Interesting excerpts (emphasis mine; most internal references removed):
The earliest, most secure archaeological evidence of human occupation of the region comes from the artefact-rich, high-latitude (~70° N) Yana RHS site dated to ~31.6 kya (…)
The Yana RHS human remains represent the earliest direct evidence of human presence in northeastern Siberia, a population we refer to as “Ancient North Siberians” (ANS). Both Yana RHS individuals were unrelated males, and belong to mitochondrial haplogroup U, predominant among ancient West Eurasian hunter-gatherers, and to Y chromosome haplogroup P1, ancestral to haplogroups Q and R, which are widespread among present-day Eurasians and Native Americans.
Symmetry tests using f4 statistics reject tree-like clade relationships with both Early West Eurasians (EWE; Sunghir) and Early East Asians (EEA; Tianyuan); however, Yana is genetically closer to EWE, despite its geographic location in northeastern Siberia
Using admixture graphs (qpGraph) and outgroup-based estimation of mixture proportions (qpAdm), we find that Yana can be modelled as EWE with ~25% contribution from EEA
Among all ancient individuals, Yana shares the most genetic drift with Mal’ta, and f4 statistics show that Mal’ta shares more alleles with Yana than with EWE (e.g. f4(Mbuti,Mal’ta;Sunghir,Yana) = 0.0019, Z = 3.99). Mal’ta and Yana also exhibit a similar pattern of genetic affinities to both EWE and EEA, consistent with previous studies.The ANE lineage can thus be considered a descendant of the ANS lineage, demonstrating that by 31.6 kya early representatives of this lineage were widespread across northern Eurasia, including far northeastern Siberia.
(…) the 9.8 kya Kolyma1 individual, representing a group we term “Ancient Paleosiberians” (AP). Our results indicate that AP are derived from a first major genetic shift observed in the region. Principal component analysis (PCA), outgroup f3-statistics and mtDNA and Y chromosome haplogroups (G1b and Q1a1a, respectively) demonstrate a close affinity between AP and present-day Koryaks, Itelmen and Chukchis, as well as with Native Americans.
For both AP and Native Americans, ANS ancestry appears more closely related to Mal’ta than Yana, therefore rejecting a direct contribution of Yana to later AP or Native American groups.
Lake Baikal Neolithic – Bronze Age
(…) the newly reported genomes from Ust’Belaya and recently published neighbouring Neolithic and Bronze Age sites show a succession of three distinct genetic ancestries over a ~6 ky time span. The earliest individuals show predominantly East Asian ancestry, closely related to the ancient individuals from DGC. In the early Bronze Age (BA), we observe a resurgence of AP ancestry (up to ~50% ancestry fraction), as well as influence of West Eurasian Steppe ANE ancestry represented by the early BA individuals from Afanasievo in the Altai region (~10%) This is consistent with previous reports of gene flow from an unknown ANE-related source into Lake Baikal hunter-gatherers.
Our results suggest a southward expansion of AP as a possible source, which is also consistent with the replacement of Y chromosome lineages observed at Lake Baikal, from predominantly haplogroup N in the Neolithic to haplogroup Q in the BA. Finally, the most recent individual from Ust’Belaya, dated to ~600 years ago, falls along the Neosiberian cline, similar to the ~760 year-old ‘Young Yana’ individual from northeastern Siberia, demonstrating the widespread distribution of Neosiberian ancestry in the most recent epoch.
At the western edge of northern Eurasia, genetic and strontium isotope data from ancient individuals at the Levänluhta site documents the presence of Saami ancestry in Southern Finland in the Late Holocene 1.5 kya. This ancestry component is currently limited to the northern fringes of the region, mirroring the pattern observed for AP ancestry in northeastern Siberia. However, while the ancient Saami individuals harbour East Asian ancestry, we find that this is better modelled by DGC rather than AP, suggesting that AP influence was likely restricted to the eastern side of the Urals. Comparison of ancient Finns and Saami with their present-day counterparts reveals additional gene flow over the past 1.6 kya, with evidence for West Eurasian admixture into modern Saami. The ancient Finn from Levänluhta shows lower Siberian ancestry than modern Finns .
Finnish_IA (ca. 350 AD) is probably a Saami-speaking individual, just like the newly reported Saami_IA from Levänluhta ca. 400-600 AD (since Fennic peoples were then likely around the Gulf of Finland).
NOTE. The conflicting strontium isotope data on certain samples from the supplementary material hint at possible external origin of some of the previously reported (and possibly the newly reported) Saami Iron Age individuals, possibly hundreds of km. to the southeast of Levänluhta (coast of the Bothnian Sea), which could imply the start of a Finnic push to the north. It is unclear which samples they are referring to, and it is also unclear which interpretation of the archaeological data is more likely, though.
These new samples give us some more information than what we already had, because Finnish_IA implies that there was few East Asian admixture (if any at all) in west Finland during the Roman Iron Age, which pushes still farther forward in time the expected appearance of Siberian ancestry among Saamic (first) and Fennic populations (later). It is unclear whether this East Asian ancestry found in Finnish_IA is actually related to DGC, or it is rather related to the ENA-like ancestry found already in Baltic hunter-gatherers (i.e. in some EHG samples from Karelia), for which Baikal_EN is a good proxy in Lazaridis et al. (2018).
The paper finds thus increased (probably the actual) Siberian ancestry in modern Finns compared to this Iron Age Saami individual. Coupled with the new Saami Iron Age samples (and the previously reported three) from between one to three centuries later – showing the start of Siberian ancestry influx – , we can begin to establish when the expansion of Siberian ancestry happened in central Finland, and thus quite likely when the Saami began to expand to the north and east and admix with Palaeo-Laplandic peoples.
One sample of haplogroup N1a1a1a1a4a1-M1982, Yana_MED, is found in the Arctic region (north-eastern Yakutia) ca. 1100 AD. Since it is derived from N1a1a1a1a-L392, it might be a surprise for some to find it in a clearly non-Uralic speaking environment at the same time other subclades of this haplogroup were admixing in the west with well-established Finno-Saamic, Volga-Finnic, Ugric, and Samoyedic populations…
On the growing doubts that these data – contradicting the CWC=IE theory – are creating among geneticists (from the supplementary materials):
The Proto-Saami language evolved in southern Finland and Karelia in the Early Iron Age, an area now host to Finnish and the closely related Karelian, but with Saami toponyms showing that the latter two languages are intrusive here (Saarikivi 2004). Saami-speaking populations are thought to have retreated to Lapland during the Middle Iron Age (300–800 AD), where it diverged into the modern Saami dialects. Genetically, the northward retreat of the Saami language correlates with the documented decrease of Saami ancestry in Southern Finland between the Iron Age and the modern period (cf. Lamnidis et al. 2018).
On the way to Lapland, the Saami replaced at least two linguistically obscure groups. This can be inferred from 1) an influx of non-Uralic loanwords into Proto-Saami in the Finnish Lakeland area, and 2) an influx of non-Uralic, non-Germanic words into Saami dialects in Lapland (Aikio 2012). Both of these borrowing events imply contact with non-Saami-speaking groups, e.g. non-Uralic-speaking hunter-gatherers that may have left a genetic and linguistic footprint on modern Saami populations.
The linguistic prehistory of Finland thus does not allow for a straightforward interpretation of the genetic data. The detection of East Asian ancestry in the genetically Saami individual is indicative of a population movement from the east (cf. Lamnidis et al. 2018, Rootsi et al. 2007), one that given the affinities with the ~7.6 ky old individuals from the Devil’s Gate Cave may have been a western extension of the Neosiberian turnover. However, it remains unclear whether this gene flow should be associated with the arrival of Uralic speakers, thus providing further support for a Uralic homeland in Eastern Eurasia, or with an earlier immigration of pre-Uralic, so-called “Paleo-Lakelandic” groups.
I think the genetic interpretation is already straightforward, though. Just another sneak peek at how this late admixture with non-Uralians (mainly Palaeo-Lakelandic and Palaeo-Laplandic peoples from Lovozero and related asbestos ware cultures) is going to unfold among expanding Saami-speaking populations after this newly published Finnish_IA sample, i.e. during the late Iron Age, thanks to Lamnidis et al. (2018):
Also, still no trace of R1a in far East Asia (reported as M17 ca. 5300 BC near Lake Baikal by Moussa et al. 2016), so I still have doubts about my previous assessment that R1a split into M17 (and thus also M417) in Siberia, with those expanding hunter-gatherer pottery.
Inhumation and cremation burials were both common in Iron Age Estonia; however, the pattern which burials were prevalent has regional as well temporal peculiarities. In Estonia, cremation burials appear in the Late Bronze Age (1100–500 BC), for example, in stone-cist graves and ship graves, although inhumation is still characteristic of the period [28, 18]. Cremation burials have occasionally been found beneath the Late Bronze Age cists and the Early Iron Age (500 BC–450 AD) tarand graves [30, 28]. In south-eastern Estonia, including Setumaa, the tradition to bury cremated human remains in pit graves also appeared in the Bronze Age and lasted during the Pre-Roman period (500 BC–50 AD) and the Roman Iron Age (50–450 AD), and even up to the medieval times [30, 23, 33, 9]. During the Early Iron Age, cremations appear in cairn graves and have occasionally been found in many Pre-Roman early tarand graves where they appear with inhumations [27, 28, 19, 20, 21, 22, 24]. In Roman Iron Age tarand graves, cremation as well inhumation were practiced [28, 36, 37]; however, cremation was the prevailing burial practice during the Roman Iron Age, for example, in tarand graves in south-eastern Estonia [30, 28]. Roman Iron Age (50 AD–450 AD) burial sites have not been found in continental west Estonia [28, 38]). At the beginning of the Middle Iron Age (450–800 AD), burial sites, for example stone graves without a formal structure, like Maidla I, Lihula and Ehmja ‘Varetemägi’, appear in Läänemaa, west Estonia; in these graves cremations as well inhumations have been found [39, 48]. Like underground cremation burial, the stone grave without a formal structure was the most common grave type during the Late Iron Age (800– 1200 AD) in west Estonia [39, 35, 48]. In south-eastern and eastern Estonia, sand barrows with cremation burials appeared at the beginning of the Middle Iron Age. Cremation barrows are attributed to the Culture of Long Barrows and are most numerous in the villages Laossina and Rõsna in northern Setomaa, on the western shore of Lake Peipsi [8, 48]. Apparently during the Iron Age, the practiced burial customs varied in Estonia.
Three Iron Age cremation graves from south-eastern Estonia and four graves including cremations as well inhumations from western Estonia were analysed by osteological and palaeodemographic methods in order to estimate the age and sex composition of burial sites, and to propose some possible demographic figures and models for living communities.
The crude birth/death rate estimated on the basis of juvenility indices varied between 55.1‰ and 60.0‰ (58.5‰ on average) at Rõsna village in south-eastern Estonia in the Middle Iron Age. The birth/death rates based on juvenility indices for south eastern graves varied to a greater degree. The estimated crude birth/death rate was somewhat lower (38.9‰) at Maidla in the Late Iron Age and extremely high (92.1‰) at Maidla in the Middle Iron Age, which indicates an unsustainable community. High crude birth/death rates are also characteristic of Poanse tarand graves from the Pre-Roman Iron Age – 92.3‰ for the 1st grave and 69.6‰for the 2nd grave. Expectedly, newborn life expectancies are extremely low in both communities – 10.8 years at Poanse I and 14.4 years at Poanse II. Most likely, both Maidla I and Poanse I were unsustainable communities.
According to the main model where the given period of grave usage is 150 years, the burial grounds were most likely exploited by communities of 3–14 people. In most cases, this corresponds to one family or household. In comparison with other graves, Maidla II stone grave in western Estonia and Rõsna-Saare I barrow cemetery in south-eastern Estonia could have been used by a somewhat larger community, which may mean an extended family, a larger household or usage by two nuclear families.
While the demographic situation in the Gulf of Finland during the Iron Age is not well known – and demography is always difficult to estimate based on burials, especially when cremation is prevalent – , there is a clear genetic bottleneck in Finns, which has been estimated precisely to this period, coincident with the expansion of Proto-Fennic.
The infiltration of N1c lineages in Estonia – the homeland of Proto-Fennic – happened during the Iron Age – as of yet found in 3 out of 5 sampled Tarand graves – , while the previous period was dominated by 100% R1a and Corded Ware + Baltic HG ancestry. With the Iron Age, a slight shift towards Corded Ware ancestry can be seen, which probably signals the arrival of warrior-traders from the Alanino culture, close to the steppe. They became integrated through alliances and intermarriages in a culture of chiefdoms dominated by hill forts. Their origin in the Mid-Volga area is witnessed by their material culture, such as Tarand-like graves (read here a full account of events).
This new political structure, reminiscent of the chiefdom system in Sintashta (with a similar fast life history causing a bottleneck of R1a-Z645 lineages), coupled with the expansion of Fennic (and displaced Saamic) peoples to the north, probably caused the spread of N1c-L392 among Finnic peoples. The linguistic influence of these early Iron Age trading movements from the Middle Volga region can be seen in similarities between Fennic and Mordvinic, which proves that the Fenno-Saamic community was by then not only separated linguistically, but also physically (unlike the period of long-term Palaeo-Germanic influence, where loanwords could diffuse from one to the other).
NOTE. Either this, or the alternative version: an increase in Corded Ware ancestry in Estonia during the Iron Age marks the arrival of the first Fennic speakers ca. 800 BC or later, splitting from Mordvinic? A ‘Mordvin-Fennic’ group in the Volga, of mainly Corded Ware ancestry…?? Which comes in turn from a ‘Volga-Saamic’ population of Siberian ancestry in the Artic region??? And, of course, Palaeo-Germanic widely distributed in North-Eastern Europe with R1a during the Bronze Age! Whichever model you find more logical.
Marital structure. The intensity of interethnic marriages puts the existence of the Ulchi population at risk. The colorful ethnic composition of the Ulchi settlements is reflected in the marriage structure [see featured image]. We found that the proportion of single-ethnic marriages of the Ulchi is on average 51%. The greatest number of such marriages takes place in the village of Bulava. Marriages of Ulchi with Russians are in second place. Marriages with indigenous peoples of the Far East, Nanais, Nivkhs, Evenks, and others, are in third place. Thus, almost half of the Ulchi marriages are with representatives of other nationalities. Such a significant level of interethnic mixing makes it possible to talk about intense processes of assimilation of this indigenous people and puts to the forefront the problem of loss of the unique gene pool of the Ulchi.
Haplogroup C (its branch M48) was genotyped for its five subbranches with markers M86, B470, F13686, B93, and the marker at position 16645386 (GRCh37), which was found by our team for the first time. Variant B93 is rare in the Ulchi, and 14 samples (that is, more than a quarter of the entire gene pool of the Ulchi, Fig. 2) belong to M86 and its subvariants. Therefore, we genotyped STR markers of C-M86 carriers for the Ulchi and neighboring Amur populations and analyzed the relationships of detected haplotypes on the phylogenetic network (Fig. 3, STR haplotypes are available from authors upon request).
(…) On the network, different clusters are associated with different populations: most Mongols belong to F13686, all Evenks of the Amur River region with this haplogroup form a subcluster within F13686, and part of Upper Nanais is the basis of cluster B470.
An estimate of the age of the entire haplogroup C-F12355 obtained from the data of genome-wide sequencing of seven specimens is 2400 ± 500 years (O.P. Balanovsky, unpublished data). That is, the common ancestor of all the studied representatives of various peoples with this haplogroup lived not so long ago, the first millennium BC. The formation time of cluster F13686 is somewhat later: 1990 ± 600 years.
(…) obvious traces of the interaction of the gene pool of the Ulchi with neighboring and remote peoples of the Far East and Central Asia in the time range of the last one to three thousand years were revealed. This shows that the results of work  on the similarity of the gene pool of the ancient (age of 7500 years) Neolithic genomes of the Amur River region to the Ulchi probably indicate not the uniqueness of the Ulchi, but the fact that this ancient gene pool was preserved in a vast circle of populations of the Far East interwoven with gene flows both with each other and, to a lesser extent, with populations of Central Asia.
The expansion of C2b1a2a-M86 (among many basal C2-M217 samples) is thus possibly associated with the spread of Tungusic, which puts C2b1a at the root of the Micro-Altaic expansion, with a formation date ca. 12700 BC, TMRCA 12500 BC (and not only Mongolian). This shows that Micro-Altaic is connected with a local population which shows a clear continuity since at least 3500 BC. This, however, tells us little about the origin of the language.
That leaves the ancestral N lineages found among Far East Asians as Palaeo-Siberian in origin, and their late expansions to the west not particularly linked with any of the known Palaeo-Siberian ethnolinguistic groups, let alone a supposed “Uralo-Altaic” language…
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.
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.
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.
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?
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).
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.
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:
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).
A split event, where the ANE component of Yamnaya splits from Sidelkino. This was inferred at about about 11 kya.
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.
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.
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.
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
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.
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:
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.
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?
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.
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:
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.
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.
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.
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.
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:
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.
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…
I read from time to time that “we have not sampled Uralic speakers yet”, and “we are waiting to see when Uralic-speaking peoples are sampled”. Are we, though?
Proto-language homelands are based on linguistic data, such as guesstimates for dialectal evolution, loanwords and phonetic changes for language contacts, toponymy for ancient territories, etc. depending on the available information. The trace is then followed back, using available archaeological data, from the known historic speakers and territory to the appropriate potential prehistoric cultures. Only then can genetic analyses help us clarify the precise prehistoric population movements that better fit the models.
The linguistic homeland
We thought – using linguistic guesstimates and fitting prehistoric cultures and their expansion – that Yamna was the Late Proto-Indo-European culture, so when Yamna was sampled, we had Late Proto-Indo-Europeans sampled. Simple deduction.
We thought that north-eastern Europe was a Uralic-speaking area during the Neolithic:
For those supporting a western continuity (and assuming CWC was Indo-European), the language was present at least since the Comb Ware culture, potentially since the Mesolithic.
For those supporting a late introduction into Finland, Uralic expanded the latest with Abashevo-related movements after its incorporation of Volosovo and related hunter-gatherers.
The expansion to the east must have happened through progressive infiltrations with Seima-Turbino / Andronovo-related expansions.
Finding the linguistic homeland going backwards can be described today as follows:
I. Proto-Fennic homeland
Based on the number of Baltic loanwords, not attested in the more eastern Uralic branches (and reaching only partially Mordvinic), the following can be said about western Finno-Permic languages (Junttila 2014):
The Volga-Kama Basin lies still too far east to be included in a list of possible contact locations. Instead, we could look for the contact area somewhere between Estonia in the west and the surroundings of Moscow in the east, a zone with evidence of Uralic settlement in the north and Baltic on the south side.
The only linguistically well-grounded version of the Stone Age continuation theory was presented by Mikko Korhonen in 1976. Its validity, however, became heavily threatened when Koivulehto 1983a-b proved the existence of a Late Proto-Indo-European or Pre-Baltic loanword layer in Saami, Finnic, and Mordvinic. Since this layer must precede the Baltic one and it was presumably acquired in the Baltic Sea region, Koivulehto posited it on the horizon of the Battle Axe period. This forces a later dating for the Baltic–Finnic contacts.
Today the Battle Axe culture is dated at 3200 to 3000 BC, a period far too remote to correspond linguistically with Proto-Baltic (Kallio 1998a).
Since the Baltic contacts began at a very initial phase of Proto-Finnic, the language must have been relatively uniform at that time. Hence, if we consider that the layer of Baltic loanwords may have spread over the Gulf of Finland at that time, we could also insist that the whole of the Proto-Finnic language did so.
II. Proto-Finno-Saamic homeland
The evidence of continued Palaeo-Germanic loanwords (from Pre- to Proto-Germanic stages) is certainly the most important data to locate the Finno-Saamic homeland, and from there backwards into the true Uralic homeland. Following Kallio (2017):
(…) the loanword evidence furthermore suggests that the ancestors of Finnic and Saamic had at least phonologically remained very close to Proto-Uralic as late as the Bronze Age (ca. 1700–500 BC). In particular, certain loanwords, whose Baltic and Germanic sources point to the first millennium BC, after all go back to the Finno-Saamic proto-stage, which is phonologically almost identical to the Uralic proto-stage (see especially the table in Sammallahti 1998: 198–202). This being the case, Dahl’s wave model could perhaps have some use in Uralic linguistics, too.
The presence of Pre-Germanic loanwords points rather to the centuries around the turn of the 2nd – 1st millennium BC or earlier. Proto-Germanic words must have been borrowed before the end of Germanic influence in the eastern Baltic at the beginning of the Iron Age, which sets a clear terminus ante quem ca. 800 BC.
(…) the earliest Indo-European loanwords in the Uralic languages (…) show that Proto-Uralic cannot have been spoken much earlier than Proto-Indo-European dated about 3500 BC (Koivulehto 2001: 235, 257). As the same loanword evidence naturally also shows that the Uralic and Indo-European homelands were not located far from one another, the Uralic homeland can most likely be located in the Middle and Upper Volga region, right north of the Indo-European homeland*. From the beginning of the Subneolithic period about 5900 BC onwards, this region was an important innovation centre, from where several cultural waves spread to the Finnish Gulf area, such as the Sperrings Ware wave about 4900 BC, the Combed Ware wave about 3900 BC, and the Netted Ware wave about 1900 BC (Carpelan & Parpola 2001: 78–90).
The mainstream position is nowadays trying to hold together the traditional views of Corded Ware as Indo-European, and a Uralic Fennoscandia during the Bronze Age.
The following is an example of how this “Volosovo/Forest Zone hunter-gatherer theory” of Uralic origins looks like, as a ‘mixture’ of cultures and languages that benefits from the lack of genetic data for certain regions and periods (taken from Parpola 2018):
The Corded Ware (or Battle Axe) culture intruded into the Eastern Baltic and coastal Finland already around 3100 BCE. The continuity hypothesis maintains that the early Proto-Finnic speakers of the coastal regions, who had come to Finland in the 4th millennium BCE with the Comb-Pitted Ware, coexisted with the Corded Ware newcomers, gradually adopting their pastoral culture and with it a number of NW-IE loanwords, but assimilating the immigrants linguistically.
The fusion of the Corded Ware and the local Comb-Pitted Ware culture resulted into the formation of the Kiukais culture (c. 2300–1500) of southwestern Finland, which around 2300 received some cultural impulses from Estonia, manifested in the appearance of the Western Textile Ceramic (which is different from the more easterly Textile Ceramic or Netted Ware, and which is first attested in Estonia c. 2700 BCE, cf. Kriiska & Tvauri 2007: 88), and supposed to have been accompanied by an influx of loanwords coming from Proto-Baltic. At the same time, the Kiukais culture is supposed to have spread the custom of burying chiefs in stone cairns to Estonia.
The coming of the Corded Ware people and their assimilation created a cultural and supposedly also a linguistic split in Finland, which the continuity hypothesis has interpreted to mean dividing Proto-Saami-Finnic unity into its two branches. Baltic Finnic, or simply Finnic, would have emerged in the coastal regions of Finland and in the northern East Baltic, while preforms of Saami would have been spoken in the inland parts of Finland.
The Nordic Bronze Age culture, correlated above with early Proto-Germanic, exerted a strong influence upon coastal Finland and Estonia 1600–700 BCE. Due to this, the Kiukais culture was transformed into the culture of Paimio ceramics (c. 1600–700 BCE), later continued by Morby ceramics (c. 700 BCE – 200 CE). The assumption is that clear cultural continuity was accompanied by linguistic continuity. Having assimilated the language of the Germanic traders and relatively few settlers of the Bronze Age, the language of coastal Finland is assumed to have reached the stage of Proto-Finnish at the beginning of the Christian era. In Estonia, the Paimio ceramics have a close counterpart in the contemporaneous Asva ceramics.
I will not comment on Siberian or Central Asian homeland proposals, because they are obviously not mainstream, still less today when we know that Uralic was certainly in contact with Proto-Indo-European, and then with Pre- and Proto-Indo-Iranian, as supported even by the Copenhagen group in Damgaard et al. (2018).
This is what Kallio (2017) has to say about the agendas behind such proposals:
Interestingly, the only Uralicists who generally reject the Central Russian homeland are the Russian ones who prefer the Siberian homeland instead. Some Russians even advocate that the Central Russian homeland is only due to Finnish nationalism or, as one of them put it a bit more tactfully, “the political and ideological situation in Finland in the first decades of the 20th century” (Napolskikh 1995: 4).
Still, some Finns (and especially those who also belong to the “school who wants it large and wants it early”) simultaneously advocate that exactly the same Central Russian homeland is due to Finnlandisierung (Wiik 2001: 466).
Hence, for those of you willing to learn about fringe theories not related to North-Eastern Europe, you also have then the large and early version of the Uralic homeland, with Wiik’s Palaeolithic continuity of Uralic peoples spread over all of eastern and central Europe (hence EHG and R1a included):
These fringe Finnish theories look a lot like the Corded Ware expansion… Better not go the Russian or Finnish nationalist ways? Agreed then, let’s discuss only rational proposals based on current data.
The archaeological homeland
For a detailed account of the Corded Ware expansion with Battle Axe, Fatyanovo-Balanovo, and Abashevo groups into the area, you can read my recent post on the origin of R1a-Z645.
1. Textile ceramics
During the 2nd millennium BC, textile impressions appear in pottery as a feature across a wide region, from the Baltic area through the Volga to the Urals, in communities that evolve from late Corded Ware groups without much external influence.
While it has been held that this style represents a north-west expansion from the Volga region (with the “Netted Ware” expansion), there are actually at least two original textile styles, one (earlier) in the Gulf of Finland, common in the Kiukainen pottery, which evolves into the Textile ware culture proper, and another which seems to have an origin in the Middle Volga region to the south-east.
The Netted ware culture is the one that apparently expands into inner Finland – a region not densely occupied by Corded Ware groups until then. There are, however, no clear boundaries between groups of both styles; textile impressions can be easily copied without much interaction or population movement; and the oldest textile ornamentation appeared on the Gulf of Finland. Hence the tradition of naming all as groups of Textile ceramics.
The fact that different adjacent groups from the Gulf of Finland and Forest Zone share similar patterns making it very difficult to differentiate between ‘Netted Ware’ or ‘Textile Ware’ groups points to:
close cultural connections that are maintained through the Gulf of Finland and the Forest Zone after the evolution of late Corded Ware groups; and
no gross population movements in the original Battle Axe / Fatyanovo regions, except for the expansion of Netted Ware to inner Finland, Karelia, and the east, where the scattered Battle Axe finds and worsening climatic conditions suggest most CWC settlements disappeared at the end of the 3rd millennium BC and recovered only later.
NOTE. This lack of population movement – or at least significant replacement by external, non-CWC groups – is confirmed in genetic investigation by continuity of CWC-related lineages (see below).
The technology present in Textile ceramics is in clear contrast to local traditions of sub-Neolithic Lovozero and Pasvik cultures of asbestos-tempered pottery to the north and east, which point to a different tradition of knowledge and learning network – showing partial continuity with previous asbestos ware, since these territories host the main sources of asbestos. We have to assume that these cultures of northern and eastern Fennoscandia represent Palaeo-European (eventually also Palaeo-Siberian) groups clearly differentiated from the south.
The Chirkovo culture (ca. 1800-700 BC) forms on the middle Volga – at roughly the same time as Netted Ware formed to the west – from the fusion of Abashevo and Balanovo elites on Volosovo territory, and is also related (like Abashevo) to materials of the Seima-Turbino phenomenon.
Bronze Age ethnolinguistic groups
In the Gulf of Finland, Kiukainen evolves into the Paimio ceramics (in Finland) — Asva Ware (in Estonia) culture, which lasts from ca. 1600 to ca. 700 BC, probably representing an evolving Finno-Saamic community, while the Netted Ware from inner Finland (the Sarsa and Tomitsa groups) and the groups from the Forest Zone possibly represent a Volga-Finnic community.
NOTE. Nevertheless, the boundaries between Textile ceramic groups are far from clear, and inner Finland Netted Ware groups seem to follow a history different from Netted Ware groups from the Middle and Upper Volga, hence they could possibly be identified as an evolving Pre-Saamic community.
Based on language contacts, with Early Baltic – Early Finnic contacts starting during the Iron Age (ca. 500 BC onwards), this is a potential picture of the situation at the end of this period, when Germanic influence on the coast starts to fade, and Lusatian culture influence is stronger:
The whole Finno-Permic community remains thus in close contact, allowing for the complicated picture that Kallio mentions as potentially showing Dahl’s wave model for Uralic languages.
Genetic data shows a uniform picture of these communities, with exclusively CWC-derived ancestry and haplogroups. So in Mittnik et al. (2018) all Baltic samples show R1a-Z645 subclades, while the recent session on Estonian populations in ISBA 8 (see programme in PDF) clearly states that:
[Of the 24 Bronze Age samples from stone-cist graves] all 18 Bronze Age males belong to R1a.
Regarding non-Uralic substrates found in Saami, supposedly absorbed during the expansion to the north (and thus representing languages spoken in northern Fennoscandia during the Bronze Age) this is what Aikio (2012) has to say:
The Saami substrate in the Finnish dialects thus reveals that also Lakeland Saami languages had a large number of vocabulary items of obscure origin. Most likely many of these words were substrate in Lakeland Saami, too, and ultimately derive from languages spoken in the region before Saami. In some cases the loan origin of these words is obvious due to their secondary Proto-Saami vowel combinations such as *ā–ë in *kāvë ‘bend; small bay’ and *šāpšë ‘whitefish’. This substrate can be called ‘Palaeo-Lakelandic’, in contrast to the ‘Palaeo-Laplandic’ substrate that is prominent in the lexicon of Lapland Saami. As the Lakeland Saami languages became extinct and only fragments of their lexicon can be reconstructed via elements preserved in Finnish place-names and dialectal vocabulary, we are not in a position to actually study the features of this Palaeo-Lakelandic substrate. Its existence, however, appears evident from the material above.
If we wanted to speculate further, based on the data we have now, it is very likely that two opposing groups will be found in the region:
A) The central Finnish group, in this hypothesis the Palaeo-Lakelandic group, made up of the descendants of the Mesolithic pioneers of the Komsa and Suomusjärvi cultures, and thus mainly Baltic HG / Scandinavian HG ancestry and haplogroups I / R1b(xM269) (see more on Scandinavian HG).
B) Lapland and Kola were probably also inhabited by similar Mesolithic populations, until it was eventually assimilated by expanding Siberian groups (of Siberian ancestry and N1c-L392 lineages) from the east – entering the region likely through the Kola peninsula – , forming the Palaeo-Laplandic group, which was in turn later replaced by expanding Proto-Saamic groups.
Siberian ancestry appears first in Fennoscandia at Bolshoy Oleni Ostrov ca. 1520 BC, with haplogroup N1c-L392 (2 samples, BOO002 and BOO004), and with Siberian ancestry. This is their likely movement in north-eastern Europe, from Lamnidis et al (2018):
The large Siberian component in the Bolshoy individuals from the Kola Peninsula provides the earliest direct genetic evidence for an eastern migration into this region. Such contact is well documented in archaeology, with the introduction of asbestos-mixed Lovozero ceramics during the second millenium BC, and the spread of even-based arrowheads in Lapland from 1,900 BCE. Additionally, the nearest counterparts of Vardøy ceramics, appearing in the area around 1,600-1,300 BCE, can be found on the Taymyr peninsula, much further to the east. Finally, the Imiyakhtakhskaya culture from Yakutia spread to the Kola Peninsula during the same period.
Obviously, these groups of asbestos-tempered ware are not connected to the Uralic expansion. From the same paper:
The fact that the Siberian genetic component is consistently shared among Uralic-speaking populations, with the exceptions of Hungarians and the non-Uralic speaking Russians, would make it tempting to equate this component with the spread of Uralic languages in the area. However, such a model may be overly simplistic. First, the presence of the Siberian component on the Kola Peninsula at ca. 4000 yBP predates most linguistic estimates of the spread of Uralic languages to the area. Second, as shown in our analyses, the admixture patterns found in historic and modern Uralic speakers are complex and in fact inconsistent with a single admixture event. Therefore, even if the Siberian genetic component partly spread alongside Uralic languages, it likely presented only an addition to populations carrying this component from earlier.
2. The Early Iron Age
The Ananino culture appears in the Vyatka-Kama area, famed for its metallurgy, with traditions similar to the North Pontic area, by this time developing Pre-Sauromatian traditions. It expanded to the north in the first half of the first millennium BC, remaining in contact with the steppes, as shown by the ‘Scythian’ nature of its material culture.
NOTE. The Ananino culture can be later followed through its zoomorphic styles into Iron Age Pjanoborskoi and Gljadenovskoi cultures, later to Ural-Siberian Middle Age cultures – Itkuska, Ust’-Poluiska, Kulaiska cultures –, which in turn can be related as prototypes of medieval Permian styles.
At the same time as the Ananino culture begins to expand ca. 1000 BC, the Netted Ware tradition from the middle Oka expanded eastwards into the Oka-Vyatka interfluve of the middle Volga region, until then occupied by the Chirkovo culture. Eventually the Akozino or Akhmylovo group (ca. 800-300 BC) emerged from the area, showing a strong cultural influence from the Ananino culture, by that time already expanding into the Cis-Urals region.
The Akozino culture remains nevertheless linked to the western Forest Zone traditions, with long-ranging influences from as far as the Lusatian culture in Poland (in metallurgical techniques), which at this point is also closely related with cultures from Scandinavia (read more on genetics of the Tollense Valley).
Different materials from Akozino reach Fennoscandia late, at the end of the Bronze Age and beginning of the Early Iron Age, precisely when the influence of the Nordic Bronze Age culture on the Gulf of Finland was declining.
This is a period when Textile ceramic cultures in north-eastern Europe evolve into well-armed chiefdom-based groups, with each chiefdom including thousands or tens of thousands, with the main settlements being hill forts, and those in Fennoscandia starting ca. 1000-400 BC.
Mälar-type celts and Ananino-type celts appear simultaneously in Fennoscandia and the Forest Zone, with higher concentrations in south-eastern Sweden (Mälaren) and the Volga-Kama region, supporting the existence of a revived international trade network.
The Paimio—Asva Ware culture evolves (ca. 700-200 BC) into the Morby (in Finland) — Ilmandu syle (in Estonia, Latvia, and Mälaren) culture. The old Paimio—Asva tradition continues side by side with the new one, showing a clear technical continuity with it, but with ornamentation compared to the Early Iron Age cultures of the Upper Volga area. This new south-eastern influence is seen especially in:
Akozino-Mälar axes (ca. 800-500 BC): introduced into the Baltic area in so great numbers – especially south-western Finland, the Åland islands, and the Mälaren area of eastern Sweden – that it is believed to be accompanied by a movement of warrior-traders of the Akozino-Akhmylovo culture, following the waterways that Vikings used more than a thousand years later. Rather than imports, they represent a copy made with local iron sources.
Tarand graves (ca. 500 BC – AD 400): these ‘mortuary houses’ appear in the coastal areas of northern and western Estonia and the islands, at the same time as similar graves in south-western Finland, eastern Sweden, northern Latvia and Courland. Similar burials are found in Akozino-Akhmylovo, with grave goods also from the upper and middle Volga region, while grave goods show continuity with Textile ware.
The use of asbestos increases in mainland Finnish wares with Kjelmøy Ware (ca. 700 BC – AD 300), which replaced the Lovozero Ware; and in the east in inner Finland and Karelia with the Luukonsaari and Sirnihta wares (ca. 700-500 BC – AD 200), where they replaced the previous Sarsa-Tomitsa ceramics.
The Gorodets culture appears during the Scythian period in the forest-steppe zone north and west of the Volga, shows fortified settlements, and there are documented incursions of Gorodets iron makers into the Samara valley, evidenced by deposits of their typical pottery and a bloom or iron in the region.
Iron Age ethnolinguistic groups
According to (Koryakova and Epimakhov 2007):
It is commonly accepted by archaeology, ethnography, and linguistics that the ancestors of the Permian peoples (the Udmurts, Komi-Permians, and Komi-Zyryans) left the sites of Ananyino cultural intercommunity.
Certain innovations shared between Proto-Fennic (identified with the Gulf of Finland) and Proto-Mordvinic (from the Gorodets culture) point to their close contact before the Proto-Fennic expansion, and thus to the identification of Gorodets as Proto-Mordvinic, hence Akozino as Volgaic (Parpola 2018):
the 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 *kümmen.
The presence of loanwords of non-Uralic origin, related to farming and trees, potentially Palaeo-European in nature (hence possibly from Siberian influence in north-eastern Europe).
The introduction of a strongly hierarchical chiefdom system can quickly change the pre-existing social order and lead to a major genetic shift within generations, without a radical change in languages, as shown in Sintashta-Potapovka compared to the preceding Poltavka society (read more about Sintashta).
Fortified settlements in the region represented in part visiting warrior-traders settled through matrimonial relationships with local chiefs, eager to get access to coveted goods and become members of a distribution network that could guarantee them even military assistance. Such a system is also seen synchronously in other cultures of the region, like the Nordic Bronze Age and Lusatian cultures (Parpola 2013).
The most likely situation is that N1c subclades were incorporated from the Circum-Artic region during the Anonino (Permic) expansion to the north, later emerged during the formation of the Akozino group (Volgaic, under Anonino influence), and these subclades in turn infiltrated among the warrior traders that spread all over Fennoscandia and the eastern Baltic (mainly among Fennic, Saamic, Germanic, and Balto-Slavic peoples), during the age of hill forts, creating alliances partially based on exogamy strategies (Parpola 2013).
Over the course of these events, no language change is necessary in any of the cultures involved, since the centre of gravity is on the expanding culture incorporating new lineages:
first on the Middle Volga, when Ananino expands to the north, incorporatinig N1c lineages from the Circum-Artic region.
then with the expansion of the Akozino-Akhmylovo culture into Ananino territory, admixing with part of its population;
then on the Baltic region, when materials are imported from Akozino into Fennoscandia and the eastern Baltic (and vice versa), with local cultures being infiltrated by foreign (Akozino) warrior-traders and their materials;
and later with the different population movements that led eventually to a greater or lesser relevance of N1c in modern Finno-Permic populations.
To argue that this infiltration and later expansion of lineages changed the language in one culture in one of these events seems unlikely. To use this argument of “opposite movement of ethnic and language change” for different successive events, and only on selected regions and cultures (and not those where the greatest genetic and cultural impact is seen, like e.g. Sweden for Akozino materials) is illogical.
NOTE. Notice how I write here about “infiltration” and “lineages”, not “migration” or “populations”. To understand that, see below the next section on autosomal studies to compare Bronze Age, Iron Age, Medieval and Modern Estonians, and see how little the population of Estonia (homeland of Proto-Fennic and partially of Proto-Finno-Saamic) has changed since the Corded Ware migrations, suggesting genetic continuity and thus mostly close inter-regional and intra-regional contacts in the Forest Zone, hence a very limited impact of the absorbed N1c lineages (originally at some point incorporated from the Circum-Artic region). You can also check on the most recent assessment of R1a vs. N1c in modern Uralic populations.
Iron Age and later populations
From the session on Estonian samples on ISBA 8, by Tambets et al.:
[Of the 13 samples from the Iron Age tarand-graves] We found that the Iron Age individuals do in fact carry chrY hg N3 (…) Furthermore, based on their autosomal data, all of the studied individuals appear closer to hunter-gatherers and modern Estonians than Estonian CWC individuals do.
EDIT (16 OCT) A recent abstract with Saag as main author (Tambets second) cites 3 out of 5 sampled Iron Age individuals as having haplogroup N3.
Looking at the plot, the genetic inflow marking the change from the Bronze Age to the Iron Age looks like an obvious expansion of nearby peoples with CWC-related ancestry, i.e. likely from the south-east, near the Middle Volga, where influence of steppe peoples is greater (hence likely Akozino) into a Proto-Fennic population already admixed (since the arrival of Corded Ware groups) with Comb Ware-like populations.
All of these groups were probably R1a-Z645 (likely R1a-Z283) since the expansion of Corded Ware peoples, with an introduction of some N1c lineages precisely during this Iron Age period. This infiltration of N1c-L392 with Akozino is obviously not directly related to Siberian cultures, given what we know about the autosomal description of Estonian samples.
Rather, N1c-L392 lineages were likely part of the incoming (Volgaic) Akozino warrior-traders, who settled among developing chiefdoms based on hill fort settlements of cultures all over the Baltic area, and began to appear thus in some of the new tarand graves associated with the Iron Age in north-eastern Europe.f
A good way to look at this is to realize that no new cluster appears compared to the data we already have from Baltic LN and BA samples from Mittnik et al. (2018), so the Estonian BA and IA clusters must be located (in a proper PCA) in the cline from Pit-Comb Ware culture through Baltic BA to Corded Ware groups:
This genetic continuity from Corded Ware (the most likely Proto-Uralic homeland) to the Proto-Fennic and Proto-Saamic communities in the Gulf of Finland correlates very well with the known conservatism of Finno-Saamic phonology, quite similar to Finno-Ugric, and both to Proto-Uralic (Kallio 2017): The most isolated region after the expansion of Corded Ware peoples, the Gulf of Finland, shielded against migrations for almost 1,500 years, is then the most conservative – until the arrival of Akozino influence.
Only later would certain regions (like Finland or Lappland) suffer Y-DNA bottlenecks and further admixture events associated with population displacements and expansions, such as the spread of Fennic peoples from their Estonian homeland (evidenced by the earlier separation of South Estonian) to the north and east:
The initial Proto-Fennic expansion was probably coupled with the expansion of Proto-Saami to the north, with the Kjelmøy Ware absorbing the Siberian population of Lovozero Ware, and potentially in inner Finland and Karelia with the Luukonsaari and Sirnihta wares (Carpelan and Parpola 2017).
This Proto-Saami population expansion from the mainland to the north, admixing with Lovozero-related peoples, is clearly reflected in the late Iron Age Saamic samples from Levänluhta (ca. 400-800 AD), as a shift (of 2 out of 3 samples) to Siberian-like ancestry from their original CWC_Baltic-like situation (see PCA from Lamnidis et al. 2018 above).
Also, Volgaic and Permic populations from inner Finland and the Forest Zone to the Cis-Urals and Circum-Artic regions probably incorporate Siberian ancestry and N1c-L392 lineages during these and later population movements, while the westernmost populations – Estonian, Mordvinic – remain less admixed (see PCA from Tambets et al. 2018 below).
We also have data of N1c-L392 in Nordic territory in the Middle Ages, proving its likely strong presence in the Mälaren area since the Iron Age, with the arrival of Akozino warrior traders. Similarly, it is found among Balto-Slavic groups along the eastern Baltic area. Obviously, no language change is seen in Nordic Bronze Age and Lusatian territory, and none is expected in Estonian or Finnish territory, either.
Therefore, no “N1c-L392 + Siberian ancestry” can be seen expanding Finno-Ugric dialects, but rather different infiltrations and population movements with limited effects on ancestry and Y-DNA composition, depending on the specific period and region.
An issue never resolved
Because N1c-L392 subclades & Siberian ancestry, which appear in different proportions and with different origins among some modern Uralic peoples, do not appear in cultures supposed to host Uralic-speaking populations until the Iron Age, people keep looking into any direction to find the ‘true’ homeland of those ‘Uralic N1c peoples’? Kind of a full circular reasoning, anyone? The same is valid for R1a & steppe ancestry being followed for ‘Indo-Europeans’, or R1b-P312 & Neolithic farmer ancestry being traced for ‘Basques’, because of their distribution in modern populations.
I understand the caution of many pointing to the need to wait and see how samples after 2000 BC are like, in every single period, from the middle and upper Volga, Kama, southern Finland, and the Forest Zone between Fennoscandia and the steppe. It’s like waiting to see how people from Western Yamna and the Carpathian Basin after 3000 BC look like, to fill in what is lacking between East Yamna and Bell Beakers, and then between them and every single Late PIE dialect.
But the answer for Yamna-Bell Beaker-Poltavka peoples during the Late PIE expansion is always going to be “R1b-L23, but with R1a-Z645 nearby” (we already have a pretty good idea about that); and the answer for the Forest Zone and northern Cis- and Trans-Urals area – during the time when Uralic languages are known to have already been spoken there – is always going to be “R1a-Z645, but with haplogroup N nearby”, as is already clear from the data on the eastern Baltic region.
So, without a previously proposed model as to where those amateurs expressing concern about ‘not having enough data’ expect to find those ‘Uralic peoples’, all this waiting for the right data looks more like a waiting for N1c and Siberian ancestry to pop up somewhere in the historic Uralic-speaking area, to be able to say “There! A Uralic-speaking male!”. Not a very reasonable framework to deal with prehistoric peoples and their languages, I should think.
But, for those who want to do that, let me break the news to you already:
And here it is, an appropriate fantasy description of the ethnolinguistic groups from the region. You are welcome:
During the Bronze Age, late Corded Ware groups evolve as the western Textile ware Fennic Balto-Slavic group in the Gulf of Finland; the Netted Ware Saamic Balto-Slavic group of inner Finland; the south Netted Ware / Akozino Volgaic Balto-Slavic groups of the Middle Volga; and the Anonino Permic Balto-Slavic group in the north-eastern Forest Zone; all developing still in close contact with each other, allowing for common traits to permeate dialects.
These Balto-Slavic groups would then incorporate west of the Urals during and after the Iron Age (ca. 800-500 BC first, and also later during their expansion to the north) limited ancestry and lineages from eastern European hunter-gatherer groups of Palaeo-European Fennic and Palaeo-Siberian Volgaic and Permic languages from the Circum-Artic region, but they adopted nevertheless the language of the newcomers in every single infiltration of N1c lineages and/or admixture with Siberian ancestry. Oh and don’t forget the Saamic peoples from central Sweden, of course, the famous N1c-L392 ‘Rurikid’ lineages expanding Saamic to the north and replacing Proto-Germanic…
The current model for those obsessed with modern Y-DNA is, therefore, that expanding Neolithic, Bronze Age and Iron Age cultures from north-eastern Europe adopted the languages of certain lineages originally from sub-Neolithic (Scandinavian and Siberian) hunter-gatherer populations of the Circum-Artic region; lineages that these cultures incorporated unevenly during their expansions. Hmmmm… Sounds like an inverse Western movie, where expanding Americans end up speaking Apache, and the eastern coast speaks Spanish until Italian migrants arrive and make everyone speak English… or something. A logic, no-nonsense approach to ethnolinguistic identification.
I kid you not, this is the kind of models we are going to see very soon. In 2018 and 2019, with ancient DNA able to confirm or reject archaeological hypotheses based on linguistic data, people will keep instead creating new pet theories to support preconceived ideas based on the Y-DNA prevalent among modern populations. That is, information available in the 2000s.
So what’s (so much published) ancient DNA useful for, exactly?
[Next post on the subject: Corded Ware—Uralic (III): Seima-Turbino and the Ugric and Samoyedic expansion]
I was reading The Bronze Age Landscape in the Russian Steppes: The Samara Valley Project (2016), and I was really surprised to find the following excerpt by David W. Anthony:
The Samara Valley links the central steppes with the western steppes and is a north-south ecotone between the pastoral steppes to the south and the forest-steppe zone to the north [see figure below]. The economic contrast between pastoral steppe subsistence, with its associated social organizations, and forest-zone hunting and fishing economies probably explains the shifting but persistent linguistic border between forest-zone Uralic languages to the north (today largely displaced by Russian) and a sequence of steppe languages to the south, recently Turkic, before that Iranian, and before that probably an eastern dialect of Proto-Indo-European (Anthony 2007). The Samara Valley represents several kinds of borders, linguistic, cultural, and ecological, and it is centrally located in the Eurasian steppes, making it a critical place to examine the development of Eurasian steppe pastoralism.
Khokhlov (translated by Anthony) further insists on the racial and ethnic divide between both populations, Abashevo to the north, and Poltavka to the south, during the formation of the Abashevo – Sintashta-Potapovka community that gave rise to Proto-Indo-Iranians:
Among all cranial series in the Volga-Ural region, the Potapovka population represents the clearest example of race mixing and probably ethnic mixing as well. The cultural advancements seen in this period might perhaps have been the result of the mixing of heterogeneous groups. Such a craniometric observation is to some extent consistent with the view of some archaeologists that the Sintashta monuments represent a combination of various cultures (principally Abashevo and Poltavka, but with other influences) and therefore do not correspond to the basic concept of an archaeological culture (Kuzmina 2003:76). Under this option, the Potapovka-Sintashta burial rite may be considered, first, a combination of traits to guarantee the afterlife of a selected part of a heterogeneous population. Second, it reflected a kind of social “caste” rather than a single population. In our view, the decisive element in shaping the ethnic structure of the Potapovka-Sintashta monuments was their extensive mobility over a fairly large geographic area. They obtained knowledge of various cultures from the populations with whom they interacted.
Interesting is also this excerpt about the predominant population in the Abashevo – Sintashta-Potapovka admixture (which supports what Chetan said recently, although this does not seemed backed by Y-DNA haplogroups found in the richest burials), coupled with the sign of incoming “Uraloid” peoples from the east, found in both Sintashta and eastern Abashevo:
The socially dominant anthropological component was Europeoid, possibly the descendants of Yamnaya. The association of craniofacial types with archaeological cultures in this period is difficult, primarily because of the small amount of published anthropological material of the cultures of steppe and forest belt (Balanbash, Vol’sko-Lbishche) and the eastern and southern steppes (Botai-Tersek). The crania associated with late MBA western Abashevo groups in the Don-Volga forest zone were different from eastern Abashevo in the Urals, where the expression of the Old Uraloid craniological complex was increased. Old Uraloid is found also on a single skull of Vol’sko-Lbishche culture (Tamar Utkul VII, Kurgan 4). Potentially related variants, including Mongoloid features, could be found among the Seima-Turbino tribes of the forest-steppe zone, who mixed with Sintashta and Abashevo. In the Sintashta Bulanova cemetery from the western Urals, some individuals were buried with implements of Seima-Turbino type (Khalyapin 2001; Khokhlov 2009; Khokhlov and Kitov 2009). Previously, similarities were noted between some individual skulls from Potapovka I and burials of the much older Botai culture in northern Kazakhstan (Khokhlov 2000a). Botai-Tersek is, in fact, a growing contender for the source of some “eastern” cranial features.
The wave of peoples associated with “eastern” features can be seen in genetics in the Sintashta outliers from Narasimhan et al. (2018), and it probably will be eventually seen in Abashevo, too. These may be related to the Seima-Turbino international network – but most likely it is directly connected to Sintashta through the starting Andronovo and Seima-Turbino horizons, by admixing of prospective groups and small-scale back-migrations.
Corded Ware – Yamna similarities?
So, if peoples of north-eastern Europe have been assumed for a long time to be Uralic speakers, what is happening with the Corded Ware = IE obsession? Is it Gimbutas’ ghost possessing old archaeologists? Probably not.
It is about certain cultural similarities evident at first sight, which have been traditionally interpreted as a sign of cultural diffusion or migration. Not dissimilar to the many Bell Beaker models available, where each archaeologist is pushing certain differences, mixing what seemed reasonable, what still might seem reasonable, and what certainly isn’t anymore after the latest ancient DNA data.
The initial models of Gimbutas, Kristiansen, or Anthony – which are known to many today – were enunciated in the infancy of archaeological studies in the regions, during and just after the fall of the USSR, and before many radiocarbon dates that we have today were published (with radiocarbon dating being still today in need of refinement), so it is only logical that gross mistakes were made.
We have similar gross mistakes related to the origins of Bell Beakers, and studying them was certainly easier than studying eastern data.
Gimbutas believed – based mainly on Kurgan-like burials – that Bell Beaker formed from a combination of Yamna settlers with the Vučedol culture, so she was not that far from the truth.
The expansion of Corded Ware from peoples of the North Pontic forest-steppe area, proposed by Gimbutas and later supported also by Kristiansen (1989) as the main Indo-European expansion – , is probably also right about the approximate origins of the culture. Only its ‘Indo-European’ nature is in question, given the differences with Khvalynsk and Yamna evolution.
Anthony only claimed that Yamna migrants settled in the Balkans and along the Danube into the Hungarian steppes. He never said that Corded Ware was a Yamna offshoot until after the first genetic papers of 2015 (read about his newest proposal). He initially claimed that only certain neighbouring Corded Ware groups “adopted” Indo-European (through cultural diffusion) because of ‘patron-client’ relationships, and was never preoccupied with the fate of Corded Ware and related cultures in the east European forest zone and Finland.
So none of them was really that far from the true picture; we might say a lot people are more way off the real picture today than the picture these three researchers helped create in the 1990s and 2000s. Genetics is just putting the last nail in the coffin of Corded Ware as a Yamna offshoot, instead of – as we believed in the 2000s – to Vučedol and Bell Beaker.
So let’s revise some of these traditional links between Corded Ware and Yamna with today’s data:
Even more than genetics – at least until we have an adequate regional and temporary sampling – , archaeological findings lead what we have to know about both cultures.
It is essential to remember that Corded Ware, starting ca. 3000/2900 BC in east-central Europe, has been proposed to be derived from Early Yamna, which appeared suddenly in the Pontic-Caspian steppes ca. 3300 BC (probably from the late Repin expansion), and expanded to the west ca. 3000.
The question at hand, therefore, is if Corded Ware can be considered an offshoot of the Late PIE community, and thus whether the CWC ethnolinguistic community – proven in genetics to be quite homogeneous – spoke a Late PIE dialect, or if – alternatively – it is derived from other neighbouring cultures of the North Pontic region.
NOTE. The interpretation of an Indo-Slavonic group represented by a previous branching off of the group is untenable with today’s data, since Indo-Slavonic – for those who support it – would itself be a branch of Graeco-Aryan, and Palaeo-Balkan languages expanded most likely with West Yamna (i.e. R1b-L23, mainly R1b-Z2103) to the south.
The convoluted alternative explanation would be that Corded Ware represents an earlier, Middle PIE branch (somehow carrying R1a??) which influences expanding Late PIE dialects; this has been recently supported by Kortlandt, although this simplistic picture also fails to explain the Uralic problem.
❔ Kurgans: The Yamna tradition was inherited from late Repin, in turn inherited from Khvalynsk-Novodanilovka proto-Kurgans. As for the CWC tradition, it is unclear if the tumuli were built as a tradition inherited from North and West Pontic cultures (in turn inherited or copied from Khvalynsk-Novodanilovka), such as late Trypillia, late Kvityana, late Dereivka, late Sredni Stog; or if they were built because of the spread of the ‘Transformation of Europe’, set in motion by the Early Yamna expansion ca. 3300-3000 BC (as found in east-central European cultures like Coţofeni, Lizevile, Șoimuș, or the Adriatic Vučedol). My guess is that it inherits an older tradition than Yamna, with an origin in east-central Europe, because of the mound-building distribution in the North Pontic area before the Yamna expansion, but we may never really know.
❌ Burial rite: Yamna features (with regional differences) single burials with body on its back, flexed upright knees, poor grave goods, common orientation east-west (heads to the west) inherited from Repin, in turn inherited from Khvalynsk-Novodanilovka. CWC tradition – partially connected to Złota and surrounding east-central European territories (in turn from the Khvalynsk-Novodanilovka expansion) – features single graves, body in fetal position, strict gender differentiation – men on the right, women on the left -, looking to the south, graves with standardized assemblages (objects representing affirmation of battle, hunting, and feasting). The burial rites clearly represent different ideologies.
❌ Corded decoration: Corded ware decoration appears in the Balkans during the 5th millennium, and represents a simple technique whereby a cord is twisted, or wrapped around a stick, and then pressed directly onto the fresh surface of a vessel leaving a characteristic decoration. It appears in many groups of the 5th and 4th millennium BC, but it was Globular Amphorae the culture which popularized the drinking vessels and their corded ornamentation. It appears thus in some regional groups of Yamna, but it becomes the standard pottery only in Corded Ware (especially with the A-horizon), which shows continuity with GAC pottery.
❌ Economy: Yamna expands from Repin (and Repin from Khvalynsk-Novodanilovka) as a nomadic or semi-nomadic purely pastoralist society (with occasional gathering of wild seeds), which naturally thrives in the grasslands of the Pontic-Caspian, lower Danube and Hungarian steppes. Corded Ware shows agropastoralism (as late Eneolithic forest-steppe and steppe groups of eastern Europe, such as late Trypillian, TRB, and GAC groups), inhabits territories north of the loess line, with heavy reliance of hunter-gathering depending on the specific region.
❌ Cattle herding: Interestingly, both west Yamna and Corded Ware show more reliance on cattle herding than other pastoralist groups, which – contrasted with the previous Eneolithic herding traditions of the Pontic-Caspian steppe, where sheep-goats predominate – make them look alike. However, the cattle-herding economy of Yamna is essential for its development from late Repin and its expansion through the steppes (over western territories practising more hunter-gathering and sheep-goat herding economy), and it does not reach equally the Volga-Ural region, whose groups keep some of the old subsistence economy (read more about the late Repin expansion). Corded Ware, on the other hand, inherits its economic strategy from east European groups like TRB, GAC, and especially late Trypillian communities, showing a predominance of cattle herding within an agropastoral community in the forest-steppe and forest zones of Volhynia, Podolia, and surrounding forest-steppe and forest regions.
❔ Horse riding: Horse riding and horse transport is proven in Yamna (and succeeding Bell Beaker and Sintashta), assumed for late Repin (essential for cattle herding in the seas of grasslands that are the steppes, without nearby water sources), quite likely during the Khvalynsk expansion (read more here), and potentially also for Samara, where the predominant horse symbolism of early Khvalynsk starts. Corded Ware – like the north Pontic forest-steppe and forest areas during the Eneolithic – , on the other hand, does not show a strong reliance on horse riding. The high mobility and short-term settlements characteristic of Corded Ware, that are often associated with horse riding by association with Yamna, may or may not be correct, but there is no need for horses to explain their herding economy or their mobility, and the north-eastern European areas – the one which survived after Bell Beaker expansion – did certainly not rely on horses as an essential part of their economy.
NOTE: I cannot think of more supposed similarities right now. If you have more ideas, please share in the comments and I will add them here.
✅ EHG: This is the clearest link between both communities. We thought it was related to the expansion of ANE-related ancestry to the west into WHG territory, but now it seems that it will be rather WHG expanding into ANE territory from the Pontic-Caspian region to the east (read more on recent Caucasus Neolithic, on , and on Caucasus HG).
NOTE. Given how much each paper changes what we know about the Palaeolithic, the origin and expansion of the (always developing) known ancestral components and specific subclades (see below) is not clear at all.
❔ CHG: This is the key link between both cultures, which will delimit their interaction in terms of time and space. CHG is intermediate between EHG and Iran N (ca. 8000 BC). The ancestry is thus linked to the Caucasus south of the steppe before the emergence of North Pontic (western) and Don-Volga-Ural (eastern) communities during the Mesolithic. The real question is: when we have more samples from the steppe and the Caucasus during the Neolithic, how many CHG groups are we going to find? Will the new specific ancestral components (say CHG1, CHG2, CHG3, etc.) found in Yamna (from Khvalynsk, in the east) and Corded Ware (probably from the North Pontic forest-steppe) be the same? My guess is, most likely not, unless they are mediated by the Khvalynsk-Novodanilovka expansion (read more on CHG in the Caucasus).
❌ WHG/EEF: This is the obvious major difference – known today – in the formation of both communities in the steppe, and shows the different contacts that both groups had at least since the Eneolithic, i.e. since the expansion of Repin with its renewed Y-DNA bottleneck, and probably since before the early Khvalynsk expansion (read more on Yamna-Corded Ware differences contrasting with Yamna-Afanasevo, Yamna-Bell Beaker, and Yamna-Sintashta similarities).
NOTE 1. Some similarities between groups can be seen depending on the sampled region; e.g. Baltic groups show more similarities with southern Pontic-Caspian steppe populations, probably due to exogamy.
NOTE 2. We have this information on the differences in “steppe ancestry” between Yamna and Corded Ware, compared to previous studies, because now we have more samples of neighbouring, roughly contemporaneous Eneolithic groups, to analyse the real admixture processes. This kind of fine scale studies is what is going to show more and more differences between Khvalynsk-Yamna and Sredni Stog-Corded Ware as more data pours in. The evolution of both communities in archaeology and in PCA (see below) is probably witness to those differences yet to be published.
❌ R1: Even though some people try very hard to think in terms of “R1” vs. (Caucasus) J or G or any other upper clade, this is plainly wrong. It is possible, given what we know now, that Q1a2-M242 expanded ANE ancestry to the west ca. 13000 BC, while R1b-P279 expanded WHG ancestry to the east with the expansion of post-Swiderian cultures, creating EHG as a WHG:ANE cline. The role of R1a-M459 is unknown, but it might be related to any of these migrations, or others (plural) along northern Eurasia (read more on the expansion of R1b-P279, on Palaeolithic Q1a2, and on R1a-M417).
NOTE. I am inclined to believe in a speculative Mesolithic-Early Neolithic community involving Eurasiatic movements accross North Eurasia, and Indo-Uralic movements in its western part, with the last intense early Uralic-PIE contacts represented by the forming west (Mariupol culture) and east (Don-Volga-Ural cultures, including Samara) communities developing side by side. Before their known Eneolithic expansions, no large-scale Y-DNA bottleneck is going to be seen in the Pontic-Caspian steppe, with different (especially R1a and R1b subclades) mixed among them, as shown in North Pontic Neolithic, Samara HG, and Khvalynsk samples.
Corded Ware and ‘steppe ancestry’
If we take a look at the evolution of Corded Ware cultures, the expansion of Bell Beakers – dominated over most previous European cultures from west to east Europe – influenced the development of the whole European Bronze Age, up to Mierzanowice and Trzciniec in the east.
The only relevant unscathed CWC-derived groups, after the expansion of Sintashta-Potapovka as the Srubna-Andronovo horizon in the Eurasian steppes, were those of the north-eastern European forest zone: between Belarus to the west, Finland to the north, the Urals to the east, and the forest-steppe region to the south. That is, precisely the region supposed to represent Uralic speakers during the Bronze Age.
This inconsistency of steppe ancestry and its relation with Uralic (and Balto-Slavic) peoples was observed shortly after the publication of the first famous 2015 papers by Paul Heggarty, of the Max-Planck Institute for Evolutionary Anthropology (read more):
Haak et al. (2015) make much of the high Yamnaya ancestry scores for (only some!) Indo-European languages. What they do not mention is that those same results also include speakers of other languages among those with the highest of all scores for Yamnaya ancestry. Only these are languages of the Uralic family, not Indo-European at all; and their Yamnaya-ancestry signals are far higher than in many branches of Indo-European in (southern) Europe. Estonian ranks very high, while speakers of the very closely related Finnish are curiously not shown, and nor are the Saami. Hungarian is relevant less directly since this language arrived only c. 900 AD, but also high.
These data imply that Uralic-speakers too would have been part of the Yamnaya > Corded Ware movement, which was thus not exclusively Indo-European in any case. And as well as the genetics, the geography, chronology and language contact evidence also all fit with a Yamnaya > Corded Ware movement including Uralic as well as Balto-Slavic.
Both papers fail to address properly the question of the Uralic languages. And this despite — or because? — the only Uralic speakers they report rank so high among modern populations with Yamnaya ancestry. Their linguistic ancestors also have a good claim to have been involved in the Corded Ware and Yamnaya cultures, and of course the other members of the Uralic family are scattered across European Russia up to the Urals.
NOTE. Although the author was trying to support the Anatolian hypothesis – proper of glottochronological studies often published from the Max Planck Institute – , the question remains equally valid: “if Proto-Indo-European expands with Corded Ware and steppe ancestry, what is happening with Uralic peoples?”
For my part, I claimed in my draft that ancestral components were not the only relevant data to take into account, and that Y-DNA haplogroups R1a and R1b (appearing separately in CWC and Yamna-Bell Beaker-Afanasevo), together with their calculated timeframes of formation – and therefore likely expansion – did not fit with the archaeological and linguistic description of the spread of Proto-Indo-European and its dialects.
In fact, it seemed that only one haplogroup (R1b-M269) was constantly and consistenly associated with the proposed routes of Late PIE dialectal expansions – like Anthony’s second (Afanasevo) and third (Lower Danube, Balkan) waves. What genetics shows fits seamlessly with Mallory’s association of the North-West Indo-European expansion with Bell Beakers (read here how archaeologists were right).
More precise inconsistencies were observed after the publication of Olalde et al. (2017) and Mathieson et al. (2017), by Volker Heyd in Kossinna’s smile (2017). Letting aside the many details enumerated (you can read a summary in my latest draft), this interesting excerpt is from the conclusion:
Simple solutions to complex problems are never the best choice, even when favoured by politicians and the media. Kossinna also offered a simple solution to a complex prehistoric problem, and failed therein. Prehistoric archaeology has been aware of this for a century, and has responded by becoming more differentiated and nuanced, working anthropologically, scientifically and across disciplines (cf. Müller 2013; Kristiansen 2014), and rejecting monocausal explanations. The two aDNA papers in Nature, powerful and promising as they are for our future understanding, also offer rather straightforward messages, heavily pulled by culture-history and the equation of people with culture. This admittedly is due partly to the restrictions of the medium that conveys them (and despite the often relevant additional detail given as supplementary information, which is unfortunately not always given full consideration).
While I have no doubt that both papers are essentially right, they do not reflect the complexity of the past. It is here that archaeology and archaeologists contributing to aDNA studies find their role; rather than simply handing over samples and advising on chronology, and instead of letting the geneticists determine the agenda and set the messages, we should teach them about complexity in past human actions and interactions. If accepted, this could be the beginning of a marriage made in heaven, with the blessing smile of Gustaf Kossinna, and no doubt Vere Gordon Childe, were they still alive, in a reconciliation of twentieth- and twenty-first-century approaches. For us as archaeologists, it could also be the starting point for the next level of a new archaeology.
The question was made painfully clear with the publication of Olalde et al. (2018) & Mathieson et al. (2018), where the real route of Yamna expansion into Europe was now clearly set through the steppes into the Carpathian basin, later expanded as Bell Beakers.
Studies analysing mtDNA and Y chromosome markers have revealed a sex-specific admixture pattern of admixture of Southeast and South Asian ancestry components for Munda speakers. While close to 100% of mtDNA lineages present in Mundas match those in other Indian populations, around 65% of their paternal genetic heritage is more closely related to Southeast Asian than South Asian variation. Such a contrasting distribution of maternal and paternal lineages among the Munda speakers is a classic example of ‘father tongue hypothesis’. However, the temporality of this expansion is contentious. Based on Y-STR data the coalescent time of Indian O2a-M95 haplogroup was estimated to be >10 KYA. Recently, the reconstructed phylogeny of 8.8 Mb region of Y chromosome data showed that Indian O2a-M95 lineages coalesce within a clade nested within East/Southeast Asian within the last ~5-7 KYA. This date estimate sets the upper boundary for the main episode of gene flow of Y chromosomes from Southeast Asia to India.
Admixture proportions suggest a novel scenario
Regardless of which West Asian population we used, we found that Munda speakers can be described on average as a mixture of ~19% Southeast Asian, 15% West Asian and 66% Onge (South Asian) components. Alternatively, the West and South Asian components of Munda could be modelled using a single South Asian population (Paniya), accounting on average to 77% of the Munda genome. When rescaling the West and South Asian (Onge) components to 1 to explore the Munda genetic composition prior to the introduction of the Southeast Asian component, we note that the West Asian component is lower (~19%) in Munda compared to Paniya (27%) (Supplementary Table S4: *Average_Lao=0). Consistently with qpGraph analyses in Narasimhan et al. (2018), this may point to an initial admixture of a Southeast Asian substrate with a South Asian substrate free of any West Asian component, followed by the encounter of the resulting admixed population with a Paniya-like population. Such a scenario would imply an inverse relationship between the Southeast and West Asian relative proportions in Munda or, in other words, the increase of Southeast Asian component should cause a greater reduction of the West Asian compared to the reduction in the South Asian component in Munda.
Dating the admixture event
In this study, we have replicated a result previously reported in Chaubey et al. (2011)7 that the Mundas lack one ancestral component (k2) that is characteristic to Indian Indo-European and Dravidian speaking populations. If this component came to India through one of the Indo-Aryan migrations then it would be fair to presume that the Munda admixture happened before this component reached India or at least before it spread all over the country. However, the admixture time computed here, falls in the exact same timeframe as the ANI-ASI mixture has been estimated to have happened in India through which the k2 component probably spread. Therefore, we propose that if the Munda admixture happened at the same time, it is possible for it to have happened in the eastern part of the country, east of Bangladesh, and later when populations from East Asia moved to the area, the Mundas migrated towards central India. Such a scenario, which may be further clarified by ancient DNA analyses, seems to be further supported by the fact that Mundas harbor a smaller fraction of West Asian ancestry compared to contemporary Paniya (Supplementary Table S4) and cannot therefore be seen as a simple admixture product of Southern Indian populations with incoming Southeast Asian ancestries.
Linguistics and genome-wide data
(…) by and large, the linguistic classification justifies itself but Kharia and Juang do not fit in this simplification perfectly.
Once again, with the current level of detail in genetic studies, there is often no clear dialectal division possible for certain groups without fine-scale population studies, and the help from linguistics and archaeology.
A total of 286 samples of Uralic-speaking individuals, of those 121 genotyped in this study, were analysed in the context of 1514 Eurasian samples (including 14 samples published for the first time) based on whole genome single nucleotide polymorphisms (SNPs) (Additional file 1: Table S1). All these samples, together with the larger sample set of Uralic speakers, were characterized for mtDNA and chrY markers.
The question as which material cultures may have co-spread together with proto-Uralic and Uralic languages depends on the time estimates of the splits in the Uralic language tree. Deeper age estimates (6,000 BP) of the Uralic language tree suggest a connection between the spread of FU languages from the Volga River basin towards the Baltic Sea either with the expansion of the Neolithic culture of Combed Ware, e.g. [6, 7, 17, 26] or with the Neolithic Volosovo culture . Younger age estimates support a link between the westward dispersion of Proto-Finno-Saamic and eastward dispersion of Proto-Samoyedic with a BA Sejma-Turbino (ST) cultural complex [14, 18, 27, 28] that mediated the diffusion of specific metal tools and weapons from the Altai Mountains over the Urals to Northern Europe or with the Netted Ware culture , which succeeded Volosovo culture in the west. It has been suggested that Proto-Uralic may have even served as the lingua franca of the merchants involved in the ST phenomenon . All these scenarios imply that material culture of the Baltic Sea area in Europe was influenced by cultures spreading westward from the periphery of Europe and/or Siberia. Whether these dispersals involved the spread of both languages and people remains so far largely unknown.
The population structure of Uralic speakers
To contextualize the autosomal genetic diversity of Uralic speakers among other Eurasian populations (Additional file 1: Table S1), we first ran the principal component (PC) analysis (Fig. 2a, Additional file 3: Figure S1). The first two PCs (Fig. 2a, Additional file 3: Figure S1A) sketch the geography of the Eurasian populations along the East-West and North-South axes, respectively. The Uralic speakers, along with other populations speaking Slavic and Turkic languages, are scattered along the first PC axis in agreement with their geographic distribution (Figs. 1 and 2a) suggesting that geography is the main predictor of genetic affinity among the groups in the given area. Secondly, in support of this, we find that FST-distances between populations (Additional file 3: Figure S2) decay in correlation with geographical distance (Pearson’s r = 0.77, p < 0.0001). On the UPGMA tree based on these FST-distances (Fig. 2b), the Uralic speakers cluster into several different groups close to their geographic neighbours.
We next used ADMIXTURE , which presents the individuals as composed of inferred genetic components in proportions that maximize Hardy-Weinberg and linkage equilibrium in the overall sample (see the ‘Methods’ section for choice of presented K). Overall, and specifically at lower values of K, the genetic makeup of Uralic speakers resembles that of their geographic neighbours. The Saami and (a subset of) the Mansi serve as exceptions to that pattern being more similar to geographically more distant populations (Fig. 3a, Additional file 3: S3). However, starting from K = 9, ADMIXTURE identifies a genetic component (k9, magenta in Fig. 3a, Additional file 3: S3), which is predominantly, although not exclusively, found in Uralic speakers. This component is also well visible on K = 10, which has the best cross-validation index among all tests (Additional file 3: S3B). The spatial distribution of this component (Fig. 3b) shows a frequency peak among Ob-Ugric and Samoyed speakers as well as among neighbouring Kets (Fig. 3a). The proportion of k9 decreases rapidly from West Siberia towards east, south and west, constituting on average 40% of the genetic ancestry of FU speakers in Volga-Ural region (VUR) and 20% in their Turkic-speaking neighbours (Bashkirs, Tatars, Chuvashes; Fig. 3a). The proportion of this component among the Saami in Northern Scandinavia is again similar to that of the VUR FU speakers, which is exceptional in the geographic context. It is also notable that North Russians, sampled from near the White Sea, differ from other Russians by sporting higher proportions of k9 (10–15%), which is similar to the values we observe in their Finnic-speaking neighbours. Notably, Estonians and Hungarians, who are geographically the westernmost Uralic speakers, virtually lack the k9 cluster membership.
We also tested the different demographic histories of female and male lineages by comparing outgroup f3 results for autosomal and X chromosome (chrX) data for pairs of populations (Estonians, Udmurts or Khanty vs others) with high versus low probability to share their patrilineal ancestry in chrY hg N (see the ‘Methods’ section, Additional file 3: Figure S13). We found a minor but significant excess of autosomal affinity relative to chrX for pairs of populations that showed a higher than 10% chance of two randomly sampled males across the two groups sharing their chrY ancestry in hg N3-M178, compared to pairs of populations where such probability is lower than 5% (Additional file 3: Figure S13).
In sum, these results suggest that most of the Uralic speakers may indeed share some level of genetic continuity via k9, which, however, also extends to the geographically close Turkic speakers.
We found that it is the admixture with the Siberians that makes the Western Uralic speakers different from the tested European populations (Additional file 3: Figure S4A-F, H, J, L). Differentiating between Estonians and Finns, the Siberians share more derived alleles with Finns, while the geographic neighbours of Estonians (and Finns) share more alleles with Estonians (Additional file 3: Figure S4M). Importantly, Estonians do not share more derived alleles with other Finnic, Saami, VUR FU or Ob-Ugric-speaking populations than Latvians (Additional file 3: Figure S4O). The difference between Estonians and Latvians is instead manifested through significantly higher levels of shared drift between Estonians and Siberians on the one hand and Latvians and their immediate geographic neighbours on the other hand. None of the Uralic speakers, including linguistically close Khanty and Mansi, show significantly closer affinities to the Hungarians than any non-FU population from NE Europe (Additional file 3: Figure S4R).
Time of Siberian admixture
The time depth of the Globetrotter (Fig. 5b) inferred admixture events is relatively recent—500–1900 AD (see also complementary ALDER results, in Additional file 13: Table S12 and Additional file 3: Figure S7)—and agrees broadly with the results reported in Busby et al. . A more detailed examination of the ALDER dates, however, reveals an interesting pattern. The admixture events detected in the Baltic Sea region and VUR Uralic speakers are the oldest (800–900 AD or older) followed by those in VUR Turkic speakers (∼1200–1300 AD), while the admixture dates for most of the Siberian populations (>1500 AD) are the most recent (Additional file 3: Figure S7). The West Eurasian influx into West Siberia seen in modern genomes was thus very recent, while the East Eurasian influx into NE Europe seems to have taken place within the first millennium AD (Fig. 5b, Additional file 3: Figure S7).
Affinities of the Uralic speakers with ancient Eurasians
We next calculated outgroup f3-statistics  to estimate the extent of shared genetic drift between modern and ancient Eurasians (Additional file 14: Table S13, Additional file 3: Figures S8-S9). Consistent with previous reports [45, 50], we find that the NE European populations including the Uralic speakers share more drift with any European Mesolithic hunter-gatherer group than Central or Western Europeans (Additional file 3: Figure S9A-C). Contrasting the genetic contribution of western hunter-gatherers (WHG) and eastern hunter-gatherers (EHG), we find that VUR Uralic speakers and the Saami share more drift with EHG. Conversely, WHG shares more drift with the Finnic and West European populations (Additional file 3: Figure S9A). Interestingly, we see a similar pattern of excess of shared drift between VUR and EHG if we substitute WHG with the aDNA sample from the Yamnaya culture (Additional file 3: Figure S9D). As reported before [2, 45], the genetic contribution of European early farmers decreases along an axis from Southern Europe towards the Ural Mountains (Fig. 6, Additional file 3: Figure S9E-F).
We then used the qpGraph software  to test alternative demographic scenarios by trying to fit the genetic diversity observed in a range of the extant Finno-Ugric populations through a model involving the four basic European ancestral components: WHG, EHG, early farmers (LBK), steppe people of Yamnaya/Corded Ware culture (CWC) and a Siberian component (Fig. 6, Additional file 3: Figure S10). We chose the modern Nganasans to serve as a proxy for the latter component because we see least evidence for Western Eurasian admixture (Additional file 3: Figure S3) among them. We also tested the Khantys for that proxy but the model did not fit (yielding f2-statistics, Z-score > 3). The only Uralic-speaking population that did not fit into the tested model with five ancestral components were Hungarians. The qpGraph estimates of the contributions from the Siberian component show that it is the main ancestry component in the West Siberian Uralic speakers and constitutes up to one third of the genomes of modern VUR and the Saami (Fig. 6). It drops, however, to less than 10% in most of NE Europe, to 5% in Estonians and close to zero in Latvians and Lithuanians.
One of the notable observations that stands out in the fineSTRUCTURE analysis is that neither Hungarians nor Estonians or Mordovians form genetic clusters with other Uralic speakers but instead do so with a broad spectrum of geographically adjacent samples. Despite the documented history of the migration of Magyars  and their linguistic affinity to Khantys and Mansis, who today live east of the Ural Mountains, there is nothing in the present-day gene pool of the sampled Hungarians that we could tie specifically to other Uralic speakers.
Perhaps even more surprisingly, we found that Estonians, who show close affinities in IBD analysis to neighbouring Finnic speakers and Saami, do not share an excess of IBD segments with the VUR or Siberian Uralic speakers. This is eIn this context, it is important to remind that the limited (5%, Fig. 6) East Eurasian impact in the autosomal gene pool of modern Estonians contrasts with the fact that more than 30% of Estonian (but not Hungarian) men carry chrY N3 that has an East Eurasian origin and is very frequent among NE European Uralic speakers . However, the spread of chrY hg N3 is not language group specific as it shows similar frequencies in Baltic-speaking Latvians and Lithuanians, and in North Russians, who in all our analyses are very similar to Finnic-speakers. The latter, however, are believed to have either significantly admixed with their Uralic-speaking neighbours or have undergone a language shift from Uralic to Indo-European .ven more striking considering that the immediate neighbours—Finns, Vepsians and Karelians—do.
With some exceptions such as Estonians, Hungarians and Mordovians, both IBD sharing and Globetrotter results suggest that there are detectable inter-regional haplotype sharing ties between Uralic speakers from West Siberia and VUR, and between NE European Uralic speakers and VUR. In other words, there is a fragmented pattern of haplotype sharing between populations but no unifying signal of sharing that unite all the studied Uralic speakers.
The paper is obviously trying to find a “N1c/Siberian ancestry = Uralic” link, but it shows (as previous papers using ancient DNA) that this identification is impossible, because it is not possible to identify “N1c=Siberian ancestry”, “N1c=Uralic”, or “Siberian ancestry = Uralic”. In fact, the arrival of N subclades and Siberian ancestry are late, both events (probably multiple stepped events) are unrelated to each other, and represent east-west demic diffusion waves (as well as founder effects) that probably coincide in part with the Scythian and Turkic (or associated) expansions, i.e. too late for any model of Proto-Uralic or Proto-Finno-Ugric expansion.
On the other hand, it shows interesting data regarding ancestry of populations that show increased Siberian influence, such as those easternmost groups admixed with Yeniseian-like populations (Samoyedic), those showing strong founder effects (Finnic), or those isolated in the Circum-Artic region with neighbouring Siberian peoples in Kola (Saami). All in all, Hungarians, Estonians and Mordovians seem to show the original situation better than the other groups, which is also reflected in part in Y-DNA, conserved as a majority of R1a lineages precisely in these groups. Just another reminder that CWC-related ancestry is found in every single Uralic group, and that it represents the main ancestral component in all non-Samoyedic groups.
The qpGraph shows the ancestor of Yamna (likely Khvalynsk) and Corded Ware stemming as different populations from a common (likely Neolithic) node – whose difference is based on the proportion of Anatolian-related ancestry – , that is, probably before the Indo-Hittite expansion; and ends with CWC groups forming the base for all Uralic peoples. Below is a detail of the qpGraph on the left, and my old guess (2017) on the right, for comparison:
#EDIT (22 sep 2018): I enjoyed re-reading it, and found this particular paragraph funny:
Despite the documented history of the migration of Magyars  and their linguistic affinity to Khantys and Mansis, who today live east of the Ural Mountains, there is nothing in the present-day gene pool of the sampled Hungarians that we could tie specifically to other Uralic speakers.