called Luwiya. Hittite laws and other documents also contain references to translations into “Luwian language.” Accordingly, Luwian was spoken in various dialects throughout southern and western Anatolia. The language belongs to the Anatolian branch of Indo-European languages. It was recorded in Akkadian cuneiform on the one hand, but also in its own hieroglyphic script, one that was used over a timespan of at least 1400 years (2000–600 BCE). Luwian hieroglyphic ranks, therefore, as the first script in which an Indo-European language is transcribed. The people using this script and speaking a Luwian language lived during the Bronze and Early Iron Age in Asia Minor and northern Syria.

luwian-language
The region where Luwian was spoken at the end of the Bronze Age was much larger than the one where
Hittite was spoken.

Highly recommended for anyone interested in assessing where and when samples with steppe admixture and Y-DNA haplogroup R1b-M269 might appear next in Anatolian samples – and how to interpret them correctly.

Related:

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

mycenaean-minoan

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

Abstract:

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

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

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

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

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

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

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

Related:

Featured map: samples studied, from the article.