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

Proto-Tocharians: From Afanasievo to the Tarim Basin through the Tian Shan

tocharians-early-eneolithic

A reader commented recently that there is little information about Indo-Europeans from Central and East Asia in this blog. Regardless of the scarce archaeological data compared to European prehistory, I think it is premature to write anything detailed about population movements of Indo-Iranians in Asia, especially now that we are awaiting the updates of Narasimhan et al (2018).

Furthermore, there was little hope that Tocharians would be different than neighbouring Andronovo-like populations (see a recent post on my predicted varied admixture of Common Tocharians), so the history of both unrelated Late PIE languages would have had to be explained by the admixture of Afanasievo-related groups with peoples of Andronovo descent and their acculturation.

However, data reported recently by Ning, Wang et al. Current Biology (2019) confirmed that peoples of mainly Afanasievo ancestry – as opposed to those of Corded Ware-related ancestry expanding with the Srubna-Andronovo horizon – spread the Tocharian branch of Proto-Indo-European from the Altai into the Tian Shan area, surviving essentially unadmixed into the Early Iron Age.

This genetic continuity of Tocharians will no doubt help us disentangle a great part the ethnolinguistic history of speakers of the Tocharian branch of Proto-Indo-European, from Pre-Proto-Tocharians of Afanasievo to Common Tocharians of the Late Bronze Age/Iron Age eastern Tian Shan.

NOTE. Tocharian’s isolation from the rest of Late PIE dialects and its early and intense language contacts have always been the key to support an early migration and physical separation of the group, hence the traditional association with Afanasievo, a late Repin/early Yamna offshoot. Even with the current incomplete archaeological and genetic picture, there is no other option left for the expansion of Tocharian.

It is not possible to use the currently available ancestry data to map the evolution of Afanasievo ancestry, lacking a proper geographical and temporal transect of Central and East Asian groups. In spite of this, Ning, Wang, et al. (2019) is a huge leap forward, discarding some archaeological models, and leaving only a few potential routes by which Tocharians may have spread southward from the Altai.

NOTE. I have updated the maps of prehistoric cultures accordingly, with colours – as always – reflecting the language/ancestry evolution of the different groups, even though the archaeological data of some groups of Xinjiang remains scarce, so their ethnolinguistic attribution – and the colours picked for them – remain tentative.

xinjiang-andronovo-xiaohe-horizon-bronze-iron-age
A rough timeline of related archaeological sites from North Eurasia. Image modified from Yang (2019).

Tocharians

The recent book Ancient China and its Eurasian Neighbors. Artifacts, Identity and Death in the Frontier, 3000–700 BCE, by Linduff, Sun, Cao, and Liu, Cambridge University Press (2017) offers an interesting summary of the introduction of metalworking into western China.

Here are some relevant excerpts (emphasis mine):

Although [the Xinjiang] route is not uniformly agreed upon (Shelach-Lavi 2009: 134–46), this western transmission has been thought to have passed through eastern Kazakhstan, especially as it is manifest in Semireiche, with Yamnaya, Afanasievo (copper) and Andronovo (tin bronze) peoples (Mei 2000: Fig. 3). From Xinjiang this knowledge has been thought to have traveled through the Gansu Corridor via the Qijia peoples (Bagley 1999) and then into territories controlled by dynastic China. The dating of this process is still a problem, as the sites and their contents in Xinjiang are consistently later than those in Gansu, suggesting that the point of contact was in Gansu and that the knowledge then spread from there westward.

1. Eneolithic Altai

tocharians-chalcolithic-eneolithic
Afanasievo expansion ca. 3300-2600 BC. See full culture and ancient DNA maps.

The Afanasievo sites, as they are identified in Mongolia, for instance, make up an Eneolithic culture analogous to that of southern Siberia (3100/2500–2000 BCE) in the Upper Yenissei Valley that is characterized by copper tools and an economy reliant on horse, sheep and cattle breeding as well as hunting. (…) The Afanasievo is best known through study of its burials, which typically include groups of round barrows (kurgans), each up to 12 m in diameter with a stone kerb and covering a central pit grave containing multiple inhumations. In their Siberian context, burial pottery types and styles have suggested contacts with the slightly earlier Kelteminar culture of the Aral and Caspian Sea area.

The Afanasievo culture monuments, located in the northern Altai and in the Minusinsk Basin (the western Sayan), have been seen as analogous evidence for cross-Eurasian exchange. These complexes contain small collections of metal, and many of the items are made of brass, although golden, silver and iron ornaments were also identified. A mere one-fourth of these objects are tools and ornaments, while the rest consist of unshaped remains and semi-manufactured objects. Its metallurgical tradition has recently been dated by Chernykh to as early as 3100 to 2700 BCE (1992),making it more compatible chronologically with the early brass-using sites in Shaanxi mentioned above. Kovalev and Erdenebaatar have excavated barrows in Bayan-Ulgii, Mongolia, that have been carbon-dated to the first half of the third millennium BCE and associated by ceramic types and styles and burial patterns with the Afanasievo (Kovalev and Erdenebaatar 2009: 357–58). These mounded kurgans were covered with stone and housed rectangular, wooden-faced tombs that included Afanasievo-type bronze awls, plates and small “leaf-shaped” knife blades (Kovalev and Erdenebaatar 2009: Figs. 6 and 7).

They also excavated sites belonging to the more recently identified Chemurchek archaeological culture, located in the foothills of the Mongolian Altai (Kovalev 2014, 2015) (Fig. 2.6). These sites are carbon-dated to the same period as the Afanasievo burials or to c. 3100/2500–1800 BCE (six barrows in Khovd aimag and four in Bayan-Ulgo aimag). In the rectangular stone kerbed Chemurchek slab burials (Ulaaanhus sum, Bayan-ul’gi aimag and so forth), bronze items included awls; and at Khovd aimag, Bulgan sum, in addition to stone sculptures, three lead and one bronze ring were excavated (Kovalev and Erdenebaatar 2009: Figs. 2 and 3; Fig. 2.6). Although we will not know if they were produced locally until much further investigation is undertaken, these discoveries do document knowledge of various uses and types of metal objects in western and south central Mongolia. The types of metal items thus far recovered are simple tools (awls) and rings (ornamental?) not unlike those associated with Andronovo archaeological cultures as well.

This is a complex circumstance where archaeological evidence is not complete, but raises very important questions about transmission of metallurgical knowledge to and from areas in present-day China. In the 1970s some Afanasievo mounds were excavated in Central Mongolia by a Soviet–Mongolian expedition led by V. V. Volkov and E. A. Novgorodova (Novgorodova 1989: 81–85). Unfortunately, these mounds did not yield metal objects, only ceramics, but they show that the Afanasievo culture with the Eneolithic metallurgical tradition of manufacturing pure copper items had already moved east at least far as central Mongolia. In 2004, Kovalev and Erdenebaatar investigated a large Afanasievo mound, Kulala ula, in the extreme northwest of Mongolia, near the Russian border (Kovalev and Erdenebaatar 2009). There they found a copper knife and awl (Fig. 2.5). There are five C14 dates on wood, coal and human bones from this mound, which belong to the period 2890–2570 BCE. This shows that the Afanasievo culture were carriers of technology and produced artifacts in the first half of the third millennium BCE and that they also moved south along the foothills of the Mongolian Altai. Afanasievo culture in Altai and the Minusinsk basin is dated by C14 to 3600–2500 BCE (Svyatko et al. 2009; Polyakov 2010). In the north of Xinjiang in the Altai district, several typical egg-shaped vessels and two censers of Afanasievo types were found. Some of these have been obtained from the stone boxes (chambers of megalithic graves of the Chemurchek culture) (Kovalev 2011). Thus, the Afanasievo tradition of pure copper metallurgy must have spread to the northern foothills of the Tienshan Mountains no later than the mid-third millennium BCE. The links with Afanasievo and local cultures adjacent to and south of the mountains into present-day China can now be assumed.

tocharians-chalcolithic-late
Afanasievo – Chemurchek evolution ca. 2600-2200 BC. See full culture and ancient DNA maps.

2. Bronze Age Altai

Kovalev and Erdenebaatar (2014a) and later Tishkin, Grushin, Kovalev and Munkhbayar (2015) in Western Mongolia conducted large-scale excavations of megalithic barrows of the Chemurchek culture (dated about 2600–1800 BCE). This peculiar culture appeared in Dzungaria and the Mongolian Altai in the second quarter of the third millennium BCE and for some time existed together with the late Afanasievo culture, as evidenced by the findings of Afanasievo ceramics in Chemurchek graves, in the stone boxes. Unfortunately, in China we do not yet know of any metal object related,without doubt, to the Chemurchek culture. Kovalev, Erdenebaatar, Tishkin and Grushin found several leaden ear rings and one ring of tin bronze in three excavated Chemurchek stone boxes (Kovalev and Erdenebaatar 2014a; Tishkin et al. 2015). Such lead rings are typical for Elunino culture,which occupied the entire West Altai after 2400–2300 BCE (Tishkin et al. 2015). This culture had developed a tradition of bronze metallurgy with various dopants, primarily tin. Thus, the tradition of bronze metallurgy as early as this time could have penetrated the Mongolian Altai far to the south. In addition, in the Hadat ovoo Chemurchek stone box, Kovalev and Erdenebaatar discovered stone vessels refurbished with the help of copper “patches,” indicating the presence there of metallurgical production (Fig. 2.7) (Kovalev and Erdenebaatar 2014a). In one of the secondary

Chemurchek graves unearthed by Kovalev and Erdenebaatar in Bayan-Ulgi (2400–2220 BCE), a bronze awl was found (Kovalev and Erdenebaatar 2009). Kovalev and Erdenebaatar also discovered a new culture in the territory of Mongolia (Map 2.3), one that begins immediately after Chemurchek – Munkh-Khairkhan culture (Kovalev and Erdenebaatar 2009, 2014b). To date, about 17 mounds of this culture have been excavated in Khovd, Zavkhan, Khovsgol, Bulgan aimag of Mongolia. This culture dates from about 1800 to 1500 BCE, that is, contemporary with the Andronovo culture. Therefore, the Andronovo culture does not extend far into the territory of Mongolia. Three knives without dedicated handles or stems and five awls have been found in the Munkh-Khairkhan culture mounds (Fig. 2.8). All these products are made of tin bronze. (…) Additionally, eight Late Bronze Age burials (c. 1400–1100 BCE) were unearthed in the Bulgan sum of Khovd aimag and belong to another previously unknown culture called Baitag. And in the Gobi Altai, a new group of “Tevsh” sites dating to the Late Bronze Age were defined in Bayankhongor and South Gobi aimags (Miyamoto and Obata 2016: 42–50). From these Tevsh and Baitag sites, we see the expansion of burial goods to include beads of semiprecious stones (carnelian), bronze beads, buttons and rings and even the famous elaborate golden hair ornaments (Tevsh uul;Bogd sum;Uverkhanagia aimag) from the Baitag barrows (Kovalev and Erdenebaatar 2009: Fig. 5; Miyamoto and Obata 2016).

2.1. Chemurchek

About the Chemurchek culture, from A re-analysis of the Qiemu’erqieke (Shamirshak) cemeteries, Xinjiang, China, by Jia and Betts JIES (2010) 38(4):

The major characteristics of Qiemu’erqieke Phase I include:

  1. Burials with two orientations of approximately 20° or 345°.
  2. Rectangular enclosures built using large stone slabs. The size of the enclosure varies from a maximum of 28 x 30 m.*to a minimum of 10.5 x 4.4 m. (Figure 8, Table 2).
  3. *The stone enclosure located near Hayinar is the largest one at approximately 30 x 40 m. based on pacing of the site during a visit by the authors in 2008.

  4. Almost life-sized anthropomorphic stone stelae erected along one side of the stone enclosures (Lin Yun 2008).
  5. Single enclosures tend to contain one or more than one burial, all or some with stone cist coffins.
  6. The cist coffin is usually constructed using five large stone slabs, four for the sides and one on top, leaving bare earth at the base (Zhang Yuzhong 2007). Sometimes the insides of the slabs have simple painted designs (Zhang Yuzhong 2005).
  7. Primary and secondary burials occur in the same grave.
  8. Some decapitated bodies (up to 20) may be associated with the main burial in one cist.
  9. Bodies are commonly placed on the back or side with the legs drawn up.
  10. Grave goods include stone and bronze arrowheads, handmade gray or brown round-bottomed ovoid jars, and small numbers of flat-bottomed jars (Fig. 7).
  11. Clay lamps appear to occur together with roundbottomed jars.
  12. Complex incised decoration on ceramics is common but some vessels are undecorated.
  13. The stone vessels are distinctive for the high quality of manufacture.
  14. Stone moulds indicate relatively sophisticated metallurgical expertise.
  15. Artefacts made from pure copper occur.
  16. Sheep knucklebones (astragali) imply a tradition (as in historical and modern times) of keeping knucklebones for ritual or other purposes. They also indicate the herding of domestic sheep as part of the subsistence economy.
tocharians-bronze-age-early
Chemurchek culture ca. 2200-1750 BC. See full culture and ancient DNA maps.

Chemurchek dating

Available evidence suggests that the date range for Qiemu’erqieke Phase I should fall from the later third into the early second millennium BC. There are several reasons to suggest that the time span is around the early second millennium BC. Lin Yun (2008) (…) maintains that the bronze artefacts found in Phase I show a greater sophistication in the level of copper alloy technology than that of the pure copper artefacts common to the Afanasievo tradition. On this basis it might be suggested that the Afanasievo could be considered to be Chalcolithic with a time span across much of the third millennium BC ( Gorsdorf et al. 2004: 86, Fig. 1). Qiemu’erqieke Phase I, however, should more properly be considered as Bronze Age.

Lin Yun also used the bronze arrowhead from burial Ml 7 to narrow down the date of Qiemu’erqieke Phase I. Two arrowheads were found in this burial, one of them leaf shaped with a single barb on the back (Fig. 7:4). A similar arrowhead, together with its casting mould, has been found at the Huoshaogou site of Siba tradition (Li Shuicheng 2005, Sun Shuyun and Han Rufen 1997), in Gansu province, northwest China, dated around 2000-1800 BC (Li Shuicheng and Shui Tao 2000) . This supports a date in the early second millennium BC for the Qiemu’erqieke arrowhead. The painted, round-bottomed jar from the Tianshanbeilu cemetery Qia Weiming, Betts and Wu Xinhua 2008: Fig. 7, bottom left) has been considered as a hybrid between the Upper Yellow River Bronze Age cultures of Siba in northwest China and the steppe tradition of Qiemu’erqieke in west Siberia (Li Shuicheng 1999). If this assumption is correct, the date of Tianshanbeilu, around 2000 BC, can be used as a reference for Qiemu’erqieke Phase I (Jia Weiming, Betts and Wu Xinhua 2008, Lin Yun 2008, Li Shuicheng 1999). Stone arrowheads found in Qiemu’erqieke Phase I also imply that the date is likely to fall within the earlier part of the Bronze Age as no such stone arrowheads have yet been found elsewhere in sites of the Bronze Age in Xinlang dated after the beginning of the second millennium BC.*
*For example Chawuhu and Xiaohe cemeteries (Xinjiang Institute of Archaeology 1999, 2003).

pottery-afanasevo-chemurchek
Pottery of Afanasevo and East European traits from the Chemurchek complex. Image modified from Kovalev (2017).

(…) Pottery “oil burners” (goblet-like ceramic vessels, possibly lamps) have been found in three traditions: Afanasievo (Gryaznov and Krizhevskaya 1986:21), Okunevo and Qiemu’erqieke. It is believed that this oil-burner found in Siberia and the Altai is a heritage from the Yamnaya and Catacomb
cultures (Sulimirski 1970: 225, 425; Shishlina 2008:46) in the Caspian steppe further to the west, but does not seem to exist in known Andronovo cultures.
The oil-burner tends to disappear after around 2300 BC during the mid-Okunevo period. It is, however, possible that the tradition continues longer in the Qiemu’erqieke sites.

The construction of the stone enclosures also reveals a close connection between Qiemu’erqieke Phase I and the mid and late Okunevo tradition (Sokolova 2007). Slab built stone enclosures emerged in both the Okunevo and Afanasievo traditions (Gryaznov and Krizhevskaya 1986:15-23, Kovalev 2008, Sokolova 2007, Anthony 2007:310, Koryakova and Epimakhov 2007). In the early Afanasievo the enclosure is circular with no cist coffin (Anthony 2007:310, Gryaznov and Krizhevskaya 1986:20), but in the early stage of the Okunevo square stone enclosures with a single cist burial are dominant. Square or rectangular stone enclosures are a marked feature of Qiemu’erqieke Phase I, suggesting temporal relationships between Qiemu’erqieke Phase I and the Okunevo. In Okunevo chronological group II, possibly with influence from the Anfanasievo, circular stone enclosures appeared in combination with rectangular enclosures within individual cemeteries, referred to by Sokolova (2007: table 2) as hybrid examples. By Okunevo chronological group III, rectangular stone slab enclosures with multi-burials emerged again. This is the dominant form in Qiemu’erqieke Phase I. Okunevo burial traditions changed again to single cist burials in the late stage around chronological group V ( Sokol ova 2007). A specific mortuary rite of decapitated burials exists in both the Qiemu’erqieke and Okunevo traditions (Sokolova 2007, Chen Kwang-tzuu and Hiebert 1995), as does the occasional occurrence of painted designs on the interior of the slabs forming the cists ( e.g., Khavrin 1997: 70, fig. 4; 77: tab. IV.5). Based on these comparisons, the date of Qiemu’erqieke Phase I may well parallel that of the Okunevo from at least chronological group II around 2400 BC (Gorsdorf et al. 2004: fig. 1).

khuh-udzuur-barrow
Khuh Udzuuriin I-1 elite barrow (ca. 2470-2190 BC). Modified from Image modified from Kovalev (2014).

In addition to the pottery making tradition, the anthropomorphic stone stelae may also have earlier antecedents. In the Okunevo assemblage there are anthropomorphic stelae that are longer, thinner and more abstract than those of Qiemu’erqieke. There is no indication of such stelae in the Afanasievo tradition (Gryaznov and Krizhevskaya 1986:15-23). However, further to the west, anthropomorphic stone stelae are associated with the Kemi-Oba and Yamnya cultures around the third millennium BC (Telegin and Mallory 1994; Figure 13). Some major characteristics of these stelae such as the icons on the front face of the stelae (Telegin and Mallory 1994:8-9) also appear on stelae found in Qiemu’erqieke Phase I. Recalling the oil burners that may have been inherited from the Yamnya culture and which are found in the Afansievo, Okunevo and Qiemu’erqieke Phase I, it migh t be possible to speculate that Qiemu’erqieke Phase I has its origins even earlier than the first half of the third millennium BC. This idea has also been suggested by Kovalev ( 1999).

Despite the affinities with the Okunevo cultural tradition, Qiemu’erqieke Phase I appears to be a discrete regional variant. The ceramic assemblage shows traits unique to this cluster of sites, while the anthropomorphic stelae are also distinctive markers of this tradition.

khuh-udzuur-stela
Khuh Udzuur anthropomorphic stone stela, oriented toward the south – south-east. Image modified from Kovalev (2014).

3. Bronze Age Xinjiang

I recently reported on this blog the description of Xiaohe and Gumugou cemeteries from interesting Master’s thesis Shifting Memories: Burial Practices and Cultural Interaction in Bronze Age China: A study of the Xiaohe-Gumugou cemeteries in the Tarim Basin, by Yunyun Yang, Uppsala University, Department of Archaeology and Ancient History (2019).

It also offered a full summary of findings from prehistoric sites of Xinjiang related to the arrival of a cultural package from the Altai region, ultimately connected to Afanasievo. Relevant excerpts include the following (emphasis mine):

In Bronze Age Xinjiang, burials were diverse but also show some common features between different geographic sections. The main three mountains, including Kunlun Mountains, Tian Shan (mountains) and Altai Mountains, enclose the Tarim Basin, and the Dzungaria Basin, but leave the eastern part of the Tarim Basin and the south-eastern part of the Dzungaria Basin open (with easy access to the surroundings). The Hami Basin is located at the transitional area, connecting the two basins. Burials are mainly spread along the edge of the mountain ranges.

xinjiang-afanasievo-andronovo-bmac-tian-shan
An assumption of the spreading/expansion routes stone burial construct.

3.1. The Lop Nur region

In the Lop Nur region, the Xiaohe cemetery (2000-1450 BCE) and the Gumugou cemetery (1900-1800 BCE) had many common features shared, and so is the Keliyahe northern cemetery:

  • Cemeteries were located in sandy areas;
  • Rectangular/boat-shaped wooden coffins with monuments of wooden planks or poles;
  • Coffins had no bottoms;
  • The dead were placed lying straight on the back;
  • The dead were commonly buried in single graves.

The Gumugou cemetery contained six special sun-radiating-spokes burial pattern in addition to the normal burials, which were similar to the wooden coffin graves of the Xiaohe cemetery.

NOTE. For more on Xiaohe and Gumugou, see the recent post on Proto-Tocharians. See other papers on the Andronovo horizon for other Early to Middle Bronze Age cultural groups less clearly associated with the Xiaohe horizon, like Hazandu, Xintala, or the Chust culture.

From Shuicheng (2006):

An assemblage of early bronzes had been recovered from northwestern Xinjiang and the periphery of Dzungaria 准噶尔 Basin. It comprises a variety of utilitarian tools and weapons, and a small number of apparels. These artifacts bear the stamps of Andronovo Culture in form, artifact type and decorative pattern. The metallographic analysis on selected artifacts indicates that they comprise mainly of tin-bronzes that contain 2–10% of tin. Moreover, the chemical compositions of these artifacts are similar to that of the Andronovo Culture. Latter date (first half of the 1st millennium BC) artifacts of the assemblage include a small number of arsenic bronzes. In all, during the period between the mid-2nd and mid-1st millennium BC, copper and bronze artifacts coexisted in this region, albeit tin-bronze comprised the majority. The composition of alloy did not show significant change over time. Some colleagues pointed out that the Nulasai 奴拉赛 site at Nileke 尼勒克 County in the Yili 伊犁 River basin of Xinjiang was the pioneer in the use of “sulphuric ore–ice copper–copper”technology. It is also the only early smelting site in Euro-Asia that arsenic ore was added to deliberately produce an alloy

tocharians-bronze-age-middle
Prehistoric cultures of Xinjiang during the Middle Bronze Age. See full culture and ancient DNA maps.

3.2. The Hami Basin-the Balikun Grassland

From Yang (2019):

The Hami Basin-the Balikun Grassland area is located at the eastern part of Tian Shan. The area is divided in a northern basin and a southern basin by the east-west stretch of the Tian Shan. In the Hami Basin-the Balikun Grassland area, the main type of burials were earth-pit graves in the early Bronze Age, and burials of stone-pit with barrows became more common in the late Bronze Age. The Hami-Tianshan-Beilu cemetery is a representative of the earth-pit graves. The features of the Hami-Tianshan-Beilu cemetery (2000-1500 bce) here were:

  • Rectangular earth pit graves;
  • The dead were often in a hocker position lying on one side;
  • Commonly a single dead in one grave.
balikun-grassland
The Balikun grassland today (source).

The Hami-Wubu cemetery (earlier than 1000 bce) and the Yanbulake cemetery (1200-600 bce) are representatives of another common earth-pit graves. Common features here were:

  • Rectangular earth pits, with two storeys and/or roofed with wooden boards;
  • The dead was placed in a hocker position lying on one side;
  • Mostly a single dead in one grave.

Later there appeared more stone-pit graves in this area, and the features can be summarized as:

  • Round burial mounds, commonly constructed by stones or a mix of stones and earth;
  • Burial mounds with a sunken top or a normal (dome) top;
  • The diameter of the burial mounds varied between 3 and 25.4 m (but not necessarily limited in this scope);
  • Circular or rectangular stone kerbs;
  • Rectangular stone pits, constructed by earth, or stones, or a mix of earth and stones;
  • Rectangular stone pits contained wooden coffins (represented by the Yiwu Baiqi’er cemetery).
hami-basin-balikun-grassland-iron-age-burials
Some representatives of stone burials in the Hami Basin – the Balikun Grassland in the Iron Age (Adapted from: Xinjiang 2011, 29-41). Image modified from Yang (2019).

In the Hami Basin, the Bronze Age cemeteries show common burial features like earth pits and hocker position of the dead. With similar pottery styles in the Hami-Tianshan-Beilu cemetery to those in the Machang and Siba cultures (Xinjiang 2011: 17), it suggests possible cultural influence or people’s migrating from the Hexi Corridor in the east.

In the Balikun Grassland, burials in an earlier time contained mostly earth-pit graves but also a small number of stone-pit graves. The pebbles were imbedded in the floors and the walls of the graves in a rectangular shape, e.g. the Balikun-Nanwan cemetery (1600-1000 bce). In a later time, there appeared huge burial mounds with a sunken top, and with the diameters of the burial mounds varying from 3 to 25.4 m, e.g. the Balikun-Dongheigou cemetery and the Balikun-Heigouliang cemetery. The Yiwu-Bai’erqi and the Yiwu-Kuola cemeteries contained either round stone burial mounds or circular stone kerbs on the ground surface. Considering the three burial elements including burial mounds, stone pits and circular kerbs, the later period cemeteries in the Balikun Grassland were actually similar to cemeteries from the southern edge of the Altai Mountain area.

From Shuicheng (2006):

The Nanwan 南湾 cemetery site at Kuisu 奎苏 Town, Balikun 巴里坤 (1600–1100 BC) also yielded an assemblage of early bronzes. The style of its early phase artifacts is similar to that of the burials distributed in the North Tianshan Route. Some sorts of cultural connection should have existed between the two.

The dates of Yanbulake 焉不拉克 Culture (1300–700 BC) are comparatively late. Its metallurgy was a continuation of the western China tradition. Artifact types include a variety of utilitarian tools, weapons and apparels.

tocharians-bronze-age-late
Prehistoric cultures of Xinjiang during the Late Bronze Age. See full culture and ancient DNA maps.

3.3. The Turpan Basin-the middle part of Tian Shan

From Yang (2019):

Turpan Basin is located at the western part of the Hami Basin, and lies at the southern edge of the eastern Tian Shan. In the Turpan Basin-the middle part of Tian Shan area, the main representative of the Bronze Age cemeteries is the Yanghai Nr.1 cemetery. The features here were:

  • Elliptic earth pit graves, commonly covered by round logs on the top;
  • Some graves contained burial beds made of round logs or reeds;
  • The dead were mainly placed lying straight on the back;
  • Mostly a single dead in one grave.

In Iron Age, the stone burials became dominant, but the stone burials varied in different regions of the Turpan Basin-the middle part of Tian Shan area. Graves containing burial mounds, stone pit, and circular stone kerbs are represented by the Shanshan-Ertanggou cemetery, the Tuokexun-Alagou cemetery, the Urumqi-Chaiwobu cemetery and the Urumqi-Yizihu-Sayi cemetery, etc. The stone funeral construction features here are similar to those contemporary cemeteries in the Hami Basin-the Balikun Grassland area.

3.4. The southern edge of the western and middle part of Tian Shan

In the southern edge of the western and middle part of Tian Shan area, the main representatives of the late Bronze Age cemeteries are the Hejing-Chawuhu Nr.4 cemetery (around 1000-500 bce), the Hejing-Xiaoshankou cemetery, the Baicheng-cemetery, etc. The main burial features of the late Bronze Age and the early Iron Age cemeteries (see Fig.12) here were:

  • Burial mounds, constructed by stones or a mix of stones and earth;
  • Irregular circular or rectangular stone kerbs;
  • Stone pit graves in a bell-shape or a rectangular shape;
  • Stone pit graves constructed by imbedding pebbles or stone slabs in walls and floors;
  • The dead were often placed lying on their back with bent legs;
  • The dead were commonly reburied a second time with multiple burials.

From the late Bronze Age to the early Iron Age in this area, the burial traditions tended to be in a more varied way. In the stone burials with stone kerbs, there is a mixture of stone pit and earth pit graves. The burial features of the Iron Age cemeteries in this section were similar to those contemporary both in the Hami Basin-the Balikun Grassland area and in the Turpan Basin-the middle part of Tian Shan area.

From Shuicheng (2006):

The Chawuhu 察吾呼 Culture (1100–500 BC) distributes on the foothills between the middle section of the Tianshan Mountain Ranges and Tarim River. Its bronze assemblage comprises a variety of weapons, utilitarian tools and small apparels. They show no apparent temporal change in form and type through the four cultural phases. In addition, bronzes bear the Chawuhu characteristics were found in Hejing 和静, Baicheng 拜城 and Luntai 轮台 (Bügür). Yet, sites distributed along the Tarim River, such as Heshuo 和硕, Kuga 库车and Aksu 阿克苏, yielded remains of a bronze culture different from that of Chawuhu. Bronzes recovered include double-eared socketed axe, arrowheads, awls, knives, needles and bracelets. Their absolute dates have been estimated to be earlier than that of Chawuhu.

tocharians-iron-age-early
Prehistoric cultures of Xinjiang during the Early Iron Age. See full culture and ancient DNA maps

3.5. The Pamir Plateau

From Yang (2019):

A typical Bronze Age cemetery from the Pamir Plateau area is the Tashenku’ergan-Xiabandi cemetery (around 1000-500 bce). The burial features here were:

  • Mainly inhumations, but also a few cremations;
  • Burial mounds, constructed of stones;
  • Irregular circular or rectangular stone kerbs;
  • Mostly a single dead in one grave;
  • The dead was placed in a hocker position lying on one side.

The adoption of burial customs from the east supports the migration of Afanasievo-related peoples from the Tian Shan up to the Pamir Plateau, strongly influencing the findings of the Xiabandi cemetery, which has been dated from an early Bronze Age phase (ca. 1500-300 BC) to a late date up to ca. 600 BC.

While it is today unclear how far the Afanasievo admixture reached into the western Xinjiang, it seems that the Pamir Plateau remained culturally connected to neighbouring Andronovo-related cultures in pottery and metallurgical innovations, hence their language probably belonged – during most part of the Bronze and Iron Ages – to the Indo-Iranian branch, even though specific dialects might have changed with each new attested group.

In particular, it is possible that the early Andronovo groups related to the Xiaohe Horizon spoke Indo-Aryan or West Iranian dialects, while Saka-related groups replaced them – or an intermediate Tocharian-speaking group – with East Iranian dialects. A close interaction with West Iranian would justify the known ancient borrowings of Tocharian, although they could also be explained by contacts with Chust-related groups farther west. For more on this, see Ged Carling’s work on the different layers of Iranian loans.

Xinjiang BA/IA Summary

From Yang (2019):

In the early Bronze Age, there are distinct regional differences in the burial customs in and surrounding the Tarim Basin. At the southern edge of the Altai Mountains area, the burial customs included stone burial mounds, stone pit graves, circular or rectangular stone kerbs and stone human sculptures; the dead were placed lying straight on the back. In the Hami Basin-the Balikun Grassland area, the burial customs included earth pit graves; the dead were placed in a hocker position lying on one side. In the Turpan Basin-the middle part of Tian Shan area, the burial customs included earth pit graves; the dead were placed lying straight on the back. In the Lop Nur region, the burial customs included wooden coffins buried in sand; the dead were placed lying straight on the back.

But from the late Bronze Age to the early Iron Age, there was a common shift in burial customs from earth pit graves to stone burials in the Hami Basin-the Balikun Grassland area and in the Turpan Basin-the middle part of Tian Shan area. The main features of the stone burials include stone burial mounds, circular or rectangular stone kerbs, and the stone pit graves in the cemeteries. Similar stone burial customs commonly appeared at the southern edge of the western and middle part of Tian Shan area and the Pamir Plateau area in Iron Age. The burial features in most areas are in a mixture of both the earth pit graves and stone pit graves, especially in the Hami Basin-the Balikun Grassland area and the Turpan Basin-the middle part of Tian Shan area.

xinjiang-bronze-age-iron-age

From Shuicheng (2006):

Historians of metallurgy conducted metallographic analyses on a sample of 234 metal specimens recovered from 16 localities in eastern Xinjiang. They concluded that the metallurgic industry in eastern Xinjiang could be roughly partitioned into three developmental phases. The early phase is represented by the burials distributed in the North Tianshan Route. The majority of the metal assemblage was tin-bronzes; however, copper and arsenic-bronzes maintained considerable proportions. The middle phase is represented by the burials at Yanbulake. During this phase, tin-bronze still maintained the majority; the proportion of arsenic-bronze increased, and some of them were high arsenic-bronzes. The late phase is represented by the burials at Heigouliang 黑沟梁. The composition of lead increased in the bronze alloy in the expense of arsenic. In addition, this phase witnessed the appearance of high tin-bronze that composed up to 16% of tin and the appearance of brass, that is, an alloy of copper and zinc. The bronze alloy consistently contained significant amount of impurities regardless of temporal difference. Casting and forging technologies coexisted throughout the three phases. The early bronzes (2000–500 BC) of eastern Xinjiang, in general, contained arsenic; however, the composition of arsenic was usually under 8%, but a few artifacts contained more than 20% arsenic. In all, arsenic had long been used in the alloy-forming of the early bronzes in eastern Xinjiang. Consequently, arsenic-bronzes were widely found in the prehistoric archaeology of the region. The artifact types, chemical compositions and manufacture techniques of the bronze assemblage of the burials of the North Tianshan Route are similar to those of Siba Culture, indicating that eastern Xinjiang had played a significant role in the East-West interactions.

An assemblage of early bronzes had been recovered from northwestern Xinjiang and the periphery of Dzungaria 准噶尔 Basin. It comprises a variety of utilitarian tools and weapons, and a small number of apparels. These artifacts bear the stamps of Andronovo Culture in form, artifact type and decorative pattern. The metallographic analysis on selected artifacts indicates that they comprise mainly of tin-bronzes that contain 2–10% of tin. Moreover, the chemical compositions of these artifacts are similar to that of the Andronovo Culture. Latter date (first half of the 1st millennium BC) artifacts of the assemblage include a small number of arsenic-bronzes. In all, during the period between the mid-2nd and mid-1st millennium BC, copper and bronze artifacts coexisted in this region, albeit tin-bronze comprised the majority.

tocharians-iron-age-late
Prehistoric cultures of Xinjiang during the Late Iron Age. See full culture and ancient DNA maps.

Tocharians in population genomics

Prehistoric population movements between the Altai and the Tian Shan are difficult to pinpoint, not the least because of the division of these territories among three different countries and their archaeological teams, only recently (more) open to the international scholarship.

The available schematic archaeological picture, where migrations could only be roughly inferred, has been recently updated to a great extent by Ning, Wang et al. (2019), whose genetic analysis of the samples is as thorough as anyone could have asked for, with a level of detail which matches the complex genetic picture of the region by the Iron Age.

As a summary, here is what they described about the samples from Shirenzigou (ca 400-200 BC), corresponding to the Iron Age populations of the Hami Basin-the Balikun Grassland area, and closely related to the preceding Yanbulake Culture:

As shown in Figure S3, the Steppe_MLBA populations including Srubnaya, Andronovo, and Sintashta were shifted toward farming populations compared with Yamnaya groups and the Shirenzigou samples. This observation is consistent with ADMIXTURE analysis that Steppe_MLBA populations have an Anatolian and European farmer-related component that Yamnaya groups and the Shirenzigou individuals do not seem to have. The analysis consistently suggested Yamnaya-related Steppe populations were the better source in modeling the West Eurasian ancestry in Shirenzigou.

biplot-yamnaya-tocharians-shirenzigou
Biplot of f3-outgroup tests illustrating the Kostenki14 and Anatolia_N like ancestries in Shirenzigou individuals. Most Shirenzigou individuals were on a cline with Yamnaya and European hunter-gatherer groups, lacking the European farmer ancestry as compared to the Steppe_MLBA populations such as Andronovo, Srubnaya and Sintashta [S1-S5]. Horizontal and vertical bars represent ± 3 standard errors, corresponding to form of outgroup f3 tests on the x axis and y axis respectively.

We continued to use qpAdm to estimate the admixture proportions in the Shirenzigou samples by using different pairs of source populations, such as Yamnaya_Samara, Afanasievo, Srubnaya, Andronovo, BMAC culture (Bustan_BA and Sappali_ Tepe_BA) and Tianshan_Hun as the West Eurasian source and Han, Ulchi, Hezhen, Shamanka_EN as the East Eurasian source. In all cases, Yamnaya, Afanasievo, or Tianshan_Hun always provide the best model fit for the Shirenzigou individuals, while Srubnaya, Andronovo, Bustan_BA and Sappali_Tepe_BA only work in some cases.

p-values-shirenzigou-samples-han-chinese
Table S2. P values in modelling a two-way (P=rank 1) admixture in Shirenzigou samples using each of the four populations (Bustan_BA, Sappali_Tepe_BA, Andronovo.SG, Srubnaya) together with Han Chinese as two sources [S6], Related to Figure 2. We used the following set of outgroups populations: Dinka, Ust_Ishim, Kostenki14, Onge, Papuan, Australian, Iran_N, EHG, LBK_EN.

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

In the PCA, ADMIXTURE, outgroup f3 statistics [see Figure S4], as well as f4 statistics (Table S3), we observed the Shirenzigou individuals were closer to the present day Tungusic and Mongolic-speaking populations in northern Asia than to the populations in central and southern China, suggesting the northern populations might contribute more to the Shirenzigou individuals. Based on this, we then modeled Shirenzigou as a three-way admixture of Yamnaya_Samara, Ulchi (or Hezhen) and Han to infer the source from the East Eurasia side that contributed to Shirenzigou. We found the Ulchi or Hezhen and Han-related ancestry had a complicated and unevenly distribution in the Shirenzigou samples. The most Shirenzigou individuals derived the majority of their East Eurasian ancestry from Ulchi or Hezhen-related populations, while the following two individuals M820 and M15-2 have more Han related than Ulchi/ Hezhen-related ancestry

It is unclear whether the Chemurchek population will show a sizeable local contribution from neighbouring groups. The fact that Okunevo shows 20% Yamnaya-related ancestry strongly supports the nature of neighbouring stone-grave-building peoples of the Altai and the northern Tian Shan as mostly Afanasievo-like, and the apparent lack of contributions of Srubna/Andronovo-like ancestry in the early Hami-Balikun stone burial builders also speaks for radical population replacement events reaching the areas south of Tian Shan, at least initially.

While ancestry cannot settle linguistic questions, it seems that nomads of the Gansu and Qinghai grasslands retained an ancestry close to Andronovo, whereas nomads of the Hami Basin-Balikun grasslands and related populations of Xinjiang remained closely related to Afanasievo. This doesn’t preclude that the ancestors of the Yuezhi became acculturated under the influence of peoples from eastern Xinjiang, but all data combined suggest an isolation of both populations – relative to other groups and to each other – and it is therefore more likely that they spoke Indo-Iranian-related languages rather than a language of the Tocharian branch.

Haplogroups

In an interesting twist of events, despite the initially reported hg. R1b and Q, Tocharians from Shirenzigou actually show a haplogroup diversity comparable to that attested in other late Iron Age populations: a similar diversity is seen, for example, among Germanic, Baltic, and Balto-Finnic peoples of the Baltic region; among East Germanic or Scythians of the north Pontic region; or among Mediterranean peoples sampled to date. Iron Age peoples show thus a complex sociopolitical setting that overcame the previous patrilineal homogeneity of Bronze Age expansions.

tocharians-pca
PCA and ADMIXTURE for Shirenzigou Samples. Modified from the original to include in black squares samples related to Yamnaya. Modified from the paper to include labels of modern populations and a dotted lines with the cline formed by Shirenzigou, from (Yamnaya-like) Afanasievo to Central and East Asian-like populations. In red circles, samples with best fit for Andronovo-like ancestry. In green circles, samples with Han-related admixture.

M15-2 (with Han-related ancestry) is of the rare haplogroup Q1a-M120, while the samples with highest Steppe_MLBA-related ancestry are of hg. R1b-PH155, which points to their recent origin among Yuezhi, or to Hun-related populations showing an admixture related to the proto-historic nomads of the Gansu and Qinghai grasslands.

The expansion of Chemurchek-related peoples was probably associated more with hg. Q1a (dubious if it’s a Pre-ISOGG 2017 nomenclature, hence possibly Q1b), a haplogroup that might be found in Khvalynsk as a “significant minority” according to Anthony (2019), and it might also be attested in sampled individuals from Afanasievo in its late phase. This might be, therefore, a case similar to the early expansion of Indo-Europeans with R1b-V1636 lineages through the Volga – North Caucasus region, and of the later expansion with I2a-L699 lineages into the Balkans.

Haplogroup Q1a2-M25 is found in individual X3, whose Steppe ancestry is likely a combination of Afanasievo plus Andronovo-like ancestry heavily admixed with Hezhen/Ulchi-like populations, in line with the expected recent contacts with the neighbouring Xiongnu, Yuezhi, and other population movements affecting eastern Xinjiang.

Sample M4, which packs the most Afanasievo-like ancestry, is of hg. R1a-Z645, which – like sample M8R1 of hg. O – is most likely related to haplogroup resurgence events of local populations, which left the predominant Afanasievo-like admixture brought by builders of stone burials essentially intact, evidenced by the almost 100% of R1a found in the Xiaohe cemetery – and in most of the early Andronovo horizon – and among expanding Kangju and Wusun, as well as by the prevalence of hg. O among sampled East Asian populations.

A question that will only be answered with more samples is how and when the prevalent R1b-L23 and Q1b lineages among Afanasievo-related peoples began to be replaced to reach the high variability seen in Shirenzigou. Given the pastoralist nature of peoples around Tian Shan, the succeeding expansions of Proto-Tocharians, and the late isolation of different Common Tocharian groups, it is more than likely that this variability represents a late and local phenomenon within Xinjiang itself.

tocharians-antiquity
Peoples of Xinjiang during Antiquity. See full culture and ancient DNA maps.

Conclusion

Tocharians are one of the main pillars that confirm the Late Proto-Indo-European homeland of the R1b-rich populations of the Don-Volga region. There is already:

Just like the East Bell Beaker expansion from Yamnaya Hungary has confirmed that Corded Ware peoples did not partake in spreading Indo-European languages (spreading Uralic languages instead), data on the expansion of Tocharian speakers from Afanasievo to the Tian Shan was always there; population genomics is merely helping to connect the dots.

In summary, genetic research is supporting the expected linguistic expansions of the Neolithic and Bronze Age step by step, slowly but surely.

Related

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

The genetic makings of South Asia – IVC as Proto-Dravidian

south-asian-language-families

Review (behind paywall) The genetic makings of South Asia, by Metspalu, Monda, and Chaubey, Current Opinion in Genetics & Development (2018) 53:128-133.

Interesting excerpts (emphasis mine):

(…) the spread of agriculture in Europe was a result of the demic diffusion of early Anatolian farmers, it was discovered that the spread of agriculture to South Asia was mediated by a genetically completely different farmer population in the Zagros mountains in contemporary Iran (IF). The ANI-ASI cline itself was interpreted as a mixture of three components genetically related to Iranian agriculturalists, Onge and Early and Middle Bronze Age Steppe populations (Steppe_EMBA).

The first ever autosomal aDNA from South Asia comes from Northern Pakistan (Swat Valley, early Iron Age). This study presented altogether 362 aDNA samples from the broad South and Central Asia and contributes substantially to our understanding of the evolutionary past of South and Central Asia. The study redefines the three genetic strata that form the basis of the Indian Cline. The Indus Periphery (IP) component is composed of (varying proportions of): first, IF, second, Ancient Ancestral South Asians (AASI), which represents an ancient branch of human genetic variation in Asia arising from a population split contemporaneous with the splits of East Asian, Onge and Australian Aboriginal ancestors and third, West_Siberian Hunter gatherers (WS_HG).

The authors argue that IP could have formed the genetic base of the Indus Valley Civilization (IVC). Upon the collapse of the IVC IP contributes to the formation of both ASI and ANI. ASI is formed as IP admixes further with AASI. ANI in turn forms when IP admixes with the incoming Middle and Late Bronze Age Steppe (Steppe_MLBA) component, (rather than the Steppe_EMBA groups suggested earlier)

ane-whg-ehg-chg-wshg-steppe
A sketch of the peopling history of South Asia. Depicting the full complexity of available reconstructions is not attempted. Placing of population labels does not indicate precise geographic location or range of the population in question. Rather we aim to highlight the essentials of the recent advancements in the field. We divide the scenario into three time horizons: Panels (a) before 10 000 BCE (pre agriculture era.); (b) 10 000 BCE to 3000 BCE (agriculture era) and (c) 3000 BCE to prehistoric era/modern era. (iron age).

Dating of the arrival of the Austro-Asiatic speakers in South Asia-based on Y chromosome haplogroup O2a1-M95 expansion estimates yielded dates between 3000 and 2000 BCE [30]. However, admixture LD decay-based approach on genome-wide data suggests the admixture between South Asian and incoming Austro-Asiatic speakers occurred slightly later between 1800 and 0 BCE (Tätte et al. submitted). It is interesting that while the mtDNA variants of the Mundas are completely South Asian, the Y chromosome variation is dominated at >60% by haplogroup O2a which is phylogeographically nested in East Asian-specific paternal lineages.

In India, the speakers of Tibeto-Burman (TB) languages live in the Seven Sisters States in Northeast India and in the very north of the country. Genetically they show a clear East Asian origin and around 20% of subsequent admixture with South Asians within the last 1000 years.The genetic flavour of East Asia in TB is different from that in Munda speakers as the best surrogates for the East Asian admixing component are contemporary Han Chinese.

I found the simplistic migration maps especially interesting to illustrate ancient population movements. The emergence of EHG is supposed to involve a WHG:ANE cline, though, and this isn’t clear from the map. Also, there is new information on what may be at the origin of WHG and Anatolian hunter-gatherers.

From the recent Reich’s session on South Asia at ISBA 8:

ani-asi-steppe-cline
– Tale of three clines, with clear indication that “Indus Periphery” samples drawn from an already-cosmopolitan and heterogeneous world of variable ASI & Iranian ancestry. (I know how some people like to pore over these pictures – so note red dots = just dummy data for illustration.)
– Some more certainty about primary window of steppe ancestry injection into S. Asia: 2000-1500 BC
Alexander M. Kim

Featured image: map of South Asian languages from http://llmap.org.

Related

Evolution of Steppe, Neolithic, and Siberian ancestry in Eurasia (ISBA 8, 19th Sep)

jena-isba8

Some information is already available from ISBA 8 (see programme in PDF), thanks to the tweets from Alexander M. Kim.

Official abstracts are listed first (emphasis mine), then reports and images with link to Kim’s tweets. Here is the list for quick access:

Updates (17:00 CET):

Turkic and Hunnic expansions

Tracing the origin and expansion of the Turkic and Hunnic confederations, by Flegontov et al.

Turkic-speaking populations, now spread over a vast area in Asia, are highly heterogeneous genetically. The first confederation unequivocally attributed to them was established by the Göktürks in the 6th c. CE. Notwithstanding written resources from neighboring sedentary societies such as Chinese, Persian, Indian and Eastern Roman, earlier history of the Turkic speakers remains debatable, including their potential connections to the Xiongnu and Huns, which dominated the Eurasian steppe in the first half of the 1st millennium CE. To answer these questions, we co-analyzed newly generated human genome-wide data from Central Asia (the 1240K panel), spanning the period from ca. 3000 to 500 YBP, and the data published by de Barros Damgaard et al. (137 ancient human genomes from across the Eurasian steppes, Nature, 2018). Firstly, we generated a PCA projection to understand genetic affinities of ancient individuals with respect to present-day Tungusic, Mongolic, Turkic, Uralic, and Yeniseian-speaking groups. Secondly, we modeled hundreds of present-day and few ancient Turkic individuals using the qpAdm tool, testing various modern/ancient Siberian and ancient West Eurasian proxies for ancestry sources.

A majority of Turkic speakers in Central Asia, Siberia and further to the west share the same ancestry profile, being a mixture of Tungusic or Mongolic speakers and genetically West Eurasian populations of Central Asia in the early 1st millennium CE. The latter are themselves modelled as a mixture of Iron Age nomads (western Scythians or Sarmatians) and ancient Caucasians or Iranian farmers. For some Turkic groups in the Urals and the Altai regions and in the Volga basin, a different admixture model fits the data: the same West Eurasian source + Uralic- or Yeniseian-speaking Siberians. Thus, we have revealed an admixture cline between Scythians and the Iranian farmer genetic cluster, and two further clines connecting the former cline to distinct ancestry sources in Siberia. Interestingly, few Wusun-period individuals harbor substantial Uralic/Yeniseian-related Siberian ancestry, in contrast to preceding Scythians and later Turkic groups characterized by the Tungusic/Mongolic-related ancestry. It remains to be elucidated whether this genetic influx reflects contacts with the Xiongnu confederacy. We are currently assembling a collection of samples across the Eurasian steppe for a detailed genetic investigation of the Hunnic confederacies.

jeong-population-clines
Three distinct East/West Eurasian clines across the continent with some interesting linguistic correlates, as earlier reported by Jeong et al. (2018). Alexander M. Kim.
siberian-genetic-component-chronology
Very important observation with implication of population turnover is that pre-Turkic Inner Eurasian populations’ Siberian ancestry appears predominantly “Uralic-Yeniseian” in contrast to later dominance of “Tungusic-Mongolic” sort (which does sporadically occur earlier). Alexander M. Kim

New interesting information on the gradual arrival of the “Uralic-Yeniseian” (Siberian) ancestry in eastern Europe with Iranian and Turkic-speaking peoples. We already knew that Siberian ancestry shows no original relationship with Uralic-speaking peoples, so to keep finding groups who expanded this ancestry eastwards in North Eurasia should be no surprise for anyone at this point.

Central Asia and Indo-Iranian

The session The Genomic Formation of South and Central Asia, by David Reich, on the recent paper by Narasimhan et al. (2018).

bmac-reich
One important upside of dense genomic sampling at single localities – greater visibility of outliers and better constraints on particular incoming ancestries’ arrival times. Gonur Tepe as a great case study of this. Alexander M. Kim
ani-asi-steppe-cline
– Tale of three clines, with clear indication that “Indus Periphery” samples drawn from an already-cosmopolitan and heterogeneous world of variable ASI & Iranian ancestry. (I know how some people like to pore over these pictures – so note red dots = just dummy data for illustration.)
– Some more certainty about primary window of steppe ancestry injection into S. Asia: 2000-1500 BC
Alexander M. Kim

British Isles

Ancient DNA and the peopling of the British Isles – pattern and process of the Neolithic transition, by Brace et al.

Over recent years, DNA projects on ancient humans have flourished and large genomic-scale datasets have been generated from across the globe. Here, the focus will be on the British Isles and applying aDNA to address the relative roles of migration, admixture and acculturation, with a specific focus on the transition from a Mesolithic hunter-gatherer society to the Neolithic and farming. Neolithic cultures first appear in Britain ca. 6000 years ago (kBP), a millennium after they appear in adjacent areas of northwestern continental Europe. However, in Britain, at the margins of the expansion the pattern and process of the British Neolithic transition remains unclear. To examine this we present genome-wide data from British Mesolithic and Neolithic individuals spanning the Neolithic transition. These data indicate population continuity through the British Mesolithic but discontinuity after the Neolithic transition, c.6000 BP. These results provide overwhelming support for agriculture being introduced to Britain primarily by incoming continental farmers, with surprisingly little evidence for local admixture. We find genetic affinity between British and Iberian Neolithic populations indicating that British Neolithic people derived much of their ancestry from Anatolian farmers who originally followed the Mediterranean route of dispersal and likely entered Britain from northwestern mainland Europe.

british-isles
Millennium of lag between farming establishment in NW mainland Europe & British Isles. Only 25 Mesolithic human finds from Britain. Alexander M. Kim.
british-admixture
– Evidently no resurgence of hunter-gatherer ancestry across Neolithic
– Argument for at least two geographically distinct entries of Neolithic farmers
Alexander M. Kim.

MN Atlantic / Megalithic cultures

Genomics of Middle Neolithic farmers at the fringe of Europe, by Sánchez Quinto et al.

Agriculture emerged in the Fertile Crescent around 11,000 years before present (BP) and then spread, reaching central Europe some 7,500 years ago (ya.) and eventually Scandinavia by 6,000 ya. Recent paleogenomic studies have shown that the spread of agriculture from the Fertile Crescent into Europe was due mainly to a demic process. Such event reshaped the genetic makeup of European populations since incoming farmers displaced and admixed with local hunter-gatherers. The Middle Neolithic period in Europe is characterized by such interaction, and this is a time where a resurgence of hunter-gatherer ancestry has been documented. While most research has been focused on the genetic origin and admixture dynamics with hunter-gatherers of farmers from Central Europe, the Iberian Peninsula, and Anatolia, data from farmers at the North-Western edges of Europe remains scarce. Here, we investigate genetic data from the Middle Neolithic from Ireland, Scotland, and Scandinavia and compare it to genomic data from hunter-gatherers, Early and Middle Neolithic farmers across Europe. We note affinities between the British Isles and Iberia, confirming previous reports. However, we add on to this subject by suggesting a regional origin for the Iberian farmers that putatively migrated to the British Isles. Moreover, we note some indications of particular interactions between Middle Neolithic Farmers of the British Isles and Scandinavia. Finally, our data together with that of previous publications allow us to achieve a better understanding of the interactions between farmers and hunter-gatherers at the northwestern fringe of Europe.

megalithic-europe
-Novel genomic data from 21 individuals from 6 sites.
– “Megalithic” individuals not systematically diff. from geographically proximate “non-megalithic” burials
– Mild evidence for over-representation of males in some British Isles megalithic tombs
– Megalithic tombs in W & N Neolithic Europe may have link to kindred structures
Alexander M. Kim

Central European Bronze Age

Ancient genomes from the Lech Valley, Bavaria, suggest socially stratified households in the European Bronze Age, by Mittnik et al.

Archaeogenetic research has so far focused on supra-regional and long-term genetic developments in Central Europe, especially during the third millennium BC. However, detailed high-resolution studies of population dynamics in a microregional context can provide valuable insights into the social structure of prehistoric societies and the modes of cultural transition.

Here, we present the genomic analysis of 102 individuals from the Lech valley in southern Bavaria, Germany, which offers ideal conditions for such a study. Several burial sites containing rich archaeological material were directly dated to the second half of the 3rd and first half of the 2nd millennium BCE and were associated with the Final Neolithic Bell Beaker Complex and the Early and Middle Bronze Age. Strontium isotope data show that the inhabitants followed a strictly patrilocal residential system. We demonstrate the impact of the population movement that originated in the Pontic-Caspian steppe in the 3rd millennium BCE and subsequent local developments. Utilising relatedness inference methods developed for low-coverage modern DNA we reconstruct farmstead related pedigrees and find a strong association between relatedness and grave goods suggesting that social status is passed down within families. The co-presence of biologically related and unrelated individuals in every farmstead implies a socially stratified complex household in the Central European Bronze Age.

lech-bavaria
Diminishing steppe ancestry and resurgent Neolithic ancestry over time. Alexander M. Kim

Notice how the arrival of Bell Beakers, obviously derived from Yamna settlers in Hungary, and thus clearly identified as expanding North-West Indo-Europeans all over Europe, marks a decrease in steppe ancestry compared to Corded Ware groups, in a site quite close to the most likely East BBC homeland. Copenhagen’s steppe ancestry = Indo-European going down the toilet, step by step…

UPDATES

Russian Far East populations

Gene geography of the Russian Far East populations – faces, genome-wide profiles, and Y-chromosomes, by Balanovsky et al.

Russian Far East is not only a remote area of Eurasia but also a link of the chain of Pacific coast regions, spanning from East Asia to Americas, and many prehistoric migrations are known along this chain. The Russian Far East is populated by numerous indigenous groups, speaking Tungusic, Turkic, Chukotko-Kamchatka, Eskimo-Aleut, and isolated languages. This linguistic and geographic variation opens question about the patterns of genetic variation in the region, which was significantly undersampled and received minor attention in the genetic literature to date. To fill in this gap we sampled Aleuts, Evenks, Evens, Itelmens, Kamchadals, Koryaks, Nanais, Negidals, Nivkhs, Orochi, Udegeis, Ulchi, and Yakuts. We also collected the demographic information of local populations, took physical anthropological photos, and measured the skin color. The photos resulted in the “synthetic portraits” of many studied groups, visualizing the main features of their faces.

north-eurasia

far-east-pca
Impressive North Eurasian biobank including 30,500 individual samples with broad consent, some genealogical info, phenotypic data. Alexander M. Kim

Finland AD 5th-8th c.

Sadly, no information will be shared on the session A 1400-year transect of ancient DNA reveals recent genetic changes in the Finnish population, by Salmela et al. We will have to stick to the abstract:

Objectives: Our objective was to use aDNA to study the population history of Finland. For this aim, we sampled and sequenced 35 individuals from ten archaeological sites across southern Finland, representing a time transect from 5th to 18th century.

Methods: Following genomic DNA extraction and preparation of indexed libraries, the samples were enriched for 1,2 million genomewide SNPs using in-solution capture and sequenced on an Illumina HighSeq 4000 instrument. The sequence data were then compared to other ancient populations as well as modern Finns, their geographical neighbors and worldwide populations. Authenticity testing of the data as well as population history inference were based on standard computational methods for aDNA, such as principal component analysis and F statistics.

Results: Despite the relatively limited temporal depth of our sample set, we are able to see major genetic changes in the area, from the earliest sampled individuals – who closely resemble the present-day Saami population residing markedly further north – to the more recent ancient individuals who show increased affinity to the neighboring Circum-Baltic populations. Furthermore, the transition to the present-day population seems to involve yet another perturbation of the gene pool.

So, most likely then, in my opinion – although possibly Y-DNA will not be reported – Finns were in the Classical Antiquity period mostly R1a with secondary N1c in the Circum-Baltic region (similar to modern Estonians, as I wrote recently), while Saami were probably mostly a mix of R1a-Z282 and I1 in southern Finland. That’s what the first transition after the 5th c. probably reflects, the spread of Finns (with mainly N1c lineages) to the north, while the more recent transition shows probably the introduction of North Germanic ancestry (and thus also R1b-U106, R1a-Z284, and I1 lineages) in the west.

Dairying in ancient Mongolia

The History of Dairying in ancient Mongolia, by Wilkin et al.

The use of mass spectrometry based proteomics presents a novel method for investigating human dietary intake and subsistence strategies from archaeological materials. Studies of ancient proteins extracted from dental calculus, as well as other archaeological material, have robustly identified both animal and plant-based dietary components. Here we present a recent case study using shotgun proteomics to explore the range and diversity of dairying in the ancient eastern Eurasian steppe. Contemporary and prehistoric Mongolian populations are highly mobile and the ephemerality of temporarily occupied sites, combined with the severe wind deflation common across the steppes, means detecting evidence of subsistence can be challenging. To examine the time depth and geographic range of dairy use in Mongolia, proteins were extracted from ancient dental calculus from 32 individuals spanning burial sites across the country between the Neolithic and Mongol Empire. Our results provide direct evidence of early ruminant milk consumption across multiple time periods, as well as a dramatic increase in the consumption of horse milk in the late Bronze Age. These data provide evidence that dairy foods from multiple species were a key part of subsistence strategies in prehistoric Mongolia and add to our understanding of the importance of early pastoralism across the steppe.

The confirmation of the date 3000-2700 BC for dairying in the eastern steppe further supports what was already known thanks to archaeological remains, that the pastoralist subsistence economy was brought for the first time to the Altai region by expanding late Khvalynsk/Repin – Early Yamna pastoralists that gave rise to the Afanasevo culture.

Neolithic transition in Northeast Asia

Genomic insight into the Neolithic transition peopling of Northeast Asia, by C. Ning

East Asian representing a large geographic region where around one fifth of the world populations live, has been an interesting place for population genetic studies. In contrast to Western Eurasia, East Asia has so far received little attention despite agriculture here evolved differently from elsewhere around the globe. To date, only very limited genomic studies from East Asia had been published, the genetic history of East Asia is still largely unknown. In this study, we shotgun sequenced six hunter-gatherer individuals from Houtaomuga site in Jilin, Northeast China, dated from 12000 to 2300 BP and, 3 farming individuals from Banlashan site in Liaoning, Northeast China, dated around 5300 BP. We find a high level of genetic continuity within northeast Asia Amur River Basin as far back to 12000 BP, a region where populations are speaking Tungusic languages. We also find our Compared with Houtaomuga hunter-gatherers, the Neolithic farming population harbors a larger proportion of ancestry from Houtaomuga related hunter-gathers as well as genetic ancestry from central or perhaps southern China. Our finding further suggests that the introduction of farming technology into Northeast Asia was probably introduced through demic diffusion.

A detail of the reported haplogroups of the Houtaomuga site:

houtaomuga-site-y-dna-mtdna

Y-DNA in Northeast Asia shows thus haplogroup N1b1 ~5000 BC, probably representative of the Baikal region, with a change to C2b-448del lineages before the Xiongnu period, which were later expanded by Mongols.

A study of genetic diversity of three isolated populations in Xinjiang using Y-SNP

indo-european-indo-iranian-migrations

New open access paper (in Chinese) A study of genetic diversity of three isolated populations in Xinjiang using Y-SNP, by liu et al. Acta Anthropologica Sinitica (2018)

Abstract:

The Keriyan, Lopnur and Dolan peoples are isolated populations with sparse numbers living in the western border desert of our country. By sequencing and typing the complete Y-chromosome of 179 individuals in these three isolated populations, all mutations and SNPs in the Y-chromosome and their corresponding haplotypes were obtained. Types and frequencies of each haplotype were analyzed to investigate genetic diversity and genetic structure in the three isolated populations. The results showed that 12 haplogroups were detected in the Keriyan with high frequencies of the J2a1b1 (25.64%), R1a1a1b2a (20.51%), R2a (17.95%) and R1a1a1b2a2 (15.38%) groups. Sixteen haplogroups were noted in the Lopnur with the following frequencies: J2a1 (43.75%), J2a2 (14.06%), R2 (9.38%) and L1c (7.81%). Forty haplogroups were found in the Dolan, noting the following frequencies: R1b1a1a1 (9.21%), R1a1a1b2a1a (7.89%), R1a1a1b2a2b (6.58%) and C3c1 (6.58%). These data show that these three isolated populations have a closer genetic relationship with the Uygur, Mongolian and Sala peoples. In particular, there are no significant differences in haplotype and frequency between the three isolated populations and Uygur (f=0.833, p=0.367). In addition, the genetic haplotypes and frequencies in the three isolated populations showed marked Eurasian mixing illustrating typical characteristics of Central Asian populations.

population-distribution-map
Figure 1. The populations distribution map. Left: Uluru. Center: Dali Yabuyi. Right: Kaerqu.

My knowledge of written Chinese is almost zero, so here are some excerpts with the help of Google Translate:

The source of 179 blood samples used in the study is shown in Figure 1. The Keriyan blood samples were collected from Dali Yabuyi Township, Yutian County (39 samples). The blood samples of the Lopnur people were collected from Kaerqu Township, Yuli County (64 cases); the blood samples of the Dolan people were collected from the town of Uluru, Awati County (76).

haplotype-frequency-uighur
Columns one and two are the Keriyan haplotypes and frequencies, respectively; the third and fourth columns are the Lopnur haplotypes and frequencies; the last four columns are the Daolang haplotypes and frequencies.

The composition and frequency of the Keriyan people’s haplogroup are closest to those of the Uighurs, and both Principal Component Analysis and Phylogenetic Tree Analysis show that their kinship is recent. We initially infer that the Keriyan are local desert indigenous people. They have a connection with the source of the Uighurs. Chen et al. [42] studied the patriarchal and maternal genetic analysis of the Keriyan people and found that they are not descendants of the Tibetan ethnic group in the West. The Keriyan people are a mixed group of Eastern and Western Europeans, which may originate from the local Vil group. Duan Ranhui [43] and other studies have shown that the nucleotide variability and average nucleotide differences in the Keriyan population are between the reported Eastern and Western populations. The phylogenetic tree also shows that the populations in Central Asia are between the continental lineage of the eastern population and the European lineage of the western population, and the genetic distance between the Keriyan and the Uighurs is the closest, indicating that they have a close relationship.

y-chromosome-pca

Regarding the origin of the Lopnur people, Purzhevski judged that it was a mixture of Mongolians and Aryans according to the physical characteristics of the Lopnur people. In 1934, the Sino-Swiss delegation discovered the famous burials of the ancient tombs in the Peacock River. After research, they were the indigenous people before the Loulan period; the researcher Yang Lan, a researcher at the Institute of Cultural Relics of the Chinese Academy of Social Sciences, said that the Lopnur people were descendants of the ancient “Landan survivors”. However, the Loulan people speaking an Indo-European language, and the Lopnur people speaking Uyghur languages contradict this; the historical materials of the Western Regions, “The Geography of the Western Regions” and “The Western Regions of the Ming Dynasty” record the Uighurs who lived in Cao Cao in the late 17th and early 18th centuries. Because of the occupation of the land by the Junggar nobles and their oppression, they fled. Some of them were forced to move to the Lop Nur area. There are many similar archaeological discoveries and historical records. We have no way to determine their accuracy, but they are at different times, and there is a great difference in what is heard in the same region. (…) The genetic characteristics of modern Lopnur people are the result of the long-term ethnic integration of Uyghurs, Mongols, and Europeans. This is also consistent with the similarity of the genetic structure of the Y chromosome of Lopnur in this study with the Uighurs and Mongolians. For example, the frequency of J haplogroup is as high as 59.37%, while J and its downstream sub-haplogroup are mainly distributed in western Europe, West Asia and Central Asia; the frequency of O, R haplogroup is close to that of Mongolians.

y-chromosome-frequency
1) KA: Keriya, LB: Rob, DL: Daolang, HTW: Hetian Uygur, HTWZ: and Uygur, TLFW: Turpan Uighur, HZ: Hui, HSKZ: Kazakh, WZBKZ: Wuhuan Others, TJKZ: Tajik, KEKZZ: Kirgiz, TTEZ: Tatar, ELSZ: Russian XBZ: Xibo, MGZ: Mongolian, SLZ: Salar, XJH: Xinjiang Han, GSH: Gansu Han, GDH: Guangdong Han SCH: Sichuan Han. 2) Reference population data source literature 19-22. After the population names in the table have been marked, all the shorthands in the text are referred to in this table. 3) Because the degree of haplotypes of each reference population is different to each sub-group branch, the sub-group branches under the same haplogroup are merged when the population haplogroup data is aggregated, for example: for haplogroup G Some people are divided into G1a and G2a levels, others are assigned to G1, G2, and G3, while some people can only determine G this time. Therefore, each subgroup is merged into a single group G.

According to Ming History·Western Biography, the Mongolians originated from the Mobei Plateau and later ruled Asia and Eastern Europe. Mongolia was established, and large areas of southern Xinjiang and Central Asia were included. Later, due to the Mongolian king’s struggle for power, it fell into a long-term conflict. People of the land fled to avoid the war, and the uninhabited plain of the lower reaches of the Yarkant River naturally became a good place to live. People from all over the world gathered together and called themselves “Dura” and changed to “Dang Lang”. The long-term local Uyghur exchanges that entered the southern Mongolian monks and “Dura” were gradually assimilated [44]. According to the report, locals wore Mongolian clothes, especially women who still maintained a Mongolian face [45]. In 1976, the robes and waistbands found in the ancient time of the Daolang people in Awati County were very similar to those of the ancients. Dalang Muqam is an important part of Daolang culture. It is also a part of the Uyghur Twelve Muqam, and it retains the ancient Western culture, but it also contains a larger Mongolian culture and relics. The above historical records show that the Daolang people should appear in the Chagatai Khanate and be formed by the integration of Mongolian and Uighur ethnic groups. Through our research, we also found that the paternal haplotype of the Daolang people is contained in both Uygur and Mongolian, and the main haplogroups are the same, whereas the frequencies are different (see Table 3). The principal component analysis and the NJ analysis are also the same. It is very close to the Uyghur and the Mongolian people, which establishes new evidence for the “mixed theory” in molecular genetics.

main-haplogroup-uighur
Genetic relationship between the three isolated populations: the Uygur and the Mongolian is the closest, and the main haplogroup can more intuitively compare the source composition of the genetic structure of each population. Haplogroups C, D, and O are mainly distributed in Asia as the East Asian characteristic haplogroup; haplogroups G, J, and R are mainly distributed in continental Europe, and the high frequency distribution is in Europe and Central Asia.

If the nomenclature follows a recent ISOGG standard, it appears that:

The presence of exclusively R1a-Z93 subclades and the lack of R1b-M269 samples is compatible with the expansion of R1a-Z93 into the area with Proto-Tocharians, at the turn of the 3rd-2nd millennium BC, as suggested by the Xiaohe samples, supposedly R1a(xZ93).

Now that it is obvious from ancient DNA (as it was clear from linguistics) that Pre-Tocharians separated earlier than other Late PIE peoples, with the expansion of late Khvalynsk/Repin into the Altai, at the end of the 4th millennium, these prevalent R1a (probably Z93) samples may be showing a replacement of Pre-Tocharian Y-DNA with the Andronovo expansion already by 2000 BC.

Lacking proper assessment of ancient DNA from Proto-Tocharians, this potential early Y-DNA replacement is still speculative*. However, if that is the case, I wonder what the Copenhagen group will say when supporting this, but rejecting at the same time the more obvious Y-DNA replacement in East Yamna / Poltavka in the mid-3rd millennium with incoming Corded Ware-related peoples. I guess the invention of an Indo-Tocharian group may be near…

*NOTE. The presence of R1b-M269 among Proto-Tocharians, as well as the presence of R1b-M269 among Tarim Basin peoples in modern and ancient times is not yet fully discarded. The prevalence of R1a-Z93 may also be the sign of a more recent replacement by Iranian peoples, before the Mongolian and Turkic expansions that probably brought R1b(xM269).

Also, the presence of R1b (xM269) samples in east Asia strengthens the hypothesis of a back-migration of R1b-P297 subclades, from Northern Europe to the east, into the Lake Baikal area, during the Early Mesolithic, as found in the Botai samples and later also in Turkic populations – which are the most likely source of these subclades (and probably also of Q1a2 and N1c) in the region.

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Y-DNA haplogroups of Tuvinian tribes show little effect of the Mongol expansion

uralic-turkic

Open access Estimating the impact of the Mongol expansion upon the gene pool of Tuvans, by Balanovskaya et al., Vavilov Journal of genetics and breeding (2018), 22(5):611-619.

Abstract (emphasis mine):

With a view to trace the Mongol expansion in Tuvinian gene pool we studied two largest Tuvinian clans – those in which, according to data of humanities, one could expect the highest Central Asian ancestry, connected with the Mongol expansion. Thus, the results of Central Asian ancestry in these two clans component may be used as upper limit of the Mongol influence upon the Tuvinian gene pool in a whole. According to the data of 59 Y-chromosomal SNP markers, the haplogroup spectra in these Tuvinian tribal groups (Mongush, N = 64, and Oorzhak, N = 27) were similar. On average, two-thirds of their gene pools (63 %) are composed by North Eurasian haplogroups (N*, N1a2, N3a, Q) connected with autochtonous populations of modern area of Tuvans. The Central Asian haplogroups (C2, O2) composed less then fifth part (17 %) of gene pools of the clans studied. The opposite ratio was revealed in Mongols: there were 10 % North Eurasian haplogroups and 75 % Central Asian haplogroups in their gene pool. All the results derived – “genetic portraits”, the matrix of genetic distances, the dendrogram and the multidimensional scaling plot, which mirror the genetic connections between Tuvinian clans and populations of South Siberia and East Asia, demonstrated the prominent similarity of the Tuvinian gene pools with populations from and Khakassia and Altai. It could be therefore assumed that Tuvinian clans Mongush and Oorzhak originated from autochtonous people (supposedly, from the local Samoyed and Kets substrata). The minor component of Central Asian haplogroups in the gene pool of these clans allowed to suppose that Mongol expansion did not have a significant influence upon the Tuvinan gene pool at a whole.

tuvan-clans-y-dna

Interesting excerpts:

Haplogroup C2 peaks in Central Asia (Wells et al., 2001; Zerial et al., 2003), though its variants are abundant in other peoples of Siberia and Far East. For instance, in one of Buryat clans, namely Ekhirids, hg C2 frequency is 88 % (Y-base); in Kazakhs from different regions of Kazakhstan, total occurrence of hg C2 variants averages between 17 and 81 % (Abilev et al., 2012; Zhabagin et al., 2013, 2014, 2017), in populations of the Amur River (such as Nanais, Negidals, Nivkhs, Ulchs) – between 40 and 65 %, in Evenks – up to 68 % (Y-base), in Kyrgyz people of Pamir-Alay – up to 22 %, correspondingly; of all Turkic peoples of Altai, relatively high hg C2 frequency (16 %) is detected only in Telengits (Balanovskaya et al., 2014; Balaganskaya et al., 2011a, 2016). In Tuvinian clans under the study, hg C2 frequency is rather low – 19 % in Mongush and 11 % in Oorzhak, while in Mongols it makes up almost two thirds of the entire gene pool an comprises different genetic lines (subhaplogroups).

tuvinian-y-chromosome
Y-chromosomal haplogroup spectra in gene pools of Tuvinian Oorzhak and Mongush clans and of the neighboring populations of South Siberia and Central Asia.

Haplogroup N is abundant all over North Eurasia from Scandinavia to Far East (Rootsi et al., 2007). The study on whole Y-chromosome sequencing conducted with participation of our group (Ilumäe et al., 2016) subdivided this haplogroup into several branches with their regional distribution. In gene pools of the Tuvans involved, hg N was represented by two sub-clades, namely N1a2 and N3a.

Sub-clade N1a2 peaks in populations of West Siberia (in Nganasans, frequency is 92 %) and South Siberia (in Khakas 34 %, in Tofalars 25 %) (Y-base). In Tuvans, N1a2 occurrence is nearly 16 % in Mongush and 15 % in Oorzhak clans, respectively, while in Mongols, the frequency is three times less (5 %). Hg N1a2 is supposed to display the impact of the Samoyedic component to the gene pool of Tuvinian clans (Kharkov et al., 2013).

Sub-clade N3a is major in the Oorzhak clan comprising almost half of the gene pool (45 %); it is represented by two sub-clades, namely N3a* and N3a5. The same sub-branches are specific to the Mongush clan as well, though with lower frequencies: N3a* – 9 % and N3a5 – 14 % (see Table). In Khori-Buryats from the Transbaikal region, a high frequency is observed – 82 % (Kharkov et al., 2014), while in Mongols, N3a5 occurs rather rarely (6 %). Hg N3a* was detected in populations of South Siberia only, and was widely spread in Khakas-Sagays and Shors (up to 40 %) (Ilumäe et al., 2016) (Y-base).

samoyedic
Map of distribution of Samoyedic languages (red) in the XVII century (approximate; hatching) and in the end of XX century (continuous background). Modified from Wikipedia, with the Tuva region labelled.

Within the pan-Eurasian haplogroup R1a1a, two large genetic lines (sub-haplogroups) are identified: “European” (marker M458) and “Asian” (marker Z93) the latter almost never occurring in Europe (Balanovsky, 2015) but abundant in South Siberia and northern Hindustan. In the Altai-Sayan region, high frequencies of the “Asian” branch are spread in many peoples – Shors, Tubalars, Altai-Kizhi people, Telengits, Sagays, Kyzyl Khakas, Koibals, Teleuts (Y-base) (Kharkov et al., 2009). Hg R1a1a comprises perceptible parts of gene pools of Tuvinian clans (19 % in Mongush, and 15 % in Oorzhak), though its occurrence in Mongols is much lower (6 %). Those results also count in favor of the hypothesis of autochtonous component dominance even in the gene pools of clans potentially most influenced by Mongolian ancestry. If we add R1a1a variants to the “North Eurasian” haplogroups, the “not-Central Asian” component will compose average four fifth of the entire gene pools for Tuvinian clans (in Mongush 77 %, and in Oorzhak 81 %), being only 16 % in Mongols. Such data are definitely contrary to the hypothesis of a crucial influence of the Mongol expansion upon the development of Tuvinian gene pool.

I found interesting the high proportion of R1a-Z93 subclades among Sagays in Khakhasia, which stem from a local Samoyed substratum, as described by the paper…

Featured Image: Map of Uralic and Altaic languages, from Wikipedia.

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“Steppe people seem not to have penetrated South Asia”

indo-iranian-sintashta-uralic-migrations

Open access structured abstract for The first horse herders and the impact of early Bronze Age steppe expansions into Asia from Damgaard et al. Science (2018) 360(6396):eaar7711.

Abstract (emphasis mine):

The Eurasian steppes reach from the Ukraine in Europe to Mongolia and China. Over the past 5000 years, these flat grasslands were thought to be the route for the ebb and flow of migrant humans, their horses, and their languages. de Barros Damgaard et al. probed whole-genome sequences from the remains of 74 individuals found across this region. Although there is evidence for migration into Europe from the steppes, the details of human movements are complex and involve independent acquisitions of horse cultures. Furthermore, it appears that the Indo-European Hittite language derived from Anatolia, not the steppes. The steppe people seem not to have penetrated South Asia. Genetic evidence indicates an independent history involving western Eurasian admixture into ancient South Asian peoples.

INTRODUCTION
According to the commonly accepted “steppe hypothesis,” the initial spread of Indo-European (IE) languages into both Europe and Asia took place with migrations of Early Bronze Age Yamnaya pastoralists from the Pontic-Caspian steppe. This is believed to have been enabled by horse domestication, which revolutionized transport and warfare. Although in Europe there is much support for the steppe hypothesis, the impact of Early Bronze Age Western steppe pastoralists in Asia, including Anatolia and South Asia, remains less well understood, with limited archaeological evidence for their presence. Furthermore, the earliest secure evidence of horse husbandry comes from the Botai culture of Central Asia, whereas direct evidence for Yamnaya equestrianism remains elusive.

RATIONALE
We investigated the genetic impact of Early Bronze Age migrations into Asia and interpret our findings in relation to the steppe hypothesis and early spread of IE languages. We generated whole-genome shotgun sequence data (~1 to 25 X average coverage) for 74 ancient individuals from Inner Asia and Anatolia, as well as 41 high-coverage present-day genomes from 17 Central Asian ethnicities.

damgaard-south-asia
Model-based admixture proportions for selected ancient and present-day individuals, assuming K = 6, shown with their corresponding geographical locations. Ancient groups are represented by larger admixture plots, with those sequenced in the present work surrounded by black borders and others used for providing context with blue borders. Present-day South Asian groups are represented by smaller admixture plots with dark red borders.

RESULTS
We show that the population at Botai associated with the earliest evidence for horse husbandry derived from an ancient hunter-gatherer ancestry previously seen in the Upper Paleolithic Mal’ta (MA1) and was deeply diverged from the Western steppe pastoralists. They form part of a previously undescribed west-to-east cline of Holocene prehistoric steppe genetic ancestry in which Botai, Central Asians, and Baikal groups can be modeled with different amounts of Eastern hunter-gatherer (EHG) and Ancient East Asian genetic ancestry represented by Baikal_EN.

In Anatolia, Bronze Age samples, including from Hittite speaking settlements associated with the first written evidence of IE languages, show genetic continuity with preceding Anatolian Copper Age (CA) samples and have substantial Caucasian hunter-gatherer (CHG)–related ancestry but no evidence of direct steppe admixture.

In South Asia, we identified at least two distinct waves of admixture from the west, the first occurring from a source related to the Copper Age Namazga farming culture from the southern edge of the steppe, who exhibit both the Iranian and the EHG components found in many contemporary Pakistani and Indian groups from across the subcontinent. The second came from Late Bronze Age steppe sources, with a genetic impact that is more localized in the north and west.

CONCLUSION
Our findings reveal that the early spread of Yamnaya Bronze Age pastoralists had limited genetic impact in Anatolia as well as Central and South Asia. As such, the Asian story of Early Bronze Age expansions differs from that of Europe. Intriguingly, we find that direct descendants of Upper Paleolithic hunter-gatherers of Central Asia, now extinct as a separate lineage, survived well into the Bronze Age. These groups likely engaged in early horse domestication as a prey-route transition from hunting to herding, as otherwise seen for reindeer. Our findings further suggest that West Eurasian ancestry entered South Asia before and after, rather than during, the initial expansion of western steppe pastoralists, with the later event consistent with a Late Bronze Age entry of IE languages into South Asia. Finally, the lack of steppe ancestry in samples from Anatolia indicates that the spread of the earliest branch of IE languages into that region was not associated with a major population migration from the steppe.

I think the wording of the abstract is weird, but consequent with their samples and results, so probably just clickbait / citebait for Indian journalists and social networks, or maybe a new attempt to ‘show respect for the sensibilities of Indians’ related to the artificially magnified “AIT vs. OIT” controversy, that is only present in India.

However, everything is possible, since it is brought to you by the same Danish group who proposed the Yamnaya ancestral component™, the CHG = Indo-European (and simultaneously EHG in Maykop = Anatolian??), and now also the CWC/R1a = Indo-European & Volosovo = Uralic

Here is the reaction of Narasimhan: Narasimhan has deleted the Tweet, it basically questioned the sentence that steppe people did not penetrate South Asia.

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