Eurasian steppe dominated by Iranian peoples, Indo-Iranian expanded from East Yamna

yamna-indo-iranian-expansion

The expected study of Eurasian samples is out (behind paywall): 137 ancient human genomes from across the Eurasian steppes, by de Barros Damgaard et al. Nature (2018).

Dicussion (emphasis mine):

Our findings fit well with current insights from the historical linguistics of this region (Supplementary Information section 2). The steppes were probably largely Iranian-speaking in the first and second millennia bc. This is supported by the split of the Indo-Iranian linguistic branch into Iranian and Indian33, the distribution of the Iranian languages, and the preservation of Old Iranian loanwords in Tocharian34. The wide distribution of the Turkic languages from Northwest China, Mongolia and Siberia in the east to Turkey and Bulgaria in the west implies large-scale migrations out of the homeland in Mongolia since about 2,000 years ago35. The diversification within the Turkic languages suggests that several waves of migration occurred36 and, on the basis of the effect of local languages, gradual assimilation to local populations had previously been assumed37. The East Asian migration starting with the Xiongnu accords well with the hypothesis that early Turkic was the major language of Xiongnu groups38. Further migrations of East Asians westwards find a good linguistic correlate in the influence of Mongolian on Turkic and Iranian in the last millennium39. As such, the genomic history of the Eurasian steppes is the story of a gradual transition from Bronze Age pastoralists of West Eurasian ancestry towards mounted warriors of increased East Asian ancestry—a process that continued well into historical times.

This paper will need a careful reading – better in combination with Narasimhan et al. (2018), when their tables are corrected – , to assess the actual ‘Iranian’ nature of the peoples studied. Their wide and long-term dominion over the steppe could also potentially explain some early samples from Hajji Firuz with steppe ancestry.
fku

eurasian-steppe-samples
Principal component analyses. The principal components 1 and 2 were plotted for the ancient data analysed with the present-day data (no projection bias) using 502 individuals at 242,406 autosomal SNP positions. Dimension 1 explains 3% of the variance and represents a gradient stretching from Europe to East Asia. Dimension 2 explains 0.6% of the variance, and is a gradient mainly represented by ancient DNA starting from a ‘basal-rich’ cluster of Natufian hunter-gatherers and ending with EHGs. BA, Bronze Age; EMBA, Early-to-Middle Bronze Age; SHG, Scandinavian hunter-gatherers.

For the moment, at first sight, it seems that, in terms of Y-DNA lineages:

  • R1b-Z93 (especially Z2124 subclades) dominate the steppes in the studied periods.
  • R1b-P312 is found in Hallstatt ca. 810 BC, which is compatible with its role in the Celtic expansion.
  • R1b-U106 is found in a West Germanic chieftain in Poprad (Slovakia) ca. 400 AD, during the Migration Period, hence supporting once again the expansion of Germanic tribes especially with R1b-U106 lineages.
  • A new sample of N1c-L392 (L1025) lineage dated ca. 400 AD, now from Lithuania, points again to a quite late expansion of this lineage to the region, believed to have hosted Uralic speakers for more than 2,000 years before this.
  • A sample of haplogroup R1a-Z282 (Z92) dated ca. 1300 AD in the Golden Horde is probably not quite revealing, not even for the East Slavic expansion.
  • Also, interestingly, some R1b(xM269) lineages seem to be associated with Turkic expansions from the eastern steppe dated around 500 AD, which probably points to a wide Eurasian distribution of early R1b subclades in the Mesolithic.

NOTE. I have referenced not just the reported subclades from the paper, but also (and mainly) further Y-SNP calls studied by Open Genomes. See the spreadsheet here.

Interesting also to read in the supplementary materials the following, by Michaël Peyrot (emphasis mine):

1. Early Indo-Europeans on the steppe: Tocharians and Indo-Iranians

The Indo-European language family is spread over Eurasia and comprises such branches and languages as Greek, Latin, Germanic, Celtic, Sanskrit etc. The branches relevant for the Eurasian steppe are Indo-Aryan (= Indian) and Iranian, which together form the Indo-Iranian branch, and the extinct Tocharian branch. All Indo-European languages derive from a postulated protolanguage termed Proto-Indo-European. This language must have been spoken ca 4500–3500 BCE in the steppe of Eastern Europe21. The Tocharian languages were spoken in the Tarim Basin in present-day Northwest China, as shown by manuscripts from ca 500–1000 CE. The Indo-Aryan branch consists of Sanskrit and several languages of the Indian subcontinent, including Hindi. The Iranian branch is spread today from Kurdish in the west, through a.o. Persian and Pashto, to minority languages in western China, but was in the 2nd and 1st millennia BCE widespread also on the Eurasian steppe. Since despite their location Tocharian and Indo-Iranian show no closer relationship within Indo-European, the early Tocharians may have moved east before the Indo-Iranians. They are probably to be identified with the Afanasievo Culture of South Siberia (ca 2900 – 2500 BCE) and have possibly entered the Tarim Basin ca 2000 BCE103.

The Indo-Iranian branch is an extension of the Indo-European Yamnaya Culture (ca 3000–2400 BCE) towards the east. The rise of the Indo-Iranian language, of which no direct records exist, must be connected with the Abashevo / Sintashta Culture (ca 2100 – 1800 BCE) in the southern Urals and the subsequent rise and spread of Andronovo-related Culture (1700 – 1500 BCE). The most important linguistic evidence of the Indo-Iranian phase is formed by borrowings into Finno-Ugric languages104–106. Kuz’mina (2001) identifies the Finno-Ugrians with the Andronoid cultures in the pre-taiga zone east of the Urals107. Since some of the oldest words borrowed into Finno-Ugric are only found in Indo-Aryan, Indo-Aryan and Iranian apparently had already begun to diverge by the time of these contacts, and when both groups moved east, the Iranians followed the Indo-Aryans108. Being pushed by the expanding Iranians, the Indo-Aryans then moved south, one group surfacing in equestrian terminology of the Anatolian Mitanni kingdom, and the main group entering the Indian subcontinent from the northwest.

steppe-migrations-pastoralists
Summary map. Depictions of the five main migratory events associated with the genomic history of the steppe pastoralists from 3000 bc to the present. a, Depiction of Early Bronze Age migrations related to the expansion of Yamnaya and Afanasievo culture. b, Depiction of Late Bronze Age migrations related to the Sintashta and Andronovo horizons. c, Depiction of Iron Age migrations and sources of admixture. d, Depiction of Hun-period migrations and sources of admixture. e, Depiction of Medieval migrations across the steppes.

2. Andronovo Culture: Early Steppe Iranian

Initially, the Andronovo Culture may have encompassed speakers of Iranian as well as Indo-Aryan, but its large expansion over the Eurasian steppe is most probably to be interpreted as the spread of Iranians. Unfortunately, there is no direct linguistic evidence to prove to what extent the steppe was indeed Iranian speaking in the 2nd millennium BCE. An important piece of indirect evidence is formed by an archaic stratum of Iranian loanwords in Tocharian34,109. Since Tocharian was spoken beyond the eastern end of the steppe, this suggests that speakers of Iranian spread at least that far. In the west of the Tarim Basin the Iranian languages Khotanese and Tumshuqese were spoken. However, the Tocharian B word etswe ‘mule’, borrowed from Iranian *atswa- ‘horse’, cannot derive from these languages, since Khotanese has aśśa- ‘horse’ with śś instead of tsw. The archaic Iranian stratum in Tocharian is therefore rather to be connected with the presence of Andronovo people to the north and possibly to the east of the Tarim Basin from the middle of the 2nd millennium BCE onwards110.

Since Kristiansen and Allentoft sign the paper (and Peyrot is a colleague of Kroonen), it seems that they needed to expressly respond to the growing criticism about their recent Indo-European – Corded Ware Theory. That’s nice.

They are obviously trying to reject the Corded Ware – Uralic links that are on the rise lately among Uralicists, now that Comb Ware is not a suitable candidate for the expansion of the language family.

IECWT-proponents are apparently not prepared to let it go quietly, and instead of challenging the traditional Neolithic Uralic homeland in Eastern Europe with a recent paper on the subject, they selected an older one which partially fit, from Kuz’mina (2001), now shifting the Uralic homeland to the east of the Urals (when Kuz’mina asserts it was south of the Urals).

Different authors comment later in this same paper about East Uralic languages spreading quite late, so even their text is not consistent among collaborating authors.

Also interesting is the need to resort to the questionable argument of early Indo-Aryan loans – which may have evidently been Indo-Iranian instead, since there is no way to prove a difference between both stages in early Uralic borrowings from ca. 4,500-3,500 years ago…

EDIT (10/5/2018) The linguistic supplement of the Science paper deals with different Proto-Indo-Iranian stages in Uralic loans, so on the linguistic side at least this influence is clear to all involved.

A rejection of such proposals of a late, eastern homeland can be found in many recent writings of Finnic scholars; see e.g. my references to Parpola (2017), Kallio (2017), or Nordqvist (2018).

NOTE. I don’t mind repeating it again: Uralic is one possibility (the most likely one) for the substrate language that Corded Ware migrants spread, but it could have been e.g. another Middle PIE dialect, similar to Proto-Anatolian (after the expansion of Suvorovo-Novodanilovka chiefs). I expressly stated this in the Corded Ware substrate hypothesis, since the first edition. What was clear since 2015, and should be clear to anyone now, is that Corded Ware did not spread Late PIE languages to Europe, and that some east CWC groups only spread languages to Asia after admixing with East Yamna. If they did not spread Uralic, then it was a language or group of languages phonetically similar, which has not survived to this day.

Their description of Yamna migrations is already outdated after Olalde et al. & Mathieson et al. (2018), and Narasimhan et al. (2018), so they will need to update their model (yet again) for future papers. As I said before, Anthony seems to be one step behind the current genetic data, and the IECWT seems to be one step behind Anthony in their interpretations.

At least we won’t have the Yamna -> Corded Ware -> BBC nonsense anymore, and they expressly stated that LPIE is to be associated with Yamna, and in particular the “Indo-Iranian branch is an extension of the Indo-European Yamnaya Culture (ca 3000–2400 BCE) to the East” (which will evidently show an East Yamna / Poltavka society of R1b-L23 subclades), so that earlier Eneolithic cultures have to be excluded, and Balto-Slavic identification with East Europe is also out of the way.

Related:

Ancient nomadic tribes of the Mongolian steppe dominated by a single paternal lineage

The genome of an ancient Rouran individual reveals an important paternal lineage in the Donghu population, by Li et al. Am J Phys Anthropol (2018), 1–11.

Abstract (emphasis mine):

Objectives
Following the Xiongnu and Xianbei, the Rouran Khaganate (Rouran) was the third great nomadic tribe on the Mongolian Steppe. However, few human remains from this tribe are available for archaeologists and geneticists to study, as traces of the tombs of these nomadic people have rarely been found. In 2014, the IA‐M1 remains (TL1) at the Khermen Tal site from the Rouran period were found by a Sino‐Mongolian joint archaeological team in Mongolia, providing precious material for research into the genetic imprint of the Rouran.

Materials and methods
The mtDNA hypervariable sequence I (HVS‐I) and Y‐chromosome SNPs were analyzed, and capture of the paternal non‐recombining region of the Y chromosome (NRY) and whole‐genome shotgun sequencing of TL1 were performed. The materials from three sites representing the three ancient nationalities (Donghu, Xianbei, and Shiwei) were selected for comparison with the TL1 individual.

Results
The mitochondrial haplotype of the TL1 individual was D4b1a2a1. The Y‐chromosome haplotype was C2b1a1b/F3830 (ISOGG 2015), which was the same as that of the other two ancient male nomadic samples (ZHS5 and GG3) related to the Xianbei and Shiwei, which were also detected as F3889; this haplotype was reported to be downstream of F3830 by Wei et al. (2017).

Discussion
We conclude that F3889 downstream of F3830 is an important paternal lineage of the ancient Donghu nomads. The Donghu‐Xianbei branch is expected to have made an important paternal genetic contribution to Rouran. This component of gene flow ultimately entered the gene pool of modern Mongolic‐ and Manchu‐speaking populations.

mongol-f3830-tree
The ancient males (TL1, ZHS5, and GG3) was grouped under C2b1a1b1/F3880 on the Y-DNA haplogroup C lineage using BEAST

Excerpt:

These results suggested that TL1 likely presents a close paternal relationship to the Donghu people and may have even descended from a branch of the ancient Donghu-Xianbei people, based on the conclusion that haplogroup C2b1a/F3918 can be considered the paternal branch of the ancient Donghu people (Zhang et al., 2018). The Y-chromosome phylogenetic tree showed that TL1 shared a branch with modern Mongolian-Buryats, Hezhen, Xibo, Yugur, and Kazakh, suggesting that the TL1 individual from the Rouran period should also generally present close paternal genetic relationships with modern Mongolic- and Manchu-speaking peoples.

In general, the Rouran Khaganate originated from an alliance of the ancient Eurasian steppe nomads, which disintegrated and disappeared with the progress of history. This group was complex, and its origin cannot be explained based only on one individual. However, we can trace the genetic imprint of the Rouran people through genome analysis of the TL1 individual. On the basis of the comparison with other ancient nomadic people (Donghu, Xianbei, and Shiwei) and data on modern individuals from published articles (Lippold et al., 2014; Wei et al., 2017) (Supporting Information S5), we found that they all share the same haplotype implying shared paternal ancestry between the Donghu, Xianbei and Rouran populations. Furthermore, this gene flow (mainly haplogroup C2b1a/F3918) did not stop with the disappearance of the Rouran, and a portion was instead passed on in other groups, such as the ancient Shiwei people (later than Rouran), eventually reaching the gene pool of modern Mongolic- and Manchu-speaking populations (Mongolian-Buryats, Hezhen, Xibo, et al).

Interesting to see now confirmed with ancient DNA the proposal of a C3*-DYS448del cluster as the paternal lineage defining ancient Mongolian tribes, a theory based on ancient and modern samples – since it is found in low frequency in almost all Mongolic- and Turkic-speaking populations.

This is yet another proof of how prehistoric ethnolinguistic expansions are usually accompanied by haplogroup expansion and reduction in variability.

I wonder what other ancient chiefdom-type steppe-based nomadic groups were also dominated by a single paternal lineage

Related:

How to do modern phylogeography: Relationships between clans and genetic kin explain cultural similarities over vast distances

yakut-phylogeography

A preprint paper has been published in BioRxiv, Relationships between clans and genetic kin explain cultural similarities over vast distances: the case of Yakutia, by Zvenigorosky et al (2017).

Abstract:

Archaeological studies sample ancient human populations one site at a time, often limited to a fraction of the regions and periods occupied by a given group. While this bias is known and discussed in the literature, few model populations span areas as large and unforgiving as the Yakuts of Eastern Siberia. We systematically surveyed 31,000 square kilometres in the Sakha Republic (Yakutia) and completed the archaeological study of 174 frozen graves, assembled between the 15th and the 19th century. We analysed genetic data (autosomal genotypes, Y-chromosome haplotypes and mitochondrial haplotypes) for all ancient subjects and confronted it to the study of 190 modern subjects from the same area and the same population. Ancient familial links and paternal clan were identified between graves up to 1500 km apart and we provide new data concerning the origins of the contemporary Yakut population and demonstrate that cultural similarities in the past were linked to (i) the expansion of specific paternal clans, (ii) preferential marriage among the elites and (iii) funeral choices that could constitute a bias in any ancient population study.

Even if you are not interested in the cultural and anthropological evolution of this Turkic-speaking people of the Russian Far Eastern region, the method used is an excellent example of how to use archaeology and genetics (especially Y-DNA and mtDNA data) to obtain meaningful results when investigating ancient populations.

For quite some time, probably since the first renown admixture analyses of ancient DNA samples were published, we have been living under the impression that phylogeography, or simply archaeogenetics as it was called back in the day, is not needed.

Cavalli-Sforza’s assertion that the study of modern populations could offer a clear picture of past population movements is now considered wrong, and the study of Y-DNA and mtDNA haplogroups is today mostly disregarded as of secondary importance, even among geneticists. Whole genomic investigation (and especially admixture analyses) have been leading the new wave of overconfidence in genetic results, tightly joint with the ignorance of its shortcomings (and commercial interests based on desires of ethnic identification), and haplogroups are usually just reported with other, not entirely meaningful aspects of ancient DNA analyses.

While it is undeniable that admixture analyses are offering quite interesting results, they must be carefully balanced against known archaeological and linguistic knowledge. Phylogeography – and especially Y-DNA haplogroup assessment – is quite interesting in investigating kinship and clans in patrilocal communities – i.e. most communities in prehistoric and historic periods, unless proven otherwise.

Luckily enough, there are those researchers who still strive to obtain meaningful information from haplotypes. The article referenced in this post is quite interesting due to its phylogeographic method’s applicability to ancient cultures and peoples.

When some geneticists look at simplistic prehistoric maps, like those depicting Yamna, Afanasevo, Corded Ware, and Bell Beaker cultures together, they forget that 1) cultural regions are selected more or less arbitrarily (we only have certain scattered sites for each of these cultures); 2) economic or population contacts are difficult to ascertain and to represent graphically; and 3) time periods for archaeological sites are important – in fact, they are probably THE most important aspect in assessing how accurate a map (and its “arrows” of migration or exchange) represents reality.

A careful, detailed study like this one, if applied to the Pontic-Caspian steppe, would probably reveal how R1b subclades dominated steppe clans, beginning at least during the Suvorovo-Novodanilovka expansion to the west, and certainly representing the vast majority of lineages during the internal expansion in the Early Yamna period and its later expansion east and west of the steppe…

Featured image from the article, summing up Geography, Archaeology, and Genetics of Yakutia – including Y-DNA and mtDNA haplogroups from ancient populations.

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