We know that the Caucasus Mountains formed a persistent prehistoric barrier to cultural and population movements. Nevertheless, an even more persistent frontier to population movements in Europe, especially since the Neolithic, is the Pontic-Caspian steppe – forest-steppe ecotone.
Like the Caucasus, this barrier could certainly be crossed, and peoples and cultures could permeate in both directions, but there have been no massive migrations through it. The main connection between both regions (steppe vs. forest-steppe/forest zone) was probably through its eastern part, through the Samara region in the Middle Volga.
The chances of population expansions crossing this natural barrier anywhere else seem quite limited, with a much less porous crossing region in the west, through the Dnieper-Dniester corridor.
A Persistent ecological and cultural frontier
It is very difficult to think about any culture that transgressed this persistent ecological and cultural frontier: many prehistoric and historical steppe pastoralists did appear eventually in the neighbouring forest-steppe areas during their expansions (e.g. Yamna, Scythians, or Turks), as did forest groups who permeated to the south (e.g. Comb Ware, GAC, or Abashevo), but their respective hold in foreign biomes was mostly temporary, because their cultures had to adapt to the new ecological environment. Most if not all groups originally from a different ecological niche eventually disappeared, subjected to renewed demographic pressure from neighbouring steppe or forest populations…
Before the emergence of pastoralism, the cultural contacts of the Pontic region (i.e. forest-steppes) with the Baltic were intense. In fact, the connection of the north Pontic area with the Baltic through the Dnieper-Dniester corridor and the Podolian-Volhynian region is essential to understand the spread of peoples of post-Maglemosian and post-Swiderian cultures (to the south), hunter-gatherer pottery (to the north), TRB (to the south), Late Trypillian groups (north), GAC (south), or Comb Ware (south) (see here for Eneolithic movements), and finally steppe ancestry and R1a-Z645 with Corded Ware (north). After the complex interaction of TRB, Trypillia, GAC, and CWC during the expansion of late Repin, this traditional long-range connection is lost and only emerges sporadically, such as with the expansion of East Germanic tribes.
A barrier to steppe migrations into northern Europe
One may think that this barrier was more permeable, then, in the past. However, the frontier is between steppe and forest-steppe ecological niches, and this barrier evolved during prehistory due to climate changes. The problem is, before the drought that began ca. 4000 BC and increased until the Yamna expansion, the steppe territory in the north Pontic region was much smaller, merely a strip of coastal land, compared to its greater size ca. 3300 BC and later.
This – apart from the cultural and technological changes associated with nomadic pastoralism – justifies the traditional connection of the north Pontic forest-steppes to the north, broken precisely after the expansion of Khvalynsk, as the north Pontic area became gradually a steppe region. The strips of north Pontic and Azov steppes and Crimea seem to have had stronger connections to the Northern Caucasus and Northern Caspian steppes than with the neighbouring forest-steppe areas during the Upper Palaeolithic, Mesolithic, and Neolithic.
NOTE. We still don’t know the genetic nature of Mikhailovka or Ezero, steppe-related groups possibly derived from Novodanilovka and Suvorovo close to the Black Sea (which possibly include groups from the Pannonian plains), and how they compare to neighbouring typically forest-steppe cultures of the so-called late Sredni Stog groups, like Dereivka or partly Kvityana.
Despite the Pontic-Caspian steppes and forest-steppes neighbouring each other for ca. 2,000 km, peoples from forested and steppe areas had an obvious advantage in their own regions, most likely due to the specialization of their subsistence economy. While this is visible already in Palaeolithic and Mesolithic hunter-gatherers, the arrival of the Neolithic package in the Pontic-Caspian region incremented the difference between groups, by spreading specialized animal domestication. The appearance of nomadic pastoralism adapted to the steppe, eventually including the use of horses and carts, made the cultural barrier based on the economic know-how even stronger.
Even though groups could still adapt and permeate a different territory (from steppe to forest-steppe/forest and vice-versa), this required an important cultural change, to the extent that it is eventually complicated to distinguish these groups from neighbouring ones (like north-west Pontic Mesolithic or Neolithic groups and their interaction with the steppes, Trypillia-Usatovo, Scythians-Thracians, etc.). In fact, this steppe – forest-steppe barrier is also seen to the east of the Urals, with the distinct expansion of Andronovo and Seima-Turbino/Andronovo-like horizons, which seem to represent completely different ethnolinguistic groups.
As a result of this cultural and genetic barrier, like that formed by the Northern Caucasus:
1) No steppe pastoralist culture (which after the emergence of Khvalynsk means almost invariably horse-riding, chariot-using nomadic herders who could easily pasture their cows in the huge grasslands without direct access to water) has ever been successful in spreading to the north or north-west into northern Europe, until the Mongols. No forest culture has ever been successful in expanding to the steppes, either (except for the infiltration of Abashevo into Sintashta-Potapovka).
2) Corded Ware was not an exception: like hunter-gatherer pottery before it (and like previous population movements of TRB, late Trypillia, GAC, Comb Ware or Lublin-Volhynia settlers) their movements between the north Pontic area and central Europe happened through forest-steppe ecological niches due to their adaptation to them. There is no reason to support a direct connection of CWC with true steppe cultures.
3) The so-called “Steppe ancestry” permeated the steppe – forest-steppe ecotone for hundreds of years during the 5th and early 4th millennium BC, due to the complex interaction of different groups, and probably to the aridization trend that expanded steppe (and probably forest-steppe) to the north. Language, culture, and paternal lineages did not cross that frontier, though.
EDIT (4 FEB 2019): Wang et al. is out in Nature Communications. They deleted the Yamna Hungary samples and related analyses, but it’s interesting to see where exactly they think the trajectory of admixture of Yamna with European MN cultures fits best. This path could also be inferred long ago from the steppe connections shown by the Yamna Hungary -> Bell Beaker evolution and by early Balkan samples:
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.
Interesting excerpts (emphasis mine, references have been deleted for clarity):
Ancient DNA was extracted from the Corded Ware culture individuals excavated in southeastern Poland (N = 12) and Moravia (N = 3). Late Eneolithic (N = 5) and Bronze Age human remains (N = 25) originated from western Ukraine and came from the Yampil barrow cemetery complex located in the north–western region of the Black Sea. Bronze Age individuals were associated with different archaeological cultures, including Yamnaya (N = 14), Catacomb (N = 2), Babyno (N = 4) and Noua (N = 5).
The PCA results described 50.62% of the variability and were combined with the k-means clustering (with the k value of 5 as the best representation of the data, at the average silhouette of 0.2608). Based on these results individuals associated with the western and eastern Yamnaya horizon (YAE and YAW in Fig. 2) were grouped within a cluster consisting of populations from central Eurasia and Europe (blue cluster) including people associated with eastern Corded Ware culture (CWPlM) and Baltic Corded Ware culture (CWBal). This cluster did not contain any populations linked with early Neolithic farmers (red), or hunter-gatherers (green and yellow). On the other hand, k-means clustering linked the western Corded Ware culture-associated population (CWW) with Near East and Neolithic farmer ancestry groups from western and central Europe.
Pairwise mtDNA-based FST values, visualized on MDS using the raw non-linearized FST (stress value = 0.099) (Fig. 4), also supported the PCA results and indicated that western and eastern Yamnaya horizon groups (YAW and YAE) were closer to people associated with the eastern Corded Ware culture (CWPlM) (FST = 0.00; FST = 0.01, respectively; both p > 0.05) and Baltic Corded Ware culture (CWBal) (FST = 0.00; FST = 0.00, respectively; both p > 0.05), than to populations associated with the western Corded Ware culture (CWW) (FST = 0.047 and FST = 0.059, respectively; both statistically significant p < 0.05). Western and eastern Yamnaya horizon groups also showed close genetic affinity to the Iron Age western Scythians (SCU) (FST = 0.0022 and FST = 0.006, respectively, both p > 0.05). The most distant populations to the Yamnaya horizon groups were western hunter-gatherers (HGW) (FST = 0.23 and FST = 0.15, p < 0.001).
The FST-based MDS reflected the general European population history in the post-LGM period as the three highest FST scores were detected between western hunter-gatherers (HGW) and people associated with Linear Pottery culture (LBK) (FST = 0.33, p < 0.001), between eastern hunter-gatherers (HGE) and Baltic hunter-gatherers (HGBal) (FST = 0.35, p < 0.05), and between western (HGW) and eastern hunter-gatherers (HGE) (FST = 0.36, p < 0.05). The Yamnaya horizon groups (YAE and YAW) were placed centrally between northern hunter-gatherers (HGN) and Neolithic farmers (LDN), in direct proximity to the Bronze and Iron Age populations from Eastern Europe (SCU, BARu, SRU) and close to individuals associated with eastern and Baltic Corded Ware culture.
Among the analyzed samples, we identified two Catacomb culture-associated individuals (poz220 and poz221) belonging to hg X4. They are the first ancient individuals assigned to this particular lineage. Haplogroup X4 is rare among present day populations and has been found only in one individual each from Central Europe, Balkans, Anatolia and Armenia.
Moreover, we have reported mtDNA haplotypes that might be associated with the migration from the steppe and point to genetic continuity in the north Pontic region from Bronze Age until the Iron Age. These haplotypes were assigned to hgs U5, U4, U2 and W3. MtDNA hgs U5a and U4, identified in this study among Yamnaya, Late Eneolithic and Corded Ware culture-associated individuals, have previously been found in high frequencies among northern and eastern hunter-gatherers. Moreover, they appeared in the north Pontic region in populations associated with Mesolithic (hg U5a), Eneolithic (Post-Stog) (hg U4), Yamnaya (hgs U5, U5a), Catacomb (hgs U5 and U5a) and Iron Age Scythians (hg U5a), suggesting genetic continuity of these particular mtDNA lineages in the Pontic region from, at least, the Bronze Age. Hgs U5a and U4-carrying populations were also present in the eastern steppe, along with individuals from the Yamnaya culture from Samara region, the Srubnaya and the Andronovo from Russia. Interestingly, hg U4c1 found in the Yamnaya individual (poz224) has so-far been found only in two Bell Beaker- associated individuals and one Late Bronze Age individual from Armenia, which might suggest a steppe origin for hg U4c1. A steppe origin can possibly also be assigned to hg U4a2f, found in one individual (poz282) but not reported in any other ancient populations to date, and to U5a1- the ancestral lineage of U5a1b, reported for individual poz232, which was identified not only in Corded Ware culture-associated population from central and eastern Europe, but also in representatives of Catacomb culture from the north Pontic region, Yamnaya from Bulgaria and Russia, Srubnaya and Andronovo-associated groups. Hg U2e, reported for Late Eneolithic individual (poz090), was also identified in western Corded Ware culture-associated individual and in succeeding Sintashta, Potapovka and Andronovo groups, suggesting possible genetic continuity of U2e1 in the western part of the north Pontic region.
Hgs W3a1 and W3a1a, found in two Yamnaya individuals from this study (poz208 and poz222), were also identified in Yamnaya-associated individuals from the Russia Samara region and later in Únětice and Bell Beaker groups from Germany, supporting the idea of an eastern European steppe origin of these haplotypes and their contribution to the Yamnaya migration toward the central Europe. The W3a1 lineage was not identified in Neolithic times and, thus, we assume that it appeared in the steppe region for the first time during the Bronze Age. Notably, hgs W1 and W5, which predate the Bronze Age in Europe, were found only in individuals associated with the early Neolithic farmers from Starčevo in Hungary (hg W5), early Neolithic farmers from Anatolia (hg W1-T119C), and from the Schöningen group (hg W1c) and Globular Amphora culture from Poland (hg W5).
The most recent radiocarbon dates show that Early Yamna expanded to the west with Repin settlers of the Lower Don ca. 3350/3300 BC. At the end of the 4th millennium, then, these settlers dominated over groups whose population had in turn also elevated ‘steppe ancestry’ (at least from ca. 4000 BC, as shown by Ukraine Eneolithic samples from the forest-zone), and probably replaced the male population completely, as evidenced by other Yamna and Poltavka, and later Bell Beaker, Catacomb, and Sintashta samples.
The ‘second wave’ of expansion of Yamna settlers to the west, into east-central European steppes, began probably ca. 3100/3000 BC, and – based on material culture – stemmed mainly from the North Pontic area. The Yampil Barrow Complex on the Dnieper (which I recently wrote about) seems to be part of one of the groups of western Yamna migrants: those who migrated westward from the left bank of the Dniester to the west into the Prut-Siret region, and north along the Prut.
This region is the key for population movements that gave rise to the Corded Ware culture (see another recent post on Corded Ware origins). It is quite likely that we will see a dance of late Trypillia / Usatovo, GAC, (Proto-)Corded Ware, and Yamna samples in this area. Judging by the clear-cut Y-DNA bottlenecks we are seeing in Neolithic populations, especially among steppe pastoralists, the difference between groups in recovered ancient samples will not only be clear from their culture, but also from their male lineages.
Based on the number of burials studied from the different settlement regions for West Yamna migrants, the Prut-Siret group was one of the smallest new Yamna ‘provinces’ in south-eastern Europe, and was probably overrun early, although – since kurgan findings continued into the Catacomb culture in the Yampil complex – the Dnieper region was well-enough connected to the core North Pontic area to be kept into its retreating territory by 2500 BC, as was the Danube delta, in contrast with other east-central European areas.
Taking into account that the earliest Corded Ware burials are from ca. 2900 BC (in the Single Grave culture), and that the earliest A-horizon pottery expanded from Lesser Poland (a syncretic pottery based on the previous GAC-type) a century later, it is likely that what this paper shows for Corded Ware in eastern Europe and the Baltic is what I have suggested many times (see here, or here) as the most likely reason for elevated steppe ancestry (and close PCA cluster) of the Baltic LN ‘outliers’: the exogamy of Corded Ware groups with females from Yamna or a North Pontic steppe culture with similar ancestry.
If Proto-Corded Ware populations of the North Pontic region did not have an identical “steppe ancestry” to these eastern CW groups already during the Eneolithic (which is the other possibility), I might be right in their more recent exogamy, and this could be seen in this study by the close cluster of east Corded Ware (especially Baltic) mtDNA to GAC and Yamna West groups, and distant from previous hunter-gatherer populations of the area, which suggests that expanding males from the Volhynia/Podolia region practiced exogamy mainly with southern groups.
I think this is probably related to demographic pressure imposed on other populations by the explosive expansion of pastoralists with their new subsistence economy (part of the “Secondary Products Revolution”), which the hunter-gatherer and farmer population of Europe could not keep up with (as seen later in the admixture of expanding East Bell Beakers), although studies on European prehistoric demography are scarce and too general to tell us anything relevant for this precise period and region.
The test trenches at Nadir Tepesi suggest that the Kura-Araxes occupation ended abruptly in the mid third millennium B.C.E. and that the site was then occupied or visited by a new group of people with new cultural traditions. Evidence for a significant destruction followed by the sharp discontinuity in the material culture could represent a violent termination of the Kura-Araxes occupation at Nadir Tepesi. This possibility provides one hypothesis for the end of the Kura-Araxes culture elsewhere as well in the Mughan Steppe.
It appears that there is no subsequent substantial built settlement until possibly the Late Iron Age in the region. Our intensive and extensive surveys on the Mughan Steppe did not provide evidence for settlements until long after the Kura-Araxes time. For whatever reason, settlements on the Mughan Steppe seem to have reappeared only in the Iron Age and remained sparse until the Sassanian period in late antiquity.45 Although some ceramics with parallels in the Middle and Late Bronze Age and the Iron Age were found at a few sites, they do not seem to represent settlements.
Indeed, except for the sites that may possibly contain burials, we do not know much about the Middle and Late Bronze Ages through the Iron Age in the Mughan Steppe. Similarly, archaeological investigations in the southern Caucasus do not provide information on settlements in the Middle Bronze Age.46 From a broad perspective, the abrupt and possibly violent end to the Kura-Araxes occupation at Nadir Tepesi, together with the sudden disappearance of the Kura-Araxes settlements and the scarcity of post–Kura-Araxes sites in the Mughan Steppe,47 may indicate that these changes were part of a larger phenomenon. This evidence could suggest a major sociocultural and demographic transformation at a regional level, at least in the western Caspian littoral plain, in the middle of the third millennium B.C.E. Other archaeological investigations in the southern Caucasus portray a similar picture, that of newcomers with a significantly different lifestyle and means of subsistence possibly associated with a mobile economy. Except in some elements of the ceramic traditions, evidence of continuity of Kura-Araxes traditions and their coexistence with newcomers is scarce and uncertain.48
On one hand, Puturidze argues that there is no evidence supporting the notion of a migration of people into the southern Caucasus.50 Rather, she associates all the changes in the post–Kura-Araxes period with influences from Near Eastern societies as a result of developing interactions by the end of the third millennium B.C.E. On the other hand, Kohl hypothesizes the possibility of a “push-pull process”51 in which new groups of people with wheeled carts and oxen-pulled wagons gradually moved from the steppes of the north into the southern Caucasus, and the Kura-Araxes communities subsequently moved farther south.52.
Kohl also reminds us of the evidence of increased militarism from the Early to the Late Bronze Age that is reflected in more fortified sites, new weaponry, and an iconography of war as seen on the Karashamb Cup.53 The appearance of defensive mechanisms such as fortification walls, which can be seen at Köhne Shahar, a Kura-Araxes settlement near Chaldran in Iranian Azerbaijan, further emphasizes the increase of intergroup conflicts and militarism during the Early Bronze Age, before the Kura-Araxes culture came to an end.54 Kohl argues that, while the number of Kura-Araxes settlements decreased in the southern Caucasus, archaeological research indicates that the Kura-Araxes culture spread to western Iran in the Zagros region and to the Levant.55 In Kohl’s view, as new groups of people moved in, the Kura-Araxes communities abandoned the southern Caucasus and moved farther south, where some of them already resided. Although some scholars suggest the possible movement of new groups of people from the northern steppes to the southern Caucasus,56 others associate the cultures of the post–Kura-Araxes period, especially the Trialeti.
We believe that the evidence supports a less uniform scenario. The Kura-Araxes culture may have disappeared in various ways; the transition to the post–Kura-Araxes time may not be explained by a single model. Different Kura-Araxes settlements may have ended differently. The evidence from Nadir Tepesi could support a violent end at that site, and it is possible that similar evidence will be found at other sites in the Mughan Steppe. At some sites, such as Köhne Tepesi in the Khoda Afarin Plain,58 the Kura-Araxes occupation also ended abruptly but without any sign of destruction. In other regions, there may be evidence supporting the coexistence of newcomers with Kura-Araxes communities for some period.59 The results from Gegharot60 in Armenia and recent excavations by one of the authors of this report at Köhne Shahar, do not support any of these models. At Köhne Shahar, the Kura-Araxes culture ended around the middle of the third millennium B.C.E.61 In the last phase of Kura-Araxes occupation at the site, six storage jars in one of the workshop units stood intact, five of them still carefully covered by stone slabs. The evidence from Köhne Shahar may point to a nonviolent end or a planned abandonment of the site.62
The picture continues to be somehow blurred for what happened in the Caucasus and North Iran after the Late Indo-European expansions, due to contradictory information.
With the analysis of the dataset from Narasimhan et al. (2018), it seemed that steppe peoples might have migrated into North Iran after the first Khvalynsk/Repin or Early Yamna expansions, because some samples from North Iran were reported to have steppe-related admixture.
NOTE. As I already said, the Hajji Firuz sample of R1b-Z2103 subclade (of uncertain date) clusters closer to the Iron Age sample F38 from Iran (Broushaki et al. 2016), of the same subclade, which is quite likely related to Proto-Armenian speakers, so it is possible that both belong to the same, Late Bronze Age / Iron Age group of migrants.
The other possibility, since it also clusters at a certain distance from the Hajji Firuz I4243 ‘outlier’, dated ca. 2326 BC (from the same archaeological site as other Chalcolithic samples, but being an intrusive Bronze Age burial), is that the Hajji Firuz sample is related to these hypothetical early migrations described here; or, that its date of I4243 is also not reliable…
These initial reports, coupled with archaeological descriptions of potential migrants from the steppe ending the Kura-Araxes culture, may suggest that peoples of steppe origin (or peoples with steppe admixture from the Caucasus) occupied territories further to the south (see here for potential early migration waves).
However, studies of samples from the Caucasus in Wang et al. (2018) have shown that no migrations related to EHG ancestry happened to the south, and that the minimal EHG/WHG contribution in Kura-Araxes individuals is probably part of the Anatolian farmer-related ancestry, and not from the steppe.
In fact, further contribution from Iran Chalcolithic-related ancestry was found intruding to the north during the Early Bronze Age, into Kura-Araxes and Maykop-Novosvobodnaya samples. In the Middle Bronze Age, some peoples from the North Caucasus show steppe ancestry (further to the south than the first steppe ancestry incursions of the North Caucasus piedmont), but most late Caucasus groups studied retain the ‘southern’ Armenian/Iran Chalcolithic profile.
All this casts doubts on the whole idea of intrusive steppe ancestry found in Iran Chalcolithic and Early Bronze Age (or, alternatively, on the proper dates of the Hajji Firuz ‘outliers’).
Also, the archaeological discontinuity in the region until the Iron Age, and the close relationship of Armenian to Greek (relative to other Palaeo-Balkan languages, which seem to have expanded to the south-west with Yamna settlers), does not support these hypothetical early steppe migrants as the Proto-Armenian community; earlier migrations of LPIE speakers without known modern descendants are obviously possible, but no clear archaeological or linguistic link has been offered to date to support this.
Until we have more samples with a clear archaeological and chronological context from Anatolia, Iran, and the Armenian Highlands during the Bronze and Iron Ages, and until they show clear steppe ancestry assessed in peer-reviewed papers (or at least thoroughly contrasted with other potential sources of such ancestry, and with solid statistical results), the question of intrusive steppe ancestry, and thus maybe LPIE-speakers (which may or may not be associated with Proto-Armenians) remains open.
NOTE. The Armenian question remains open, not because genetics has precedence over linguistics, but because the linguistic classification and date of separation, in this case, is not clear, and may be quite old. The fact that Palaeo-Balkan and Pre-Indo-Iranian might have separated quite early within the Khvalynsk – Volga-Ural (Early Yamna) community adds to the difficulty in assessing migration routes, although I do believe that the close similarity of Armenian with Greek among Palaeo-Balkan languages do not warrant such an early separation, and the Middle to Late Bronze Age period in the Balkans and Anatolia offers a better route for this expansion.
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.
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 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.
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.
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…
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.
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.
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”.
– There is a Yamna cultural-historical community (i.e. with a potential ethnolinguistic unity). Although many different inner groups can be distinguished (based on cultural, social, anthropological differences), one cannot divide the culture in distinct cultures.
– The Budzhak culture is likely the heir of Repin, which is compatible with its expansion westward. According to Klejn and Anthony (Usatovo), this region was connected to (and might have influenced) the Corded Ware culture. Therefore – that is my contribution, not theirs – a hidden community of R1a-M417 subclades (that a lot of people are eager to find) might have stemmed from there.
– Globular Amphorae and Corded Ware cultures are related, because GAC is actually not a uniform culture, but a ‘complex’ (i.e., in the same sense that Bell Beaker was not a culture, but a complex, as genetics has shown), and CWC is also a complex of cultures, as supported by Furholt. If that is true, the sampling of certain peoples classified as from Globular Amphorae – which some have rushed to cite as the end of the GAC-CWC connection might not be the last word, and Kristiansen’s model of long-lasting GAC-CWC connection may still be open.
Aspects I disagree with
– There is no migration, but long-lasting contacts that show up in genetics, since the Mesolithic-Neolithic transition. In my opinion, the expansion of admixture + haplogroup reduction and expansion depict clear migratory movements (although, obviously, without cultural identification no speculative ethnolinguistic grouping can be proposed). That much is obvious from Genomics, and if we are not going to accept the most basic findings as proof in favour of certain anthropological models, then Archaeology will not benefit at all from genetic studies.
– Because this is the Russian school of thought, when they talk about Proto-Indo-European they refer to a homeland dating to the Mesolithic-Neolithic transition, and therefore cultural-historical communities after that (and long-lasting contacts) refer to potential continuations of Proto-Indo-European. Following common language guesstimates, though, there is no reason to date the split of Anatolian from a common Indo-Hittite before ca- 4500 BC, since there is no reason to date a Late Proto-Indo-European beyond 4000-3000 BC, apart from controversial glottochronological studies…