Yamnaya replaced Europeans, but admixed heavily as they spread to Asia

narasimhan-spread-yamnaya-ancestry

Recent papers The formation of human populations in South and Central Asia, by Narasimhan, Patterson et al. Science (2019) and An Ancient Harappan Genome Lacks Ancestry from Steppe Pastoralists or Iranian Farmers, by Shinde et al. Cell (2019).

NOTE. For direct access to Narasimhan, Patterson et al. (2019), visit this link courtesy of the first author and the Reich Lab.

I am currently not on holidays anymore, and the information in the paper is huge, with many complex issues raised by the new samples and analyses rather than solved, so I will stick to the Indo-European question, especially to some details that have changed since the publication of the preprint. For a summary of its previous findings, see the book series A Song of Sheep and Horses, in particular the sections from A Clash of Chiefs where I discuss languages and regions related to Central and South Asia.

I have updated the maps of the Preshistory Atlas, and included the most recently reported mtDNA and Y-DNA subclades. I will try to update the Eurasian PCA and related graphics, too.

NOTE. Many subclades from this paper have been reported by Kolgeh (download), Pribislav and Principe at Anthrogenica on this thread. I have checked some out for comparison, but even if it contradicted their analyses mine would be the wrong ones. I will upload my spreadsheets and link to them from this page whenever I find the time.

caucasus-cline-narasimhan
Ancestry clines (1) before and (2) after the advent of farming. Colour modified from the original to emphasize the CHG cline: notice the apparent relevance of forest-steppe groups in the formation of this CHG mating network from which Pre-Yamnaya peoples emerged.

Indo-Europeans

I think the Narasimhan, Patterson et al. (2019) paper is well-balanced, and unexpectedly centered – as it should – on the spread of Yamnaya-related ancestry (now Western_Steppe_EMBA) as the marker of Proto-Indo-European migrations, which stretched ca. 3000 BC “from Hungary in the west to the Altai mountains in the east”, spreading later Indo-European dialects after admixing with local groups, from the Atlantic to South Asia.

I. Afanasievo

I.1. East or West PIE?

I expected Afanasievo to show (1) R1b-L23(xZ2103, xL51) and (2) R1b-L51 lineages, apart from (3) the known R1b-Z2103 ones, pointing thus to an ancestral PIE community before the typical Yamnaya bottlenecks, and with R1b-L51 supporting a connection with North-West Indo-European. The presence of some samples of hg. Q pointed in this direction, too.

However, Afanasievo samples show overwhelmingly R1b-Z2103 subclades (all except for those with low coverage), all apparently under R1b-Z2108 (formed ca. 3500 BC, TMRCA ca. 3500 BC), like most samples from East Yamnaya.

This necessarily shifts the split and spread of R1b-L23 lineages to Khvalynsk/early Repin-related expansions, in line with what TMRCA suggested, and what advances by Anthony (2019) and Khokhlov (2018) on future samples from the Reich Lab suggest.

Given the almost indistinguishable ancestry between Afanasievo and Early Yamnaya, there seems to be as of yet little potential information to support in population genomics that Pre-Tocharians were more closely related to North-West Indo-Europeans than to Graeco-Aryans, as it is proposed in linguistics based on the few shared traits between them, and the lack of innovations proper of the Graeco-Aryan community.

NOTE. A new issue of Wekʷos contains an abstract from a relevant paper by Blažek on vocabulary for ‘word’, including the common NWIE *wrdʰo-/wordʰo-, but also a new (for me, at least) Northern Indo-European one: *rēki-/*rēkoi̯-, shared by Slavic and Tocharian.

The fact that bottlenecks happened around the time of the late Repin expansion suggests that we might be able to see different clans based on the predominant lineages developing around the Don-Volga area in the 4th millennium BC. The finding of Pre-R1b-L51 in Lopatino (see below), and of a Catacomb sample of hg. R1b-Z2103(Z2105-) in the North Caucasus steppe near Novoaleksandrovskij also support a star-like phylogeny of R1b-L23 stemming from the Don-Volga area.

NOTE. Interestingly, a dismissal of a common trunk between Tocharian and North-West Indo-European would mean that shared similarities between such disparate groups could be traced back to a Common Late PIE trunk, and not to a shared (western) Repin community. For an example of such a ‘pure’ East-West dialectal division, see the diagram of Adams & Mallory (2007) at the end of the post. It would thus mean a fatal blow to Kortlandt’s Indo-Slavonic group among other hypothetical groupings (remade versions of the ancient Centum-Satem division), as well as to certain assumptions about laryngeal survival or tritectalism that usually accompany them. Still, I don’t think this is the case, so the question will remain a linguistic one, and maybe some similarities will be found with enough number of samples that differentiate Northern Indo-Europeans from the East Yamna/Catacomb-Poltavka-Balkan_EBA group.

afanasievo-y-dna
Y-chromosome haplogroups of Afanasievo samples and neighbouring groups. See full maps.

I.2. Expansion or resurgence of hg. Q1b?

Haplogroup Q1b-Y6802(xY6798) seems to be the main lineage that expanded with Afanasievo, or resurged in their territory. It’s difficult to tell, because the three available samples are family, and belong to a later period.

NOTE. I have finally put some order to the chaos of Q1a vs. Q1b subclades in my spreadsheet and in the maps. The change of ISOGG 2016 to 2017 has caused that many samples reported as of Q1 subclades from papers prepared during the 2017-2018 period, and which did not provide specific SNP calls, were impossible to define with certainty. By checking some of them I could determine the specific standard used.

In favour of the presence of this haplogroup in the Pre-Yamnaya community are:

  • The statement by Anthony (2019) that Q1a [hence maybe Q1b in the new ISOGG nomenclature] represented a significant minority among an R1b-rich community.
  • The sample found in a Sintastha WSHG outlier (see below), of hg. Q1b-Y6798, and the sample from Lola, of hg. Q1b-L717, are thus from other lineage(s) separated thousands of years from the Afanasievo subclade, but might be related to the Khvalynsk expansion, like R1b-V1636 and R1b-M269 are.

These are the data that suggest multiple resurgence events in Afanasievo, rather than expanding Q1b lineages with late Repin:

  • Overwhelming presence of R1b in early Yamnaya and Afanasievo samples; one Q1(xQ1b) sample reported in Khvalynsk.
  • The three Q1b samples appear only later, although wide CI for radiocarbon dates, different sites, and indistinguishable ancestry may preclude a proper interpretation of the only available family.
    • Nevertheless, ancestry seems unimportant in the case of Afanasievo, since the same ancestry is found up to the Iron Age in a community of varied haplogroups.
  • Another sample of hg. Q1b-Y6802(xY6798) is found in Aigyrzhal_BA (ca. 2120 BC), with Central_Steppe_EMBA (WSHG-related) ancestry; however, this clade formed and expanded ca. 14000 BC.
  • The whole Altai – Baikal area seems to be a Q1b-L54 hotspot, although admittedly many subclades separated very early from each other, so they might be found throughout North Eurasia during the Neolithic.
  • One Afanasievo sample is reported as of hg. C in Shin (2017), and the same haplogroup is reported by Hollard (2014) for the only available sample of early Chemurchek to date, from Kulala ula, North Altai (ca. 2400 BC).
afanasievo-chemurchek-y-dna
Y-chromosome haplogroups of late Afanasievo – early Chemurchek samples and neighbouring groups. See full maps.

I.3. Agricultural substrate

Evidence of continuous contacts of Central_Steppe_MLBA populations with BMAC from ca. 2100 BC on – visible in the appearance of Steppe ancestry among BMAC samples and BMAC ancestry among Steppe pastoralists – supports the close interaction between Indo-Iranian pastoralists and BMAC agriculturalists as the origin of the Asian agricultural substrate found in Proto-Indo-Iranian, hence likely related to the language of the Oxus Civilization.

Similar to the European agricultural substrate adopted by West Yamnaya settlers (both NWIE and Palaeo-Balkan speakers), Tocharian shows a few substrate terms in common with Indo-Iranian, which can be explained by contacts in different dialectal stages through phonetic reconstruction alone.

The recent Hermes et al. (2019) supports the early integration of pastoralism and millet cultivation in Central Asia (ca. 2700 BC or earlier), with the spread of agriculture to the north – through the Inner Asian Mountain Corridor – being thus unrelated to the Indo-Iranian expansions, which might support independent loans.

However, compared to the huge number of parallel shared loans between NWIE and Palaeo-Balkan languages in the European substratum, Indo-Iranians seem to have been the first borrowers of vocabulary from Asian agriculturalists, while Proto-Tocharian shows just one certain related word, with phonetic similarities that warrant an adoption from late Indo-Iranian dialects.

chemurchek-sintashta-bmac
Y-chromosome haplogroups of Sintashta, Central Asia, and neighbouring groups in the Early Bronze Age. See full maps.

The finding of hg. (pre-)R1b-PH155 in a BMAC sample from Dzharkutan (to the west of Xinjiang) together with hg. R1b in a sample from Central Mongolia previously reported by Shin (2017) support the widespread presence of this lineage to the east and west of Xinjiang, which means it might have become incorporated to Indo-Iranian migrants into the Xiaohe horizon, to the Afanasievo-Chemurchek-derived groups, or the later from the former. In other words, the Island Biogeography Theory with its explanation of founder effects might be, after all, applicable to the whole Xinjiang area, not only during the Chemurchek – Tianshan-Beilu – Xiaohe interaction.

Of course, there is no need for too complicated models of haplogroup resurgence events in Central and South Asia, seeing how the total amount of hg. R1a-L657 (today prevalent among Indo-Aryan speakers from South Asia) among ancient Western/Central_Steppe_MLBA-related samples amounts to a total of 0, and that many different lineages survived in the region. Similar cases of haplogroup resurgence and Y-DNA bottleneck events are also found in the Central and Eastern Mediterranean, and in North-Eastern Europe. From the paper:

[It] could reflect stronger ecological or cultural barriers to the spread of people in South Asia than in Europe, allowing the previously established groups more time to adapt and mix with incoming groups. A second difference is the smaller proportion of Steppe pastoralist– related ancestry in South Asia compared with Europe, its later arrival by ~500 to 1000 years, and a lower (albeit still significant) male sex bias in the admixture (…).

Y-chromosome haplogroups of samples from the Srubna-Andronovo and Andronovo-related horizon, Xiaohe, late BMAC, and neighbouring groups. See full maps.

II. R1b-Beakers replaced R1a-CWC peoples

II.1. R1a-M417-rich Corded Ware

Newly reported Corded Ware samples from Radovesice show hg. R1a-M417, at least some of them xZ645, ‘archaic’ lineages shared with the early Bergrheinfeld sample (ca. 2650 BC) and with the coeval Esperstedt family, hence supporting that it eventually became the typical Western Corded Ware lineage(s), probably dominating over the so-called A-horizon and the Single Grave culture in particular. On the other hand, R1a-Z645 was typical of bottlenecks among expanding Eastern Corded Ware groups.

Interestingly, it is supported once again that known bottlenecks under hg. R1a-M417 happened during the Corded Ware expansion, evidenced also by the remarkable high variability of male lineages among early Corded Ware samples. Similarly, these Corded Ware samples from Bohemia form part of the typical ‘Central European’ cluster in the PCA, which excludes once again not only the ‘official’ Espersted outlier I1540, but also the known outlier with Yamnaya ancestry.

NOTE. The fact that Esperstedt is closely related geographically and in terms of ancestry to later Únětice samples further complicates the assumption that Únětice is a mixture of Bell Beakers and Corded Ware, being rather an admixture of incoming Bell Beakers with post-Yamnaya vanguard settlers who admixed with Corded Ware (see more on the expansion of Yamnaya ancestry). In other words, Únětice is rather an admixture of Yamnaya+EEF with Yamnaya+(CWC+EEF).

Y-chromosome haplogroups of samples from Catacomb, Poltavka, Balkan EBA, and Bell Beaker, as well as neighbouring groups. See full maps.

On Ukraine_Eneolithic I6561

If the bottlenecks are as straightforward as they appear, with a star-like phylogeny of R1a-M417 starting with the Pre-Corded Ware expansion, then what is happening with the Alexandria sample, so precisely radiocarbon dated to ca. 4045-3974 BC? The reported hg. R1a-M417 was fully compatible, while R1a-Z645 could be compatible with its date, but the few positive SNPs I got in my analysis point indeed to a potential subclade of R1a-Z94, and I trust more experienced hobbyists in this ‘art’ of ascertaining the SNPs of ancient samples, and they report hg. R1a-Z93 (Z95+, Y26+, Y2-).

Seeing how Y-DNA bottlenecks worked in Yamnaya-Afanasievo and in Corded Ware and related groups, and if this sample really is so deep within R1a-Z93 in a region that should be more strongly affected by the known Neolithic Y-chromosome bottlenecks and forest-steppe ecotone, someone from the lab responsible for this sample should check its date once again, before more people keep chasing their tails with an individual that (based on its derived SNPs’ TMRCA) might actually be dated to the Bronze Age, where it could make much more sense in terms of ancestry and position in the PCA.

EDIT (14 SEP 2019): … and with the fact that he is the first individual to show the genetic adaptation for lactase persistence (I3910-T), which is only found later among Bell Beakers, and much later in Sintashta and related Steppe_MLBA peoples (see comments below).

This is also evidenced by the other Ukraine_Eneolithic (likely a late Yamnaya) sample of hg. R1b-Z2103 from Dereivka (ca. 2800 BC) and who – despite being in a similar territory 1,000 years later – shows a wholly diluted Yamnaya ancestry under typically European HG ancestry, even more so than other late Sredni Stog samples from Dereivka of ca. 3600-3400 BC, suggesting a decrease in Steppe ancestry rather than an increase – which is supposedly what should be expected based on the ancestry from Alexandria…

Like the reported Chalcolithic individual of Hajji Firuz who showed an apparently incompatible subclade and Yamnaya ancestry at least some 1,000 years before it should, and turned out to be from the Iron Age (see below), this may be another case of wrong radiocarbon dating.

NOTE. It would be interesting, if this turns out to be another Hajji Firuz-like error, to check how well different ancestry models worked in whose hands exactly, and if anyone actually pointed out that this sample was derived, and not ancestral, to many different samples that were used in combination with it. It would also be a great control to check if those still supporting a Sredni Stog origin for PIE would shift their preference even more to the north or west, depending on where the first “true” R1a-M417 samples popped up. Such a finding now could be thus a great tool to discover whether haplogroup-based bias plays a role in ancestry magic as related to the Indo-European question, i.e. if it really is about “pure statistics”, or there is something else to it…

II.1. R1b-L51-rich Bell Beakers

The overwhelming majority of R1b-L51 lineages in Radovesice during the Bell Beaker period, just after the sampled Corded Ware individuals from the same site, further strengthen the hypothesis of an almost full replacement of R1a-M417 lineages from Central Europe up to southern Scandinavia after the arrival of Bell Beakers.

Yet another R1b-L151* sample has popped up in Central Europe, in the individual classified as Bilina_BA (ca. 2200-800 BC), which clusters with Bell Beakers from Bohemia, with the outlier from Turlojiškė, and with Early Slavs, suggesting once again that a group of central-east European Beakers represented the Pre-Proto-Balto-Slavic community before their spread and admixture events to the east.

The available ancient distribution of R1b-L51*, R1b-L52* or R1b-L151* is getting thus closer to the most likely origin of R1b-L51 in the expansion of East Bell Beakers, who trace their paternal ancestors to Yamnaya settlers from the Carpathian Basin:

NOTE. Some of these are from other sources, and some are samples I have checked in a hurry, so I may have missed some derived SNPs. If you send me a corrected SNP call to dismiss one of these, or more ‘archaic’ samples, I’ll correct the map accordingly. See also maps of modern distributionof R1b-M269 subclades .

r1b-l51-ancient-europe
Distribution of ‘archaic’ R1b-L51 subclades in ancient samples, overlaid over a map of Yamnaya and Bell Beaker migrations. In blue, Yamnaya Pre-L51 from Lopatino (not shown) and R1b-L52* from BBC Augsburg. In violet, R1b-L51 (xP312,xU106) from BBC Prague and Poland. In maroon, hg. R1b-L151* from BBC Hungary, BA Bohemia, and (not shown) a potential sample from BBC at Mondelange, which is certainly xU106, maybe xP312. Interestingly, the earliest sample of hg. R1b-U106 (a lineage more proper of northern Europe) has been found in a Bell Beaker from Radovesice (ca. 2350 BC), between two of these ‘archaic’ R1b-L51 samples; and a sample possibly of hg. R1b-ZZ11+ (ancestral to DF27 and U152) was found in a Bell Beaker from Quedlinburg, Germany (ca. 2290 BC), to the north-west of Bohemia. The oldest R1b-U152 are logically from Central Europe, too.

III. Proto-Indo-Iranian

Before the emergence of Proto-Indo-Iranian, it seems that Pre-Proto-Indo-Iranian-speaking Poltavka groups were subjected to pressure from Central_Steppe_EMBA-related peoples coming from the (south-?)east, such as those found sampled from Mereke_BA. Their ‘kurgan’ culture was dated correctly to approximately the same date as Poltavka materials, but their ancestry and hg. N2(pre-N2a) – also found in a previous sample from Botai – point to their intrusive nature, and thus to difficulties in the Pre-Proto-Indo-Iranian community to keep control over the previous East Yamnaya territory in the Don-Volga-Ural steppes.

We know that the region does not show genetic continuity with a previous period (or was not under this ‘eastern’ pressure) because of an Eastern Yamnaya sample from the same site (ca. 3100 BC) showing typical Yamnaya ancestry. Before Yamnaya, it is likely that Pre-Yamnaya ancestry formed through admixture of EHG-like Khvalynsk with a North Caspian steppe population similar to the Steppe_Eneolithic samples from the North Caucasus Piedmont (see Anthony 2019), so we can also rule out some intermittent presence of a Botai/Kelteminar-like population in the region during the Khvalynsk period.

It is very likely, then, that this competition for the same territory – coupled with the known harsher climate of the late 3rd millennium BC – led Poltavka herders to their known joint venture with Abashevo chiefs in the formation of the Sintashta-Potapovka-Filatovka community of fortified settlements. Supporting these intense contacts of Poltavka herders with Central Asian populations, late ‘outliers’ from the Volga-Ural region show admixture with typical Central_Steppe_MLBA populations: one in Potapovka (ca. 2220 BC), of hg. R1b-Z2103; and four in the Sintashta_MLBA_o1 cluster (ca. 2050-1650 BC), with two samples of hg. R1b-L23 (one R1b-Z2109), one Q1b-L56(xL53), one Q1b-Y6798.

central-steppe-pastoralists
Outlier analysis reveals ancient contacts between sites. We plot the average of principal component 1 (x axis) and principal component 2 (y axis) for the West Eurasian and All Eurasian PCA plots (…). In the Middle to Late Bronze Age Steppe, we observe, in addition to the Western_Steppe_MLBA and Central_Steppe_MLBA clusters (indistinguishable in this projection), outliers admixed with other ancestries. The BMAC-related admixture in Kazakhstan documents northward gene flow onto the Steppe and confirms the Inner Asian Mountain Corridor as a conduit for movement of people.

Similar to how the Sintashta_MLBA_o2 cluster shows an admixture with central steppe populations and hg. R1a-Z645, the WSHG ancestry in those outliers from the o1 cluster of typically (or potentially) Yamnaya lineages show that Poltavka-like herders survived well after centuries of Abashevo-Poltavka coexistence and admixture events, supporting the formation of a Proto-Indo-Iranian community from the local language as pronounced by the incomers, who dominated as elites over the fortified settlements.

The Proto-Indo-Iranian community likely formed thus in situ in the Don-Volga-Ural region, from the admixture of locals of Yamnaya ancestry with incomers of Corded Ware ancestry – represented by the ca. 67% Yamnaya-like ancestry and ca. 33% ancestry from the European cline. Their community formed thus ca. 1,000 years later than the expansion of Late PIE ca. 3500 BC, and expanded (some 500 years after that) a full-fledged Proto-Indo-Iranian language with the Srubna-Andronovo horizon, further admixing with ca. 9% of Central_Steppe_EMBA (WSHG-related) ancestry in their migration through Central Asia, as reported in the paper.

IV. Armenian

The sample from Hajji Firuz, of hg. R1b-Z2103 (xPF331), has been – as expected – re-dated to the Iron Age (ca. 1193-1019 BC), hence it may offer – together with the samples from the Levant and their Aegean-like ancestry rapidly diluted among local populations – yet another proof of how the Late Bronze Age upheaval in Europe was the cause of the Armenian migration to the Armenoid homeland, where they thrived under the strong influence from Hurro-Urartian.

middle-east-armenia-y-dna
Y-chromosome haplogroups of the Middle East and neighbouring groups during the Late Bronze Age / Iron Age. See full maps.

Indus Valley Civilization and Dravidian

A surprise came from the analysis reported by Shinde et al. (2019) of an Iran_N-related IVC ancestry which may have split earlier than 10000 BC from a source common to Iran hunter-gatherers of the Belt Cave.

For the controversial Elamo-Dravidian hypothesis of the Muscovite school, this difference in ancestry between both groups (IVC and Iran Neolithic) seems to be a death blow, if population genomics was even needed for that. Nevertheless, I guess that a full rejection of a recent connection will come down to more recent and subtle population movements in the area.

EDIT (12 SEP): Apparently, Iosif Lazaridis is not so sure about this deep splitting of ‘lineages’ as shown in the paper, so we may be talking about different contributions of AME+ANE/ENA, which means the Elamo-Dravidian game is afoot; at least in genomics:

I shared the idea that the Indus Valley Civilization was linked to the Proto-Dravidian community, so I’m inclined to support this statement by Narasimhan, Patterson, et al. (2019), even if based only on modern samples and a few ancient ones:

The strong correlation between ASI ancestry and present-day Dravidian languages suggests that the ASI, which we have shown formed as groups with ancestry typical of the Indus Periphery Cline moved south and east after the decline of the IVC to mix with groups with more AASI ancestry, most likely spoke an early Dravidian language.

india-steppe-indus-valley-andamanese-ancestry
Natural neighbour interpolation of qpAdm results – Maximum A Posteriori Estimate from the Hierarchical Model (estimates used in the Narasimhan, Patterson et al. 2019 figures) for Central_Steppe_MLBA-related (left), Indus_Periphery_West-related (center) and Andamanese_Hunter-Gatherer-related ancestry (right) among sampled modern Indian populations. In blue, peoples of IE language; in red, Dravidian; in pink, Tibeto-Burman; in black, unclassified. See full image.

I am wary of this sort of simplistic correlation with modern speakers, because we have seen what happened with the wrong assumptions about modern Balto-Slavic and Finno-Ugric speakers and their genetic profile (see e.g. here or here). In fact, I just can’t differentiate as well as those with deep knowledge in South Asian history the social stratification of the different tribal groups – with their endogamous rules under the varna and jati systems – in the ancestry maps of modern India. The pattern of ancestry and language distribution combined with the findings of ancient populations seem in principle straightforward, though.

Conclusion

The message to take home from Shinde et al. (2019) is that genomic data is fully at odds with the Anatolian homeland hypothesis – including the latest model by Heggarty (2014)* – whose relevance is still overvalued today, probably due in part to the shift of OIT proponents to more reasonable Out-of-Iran models, apparently more fashionable as a vector of Indo-Aryan languages than Eurasian steppe pastoralists?
*The authors listed this model erroneously as Heggarty (2019).

The paper seems to play with the occasional reference to Corded Ware as a vector of expansion of Indo-European languages, even after accepting the role of Yamnaya as the most evident population expanding Late PIE to western Europe – and the different ancestry that spread with Indo-Iranian to South Asia 1,000 years later. However, the most cringe-worthy aspect is the sole citation of the debunked, pseudoscientific glottochronological method used by Ringe, Warnow, and Taylor (2002) to support the so-called “steppe homeland”, a paper and dialectal scheme which keeps being referenced in papers of the Reich Lab, probably as a consequence of its use in Anthony (2007).

On the other hand, these are the equivalent simplistic comments in Narasimhan, Patterson et al. (2019):

The Steppe ancestry in South Asia has the same profile as that in Bronze Age Eastern Europe, tracking a movement of people that affected both regions and that likely spread the unique features shared between Indo-Iranian and Balto-Slavic languages. (…), which despite their vast geographic separation share the “satem” innovation and “ruki” sound laws.

mallory-adams-tree
Indo-European dialectal relationships, from Mallory and Adams (2006).

The only academic closely related to linguistics from the list of authors, as far as I know, is James P. Mallory, who has supported a North-West Indo-European dialect (including Balto-Slavic) for a long time – recently associating its expansion with Bell Beakers – opposed thus to a Graeco-Aryan group which shared certain innovations, “Satemization” not being one of them. Not that anyone needs to be a linguist to dismiss any similarities between Balto-Slavic and Indo-Iranian beyond this phonetic trend, mind you.

Even Anthony (2019) supports now R1b-rich Pre-Yamnaya and Yamnaya communities from the Don-Volga region expanding Middle and Late Proto-Indo-European dialects.

So how does the underlying Corded Ware ancestry of eastern Europe (where Pre-Balto-Slavs eventually spread to from Bell Beaker-derived groups) and of the highly admixed (“cosmopolitan”, according to the authors) Sintashta-Potapovka-Filatovka in the east relate to the similar-but-different phonetic trends of two unrelated IE dialects?

If only there was a language substrate that could (as Shinde et al. put it) “elegantly” explain this similar phonetic evolution, solving at the same time the question of the expansion of Uralic languages and their strong linguistic contacts with steppe peoples. Say, Eneolithic populations of mainly hunter-fisher-gatherers from the North Pontic forest-steppes with a stronger connection to metalworking

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

Scytho-Siberians of Aldy-Bel and Sagly, of haplogroup R1a-Z93, Q1b-L54, and N

iron-age-sakas-aldy-bel-scythians

Recently, a paper described Eastern Scythian groups as “Uralic-Altaic” just because of the appearance of haplogroup N in two Pazyryk samples.

This simplistic identification is contested by the varied haplogroups found in early Altaic groups, by the early link of Cimmerians with the expansion of hg. N and Q, by the link of N1c-L392 in north-eastern Europe with Palaeo-Laplandic, and now (paradoxically) by the clear link between early Mongolic expansion and N1c-L392 subclades.

A new paper (behind paywall) offers insight into the prevalent presence of R1a-Z93 among eastern Scytho-Siberian groups (most likely including Samoyedic speakers in the forest-steppes), and a new hint to the westward expansion of haplogroups Q and N (probably coupled with the so-called “Siberian ancestry”) from the east with different groups of Iron Age steppe nomads:

Genetic kinship and admixture in Iron Age Scytho-Siberians, by Mary et al. Human Genetics (2019).

Interesting excerpts (emphasis mine):

From an archeological and historical point of view, the term “Scythians” refers to Iron Age nomadic or seminomadic populations characterized by the presence of three types of artifacts in male burials: typical weapons, specific horse harnesses and items decorated in the so-called “Animal Style”. This complex of goods has been termed the “Scythian triad” and was considered to be characteristic of nomadic groups belonging to the “Scythian World” (Yablonsky 2001). This “Scythian World” includes both the Classic (or European) Scythians from the North Pontic region (7th–3th century BC) and the Southern Siberian (or Asian) populations of the Scythian period (also called Scytho-Siberians). These include, among others, the Sakas from Kazakhstan, the Tagar population from the Minusinsk Basin (Republic of Khakassia), the Aldy-Bel population from Tuva (Russian Federation) and the Pazyryk and Sagly cultures from the Altai Mountains.

mtdna-scytho-siberians
Proportions of Scythian mtDNA haplogroups. Western (blue) and eastern (pink) Eurasian lineages are equally distributed in the Arzhan Scytho-Siberian sample. The U5a2a1 haplogroup shared between the two Scythian groups studied is in bold

In this work, we first aim to address the question of the familial and social organization of Scytho-Siberian groups by studying the genetic relationship of 29 individuals from the Aldy-Bel and Sagly cultures using autosomal STRs. (…) were obtained from 5 archeological sites located in the valley of the Eerbek river in Tuva Republic, Russia (Fig. 1). All the mounds of this archeological site were excavated but DNA samples were not collected from all of them. 14C dates mainly fall within the Hallstatt radiocarbon calibration plateau (ca. 800–400 cal BC) where the chronological resolution is poor. Only one date falls on an earlier segment of calibration curve: Le 9817–2650 ± 25 BP, i.e. 843–792 cal BC with a probability of 94.3% (using the OxCal v4.3.2 program). This sample (Bai-Dag 8, Kurgan 1, grave 10) is not from one of the graves studied but was used to date the kurgan as a whole.

Y-chromosome haplogroups were first assigned using the ISOGG 2018 nomenclature. In order to improve the precision of haplogroup definition, we also analyzed a set of Y-chromosome SNP (Supplementary Table 2). Nine samples belonged to the R1a-M513 haplogroup (defined by marker M513) and two of these nine samples were characterized as belonging to the R1a1a1b2-Z93 haplogroup or one of its subclades. Six samples belonged to the Q1b1a-L54 haplogroup and five of these six samples belonged to the Q1b1a3-L330 subclade. One sample belonged to the N-M231 haplogroup.

haplogroups-scythian-siberians

The distribution of these haplogroups in the population must be confronted with the prevalence of kinship among the samples. Although five individuals belonged to haplogroup Q1b1a3-L330, three of them (ARZ-T18, ARZ-T19 and ARZ-T20) were paternally related (Fig. 2). It must, therefore, be considered that haplogroup Q1b1a3-L330 is present in three independent instances (given that the remaining two instances exhibit no close familial relationship with other samples or one another). All five were buried on the Eki-Ottug 1 archaeological site (although in two different kurgans).

In the same way, although two groups, of two and three individuals, shared haplotypes belonging to the R1a-M513 haplogroup, these groups likely include a father/son pair (ARZ-T2 and ARZ-T12). Therefore, among nine R1a-M513 men, we found six independent haplotypes, one being present in two independent instances. All R1a-M513 haplotypes, however, including those attributed to the R1a1a1b2-Z93 subclade, only differed by one-step mutations, across 5 loci at most. All R1a-M513 individuals were buried on the same site, Eki-Ottug 2, in a single Kurgan.

y-haplogroups-r1a-n-q1b

Haplogroup R1a-M173 was previously reported for 6 Scytho-Siberian individuals from the Tagar culture (Keyser et al. 2009) and one Altaian Scytho-Siberian from the Sebÿstei site (Ricaut et al. 2004a), whereas haplogroup R1a1a1b2-Z93 (or R1a1a1b-S224) was described for one Scythian from Samara (Mathieson et al. 2015) and two Scytho-Siberians from Berel and the Tuva Republic (Unterländer et al. 2017). On the contrary, North Pontic Scythians were found to belong to the R1b1a1a2 haplogroup (Krzewińska et al. 2018), showing a distinction between the two groups of Scythians. (…) The absence of R1b lineages in the Scytho-Siberian individuals tested so far and their presence in the North Pontic Scythians suggest that these 2 groups had a completely different paternal lineage makeup with nearly no gene flow from male carriers between them.

The seven other male individuals studied in this work were found to carry Eastern Eurasian Y haplogroups Q1b1a and one of its subclades (n = 6) and N (n = 1). Haplogroup Q1b1a-L54 was previously described in four males from the Bronze Age in the Altai Mountains (Hollard et al. 2014, 2018) and was clearly associated with Siberian populations (Regueiro et al. 2013).

The N-M231 haplogroup emerged from haplogroup K in Southern Asia around 21,000 years BCE, maybe in Southern China (Shi et al. 2013; Ilumäe et al. 2016). Previous studies attested to its presence in samples from Neolithic and Bronze Age in China (Li et al. 2011; Cui et al. 2013). Waves of northwestern expansion of this haplogroup are described as beginning during the Paleolithic period (Derenko et al. 2006; Shi et al. 2013) but traces of this expansion in archeological samples were reported only in two Scytho-Siberian males from the Altai (Pilipenko et al. 2015).

The sample of haplogroup N comes from the Aldy-Bel culture (ARZ-T15), from the Eerbek site, but has no radiocarbon date. All Q1b-L330 samples come from the Sagly culture, and three are paternally related. The other Q1b-L54 sample is from other tombs in one kurgan at Aldy Bel.

It seems that – exactly as expected – different waves of steppe nomads brought different lineages at a time (the Iron Age) when many regions incorporated different eastern lineages without necessarily changing language. Just like the expansion of N among Ugrians and Samoyeds, and N1c among Finno-Permic peoples, and like many other lineages expanding with federation-like groups in eastern, central, and western Europe

Related

R1a-Z280 and R1a-Z93 shared by ancient Finno-Ugric populations; N1c-Tat expanded with Micro-Altaic

Two important papers have appeared regarding the supposed link of Uralians with haplogroup N.

Avars of haplogroup N1c-Tat

Preprint Genetic insights into the social organisation of the Avar period elite in the 7th century AD Carpathian Basin, by Csáky et al. bioRxiv (2019).

Interesting excerpts (emphasis mine):

After 568 AD the Avars settled in the Carpathian Basin and founded the Avar Qaganate that was an important power in Central Europe until the 9th century. Part of the Avar society was probably of Asian origin, however the localisation of their homeland is hampered by the scarcity of historical and archaeological data.

Here, we study mitogenome and Y chromosomal STR variability of twenty-six individuals, a number of them representing a well-characterised elite group buried at the centre of the Carpathian Basin more than a century after the Avar conquest.

The Y-STR analyses of 17 males give evidence on a surprisingly homogeneous Y chromosomal composition. Y chromosomal STR profiles of 14 males could be assigned to haplogroup N-Tat (also N1a1-M46). N-Tat haplotype I was found in four males from Kunpeszér with identical alleles on at least nine loci. The full Y-STR haplotype I, reconstructed from AC17 with 17 detected STRs, is rare in our days. Only nine matches were found among haplotypes in YHRD database, such as samples from the Ural Region, Northern Europe (Estonia, Finland), and Western Alaska (Yupiks). We performed Median Joining (MJ) network analysis using N-Tat haplotypes with ten shared STR loci (Fig. 3, Table S9). All modern N-Tat samples included in the network had derived allele of L708 as well. Haplotype I (Cluster 1 in Fig. 3) is shared by eight populations on the MJ network among the 24 identical haplotypes. Cluster 1 represents the founding lineage, as it is described in Siberian populations, because this haplotype is shared by the most populations and it is more diverse than Cluster 2.

Nine males share N-Tat haplotype II (on a minimum of eight detected alleles), all of them buried in the Danube-Tisza Interfluve. We found 30 direct matches of this N-Tat haplotype II in the YHRD database, using the complete 17 STR Y-filer profile of AC1, AC12, AC14, AC15, AC19 samples. Most hits came from Mongolia (seven Buryats and one Khalkh) and from Russia (six Yakuts), but identical haplotypes also occur in China (five in Xinjiang and four in Inner Mongolia provinces). On the MJ network, this haplotype II is represented by Cluster 2 and is composed of 45 samples (including 32 Buryats) from six populations (Fig. 3).

y-str-haplogroup-n-mongolian-ugrians
Median Joining network of 162 N-Tat Y-STR haplotypes Allelic information of ten Y-STR loci were used for the network. Only those Avar samples were included, which had results for these ten Y-STR loci. The founder haplotype I (Cluster 1) is shared by eight populations including three Mongolian, three Székely, three northern Mansi, two southern Mansi, two Hungarian, eight Khanty, one Finn and two Avar (AC17, AC26) chromosomes. Haplotype II (Cluster 2) includes 45 haplotypes from six populations studied: 32 Buryats, two Mongolians, one Székely, one Uzbek, one Uzbek Madjar, two northern Mansi and six Avars (AC1, AC12, AC14, AC15, AC19 and KSZ 37). Haplotype III (indicated by a red arrow) is AC8. Information on the modern reference samples is seen in Table S9.

A third N-Tat lineage (type III) was represented only once in the Avar dataset (AC8), and has no direct modern parallels from the YHRD database. This haplotype on the MJ network (see red arrow in Fig. 3) seems to be a descendent from other haplotype cluster that is shared by three populations (two Buryat from Mongolia, three Khanty and one Northern Mansi samples). This haplotype cluster also differs one molecular step (locus DYS393) from haplotype II. We classified the Avar samples to downstream subgroup N-F4205 within the N-Tat haplogroup, based on the results of ours and Ilumäe et al.18 and constructed a second network (Fig. S4). The N-F4205 network results support the assumption that the N-Tat Avar samples belong to N-F4205 subgroup (see SI chapter 1d for more details).

Based on our calculation, the age of accumulated STR variance (TMRCA) within N-Tat lineage for all samples is 7.0 kya (95% CI: 4.9 – 9.2 kya), considering the core haplotype (Cluster 1) to be the founding lineage. Y haplogroup N-Tat was not detected by large scale Eurasian ancient DNA studies but it occurs in late Bronze Age Inner Mongolia and late medieval Yakuts, among them N-Tat has still the highest frequency.

Two males (AC4 and AC7) from the Transtisza group belong to two different haplotypes of Y-haplogroup Q1. Both Q1a-F1096 and Q1b-M346 haplotypes have neither direct nor one step neighbour matches in the worldwide YHRD database. A network of the Q1b-M346 haplotype shows that this male had a probable Altaian or South Siberian paternal genetic origin.

EDIT (5 APR 2019): The paper offers an interesting late sample before the arrival of Hungarian conquerors, although we don’t know which precise lineage the sample belongs to:

One sample in our dataset (HC9) comes from this population, and both his mtDNA (T1a1b) and Y chromosome (R1a) support Eastern European connections. (…) Furthermore, we excluded sample HC9 from population-genetic statistical analyses because it belongs to a later period (end of 7th – early 9th centuries)

Apparently, then, results are consistent with what was already known from studies of modern populations:

According to Ilumäe et al. study, the frequency peak of N-F4205 (N3a5-F4205) chromosomes is close to the Transbaikal region of Southern Siberia and Mongolia, and we conclude that most Avar N-Tat chromosomes probably originated from a common source population of people living in this area, completely in line with the results of Ilumäe et al.

haplogroup_n1
Geographic-Distribution Map of hg N3 from Ilumäe et al.

Finno-Ugrians share haplogroup R1a-Z280

Another paper, behind paywall, Genetic history of Bashkirian Mari and Southern Mansi ethnic groups in the Ural region, by Dudás et al. Molecular Genetics and Genomics (2019).

Interesting excerpts (emphasis mine):

Y‑chromosome diversity

The most frequent haplogroups of the Bashkirian Maris were N1b-P43 (42%), R1a-Z280 (16%), R1a-Z93 (16%), N1c-Tat (13%), and J2-M172 (7%). Furthermore, subgroup R1b-M343 accounted for 4% and I2a-P37 covered 2% of the lineages. None of the Mari N1c Y chromosomes belonged to the N1c subgroups investigated (L1034, VL29, Z1936).

In the case of the Southern Mansi males, the most frequent haplogroups were N1b-P43 (33%), N1c-L1034 (28%) and R1a-Z280 (19%). The frequencies of the remaining haplogroups were as follows: R1a-M458 (6%), I1-L22 (3%), I2a-P37 (3%), and R1b-P312 (3%). The haplotype and haplogroup diversities of the Bashkirian Mari group were 0.9929 and 0.7657, whereas these values for the Southern Mansi were 0.9984 and 0.7873, respectively. The results show that, in both populations, haplotypes are much more diverse than haplogroups.

bashkir-mari-southern-mansi
Haplogroup frequencies of the Bashkirian Mari and the Southern Mansi ethnic groups in Ural region

Genetic structure

(..) the studied Bashkirian Mari and Southern Mansi population groups formed a compact cluster along with two Khanty, Northern Mansi, Mari, and Estonian populations based on close Fst-genetic distances (< 0.05), with nonsignificant p values (p > 0.05) except for the Estonian population. All of these populations belong to the Finno-Ugric language family. Interestingly, the other Mansi population studied by Pimenoff et al. (2008) (pop # 38) was located a great distance from the Southern Mansi group (0.268). In addition, the Bashkir population (pop # 6) did not show a close genetic affinity to the Bashkirian Mari group (0.194), even though it is the host population. However, the Russian population from the Eastern European region of Russia (pop # 49) showed a genetic distance of 0.055 with the Southern Mansi group. All Hungarian speaking populations (pops 13, 22, 23, 24, 50, and 51) showed close genetic affinities to each other and to the neighbouring populations, but not to the two studied populations.

y-dna-hungarians-ugric-mansi
Multidimensional scaling (MDS) plot constructed on Fstgenetic distances of Y haplogroup frequencies of 63 populations compared. The haplogroup frequency data used for population comparison together with references are seen in Online Resource 2 (ESM_2). Pairwise Fst-genetic distances and p values between 63 populations were calculated as shown in Online Resource 3 (ESM_3) Fig. 4 Multidimensional scaling (MDS) plot constructed on Rstgenetic distances of 10 STR-based Y haplotype frequencies of 21 populations compared. Image modified to include labels of modern populations.

Phylogenetic analysis

Median-joining networks were constructed for:

N-P43 (earlier N1b):

(…) TMRCA estimates for this haplogroup were made for all P43 samples (n = 157) 8.7 kya (95% CI 6.7–10.8 kya), for the N-P43 Asian.

N1c-Tat:

(…) 75% of Buryats belonged to Haplotype 2, indicating that the Buryats studied by us is a young and isolated population (Bíró et al. 2015). Bashkirian Mari samples derive from Haplotype 2 via Haplotype 3 (see dark purple circles on the top of Fig. 6a). Haplotype 3 contained six males (2 Buryat, 1 Northern Mansi, and 3 Khanty samples from Pimenoff et al. 2008). The biggest Bashkirian Mari haplotype node (3 Mari samples) was positioned three mutational steps away from Haplotype 1 and the remaining Mari samples can be derived from this haplotype. Southern Mansi haplotypes were scattered within the network except for two, which formed a smaller haplotype node with two Northern Mansi and two Khanty samples from Pimenoff et al. (2008).

n1c-n-tat-uralic-ugric
Median-Joining Networks (MJ) of 153 N-Tat (a) and 26 N-L1034 (b) haplotypes constructed. The circle sizes are proportional to the haplotype frequencies. The smallest area is equivalent to one individual. For N-Tat network, we used data from Southern Mansi (n = 11), Bashkirian Mari (n = 6) samples with Hungarian (n = 12), Hungarian speaking Székely (n = 6), Northern Mansi (n = 14), Mongolian (n = 16), Buryat (n = 44), Finnish (n = 13), Uzbek Madjar (n = 2), Uzbek (n = 3), Khanty (n = 4) populations studied earlier by us (Fehér et al. 2015; Bíró et al. 2015) and Khanty (n = 18) and Mansi (n = 4) studied by Pimenoff et al. (2008)

R1a-Z280 haplotypes, shared by Maris, Mansis, and Hungarians, hence ancient Finno-Ugrians:

The founder R1a-Z280 haplotype was shared by four samples from four populations (1 Bashkirian Mari; 1 Southern Mansi; 1 Hungarian speaking Székely; and 1 Hungarian), as presented in Fig. 7 (Haplotype 1). Haplotype 2 included five males (3 Bashkirian Mari and 2 Hungarian), as it can be seen in Fig. 7. Haplotype 4 included two shared haplotypes (1 Bashkirian Mari and one Hungarian speaking Csángó). The remaining two Bashkirian Mari haplotypes differ from the founder haplotype (Haplotype 1) by two mutational steps via Hungarian or Hungarian and Bashkirian Mari shared haplotypes. Beside Haplotype 1, the remaining Southern Mansi haplotypes were shared with Hungarians (Haplotype 5 or turquoise blue and red-coloured circles above Haplotype 7) or with Hungarians and Hungarian speaking Székely group (Haplotypes 3, 5, and 6). Haplotype 7 included ten Hungarian speakers (Hungarian, Székely, and Csángó). One Hungarian and one Uzbek Khwarezm shared haplotype can be found in Fig. 7 as well (red and white-coloured circle). All the other haplotypes were scattered in the network. The age of accumulated STR variation within R1a-Z280 lineage for 93 samples is estimated to be 9.4 kya (95% CI 6.5–12.4 kya) considering Haplotype 1 (Fig. 7) to be the founder.

r1a-z280-ugrians
Median-Joining Networks (MJ) of 93 R1a-Z280 haplotypes constructed. The circle sizes are proportional to the haplotype frequencies. The smallest area is equivalent to one individual. We used haplotype data from Bashkirian Mari (n = 7), Southern Mansi (n = 7), Hungarian (n = 52), Hungarian speaking Székely (n = 11), Hungarian speaking Csángó (n = 10), Uzbek Ferghana (n = 2), Uzbek Tashkent (n = 1), Uzbek Khwarezm (n = 1) and Northern Mansi (n = 2) populations

R1a-Z93 as isolated lineages among Permic and Ugric populations:

Figure 8 depicts an MJ network of R1a-Z93* samples using 106 haplotypes from the 14 populations (Fig. 8). All of the Bashkirian Mari samples (7 haplotypes) formed a very isolated branch and differed from the one Hungarian haplotype (Fig. 8, see Haplotype 1) by seven mutational steps as well from two Uzbek Tashkent samples (see Haplotype 3). Another Hungarian sample shared two haplotypes of Uzbek Khwarezm samples in Haplotype 4. This haplotype can be derived from Haplotype 3 (Uzbek Tashkent). Haplotype 2 included one Hungarian and one Khakassian male. The remaining three Hungarian haplotypes are outliers in the network and are not shared by any sample. The other population samples included in the network either form independent clusters such as Altaians, Khakassians, Khanties, and Uzbek Madjars or were scattered in the network. The age of accumulated STR variation (TMRCA) within R1a-Z93* lineage for 106 samples is estimated as 11.6 kya (95% CI 9.3–14.0 kya) considering an Armenian haplotype (Fig. 8, “A”) to be the founder and the median haplotype.

r1a-z93-ugrians
Median-Joining Networks (MJ) of 106 R1a-Z93 haplotypes constructed. The circle sizes are proportional to the haplotype frequencies. The smallest area is equivalent to one individual. We used the next haplotype data: 7 Bashkirian Mari, 6 Khanty, 4 Uzbek Madjar, 5 Uzbek Ferghana, 9 Uzbek Tashkent, 7 Uzbek Khwarezm, 2 Mongolian, 2 Buryat, 6 Hungarian samples tested by us for this study or published earlier (Bíró et al. 2015) and populations (3 Armenian; 3 Afghan Tajik;
16 Altaian; 24 Khakassian; 12 Kyrgyz) from Underhill et al. (2015)

Comments

The results of modern populations for N (especially N1c) subclades show really wide clusters and ancient TMRCA, consistent with their known ancient and wide distribution in northern and eastern Eurasian groups, and thus with infiltration of different lineages with eastern nomads (and northern Arctic populations) coupled with later bottlenecks, as well as acculturation of groups.

EDIT (2 APR): Interesting is the specific subclade to which ancient Mongolic-speaking Avars belong (information from Yfull) N1c-F4205 (TMRCA ca. 500 BC), subclade of N1c-Y6058 (formed ca. 2800 BC, TMRCA ca. 2800 BC). This branch also gives the “European” branch N1c-CTS10760 (formed ca. 2800 BC, TMRCA ca. 2100 BC), and is subclade of a branch of N1c-L392 (formed ca. 4400 BC, TMRCA ca. 2800 BC). A northern expansion of N1c-L392 is probably represented by its branch N1c-Z1936 (formed ca. 2800, TMRCA ca. 2100 BC), the most likely candidate to appear in the Kola Peninsula in the Bronze Age as the Palaeo-Laplandic population (see here). Read more about potential routes of expansion of haplogroup N.

On the other hand, R1a-Z280 lineages form a tight cluster connecting Permic with Ugric groups, with R1a-Z93 showing early isolation (probably) between Cis-Urals and Trans-Urals regions. While both Corded Ware lineages in Finno-Ugrians are most likely related to the Abashevo expansion through Seima-Turbino and the Andronovo-like Horizon (and potentially later Eurasian expansions), a plausible hypothesis would be that Finno-Ugrians are related to an expansion of R1a-Z283 haplogroups (we already knew about the Finno-Permic connection), while the ancient connection between Permians and Hungarians with R1a-Z93 would correspond to this haplogroup’s potentially tighter link with an early Samoyedic split.

I don’t think that an explosive expansion of eastern Corded Ware groups of R1a-Z645 lineages will show a clear-cut division of haplogroups among Eastern Uralic groups, though, and culturally I doubt we will have such a clear image, either (similar to how the explosive expansion of Bell Beakers cannot be easily divided by regional/language group into R1b-L151 subclades before the known bottlenecks). Relevant in this regard are the known Z93 samples from the Árpád dynasty.

Nevertheless, this data may represent a slightly more recent wave of R1a-Z280 lineages linked to the expansion of Ugric into the Trans-Uralian region, after their split from Finno-Permic, still in close contact with Indo-Iranians in Poltavka and Sintashta-Potapovka, evident from the early and late Indo-Iranian borrowings, during a common period when Samoyedic had already separated.

Such a “Z283 over Z93” layer in the Trans-Urals (and Cis-Urals?) forest-steppes would be similar to the apparent replacement of Z284 by Z282 in the Eastern Baltic during the Bronze Age (possibly with the second or Estonian Battle Axe wave or, much more likely during later population movements). Such an early R1a-Z93 split could potentially be supported also by the separation into bottlenecks under “Northern” (R1a-Z283) Finno-Ugric-speaking Abashevo-related groups and “Southern” (R1a-Z93) acculturated Indo-Iranian-speaking Abashevo migrants developing Sintashta-Potapovka admixing with Poltavka R1b-Z2103 herders.

r1a-z282-z280-z2125-distribution
Modified image, from Underhill et al. (2015). Spatial frequency distributions of Z282 (green) and Z93 (blue) affiliated haplogroups.. Notice the potential Finno-Ugric-associated distribution of Z282 (especially R1a-M558, a Z280 subclade), the expansion of R1a-Z2123 subclades with Central Asian forest-steppe groups.

Conclusion

Let’s review some of the most common myths about Hungarians (and Finno-Ugrians in general) repeated ad nauseam, side by side with my assertions:

❌ N (especially N1c-Tat) in ancient and modern samples represent the True Uralic™ N1c peoples including Magyar tribes? Nope.

✅ Ancient N (especially N1c-Tat) lineages among Uralic populations expanded relatively recently, and differently in different regions (including eastern steppe nomads and northern arctic populations) not associated with a particular language or language group? Yep (read the series on Corded Ware = Uralic expansion).

❌ Modern Hungarian R1a-Z280 lineages represent the majority of the native population, poor Slavic ‘peasants’ from the Carpathian Basin, forcibly acculturated by a minority of bad bad Hungarian hordes? Nope.

✅ Modern Hungarian R1a-Z280 subclades represent Ugric lineages in common with ancient R1a-Z645 Finno-Ugric populations from north-eastern Europe and the Trans-Urals? Yep (see Avars and Ugrians).

❌ Modern Hungarian R1a-Z93 lineages represent acculturated Iranian/Turkic peoples from the steppes? Not likely.

✅ Modern Hungarian R1a-Z93 lineages represent a remnant of the expansion of Corded Ware to the east, potentially more clearly associated with Samoyedic? Much more likely.

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

Sooo, the theory of a “diluted” Y-DNA in Modern Hungarians from originally fully N-dominated conquerors subjugating native R1a-Z280 Slavs from the Carpathian Basin is not backed up by genetic studies? The ethnic Iranian-Turkic R1a-Z93 federation in the steppes that ended up speaking Magyar is not real?? Who would’ve thunk.

Another true story whose rejection in genetics could not be predicted, like, not at all.

Totally unexpected, too, the drift of “R1a=IE” fans with the newest genetic findings towards a Molgen-like “Yamna/R1b = Vasconic-Caucasian”, “N1c = Uralic-Altaic”, and “R1a = the origin of the white world in Mother Russia”. So much for the supposed interest in “Steppe ancestry” and fancy statistics.

Related

Yekaterinovsky Cape, a link between the Samara culture and early Khvalynsk

ekaterinovsky-cape

We already had conflicting information about the elite individual from the Yekaterinovsky Cape and the materials of his grave, which seemed quite old:

For the burial of 45 in the laboratory of the University of Pennsylvania, a 14C date was obtained: PSUAMS-2880 (Sample ID 16068)> 30 kDa gelatin Russia. 12, Ekaterinovka Grave 45 14C age (BP) 6325 ± 25 δ 13C (‰) –23.6 δ15 N (‰) 14.5. The results of dating suggest chronological proximity with typologically close materials from Yasinovatsky and Nikolsky burial grounds (Telegini et al. 2001: 126). The date obtained also precedes the existing dates for the Khvalynsk culture (Morgunova 2009: 14–15), which, given the dominance of Mariupol traits of the burial rite and inventory, confirms its validity. However, the date obtained for human bones does not exclude the possibility of a “reservoir effect” when the age can increase three or more centuries (Shishlin et al. 2006: 135–140).

Now the same date is being confirmed by the latest study published on the site, by Korolev, Kochkina, and Stachenkov (2019) and it seems it is really going to be old. Abstract (in part the official one, in part newly translated for clarity):

For the first time, pottery of the Early Eneolithic burial ground Ekaterinovsky Cape is published. Ceramics were predominantly located on the sacrificial sites in the form of compact clusters of fragments. As a rule, such clusters were located above the burials, sometimes over the burials, some were sprinkled with ocher. The authors have identified more than 70 vessels, some of which have been partially reconstructed. Ceramic was made with inclusion of the crushed shell into molding mass. The rims of vessels had the thickened «collar»; the bottoms had a rounded shape. The ornament was located on the rims and the upper part of the potteries. Fully decorated vessels are rare. The vessels are ornamented with prints of comb and rope stamps, with small pits. A particularity of ceramics ornamentation is presented by the imprints of soft stamps (leather?) or traces of leather form for the making of vessels. The ornamentation, made up of «walking comb» and incised lines, was used rarely as well as the belts of pits made decoration under «collar» of a rim. Some features of the ceramics decoration under study relate it with ceramics of the Khvalynsk culture. The ceramics of Ekaterinovsky Cape burial ground is attributed by the authors to the Samara culture. The ceramic complex under study has proximity to the ceramics from Syezzhe burial ground and the ceramics of the second phase of Samara culture. The chronological position is determined by the authors as a later period than the ceramics from the Syezzhe burial ground, and earlier than the chronological position of ceramics of the Ivanovka stage of the Samara culture and the Khvalynsk culture.

ekaterinovsky-cape-pottery
Ceramics from Ekaterinovsky Cape burial ground. 1–2, 4–5, 7–11 – ceramics from aggregations; 3, 6 – ceramics from the cultural layer.

More specifically:

Based on ceramic fragments from a large vessel from a cluster of sq.m. 14, the date received was: SPb-2251–5673 ± 120 BP. The second date was obtained in fragments from the aggregation [see picture above] from the cluster of sq.m. 45–46: SPb-2252–6372 ± 100 BP. The difference in dating indicates that the process of determining the chronology of the burial ground is far from complete, although we note that the earlier date almost coincided with the date obtained from the human bone from individual 45 (Korolev, Kochkina, Stashenkov, 2018, p. 300).

Therefore, the ceramics of the burial ground Ekaterinovsky Cape possess an originality that determines the chronological position of the burial ground between the earliest materials of the burial type in Syezzhe and the Khvalynsk culture. Techno-typological features of dishes make it possible to attribute it to the Samara culture at the stage preceding the appearance of Ivanovska-Khvalynsk ceramics.

It seems that this site showed cultural influences from the upstream region near the Kama-Vyatka interfluve, too, according to Korolev, Kochkina, Stashenkov, and Khokhlov (2018):

In 2017, excavation of burial ground Ekaterinovsky Cape were continued, located in the area of the confl uence of the Bezenchuk River in the Volga River. During the new excavations, 14 burials were studied. The skeleton of the buried were in a position elongated on the back, less often – crooked on the back with knees bent at the knees. In one burial (No. 90), a special position of the skeleton was recorded. In the burial number 90 in the anatomical order, parts of the male skeleton. This gave grounds for the reconstruction of his original position in a semi-sitting position with the support of elbows on the bottom of the pit. Noteworthy inventory: on the pelvic bones on the left lay a bone spoon, near the right humerus, the pommel of a cruciform club was found. A conclusion is made about the high social status of the buried. The results of the analysis of the burial allow us to outline the closest circle of analogies in the materials of Khvalynsky I and Murzikhinsky burial grounds.

Important sites mentioned in both papers and in this text:

To sum up, it seems that the relative dates we have used until now have to be corrected: older Khvalynsk I Khvalynsk II individuals, supposedly dated ca. 5200-4000 BC (most likely after 4700 BC), and younger Yekaterinovsky individuals, supposedly of the fourth quarter of the 5th millennium (ca. 4250-4000 BC), are possibly to be considered, in fact, roughly reversed, if not chronologically, at least culturally speaking.

Interestingly, this gives a new perspective to the presence of a rare fish- or reptile-headed pommel-scepter, which would be natural in a variable period of expansion of the horse and horse-related symbolism, a cultural trait rooted in the Samara culture attested in Syezzhe before the unification of the symbol of power under the ubiquitous Khvalynsk-Suvorovo horse-headed scepters and related materials.

ekaterinovsky-cape-pommel-mace
Ekaterinovsky Cape Burial Ground. Inventory of the burial no 90: 1, 2 – stone pommel of the mace; 3, 4 – bone article.

The Khvalynsk chieftain

If the reported lineages from Yekaterinovsky Cape are within the R1b-P297 tree, but without further clades, as Yleaf comparisons may suggest, there is not much change to what we have, and R1b-M269 could actually represent a part of the local population, but also incomers from the south (e.g. the north Caspian steppe hunter-gatherers like Kairshak), the east (with hunter-gatherer pottery), or the west near the Don River (in contact with Mariupol-related cultures, as the authors inferred initially from material culture).

Just like R1a-M417 became incorporated into the Sredni Stog groups after the Novodanilovka-Suvorovo expansion, probably as incoming hunter-gatherer pottery groups from the north admixing with peoples of “Steppe ancestry”, R1b-M269 lineages might have expanded explosively only during the Repin expansion, and maybe (like R1b-L51 later) they formed just a tiny part of the clans that dominated the steppe during the Khvalynsk-Novodanilovka community.

On the other hand, the potential finding of various R1b-M269/L23 samples in Yekaterinovsky Cape (including an elite individual) would suggest now, as it was supported in the original report by Mathieson et al. (2015), that these ancient R1b lineages found in the Volga – Ural region are in fact most likely all R1b-M269 without enough coverage to obtain proper SNP calls, which would simplify the picture of Neolithic expansions (yet again). From the supplementary materials:

10122 / SVP35 (grave 12). Male (confirmed genetically), age 20-30, positioned on his back with raised knees, with 293 copper artifacts, mostly beads, amounting to 80% of the copper objects in the combined cemeteries of Khvalynsk I and II. Probably a high-status individual, his Y-chromosome haplotype, R1b1, also characterized the high-status individuals buried under kurgans in later Yamnaya graves in this region, so he could be regarded as a founder of an elite group of patrilineally related families. His MtDNA haplotype H2a1 is unique in the Samara series.

khvalynsk-cemetery
Khvalynsk cemetery and grave gifts. Grave 90 contained copper beads and rings, a harpoon, flint blades, and a bird-bone tube. Both graves (90 and 91) were partly covered by Sacrificial Deposit 4 with the bones from a horse, a sheep, and a cow. Center: grave goods from the Khvalynsk cemetery-copper rings and bracelets, polished stone mace heads, polished stone bracelet, Cardium shell ornaments, boars tusk chest ornaments, flint blades, and bifiacial projectile points. Bottom: shell-tempered pottery from the Khvalynsk cemetery. After Agapov, Vasiliev, and Pestrikova 1990; and Ryndina 1998, Figure 31. Modified from Anthony (2007).

This remarkable Khvalynsk chieftain, whose rich assemblage may correspond to the period of domination of the culture all over the Pontic-Caspian steppes, has been consistently reported as of hg. R1b-L754 in all publications, including Wang et al. (2018/2019) tentative SNP calls in the supplementary materials (obtained with Yleaf, as the infamous Narasimhan et al. 2018 samples), but has been variously reported by amateurs as within the R1b-M73, R1b-V88, or (lately) R1b-V1636 trees, which makes it unlikely that quality of the sample is allowing for a proper SNP call.

The fact that Mathieson et al. (2015) considered it a member of the R1b-M269 clans appearing later in Yamna seems on point right now, especially if samples from Yekaterinovka are all within this tree. The relevance of R1b-L23 in the expansion of Repin and Yamna is reminiscent of the influence of successful clans among Yamna offshoots, such as Bell Beakers, and among Bell Beaker offshoots during the Bronze Age all over Europe.

Taking these younger expansions as example, it seems quite likely based on cultural links that (at least part of) the main clans of Khvalynsk were of R1b-M269 lineage, stemming from a R1b-dominated Samara culture, in line with the known succeeding expansions and the expected strictly patriarcal and patrilineal society of Proto-Indo-Europeans, which would have exacerbated the usual reduction in Y-chromosome haplogroup variability that happens during population expansions, and the aversion towards foreign groups while the culture lasted.

pontic-steppe-neolithic
Cultures of the Pontic-Caspian steppes and forest-steppes and surrounding areas during the Neolithic.

The finding of R1b-L23 in Yekaterinovka, associated with the Samara culture, before or during the Khvalynsk expansion, and close to the Khvalynsk site, would make this Khvalynsk chieftain most likely a member of the M269 tree (paradoxically, the only R1b-L754 branch amateurs have not yet reported for it). Similarly, the sample of a “Samara hunter-gatherer” of Lebyazhinka, of hg. R1b-P297, could also be under this tree, just like most R1b-M269 from Yamna are downstream from R1b-L23, and most reported R1b-M269 or R1b-L23 from Bell Beakers are under R1b-L151.

On the other hand, we know of the shortcomings of attributing a haplogroup expansion to the best known rulers, such as the famous lineages previously wrongly attributed to Niall of the Nine Hostages or Genghis Khan. The known presence of R1b-V1636 up to modern Greeks would be in line with an ancient steppe expansion that we know will show up during the Neolithic, although it could also be a sign of a more recent migration from the Caucasus. The presence of a sister clade of R1b-L23, R1b-PF7562, among modern Balkan populations, may also be attributed to a pre-Yamna steppe expansion.

y-dna-khvalynsk
Y-DNA samples from Khvalynsk and neighbouring cultures. See full version here.

On SNP calls

I reckon that even informal reports on SNP calls, like any other analyses, should be offered in full: not only with a personal or automatic estimation of the result, but with a detailed explanation of the good, dubious, and bad calls, alternatives to that SNP estimation, and a motivated reasoning of why one branch should be preferred over others. Downloading a sample and giving an instruction using a free software tool is never enough, as it became crystal clear recently for the hilariously biased and flawed qpAdm reports on Dutch Bell Beakers as the ‘missing link’ between Corded Ware and Bell Beakers…

Another example I can recall is the report of a R1a-Z93 subclade in the R1a-M417 sample ca. 4000 BC from Alexandria, which seems rather unlikely, seeing how this subclade must have split and expanded explosively with R1a-Z645 to the east with eastern Corded Ware groups, i.e. 1,000 years later, just like Z282 lineages expanded mainly to the north-east. But then again, as with the Khvalynsk chieftain, I have only seen indirect reports of that supposed SNP (including Y26+!), so we should just stick with its officially reported R1a-M417 lineage. This upstream haplogroup was, in fact, repeated with Yleaf’s tentative estimates in Wang et al. (2019) supplementary materials…

The combination of inexperienced, biased, or simply careless design, analyses, and reports, including SNP calls and qpAdm analyses (whether in forums or publications), however well-intentioned (or not) they might be, are hindering a proper analysis of data, adding to the difficulties we already have due to the scarcity of samples, their limited coverage, and the lack of proper context.

Some people like to repeat ad nauseam that archaeology and/or linguistics are ‘not science’ whenever they don’t fit their beliefs and myths based on haplogroup and/or ancestry. But it’s becoming harder and harder to rely on certain genetic data, too, and on their infinite changing interpretations, much more than it is to rely on linguistic and archaeological research, including data, assessments, and discussions that are open for anyone to review…if one is truly interested in them.

Aquitanians and Iberians of haplogroup R1b are exactly like Indo-Iranians and Balto-Slavs of haplogroup R1a

eba-indo-iranian-balto-slavs

The final paper on Indo-Iranian peoples, by Narasimhan and Patterson (see preprint), is soon to be published, according to the first author’s Twitter account.

One of the interesting details of the development of Bronze Age Iberian ethnolinguistic landscape was the making of Proto-Iberian and Proto-Basque communities, which we already knew were going to show R1b-P312 lineages, a haplogroup clearly associated during the Bell Beaker period with expanding North-West Indo-Europeans:

From the Bronze Age (~2200–900 BCE), we increase the available dataset from 7 to 60 individuals and show how ancestry from the Pontic-Caspian steppe (Steppe ancestry) appeared throughout Iberia in this period, albeit with less impact in the south. The earliest evidence is in 14 individuals dated to ~2500–2000 BCE who coexisted with local people without Steppe ancestry. These groups lived in close proximity and admixed to form the Bronze Age population after 2000 BCE with ~40% ancestry from incoming groups. Y-chromosome turnover was even more pronounced, as the lineages common in Copper Age Iberia (I2, G2, and H) were almost completely replaced by one lineage, R1b-M269.

iberia-admixture-y-dna
Proportion of ancestry derived from central European Beaker/Bronze Age populations in Iberians from the Middle Neolithic to the Iron Age (table S15). Colors indicate the Y-chromosome haplogroup for each male. Red lines represent period of admixture. Modified from Olalde et al. (2019).

The arrival of East Bell Beakers speaking Indo-European languages involved, nevertheless, the survival of the two non-IE communities isolated from each other – likely stemming from south-western France and south-eastern Iberia – thanks to a long-lasting process of migration and admixture. There are some common misconceptions about ancient languages in Iberia which may have caused some wrong interpretations of the data in the paper and elsewhere:

NOTE. A simple reading of Iberian prehistory would be enough to correct these. Two recent books on this subject are Villar’s Indoeuropeos, iberos, vascos y otros parientes and Vascos, celtas e indoeuropeos. Genes y lenguas.

Iberian languages were spoken at least in the Mediterranean and the south (ca. “1/3 of Iberia“) during the Bronze Age.

Nope, we only know the approximate location of Iberian culture and inscriptions from the Late Iron Age, and they occupy the south-eastern and eastern coastal areas, but before that it is unclear where they were spoken. In fact, it seems evident now that the arrival of Urnfield groups from the north marks the arrival of Celtic-speaking peoples, as we can infer from the increase in Central European admixture, while the expansion of anthropomorphic stelae from the north-west must have marked the expansion of Lusitanian.

Vasconic was spoken in both sides of the Pyrenees, as it was in the Middle Ages.

Wrong. One of the worst mistakes I am seeing in many comments since the paper was published, although admittedly the paper goes around this problem talking about “Modern Basques”. Vasconic toponyms appear south of the Pyrenees only after the Roman conquests, and tribes of the south-western Pyrenees and Cantabrian regions were likely Celtic-speaking peoples. Aquitanians (north of the western Pyrenees) are the only known ancient Vasconic-speaking population in proto-historic times, ergo the arrival of Bell Beakers in Iberia was most likely accompanied by Indo-European languages which were later replaced by Celtic expanding from Central Europe, and Iberian expanding from south-east Iberia, and only later with Latin and Vasconic.

Ligurian is non-Indo-European, and Lusitanian is Celtic-like, so Iberia must have been mostly non-Indo-European-speaking.

The fragmentary material available on Ligurian is enough to show that phonetically it is a NWIE dialect of non-Celtic, non-Italic nature, much like Lusitanian; that is, unless you follow laryngeals up to Celtic or Italic, in which case you can argue anything about this or any other IE language, as people who reconstruct laryngeals for Baltic in the common era do.

EDIT (19 Mar 2019): It was not clear enough from this paragraph, because Ligurian-like languages in NE Iberia is just a hypothesis based on the archaeological connection of the whole southern France Bell Beaker region. My aim was to repeat the idea that Old European hydro-toponymy is older in NE Iberia (as almost anywhere in Iberia) than Iberian toponymy, so the initial hypothesis is that:

  1. a Palaeo-European language (as Villar puts it) expanded into most regions of Iberia in ancient times (he considered at some point the Mesolithic, but that is obviously wrong, as we know now); then
  2. Celts expanded at least to the Ebro River Basin; then
  3. Iberians expanded to the north and replaced these in NE Iberia; and only then
  4. after the Roman invasion, around the start of the Common Era, appear Vasconic toponyms south of the Pyrenees.

Lusitanian obviously does not qualify as Celtic, lacking the most essential traits that define Celticness…Unless you define “(Para-)Celtic” as Pre-Proto-Celtic-like, or anything of the sort to support some Atlantic continuity, in which case you can also argue that Pre-Italic or Pre-Germanic are Celtic, because you would be essentially describing North-West Indo-European

If Basques have R1b, it’s because of a culture of “matrilocality” as opposed to the “patrilocality” of Indo-Europeans

So wrong it hurts my eyes every time I read this. Not only does matrilocality in a regional group have few known effects in genetics, but there are many well-documented cases of population replacement (with either ancestry or Y-DNA haplogroups, or both) without language replacement, without a need to resort to “matrilineality” or “matrilocality” or any other cultural difference in any of these cases.

In fact, it seems quite likely now that isolated ancient peoples north of the Pyrenees will show a gradual replacement of surviving I2a lineages by neighbouring R1b, while early Iberian R1b-DF27 lineages are associated with Lusitanians, and later incoming R1b-DF27 lineages (apart from other haplogroups) are most likely associated with incoming Celts, which must have remained in north-central and central-east European groups.

NOTE. Notice how R1a is fully absent from all known early Indo-European peoples to date, whether Iberian IE, British IE, Italic, or Greek. The absence of R1a in Iberia after the arrival of Celts is even more telling of the origin of expanding Celts in Central Europe.

I haven’t had enough time to add Iberian samples to my spreadsheet, and hence neither to the ASoSaH texts nor maps/PCAs (and I don’t plan to, because it’s more efficient for me to add both, Asian and Iberian samples, at the same time), but luckily Maciamo has summed it up on Eupedia. Or, graphically depicted in the paper for the southeast:

iberia-haplogroups
Y chromosome haplogroup composition of individuals from southeast Iberia during the past 2000 years. The general Iberian Bronze and Iron Age population is included for comparison. Modified from Olalde et al. (2019).

Does this continued influx of Y-DNA haplogroups in Iberia with different cultures represent permanent changes in language? Are, therefore, modern Iberian languages derived from Lusitanian, Sorothaptic/Celtic, Greek, Phoenician, East or West Germanic, Hebrew, Berber, or Arabic languages? Obviously not. Same with Italy (see the recent preprint on modern Italians by Raveane et al. 2018), with France, with Germany, or with Greece.

If that happens in European regions with a known ancient history, why would the recent expansions and bottlenecks of R1b in modern Basques (or N1c around the Baltic, or R1a in Slavs) in the Middle Ages represent an ancestral language surviving into modern times?

Indo-Iranians

If something is clear from Narasimhan, Patterson, et al. (2018), is that we know finally the timing of the introduction and expansion of R1a-Z645 lineages among Indo-Iranians.

We could already propose since 2015 that a slow admixture happened in the steppes, based on archaeological finds, due to settlement elites dominating over common peoples, coupled with the known Uralic linguistic traits of Indo-Iranian (and known Indo-Iranian influence on Finno-Ugric) – as I did in the first version of the Indo-European demic diffusion model.

The new huge sampling of Sintashta – combined with that of Catacomb, Poltavka, Potapovka, Andronovo, and Srubna – shows quite clearly how this long-term admixture process between Uralic peoples and Indo-Iranians happened between forest-steppe CWC (mainly Abashevo) and steppe groups. The situation is not different from that of Iberia ca. 2500-2000 BC; from Narasimhan, Patterson, et al. (2018):

We combined the newly reported data from Kamennyi Ambar 5 with previously reported data from the Sintashta 5 individuals (10). We observed a main cluster of Sintashta individuals that was similar to Srubnaya, Potapovka, and Andronovo in being well modeled as a mixture of Yamnaya-related and Anatolian Neolithic (European agriculturalist-related) ancestry.

Even with such few words referring to one of the most important data in the paper about what happened in the steppes, Wang et al. (2018) help us understand what really happened with this simplistic concept of “steppe ancestry” regarding Yamna vs. Corded Ware differences:

anatolia-neolithic-steppe-eneolithic
Image modified from Wang et al. (2018). Marked are: 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 1128 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 (see also Supplementary Tables 10, 14 and 20).”

As with Iberia (or any prehistoric region), the details of how exactly this language change happened are not evident, but we only need a plausible explanation coupled with archaeology and linguistics. Poltavka, Potapovka, and Sintashta samples – like the few available Iberian ones ca. 2500-2000 BC – offer a good picture of the cohabitation of R1b-L23 (mainly Z2103) and R1a-Z645 (mainly Z93+): a glimpse at the likely presence of R1a-Z93 within settlements – which must have evolved as the dominant elites – in a society where the majority of the population was initially formed by nomad herders (probably most R1b-Z2103), who were usually buried outside of the main settlements.

Will the upcoming Narasimhan, Patterson et al. (2019) deal with this problem of how R1a-M417 replaced R1b-M269, and how the so-called “Steppe_MLBA” (i.e. Corded Ware) ancestry admixed with “Steppe_EMBA” (i.e. Yamnaya) ancestry in the steppes, and which one of their languages survived in the region (that is, the same the Reich Lab has done with Iberia)? Not likely. The ‘genetic wars’ in Iberia deal with haplogroup R1b-P312, and how it was neither ‘native’ nor associated with Basques and non-Indo-European peoples in general. The ‘genetic wars’ in South Asia are concerned with the steppe origin of R1a, to prove that it is not a ‘native’ haplogroup to India, and thus neither are Indo-Aryan languages. To each region a politically correct account of genetic finds, with enough care not to fully dismiss national myths, it seems.

NOTE. Funnily enough, these ‘genetic wars’ are the making of geneticists since the 1990s and 2000s, so we are still in the midst of mostly internal wars caused by what they write. Just as genetic papers of the 2020s will most likely be a reaction to what they are writing right now about “steppe ancestry” and R1a. You won’t find much change to the linguistic reconstruction in this whole period, except for the most multicolored glottochronological proposals…

The first author of the paper has engaged, as far as I could see in Twitter, in dialogue with Hindu nationalists who try to dismiss the arrival of steppe ancestry and R1a into South Asia as inconclusive (to support the potential origin of Sanskrit millennia ago in the Indus Valley Civilization). How can geneticists deal with the real problem here (the original ethnolinguistic group expanding with Corded Ware), when they have to fend off anti-steppists from Europe and Asia? How can they do it, when they themselves are part of the same societies that demand a politically correct presentation of data?

This is how the data on the most likely Indo-Iranian-speaking region should be presented in an ideal world, where – as in the Iberia paper – geneticists would look closely to the Volga-Ural region to discover what happened with Proto-Indo-Iranians from their earliest to their latest stage, instead of constantly looking for sites close to the Indus Valley to demonstrate who knows what about modern Indian culture:

indo-iranian-admixture-similar-iberians
Tentative map of the Late PIE and Indo-Iranian community in the Volga-Ural steppes since the Eneolithic. Proportion of ancestry derived from central European Corded Ware peoples. Colors indicate the Y-chromosome haplogroup for each male. Red lines represent period of admixture. Modified from Olalde et al. (2019).

Now try and tell Hindu nationalists that Sanskrit expanded from an Early Bronze Age steppe community of R1b-rich nomadic herders that spoke Pre-Indo-Iranian, which was dominated and eventually (genetically) mostly replaced by elite Uralic-speaking R1a peoples from the Russian forest, hence the known phonetic (and some morphological) traits that remained. Good luck with the Europhobic shitstorm ahead..

Balto-Slavic

Iberian cultures, already with a majority of R1b lineages, show a clear northward expansion over previously Urnfield-like groups of north-east Iberia and Mediterranean France (which we now know probably represent the migration of Celts from central Europe). Similarly, Eastern Balts already under a majority of R1a lineages expanded likely into the Baltic region at the same time as the outlier from Turlojiškė (ca. 1075 BC), which represents the first obvious contacts of central-east Europe with the Baltic.

Iberia shows a more recent influx of central and eastern Mediterranean peoples, one of which eventually succeeded in imposing their language in Western Europe: Romans were possibly associated mainly with R1b-U152, apart from many other lineages. Proto-Slavs probably expanded later than Celts, too, connected to the disintegration of the Lusatian culture, and they were at some point associated with R1a-M458 and R1a-Z280(xZ92) lineages, apart from others already found in Early Slavs.

pca-balto-slavs-tollense-valley
PCA of central-eastern European groups which may have formed the Balto-Slavic-speaking community derived from Bell Beaker, evident from the position ‘westwards’ of CWC in the PCA, and surrounding cultures. Left: Early Bronze Age. Right: Tollense Valley samples.

This parallel between Iberia and eastern Europe is no coincidence: as Europe entered the Bronze Age, chiefdom-based systems became common, and thus the connection of ancestry or haplogroups with ethnolinguistic groups became weaker.

What happened earlier (and who may represent the Pre-Balto-Slavic community) will be clearer when we have enough eastern European samples, but basically we will be able to depict this admixture of NWIE-speaking BBC-derived peoples with Uralic-speaking CWC-derived groups (since Uralic is known to have strongly influenced Balto-Slavic), similar to the admixture found in Indo-Iranians, more or less like this:

iberian-admixture-balto-slavic
Tentative map of the North-West Indo-European and Balto-Slavic community in central-eastern Europe since the East Bell Beaker expansion. Proportion of ancestry derived from Corded Ware peoples. Colors indicate the Y-chromosome haplogroup for each male. Red lines represent period of admixture. Modified from Olalde et al. (2019).

The Early Scythian period marked a still stronger chiefdom-based system which promoted the creation of alliances and federation-like groups, with an earlier representation of the system expanding from north-eastern Europe around the Baltic Sea, precisely during the spread of Akozino warrior-traders (in turn related to the Scythian influence in the forest-steppes), who are the most likely ancestors of most N1c-V29 lineages among modern Germanic, Balto-Slavic, and Volga-Finnic peoples.

Modern haplogroup+language = ancient ones?

It is not difficult to realize, then, that the complex modern genetic picture in Eastern Europe and around the Urals, and also in South Asia (like that of the Aegean or Anatolia) is similar to the Iron Age / medieval Iberian one, and that following modern R1a as an Indo-European marker just because some modern Indo-European-speaking groups showed it was always a flawed methodology; as flawed as following R1b for ancient Vasconic groups, or N1c for ancient Uralic groups.

Why people would argue that haplogroups mean continuity (e.g. R1b with Basques, N1c with Finns, R1a with Slavs, etc.) may be understood, if one lives still in the 2000s. Just like why one would argue that Corded Ware is Indo-European, because of Gimbutas’ huge influence since the 1960s with her myth of “Kurgan peoples”. Not many denied these haplogroup associations, because there was no reason to do it, and those who did usually aligned with a defense of descriptive archaeology.

However, it is a growing paradox that some people interested in genetics today would now, after the Iberian paper, need to:

  • accept that ancient Iberians and probably Aquitanians (each from different regions, and probably from different “Basque-Iberian dialects” in the Chalcolithic, if both were actually related) show eventually expansions with R1b-L23, the haplogroup most obviously associated with expanding Indo-Europeans;
  • acknowledge that modern Iberians have many different lineages derived from prehistoric or historic peoples (Celts, Phoenicians, Greeks, Romans, Jews, Goths, Berbers, Arabs), which have undergone different bottlenecks, the last ones during the Reconquista, but none of their languages have survived;
  • realize that a similar picture is to be found everywhere in central and western Europe since the first proto-historic records, with language replacement in spite of genetic continuity, such as the British Isles (and R1b-L21 continuity) after the arrival of Celts, Romans, Anglo-Saxons, Vikings, or Normans;
  • but, at the same time, continue blindly asserting that haplogroup R1a + “steppe ancestry” represent some kind of supernatural combination which must show continuity with their modern Indo-Iranian or Balto-Slavic language from time immemorial.
sintashta-y-dna
Replacement of R1b-L23 lineages during the Early Bronze Age in eastern Europe and in the Eurasian steppes: emergence of R1a in previous Yamnaya and Bell Beaker territories. Modified from EBA Y-DNA map.

Behave, pretty please

The ‘conservative’ message espoused by some geneticists and amateur genealogists here is basically as follows:

  • Let’s not rush to new theories that contradict the 2000s, lest some people get offended by granddaddy not being these pure whatever wherever as they believed, and let’s wait some 5, 10, or 20 years, as long as necessary – to see if some corner of the Yamna culture shows R1a, or some region in north-eastern Europe shows N1c, or some Atlantic Chalcolithic sample shows R1b – to challenge our preferred theories, if we actually need to challenge anything at all, because it hurts too much.
  • Just don’t let many of these genetic genealogists or academics of our time be unhappy, pretty please with sugar on top, and let them slowly adapt to reality with more and more pet theories to fit everything together (past theories + present data), so maybe when all of them are gone, within 50 or 70 years, society can smoothly begin to move on and propose something closer to reality, but always as politically correct as possible for the next generations.
  • For starters, let’s discuss now (yet again) that Bell Beakers may not have been Indo-European at all, despite showing (unlike Corded Ware) clearly Yamna male lineages and ancestry, because then Corded Ware and R1a could not have been Indo-European and that’s terrible, so maybe Bell Beakers are too brachycephalic to speak Indo-European or something, or they were stopped by the Fearsome Tisza River, or they are not pure Dutch Single Grave in The South hence not Indo-European, or whatever, and that’s why Iron Age Iberians or Etruscans show non-Indo-European languages. That’s not disrespectful to the history of certain peoples, of course not, but talking about the evident R1a-Uralic connection is, because this is The South, not The North, and respect works differently there.
  • Just don’t talk about how Slavs and Balts enter history more than 1,500 years later than Indo-European peoples in Western and Southern Europe, including Iberia, and assume a heroic continuity of Balts and Slavs as pure R1a ‘steppe-like’ peoples dominating over thousands of kms. in the Baltic, Fennoscandia, eastern Europe, and northern Asia for 5,000 years, with multiple Balto-Slavs-over-Balto-Slavs migrations, because these absolute units of Indo-European peoples were a trip and a half. They are the Asterix and Obelix of white Indo-European prehistory.
  • Perhaps in the meantime we can also invent some new glottochronological dialectal scheme that fits the expansion of Sredni Stog/Corded Ware with (Germano-?)Indo-Slavonic separated earlier than any other Late PIE dialect; and Finno-Volgaic later than any other Uralic dialect, in the Middle Ages, with N1c.
balto-slavic-pca
Genetic structure of the Balto-Slavic populations within a European context according to the three genetic systems, from Kushniarevich et al. (2015). Pure Balto-Slavs from…hmm…yeah this…ancient…region…or people…cluster…Whatever, very very steppe-like peoples, the True Indo-Europeans™, so close to Yamna…almost as close as Finno-Ugrians.

To sum up: Iberia, Italy, France, the British Isles, central Europe, the Balkans, the Aegean, or Anatolia, all these territories can have a complex history of periodic admixture and language replacement everywhere, but some peoples appearing later than all others in the historical record (viz. Basques or Slavs) apparently cannot, because that would be shameful for their national or ethnic myths, and these should be respected.

Ignorance of the own past as a blank canvas to be filled in with stupid ethnolinguistic continuity, turned into something valuable that should not be challenged. Ethnonationalist-like reasoning proper of the 19th century. How can our times be called ‘modern’ when this kind of magical thinking is still prevalent, even among supposedly well-educated people?

Related

ASoSaH Reread (II): Y-DNA haplogroups among Uralians (apart from R1a-M417)

corded-ware-yamna-ancestry

This is mainly a reread of from Book Two: A Game of Clans of the series A Song of Sheep and Horses: chapters iii.5. Early Indo-Europeans and Uralians, iv.3. Early Uralians, v.6. Late Uralians and vi.3. Disintegrating Uralians.

“Sredni Stog”

While the true source of R1a-M417 – the main haplogroup eventually associated with Corded Ware, and thus Uralic speakers – is still not known with precision, due to the lack of R1a-M198 in ancient samples, we already know that the Pontic-Caspian steppes were probably not it.

We have many samples from the north Pontic area since the Mesolithic compared to the Volga-Ural territory, and there is a clear prevalence of I2a-M223 lineages in the forest-steppe area, mixed with R1b-V88 (possibly a back-migration from south-eastern Europe).

R1a-M459 (xR1a-M198) lineages appear from the Mesolithic to the Chalcolithic scattered from the Baltic to the Caucasus, from the Dniester to Samara, in a situation similar to haplogroups Q1a-M25 and R1b-L754, which supports the idea that R1a, Q1a, and R1b expanded with ANE ancestry, possibly in different waves since the Epipalaeolithic, and formed the known ANE:EHG:WHG cline.

y-dna-khvalynsk
Y-DNA samples from Khvalynsk and neighbouring cultures. See full version.

The first confirmed R1a-M417 sample comes from Alexandria, roughly coinciding with the so-called steppe hiatus. Its emergence in the area of the previous “early Sredni Stog” groups (see the mess of the traditional interpretation of the north Pontic groups as “Sredni Stog”) and its later expansion with Corded Ware supports Kristiansen’s interpretation that Corded Ware emerged from the Dnieper-Dniester corridor, although samples from the area up to ca. 4000 BC, including the few Middle Eneolithic samples available, show continuity of hg. I2a-M223 and typical Ukraine Neolithic ancestry.

NOTE. The further subclade R1a-Z93 (Y26) reported for the sample from Alexandria seems too early, given the confidence interval for its formation (ca. 3500-2500 BC); even R1a-Z645 could be too early. Like the attribution of the R1b-L754 from Khvalynsk to R1b-V1636 (after being previously classifed as of Pre-V88 and M73 subclade), it seems reasonable to take these SNP calls with a pinch of salt: especially because Yleaf (designed to look for the furthest subclade possible) does not confirm for them any subclade beyond R1a-M417 and R1b-L754, respectively.

The sudden appearance of “steppe ancestry” in the region, with the high variability shown by Ukraine_Eneolithic samples, suggests that this is due to recent admixture of incoming foreign peoples (of Ukraine Neolithic / Comb Ware ancestry) with Novodanilovka settlers.

The most likely origin of this population, taking into account the most common population movements in the area since the Neolithic, is the infiltration of (mainly) hunter-gatherers from the forest areas. That would confirm the traditional interpretation of the origin of Uralic speakers in the forest zone, although the nature of Pontic-Caspian settlers as hunter-gatherers rather than herders make this identification today fully unnecessary (see here).

EDIT (3 FEB 2019): As for the most common guesstimates for Proto-Uralic, roughly coinciding with the expansion of this late Sredni Stog community (ca. 4000 BC), you can read the recent post by J. Pystynen in Freelance Reconstruction, Probing the roots of Samoyedic.

eneolithic-ukraine-corded-ware
Late Sredni Stog admixture shows variability proper of recent admixture of forest-steppe peoples with steppe-like population. See full version here.

NOTE. Although my initial simplistic interpretation (of early 2017) of Comb Ware peoples – traditionally identified as Uralic speakers – potentially showing steppe ancestry was probably wrong, it seems that peoples from the forest zone – related to Comb Ware or neighbouring groups like Lublyn-Volhynia – reached forest-steppe areas to the south and eventually expanded steppe ancestry into east-central Europe through the Volhynian Upland to the Polish Upland, during the late Trypillian disintegration (see a full account of the complex interactions of the Final Eneolithic).

The most interesting aspect of ascertaining the origin of R1a-M417, given its prevalence among Uralic speakers, is to precisely locate the origin of contacts between Late Proto-Indo-European and Proto-Uralic. Traditionally considered as the consequence of contacts between Middle and Upper Volga regions, the most recent archaeological research and data from ancient DNA samples has made it clear that it is Corded Ware the most likely vector of expansion of Uralic languages, hence these contacts of Indo-Europeans of the Volga-Ural region with Uralians have to be looked for in neighbours of the north Pontic area.

sredni-stog-repin-contacts
Sredni Stog – Repin contacts representing Uralic – Late Indo-European contacts were probably concentrated around the Don River.

My bet – rather obvious today – is that the Don River area is the source of the earliest borrowings of Late Uralic from Late Indo-European (i.e. post-Indo-Anatolian). The borrowing of the Late PIE word for ‘horse’ is particularly interesting in this regard. Later contacts (after the loss of the initial laryngeal) may be attributed to the traditionally depicted Corded Ware – Yamna contact zone in the Dnieper-Dniester area.

NOTE. While the finding of R1a-M417 populations neighbouring R1b-L23 in the Don-Volga interfluve would be great to confirm these contacts, I don’t know if the current pace of more and more published samples will continue. The information we have right now, in my opinion, suffices to support close contacts of neighbouring Indo-Europeans and Uralians in the Pontic-Caspian area during the Late Eneolithic.

Classical Corded Ware

After some complex movements of TRB, late Trypillia and GAC peoples, Corded Ware apparently emerged in central-east Europe, under the influence of different cultures and from a population that probably (at least partially) stemmed from the north Pontic forest-steppe area.

Single Grave and central Corded Ware groups – showing some of the earliest available dates (emerging likely ca. 3000/2900 BC) – are as varied in their haplogroups as it is expected from a sink (which does not in the least resemble the Volga-Ural population):

Interesting is the presence of R1b-L754 in Obłaczkowo, potentially of R1b-V88 subclade, as previously found in two Central European individuals from Blätterhole MN (ca. 3650 and 3200 BC), and in the Iron Gates and north Pontic areas.

Haplogroups I2a and G have also been reported in early samples, all potentially related to the supposed Corded Ware central-east European homeland, likely in southern Poland, a region naturally connected to the north Pontic forest-steppe area and to the expansion of Neolithic groups.

corded-ware-haplogroups
Y-DNA samples from early Corded Ware groups and neighbouring cultures. See full version.

The true bottlenecks under haplogroup R1a-Z645 seem to have happened only during the migration of Corded Ware to the east: to the north into the Battle Axe culture, mainly under R1a-Z282, and to the south into Middle Dnieper – Fatyanovo-Balanovo – Abashevo, probably eventually under R1a-Z93.

This separation is in line with their reported TMRCA, and supports the split of Finno-Permic from an eastern Uralic group (Ugric and Samoyedic), although still in contact through the Russian forest zone to allow for the spread of Indo-Iranian loans.

This bottleneck also supports in archaeology the expansion of a sort of unifying “Corded Ware A-horizon” spreading with people (disputed by Furholt), the disintegrating Uralians, and thus a source of further loanwords shared by all surviving Uralic languages.

Confirming this ‘concentrated’ Uralic expansion to the east is the presence of R1a-M417 (xR1a-Z645) lineages among early and late Single Grave groups in the west – which essentially disappeared after the Bell Beaker expansion – , as well as the presence of these subclades in modern Central and Western Europeans. Central European groups became thus integrated in post-Bell Beaker European EBA cultures, and their Uralic dialect likely disappeared without a trace.

NOTE. The fate of R1b-L51 lineages – linked to North-West Indo-Europeans undergoing a bottleneck in the Yamna Hungary -> Bell Beaker migration to the west – is thus similar to haplogroup R1a-Z645 – linked to the expansion of Late Uralians to the east – , hence proving the traditional interpretation of the language expansions as male-driven migrations. These are two of the most interesting genetic data we have to date to confirm previous language expansions and dialectal classifications.

It will be also interesting to see if known GAC and Corded Ware I2a-Y6098 subclades formed eventually part of the ancient Uralic groups in the east, apart from lineages which will no doubt appear among asbestos ware groups and probably hunter-gatherers from north-eastern Europe (see the recent study by Tambets et al. 2018).

Corded Ware ancestry marked the expansion of Uralians

Sadly, some brilliant minds decided in 2015 that the so-called “Yamnaya ancestry” (now more appropriately called “steppe ancestry”) should be associated to ‘Indo-Europeans’. This is causing the development of various new pet theories on the go, as more and more data contradicts this interpretation.

There is a clear long-lasting cultural, populational, and natural barrier between Yamna and Corded Ware: they are derived from different ancestral populations, which show clearly different ancestry and ancestry evolution (although they did converge to some extent), as well as different Y-DNA bottlenecks; they show different cultures, including those of preceding and succeeding groups, and evolved in different ecological niches. The only true steppe pastoralists who managed to dominate over grasslands extending from the Upper Danube to the Altai were Yamna peoples and their cultural successors.

corded-ware-yamna-pca
Corded Ware admixture proper of expanding late Sredni Stog-like populations from the forest-steppe. See full version here.

NOTE. You can also read two recent posts by FrankN in the blog aDNA era, with detailed information on the Pontic-Caspian cultures and the formation of “steppe ancestry” during the Palaeolithic, Mesolithic and Neolithic: How did CHG get into Steppe_EMBA? Part 1: LGM to Early Holocene and How did CHG get into Steppe_EMBA? Part 2: The Pottery Neolithic. Unlike your typical amateur blogger on genetics using few statistical comparisons coupled with ‘archaeolinguoracial mumbo jumbo’ to reach unscientific conclusions, these are obviously carefully redacted texts which deserve to be read.

I will not enter into the discussion of “steppe ancestry” and the mythical “Siberian ancestry” for this post, though. I will just repost the opinion of Volker Heyd – an archaeologist specialized in Yamna Hungary and Bell Beakers who is working with actual geneticists – on the early conclusions based on “steppe ancestry”:

[A]rchaeologist Volker Heyd at the University of Bristol, UK, disagreed, not with the conclusion that people moved west from the steppe, but with how their genetic signatures were conflated with complex cultural expressions. Corded Ware and Yamnaya burials are more different than they are similar, and there is evidence of cultural exchange, at least, between the Russian steppe and regions west that predate Yamnaya culture, he says. None of these facts negates the conclusions of the genetics papers, but they underscore the insufficiency of the articles in addressing the questions that archaeologists are interested in, he argued. “While I have no doubt they are basically right, it is the complexity of the past that is not reflected,” Heyd wrote, before issuing a call to arms. “Instead of letting geneticists determine the agenda and set the message, we should teach them about complexity in past human actions.

Related

R1a-Z280 lineages in Srubna; and first Palaeo-Balkan R1b-Z2103?

herodotus-world-map

Scythian samples from the North Pontic area are far more complex than what could be seen at first glance. From the new Y-SNP calls we have now thanks to the publications at Molgen (see the spreadsheet) and in Anthrogenica threads, I think this is the basis to work with:

NOTE. I understand that writing a paper requires a lot of work, and probably statistical methods are the main interest of authors, editors, and reviewers. But it is difficult to comprehend how any user of open source tools can instantly offer a more complex assessment of the samples’ Y-SNP calls than professionals working on these samples for months. I think that, by now, it should be clear to everyone that Y-DNA is often as important (sometimes even more) than statistical tools to infer certain population movements, since admixture can change within few generations of male-biased migrations, whereas haplogroups can’t…

Srubna

Srubna-Andronovo samples are as homogeneous as they always were, dominated by R1a-Z645 subclades and CWC-related (steppe_MLBA) ancestry.

The appearance of one (possibly two) R-Z280 lineages in this mixed Srubna-Alakul region of the southern Urals and this early (1880-1690 BC, hence rather Pokrovka-Alakul) points to the admixture of R1a-Z93 and R1a-Z280 already in Abashevo, which also explains the wide distribution of both subclades in the forest zones of Central Asia.

If Abashevo is the cornerstone of the Indo-Iranian / Uralic community, as it seems, the genetic admixture would initially be quite similar, undergoing in the steppes a reduction to haplogroup R1a-Z93 (obviously not complete), at the same time as it expanded to the west with Pokrovka and Srubna, and to the east with Petrovka and Andronovo. To the north, similar reductions will probably be seen following the Seima-Turbino phenomenon.

NOTE. Another R1a-Z280 has been found in the recent sample from Bronze Age Poland (see spreadsheet). As it appears right now in ancient and modern DNA, there seems to be a different distribution between subclades:

  • R1a-Z280 (formed ca. 2900 BC, TMRCA ca. 2600 BC) appears mainly distributed today to the east, in the forest and steppe regions, with the most ‘successful’ expansions possibly related to the spread of Abashevo- and Battle Axe-related cultures (Indo-Iranian and Uralic alike).
  • R1a-M458 (formed ca. 2700, TMRCA ca. 2700 BC) appears mainly distributed to the north, from central Europe to the east – but not in the steppe in aDNA, with the most ‘successful’ expansions to the west.

M458 lineages seem thus to have expanded in the steppe in sizeable numbers only after the Iranian expansions (see a map of modern R1a distributions) i.e. possibly with the expansion of Slavs, which supports the model whereby cultures from central-east Europe (like Trzciniec and Lusatian), accompanied mainly by M458 lineages, were responsible for the expansion of Proto-Balto-Slavic (and later Proto-Slavic).

The finding of haplogroup R1a-Z93, among them one Z2123, is no surprise at this point after other similar Srubna samples. As I said, the early Srubna expansion is most likely responsible for the Szólád Bronze Age sample (ca. 2100-1700 BC), and for the Balkans BA sample (ca. 1750-1625 BC) from Merichleri, due to incursions along the central-east European steppe.

cheek-pieces
Map of decorated bone/antler bridle cheek-pieces and whip handle equivalents. They are often local translations that remained faithful to the originals (from data in Piggott, 1965; Kristiansen & Larsson, 2005; David, 2007). Image from Vandkilde (2014).

Cimmerians

Cimmerian samples from the west show signs of continuity with R1a-Z93 lineages. Nevertheless, the sample of haplogroup Q1a-Y558, together with the ‘Pre-Scythian’ sample of haplogroup N (of the Mezőcsát Culture) in Hungary ca. 980-830 BC, as well as their PCA, seem to depict an origin of these Pre-Scythian peoples in populations related to the eastern Central Asian steppes, too.

NOTE. I will write more on different movements (unrelated to Uralic expansions) from Central and East Asia to the west accompanied by Siberian ancestry and haplogroup N with the post of Ugric-Samoyedic expansions.

Scythians

The Scythian of Z2123 lineage ca. 375-203 BC from the Volga (in Mathieson et al. 2015), together with the sample scy193 from Glinoe (probably also R1a-Z2123), without a date, as well as their common Steppe_MLBA cluster, suggest that Scythians, too, were at first probably quite homogeneous as is common among pastoralist nomads, and came thus from the Central Asian steppes.

The reduction in haplogroup variability among East Iranian peoples seems supported by the three new Late Sarmatian samples of haplogroup R1a-Z2124.

Approximate location of Glinoe and Glinoe Sad (with Starosilya to the south, in Ukrainian territory):

This initial expansion of Scythians does not mean that one can dismiss the western samples as non-Scythians, though, because ‘Scythian’ is a cultural attribution, based on materials. Confirming the diversity among western Scythians, a session at the recent ISBA 8:

Genetic continuity in the western Eurasian Steppe broken not due to Scythian dominance, but rather at the transition to the Chernyakhov culture (Ostrogoths), by Järve et al.

The long-held archaeological view sees the Early Iron Age nomadic Scythians expanding west from their Altai region homeland across the Eurasian Steppe until they reached the Ponto-Caspian region north of the Black and Caspian Seas by around 2,900 BP. However, the migration theory has not found support from ancient DNA evidence, and it is still unclear how much of the Scythian dominance in the Eurasian Steppe was due to movements of people and how much reflected cultural diffusion and elite dominance. We present new whole-genome results of 31 ancient Western and Eastern Scythians as well as samples pre- and postdating them that allow us to set the Scythians in a temporal context by comparing the Western Scythians to samples before and after within the Ponto-Caspian region. We detect no significant contribution of the Scythians to the Early Iron Age Ponto-Caspian gene pool, inferring instead a genetic continuity in the western Eurasian Steppe that persisted from at least 4,800–4,400 cal BP to 2,700–2,100 cal BP (based on our radiocarbon dated samples), i.e. from the Yamnaya through the Scythian period.

(…) Our results (…) support the hypothesis that the Scythian dominance was cultural rather than achieved through population replacement.

Detail of the slide with admixture of Scythian groups in Ukraine:

scythians-admixture

The findings of those 31 samples seem to support what Krzewińska et al. (2018) found in a tiny region of Moldavia-south-western Ukraine (Glinoi, Glinoi Sad, and Starosilya).

The question, then, is as follows: if Scythian dominance was “cultural rather than achieved through population replacement”…Where are the R1b-Z2103 from? One possibility, as I said in the previous post, is that they represent pockets of Iranian R1b lineages in the steppes descended from eastern Yamna, given that this haplogroup appears in modern populations from a wide region surrounding the steppes.

The other possibility, which is what some have proposed since the publication of the paper, is that they are related to Thracians, and thus to Palaeo-Balkan populations. About the previously published Thracian individuals in Sikora et al. (2014):

thracian-samples
Geographic origin of ancient samples and ADMIXTURE results. (A) Map of Europe indicating the discovery sites for each of the ancient samples used in this study. (B) Ancestral population clusters inferred using ADMIXTURE on the HGDP dataset, for k = 6 ancestral clusters. The width of the bars of the ancient samples was increased to aid visualization. https://doi.org/10.1371/journal.pgen.1004353.g001

For the Thracian individuals from Bulgaria, no clear pattern emerges. While P192-1 still shows the highest proportion of Sardinian ancestry, K8 more resembles the HG individuals, with a high fraction of Russian ancestry.

Despite their different geographic origins, both the Swedish farmer gok4 and the Thracian P192-1 closely resemble the Iceman in their relationship with Sardinians, making it unlikely that all three individuals were recent migrants from Sardinia. Furthermore, P192-1 is an Iron Age individual from well after the arrival of the first farmers in Southeastern Europe (more than 2,000 years after the Iceman and gok4), perhaps indicating genetic continuity with the early farmers in this region. The only non-HG individual not following this pattern is K8 from Bulgaria. Interestingly, this individual was excavated from an aristocratic inhumation burial containing rich grave goods, indicating a high social standing, as opposed to the other individual, who was found in a pit.

pca-thracians

The following are excerpts from A Companion to Ancient Thrace (2015), by Valeva, Nankov, and Graninger (emphasis mine):

Thracian settlements from the 6th c. BC on:

(…) urban centers were established in northeastern Thrace, whose development was linked to the growth of road and communication networks along with related economic and distributive functions. The early establishment of markets/emporia along the Danube took place toward the middle of the first millennium BCE (Irimia 2006, 250–253; Stoyanov in press). The abundant data for intensive trade discovered at the Getic village in Satu Nou on the right bank of the Danube provides another example of an emporion that developed along the main artery of communication toward the interior of Thrace (Conovici 2000, 75–76).

Undoubtedly the most prominent manifestation of centralization processes and stratification in the settlement system of Thrace arrives with the emergence of political capitals – the leading urban centers of various Thracian political formations.

getic-thracian
Image from Volf at Vol_Vlad LiveJournal.

Their relationships with Scythians and Greeks

The Scythian presence south of the Danube must be balanced with a Thracian presence north of the river. We have observed Getae there in Alexander’s day, settled and raising grain. For Strabo the coastlands from the Danube delta north as far as the river and Greek city of Tyras were the Desert of the Getae (7.3.14), notable for its poverty and tracklessness beyond the great river. He seems to suggest also that it was here that Lysimachus was taken alive by Dromichaetes, king of the Getae, whose famous homily on poverty and imperialism only makes sense on the steppe beyond the river (7.3.8; cf. Diod. 21.12; further on Getic possessions above the Danube, Paus. 1.9 with Delev 2000, 393, who seems rather too skeptical; on poverty, cf. Ballesteros Pastor 2003). This was the kind of discourse more familiarly found among Scythians, proud and blunt in the strength of their poverty. However, as Herodotus makes clear, simple pastoralism was not the whole story as one advanced round into Scythia. For he observes the agriculture practiced north and west of Olbia. These were the lands of the Alizones and the people he calls the Scythian Ploughmen, not least to distinguish them from the Royal Scythians east of Olbia, in whose outlook, he says, these agriculturalist Scythians were their inferiors, their slaves (Hdt. 4.20). The key point here is that, as we began to see with the Getan grain-fields of Alexander’s day, there was scope for Thracian agriculturalists to maintain their lifestyles if they moved north of the Danube, the steppe notwithstanding. It is true that it is movement in the other direction that tends to catch the eye, but there are indications in the literary tradition and, especially, in the archaeological record that there was also significant movement northward from Thrace across the Danube and the Desert of the Getae beyond it.

Greek literary sources were not much concerned with Thracian migration into Scythia, but we should observe the occasional indications of that process in very different texts and contexts. At the level of myth, it is to be remembered that Amazons were regularly considered to be of Thracian ethnicity from Archaic times onward and so are often depicted in Thracian dress in Greek art (Bothmer 1957; cf. Sparkes 1997): while they are most familiar on the south coast of the Black Sea, east of Sinope, they were also located on the north coast, especially east of the Don (the ancient Tanais). Herodotus reports an origin-story of the Sauromatians there, according to which this people had been created by the union of some Scythian warriors with Amazons captured on the south coast and then washed up on the coast of Scythia (4.110). While the story is unhistorical, it is not without importance. First, it reminds us that passage north from the Danube was not the only way that Thracians, Thracian influence, and Thracian culture might find their way into Scythia. There were many more and less circuitous routes, especially by sea, that could bring Thrace into Scythia. Secondly, the myth offered some ideological basis for the Sauromatian settlement in Thrace that Strabo records, for Sauromatians might claim a Thracian origin through their Amazon forebears. Finally, rather as we saw that Heracles could bring together some of the peoples of the region, we should also observe that Ares, whose earthly home was located in Thrace by a strong Greek and Roman tradition, seems also to have been a deity of special significance and special cult among the Scythians. So much was appropriate, especially from a Classical perspective, in associations between these two peoples, whose fame resided especially in their capacity for war.

skythen
Scythians: cultures and findings (ca. 7th-4th/3rd c. BC). Greek colonies marked with concentric circles.

This broad picture of cultural contact, interaction, and osmosis, beyond simple conflict, provides the context for a range of archaeological discoveries, which – if examined separately – may seem to offer no more than a scatter of peculiarities. Here we must acknowledge especially the pioneering work of Melyukova, who has done most to develop thinking on Thracian–Scythian interaction. As she pointed out, we have a good example of Thracian–Scythian osmosis as early as the mid-seventh century bce at Tsarev Brod in northeastern Bulgaria, where a warrior’s burial combines elements of Scythian and Thracian culture (Melyukova 1965). For, while the manner of his burial and many of the grave goods find parallels in Scythia and not Thrace, there are also goods which would be odd in a Scythian burial and more at home in a Thracian one of this period (notably a Hallstatt vessel, an iron knife, and a gold diadem). Also interesting in this regard are several stone figures found in the Dobrudja which resemble very closely figures of this kind (baby) known from Scythia (Melyukova 1965, 37–38). They range in date from perhaps the sixth to the third centuries bce, and presumably were used there – as in Scythia – to mark the burials of leading Scythians deposited in the area. Is this cultural osmosis? We should probably expect osmosis to occur in tandem with the movement of artefacts, so that only good contexts can really answer such questions from case to case. However, the broad pattern is indicated by a range of factors. Particularly notable in this regard is the observable development of a Thraco-Scythian form of what is more familiar as “Scythian animal style,” a term which – it must be understood – already embraces a range of types as we examine the different examples of the style across the great expanse from Siberia to the western Ukraine. As Melyukova observes, Thrace shows both items made in this style among Scythians and, more numerous and more interesting, a Thracian tendency to adapt that style to local tastes, with observable regional distinctions within Thrace itself. Among the Getae and Odrysians the adaptation seems to have been at its height from the later fifth century to the mid-third century (Melyukova 1965, 38; 1979).

The absence of local animal style in Bulgaria before the fifth century bce confirms that we have cultural influences and osmosis at work here, though that is not to say that Scythian tradition somehow dominated its Thracian counterpart, as has been claimed (pace Melyukova 1965, 39; contrast Kitov 1980 and 1984). Of particular interest here is the horse-gear (forehead-covers, cheek-pieces, bridle fittings, and so on) which is found extensively in Romania and Bulgaria as well as in Scythia, both in hoarded deposits and in burials. This exemplifies the development of a regional animal style, not least in silver and bronze, which problematizes the whole issue of the place(s) of its production. Accordingly, the regular designation as “Thracian” of horse-gear from the rich fourth century Scythian burial of Oguz in the Ukraine becomes at least awkward and questionable (further, Fialko 1995). And let us be clear that this is no minor matter, nor even part of a broader debate about the shared development of toreutics among Thracians and Scythians (e.g., Kitov 1980 and 1984). A finely equipped horse of fine quality was a strong statement and striking display of wealth and the power it implied

(…) while Thracian pottery appears at Olbia, Scythian pottery among Thracians is largely confined to the eastern limits of what should probably be regarded as Getic territory, namely the area close to the west of the Dniester, from the sixth century bce. Rather exceptional then is the Scythian pottery noted at Istros, which has been explained as a consequence of the Scythian pursuit of the withdrawing army of Darius and, possibly, a continued Scythian grip on the southern Danube in its aftermath (Melyukova 1965, 34). The archaeology seems to show us, therefore, that the elite Thracians and Scythians were more open to adaptation and acculturation than were their lesser brethren.

palaeo-balkan-languages
Paleo-Balkan languages in Eastern Europe between 5th and 1st century BC. From Wikipedia.

Conclusion

(…) we see distinct peoples and organizations, for example as Sitalces’ forces line up against the Scythians. Much more striking, however, against that general background, are the various ways in which the two peoples and their elites are seen to interact, connect, and share a cultural interface. We see also in Scyles’ story how the Greek cities on the coast of Thrace and Scythia played a significant role in the workings of relationships between the two peoples. It is not simply that these cities straddled the Danube, but also that they could collaborate – witness the honors for Autocles, ca. 300 bce (SEG 49.1051; Ochotnikov 2006) – and were implicated with the interactions of the much greater non-Greek powers around them. At the same time, we have seen the limited reality of familiar distinctions between settled Thracians and nomadic Scythians and the limited role of the Danube too in dividing Thrace and Scythia. The interactions of the two were not simply matters of dynastic politics and the occasional shared taste for artefacts like horse-gear, but were more profoundly rooted in the economic matrix across the region, so that “Scythian” nomadism might flourish in the Dobrudja and “Thracian-style” agriculture and settlement can be traced from Thrace across the Danube as far as Olbia. All of that offers scant justification for the Greek tendency to run together Thracians and Scythians as much the same phenomenon, not least as irrational, ferocious, and rather vulgar barbarians (e.g., Plato, Rep. 435b), because such notions were the result of ignorance and chauvinism. However, Herodotus did not share those faults to any degree, so that we may take his ready movement from Scythians to Thracians to be an indication of the importance of interaction between the two peoples whom he had encountered not only as slaves in the Aegean world, but as powerful forces in their own lands (e.g., Hdt. 4.74, where Thracian usage is suddenly brought into his account of Scythian hemp). Similarly, Thucydides, who quite without need breaks off his disquisition on the Odrysians to remark upon political disunity among the Scythians (Thuc. 2.97, a favorite theme: cf. Hdt. 4.81; Xen., Cyr. 1.1.4). As we have seen throughout this discussion, there were many reasons why Thracians might turn the thoughts of serious writers to Scythians and vice versa.

It seems, following Sikora et al. (2014), that Thracian ‘common’ populations would have more Anatolian Neolithic ancestry compared to more ‘steppe-like’ samples. But there were important differences even between the two nearby samples published from Bulgaria, which may account for the close interaction between Scythians and Thracians we see in Krzewińska et al. (2018), potentially reflected in the differences between the Central, Southern and the South-Central clusters (possibly related to different periods rather than peoples??).

If these R1b-Z2103 were descended from Thracian elites, this would be the first proof of Palaeo-Balkan populations showing mainly R1b-Z2103, as I expect. Their appearance together with haplogroup I2a2a1b1 (also found in Ukraine Neolithic and in the Yamna outlier from Bulgaria) seem to support this regional continuity, and thus a long-lasting cultural and ethnic border roughly around the Danube, similar to the one found in the northern Caucasus.

However, since these samples are some 2,500 years younger than the Yamna expansion to the south, and they are archaeologically Scythians, it is impossible to say. In any case, it would seem that the main expansion of R1a-Z645 lineages to the south of the Danube – and therefore those found among modern Greeks – was mediated by the Slavic expansions centuries later.

krzewinska-scythians-pca
Modified image from Krzewińska et al. (2018), with added Y-DNA haplogroups to each defined Scythian cluster and Sarmatians. Principal component analysis (PCA) plot visualizing 35 Bronze Age and Iron Age individuals presented in this study and in published ancient individuals in relation to modern reference panel from the Human Origins data set. See image with population references.

On the Northern cluster there is a sample of haplogroup R1b-P312 which, given its position on the PCA (apparently even more ‘modern Celtic’-like than the Hallstatt_Bylany sample from Damgaard et al. 2018), it seems that it could be the product of the previous eastward Hallstatt expansion…although potentially also from a recent one?:

Especially important in the archaeology of this interior is the large settlement at Nemirov in the wooded steppe of the western Ukraine, where there has been considerable excavation. This settlement’s origins evidently owe nothing significant to Greek influence, though the early east Greek pottery there (from ca. 650 bce onward: Vakhtina 2007) and what seems to be a Greek graffito hint at its connections with the Greeks of the coast, especially at Olbia, which lay at the estuary of the River Bug on whose middle course the site was located (Braund 2008). The main interest of the site for the present discussion, however, is its demonstrable participation in the broader Hallstatt culture to its west and south (especially Smirnova 2001). Once we consider Nemirov and the forest steppe in connection with Olbia and the other locations across the forest steppe and coastal zone, together with the less obvious movements across the steppe itself, we have a large picture of multiple connectivities in which Thrace bulks large.

scythian-peoples-balkans
Early Iron Age cultures of the Carpathian basin ca. 7-6th century BC, including steppe-related groups. Ďurkovič et al. (2018).

While the above description of clear-cut R1a-Steppe and R1b-Balkans is attractive (and probably more reliable than admixture found in scattered samples of unclear dates), the true ancient genetic picture is more complicated than that:

  • There is nothing in the material culture of the published western Scythians to distinguish the supposed Thracian elites.
  • We have the sample I0575, an Early Sarmatian from the southern Urals (one of the few available) of haplogroup R1b-Z2106, which supports the presence of R1b-Z2103 lineages among Eastern Iranian-speaking peoples.
  • We also have DA30, a Sarmatian of I2b lineage from the central steppes in Kazakhstan (ca. 47 BC – 24 AD).
  • Other Sarmatian samples of haplogroup R remain undefined.
  • There is R1a-Z93 in a late Sarmatian-Hun sample, which complicates the picture of late pastoralist nomads further.

Therefore, the possibility of hidden pockets of Iranian peoples of R1b-Z2103 (maybe also R1b-P312) lineages remains the best explanation, and should not be discarded simply because of the prevalent haplogroups among modern populations, or because of the different clusters found, or else we risk an obvious circular reasoning: “this sample is not (autosomically or in prevalent haplogroups) like those we already had from the steppe, ergo it is not from this or that steppe culture.” Hopefully, the upcoming paper by Järve et al. will help develop a clearer genetic transect of Iranian populations from the steppes.

All in all, the diversity among western Scythians represents probably one of the earliest difficult cases of acculturation to be studied with ancient DNA (obviously not the only one), since Scythians combine unclear archaeological data with limited and conflicting proto-historical accounts (also difficult to contrast with the wide confidence intervals of radiocarbon dates) with different evolving clusters and haplogroups – especially in border regions with strong and continued interactions of cultures and peoples.

With emerging complex cases like these during the Iron Age, I am happy to see that at least earlier expansions show clearer Y-DNA bottlenecks, or else genetics would only add more data to argue about potential cultural diffusion events, instead of solving questions about proto-language expansions once and for all…

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