Interesting excerpts, referring mainly to Uralic peoples (emphasis mine):
A model-based clustering analysis using ADMIXTURE shows a similar pattern (Fig. 2b and Supplementary Fig. 3). Overall, the proportions of ancestry components associated with Eastern or Western Eurasians are well correlated with longitude in inner Eurasians (Fig. 3). Notable outliers include known historical migrants such as Kalmyks, Nogais and Dungans. The Uralic- and Yeniseian-speaking populations, as well as Russians from multiple locations, derive most of their Eastern Eurasian ancestry from a component most enriched in Nganasans, while Turkic/Mongolic speakers have this component together with another component most enriched in populations from the Russian Far East, such as Ulchi and Nivkh (Supplementary Fig. 3). Turkic/Mongolic speakers comprising the bottom-most cline have a distinct Western Eurasian ancestry profile: they have a high proportion of a component most enriched in Mesolithic Caucasus hunter-gatherers and Neolithic Iranians and frequently harbour another component enriched in present-day South Asians (Supplementary Fig. 4). Based on the PCA and ADMIXTURE results, we heuristically assigned inner Eurasians to three clines: the ‘forest-tundra’ cline includes Russians and all Uralic and Yeniseian speakers; the ‘steppe-forest’ cline includes Turkic- and Mongolic-speaking populations from the Volga and Altai–Sayan regions and Southern Siberia; and the ‘southern steppe’ cline includes the rest of the populations.
For the forest-tundra populations, the Nganasan + Srubnaya model is adequate only for the two Volga region populations, Udmurts and Besermyans (Fig. 5 and Supplementary Table 8).
For the other populations west of the Urals, six from the northeastern corner of Europe are modelled with additional Mesolithic Western European hunter-gatherer (WHG) contribution (8.2–11.4%; Supplementary Table 8), while the rest need both WHG and early Neolithic European farmers (LBK_EN; Supplementary Table 2). Nganasan-related ancestry substantially contributes to their gene pools and cannot be removed from the model without a significant decrease in the model fit (4.1–29.0% contribution; χ2 P ≤ 1.68 × 10−5; Supplementary Table 8).
NOTE. It doesn’t seem like Hungarians can be easily modelled with Nganasan ancestry, though…
For the 4 populations east of the Urals (Enets, Selkups, Kets and Mansi), for which the above models are not adequate, Nganasan + Srubnaya + AG3 provides a good fit (χ2 P ≥ 0.018; Fig. 5 and Supplementary Table 8). Using early Bronze Age populations from the Baikal Lake region (‘Baikal_EBA’; Supplementary Table 2) as a reference instead of Nganasan, the two-way model of Baikal_EBA + Srubnaya provides a reasonable fit (χ2 P ≥ 0.016; Supplementary Table 8) and the three-way model of Baikal_EBA + Srubnaya + AG3 is adequate but with negative AG3 contribution for Enets and Mansi (χ2 P ≥ 0.460; Supplementary Table 8).
Bronze/Iron Age populations from Southern Siberia also show a similar ancestry composition with high ANE affinity (Supplementary Table 9). The additional ANE contribution beyond the Nganasan + Srubnaya model suggests a legacy from ANE-ancestry-rich clines before the Late Bronze Age.
Even among the earliest available inner Eurasian genomes, east–west connectivity is evident. These, too, form a longitudinal cline, characterized by the easterly increase of a distinct ancestry, labelled Ancient North Eurasian (ANE), lowest in western European hunter-gatherers (WHG) and highest in Palaeolithic Siberians from the Baikal region. Flow-through from this ANE cline is seen in steppe populations until at least the Bronze Age, including the world’s earliest known horse herders — the Botai. However, this is eroded over time by migration from west and east, following agricultural adoption on the continental peripheries (Fig. 1b,c).
Strikingly, Jeong et al. model the modern upper steppe cline as a simple two-way mixture between western Late Bronze Age herders and Northeast Asians (Fig. 1c), with no detectable residue from the older ANE cline. They propose modern steppe peoples were established mainly through migrations post-dating the Bronze Age, a sequence for which has been recently outlined using ancient genomes. In contrast, they confirm a substantial ANE legacy in modern Siberians of the northernmost cline, a pattern mirrored in excesses of WHG ancestry west of the Urals (Fig. 1b). This marks the inhospitable biome as a reservoir for older lineages, an indication that longstanding barriers to latitudinal movement may indeed be at work, reducing the penetrance of gene flows further south along the steppe.
Given the findings as reported in the paper, I think it should be much easier to describe different subclines in the “northernmost cline” than in the much more recent “Turkic/Mongolic cline”, which is nevertheless subdivided in this paper in two clines. As an example, there are at least two obvious clines with “Nganasan-related meta-populations” among Uralians, which converge in a common Steppe MLBA (i.e. Corded Ware) ancestry – one with Palaeo-Laplandic peoples, and another one with different Palaeo-Siberian populations:
The inclusion of certain Eurasian groups (or lack thereof) in the PCA doesn’t help to distinguish these subclines visually, and I guess the tiny “Naganasan-related” ancestral components found in some western populations (e.g. the famous ~5% among Estonians) probably don’t lend themselves easily to further subdivisions. Notice, nevertheless, the different components of the Eastern Eurasian source populations among Finno-Ugrians:
Also remarkable is the lack of comparison of Uralic populations with other neighbouring ones, since the described Uralic-like ancestry of Russians was already known, and is most likely due to the recent acculturation of Uralic-speaking peoples in the cradle of Russians, right before their eastward expansions.
A comparison of Estonians and Finns with Balts, Scandinavians, and Eastern Europeans would have been more informative for the division of the different so-called “Nganasan-like meta-populations”, and to ascertain which one of these ancestral peoples along the ancient WHG:ANE cline could actually be connected (if at all) to the Cis-Urals.
Because, after all, based on linguistics and archaeology, geneticists are not supposed to be looking for populations from the North Asian Arctic region, for “Siberian ancestry”, or for haplogroup N1c – despite previous works by their peers – , but for the Bronze Age Volga-Kama region…
NOTE. I understand that writing a paper requires a lot of work, and probably statistical methods are the main interest of authors, editors, and reviewers. But it is difficult to comprehend how any user of open source tools can instantly offer a more complex assessment of the samples’ Y-SNP calls than professionals working on these samples for months. I think that, by now, it should be clear to everyone that Y-DNA is often as important (sometimes even more) than statistical tools to infer certain population movements, since admixture can change within few generations of male-biased migrations, whereas haplogroups can’t…
Srubna-Andronovo samples are as homogeneous as they always were, dominated by R1a-Z645 subclades and CWC-related (steppe_MLBA) ancestry.
The appearance of one (possibly two) R-Z280 lineages in this mixed Srubna-Alakul region of the southern Urals and this early (1880-1690 BC, hence rather Pokrovka-Alakul) points to the admixture of R1a-Z93 and R1a-Z280 already in Abashevo, which also explains the wide distribution of both subclades in the forest zones of Central Asia.
If Abashevo is the cornerstone of the Indo-Iranian / Uralic community, as it seems, the genetic admixture would initially be quite similar, undergoing in the steppes a reduction to haplogroup R1a-Z93 (obviously not complete), at the same time as it expanded to the west with Pokrovka and Srubna, and to the east with Petrovka and Andronovo. To the north, similar reductions will probably be seen following the Seima-Turbino phenomenon.
NOTE. Another R1a-Z280 has been found in the recent sample from Bronze Age Poland (see spreadsheet). As it appears right now in ancient and modern DNA, there seems to be a different distribution between subclades:
R1a-Z280 (formed ca. 2900 BC, TMRCA ca. 2600 BC) appears mainly distributed today to the east, in the forest and steppe regions, with the most ‘successful’ expansions possibly related to the spread of Abashevo- and Battle Axe-related cultures (Indo-Iranian and Uralic alike).
R1a-M458 (formed ca. 2700, TMRCA ca. 2700 BC) appears mainly distributed to the north, from central Europe to the east – but not in the steppe in aDNA, with the most ‘successful’ expansions to the west.
M458 lineages seem thus to have expanded in the steppe in sizeable numbers only after the Iranian expansions (see a map of modern R1a distributions) i.e. possibly with the expansion of Slavs, which supports the model whereby cultures from central-east Europe (like Trzciniec and Lusatian), accompanied mainly by M458 lineages, were responsible for the expansion of Proto-Balto-Slavic (and later Proto-Slavic).
The finding of haplogroup R1a-Z93, among them one Z2123, is no surprise at this point after other similar Srubna samples. As I said, the early Srubna expansion is most likely responsible for the Szólád Bronze Age sample (ca. 2100-1700 BC), and for the Balkans BA sample (ca. 1750-1625 BC) from Merichleri, due to incursions along the central-east European steppe.
Cimmerian samples from the west show signs of continuity with R1a-Z93 lineages. Nevertheless, the sample of haplogroup Q1a-Y558, together with the ‘Pre-Scythian’ sample of haplogroup N (of the Mezőcsát Culture) in Hungary ca. 980-830 BC, as well as their PCA, seem to depict an origin of these Pre-Scythian peoples in populations related to the eastern Central Asian steppes, too.
NOTE. I will write more on different movements (unrelated to Uralic expansions) from Central and East Asia to the west accompanied by Siberian ancestry and haplogroup N with the post of Ugric-Samoyedic expansions.
The Scythian of Z2123 lineage ca. 375-203 BC from the Volga (in Mathieson et al. 2015), together with the sample scy193 from Glinoe (probably also R1a-Z2123), without a date, as well as their common Steppe_MLBA cluster, suggest that Scythians, too, were at first probably quite homogeneous as is common among pastoralist nomads, and came thus from the Central Asian steppes.
The reduction in haplogroup variability among East Iranian peoples seems supported by the three new Late Sarmatian samples of haplogroup R1a-Z2124.
Approximate location of Glinoe and Glinoe Sad (with Starosilya to the south, in Ukrainian territory):
This initial expansion of Scythians does not mean that one can dismiss the western samples as non-Scythians, though, because ‘Scythian’ is a cultural attribution, based on materials. Confirming the diversity among western Scythians, a session at the recent ISBA 8:
Genetic continuity in the western Eurasian Steppe broken not due to Scythian dominance, but rather at the transition to the Chernyakhov culture (Ostrogoths), by Järve et al.
The long-held archaeological view sees the Early Iron Age nomadic Scythians expanding west from their Altai region homeland across the Eurasian Steppe until they reached the Ponto-Caspian region north of the Black and Caspian Seas by around 2,900 BP. However, the migration theory has not found support from ancient DNA evidence, and it is still unclear how much of the Scythian dominance in the Eurasian Steppe was due to movements of people and how much reflected cultural diffusion and elite dominance. We present new whole-genome results of 31 ancient Western and Eastern Scythians as well as samples pre- and postdating them that allow us to set the Scythians in a temporal context by comparing the Western Scythians to samples before and after within the Ponto-Caspian region. We detect no significant contribution of the Scythians to the Early Iron Age Ponto-Caspian gene pool, inferring instead a genetic continuity in the western Eurasian Steppe that persisted from at least 4,800–4,400 cal BP to 2,700–2,100 cal BP (based on our radiocarbon dated samples), i.e. from the Yamnaya through the Scythian period.
(…) Our results (…) support the hypothesis that the Scythian dominance was cultural rather than achieved through population replacement.
Detail of the slide with admixture of Scythian groups in Ukraine:
The findings of those 31 samples seem to support what Krzewińska et al. (2018) found in a tiny region of Moldavia-south-western Ukraine (Glinoi, Glinoi Sad, and Starosilya).
The question, then, is as follows: if Scythian dominance was “cultural rather than achieved through population replacement”…Where are the R1b-Z2103 from? One possibility, as I said in the previous post, is that they represent pockets of Iranian R1b lineages in the steppes descended from eastern Yamna, given that this haplogroup appears in modern populations from a wide region surrounding the steppes.
The other possibility, which is what some have proposed since the publication of the paper, is that they are related to Thracians, and thus to Palaeo-Balkan populations. About the previously published Thracian individuals in Sikora et al. (2014):
For the Thracian individuals from Bulgaria, no clear pattern emerges. While P192-1 still shows the highest proportion of Sardinian ancestry, K8 more resembles the HG individuals, with a high fraction of Russian ancestry.
Despite their different geographic origins, both the Swedish farmer gok4 and the Thracian P192-1 closely resemble the Iceman in their relationship with Sardinians, making it unlikely that all three individuals were recent migrants from Sardinia. Furthermore, P192-1 is an Iron Age individual from well after the arrival of the first farmers in Southeastern Europe (more than 2,000 years after the Iceman and gok4), perhaps indicating genetic continuity with the early farmers in this region. The only non-HG individual not following this pattern is K8 from Bulgaria. Interestingly, this individual was excavated from an aristocratic inhumation burial containing rich grave goods, indicating a high social standing, as opposed to the other individual, who was found in a pit.
The following are excerpts from A Companion to Ancient Thrace (2015), by Valeva, Nankov, and Graninger (emphasis mine):
Thracian settlements from the 6th c. BC on:
(…) urban centers were established in northeastern Thrace, whose development was linked to the growth of road and communication networks along with related economic and distributive functions. The early establishment of markets/emporia along the Danube took place toward the middle of the first millennium BCE (Irimia 2006, 250–253; Stoyanov in press). The abundant data for intensive trade discovered at the Getic village in Satu Nou on the right bank of the Danube provides another example of an emporion that developed along the main artery of communication toward the interior of Thrace (Conovici 2000, 75–76).
Undoubtedly the most prominent manifestation of centralization processes and stratification in the settlement system of Thrace arrives with the emergence of political capitals – the leading urban centers of various Thracian political formations.
Their relationships with Scythians and Greeks
The Scythian presence south of the Danube must be balanced with a Thracian presence north of the river. We have observed Getae there in Alexander’s day, settled and raising grain. For Strabo the coastlands from the Danube delta north as far as the river and Greek city of Tyras were the Desert of the Getae (7.3.14), notable for its poverty and tracklessness beyond the great river. He seems to suggest also that it was here that Lysimachus was taken alive by Dromichaetes, king of the Getae, whose famous homily on poverty and imperialism only makes sense on the steppe beyond the river (7.3.8; cf. Diod. 21.12; further on Getic possessions above the Danube, Paus. 1.9 with Delev 2000, 393, who seems rather too skeptical; on poverty, cf. Ballesteros Pastor 2003). This was the kind of discourse more familiarly found among Scythians, proud and blunt in the strength of their poverty. However, as Herodotus makes clear, simple pastoralism was not the whole story as one advanced round into Scythia. For he observes the agriculture practiced north and west of Olbia. These were the lands of the Alizones and the people he calls the Scythian Ploughmen, not least to distinguish them from the Royal Scythians east of Olbia, in whose outlook, he says, these agriculturalist Scythians were their inferiors, their slaves (Hdt. 4.20). The key point here is that, as we began to see with the Getan grain-fields of Alexander’s day, there was scope for Thracian agriculturalists to maintain their lifestyles if they moved north of the Danube, the steppe notwithstanding. It is true that it is movement in the other direction that tends to catch the eye, but there are indications in the literary tradition and, especially, in the archaeological record that there was also significant movement northward from Thrace across the Danube and the Desert of the Getae beyond it.
Greek literary sources were not much concerned with Thracian migration into Scythia, but we should observe the occasional indications of that process in very different texts and contexts. At the level of myth, it is to be remembered that Amazons were regularly considered to be of Thracian ethnicity from Archaic times onward and so are often depicted in Thracian dress in Greek art (Bothmer 1957; cf. Sparkes 1997): while they are most familiar on the south coast of the Black Sea, east of Sinope, they were also located on the north coast, especially east of the Don (the ancient Tanais). Herodotus reports an origin-story of the Sauromatians there, according to which this people had been created by the union of some Scythian warriors with Amazons captured on the south coast and then washed up on the coast of Scythia (4.110). While the story is unhistorical, it is not without importance. First, it reminds us that passage north from the Danube was not the only way that Thracians, Thracian influence, and Thracian culture might find their way into Scythia. There were many more and less circuitous routes, especially by sea, that could bring Thrace into Scythia. Secondly, the myth offered some ideological basis for the Sauromatian settlement in Thrace that Strabo records, for Sauromatians might claim a Thracian origin through their Amazon forebears. Finally, rather as we saw that Heracles could bring together some of the peoples of the region, we should also observe that Ares, whose earthly home was located in Thrace by a strong Greek and Roman tradition, seems also to have been a deity of special significance and special cult among the Scythians. So much was appropriate, especially from a Classical perspective, in associations between these two peoples, whose fame resided especially in their capacity for war.
This broad picture of cultural contact, interaction, and osmosis, beyond simple conflict, provides the context for a range of archaeological discoveries, which – if examined separately – may seem to offer no more than a scatter of peculiarities. Here we must acknowledge especially the pioneering work of Melyukova, who has done most to develop thinking on Thracian–Scythian interaction. As she pointed out, we have a good example of Thracian–Scythian osmosis as early as the mid-seventh century bce at Tsarev Brod in northeastern Bulgaria, where a warrior’s burial combines elements of Scythian and Thracian culture (Melyukova 1965). For, while the manner of his burial and many of the grave goods find parallels in Scythia and not Thrace, there are also goods which would be odd in a Scythian burial and more at home in a Thracian one of this period (notably a Hallstatt vessel, an iron knife, and a gold diadem). Also interesting in this regard are several stone figures found in the Dobrudja which resemble very closely figures of this kind (baby) known from Scythia (Melyukova 1965, 37–38). They range in date from perhaps the sixth to the third centuries bce, and presumably were used there – as in Scythia – to mark the burials of leading Scythians deposited in the area. Is this cultural osmosis? We should probably expect osmosis to occur in tandem with the movement of artefacts, so that only good contexts can really answer such questions from case to case. However, the broad pattern is indicated by a range of factors. Particularly notable in this regard is the observable development of a Thraco-Scythian form of what is more familiar as “Scythian animal style,” a term which – it must be understood – already embraces a range of types as we examine the different examples of the style across the great expanse from Siberia to the western Ukraine. As Melyukova observes, Thrace shows both items made in this style among Scythians and, more numerous and more interesting, a Thracian tendency to adapt that style to local tastes, with observable regional distinctions within Thrace itself. Among the Getae and Odrysians the adaptation seems to have been at its height from the later fifth century to the mid-third century (Melyukova 1965, 38; 1979).
The absence of local animal style in Bulgaria before the fifth century bce confirms that we have cultural influences and osmosis at work here, though that is not to say that Scythian tradition somehow dominated its Thracian counterpart, as has been claimed (pace Melyukova 1965, 39; contrast Kitov 1980 and 1984). Of particular interest here is the horse-gear (forehead-covers, cheek-pieces, bridle fittings, and so on) which is found extensively in Romania and Bulgaria as well as in Scythia, both in hoarded deposits and in burials. This exemplifies the development of a regional animal style, not least in silver and bronze, which problematizes the whole issue of the place(s) of its production. Accordingly, the regular designation as “Thracian” of horse-gear from the rich fourth century Scythian burial of Oguz in the Ukraine becomes at least awkward and questionable (further, Fialko 1995). And let us be clear that this is no minor matter, nor even part of a broader debate about the shared development of toreutics among Thracians and Scythians (e.g., Kitov 1980 and 1984). A finely equipped horse of fine quality was a strong statement and striking display of wealth and the power it implied
(…) while Thracian pottery appears at Olbia, Scythian pottery among Thracians is largely confined to the eastern limits of what should probably be regarded as Getic territory, namely the area close to the west of the Dniester, from the sixth century bce. Rather exceptional then is the Scythian pottery noted at Istros, which has been explained as a consequence of the Scythian pursuit of the withdrawing army of Darius and, possibly, a continued Scythian grip on the southern Danube in its aftermath (Melyukova 1965, 34). The archaeology seems to show us, therefore, that the elite Thracians and Scythians were more open to adaptation and acculturation than were their lesser brethren.
(…) we see distinct peoples and organizations, for example as Sitalces’ forces line up against the Scythians. Much more striking, however, against that general background, are the various ways in which the two peoples and their elites are seen to interact, connect, and share a cultural interface. We see also in Scyles’ story how the Greek cities on the coast of Thrace and Scythia played a significant role in the workings of relationships between the two peoples. It is not simply that these cities straddled the Danube, but also that they could collaborate – witness the honors for Autocles, ca. 300 bce (SEG 49.1051; Ochotnikov 2006) – and were implicated with the interactions of the much greater non-Greek powers around them. At the same time, we have seen the limited reality of familiar distinctions between settled Thracians and nomadic Scythians and the limited role of the Danube too in dividing Thrace and Scythia. The interactions of the two were not simply matters of dynastic politics and the occasional shared taste for artefacts like horse-gear, but were more profoundly rooted in the economic matrix across the region, so that “Scythian” nomadism might flourish in the Dobrudja and “Thracian-style” agriculture and settlement can be traced from Thrace across the Danube as far as Olbia. All of that offers scant justification for the Greek tendency to run together Thracians and Scythians as much the same phenomenon, not least as irrational, ferocious, and rather vulgar barbarians (e.g., Plato, Rep. 435b), because such notions were the result of ignorance and chauvinism. However, Herodotus did not share those faults to any degree, so that we may take his ready movement from Scythians to Thracians to be an indication of the importance of interaction between the two peoples whom he had encountered not only as slaves in the Aegean world, but as powerful forces in their own lands (e.g., Hdt. 4.74, where Thracian usage is suddenly brought into his account of Scythian hemp). Similarly, Thucydides, who quite without need breaks off his disquisition on the Odrysians to remark upon political disunity among the Scythians (Thuc. 2.97, a favorite theme: cf. Hdt. 4.81; Xen., Cyr. 1.1.4). As we have seen throughout this discussion, there were many reasons why Thracians might turn the thoughts of serious writers to Scythians and vice versa.
It seems, following Sikora et al. (2014), that Thracian ‘common’ populations would have more Anatolian Neolithic ancestry compared to more ‘steppe-like’ samples. But there were important differences even between the two nearby samples published from Bulgaria, which may account for the close interaction between Scythians and Thracians we see in Krzewińska et al. (2018), potentially reflected in the differences between the Central, Southern and the South-Central clusters (possibly related to different periods rather than peoples??).
If these R1b-Z2103 were descended from Thracian elites, this would be the first proof of Palaeo-Balkan populations showing mainly R1b-Z2103, as I expect. Their appearance together with haplogroup I2a2a1b1 (also found in Ukraine Neolithic and in the Yamna outlier from Bulgaria) seem to support this regional continuity, and thus a long-lasting cultural and ethnic border roughly around the Danube, similar to the one found in the northern Caucasus.
However, since these samples are some 2,500 years younger than the Yamna expansion to the south, and they are archaeologically Scythians, it is impossible to say. In any case, it would seem that the main expansion of R1a-Z645 lineages to the south of the Danube – and therefore those found among modern Greeks – was mediated by the Slavic expansions centuries later.
On the Northern cluster there is a sample of haplogroup R1b-P312 which, given its position on the PCA (apparently even more ‘modern Celtic’-like than the Hallstatt_Bylany sample from Damgaard et al. 2018), it seems that it could be the product of the previous eastward Hallstatt expansion…although potentially also from a recent one?:
Especially important in the archaeology of this interior is the large settlement at Nemirov in the wooded steppe of the western Ukraine, where there has been considerable excavation. This settlement’s origins evidently owe nothing significant to Greek influence, though the early east Greek pottery there (from ca. 650 bce onward: Vakhtina 2007) and what seems to be a Greek graffito hint at its connections with the Greeks of the coast, especially at Olbia, which lay at the estuary of the River Bug on whose middle course the site was located (Braund 2008). The main interest of the site for the present discussion, however, is its demonstrable participation in the broader Hallstatt culture to its west and south (especially Smirnova 2001). Once we consider Nemirov and the forest steppe in connection with Olbia and the other locations across the forest steppe and coastal zone, together with the less obvious movements across the steppe itself, we have a large picture of multiple connectivities in which Thrace bulks large.
While the above description of clear-cut R1a-Steppe and R1b-Balkans is attractive (and probably more reliable than admixture found in scattered samples of unclear dates), the true ancient genetic picture is more complicated than that:
There is nothing in the material culture of the published western Scythians to distinguish the supposed Thracian elites.
We have the sample I0575, an Early Sarmatian from the southern Urals (one of the few available) of haplogroup R1b-Z2106, which supports the presence of R1b-Z2103 lineages among Eastern Iranian-speaking peoples.
We also have DA30, a Sarmatian of I2b lineage from the central steppes in Kazakhstan (ca. 47 BC – 24 AD).
Other Sarmatian samples of haplogroup R remain undefined.
There is R1a-Z93 in a late Sarmatian-Hun sample, which complicates the picture of late pastoralist nomads further.
Therefore, the possibility of hidden pockets of Iranian peoples of R1b-Z2103 (maybe also R1b-P312) lineages remains the best explanation, and should not be discarded simply because of the prevalent haplogroups among modern populations, or because of the different clusters found, or else we risk an obvious circular reasoning: “this sample is not (autosomically or in prevalent haplogroups) like those we already had from the steppe, ergo it is not from this or that steppe culture.” Hopefully, the upcoming paper by Järve et al. will help develop a clearer genetic transect of Iranian populations from the steppes.
All in all, the diversity among western Scythians represents probably one of the earliest difficult cases of acculturation to be studied with ancient DNA (obviously not the only one), since Scythians combine unclear archaeological data with limited and conflicting proto-historical accounts (also difficult to contrast with the wide confidence intervals of radiocarbon dates) with different evolving clusters and haplogroups – especially in border regions with strong and continued interactions of cultures and peoples.
With emerging complex cases like these during the Iron Age, I am happy to see that at least earlier expansions show clearer Y-DNA bottlenecks, or else genetics would only add more data to argue about potential cultural diffusion events, instead of solving questions about proto-language expansions once and for all…
Interesting excerpts (emphasis mine, some links to images and tables deleted for clarity):
Late Bronze Age (LBA) Srubnaya-Alakulskaya individuals carried mtDNA haplogroups associated with Europeans or West Eurasians (17) including H, J1, K1, T2, U2, U4, and U5 (table S3). In contrast, the Iron Age nomads (Cimmerians, Scythians, and Sarmatians) additionally carried mtDNA haplogroups associated with Central Asia and the Far East (A, C, D, and M). The absence of East Asian mitochondrial lineages in the more eastern and older Srubnaya-Alakulskaya population suggests that the appearance of East Asian haplogroups in the steppe populations might be associated with the Iron Age nomads, starting with the Cimmerians.
#UPDATE (5 OCT 2018): Some Y-SNP calls have been published in a Molgen thread, with:
Srubna samples have possibly two R1a-Z280, three R1a-Z93.
Cimmerians may not have R1b: cim357 is reported as R1a.
Some Scythians have low coverage to the point where it is difficult to assign even a reliable haplogroup (they report hg I2 for scy301, or E for scy197, probably based on some shared SNPs?), but those which can be reliably assigned seem R1b-Z2103 [hence probably the use of question marks and asterisks in the table, and the assumption of the paper that all Scythians are R1b-L23]:
The most recent subclade is found in scy305: R1b-Z2103>Z2106 (Z2106+, Y12538/Z8131+)
scy304: R1b-Z2103 (M12149/Y4371/Z8128+).
scy009: R1b-P312>U152>L2 (P312+, U152?, L2+)?
Sarmatians are apparently all R1a-Z93 (including tem002 and tem003);
Srubnaya-Alakulskaya individuals exhibited genetic affinity to northern and northeastern present-day Europeans, and these results were also consistent with outgroup f3 statistics.
The Cimmerian individuals, representing the time period of transition from Bronze to Iron Age, were not homogeneous regarding their genetic similarities to present-day populations according to the PCA. F3 statistics confirmed the heterogeneity of these individuals in comparison with present-day populations
The Scythians reported in this study, from the core Scythian territory in the North Pontic steppe, showed high intragroup diversity. In the PCA, they are positioned as four visually distinct groups compared to the gradient of present-day populations:
A group of three individuals (scy009, scy010, and scy303) showed genetic affinity to north European populations (…).
A group of four individuals (scy192, scy197, scy300, and scy305) showed genetic similarities to southern European populations (…).
A group of three individuals (scy006, scy011, and scy193) located between the genetic variation of Mordovians and populations of the North Caucasus (…). In addition, one Srubnaya-Alakulskaya individual (kzb004), the most recent Cimmerian (cim357), and all Sarmatians fell within this cluster. In contrast to the Scythians, and despite being from opposite ends of the Pontic-Caspian steppe, the five Sarmatians grouped close together in this cluster.
A group of three Scythians (scy301, scy304, and scy311) formed a discrete group between the SC and SE and had genetic affinities to present-day Bulgarian, Greek, Croatian, and Turkish populations (…).
Finally, one individual from a Scythian cultural context (scy332) is positioned outside of the modern West Eurasian genetic variation (Fig. 1C) but shared genetic drift with East Asian populations.
The presence of an SA component (as well as finding of metals imported from Tien Shan Mountains in Muradym 8) could therefore reflect a connection to the complex networks of the nomadic transmigration patterns characteristic of seasonal steppe population movements. These movements, although dictated by the needs of the nomads and their animals, shaped the economic and social networks linking the outskirts of the steppe and facilitated the flow of goods between settled, semi-nomadic, and nomadic peoples. In contrast, all Cimmerians carried the Siberian genetic component. Both the PCA and f4 statistics supported their closer affinities to the Bronze Age western Siberian populations (including Karasuk) than to Srubnaya. It is noteworthy that the oldest of the Cimmerians studied here (cim357) carried almost equal proportions of Asian and West Eurasian components, resembling the Pazyryks, Aldy-Bel, and Iron Age individuals from Russia and Kazakhstan (12). The second oldest Cimmerian (cim358) was also the only one with both uniparental markers pointing toward East Asia. The Q1* Y chromosome sublineage of Q-M242 is widespread among Asians and Native Americans and is thought to have originated in the Altai Mountains (24)
In contrast to the eastern steppe Scythians (Pazyryks and Aldy-Bel) that were closely related to Yamnaya, the western North Pontic Scythians were instead more closely related to individuals from Afanasievo and Andronovo groups. Some of the Scythians of the western Pontic-Caspian steppe lacked the SA and the East Eurasian components altogether and instead were more similar to a Montenegro Iron Age individual (3), possibly indicating assimilation of the earlier local groups by the Scythians.
Toward the end of the Scythian period (fourth century CE), a possible direct influx from the southern Ural steppe zone took place, as indicated by scy332. However, it is possible that this individual might have originated in a different nomadic group despite being found in a Scythian cultural context.
I am surprised to find this new R1b-L23-based bottleneck in Eastern Iranian expansions so late, but admittedly – based on data from later times in the Pontic-Caspian steppe near the Caucasus – it was always a possibility. The fact that pockets of R1b-L23 lineages remained somehow ‘hidden’ in early Indo-Iranian communities was clear already since Narasimhan et al. (2018), as I predicted could happen, and is compatible with the limited archaeological data on Sintashta-Potapovka populations outside fortified settlements. I already said that Corded Ware was out of Indo-European migrations then, this further supports it.
Even with all these data coming just from a north-west Pontic steppe region (west of the Dnieper), these ‘Cimmerians’ – or rather the ‘Proto-Scythian’ nomadic cultures appearing before ca. 800 BC in the Pontic-Caspian steppes – are shown to be probably formed by diverse peoples from Central Asia who brought about the first waves of Siberian ancestry (and Asian lineages) seen in the western steppes. You can read about a Cimmerian-related culture, Anonino, key for the evolution of Finno-Permic peoples.
Also interesting about the Y-DNA bottleneck seen here is the rejection of the supposed continuous western expansions of R1a-Z645 subclades with steppe tribes since the Bronze Age, and thus a clearest link of the Hungarian Árpád dynasty (of R1a-Z2123 lineage) to either the early Srubna-related expansions or – much more likely – to the actual expansions of Hungarian tribes near the Urals in historic times.
NOTE. I will add the information of this paper to the upcoming post on Ugric and Samoyedic expansions, and the late introduction of Siberian ancestry to these peoples.
A few interesting lessons to be learned:
Remember the fantasy story about that supposed steppe nomadic pastoralist society sharing different Y-DNA lineages? You know, that Yamna culture expanding with R1b from Khvalynsk-Repin into the whole Pontic-Caspian steppes and beyond, developing R1b-dominated Afanasevo, Bell Beaker, and Poltavka, but suddenly appearing (in the middle of those expansions through the steppes) as a different culture, Corded Ware, to the north (in the east-central European forest zone) and dominated by R1a? Well, it hasn’t happened with any other steppe migration, so…maybe Proto-Indo-Europeans were that kind of especially friendly language-teaching neighbours?
Remember that ‘pure-R1a’ Indo-Slavonic society emerged from Sintashta ca. 2100 BC? (or even Graeco-Aryan??) Hmmmm… Another good fantasy story that didn’t happen; just like a central-east European Bronze Age Balto-Slavic R1a continuitydidn’t happen, either. So, given that cultures from around Estonia are those showing the closest thing to R1a continuity in Europe until the Iron Age, I assume we have to get ready for the Gulf of Finland Balto-Slavic soon.
Remember that ‘pure-R1a’ expansion of Indo-Europeans based on the Tarim Basin samples? This paper means ipso facto an end to the Tarim Basin – Tocharian artificial controversy. The Pre-Tocharian expansion is represented by Afanasevo, and whether or not (Andronovo-related) groups of R1a-Z645 lineages replaced part or eventually all of its population before, during, or after the Tocharian expansion into the Tarim Basin, this does not change the origin of the language split and expansion from Yamna to Central Asia; just like this paper does not change the fact that these steppe groups were Proto-Iranian (Srubna) and Eastern Iranian (Scythian) speakers, regardless of their dominant haplogroup.
Do you smell that fresher air? It’s the Central and East European post-Communist populist and ethnonationalist bullshit (viz. pure blondR1a-based Pan-Nordicism / pro-Russian Pan-Slavism / Pan-Eurasianism, as well as Pan-Turanism and similar crap from the 19th century) going down the toilet with each new paper.
#EDIT (5 OCT 2018): It seems I was too quick to rant about the consequences of the paper without taking into account the complexity of the data presented. Not the first time this impulsivity happens, I guess it depends on my mood and on the time I have to write a post on the specific work day…
While the data on Srubna, Cimmerians, and Sarmatians shows clearer Y-DNA bottlenecks (of R1a-Z645 subclades) with the new data, the Scythian samples remain controversial, because of the many doubts about the haplogroups (although the most certain cases are R1b-Z2103), their actual date, and cultural attribution. However, I doubt they belong to other peoples, given the expansionist trends of steppe nomads before, during, and after Scythians (as shown in statistical analyses), so most likely they are Scythian or ‘Para-Scythian’ nomadic groups that probably came from the east, whether or not they incorporated Balkan populations. This is further supported by the remaining R1b-P312 and R1b-Z2103 populations in and around the modern Eurasian steppe region.
You can find an interesting and detailed take on the data published (in Russian) at Vol-Vlad’s LiveJournal (you can read an automatic translation from Google). I think that post is maybe too detailed in debunking all information associated to the supposed Scythians – to the point where just a single sample seems to be an actual Scythian (?!) -, but is nevertheless interesting to read the potential pitfalls of the study.
Previous research at KA-5 was carried out by A. V. Epimakhov in 1994–1995 and 2002–2003 and resulted in the excavation of three Sintashta culture barrows (kurgans) that produced 35 burial pits and a reported 100 skeletons (Epimakhov, 2002, 2005; Epimakhov et al., 2005; Razhev and Epimakhov, 2004). Seven AMS radiocarbon dates on human remains from the cemetery yielded a date range of 2040–1730 cal. BC (2 sigma), which placed the cemetery within the Sintashta phase of the regional Bronze Age (Hanks et al., 2007). Twelve recently obtained AMS radiocarbon dates, taken from short-lived wood and charcoal species recovered from the Kamennyi Ambar settlement, have provided a date range of 2050–1760 cal. BC (2 sigma). Importantly, these dates confirm the close chronological relationship between the settlement and cemetery for the Middle Bronze Age phase and discount the possibility of a freshwater reservoir effect influencing the earlier dating of the human remains from the Kamennyi Ambar 5 cemetery (Epimakhov and Krause, 2013).
Sintashta cemeteries frequently yield fewer than six barrow complexes and the number of skeletons recovered represents a fraction of the total population that would have inhabited the settlements (Judd et al., 2018; Johnson and Hanks, 2012). Scholars have suggested that only members of higher status were afforded interment in these cemeteries and that principles of social organization structured placement of individuals within central or peripheral grave pits (Fig. 2) (Koryakova and Epimakhov, 2007: 75–81). In comparison with other Sintashta cemeteries that have been excavated, KA-5 provides one of the largest skeletal inventories currently available for study.
The KA-5 (MBA), Bestamak (MBA) and Lisakovsk (LBA) datasets exhibited a wide range of δ13C and δ15N values for both humans and herbivores (Figs. 5 and 6 & Table 8). This diversity in isotopic signals may be evident for a variety of reasons. For example, the range of values may be associated with a broad spectrum of C3 and C4 plant diversity in the ancient site biome or herbivore grazing patterns that included more diverse environmental niche areas in the microregion around the sampled sites. Herders also may have chosen to graze animals in niche areas due to recognized territorial boundaries between settlements and concomitant patterns of mobility. Importantly, data from Bolshekaragansky represents humans with lower δ15N values that are more closely associated with δ15N values of the sampled domestic herbivores (Fig. 6). When the archaeological evidence from associated settlement sites is considered, Bolshekaragansky, Bestamak, Lisakovsk and KA-5 have been assumed to represent populations that shared similar forms of pastoral subsistence economies with significant dietary reliance upon domesticated herbivore meat and milk. Human diets have δ13C values closely related to those of local herbivores in terms of the slope of the trendline and range of values (Fig. 6). Comparatively, the cemetery of Bolshekaragansky (associated with the Arkaim settlement) reflects individuals with trend lines closer to those of cattle and caprines and may indicate a stronger reliance on subsistence products from these species with less use of wild riverine and terrestrial resources. The site of Čiča is significantly different with elevated human δ15N isotopic values and depleted δ13C values indicative of a subsistence regime more closely associated with the consumption of freshwater resources, such as fish. The stable isotopic data in this instance is strongly supported by zooarchaeological evidence recovered from the Čiča settlement and also is indicative of significant diachronic changes from the LBA phases through the Iron Age (Fig. 6).
(…) The isotopic results from KA-5, and recent botanical and archaeological studies from the Kamennyi Ambar settlement, have not produced any evidence for the production or use of domesticated cereals. While this does not definitively answer the question as to whether Sintashta populations engaged in agriculture and/or utilized agricultural products, it does call into serious question the ubiquity of such practices across the region and correlates well with recent archaeological, bioarchaeological, and isotopic studies of human and animal remains from the Southwestern Urals region and Samara Basin (Anthony et al., 2016; Schulting and Richards, 2016). The results substantiate a broader spectrum subsistence diet that in addition to the use of domesticated animal products also incorporated wild flora, wild fauna and fish species. These findings further demonstrate the need to draw on multiple methods and datasets for the reconstruction of late prehistoric subsistence economies in the Eurasian steppes. When possible, this should include datasets from both settlements and associated cemeteries.
Variability in subsistence practices in the central steppes region has been highlighted by other scholars and appears to be strongly correlated with local environmental conditions and adaptations. More comprehensive isotopic studies of human, animal and fish remains are of fundamental importance to achieve more robust and empirically substantiated reconstructions of local biomes and to aid the refinement of regional and micro-regional economic subsistence models. This will allow for a fuller understanding of key diachronic shifts within dietary trends and highlight regional variation of such practices. Ultimately, this will more effectively index the diverse social and environmental variables that contributed to late prehistoric lifeways and the economic strategies employed by these early steppe communities.
Social organization of Sintashta-Petrovka
Interesting to remember now the recent article by Chechushkov et al. (2018) about the social stratificaton in Sintashta-Petrovka, and how it must have caused the long-lasting, peaceful admixture process that led to the known almost full replacement of R1b-L23 (mostly R1b-Z2103) by R1a-Z645 (mostly R1a-Z93) subclades in the North Caspian steppe, coinciding with the formation of the Proto-Indo-Iranian community and language (read my thoughts on this after Damgaard et al. 2018).
Here is another relevant excerpt from Chechushkov et al. (2018), translated from Russian:
The analysis suggests that the Sintashta-Petrovka societies had a certain degree of social stratification, expressed both in selective funeral rituals and in the significant difference in lifestyle between the elite and the immediate producers of the product. The data obtained during the field study suggest that the elite lived within the fortifications, while a part of the population was outside their borders, on seasonal sites, and also in stationary non-fortified settlements. Probably, traces of winter settlements can be found near the walls, while the search for summer ones is a task of a separate study. From our point of view, the elite of the early complex societies of the Bronze Age of the Eurasian steppe originated as a response to environmental challenges that created risks for cattle farming. The need to adapt the team to the harsh and changing climatic conditions created a precedent in which the settled collectives of pastoralists – hunter-gatherers could afford the content and magnificent posthumous celebration of people and their families who were not engaged in the production or extraction of an immediate product. In turn, representatives of this social group directed their efforts to the adoption of socially significant decisions, the organization of collective labor in the construction of settlement-shelters and risked their lives, acting as military leaders and fighters.
Thus, in Bronze Age steppe societies, the formation, development and decline of social complexity are directly related to the intensity of pastoralism and the development of new territories, where collectives had to survive in part a new ecological niche. At the same time, some members of the collective took upon themselves the organization of the collective’s life, receiving in return a privileged status. As soon as the conditions of the environment and management changed, the need for such functions was virtually eliminated, as a result of which the privileged members of society dissolved into the general mass, having lost their lifetime status and the right to be allocated posthumously.
Regarding the special position of the Chicha-1 samples in the change of diet and economy during the Iron Age, it is by now well known that haplogroup N must have arrived quite late to North-East Europe, and possibly not linked with the expansion of Siberian ancestry – or linked only with some waves of Siberian ancestry in the region, but not all of them. See Lamnidis et al. (2018) for more on this.
Also, the high prevalence of haplogroup N among Fennic and Siberian (Samoyedic) peoples is not related: while the latter reflects probably the native (Palaeo-Siberian) population that acquired their Uralic branch during the MLBA expansions associated with Corded Ware groups, the former points to the expansion of Fennic peoples into Saamic territory (i.e. after the Fenno-Saamic split) as the most likely period of expansion of N1c1-L392 subclades (see known recent bottlenecks among Finns, and on Proto-Finnic dialectalization).
Probably related to these late incomers are the ancient DNA samples from the Sargat culture during the Iron Age, which show the arrival of N subclades in the region, replacing most – but not all – R1a lineages (see Pilipenko et al. (2017)). Regarding the site of Chicha-1, the following are relevant excerpts about the cultural situation that could have allowed for such stepped, diachronic admixture events in Northern Eurasia, from the paper Stages in the settlement history of Chicha-1: The Results of ceramic analysis, by Molodin et al. (2008):
The stratigraphic data allows us to make the following inference: originally, the settlement was inhabited by people bearing the Late Irmen culture. Later, the people of the Baraba trend of the Suzgun culture arrived at the site (Molodin, Chemyakina, 1984: 40–62). The Baraba-Suzgun pottery demonstrates features similar to what has been reported from the sites of the transitional Bronze to Iron Age culture in the pre-taiga and taiga zones in the Irtysh basin (Potemkina, Korochkova, Stefanov, 1995; Polevodov, 2003). The major morphological types are slightly and well-profiled pots with a short throat. (…)
During the following stage of development of the site, the Chicha population increased with people who practiced cultures others than those noted in earlier collections. The ceramic materials from layer 5 provide data on possible relationships. In addition to migrants from northwestern regions practicing the Suzgun culture, there were people bearing the Krasnoozerka culture. Available data also suggests that people from the northern taiga region with the Atlym culture visited the site.
However, people from the west and southwest represent the greatest migration to the region under study. In all likelihood they moved from the northern forest-steppe zone of modern Kazakhstan and practiced the Berlik culture. The spatial distribution analysis of the Chicha-1 site suggests that the Berlik population was rather large. The Berlik people formed a single settlement with the indigenous Late Irmen people and apparently waged certain common economic activities, but preserved their own ethnic and cultural specificity (Molodin, Parzinger, 2006: 49–55). Judging by the data on the chronological sequence of deposited artifacts, migration took place roughly synchronously, hence Chicha-1 became a real cultural and economic center.
(…) In sum, the noted distribution of ceramics over the culture-bearing horizons suggests that beginning with layer 5, traditions of ceramic manufacture described above were practiced, hence the relevant population inhabited the site. Apparently, there were two predominant traditions: the local Late Irmen cultural tradition and the Berlik tradition, which was brought by the immigrants. The Late Irmen people mostly populated the citadel, while the Berlik immigrants inhabited the areas to the east and the north of the citadel.
The stratigraphic data also suggest that the Early Sargat ceramics emerged at the site likely as a part of the Late Irmen tradition (…) Early Sargat ceramics is apparently linked with the Late Irmen tradition. Artifacts associated with the Sargat culture proper have been found in several areas of Chicha-1 (e.g., in excavation area 16). However, the Sargat people appeared at the site after it had been abandoned by its previous inhabitants, and had eventually become completely desolated. This happened no earlier than the 6th cent. BC, possibly in the 5th cent. BC (in fact, the radiocarbon dates for that horizon are close to the turn of the Christian era).
This post should probably read “Consequences of Narasimhan et al. (2018),” too, since there seems to be enough data and materials published by the Copenhagen group in Nature and Science to make a proper interpretation of the data that will appear in their corrected tables.
The finding of late Khvalynsk/early Yamna migrations, identified with early LPIE migrants almost exclusively of R1b-L23 subclades is probably one of the most interesting findings in the recent papers regarding the Indo-European question.
Although there are still few samples to derive fully-fledged theories, they begin to depict a clearer idea of waves that shaped the expansion of Late Proto-Indo-European migrants in Eurasia during the 4th millennium BC, i.e. well before the expansion of North-West Indo-European, Palaeo-Balkan, and Indo-Iranian languages.
Late Khvalynsk expansions and archaic Late PIE
Like Anatolian, Tocharian has been described as having a more archaic nature than the rest of Late PIE. However, Pre-Tocharian belongs to the Late PIE trunk, clearly distinguishable phonetically and morphologically from Anatolian.
It is especially remarkable that – even though it expanded into Asia – it has more in common with North-West Indo-European, hence its classification (together with NWIE) as part of a Northern group, unrelated to Graeco-Aryan.
The linguistic supplement by Kroonen et al. accepts that peoples from the Afanasevo culture (ca. 3000-2500 BC) are the most likely ancestors of Tocharians.
NOTE. For those equating the Tarim Mummies (of R1a-Z93 lineages) with Tocharians, you have this assertion from the linguistic supplement, which I support:
An intermediate stage has been sought in the oldest so-called Tarim Mummies, which date to ca. 1800 BCE (Mallory and Mair 2000; Wáng 1999). However, also the language(s) spoken by the people(s) who buried the Tarim Mummies remain unknown, and any connection between them and the Afanasievo culture on the one hand or the historical speakers of Tocharian on the other has yet to be demonstrated (cf. also Mallory 2015; Peyrot 2017).
New samples of late Khvalynsk origin
These are are the recent samples that could, with more or less certainty, correspond to migration waves from late Khvalynsk (or early Yamna), from oldest to most recent:
The Namazga III samples from the Late Eneolithic period (in Turkmenistan), dated ca. 3360-3000 BC (one of haplogroup J), potentially showing the first wave of EHG-related steppe ancestry into South Asia. Not related to Indo-Iranian migrations.
NOTE. A proper evaluation with further samples from Narasimhan et al. (2018) is necessary, though, before we can assert a late Khvalynsk origin of this ancestry.
Afanasevo samples, dated ca. 3081-2450 BC, with all samples dated before ca. 2700 BC uniformly of R1b-Z2103 subclades, sharing a common genetic cluster with Yamna, showing together the most likely genomic picture of late Khvalynsk peoples.
NOTE 1. Anthony (2007) put this expansion from Repin ca. 3300-3000 BC, while his most recent review (2015) of his own work put its completion ca. 3000-2800. While the migration into Afanasevo may have lasted some time, the wave of migrants (based on the most recent radiocarbon dates) must be set at least before ca. 3100 BC from Khvalynsk.
NOTE 2. I proposed that we could find R1b-L51 in Afanasevo, presupposing the development of R1b-L51 and R1b-Z2103 lineages with separating clans, and thus with dialectal divisions. While finding this is still possible within Khvalynsk regions, it seems we will have a division of these lineages already ca. 4250-4000 BC, which would require a closer follow-up of the different inner late Khvalynsk groups and their samples. For the moment, we don’t have a clear connection through lineages between North-West Indo-European groups and Tocharian.
Subsequent and similar migration waves are probably to be suggested from the new sample of Karagash, beyond the Urals (attributed to the Yamna culture, hence maintaining cultural contacts after the migration waves), of R1b-Z2103 subclade, ca. 3018-2887 BC, potentially connected then to the event that caused the expansion of Yamna migrants westward into the Carpathians at the same time. Not related to Indo-Iranian migrations.
The isolated Darra-e Kur sample, without cultural adscription, ca. 2655 BC, of R1b-L151 lineage. Not related to Indo-Iranian migrations.
The Hajji Firuz samples: I4243 dated ca. 2326 BC, female, with a clear inflow of steppe ancestry; and I2327 (probably to be dated to the late 3rd millennium BC or after that), of R1b-Z2103 lineage. Not related to Indo-Iranian migrations.
NOTE. A new radiocarbon dating of I2327 is expected, to correct the currently available date of 5900-5000 BC. Since it clusters nearer to Chalcolithic samples from the site than I4243 (from the same archaeological site), it is possible that both are part of similar groups receiving admixture around this period, or maybe I2327 is from a later period, coinciding with the Iron Age sample F38 from Iran (Broushaki et al. 2016), with which it closely clusters. Also, the finding of EHG-related ancestry in Maykop samples dated ca. 3700-3000 BC (maybe with R1b-L23 subclades) offers another potential source of migrants for this Iranian group.
NOTE. Samples from Narasimhan et al. (2018) still need to be published in corrected tables, which may change the actual subclades shown here.
These late Khvalynsk / early Yamna migration waves into Asia are quite early compared to the Indo-Iranian migrations, whose ancestors can only be first identified with Volga-Ural groups of Yamna/Poltavka (ca. 3000-2400 BC), with its fully formed language expanding only with MLBA waves ca. 2300-1200 BC, after mixing with incoming Abashevo migrants.
While the authors apparently forget to reference the previous linguistic theories whereby Tocharian is more archaic than the rest of Late PIE dialects, they refer to the ca. 1,000-year gap between Pre-Tocharian and Proto-Indo-Iranian migrations, and thus their obvious difference:
The fact that Tocharian is so different from the Indo-Iranian languages can only be explained by assuming an extensive period of linguistic separation.
Potential linguistic substrates in the Middle East
A few words about relevant substrate language proposals.
What Gordon Whittaker proposes is a North-West Indo-European-related substratum in Sumerian language and texts ca. 3500 BC, which may explain some non-Sumerian, non-Semitic word forms. It is just one of many theories concerning this substratum.
This is a summary of his findings from his latest writing on the subject (a chapter of a book on Indo-European phonetics, from the series Copenhagen Studies in Indo-European):
In Sumerian and Akkadian vocabulary, the cuneiform writing system, and the names of deities and places in Southern Mesopotamia a body of lexical material has been preserved that strongly suggests influence emanating from a superstrate of Indo-European origin. his Indo-European language, which has been given the name Euphratic, is, at present, attested only indirectly through the filters of Sumerian and Akkadian. The attestations consist of words and names recorded from the mid-4th millennium BC (Late Uruk period) onwards in texts and lexical lists. In addition, basic signs that originally had a recognizable pictorial structure in proto-cuneiform preserve (at least from the early 3rd millennium on) a number of phonetic values with no known motivation in Sumerian lexemes related semantically to the items depicted. This suggests that such values are relics from the original logographic values for the items depicted and, thus, that they were inherited from a language intimately associated with the development of writing in Mesopotamia. Since specialists working on proto-cuneiform, most notably Robert K. Englund of the Cuneiform Digital Library Initiative, see little or no evidence for the presence of Sumerian in the corpus of archaic tablets, the proposed Indo-European language provides a potential solution to this problem. It has been argued that this language, Euphratic, had a profound influence on Sumerian, not unlike that exerted by Sumerian and Akkadian on each other, and that the writing system was the primary vehicle of this influence. he phonological sketch drawn up here is an attempt to chart the salient characteristics of this influence, by comparing reconstructed Indo-European lexemes with similarly patterned ones in Sumerian (and, to a lesser extent, in Akkadian).
His original model, based on phonetic values in basic proto-cuneiform signs, is quite imaginative and a very interesting read, if you have the time. His Academia.edu account hosts most of his papers on the subject.
We could speculate about the potential expansion of this substrate language with the commercial contacts between Uruk and Maykop (as I did), now probably more strongly supported because of the EHG found in Maykop samples.
NOTE. We could also put it in relation with the Anatolian language of Mari, but this would require a new reassessment of its North-West Indo-European nature.
Nevertheless, this theory is far from being mainstream, anywhere. At least today.
NOTE. The proposal remains still hypothetic, because of the flaws in the Indo-European parallels – similar to Koch’s proposal of Indo-European in Tartessian inscriptions. A comprehensive critic approach to the theory is found in Sylvie Vanséveren’s A “new” ancient Indo-European language? On assumed linguistic contacts between Sumerian and Indo-European “Euphratic”, in JIES (2008) 36:3&4.
References to Gutian are popping up related to the Hajji Firuz samples of the mid-3rd millennium.
The hypothesis was put forward by Henning (1978) in purely archaeological terms.
This is the relevant excerpt from the book:
(…) Comparativists have asserted that, in spite of its late appearance, Tokharian is a relatively archaic form of Indo-European.3 This claim implies that the speakers of this group separated from their Indo-European brethren at a comparatively early date. They should accordingly have set out on their migrations rather early, and should have appeared within the Babylonian sphere of influence also rather early. Earlier, at any rate, than the Indo-Iranians, who spoke a highly developed (therefore probably later) form of Indo-European. Moreover, as some of the Indo-Iranians after their division into Iranians and Indo-Aryans4 appeared in Mesopotamia about 1500 B.C., we should expect the Proto-Tokharians about 2000 B.C. or even earlier.
If, armed with these assumptions as our working hypothesis, we look through the pages of history, we find one nation – one nation only – that perfectly fulfills all three conditions, which, therefore, entitles us to recognize it as the “Proto-Tokharians”. Tis name was Guti; the intial is also spelled with q (a voiceless back velar or pharyngeal), but the spelling with g is the original one. The closing -i is part of the name, for the Akkadian case-endings are added to it, nom. Gutium etc. Guti (or Gutium, as some scholars prefer) was valid for the nation, considered as an entity, but also for the territory it occupied.
The text goes on to follow the invasion of Babylonia by the Guti, and further eastward expansions supposedly connected with these, to form the attested Tocharians.
Among the Gutian rulers is one Elulumesh, whose name is evidently Akkadian Elulum slightly “Gutianized” by the Gutian case(?) ending -eš.40 This Gutian ruler Elulum is obviously the same man whom we find participating in the scramble for power after the death of Shar-kali-sharrii; his name appears there in Sumerian form without mimation as Elulu.
The Gutian dynasty, from ca. 22nd c. BC appears as follows:
I don’t think we could derive a potential relation to any specific Indo-European branch from this simple suffix repeated in Gutian rulers, though.
The hypothesis of the Tocharian-like nature of the Guti (apart from the obvious error of considering them as the ancestors of Tocharians) remains not contrasted in new works since. It was cited e.g. by Gamkrelidze and Ivanov (1995) to advance their Armenian homeland, and by Mallory and Adams in their Encyclopedia (1997).
It lies therefore in the obscurity of undeveloped archaeological-linguistic hypotheses, and its connection with the attested R1b-Z2103 samples from Iran is not (yet) warranted.
Nevertheless, since we have very few samples, I think we could still see a clear genetic contribution from Yamna to Corded Ware immigrants in the North Caspian region (from Abashevo, in turn a mix of Fatyanovo/Balanovo and Catacomb/Poltavka cultures) in terms of:
Ancestral components and PCA in new Sintashta-Petrovka, Andronovo, and/or later samples – similar the ‘steppe’ drift seen in Potapovka relative to Sintashta samples, both formed by incoming Corded Ware migrants – ; and
R1b-L23 subclades, either appearing scattered during the Sintashta melting pot (of Abashevo/R1a-Z645 and East Yamna-Poltavka/R1b-Z2103 peoples), or resurging after this period, as we have seen in Pre-Balto-Slavic territory.
A lot of people seem to be looking like crazy since O&M 2018 for some sort of connection between Corded Ware and Yamna migrants in Eastern and Central Europe (wheter in SNP calls of samples published, or among almost forgotten academic papers), either to support the ideas of the 2015 papers – for those who relied on their conclusions and built (even if only mentally) far-fetched migration models around it – , or just because of some sort of absurd continuity theory involving modern R1a-Z645 subclades:
Some (the nostalgic ones?) keep looking for just one sample of R1a-Z645 in Yamna, for the same reason.
NOTE. The situation we have seen with the hundreds of samples from O&M 2018, and with the recent additional Eastern European samples, depict an unexpected absolutely clear-cut distinction in Y-DNA haplogroups between Corded Ware and Yamna/Bell Beaker: I really can’t see how the situation could be more obvious for everyone, so I doubt any further samples will make certain people change their minds. Their hope is, I guess, that just one sample may give some more oxygen to infinite pet theories, as we are still surprisingly seeing even with reactionary R1b autochthonous continuists in Western Europe…
However, looking into the most likely future for the field, what we should be expecting right now is continuity of Yamna ancestry and lineages in early Proto-Indo-Iranian territory. Since we only have a few samples from Sintashta-Petrovka, Potapovka, and Andronovo, I think there might be a sizeable number of R1b-Z2103 subclades in the territory inhabited by those who – no doubt – spread the language into Central Asia.
in the Balkans (e.g. in Vučedol or Makó-Kosihy-Čaka), including Greek and even in historical Armenian territory (potentially including Iran Iron Age sample F38 and Armenia LBA/IA RISE397, although the Mitanni may be a confounding factor here), showing the expansion of Palaeo-Balkan languages;
If we find now, as I expect, genetic continuity of east Yamna in Sintashta -> Andronovo (relative to other late Corded Ware peoples), probably including haplogroup R1b-Z2103 mixed with R1a-Z93 before its further reduction of subclades (e.g. to L657) and expansion during its subsequent spread southward…
Why exactly do we need Corded Ware to explain migrations of Late Indo-European speakers?
In other words: if we had the data we have today in 2015, would we have a need for Corded Ware to explain Indo-European migrations from the steppe? Are some people so blinded by their will to (appear to) be right in their past interpretations that they can’t just let go?
NOTE. On a side note, wouldn’t it be nice for this paper to publish some other R1b-L23 (x2103) sample – maybe even R1b-L51 – in Yamna, Andronovo, or Afanasevo territory, to end both autochthonous continuity theories (of North-Eastern and Western Europe) at the same time?
I really hope someone in David Reich’s team understands this matter, or else they will still identify Corded Ware as the (now probably ‘a’ instead) vector of expansion of Indo-European languages, and some of us will still have fun for another 2 or 3 years with such conclusions, until someone in the lab realizes that ancestry ≠ population ≠ ethnic identification ≠ language.
NOTE. It seems rather dull to read how people are discussing in the Twitterverse conventional constructs like ‘human race‘ as found in Reich’s op-ed in The New York Times, as if such grandiose semantic discussions had any practical meaning, when basic anthropological questions actually relevant for Genomics, like the essential ancestral component ≠ people tenet seem not to be of interest for anyone in the field….
Since our Indo-European demic difusion model (and its consequences for our reconstruction of North-West Indo-European) and this blog are becoming more and more popular each day – judging by the constant growth in visits in the past 6 months or so – , I guess the simplemindedness and predictability of certain geneticists is benefitting traditional anthropology directly, driving more and more amateur geneticists to look for sound academic models to answer the growing inconsistencies of genetic research.
NOTE. I am not saying the rejection of Corded Ware as spreading Indo-European is definitive. Maybe more samples within some years will depict a clear ancient expansion of Early or Middle Proto-Indo-Europeans from Khvalynsk to the forest-steppe and forest zone, and later with certain Corded Ware migrants into Central Europe, over whose territory a Late Indo-European dialect from Bell Beakers became the superstrate, as some have proposed in the past – e.g. to explain Krahe’s Old European hydronymy. I really doubt you could demonstrate such an old ethnolinguistic identification with a clear, unbroken archaeological trail, though, and we know now that this old hydronymy is probably of Late Indo-European nature (possibly even more recent).
What I am saying is: with the data we have now, it does not make any sense to keep the anthropological models invented by geneticists ex nihiloin 2015, and the hundred different alternative Late Indo-European migration models that are – born – with – each – new – paper.
These Yamna -> Corded Ware migration models didn’t have any sense for me since early 2016, but now after O&M 2017, and especially O&M 2018, I don’t think any geneticist with a little knowledge in Linguistics or Archaeology (if they are decent about their quest for truth in describing ancient European migrations) would buy them, if not for some sort of created ‘tradition’. So let’s ditch Corded Ware as Late Indo-European-speaking, let’s accept that late Corded Ware migrants should most likely be identified as early Uralic speakers, and then future data will tell if we are – again – wrong.
Please, don’t let Genomics become another pseudoscience based solely on Bioinformatics like glottochronology: let anthropologists (preferably mainstream archaeologists, but also the true Indo-Europeanists, linguists) help you interpret your raw data. Don’t deceive yourselves thinking that you have read enough about the Indo-European question, or that you know enough Indo-Europeanists (say what?) to derive your own conclusions.
Use the South Asia paper to begin expressly retracting the Corded Ware mess.
It happens so that the discussion has turned lately mainly to ancient Y-DNA haplogroups, because they help confirm previous mainstream anthropological models of cultural diffusion and migration. It is obviously not reasonable to judge prehistoric ethnolinguistic migrations from ca. 5,000 years ago based on historical nation-states and ethnic or religious concepts invented since the Middle Ages, coupled with “your” people’s main modern (or your own) paternal lineage.
EDIT (27 MAR 2018): Minor corrections and post made shorter.
It has been widely accepted that the Finno-Ugric Hungarian language, originated from proto Uralic people, was brought into the Carpathian Basin by the Hungarian Conquerors. From the middle of the 19th century this view prevailed against the deep-rooted Hungarian Hun tradition, maintained in folk memory as well as in Hungarian and foreign written medieval sources, which claimed that Hungarians were kinsfolk of the Huns. In order to shed light on the genetic origin of the Conquerors we sequenced 102 mitogenomes from early Conqueror cemeteries and compared them to sequences of all available databases. We applied novel population genetic algorithms, named Shared Haplogroup Distance and MITOMIX, to reveal past admixture of maternal lineages. Phylogenetic and population genetic analysis indicated that more than one third of the Conqueror maternal lineages were derived from Central-Inner Asia and their most probable ultimate sources were the Asian Huns. The rest of the lineages most likely originated from the Bronze Age Potapovka-Poltavka-Srubnaya cultures of the Pontic-Caspian steppe, which area was part of the later European Hun empire. Our data give support to the Hungarian Hun tradition and provides indirect evidence for the genetic connection between Asian and European Huns. Available data imply that the Conquerors did not have a major contribution to the gene pool of the Carpathian Basin, raising doubts about the Conqueror origin of Hungarian language.
Featured image, from the article: “Hypothetic origin and migration route of different components of the Hungarian Conquerors. Bluish line frames the Eurasian steppe zone, within which all presumptive ancestors of the Conquerors were found. Yellow area designates the Xiongnu Empire at its zenith from which area the East Eurasian lineages originated. Phylogeographical distribution of modern East Eurasian sequence matches (Fig. 1) well correspond to this territory, especially considering that Yakuts, Evenks and Evens lived more south in the past , and European Tatars also originated from this area. Regions where Asian and European Scythian remains were found are labeled green, pink is the presumptive range of the Srubnaya culture. Migrants of Xiongnu origin most likely incorporated descendants of these groups. The map was created using QGIS 2.18.4”.
I already wrote about the concept of outlier in Human Ancestry, so I am not going to repeat myself. This is just an update of “outliers” in recent studies, and their potential origins (here I will repeat some of the examples):
Early Khvalynsk: the three samples from the Samara region have quite different positions in PCA, from nearest to EHG (of Y-DNA haplogroup R1a) to nearest to ANE ancestry (of Y-DNA haplogroup Q). This could represent the initial consequences of the second wave of ANE ancestry – as found later in Yamna samples from a neighbouring region -, possibly brought then by Eurasian migrants related to haplogroup Q.
With only 3 samples, this is obviously just a tentative explanation of the finds. The samples can only be reasonably said to show an unstable time for the region in terms of admixture (i.e. probably migration), judging by the data on PCA.
Ukraine Eneolithic samples offer a curious example of how the concept of outlier can change radically: from the third version (May 30th) of the preprint paper of Mathieson et al. (2017), when the Ukraine Eneolithic sample with steppe ancestry (and clustering with central European samples) was the ‘outlier’, to the fourth version (September 19th), when two samples with steppe ancestry clustering close to Corded Ware samples were now the ‘normal’ ones (i.e. those representing Ukraine Eneolithic population), and the outlier was the one clustering closely with Ukraine Mesolithic samples…
This is one of the funny consequences of the wrong interpretation of the ‘yamnaya component’, that made geneticists believe at first that, out of two samples (!), the ‘outlier’ was the one with ‘yamnaya’ ancestry, because this component would have been brought by an eastern immigrant from early Khvalynsk…
West Yamna (to insist on the same question, the ‘yamnaya’ component): we have only four western Yamna samples, two of them showing Anatolian Neolithic ancestry (one of them, from Ukraine, with a strong ‘southern’ drift). On the other hand, Corded Ware migrants do not show this. So we could infer that their migrations were not coetaneous: whereas peoples of Corded Ware culture expanded ca. 3300 BC to the north – in the natural corridor to the Baltic that has been proposed for this culture in Archaeology for decades (and that is well represented by Ukraine Eneolithic samples) -, peoples of Yamna culture expanded to the west, replacing the Ukraine Eneolithic population (i.e. probably those of ‘Proto-Corded Ware culture’), and eventually mixing with Balkan populations of Anatolian Neolithic ancestry.
Potapovka, Andronovo, and Srubna: while Potapovka clusters closely to the steppe, and Andronovo (like Sintashta) clusters closely to Corded Ware (i.e. Ukraine Neolithic / Central-East European), both have certain ‘outliers’ in PCA: the former has one individual clustering closely to Corded Ware, and the latter to the steppe. Both ‘outliers’ fit well with the interpretation of the recent mixture of Corded Ware peoples with steppe populations, and they offer a different image for the evolution of populations of Potapovka and Sintashta-Petrovka, potentially influencing their language. The position of Srubna samples, nearer to Sintashta and Andronovo (but occupying the same territory as the previous Potapovka) offers the image of a late westward conquest from Corded Ware-related populations.
Iron Age Bulgaria: a sample of haplogroup R1a-z93, with more ‘yamnaya’ ancestry than any other previous sample from the Balkans. For some, it might mean continuity from an older time. However – as with the Corded Ware outlier from Esperstedt before it – it is more likely a recent migrant from the steppe. The most likely origin of this individual is therefore people from the steppe, i.e. either the Srubna culture or a related group. Its relatively close cluster in PCA to certain recent Slavic populations can be interpreted in light of the multiple back and forth migrations in the region: of steppe populations to the west (Srubna, Cimmerians, Scythians, Sarmatians,…), and of Slavic-speaking populations:
And then rapidly expanding as a Proto-Slavic-speaking community from the steppe to the west.
Well-defined outliers are, therefore, essential to understand a recent history of admixture. On the other hand, the very concept of “outlier” can be a dangerous tool – when the lack of enough samples makes their classification as as such unjustified -, leading to the wrong interpretations.