Genetic continuity among Uralic-speaking cultures in north-eastern Europe

east-europe-bronze-age

The recent study of Estonian Late Bronze Age/Iron Age samples has shown, as expected, large genetic continuity of Corded Ware populations in the East Baltic area, where West Uralic is known to have been spoken since at least the Early Bronze Age.

The most interesting news was that, unexpectedly for many, the impact of “Siberian ancestry” (whatever that actually means) was small, slow, and gradual, with slight increases found up to the Middle Ages, compatible with multiple contact events in north-eastern Europe. Haplogroup N became prevalent among Finnic populations only through late bottlenecks, as research of modern populations have long suggested, and as ancient DNA research hinted since at least 2015.

I risked to correlate the arrival of chiefs from the south-west with the infiltration of N1c-VL29 subclades during the transition to the Iron Age, coupled with that minimal “Siberian” ancestry (see e.g. here and here). Now we know that the penetration of this non-CW ancestry started, as predicted, in the Iron Age; that it was highly variable in the few samples where it appeared, with ca. 1-4%, while most Iron Age individuals show 0%; and that it was not especially linked to individuals of N1c-Vl29 lineages.

It is also basically confirmed, based on the (ancient and Modern Swedish) N1c-L550 subclades found among Iron Age Estonians, that N1c-VL29 lineages and the so-called “Siberian” ancestry will be found simultaneously around the Baltic coastal areas, and that different lineages must have suffered later founder effects among Finns, which suggests that these alliances through exogamy brought exactly as much language change in Sweden, Lithuania, or Poland, as they did in the East Baltic region…

On the other hand, the paper has also shown a potential movement of Corded Ware-derived peoples, if the change from LBA to IA samples is meaningful; in fact, even more Corded Ware-like than Baltic and Estonian BA populations. The exact origin of that movement is difficult to pinpoint, and it may not be related to the arrival of Akozino warrior-traders from the south-east, since theirs seems to be a minor impact proper of elites in a chiefdom system around the Baltic.

fortified-settlements-lba-ia
Distribution of fortified settlements (filled circles) and other hilltop sites (empty circles) of the Late Bronze Age and Pre-Roman Iron Ages in the East Baltic region. Tentative area of most intensive contacts between Baltic and Balto-Finnic communities marked with a dashed line. Image modified from (Lang 2016).

Also suggesting a potential movement is the ‘southern’ shift observed in the West and East Baltic areas, likely showing the arrival of Proto-East Baltic speakers (such as the Trzciniec outlier), as we have already discussed in this blog. The unexpected increase in Corded Ware-like ancestry in the Eastern Baltic, coupled with the expected large continuity of hg. R1a-Z283 in the homeland of Balto-Finnic expansions, gives even more support to the known complex system of exogamy along the Baltic coasts, and offers another potential reason for the rise of Baltic-speaking territories in the West Baltic: elite domination.

It is nevertheless important to understand that, even among the most “genetic continuous” regions like Estonia, not a single population in Europe is heir of some ancestral, immutable people. Not in terms of haplogroups, and not in terms of admixture. Balto-Finnic speakers, however continuous they might seem (e.g. in Southern Estonians) aren’t an exception.

After all, this blog was (re)born to fight the currently prevalent sheer stupidity surrounding the simplistic “R1a/steppe ancestry=Indo-European” association, so I wouldn’t like to see it replaced with some other stupid continuity or purity ideas within 10 to 20 years…

Late Uralic stems from East Corded Ware groups

With the currently available tools – linguistics, archaeology, and now genetics -, I don’t think there is any argument to date to question the direct connection of the Late Proto-Uralic expansion with all Eastern Corded Ware groups (i.e. Battle Axe, Fatyanovo-Balanovo, and Abashevo), and thus at least with the unifying A-horizon of Corded Ware and the bottlenecks under R1a-Z645.

NOTE. The only out-group among Corded Ware cultures is the Single Grave culture. It appears to be an early Corded Ware offshoot, reflected in their non-unitary cultural traits (distinct from later unifying waves), in their varied patrilineal clans, and in the short-lasting cultural effect in northern Europe before their complete demise under pressure of expanding Yamna/Bell Beaker peoples from the Danube. The culture’s minimal (if any) effects on succeeding peoples might be seen mostly in the (mainly phonetic) Uralic substrate found in Balto-Slavic – although this may also stem from a more eastern influence, close to the Baltic – and in the contacts of Celtic with Uralic. The huge time depth between this early hypothetic Uralic layer in northern Europe and the emergence of peoples inhabiting these territories in recorded history have no doubt been erroneously interpreted as a lack of Uralic presence in the area.

1) That connection was evident in the Yamna – CWC differences in archaeology, and especially later, with at least Fatyanovo-Balanovo and Abashevo representing the obvious replacement of the Volosovo culture before further expansions of CWC-related groups west and east of the Urals.

The mythical millennia-long continuity of Volosovo hunter-gatherers, including centuries among Corded Ware peoples, as expected lately by the Copenhagen group (and anyone who doesn’t want to question the 1960s association of Indo-European with CWC) must be rejected today in population genomics, as the recent studies of ancient and modern populations show, and as ancient DNA from the region will confirm.

2) In linguistics, the survival of Volosovo as The Uralic-speaking culture was also hardly believable. From Kallio (2015):

While we can say at least something about Uralic substrates in Northeastern Europe, non-Uralic substrates cannot at all easily be identified, because of multiple language shifts, viz. first from non-Uralic to Uralic and then from Uralic to Russian. Yet the Soviet Uralicist Boris Serebrennikov (1956, 1959) argued that there are some non-Uralic substrate toponyms in the Volga-Oka region, but his idea was never taken seriously in the west (cf. Sauvageot 1958), and it pretty soon also sank into oblivion in Russia, even though it can still occasionally pop up there in non-onomastic circles (cf. Napolskikh 1995: 18–19). However, not all the hypotheses on non-Uralic substrates in Northeastern Europe should be rejected (see e.g. Helimski 2001b).

bronze-age-early-languages-east-europe
Tentative map of the distribution of known languages in Eastern Europe during the Early Bronze Age. See full map.

Helimski (2001) argues for a non-Uralic topo-hydronomy in Northern Russia, whose population may have kept their languages up to the Common Era despite the Corded Ware expansion, which is in line with the survival of some non-Indo-European languages everywhere in Europe after the expansion of Yamna and its offshoots:

It should be borne in mind that these [Uralic] hydronyms reached us mainly through Northern Russian and, accordingly, with a tendency to phonetic-morphological adaptation and unification (for river names it is “natural” to be, like the word ‘river’ itself, feminine and to end in -a). Taking into account this circumstance, it may turn out to be non-useless for etymological identification of at least some of the hydronyms on the Finno-Ugric basis.

On the other hand, I wouldn’t exclude the possibility that some parts of this large geographical area were never (completely) Finno-Ugric. The population that created the most important part of the hydronymy of the Russian North could be finally pushed aside or assimilated only at the end of the 1st – beginning of the 2nd millennium AD, during the Russian colonization, retaining the memory of the White-Eyed Chude in its own memory.

NOTE. For more on this non-IE substrate in (especially West) Uralic, see e.g. Zhivlov (2015),

The same non-Uralic substrate is most likely behind most of the shared traits by Mordvinic and Balto-Finnic (see below).

3) In genetics, I don’t think the picture could get any clearer. I don’t know what “Steppe ancestry = Indo-European” proponents expected from 2019, if they expected anything at all (I haven’t seen any coherent model, proposal, or prediction for a long time now), but I doubt the recent results are compatible with any of their implied expectations.

corded-ware-pca-sub-neolithic-europe
Detail of the PCA of the Corded Ware expansion. See full PCA and more related files.

Notice, from the PCA above, how this Baltic Late Neolithic group shows actually a shift from Sredni Stog (see PCA with Sredni Stog) towards typical Khvalynsk-Urals-related ancestry, i.e. populations from eastern European forested regions, derived from hunter-gatherer pottery groups, as I have proposed for a very long time, since the first time a Baltic LN “outlier” appeared. It’s amazing how some amateurs can find 0.1% of any Siberian outlier’s ancestry among Uralians 4,000 years later, but fail to see the direct connection here. The esoteric uses of qpAdm, I guess…

Especially noticeable is the extra WHG-like ancestry and corresponding shift, seen especially marked in late Polish CWC samples, but also in Baltic CWC and especially in one Sweden Battle Axe sample, all of them shifting apparently closer to Pitted Ware and SHG. While that may have been interpreted as an in situ admixture in Scandinavia before, the late Polish CWC samples show likely a resurgence of local populations, so we can assume that both shifts (to SHG- and EHG-like populations) of available CWC samples around the Baltic are clearly part of the WHG:EHG continuum that will be found in the eastern European sub-Neolithic cultures, from Narva to Volosovo.

This WHG-related ancestry is clearly predominant in groups with which Battle Axe peoples admixed, based on the shift towards Pitted Ware, which – I can only guess based on modern Volga Finns – is different from the shift we will see in Netted Ware, more towards the Khvalynsk-Urals cluster. This is in line with the expansion of Battle Axe eastward through coastal areas (West to East Baltic and Finland into Sweden), while Fatyanovo peoples probably emerged from a slightly different route, but also a northern one, if one is to follow archaological similarities and their chronology.

bronze-age-europe-baltic
Detail of the PCA of European Bronze Age populations. See full PCA and more related files.

During the Iron Age, the only peoples that probably shifted strongly (based on modern populations) are West Baltic ones, getting closer to the available Late Trzciniec samples, and even closer to the Trzciniec outlier, i.e. away from the earlier Eastern Corded Ware cluster, and towards Central European groups like Czech EBA or Poland EBA, both of them clearly derived from Bell Beakers, but also admixed with (and thus shifted toward) CW-like populations.

If one looks carefully at the previous PCA on Bronze Age populations, and the next one on Iron Age clusters, it is evident that adding the Swedish LN outlier to East Baltic BA (both strongly related to Battle Axe populations) essentially gives us the continuity of East Baltic BA into the Iron Age. This cluster is continued also in two outliers from Sigtuna, a Viking town close to the Gulf of Finland, known to be an important trading site, 1,500 years later. Not much of a change around the Gulf of Finland, then:

iron-age-eastern-europe
Detail of the PCA of East and North European Iron Age populations. See full PCA and more related files.

Based on the two simplistic Uralic clines one might see described (among the many that certainly existed, from Corded Ware to different Eurasian populations), and just like BOO was for some months fashionable as “Samic”, some may be tempted to say that certain Sintashta or Srubna outliers close to the Urals mark the True Uralic™ peoples. Because, of course they do. Ghost haplogroup N and stuff. And Corded Ware never ever Uralic. Because Gimbutas, and my IE R1a grandfather.

NOTE. Funny thing here: there might be Corded Ware, Iranian, Slavic, Germanic, etc… outliers or out-groups, and they might form the widest genetic clusters ever seen, but they are all of one language, because archaeology and linguistics; however, one “outlier” (also, put your own definition of “outlier” here, let’s say 1% of whatever, and strontium isotope potentially from 100 km away) ca. 600 BC in the Baltic who (surprise!) happens to show hg. N, and he signals the first incoming True Uralic™ speaker from wherever… It won’t be the first or the last time some people resort to “the complexity of Uralic-speaking peoples” in ancestry, just to look for “hg. N = Uralic” like crazy. You only need common sense to understand that this is not how this works. Amateur genomics can’t get more embarrassing than the current “let’s look for ‘Siberian ancestry’ in every individual of haplogroup N” trend. Or maybe it can, and it will, but I can’t see it yet.

If one were to insist on looking for ‘foreign’ contributions among Iron Age Estonians, though, I think one should also check out first archaeology, and then the PC3 (or, more graphically, a 3D plot), to understand what might be happening with the many Uralic clines derived from Corded Ware, before starting to play around with bioinformatic tools to discover a teeny tiny 1% admixture of the wrong population, and rushing to build far-fetched narratives. Apparently, one of the different clines formed roughly between southern (steppe – forest-steppe) and northern (tundra-taiga) populations in Uralians is also seen in some Iron Age Estonian individuals – especially in some late samples from Ingria…This is not my main interest, so I will leave this here for others to keep wasting their time chasing the white whale of the 0.5% of True Uralic™ ancestry in ancient Baltic samples of hg. N.

pca-3d-estonians-iron-age-boo-samic
Still images of the 3D plot of Eurasian samples. Typical PC1 vs. PC2 visualization to the left, and shift of the view to PC3 on the right image. See full PCA and more related files.

An exclusive Volga-Kama homeland for Disintegrating Uralic?

Since I don’t believe in macro-regions of largely continuous ethnolinguistic communities, as I have often said about Slavic (naively associated with prehistoric tribes of Eastern Europe) or Germanic (absurdly considered to be represented by Battle Axe), it is difficult for me to believe that Battle Axe-derived cultures remained of the same Finno-Samic dialects since the Corded Ware expansion…unless we live in Westeros, where everything happens “for thousands of years”.

I have to admit, then, that the now prevalent identification among Uralicists has become quite attractive:

  • Fatyanovo-Balanovo as Finno-Permic:
    • Fatyanovo/Netted Ware with West Uralic (also called Finno-Mordvinic).
    • Balanovo/Chirkovo-Kazan with Central Uralic (Mari-Permic).
  • Abashevo, into the Andronovo-like Horizon through the Seima-Turbino phenomenon, with East Uralic (also Ugro-Samoyedic).

Exactly like the identification of Yamna Hungary – Bell Beaker transition as the North-West Indo-European homeland, it gives us simplicity and small and late ethnolinguistic communities, away from the traditionally overused big and early language territories.

This late homeland would be supported, among others, by:

  • The presence of Indo-Iranian loanwords in Finno-Permic and Ugric (probably also in Samoyedic, either lost, or – much more likely – underresearched), compatible with the immediate contact between Abashevo – Sintashta-Potapovka-Filatovka and Fatyanovo-Balanovo.
  • The supposed expansion of Netted Ware from Fatyanovo to the north-west, which may be explained as the split and expansion of Balto-Finnic and Samic ca. 1900 BC.
  • A longer-lasting Finno-Permic (West+Central Uralic) community contrasting with the early separation of East Uralic.
  • The compatibility of this late expansion with the late expansion of Pre-Germanic from Denmark with the Dagger Period, and of Balto-Slavic with Trzciniec, which puts all three dialects reaching the Baltic Sea in the EBA.

NOTE. I meant to update the linguistic text to include the most recently favoured phylogenetic tree of Uralic languages after Häkkinen (2007, 2009, 2014), which has very quickly become the new normal among Uralicists, but I don’t think I will have enough time to review the necessary papers for that. I am rushing to publish a printed edition, so the text will wind up being a mixture of “traditional” (meaning, basically, pre-2010s) description of Uralic dialects but using modern divisions; say, “West Uralic” instead of “Finno-Samic”. By the way, I am still amazed that none of my reader-haters (or any online user discussing Uralic migrations, for that matter) have come up with the questions that the new division pose, and it supports my suspicion about the complete lack of interest in linguistics of most (a)DNA fans, except for the occasional use of old and free PDFs Googled to support new narratives invented expressly for some qpAdm results…

textile-ceramics-europe-bronze-age
Textile ceramic styles and influence of Bronze Age cultures divided in clusters.

Problems with this Parpola-Carpelan’s (2012-2018) interpretation include:

  • The differentiation between Fennoscandian Textile Ceramics vs. Netted Ware, which is not warranted in archaeology. The assumption that Netted Ware expanded to the Baltic Sea (as Kallio does, following the traditional view) is thus weak, and it was probably a question of cultural contacts coupled with short-distance population movements/exchange in both directions (from the Baltic to the Volga and vice versa). In fact, the culture division relies on some fairly common and technically simple ornamentation patterns, widespread all over northern Europe, even before the Corded Ware expansion, and it is very difficult to separate certain neighboring Textile Ceramics from Netted Ware groups in southern Finland (i.e. Sarsa-Tomitsa groups).
  • The strict and radical direction described for the Netted Ware by Carpelan, as an eastward and northward expansion, within a very short time frame (ca. 1900-1800 BC), based on few radiocarbon dates, which seems to me like a very risky assumption. We know how this kind of descriptions of direction of culture expansion based on radiocarbon dates has turned out in much more complex “packages”, like the Bell Beaker culture… In fact, the earliest dates for Textile Ware are from the East Baltic, earlier than those of Netted Ware.
  • The assumption that Balto-Finnic traits shared with Mordvinic are a) late and b) meaningful for dialectalization of two closely related dialects, when it is clear that both dialects separated quite early. Phonologically Finnic is more conservative, morphologically less so, and the shared traits include a handful of non-Uralic substrate words which can’t be traced to a single common source, hence they were adopted when both languages had already separated… All in all, Finnic – Mordvinic correspondances are not even close to Italo-Celtic ones, which is clearly fully incompatible with a proposal of a Finnic separation from Mordvinic coinciding with the LBA-IA transition.

Especially problematic for Parpola’s model is the lack of genetic impact in Bronze Age or Iron Age Estonians, not reaching a significant level under any possible statistical threshold – which I am sure was quite disappointing for some of my readers -, but is in line with major archaeological continuity of groups the from region, only disturbed in cultural (and Y-chromosome) terms by the expansion of Akozino warrior-traders all over the Baltic Sea. Any proposed population movement will be very difficult to support in genetics, given the Corded Ware-derived populations that we will see in both regions, and the continued Baltic-Volga contacts since the Corded Ware expansion.

Problems with an interpretation of such a small impact in population genomics includes the similarly weak impacts and haplogroup infiltrations that can be seen among populations basically everywhere in Eurasia, during any given period, and much greater genetic impacts that are supposed to be (or that were certainly) followed by ethnolinguistic continuity.

akozino-malar-axes-fennoscandia
Distribution of the Akozino-Mälar axes according to Sergej V. Kuz’minykh (1996: 8, Abb. 2).

The Battle Axe question

From Kallio (2015), about choosing a tentative homeland for Proto-Uralic:

(…) linguistically uniform Proto-Uralic would have been spoken in the Volga-Oka region until the mid-third millennium BC when the Proto-Uralic-speaking area would have expanded to the Volga-Kama region as well. By the end of the same millennium, this expansion would have led to the earliest dialectal splits within Uralic into Finno-Mordvin, Mari-Permic, and Ugro-Samoyed. The splitting up of these three soon followed during the early second millennium BC when the Uralic-speaking area finally stretched from the Baltic Sea in the west to the Altai mountains in the east. Indeed, no matter where Proto-Uralic was spoken, the branching into the nine well-attested subgroups (viz. Finnic, Saami, Mordvin, Mari, Permic, Hungarian, Mansi, Khanty, and Samoyed) must have taken less than a millennium, because their shared phonological and morphosyntactic isoglosses are rather limited (see Salminen 2002). The traditional view that all this branching would have taken several millennia violates everything linguistic typology teaches us about the rate of language change.

The basic problem of this identification of Fatyanovo-Balanovo as West-Central Uralic and Abashevo as East Uralic is the nature of the Battle Axe culture, including the Bronze Age East Baltic and Gulf of Finland area. Even if it is accepted that Fatyanovo-Balanovo represented all Western groups, Battle Axe must have represented West Uralic-like dialects.

The ethnolinguistic identification of Battle Axe depends ultimately on the nature of contacts of Fatyanovo/Netted Ware with Battle Axe/Textile Ceramics. If both groups were close and interacted profusely, as it seems, it doesn’t seem granted that we will be able to distinguish a close Para-West Uralic dialect of Scandinavia from the actual expanding Balto-Finnic and Samic dialects, if they were actually linked to the Netted Ware expansion. Also from Kallio (2015):

No doubt the most convincing substrate theory has recently been put forward by the Saami Uralicist Ante Aikio (2004), who has not only rehabilitated but also improved the old idea of a non-Uralic substrate in Saami. His study shows that there were still non-Uralic languages spoken in Northern Fennoscandia as recently as the first millennium AD. Most of all, they were not only genetically non-Uralic but also typologically non-Uralic-looking, bearing a closer resemblance to the so-called Palaeo-European substrates (for which see e.g. Schrijver 2001; Vennemann 2003).

In comparison, the case of Finnic is much more difficult. The fact that Proto-Uralic was not spoken in the East Baltic region means that this area must have originally been non-Uralic-speaking, but so far the evidence for a non-Uralic substrate in Finnic has consisted of appellatives and proper names with no etymology (cf. Ariste 1971; Saarikivi 2004a). Contrary to the proposed substrate words in Saami, those in Finnic show no structural non-Uralisms, as if they had indeed been borrowed from some genetically related or at least typologically similar languages, as I suggested above. Also none of them is more recent than the Middle Proto-Finnic stage, which makes them at least two millennia old. All this agrees with archaeological evidence discussed earlier that the Uralicization of the East Baltic region occurred during the Bronze Age (ca. 1900–500 BC).

The discussion of the paper continues with an unsuccessful attempt to find a hypothetical ancient Indo-European substrate that Kallio believes must be associated with the expansion of Corded Ware, in line with the traditional belief. For example, the often mentioned – almost folk etymology-like, unsurprisingly popular among amateurs – ‘Neva’ as derived from IE “young” is logically rejected…Unlike Parpola, Kallio’s view seems to be confident that Netted Ware (as Textile Ware) expanded into the East Baltic, on both sides of the Gulf of Finland, already during the Bronze Age.

As it has become apparent in population genomics, none of them was right, and Textile Ceramics will essentially show – like Netted Ware – a large genetic continuity of Corded Ware peoples in the whole north-eastern Europe – despite small regional population movements, obviously -, which necessarily implies that the whole Corded Ware culture – and not only Fatyanovo-Balanovo and Abashevo – were Uralic-speaking territories.

The similarities in terms of culture and Y-DNA bottlenecks between Battle Axe and Fatyanovo-Balanovo also imply that the linguistic differences between these groups were probably not many, and became strongly divided only after their territorial division. Continued contacts between Battle Axe- and Fatyanovo-derived groups can explain the proposed contacts (Finnic with Samic, Finnic with Mordvinic) after their linguistic-but-not-physical separation.

east-european-fatyanovocwc
East European movement directions (arrows) of the representatives of the Central European Corded Ware Culture (according to I.I. Artemenko).

Battle Axe spoke “Para-Balto-Finnic”?

The Balto-Finnic-speaking nature of Battle Axe is thus supported by:

  • The lack of non-Uralic substrates in Balto-Finnic territory (Kallio 2015).
  • The early separation of Samic and Finnic from Mordvinic, and the virtual identity of Proto-West-Uralic and Proto-Uralic, which suggests that Proto-Uralic spread fast (Parpola 2012).
  • The scarce non-Uralic topo-hydronymy in the East Baltic and around the Gulf of Finland (Saarikivi 2004), comparable to that on the Upper Volga region.
  • The strong influence of a Balto-Finnic-like substrate on Pre-Germanic (or, in Kallio’s opinion, the same Scandinavian substrate influencing both Germanic and Balto-Finnic at the same time), and the continued influence of Balto-Finnic on Proto-Baltic and Proto-Slavic.
  • The continued influence of Corded Ware-derived groups in central-east Sweden in Finland and the East Baltic in terms of agricultural innovations appearing in the LBA, compatible with Schrijver’s proposal of intermediate Germanic-shifted Balto-Finnic groups and Balto-Finnic groups influenced by their pronunciation.
  • The intense Palaeo-Germanic and late Balto-Slavic / early Proto-Baltic superstrate on Balto-Finnic, which place all three dialects around the Baltic Sea since the Early Bronze Age.
  • The easy replacement of a hypothetic Para-Balto-Finnic dialect by incoming Proto-Balto-Finnic-speaking peoples (say, with textile ceramics), without much linguistic impact.

In fact, the continuous contacts of the East Baltic with the Volga, and especially the close interaction with Akozino warrior-traders just before the Tarand-grave period, could be the actual origin of the recent (if any) Finnic-Mordvinic connections that need to be traced back to the LBA-IA (maybe here the number ‘ten’), since most of them can be related to a Pit-Comb Ware culture substrate and earlier contacts through the forest zone, which Samic (due to its early split and presence to the north of the Gulf of Finland during the BA) does not share. In fact, some of them can be traced back to Balto-Finnic first

These are the most often mentioned, in order of descending relevance for a shared ancient community:

  • Noun paradigms and the form and function of individual cases.
  • The geminate *mm (foreign to Proto-Uralic before the development of Fennic under Germanic influence) and other non-Uralic consonant clusters.
  • The change of numeral *luka ‘ten’ with (non-Uralic) *kümmen.
  • The presence of loanwords of non-Uralic origin, related to farming and trees, potentially Palaeo-European in nature.

It’s not only a question of quantity. Are these shared Mordvinic – Balto-Finnic traits really more relevant than, say, those between Italo-Celtic, which are supposed to have formed a community for a very short period at the end of the 3rd millennium around the Alps? Are these traits even sufficient to propose a common early Mordvinic-Finnic group within West Uralic, rather than loose Mordvinic – Balto-Finnic contacts, i.e. contacts between East Baltic (Textile Ceramics) and Volga-Kama (Netted Ware)?

Based on the alternative (Kallio’s) view of continued contacts between Textile Ceramics groups, even without knowing anything about linguistics, you can guess that Parpola is spinning very thin when assuming that these changes suggest that Balto-Finnic may have expanded with Akozino warrior-traders, separating thus ca. 800 BC from Mordvinic…

Genetic findings now clearly help dismiss any meaningful population impact in the LBA-IA transition, although any linguist can obviously argue for linguistic change in spite of major genetic continuity. But then we are stuck in the pre-ancient DNA era, so what’s ancient DNA for.

netted-ware-textile-ceramics
Middle Bronze Age cultures of Eastern Europe.

Genetic continuity = language continuity?

In the end, it’s very difficult to say how much language continuity there is around Estonia since the arrival of Corded Ware peoples. Looking at Modern Estonians, they have been clearly influenced by recent contacts with Baltic- and Germanic-speaking peoples clustering to the south-west in the PCA. They seem to have also received contacts from north(-east)ern peoples, likely from Finland, evidenced by their shifts toward the modern Estonian cluster during and after the Middle Ages, with a slight increase in Siberian ancestry and N1c subclades associated with Lovozero Ware. How much language change did these contacts bring? Maybe an expansion of Gulf of Finland Finnic (Northern Estonian) over Inland Finnic (Southern Estonian) and Gulf of Riga Finnic (Livonian)? Difficult to know, exactly, but, in the traditional view of Balto-Finnic dialectal distribution among Uralicists like Kallio, possibly no change at all.

So, if the obvious changes in the Estonia_MA cluster relative to Estonia_IA cluster and Estonia_Modern relative to Estonia_MA do not represent radical language change…Why would Estonia_IA represent a change relative to Estonia_BA, when it is statistically basically the same? Or Estonia_BA relative to CWC_Baltic? Because of the infiltration of haplogroup N1c around the whole Baltic? Because of the occasional 1% “Siberian” ancestry in some non-locals of varied haplogroups across the whole Baltic area?

In spite of all this, the amount of special pleading we are seeing among openly Nordicist amateurs when discussing the Uralic homeland relative to the Indo-European question in genetics has become a matter of plain willful ignorance. Like the living corpses of the Anatolian homeland, the Armenian homeland, the OIT proponents, or the nativist Basque R1b association, the personal involvement in the revival of “R1a=Indo-European” and “N=Uralic” trends is just painful to watch.

[Next post in this line, if I manage to make time for it: “Genetic (dis)continuity in Central Europe“. Let’s see if early Balts and early Slavs, as well as Germanic peoples, show a cluster closer to Danubian EBA (viz. Maros), Hungary-Balkans BA, and Urnfield-related samples than their predecessors in their areas, i.e. away from East Corded Ware groups… If you want, you can enjoy for the moment the new PCAs I could get done and the tentative map of languages in the Early Bronze Age, that will probably give you the right idea about early Indo-European and Uralic population movements]

bronze-age-early-indo-european
European Early Bronze Age: tentative language map based on linguistics, archaeology, and genetics. See full map.

Related

Złota a GAC-CWC transitional group…but not the origin of Corded Ware peoples

koszyce-gac-zlota-cwc

Open access Unraveling ancestry, kinship, and violence in a Late Neolithic mass grave, by Schroeder et al. PNAS (2019).

Interesting excerpts of the paper and supplementary materials, about the Złota group variant of Globular Amphora (emphasis mine):

A special case is the so-called Złota group, which emerged around 2,900 BCE in the northern part of the Małopolska Upland and existed until 2,600-2,500 BCE. Originally defined as a separate archaeological “culture” (15), this group is mainly defined by the rather local introduction of a distinct form of burial in the area mentioned. Distinct Złota settlements have not yet been identified. Nonetheless, because of the character of its burial practices and material culture, which both retain many elements of the GAC and yet point forward to the Corded Ware tradition, and because of its geographical location, the Złota group has attracted significant archaeological attention (15, 16).

The Złota group buried their dead in a new, distinct type of funerary structure; so-called niche graves (also called catacomb graves). These structures featured an entrance shaft or pit and, below that, a more or less extensive niche, sometimes connected to the entrance area by a narrow corridor. Local limestone was used to seal off the entrance shaft and to pave the floor of the niche, on which the dead were usually placed along with grave goods. This specific and relatively sophisticated form of burial probably reflects contacts between the northern Małopolska Upland and the steppe and forest-steppe communities further to the east, who also buried their dead in a form of catacomb graves. Individual cases of the use of ochre and of deformation of skulls in Złota burials provide further indications of such a connection (15). At the same time, the Złota niche grave practice also retains central elements of the GAC funerary tradition, such as the frequent practice of multiple burials in one grave, often entailing redeposition and violation of the anatomical order of corpses, and thus differs from the catacomb grave customs found on the steppes which are strongly dominated by single graves. Nonetheless, at Złota group cemeteries single burial graves appear, and even in multiple burial graves the identity of each individual is increasingly emphasized, e.g. by careful deposition of the body and through the personal nature of grave goods (16).

globular-amphorae-corded-ware-zlota-amphorae
Correspondence analysis of amphorae from the Złota-graveyards reveals that there is no typological break between Globular Amphorae and Corded Ware Amphorae, including ‘Strichbündelamphorae’ (after Furholt 2008)

Just like its burial practices, the material culture and grave goods of the Złota group combine elements of the GAC, such as amber ornaments and central parts of the ceramic inventory, with elements also found in the Corded Ware tradition, such as copper ornaments, stone shaft-hole axes, bone and shell ornaments, and other stylistic features of the ceramic inventory. In particular, Złota group ceramic styles have been seen as a clear transitional phenomenon between classical GAC styles and the subsequent Corded Ware ceramics, probably playing a key role in the development of the typical cord decoration patterns that came to define the latter (17).

As briefly summarized above, the Złota group displays a distinct funerary tradition and combination of material culture traits, which give the clear impression of a cultural “transitional situation”. While the group also appears to have had long-distance contacts directed elsewhere (e.g. to Baden communities to the south), it is the combination of Globular Amphora traits, on the one hand, and traits found among late Yamnaya or Catacomb Grave groups to the east as well as the closely related Corded Ware groups that emerged around 2,800 BCE, on the other hand, that is such a striking feature of the Złota group and which makes it interesting when attempting to understand cultural and demographic dynamics in Central and Eastern Europe during the early 3rd millennium BCE.

catacomb-grave-ksiaznice
Catacomb grave no. 2a/06 from Książnice, Złota culture (acc. to Wilk 2013). Image from Włodarczak (2017)

Książnice (site 2, grave 3ZC), Świętokrzyskie province. This burial, a so-called niche grave of the Złota type (with a vertical entrance shaft and perpendicularly situated niche), was excavated in 2006 and contained the remains of 8 individuals, osteologically identified as three adult females and five children, positioned on limestone pavement in the niche part of the grave. Radiocarbon dating of the human remains indicates that the grave dates to 2900-2630 BCE, 95.4% probability (Dataset S1). The grave had an oval entrance shaft with a diameter of 60 cm and depth of 130 cm; the depth of the niche reached to 170 cm (both measured from the modern surface), and it also contained a few animal bones, a few flint artefacts and four ceramic vessels typical of the Złota group. Książnice is located in the western part of the Małopolska Upland, which only has a few Złota group sites but a stronger presence of other, contemporary groups (including variants of the Baden culture).

Wilczyce (site 90, grave 10), Świętokrzyskie province. A rescue excavation in 2001 uncovered a niche grave of the Złota type, which had a round entrance shaft measuring 90 cm in diameter. The grave was some 60-65 cm deep below the modern surface and the bottom of the niche was paved with thin limestone plates, on which remains of three individuals had been placed; two adults, one female and one male, and one child. Four ceramic vessels of Złota group type were deposited in the niche along with the bodies. Wilczyce is located in the Sandomierz Upland, an area with substantial presence of both the Globular Amphora culture and Złota group, as well as the Corded Ware culture from 2800 BCE.

zlota-gac-cwc
Genetic affinities of the Koszyce individuals and other GAC groups (here including Złota) analyzed in this study. (A) Principal component analysis of previously published and newly sequenced ancient individuals. Ancient genomes were projected onto modern reference populations, shown in gray. (B) Ancestry proportions based on supervised ADMIXTURE analysis (K = 3), specifying Western hunter-gatherers, Anatolian Neolithic farmers, and early Bronze Age steppe populations as ancestral source populations. LP, Late Paleolithic; M, Mesolithic; EN, Early Neolithic; MN, Middle Neolithic; LN, Late Neolithic; EBA, Early Bronze Age; PWC, Pitted Ware culture; TRB, Trichterbecherkultur/Funnelbeaker culture; LBK, Linearbandkeramik/Linear Pottery culture; GAC, Globular Amphora culture; Złota, Złota culture. Image modified to outline in red GAC and Złota groups.

To further investigate the ancestry of the Globular Amphora individuals, we performed a supervised ADMIXTURE (6) analysis, specifying typical western European hunter-gatherers (Loschbour), early Neolithic Anatolian farmers (Barcın), and early Bronze Age steppe populations (Yamnaya) as ancestral source populations (Fig. 2B). The results indicate that the Globular Amphora/Złota group individuals harbor ca. 30% western hunter-gatherer and 70% Neolithic farmer ancestry, but lack steppe ancestry. To formally test different admixture models and estimate mixture proportions, we then used qpAdm (7) and find that the Polish Globular Amphora/Złota group individuals can be modeled as a mix of western European hunter-gatherer (17%) and Anatolian Neolithic farmer (83%) ancestry (SI Appendix, Table S2), mirroring the results of previous studies.

zlota-steppe-ancestry-cwc
Table S2. qpADM results. The ancestry of most Globular Amphora/Złota group individuals
can be modelled as a two-way mixture of Mesolithic western hunter-gatherers (WHG), and early Anatolian Neolithic farmers (Barcın). The five individuals from Książnice (Złota group) show evidence for additional gene flow, most likely from an eastern source.

The lack of a direct genetic connection of Corded Ware peoples with the Złota group despite their common “steppe-like traits” – shared with Yamna – reveals, once more, how the few “Yamna-like” traits of Corded Ware do not support a direct connection with Indo-Europeans, and are the result of the expansion of the so-called steppe package all over Europe, and particularly among cultures closely related to the Khvalynsk expansion, and later under the influence of expanding Yamna peoples.

The results from Książnice may support that early Corded Ware peoples were in close contact with GAC peoples in Lesser Poland during the complex period of GAC-Trypillia-CWC interactions, and especially close to the Złota group at the beginning of the 3rd millennium BC. Nevertheless, patrilineal clans of Złota apparently correspond to Globular Amphorae populations, with the only male sample available yet being within haplogroup I2a-L801, prevalent in GAC.

NOTE. The ADMIXTURE of Złota samples in common with GAC samples (and in contrast with the shared Sredni Stog – Corded Ware “steppe ancestry”) makes the possibility of R1a-M417 popping up in the Złota group from now on highly unlikely. If it happened, that would complicate further the available picture of unusually diverse patrilineal clans found among Uralic speakers expanding with early Corded Ware groups, in contrast with the strict patrilineal and patrilocal culture of Indo-Europeans as found in Repin, Yamna and Bell Beakers.

Once again the traditional links between groups hypothesized by archaeologists – like Gimbutas and Kristiansen in this case – are wrong, as is the still fashionable trend in descriptive archaeology, of supporting 1) wide cultural relationships in spite of clear-cut inter-cultural differences (and intra-cultural uniformity kept over long distances by genetically-related groups), 2) peaceful interactions among groups based on few common traits, and 3) regional population continuities despite cultural change. These generalized ideas made some propose a steppe language shared between Pontic-Caspian groups, most of which have been proven to be radically different in culture and genetics.

gimbutas-kurgan-indo-european
The background shading indicates the tree migratory waves proposed by Marija Gimbutas, and personally checked by her in 1995. Image from Tassi et al. (2017).

Furthermore, paternal lines show once again marked bottlenecks in expanding Neolithic cultures, supporting their relevance to follow the ethnolinguistic identity of different cultural groups. The steppe- or EHG-related ancestry (if it is in fact from early Corded Ware peoples) in Książnice was thus probably, as in the case of Trypillia, in the form of exogamy with females of neighbouring groups:

The presence of unrelated females and related males in the grave is interesting because it suggests that the community at Koszyce was organized along patrilineal lines of descent, adding to the mounting evidence that this was the dominant form of social organization among Late Neolithic communities in Central Europe. Usually, patrilineal forms of social organization go hand in hand with female exogamy (i.e., the practice of women marrying outside their social group). Indeed, several studies (11, 12) have shown that patrilocal residence patterns and female exogamy prevailed in several parts of Central Europe during the Late Neolithic. (…) the high diversity of mtDNA lineages, combined with the presence of only a single Y chromosome lineage, is certainly consistent with a patrilocal residence system.

funnelbeaker-trypillia-corded-ware
Map of territorial ranges of Funnel Beaker Culture (and its settlement concentrations in Lesser Poland), local Tripolyan groups and Corded Ware Culture settlements (■) at the turn of the 4th/3rd millennia BC.

Since ancient and modern Uralians show predominantly Corded Ware ancestry, and Proto-Uralic must have been in close contact with Proto-Indo-European for a very long time – given the different layers of influence that can be distinguished between them -, it follows as logical consequence that the North Pontic forest-steppes (immediately to the west of the PIE homeland in the Don-Volga-Ural steppes) is the most likely candidate for the expansion of Proto-Uralic, accompanying the spread of Sredni Stog ancestry and a bottleneck under R1a-M417 lineages.

The early TMRCAs in the 4th millennium BC for R1a-M417 and R1a-Z645 support this interpretation, like the R1a-M417 sample found in Sredni Stog. On the other hand, the resurgence of typical GAC-like ancestry in late Corded Ware groups, with GAC lineages showing late TMRCAs in the 3rd millennium BC, proves the disintegration of Corded Ware all over Europe (except in Textile Ceramics- and Abashevo-related groups) as the culture lost its cohesion and different local patrilineal clans used the opportunity to seize power – similar to how eventually I2a-L621 infiltrated eastern (Finno-Ugrian) groups.

Related

Uralic speakers formed clines of Corded Ware ancestry with WHG:ANE populations

steppe-forest-tundra-biomes-uralic

The preprint by Jeong et al. (2018) has been published: The genetic history of admixture across inner Eurasia Nature Ecol. Evol. (2019).

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.

eurasian-clines-uralic-altaic
The first two PCs summarizing the genetic structure within 2,077 Eurasian individuals. The two PCs generally mirror geography. PC1 separates western and eastern Eurasian populations, with many inner Eurasians in the middle. PC2 separates eastern Eurasians along the northsouth cline and also separates Europeans from West Asians. Ancient individuals (color-filled shapes), including two Botai individuals, are projected onto PCs calculated from present-day individuals.

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).

west-urals-finno-ugrians-qpadm
Supplementary Table 8. QpAdm-based admixture modeling of the forest-tundra cline populations. For the 13 populations west of the Urals, we present a four-way admixture model, Nganasan+Srubnaya+WHG+LBK_EN, or its minimal adequate subset. Modified from the article, to include colors for cultures, and underlined best models for Corded Ware ancestry among Uralians.

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).

east-urals-ugric-samoyedic-qpadm
Supplementary Table 8. QpAdm-based admixture modeling of the forest-tundra cline populations. For the four populations east of the Urals, we present three admixture models: Baikal_EBA+Srubnaya, Baikal_EBA+Srubnaya+AG3 and Nganasan+Srubnaya+AG3. For each model, we present qpAdm p-value, admixture coefficient estimates and associated 5 cM jackknife standard errors (estimate ± SE). Modified from the article, to include colors for cultures, and underlined best models for Corded Ware ancestry among Uralians.

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.

bronze-age-iron-age-karasuk-mezhovska-tagar-qpadm
Supplementary Table 9. QpAdm-based admixture modeling of Bronze and Iron Age populations of southern Siberia. For ancieint individuals associated with Karasuk and Tagar cultures, Nganasan+Srubnaya model is insufficient. For all five groups, adding AG3 as the third ancestry or substituting Nganasan with Baikal_EBA with higher ANE affinity provides an adequate model. For each model, we present qpAdm p-value, admixture coefficient estimates and associated 5 cM jackknife standard errors (estimate ± SE). Models with p-value ≥ 0.05 are highlighted in bold face. Modified from the article, to include colors for cultures, and underlined best models for Corded Ware ancestry among Uralians.

Lara M. Cassidy comments the results of the study in A steppe in the right direction (you can read it here):

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.

eurasian-clines-uralic-turkic-mongol-altaic
The genomic formation of inner Eurasians. b–d, Depiction of the three main clines of ancestry identified among Inner Eurasians. Sources of admixture for each cline are represented using proxy ancient populations, both sampled and hypothesised, based on the study’s modelling results. The major eastern and western ancestries used to model each cline are shown in bold; the peripheral admixtures that gave rise to these are also shown. Additional contributions to subsections of each cline are marked with dashed lines. b, The northernmost cline, illustrating the legacy of WHG and ANE-related populations. c,d, The upper (c) and lower (d) steppe clines are shown, both of which have substantial eastern contributions related to modern Tungusic speakers. The authors propose these populations are themselves the result of an admixture between groups related to the Nganasan, whose ancestors potentially occupied a wider range, and hunter-gatherers (HGs) from the Amur River Basin. While the upper steppe cline in c can be described as a mixture between this eastern ancestry and western steppe herders, the current model for the southern steppe cline as shown in d is not adequate and is likely confounded by interactions with diverse bordering ancestries. Credit: Ecoregions 2017, Resolve https://ecoregions2017.appspot.com/

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:

siberian-clines-uralic-altaic
PCA of ancient and modern Eurasian samples. Ancient Palaeo-Laplandic, Palaeosiberian, and Altai clines drawn, with modern populations labelled. See a version with higher resolution.

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:

uralic-admixture-qpadm
Characterization of the Western and Eastern Eurasian source ancestries in inner Eurasian populations. [Modified from the paper, includes only Uralic populations]. a, Admixture f3 values are compared for different Eastern Eurasian (Mixe, Nganasan and Ulchi; green) and Western Eurasian references (Srubnaya and Chalcolithic Iranians (Iran_ChL); red). For each target group, darker shades mark more negative f3 values. b, Weights of donor populations in two sources characterizing the main admixture signal (date 1 and PC1) in the GLOBETROTTER analysis. We merged 167 donor populations into 12 groups (top right). Target populations were split into five groups (from top to bottom): Aleuts; the forest-tundra cline populations; the steppe-forest cline populations; the southern steppe cline populations; and ‘others’.

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.

west-eurasian-east-eurasian-ancestry
Supplementary Fig. 4. ADMIXTURE results qualitatively support PCA-based grouping of inner Eurasians into three clines. (A) Most southern steppe cline populations derive a higher proportion of their total Western Eurasian ancestry from a source related to Caucasus, Iran and South Asian populations. (B) Turkic- and Mongolic-speaking populations tend to derive their Eastern Eurasian ancestry more from the Devil’s Gate related one than from Nganasan-related one, while the opposite is true for Uralic- and Yeiseian-speakers. To estimate overall western Eurasian ancestry proportion, we sum up four components in our ADMIXTURE results (K=14), which are the dominant components in Neolithic Anatolians (“Anatolia_N”), Mesolithic western European hunter-gatherers (“WHG”), early Holocene Caucasus hunter-gatherers (“CHG”) and Mala from southern India, respectively. The “West / South Asian ancestry” is a fraction of it, calculated by summing up the last two components. To estimate overall Eastern Eurasian ancestry proportion, we sum up six components, most prevalent in Surui, Chipewyan, Itelmen, Nganasan, Atayal and early Neolithic Russian Far East individuals (“Devil’s Gate”). Eurasians into three clines. (A) Most southern steppe cline populations derive a higher proportion of their total Western Eurasian ancestry from a source related to Caucasus, Iran and South Asian populations. (B) Turkic- and Mongolic-speaking populations tend to derive their Eastern Eurasian ancestry more from the Devil’s Gate related one than from Nganasan-related one, while the opposite is true for Uralic- and Yeiseian-speakers. To estimate overall western Eurasian ancestry proportion, we sum up four components in our ADMIXTURE results (K=14), which are the dominant components in Neolithic Anatolians (“Anatolia_N”), Mesolithic western European hunter-gatherers (“WHG”), early Holocene Caucasus hunter-gatherers (“CHG”) and Mala from southern India, respectively. The “West / South Asian ancestry” is a fraction of it, calculated by summing up the last two components. To estimate overall Eastern Eurasian ancestry proportion, we sum up six components, most prevalent in Surui, Chipewyan, Itelmen, Nganasan, Atayal and early Neolithic Russian Far East individuals (“Devil’s Gate”).

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…

Related

The genetic and cultural barrier of the Pontic-Caspian steppe – forest-steppe ecotone

steppe-forest-steppe-biomes

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…

The Samara region in the Middle Volga may be pointed out as the true prehistoric link between forests and steppes (see David Anthony’s remarks), something reflected in its nature as a prehistoric sink in genetics. This strong forest – forest-steppe – steppe connection was seen in the Eurasian technocomplex, during the expansion of hunter-gatherer pottery, in the expansion of Abashevo peoples to the steppes (in one of the most striking cases of population admixture in the area), with Scythians (visible in the intense contacts with Ananyino), and with Turks (Volga Turks).

steppe-forest-steppe-europe
Simplified map of the distribution of steppes and forest-steppes (Pontic and Pannonian) and xeric grasslands in Eastern Central Europe (with adjoining East European ranges) with their regionalisation as used in the review (Northern—Pannonic—Pontic). Modified from Kajtoch et al. (2016).

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.

steppe-forest-steppe-migration-routes
Typical migration routes through European steppes and forest-steppes. Red line represents the persistent cultural and genetic barrier, with the latest evolution in steppe region represented by the shift from dashed line to the north. Arrows show the most common population movements. Modified from Kajtoch et al. (2016).

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:

wang-yamna-connection
Prehistoric individuals projected onto a PCA of 84 modern-day West Eurasian populations (open symbols). Dashed arrows indicate trajectories of admixture: EHG—CHG (petrol), Yamnaya—Central European MN (pink), Steppe—Caucasus (green), and Iran Neolithic—Anatolian Neolithic (brown). Modified from the original, a red circle has been added to the Yamna-Central European MN admixture.

Related

Eastern pressure blade technology in west Scandinavia associated with WHG

New interesting preprint Ancient DNA from chewing gums connects material culture and genetics of Mesolithic hunter-gatherers in Scandinavia, by Kashuba et al. (2018).

Interesting excerpts (emphasis mine):

Mitochondrial genomes from all three individuals belong to the U5a2d haplogroup. (…) The mitochondrial U5a2d haplogroup is consistent with earlier published results for ancient individuals from Scandinavia, U5a being the most common within SHG. Of the 16 Mesolithic individuals from Scandinavia published prior to our study, seven belong to the U5a haplogroup, nine share the U2 and U4 haplogroups

We divided the SHG group into two groups: SHGa and SHGb (ancient individuals found in contemporary Norway and Sweden, respectively). We based this on both the geographical distribution and the previous studies demonstrating the close relation of SHGa to EHG group and SHGb to WHG group. To further explore the demography within the SHG group, we compared the ancestry of BLE individuals within SHGa and SHGb groups. This comparison revealed a high relative shared drift between BLE individuals and the SHGb group

scandinavia-hunter-gatherer-admixture
Admixture analysis showing the major mode for K=15. The figure represents 11 runs out of 20 replicates (Greedy algorithm ran with the Jaccard distance and a 0.97 similarity threshold)

The results from Huseby Kiev allow us to finally connect the SHG group with the eastern pressure blade technology. However, the higher genetic affinity between Huseby Kiev individuals and the WHG group challenges the earlier suggested tie between eastern technology and EHG genetics. Our results suggest either early cultural transmission, or a more complex course of events involving both non- and co-dependent cultural and genetic admixture.

huseby-kiev

Seeing how culture is indeed usually associated with the expansion of a certain population, especially at such an early date, I guess this similarity with WHG of incoming eastern peoples comes from an originally EHG population expanding into a mainly WHG area in the west (similar to what happens e.g. with Bell Beakers), or being replaced later by a WHG population which adopted the culture (similar to what happened with late Corded Ware populations in central-east Europe after the expansion of Bell Beakers).

Unlike later periods, it will always be difficult to judge such ancient population movements with few samples covering thousands of years… Probably specific Y-DNA haplogroups would help differentiate between both expanding populations from east and west.

Related

Minimal gene flow from western pastoralists in the Bronze Age eastern steppes

jeong-steppes-mongolia

Open access paper Bronze Age population dynamics and the rise of dairy pastoralism on the eastern Eurasian steppe, by Jeong et al. PNAS (2018).

Interesting excerpts (emphasis mine):

To understand the population history and context of dairy pastoralism in the eastern Eurasian steppe, we applied genomic and proteomic analyses to individuals buried in Late Bronze Age (LBA) burial mounds associated with the Deer Stone-Khirigsuur Complex (DSKC) in northern Mongolia. To date, DSKC sites contain the clearest and most direct evidence for animal pastoralism in the Eastern steppe before ca. 1200 BCE.

Most LBA Khövsgöls are projected on top of modern Tuvinians or Altaians, who reside in neighboring regions. In comparison with other ancient individuals, they are also close to but slightly displaced from temporally earlier Neolithic and Early Bronze Age (EBA) populations from the Shamanka II cemetry (Shamanka_EN and Shamanka_EBA, respectively) from the Lake Baikal region. However, when Native Americans are added to PC calculation, we observe that LBA Khövsgöls are displaced from modern neighbors toward Native Americans along PC2, occupying a space not overlapping with any contemporary population. Such an upward shift on PC2 is also observed in the ancient Baikal populations from the Neolithic to EBA and in the Bronze Age individuals from the Altai associated with Okunevo and Karasuk cultures.

pca-eurasians-karasuk-khovsgol
Image modified from the article. Karasuk cluster in green, closely related to sample ARS026 in red. Principal Component Analysis (PCA) of selected 2,077 contemporary Eurasians belonging to 149 groups. Contemporary individuals are plotted using three-letter abbreviations for operational group IDs. Group IDs color coded by geographic region. Ancient Khövsgöl individuals and other selected ancient groups are represented on the plot by filled shapes. Ancient individuals are projected onto the PC space using the “lsqproject: YES” option in the smartpca program to minimize the impact of high genotype missing rate.

(…) two individuals fall on the PC space markedly separated from the others: ARS017 is placed close to ancient and modern northeast Asians, such as early Neolithic individuals from the Devil’s Gate archaeological site (22) and present-day Nivhs from the Russian far east, while ARS026 falls midway between the main cluster and western Eurasians.

Upper Paleolithic Siberians from nearby Afontova Gora and Mal’ta archaeological sites (AG3 and MA-1, respectively) (25, 26) have the highest extra affinity with the main cluster compared with other groups, including the eastern outlier ARS017, the early Neolithic Shamanka_EN, and present-day Nganasans and Tuvinians (Z > 6.7 SE for AG3). Main cluster Khövsgöl individuals mostly belong to Siberian mitochondrial (A, B, C, D, and G) and Y (all Q1a but one N1c1a) haplogroups.

mongolia-botai-ehg-ane-cline
The genetic affinity of the Khövsgöl clusters measured by outgroup-f3 and -f4 statistics. (A) The top 20 populations sharing the highest amount of >genetic drift with the Khövsgöl main cluster measured by f3(Mbuti; Khövsgöl, X). (B) The top 15 populations with the most extra affinity with each of the three Khövsgöl clusters in contrast to Tuvinian (for the main cluster) or to the main cluster (for the two outliers), measured by f4(Mbuti, X; Tuvinian/Khövsgöl, Khövsgöl/ARS017/ARS026). Ancient and contemporary groups are marked by squares and circles, respectively. Darker shades represent a larger f4 statistic.

Previous studies show a close genetic relationship between WSH populations and ANE ancestry, as Yamnaya and Afanasievo are modeled as a roughly equal mixture of early Holocene Iranian/ Caucasus ancestry (IRC) and Mesolithic Eastern European hunter-gatherers, the latter of which derive a large fraction of their ancestry from ANE. It is therefore important to pinpoint the source of ANE-related ancestry in the Khövsgöl gene pool: that is, whether it derives from a pre-Bronze Age ANE population (such as the one represented by AG3) or from a Bronze Age WSH population that has both ANE and IRC ancestry.

The amount of WSH contribution remains small (e.g., 6.4 ± 1.0% from Sintashta). Assuming that the early Neolithic populations of the Khövsgöl region resembled those of the nearby Baikal region, we conclude that the Khövsgöl main cluster obtained ∼11% of their ancestry from an ANE source during the Neolithic period and a much smaller contribution of WSH ancestry (4–7%) beginning in the early Bronze Age.

khovsgol-shamanka-sintashta
Admixture modeling of Altai populations and the Khövsgöl main cluster using qpAdm. For the archaeological populations, (A) Shamanka_EBA and (B and C) Khövsgöl, each colored block represents the proportion of ancestry derived from a corresponding ancestry source in the legend. Error bars show 1 SE. (A) Shamanka_EBA is modeled as a mixture of Shamanka_EN and AG3. The Khövsgöl main cluster is modeled as (B) a two-way admixture of Shamanka_EBA+Sintashta and (C) a three-way admixture Shamanka_EN+AG3+Sintashta.

Apparently, then, the first individual with substantial WSH ancestry in the Khövsgöl population (ARS026, of haplogroup R1a-Z2123), directly dated to 1130–900 BC, is consistent with the first appearance of admixed forest-steppe-related populations like Karasuk (ca. 1200-800 BC) in the Altai. Interestingly, haplogroup N1a1a-M178 pops up (with mtDNA U5a2d1) among the earlier Khövsgöl samples.

I will repeat what I wrote recently here: Samoyedic arrived in the Altai with Karasuk and hg R1a-Z645 + Steppe_MLBA-like ancestry, admixed with Altai populations, clustering thus within an Ancient Altai cline. Only later did N1a1a subclades infiltrate Samoyedic (and Ugric) populations, bringing them closer to their modern Palaeo-Siberian cline. The shared mtDNA may support an ancestral EHG-“Siberian” cline, or else a more recent Afanasevo-related origin.

east-uralic-clines
Modified image from Jeong et al. (2018), supplementary materials. The first two PCs summarizing the genetic structure within 2,077 Eurasian individuals. The two PCs generally mirror geography. PC1 separates western and eastern Eurasian populations, with many inner Eurasians in the middle. PC2 separates eastern Eurasians along the north-south cline and also separates Europeans from West Asians. Ancient individuals (color-filled shapes), including two Botai individuals, are projected onto PCs calculated from present-day individuals. Read more.

Also interesting, Q1a2 subclades and ANE ancestry making its appearance everywhere among ancestral Eurasian peoples, as Chetan recently pointed out.

Related

“Steppe ancestry” step by step: Khvalynsk, Sredni Stog, Repin, Yamna, Corded Ware

dzudzuana_pca-large

Wang et al. (2018) is obviously a game changer in many aspects. I have already written about the upcoming Yamna Hungary samples, about the new Steppe_Eneolithic and Caucasus Eneolithic keystones, and about the upcoming Greece Neolithic samples with steppe ancestry.

An interesting aspect of the paper, hidden among so many relevant details, is a clearer picture of how the so-called Yamnaya or steppe ancestry evolved from Samara hunter-gatherers to Yamna nomadic pastoralists, and how this ancestry appeared among Proto-Corded Ware populations.

anatolia-neolithic-steppe-eneolithic
Image modified from Wang et al. (2018). Marked are in orange: equivalent Steppe_Maykop ADMIXTURE; in red, approximate limit of Anatolia_Neolithic ancestry found in Yamna populations; in blue, Corded Ware-related groups. “Modelling results for the Steppe and Caucasus cluster. Admixture proportions based on (temporally and geographically) distal and proximal models, showing additional Anatolian farmer-related ancestry in Steppe groups as well as additional gene flow from the south in some of the Steppe groups as well as the Caucasus groups.”

Please note: arrows of “ancestry movement” in the following PCAs do not necessarily represent physical population movements, or even ethnolinguistic change. To avoid misinterpretations, I have depicted arrows with Y-DNA haplogroup migrations to represent the most likely true ethnolinguistic movements. Admixture graphics shown are from Wang et al. (2018), and also (the K12) from Mathieson et al. (2018).

1. Samara to Early Khvalynsk

The so-called steppe ancestry was born during the Khvalynsk expansion through the steppes, probably through exogamy of expanding elite clans (eventually all R1b-M269 lineages) originally of Samara_HG ancestry. The nearest group to the ANE-like ghost population with which Samara hunter-gatherers admixed is represented by the Steppe_Eneolithic / Steppe_Maykop cluster (from the Northern Caucasus Piedmont).

Steppe_Eneolithic samples, of R1b1 lineages, are probably expanded Khvalynsk peoples, showing thus a proximate ancestry of an Early Eneolithic ghost population of the Northern Caucasus. Steppe_Maykop samples represent a later replacement of this Steppe_Eneolithic population – and/or a similar population with further contribution of ANE-like ancestry – in the area some 1,000 years later.

PCA-caucasus-steppe-samara

This is what Steppe_Maykop looks like, different from Steppe_Eneolithic:

steppe-maykop-admixture

NOTE. This admixture shows how different Steppe_Maykop is from Steppe_Eneolithic, but in the different supervised ADMIXTURE graphics below Maykop_Eneolithic is roughly equivalent to Eneolithic_Steppe (see orange arrow in ADMIXTURE graphic above). This is useful for a simplified analysis, but actual differences between Khvalynsk, Sredni Stog, Afanasevo, Yamna and Corded Ware are probably underestimated in the analyses below, and will become clearer in the future when more ancestral hunter-gatherer populations are added to the analysis.

2. Early Khvalynsk expansion

We have direct data of Khvalynsk-Novodanilovka-like populations thanks to Khvalynsk and Steppe_Eneolithic samples (although I’ve used the latter above to represent the ghost Caucasus population with which Samara_HG admixed).

We also have indirect data. First, there is the PCA with outliers:

PCA-khvalynsk-steppe

Second, we have data from north Pontic Ukraine_Eneolithic samples (see next section).

Third, there is the continuity of late Repin / Afanasevo with Steppe_Eneolithic (see below).

3. Proto-Corded Ware expansion

It is unclear if R1a-M459 subclades were continuously in the steppe and resurged after the Khvalynsk expansion, or (the most likely option) they came from the forested region of the Upper Dnieper area, possibly from previous expansions there with hunter-gatherer pottery.

Supporting the latter is the millennia-long continuity of R1b-V88 and I2a2 subclades in the north Pontic Mesolithic, Neolithic, and Early Eneolithic Sredni Stog culture, until ca. 4500 BC (and even later, during the second half).

Only at the end of the Early Eneolithic with the disappearance of Novodanilovka (and beginning of the steppe ‘hiatus’ of Rassamakin) is R1a to be found in Ukraine again (after disappearing from the record some 2,000 years earlier), related to complex population movements in the north Pontic area.

NOTE. In the PCA, a tentative position of Novodanilovka closer to Anatolia_Neolithic / Dzudzuana ancestry is selected, based on the apparent cline formed by Ukraine_Eneolithic samples, and on the position and ancestry of Sredni Stog, Yamna, and Corded Ware later. A good alternative would be to place Novodanilovka still closer to the Balkan outliers (i.e. Suvorovo), and a source closer to EHG as the ancestry driven by the migration of R1a-M417.

PCA-sredni-stog-steppe

The first sample with steppe ancestry appears only after 4250 BC in the forest-steppe, centuries after the samples with steppe ancestry from the Northern Caucasus and the Balkans, which points to exogamy of expanding R1a-M417 lineages with the remnants of the Novodanilovka population.

steppe-ancestry-admixture-sredni-stog

4. Repin / Early Yamna expansion

We don’t have direct data on early Repin settlers. But we do have a very close representative: Afanasevo, a population we know comes directly from the Repin/late Khvalynsk expansion ca. 3500/3300 BC (just before the emergence of Early Yamna), and which shows fully Steppe_Eneolithic-like ancestry.

afanasevo-admixture

Compared to this eastern Repin expansion that gave Afanasevo, the late Repin expansion to the west ca. 3300 BC that gave rise to the Yamna culture was one of colonization, evidenced by the admixture with north Pontic (Sredni Stog-like) populations, no doubt through exogamy:

PCA-repin-yamna

This admixture is also found (in lesser proportion) in east Yamna groups, which supports the high mobility and exogamy practices among western and eastern Yamna clans, not only with locals:

yamnaya-admixture

5. Corded Ware

Corded Ware represents a quite homogeneous expansion of a late Sredni Stog population, compatible with the traditional location of Proto-Corded Ware peoples in the steppe-forest/forest zone of the Dnieper-Dniester region.

PCA-latvia-ln-steppe

We don’t have a comparison with Ukraine_Eneolithic or Corded Ware samples in Wang et al. (2018), but we do have proximate sources for Abashevo, when compared to the Poltavka population (with which it admixed in the Volga-Ural steppes): Sintashta, Potapovka, Srubna (with further Abashevo contribution), and Andronovo:

sintashta-poltavka-andronovo-admixture

The two CWC outliers from the Baltic show what I thought was an admixture with Yamna. However, given the previous mixture of Eneolithic_Steppe in north Pontic steppe-forest populations, this elevated “steppe ancestry” found in Baltic_LN (similar to west Yamna) seems rather an admixture of Baltic sub-Neolithic peoples with a north Pontic Eneolithic_Steppe-like population. Late Repin settlers also admixed with a similar population during its colonization of the north Pontic area, hence the Baltic_LN – west Yamna similarities.

NOTE. A direct admixture with west Yamna populations through exogamy by the ancestors of this Baltic population cannot be ruled out yet (without direct access to more samples), though, because of the contacts of Corded Ware with west Yamna settlers in the forest-steppe regions.

steppe-ancestry-admixture-latvia

A similar case is found in the Yamna outlier from Mednikarovo south of the Danube. It would be absurd to think that Yamna from the Balkans comes from Corded Ware (or vice versa), just because the former is closer in the PCA to the latter than other Yamna samples. The same error is also found e.g. in the Corded Ware → Bell Beaker theory, because of their proximity in the PCA and their shared “steppe ancestry”. All those theories have been proven already wrong.

NOTE. A similar fallacy is found in potential Sintashta→Mycenaean connections, where we should distinguish statistically that result from an East/West Yamna + Balkans_BA admixture. In fact, genetic links of Mycenaeans with west Yamna settlers prove this (there are some related analyses in Anthrogenica, but the site is down at this moment). To try to relate these two populations (separated more than 1,000 years before Sintashta) is like comparing ancient populations to modern ones, without the intermediate samples to trace the real anthropological trail of what is found…Pure numbers and wishful thinking.

Conclusion

Yamna and Corded Ware show a similar “steppe ancestry” due to convergence. I have said so many times (see e.g. here). This was clear long ago, just by looking at the Y-chromosome bottlenecks that differentiate them – and Tomenable noticed this difference in ADMIXTURE from the supplementary materials in Mathieson et al. (2017), well before Wang et al. (2018).

This different stock stems from (1) completely different ancestral populations + (2) different, long-lasting Y-chromosome bottlenecks. Their similarities come from the two neighbouring cultures admixing with similar populations.

If all this does not mean anything, and each lab was going to support some pre-selected archaeological theories from the 1960s or the 1980s, coupled with outdated linguistic models no matter what – Anthony’s model + Ringe’s glottochronological tree of the early 2000s in the Reich Lab; and worse, Kristiansen’s CWC-IE + Germano-Slavonic models of the 1940s in the Copenhagen group – , I have to repeat my question again:

What’s (so much published) ancient DNA useful for, exactly?

See also

Related

Dzudzuana, Sidelkino, and the Caucasus contribution to the Pontic-Caspian steppe

hunter-gatherer-pottery

It has been known for a long time that the Caucasus must have hosted many (at least partially) isolated populations, probably helped by geographical boundaries, setting it apart from open Eurasian areas.

David Reich writes in his book the following about India:

The genetic data told a clear story. Around a third of Indian groups experienced population bottlenecks as strong or stronger than the ones that occurred among Finns or Ashkenazi Jews. We later confirmed this finding in an even larger dataset that we collected working with Thangaraj: genetic data from more than 250 jati groups spread throughout India (…)

Rather than an invention of colonialism as Dirks suggested, long-term endogamy as embodied in India today in the institution of caste has been overwhelmingly important for millennia. (…)

The Han Chinese are truly a large population. They have been mixing freely for thousands of years. In contrast, there are few if any Indian groups that are demographically very large, and the degree of genetic differentiation among Indian jati groups living side by side in the same village is typically two to three times higher than the genetic differentiation between northern and southern Europeans. The truth is that India is composed of a large number of small populations.

There is little doubt now, based on findings spanning thousands of years, that the Mesolithic and Neolithic Caucasus hosted various very small populations, even if the ancestral components may be reduced to the few known to date (such as ANE, EHG, AME*, ENA, CHG, and other “deep” ancestral components).

NOTE. I will call the ancestral component of Dzudzuana/Anatolian hunter-gatherers Ancient Middle Easterner (AME), to give a clear idea of its likely extension during the Late Upper Palaeolithic, and to avoid using the more simplistic Dzudzuana, unless it is useful to mention these specific local samples.

dzudzuana-pca
Image modified from Lazaridis et al. (2018), including Caucasus, Don-Volga-Ural, and North Pontic Mesolithic-Neolithic populations. “Ancient West Eurasian population structure. (a) Geographical distribution of key ancient West Eurasian populations. (b) Temporal distribution of key ancient West Eurasian populations (approximate date in ky BP). (c) PCA of key ancient West Eurasians, including additional populations (shown with grey shells), in the space of outgroup f4-statistics (Methods).”

Genetic labs have a strong fixation with ancestry. I guess the use of complex statistical methods gives professionals and laymen alike the feeling of dealing with “Science”, as opposed to academic fields where you have to interpret data. I think language reveals a lot about the way people think, and the fact that ancestral components are called ‘lineages’ – while not wrong per se – is a clear symptom of the lack of interest in the true lineages: Y-DNA haplogroups.

Y-DNA bottlenecks

It has become quite clear that male-biased migrations are often the ones which can be confidently followed for actual population movements and ethnolinguistic identification, at least until the Iron Age. The frequently used Palaeolithic clusters offer a clear example of why ancestry does not represent what some people believe: They merely give a basic idea of sizeable population replacements by distant peoples.

Both concepts are important: sizeable and distant peoples. For example, during the Upper Palaeolithic in Europe there was a sizeable population replacement of the Aurignacian Goyet cluster by the Gravettian Vestonice cluster (probably from populations of far eastern Russia) coupled with the arrival of haplogroup I, although during the thousands of years that this material culture lasted, the previously expanded C1a2 lineages did not disappear, and there were probably different resurgence and admixture events.

Haplogroup I certainly expanded with the Gravettian culture to Iberia, where the Goyet ancestry did not change much – probably because of male-driven migrations -, to the extent that during the Magdalenian expansions haplogroup I expanded with an ancestry closer to Goyet, in what is called a ‘resurge’ of the Goyet cluster – even though there is a clear replacement of male lines.

The Villabruna (WHG) cluster is another good example. It probably spread with haplogroup R1b-L754, which – based on the extra ‘East Asian’ affinity of some samples and on modern samples from the Middle East – came probably from the east through a southern route, and not too long before the expansion of WHG likely from around the Black Sea, although this is still unclear. The finding of haplogroup I in samples of mostly WHG ancestry could confuse people that do not care about timing, sub-structured populations, and gene flow.

palaeolithic-expansions-reich
Image from David Reich’s Who We Are and How We Got Here. Having migrated out of Africa and the Near East, modern human pioneer populations spread throughout Eurasia (1). By at least thirty-nine thousand years ago, one group founded a lineage of European hunter-gatherers that persisted largely uninterrupted for more than twenty thousand years (2). Eventually, groups derived from an eastern branch of this founding population of European huntergatherers spread west (3), displaced previous groups, and were eventually themselves pushed out of northern Europe by the spread of glacial ice, shown at its maximum extent (top right). As the glaciers receded, western Europe was repeopled from the southwest (4) by a population that had managed to persist for tens of thousands of years and was related to an approximately thirty-five-thousand-year old individual from far western Europe. A later human migration, following the first strong warming period, had an even larger impact, with a spread from the southeast (5) that not only transformed the population of western Europe but also homogenized the populations of Europe and the Near East. At a single site—Goyet Caves in Belgium—ancient DNA from individuals spread over twenty thousand years reflects these transformations, with representatives from the Aurignacian, Gravettian, and Magdalenian periods.

NOTE. If you don’t understand why ‘clusters’ that span thousands of years don’t really matter for the many Palaeolithic population expansions that certainly happened among hunter-gatherers in Europe, just take a look at what happened with Bell Beakers expanding from Yamna into western Europe within 500 years.

If we don’t thread carefully when talking about population migrations, these terms are bound to confuse people. Just as the fixation on “steppe ancestry” – which marks the arrival in Chalcolithic Europe of peoples from the Pontic-Caspian region – has confused a lot of researchers to this day.

When I began to write about the Indo-European demic diffusion model, my concern was to find a single spot where a North-West Indo-European proto-language could have expanded from ca. 2000 BC (our most common guesstimate). Based on the 2015 papers, and in spite of their conclusions, I thought it had become clear that Corded Ware was not it, and it was rather Bell Beakers. I assumed that Uralic was spoken to the north (as was the traditional belief), and thus Corded Ware expanded from the forest zone, hence steppe ancestry would also be found there with other R1a lineages.

With the publication of Mathieson et al. (2017) and Olalde et al. (2017), I changed my mind, seeing how “steppe ancestry” did in fact appear quite late, hence it was likely to be the result of very specific population movements, probably directly from the Caucasus. Later, Mathieson published in a revision the sample from Alexandria of hg R1a-M417 (probably R1a-Z645, possibly Z93+), which further supported the idea that the migration of Corded Ware peoples started near the North Pontic forest-steppe (as I included in a the next revision).

The question remains the same I repeated recently, though: where do the extra Caucasus components (i.e. beyond EHG) of Eneolithic Ukraine/Corded Ware and Khvalynsk/Yamna come from?

Steppe ancestry: “EHG” + “CHG”?

About EHG ancestry

From Lazaridis et al. (2018):

Considering 2-way mixtures, we can model Karelia_HG as deriving 34 ± 2.8% of its ancestry from a Villabruna-related source, with the remainder mainly from ANE represented by the AfontovaGora3 (AG3) sample from Lake Baikal ~17kya.

AG3 was likely of haplogroup Q1a (as reported by YFull, see Genetiker), and probably the ANE ancestry found in Eastern Europe accompanied a Palaeolithic migration of Q1a2-M25 (formed ca. 22600 BC, TMRCA ca. 14300 BC).

NOTE. You can read more about the expansion of Q lineages during the Palaeolithic.

Combined with what we know about the Eneolithic Steppe and Caucasus populations – it is likely that ANE ancestry remained the most important component of some of the small ghost populations of the Caucasus until their emergence with the Lola culture.

pca-caucasus-dzudzuana
Image modified from Wang et al. (2018). Samples projected in PCA of 84 modern-day West Eurasian populations (open symbols). Previously known clusters have been marked and referenced. Marked and labelled are the Balkan samples referenced in this text An EHG and a Caucasus ‘clouds’ have been drawn, leaving Pontic-Caspian steppe and derived groups between them. See the original file here. To understand the drawn potential Caucasus Mesolithic cluster, see above the PCA from Lazaridis et al. (2018).

The first sample we have now attributed to the EHG cluster is Sidelkino, from the Samara region (ca. 9300 BC), mtDNA U5a2. In Damgaard et al. (Science 2018), Yamnaya could be modelled as a CHG population related to Kotias Klde (54%) and the remaining from ANE population related to Sidelkino (>46%), with the following split events:

  1. A split event, where the CHG component of Yamnaya splits from KK1. The model inferred this time at 27 kya (though we note the larger models in Sections S2.12.4 and S2.12.5 inferred a more recent split time).
  2. A split event, where the ANE component of Yamnaya splits from Sidelkino. This was inferred at about about 11 kya.
  3. A split event, where the ANE component of Yamnaya splits from Botai. We inferred this to occur 17 kya. Note that this is above the Sidelkino split time, so our model infers Yamnaya to be more closely related to the EHG Sidelkino, as expected.
  4. An ancestral split event between the CHG and ANE ancestral populations. This was inferred to occur around 40 kya.

Other samples classified as of the EHG cluster:

  • Popovo2 (ca. 6250 BC) of hg J1, mtDNA U4d – Po2 and Po4 from the same site (ca. 6550 BC) show continuity of mtDNA.
  • Karelia_HG, from Juzhnii Oleni Ostrov (ca. 6300 BC): I0211/UzOO40 (ca. 6300 BC) of hg J1(xJ1a), mtDNA U4a; and I0061/UzOO74 of hg R1a1(xR1a1a), mtDNA C1
  • UzOO77 and UzOO76 from Juzhnii Oleni Ostrov (ca. 5250 BC) of mtDNA R1b.
  • Samara_HG from Lebyanzhinka (ca. 5600 BC) of hg R1b1a, mtDNA U5a1d.

From the analysis of Lazaridis et al. (2018), we have some details about their admixture:

dzudzuana-admixture-sidelkino
Image modified from Lazaridis et al. (2018). Modeling present-day and ancient West-Eurasians. Mixture proportions computed with qpAdm (Supplementary Information section 4). The proportion of ‘Mbuti’ ancestry represents the total of ‘Deep’ ancestry from lineages that split prior to the split of Ust’Ishim, Tianyuan, and West Eurasians and can include both ‘Basal Eurasian’ and other (e.g., Sub-Saharan African) ancestry. (Left) ‘Conservative’ estimates. Each population 367 cannot be modeled with fewer admixture events than shown. (Right) ‘Speculative’ estimates. The highest number of sources (≤5) with admixture estimates within [0,1] are shown for each population. Some of the admixture proportions are not significantly different from 0 (Supplementary Information section 4).

About Anatolia_Neolithic ancestry

About the enigmatic Anatolia_Neolithic-related ancestry found in Pontic-Caspian steppe samples, this is what Wang et al. (2018) had to say:

We focused on model of mixture of proximal sources such as CHG and Anatolian Chalcolithic for all six groups of the Caucasus cluster (Eneolithic Caucasus, Maykop and Late Makyop, Maykop-Novosvobodnaya, Kura-Araxes, and Dolmen LBA), with admixture proportions on a genetic cline of 40-72% Anatolian Chalcolithic related and 28-60% CHG related (Supplementary Table 7). When we explored Romania_EN and Greece_Neolithic individuals as alternative southeast European sources (30-46% and 36-49%), the CHG proportions increased to 54-70% and 51-64%, respectively. We hypothesize that alternative models, replacing the Anatolian Chalcolithic individual with yet unsampled populations from eastern Anatolia, South Caucasus or northern Mesopotamia, would probably also provide a fit to the data from some of the tested Caucasus groups.

Also:

The first appearance of ‘Near Eastern farmer related ancestry’ in the steppe zone is evident in Steppe Maykop outliers. However, PCA results also suggest that Yamnaya and later groups of the West Eurasian steppe carry some farmer related ancestry as they are slightly shifted towards ‘European Neolithic groups’ in PC2 (Fig. 2D) compared to Eneolithic steppe. This is not the case for the preceding Eneolithic steppe individuals. The tilting cline is also confirmed by admixture f3-statistics, which provide statistically negative values for AG3 as one source and any Anatolian Neolithic related group as a second source

yamnaya-caucasus-dzudzuana
Modified image from Wang et al. (2018). In blue, Yamna-related populations. In red, Corded Ware-related populations, and two elevated Anatolia_Neolithic values in Yamna. Notice how only GAC-related admixture increases the Anatolian_N-related ancestry in the Yamna outlier from Ozero, and the late Yamna sample from Hungary, related to the homogeneous Yamna population. “Supplementary Table 14. P values of rank=3 and admixture proportions in modelling Steppe ancestry populations as a four-way admixture of distal sources EHG, CHG, Anatolian_Neolithic and WHG using 14 outgroups.Left populations: Steppe cluster, EHG, CHG, WHG, Anatolian_Neolithic. Right populations: Mbuti.DG, Ust_Ishim.DG, Kostenki14, MA1, Han.DG, Papuan.DG, Onge.DG, Villabruna, Vestonice16, ElMiron, Ethiopia_4500BP.SG, Karitiana.DG, Natufian, Iran_Ganj_Dareh_Neolithic.”

Detailed exploration via D-statistics in the form of D(EHG, steppe group; X, Mbuti) and D(Samara_Eneolithic, steppe group; X, Mbuti) show significantly negative D values for most of the steppe groups when X is a member of the Caucasus cluster or one of the Levant/Anatolia farmer-related groups (Supplementary Figs. 5 and 6). In addition, we used f- and D-statistics to explore the shared ancestry with Anatolian Neolithic as well as the reciprocal relationship between Anatolian- and Iranian farmer-related ancestry for all groups of our two main clusters and relevant adjacent regions (Supplementary Fig. 4). Here, we observe an increase in farmer-related ancestry (both Anatolian and Iranian) in our Steppe cluster, ranging from Eneolithic steppe to later groups. In Middle/Late Bronze Age groups especially to the north and east we observe a further increase of Anatolian farmer related ancestry consistent with previous studies of the Poltavka, Andronovo, Srubnaya and Sintashta groups and reflecting a different process not especially related to events in the Caucasus.

(…) Surprisingly, we found that a minimum of four streams of ancestry is needed to explain all eleven steppe ancestry groups tested, including previously published ones (Fig. 2; Supplementary Table 12). Importantly, our results show a subtle contribution of both Anatolian farmer-related ancestry and WHG-related ancestry (Fig.4; Supplementary Tables 13 and 14), which was likely contributed through Middle and Late Neolithic farming groups from adjacent regions in the West. The discovery of a quite old AME ancestry has rendered this probably unnecessary, because this admixture from an Anatolian-like ghost population could be driven even by small populations from the Caucasus.

yamna-caucasus-cwc-anatolia-neolithic
Image modified from Wang et al. (2018). Marked are: in red, approximate limit of Anatolia_Neolithic ancestry found in Yamna populations; in blue, Corded Ware-related groups. “Modelling results for the Steppe and Caucasus 1128 cluster. Admixture proportions based on (temporally and geographically) distal and proximal models, showing additional Anatolian farmer-related ancestry in Steppe groups as well as additional gene flow from the south in some of the Steppe groups as well as the Caucasus groups (see also Supplementary Tables 10, 14 and 20).”

NOTE. For a detailed account of the possibilities regarding this differential admixture in the North Pontic area in contrast to the Don-Volga-Ural region, you can read the posts Sredni Stog, Proto-Corded Ware, and their “steppe admixture”, and Corded Ware culture origins: The Final Frontier.

While it is not yet fully clear, the increased Anatolian_Neolithic-like ancestry in Ukraine_Eneolithic samples (see below) makes it unlikely that all such ancestry in Corded Ware groups comes from a GAC-related contribution. It is likely that at least part of it represents contributions from populations of the Caucasus, based on the mostly westward population movements in the steppe from ca. 4600 BC on, including the Suvorovo-Novodanilovka expansion, and especially the Kuban-Maykop expansion during the final Eneolithic into the North Pontic area.

NOTE. Since CHG-like groups from the Caucasus may have combinations of AME and ANE ancestry similar to Yamna (which may thus appear as ‘steppe ancestry’ in the North Pontic area), it is impossible to interpret with precision the following ADMIXTURE graphic:

ukraine-whg-ehg-steppe
Modified image from Mathieson et al. (2018). Supervised ADMIXTURE analysis, modelling each ancient individual (one per row) as a mixture of population clusters constrained to contain northwestern-Anatolian Neolithic (grey), Yamnaya from Samara (yellow), EHG (pink) and WHG (green) populations. Dates in parentheses indicate approximate range of individuals in each population.

North-Eastern Technocomplex

The East Asian contribution to samples from the WHG samples (like Loschbour or La Braña), as specified in Fu et al. (2016), does not seem to be related to Baikal_EN, and appears possibly (in the ADMIXTURE analysis) integrated into he Villabruna component. I guess this implies that the shared alleles with East Asians are quite early, and potentially due to the expansion of R1b-L754 from the East.

It would be interesting to know the specific material culture Sidelkino belonged to – i.e. if it was related to the expansion of the North-Eastern Technocomplex – , and its Y-DNA. The Post-Swiderian expansion into eastern Europe, probably associated with the expansion of R1b-P297 lineages (including R1b-M73, found later in Botai and in Baltic HG) is supposed to have begun during the 11th millennium BC, but migrations to the Urals and beyond are probably concentrated in the 9th millennium, so this sample is possibly slightly early for R1b.

NOTE. User Rozenfeld at Anthrogenica posted this, which I think is interesting (in case anyone wants to try a Y-SNP call):

there is something strange with Sidelkino EHG: first, its archaeological context is not described in the supplementary. Second, its sex is not listed in the supplementary tables. Third, after looking for info about this sample, I found that: “Сиделькино-3. Для снятия вопроса о половой принадлежности индивида была проведена генетическая экспертиза, выявившая принадлежность останков мужчине.”(translation: Sidelkino-3. To resolve the question about sex of the remains, the genetic analysis was conducted, which showed that remains belonged to male), source: http://static.iea.ras.ru/books/7487_Traditsii.pdf

So either they haven’t mentioned his Y-DNA in the paper for some reason, or there are more than one Sidelkino sample and the male one has not yet been published. The coverage of the Sidelkino sample from the paper is 2.9, more than enough to tell Y-DNA haplogroup.

zaliznyak-post-swiderian
The map of spreading of Post-Swiderian and Post-Krasnosillian sites in Mesolithic of Eastern Europe in the 8th millennia BC. From Zaliznyak (see here).

My speculative guess right now about specific population movements in far eastern Europe, based on the few data we have:

  • The expansion of the North-Eastern Technocomplex first around the 9th millennium BC, most likely expanded R1b-P279 ca. 11300 BC, judging by its TMRCA, with both R1b-M73 (TMRCA 5300) and R1b-M269 (TMRCA 4400 BC) info (with extra El Mirón ancestry) back, and thus Eurasiatic.
  • The expansion of haplogroup J1 to the north may have happened before or after the R1b-P279 expansion. Judging by the increase in AG3-related ancestry near Karelia compared to Baltic_HG, it is possible that it expanded just after R1b-P279 (hence possibly J1-Y6304? TMRCA 9700 BC). Its long-lasting presence in the Caucasus is supported by the Satsurblia (ca. 11300 BC) and the Dolmen BA (ca. 1300 BC) samples.
  • The expansion of R1a-M17 ca. 6600 BC is still likely to have happened from the east, based on the R1a-M17 samples found in Baikalic cultures slightly later (ca. 5300 BC). The presence of elevated Baikal_EN ancestry in Karelia HG and in Samara HG, and the finding of R1a-M417 samples in the Forest Zone after the Mesolithic suggests a connection with the expansion of Hunter-Gatherer pottery, from the Elshanka culture in the Samara region northward into the Forset Zone and westward into the North Pontic area.
  • The expansion of R1b-M73 ca. 5300 BC is likely to be associated with the emergence of a group east of the Urals (related to the later Botai culture, and potentially Pre-Yukaghir). Its presence in a Narva sample from Donkalnis (ca. 5200 BC) suggest either an early split and spread of both R1b-P297 lineages (M73 and M269) through Eastern Europe, or maybe a back-migration with hunter-gatherer pottery.
  • R1b-M269 spread successfully ca. 4400 BC (and R1b-L23 ca. 4100 BC, both based on TMRCA), and this successful expansion is probably to be associated with the Khvalynsk-Novodanilovka expansion. We already know that Samara_HG ca. 5600 was R1b1a, so it is likely that R1b-M269 appeared (or ‘resurged’) in the Volga-Ural region shortly after the expansion of R1a-M17, whose expansion through the region may be inferred by the additional AG3 and Baikal_EN ancestry. Interesting from Samara_HG compared to the previous Sidelkino sample is the introduction of more El Mirón-related ancestry, typical of WHG populations (and thus proper of Baltic groups).

NOTE. The TMRCA dates are obviously gross approximations, because a) the actual rate of mutation is unknown and b) TMRCA estimates are based on the convergence of lineages that survived. The potential finding of R1a-Z645 (possibly Z93+) in Ukraine Eneolithic (ca. 4000 BC), and the potential finding of R1b-L23 in Khvalynsk ca. 4250 BC complicates things further, in terms of dates and origins of any subclade.

The question thus remains as it was long ago: did R1b-M269 lineages expand (‘return’) from the east, near the Urals, or directly from the north? Were they already near Samara at the same time as the expansion of hunter-gatherer pottery, and were not much affected by it? Or did they ‘resurge’ from populations admixed with Caucasus-related ancestry after the expansion of R1a-M17 with this pottery (since there are different stepped expansions from the Samara region)? We could even ask, did R1a-M17 really expand from the east, i.e. are the dates on Baikalic subclades from Moussa et al. (2016) reliable? Or did R1a-M17 expand from some pockets in the Pontic-Caspian steppe, taking over the expansion of HG pottery at some point?

hunger-gatherer-pottery
Early Neolithic cultures in eastern and central Europe: 1–Yelshanian; 2–North Caspian; 3–Rakushechnyj Yar; 4–Surskian; 5–Dnieper-Donetsian; 6– Bug-Dniesterian; 7–Upper Volga; 8–Narvian; 9–Linear Pottery. White arrows: expansion of early farming; black arrows: spread of pottery-making traditions. From Dolukhanov et al. (2009).

Maglemose-related migrations

The most interesting aspect from the new paper (regarding Indo-Uralic migrations) is that Ancestral Middle Easterner ancestry will probably be a better proxy for the Anatolia_Neolithic component found in Ukraine Mesolithic to Eneolithic, and possibly also for some of the “more CHG-like” component found among Pontic-Caspian steppe populations, all likely derived from different admixture events with groups from the Caucasus.

NOTE. Even the supposed gene flow of Neolithic Iranian ancestry into the Caucasus can be put into question, since that means possibly a Dzudzuana-like population with greater “deep ancestry” proportion than the one found in CHG, which may still be found within the Caucasus.

If it was not clear already that following ‘steppe ancestry’ wherever it appears is a rather lame way of following Indo-European migrations, every single sample from the Caucasus and their admixture with Pontic-Caspian steppe populations will probably show that “steppe ancestry” is in fact formed by a variety of steppe-related ancestral components, impossible to follow coherently with a single population. Exactly what is happening already with the Siberian ancestry.

If the paper on the Dzudzuana samples has shown something, is that the expansion of an ANE-like population shook the entire Caucasus area up to the Zagros Mountains, creating this ANE – AME cline that are CHG and Iran_N, with further contributions of “deep ancestries” (probably from the south) complicating the picture further.

If this happens with few known samples, and we know of an ANE-like ghost population in the Caucasus (appearing later in the Lola culture), we can already guess that the often repeated “CHG component” found in Ukraine_Eneolithic and Khvalynsk will not be the same (except the part mediated by the Novodanilovka expansion).

This ANE-like expansion happened probably in the Late Upper Palaeolithic, and reached Northern Europe probably after the expansion of the Villabruna cluster (ca. 12000 BC), judging by the advance of AG3-like and ENA-like ancestry in later WHG samples.

The population movements during the Mesolithic and Early Neolithic in the North Pontic area are quite complicated: the extra AME ancestry is probably connected to the admixture with populations from the Caucasus, while the close similarity of Ukraine populations with Scandinavian ones (with an increase in Villabruna ancestry from Mesolithic to Neolithic samples), probably reveal population movements related to the expansion of Maglemose-related groups.

maglemose-mesolithic
Etno-cultural situation in Central and Eastern Europe in the Late Mesolithic — Early Neolithic (VI—V Mill. BC) (after Конча 2004: 201, карта 1; made after ideas by L. L. Zaliznyak). Legend: 1 — Maglemose circle in the VII Mill. BC (after Gr. Clark); 2—7 — Mesolithic cultures of the Post-Maglemose tradition, VI Mill. BC (after S. Kozłowsky, L. L. Zaliznyak): 2 — de Leyen-Wartena; 3 — Oldesloe — Godenaa; 4 — Chojnice — Peńki; 5 — Janisłavice; 6 — finds of Janisłavice artefacts outside of the main area; 7 — Donets culture; 8 — directions of the settling of Janisłavice people (after S. Kozłowsky and L. L. Zaliznyak); 9 — the south border of Mesolithic and Early Neolithic cultures of post-Swidrian and post-Arensburgian traditions; 10 — northern border of settlement of the Balkan-Danubian farmers; 11 — Bug- Dniester culture; 12 — Neolithic cultures emerged on the ethno-cultural basis of post-Maglemose: Э — Ertebölle-Ellerbeck, Н — Neman, Д — Dnieper-Donets, М — Mariupol (western variants). From Klein (2017).

These Maglemose-related groups were probably migrants from the north-west, originally from the Northern European Plains, who occupied the previous Swiderian territory, and then expanded into the North Pontic area. The overwhelming presence of I2a (likely all I2a2a1b1b) lineages in Ukraine Neolithic supports this migration.

The likely picture of Mesolithic-Neolithic migrations in the North Pontic area right now is then:

  1. Expansion of R1a-M459 from the east ca. 12000 BC – probably coupled with AG3 and also some Baikal_EN ancestry. First sample is I1819 from Vasilievka (ca. 8700 BC), another is from Dereivka ca. 6900 BC.
  2. Expansion of R1b-V88 from the Balkans in the west ca. 9700 BC, based on its TMRCA and also the Balkan hunter-gatherer population overwhemingly of this haplogroup from the 10th millennium until the Neolithic. First sample is I1734 from Vasilievka (ca. 7252 BC), which suggests that it replaced the male population there, based on their similar EHG-like adxmixture (and lack of sizeable WHG increase), and shared mtDNA U5b2, U5a2.
  3. Expansion of I2a-Y5606 probably ca. 6800 based on its TMRCA with Janislawice culture. Supporting this is the increase in WHG contribution to Neolithic samples, including the spread of U4 subclades compared to the previous period.
  4. Expansion of R1a-M17 starting probably ca. 6600 BC in the east (see above).

NOTE. The first sample of haplogroup I appears in the Mesolithic: I1763 (ca. 8100 BC) of haplogroup I2a1, probably related to an older Upper Palaeolithic expansion.

janislawice
Distribution of archeological cultures in the North Pontic Region during the Mesolithic (7th – 6th millennium BCE). Dotted, dashed and solid lines with corresponding arrows indicate alternative models of the spread of the Grebenyky culture groups. (After Bryuako IV., Samojlova TL., Eds, Drevnie kul’tury Severo-­‐Zapadnogo Prichernomor’ya, Odessa: SMIL, 2013.) Nikitin – Ivanova 2017.

Conclusion

It is becoming more and more clear with each new paper that – unless the number of very ancient samples increases – the use of Y-chromosome haplogroups remains one of the most important tools for academics; this is especially so in the steppes, in light of the diversity found in populations from the Caucasus. A clear example comes from the Yamna – Corded Ware similarities:

After the publication of the 2015 papers, it was likely that Yamna expanded with haplogroup R1b-L23, but it has only become crystal clear that Yamna expanded through the steppes into Bell Beakers, now that we have data about the strict genetic homogeneity of the whole Yamna population from west to east (including Afanasevo), in contrast with contemporary Corded Ware peoples which expanded from a different forest-steppe population.

The presence of haplogroups Q and R1a-M459 (xM17) in Khvalynsk along with a R1b1a sample, which some interpreted as being akin to modern ‘mixed’ populations in the past, is likely to point instead to a period of Khvalynsk-Novodanilovka expansion with R1b-M269, where different small populations from the steppe were being integrated into the common Khvalynsk stock, but where differences are seen in material culture surrounding their burials, as supported by the finding of R1b1 in the Kuban area already in the first half of the 5th millennium. The case would be similar to the early ‘mixed’ Icelandic population.

Only after the emergence of the Samara culture (in the second half of the 6th millennium BC), with a sample of haplogroup R1b1a, starts then the obvious connection with Early Proto-Indo-Europeans; and only after the appearance of late Sredni Stog and haplogroup R1a-M417 (ca. 4000 BC) is its connection with Uralic also clear. In previous population movements, I think more haplogroups were involved in migrations of small groups, and only some communities among them were eventually successful, expanding to be dominant, creating ever growing cultures during their expansions.

Indeed, if you think in terms of Uralic and Indo-European just as converging languages, and forget their potential genetic connection, then the genetic + linguistic picture becomes simplified, and the upper frontier of the 6th millennium BC with a division North Pontic (Mariupol) vs. Volga-Ural (Samara) is enough. However, tracing their movements backwards – with cultural expansions from west to east (with the expansion of farming), and earlier east to west (with hunter-gatherer pottery), and still earlier west to east (with the north-eastern technocomplex), offers an interesting way to prove their potential connection to macrofamilies, at least in terms of population movements.

corded-ware-uralic-qpgraph
Modified image from Tambets et al. (2018) Proportions of ancestral components in studied European and Siberian populations and the tested qpGraph model. a The qpGraph model fitting the data for the tested populations. Colour codes for the terminal nodes: pink—modern populations (‘Population X’ refers to test population) and yellow—ancient populations (aDNA samples and their pools). Nodes coloured other than pink or yellow are hypothetical intermediate populations. We putatively named nodes which we used as admixture sources using the main recipient among known populations. The colours of intermediate nodes on the qpGraph model match those on the admixture proportions panel. The NeolL (Neolithic Levant) ancestry selected in this qpGraph is likely to correspond (at least in part) to a specific Dzudzuana-like component present in the CHG-like population that admixed in the North Pontic area.

I am quite convinced right now that it would be possible to connect the expansion of R1b-L754 subclades with a speculative Nostratic (given the R1b-V88 connection with Afroasiatic, and the obvious connection of R1b-L297 with Eurasiatic). Paradoxically, the connection of an Indo-Uralic community in the steppes (after the separation of Yukaghir) with any lineage expansion (R1a-M17, R1b-M269, or even Q, I or J1) seems somehow blurrier than one year ago, possibly just because there are too many open possibilities.

David Reich says about the admixture with Neanderthals, which he helped discover:

At the conclusion of the Neanderthal genome project, I am still amazed by the surprises we encountered. Having found the first evidence of interbreeding between Neanderthals and modern humans, I continue to have nightmares that the finding is some kind of mistake. But the data are sternly consistent: the evidence for Neanderthal interbreeding turns out to be everywhere. As we continue to do genetic work, we keep encountering more and more patterns that reflect the extraordinary impact this interbreeding has had on the genomes of people living today.

I think this is a shared feeling among many of us who have made proposals about anything, to fear that we have made a gross, evident mistake, and constantly look for flaws. However, it seems to me that geneticists are more preoccupied with being wrong in their developed statistical methods, in the theoretical models they are creating, and not so much about errors in the true ancient ethnolinguistic picture human population genetics is (at least in theory) concerned about. Their publications are, after all, constantly associating genetic finds with cultures and (whenever possible) languages, so this aspect of their research should not be taken lightly.

Seeing how David Anthony or Razib Khan (among many others) have changed their previously preferred migration models as new data was published, and they continue to be respected in their own fields, I guess we can be confident that professionals with integrity are going to accept whatever new picture appears. While I don’t think that genetic finds can change what we can reconstruct with comparative grammar, I am also ready to revise guesstimates and routes of expansion of certain dialects if R1a-Z645 is shown to have accompanied Late Proto-Indo-Europeans during their expansion with Yamna, and later integrated somehow with Corded Ware.

However, taking into account the obsession of some with an ancestral, uninterrupted R1a—Indo-European association, and the lack of actual political repercussion of Neanderthal admixture, I think the most common nightmare that all genetic researchers should be worried about is to keep inflating this “Yamnaya ancestry”-based hornet’s nest, which has been constantly stirred up for the past two years, by rejecting it – or, rather, specifying it into its true complex nature.

This succession of corrections and redefinitions, coupled with the distinct Y-DNA bottleneck of each steppe population, will eventually lead to a completely different ethnolinguistic picture of the Pontic-Caspian region during the Eneolithic, which is likely to eventually piss off not only reasonable academics stubbornly attached to the CWC-IE idea, but also a part of those interested in daydreaming about their patrilineal ancestors.

Sometimes it’s better to just rip off the band-aid once and for all…

Featured image from The oldest pottery in hunter-gatherer communitiesand models of Neolithisation of Eastern Europe (2015), by Andrey Mazurkevich and Ekaterina Dolbunova.

Related

Interesting is today’s post in Ancient DNA Era: Is Male-driven Genetic Replacement always meaning Language-shift?