Let me begin this final post on the Corded Ware—Uralic connection with an assertion that should be obvious to everyone involved in ethnolinguistic identification of prehistoric populations but, for one reason or another, is usually forgotten. In the words of David Reich, in Who We Are and How We Got Here (2018):
Human history is full of dead ends, and we should not expect the people who lived in any one place in the past to be the direct ancestors of those who live there today.
Another recurrent argument – apart from “Siberian ancestry” – for the location of the Uralic homeland is “haplogroup N”. This is as serious as saying “haplogroup R1” to refer to Indo-European migrations, but let’s explore this possibility anyway:
We have now a better idea of how many ancient migrations (previously hypothesized to be associated with westward Uralic migrations) look like in genetic terms. From Damgaard et al. (Science 2018):
These serial changes in the Baikal populations are reflected in Y-chromosome lineages (Fig. SA; figs. S24 to S27, and tables S13 and SI4). MAI carries the R haplogroup, whereas the majority of Baikal_EN males belong to N lineages, which were widely distributed across Northern Eurasia (29), and the Baikal_LNBA males all carry Q haplogroups, as do most of the Okunevo_EMBA as well as some present-day Central Asians and Siberians.
The only N1c1 sample comes from Ust’Ida Late Neolithic, 180km to the north of Lake Baikal, which – together with the Bronze Age sample from the Kola peninsula, and the medieval sample from Ust’Ida – gives a good idea of the overall expansion of N subclades and Siberian ancestry among the Circum-Arctic peoples of Eurasia, speakers of Palaeo-Siberian languages.
What we should expect from Uralic peoples expanding with haplogroup N – seeing how Yamna expands with R1b-L23, and Corded Ware expands with R1a-Z645 – is to find a common subclade spreading with Uralic populations. Let’s see if it works like that for any N-X subclade, in data from Ilumäe et al. (2016):
Within the Eurasian circum-Arctic spread zone, N3 and N2a reveal a well-structured spread pattern where individual sub-clades show very different distributions:
N1a1-M46 (or N-TAT), formed ca. 13900 BC, TMRCA 9800 BC
N1a1a2-B187, formed ca. 9800 BC, TMRCA 1050 AD:
The sub-clade N3b-B187 is specific to southern Siberia and Mongolia, whereas N3a-L708 is spread widely in other regions of northern Eurasia.
N1a1a1a-L708, formed ca. 6800 BC, TMRCA 5400 BC.
N1a1a1a2-B211/Y9022, formed ca. 5400 BC, TMRCA 1900 BC:
The deepest clade within N3a is N3a1-B211, mostly present in the Volga-Uralic region and western Siberian Khanty and Mansi populations.
N1a1a1a1a-L392/L1026), formed ca. 4400 BC, TMRCA 2800 BC:
The neighbor clade, N3a3’6-CTS6967, spreads from eastern Siberia to the eastern part of Fennoscandia and the Baltic States
N1a1a1a1a1a-CTS2929/VL29, formed ca. 2100 BC, TMRCA 1600 BC:
In Europe, the clade N3a3-VL29 encompasses over a third of the present-day male Estonians, Latvians, and Lithuanians but is also present among Saami, Karelians, and Finns (Table S2 and Figure 3). Among the Slavic-speaking Belarusians, Ukrainians, and Russians, about three-fourths of their hg N3 Y chromosomes belong to hg N3a3.
In the post on Finno-Permic expansions, I depicted what seems to me the most likely way of infiltration of N1c-L392 lineages with Akozino warrior-traders into the western Finno-Ugric populations, with an origin around the Barents sea.
This includes the potential spread of (a minority of) N1c-B211 subclades due to contacts with Anonino on both sides of the Urals, through a northern route of forest and forest-steppe regions (equivalent to the distribution of Cherkaskul compared to Andronovo), given the spread of certain subclades in Ugric populations.
NOTE. An alternative possibility is the association of certain B211 subclades with a southern route of expansion with Pre-Scythian and Scythian populations, under whose influence the Ananino culture emerged -which would imply a very quick infiltration of certain groups of haplogroup N everywhere among Finno-Ugrics on both sides of the Urals – , and also the expansion of some subclades with Turkic-speaking peoples, who apparently expanded with alliances of different peoples. Both (Scythian and Turkic) populations expanded from East Asia, where haplogroup N (including N1c) was present since the Neolithic. I find this a worse model of expansion for upper clades, but – given the YFull estimates and the presence of this haplogroup among Turkic peoples – it is a possibility for many subclades.
N1a1a1a1a2-Z1936, formed ca. 2800 BC, TMRCA 2400 BC:
The only notable exception from the pattern are Russians from northern regions of European Russia, where, in turn, about two-thirds of the hg N3 Y chromosomes belong to the hg N3a4-Z1936—the second west Eurasian clade. Thus, according to the frequency distribution of this clade, these Northern Russians fit better among other non-Slavic populations from northeastern Europe. N3a4 tends to increase in frequency toward the northeastern European regions but is also somewhat unexpectedly a dominant hg N3 lineage among most Turcic-speaking Volga Tatars and South-Ural Bashkirs.
N1a1a1a1a4-M2019 (previously N3a2), formed ca. 4400 BC, TMRCA 1700 BC:
Sub-hg N3a2-M2118 is one of the two main bifurcating branches in the nested cladistic structure of N3a2’6-M2110. It is predominantly found in populations inhabiting present-day Yakutia (Republic of Sakha) in central Siberia and at lower frequencies in the Khanty and Mansi populations, which exhibit a distinct Y-STR pattern (Table S7) potentially intrinsic to an additional clade inside the sub-hg N3a2
The second widespread sub-clade of hg N is N2a. (…):
N1a2b-P43 (B523/FGC10846/Y3184), formed ca. 6800 BC, TMRCA ca. 2700 BC:
The absolute majority of N2a individuals belong to the second sub-clade, N2a1-B523, which diversified about 4.7 kya (95% CI = 4.0–5.5 kya). Its distribution covers the western and southern parts of Siberia, the Taimyr Peninsula, and the Volga-Uralic region with frequencies ranging from from 10% to 30% and does not extend to eastern Siberia (…)
The “European” branch suggested earlier from Y-STR patterns turned out to consist of two clades
N1a2b2a-Y3185/FGC10847, formed ca. 2200 BC, TMRCA 800 BC:
N2a1-L1419, spread mainly in the northern part of that region.
N1a2b2b1-B528/Y24382, formed ca. 900 BC, TMRCA ca. 900 BC:
N2a1-B528, spread in the southern Volga-Uralic region.
We also have a good idea of the distribution of haplogroup R1a-Z645 in ancient samples. Its subclades were associated with the Corded Ware expansion, and some of them fit quite well the early expansion of Finno-Permic, Ugric, and Samoyedic peoples to the east.
This is how the modern distribution of R1a among Uralians looks like, from the latest report in Tambets et al. (2018):
Among Fennic populations, Estonians and Karelians (ca. 1.1 million) have not suffered the greatest bottleneck of Finns (ca. 6-7 million), and show thus a greater proportion of R1a-Z280 than N1c subclades, which points to the original situation of Fennic peoples before their expansion. To trust Finnish Y-DNA to derive conclusions about the Uralic populations is as useful as relying on the Basque Y-DNA for the language spread by R1b-P312…
Among Volga-Finnic populations, Mordovians (the closest to the original Uralic cluster, see above) show a majority of R1a lineages (27%).
Hungarians (ca. 13-15 million) represent the majority of Ugric (and Finno-Ugric) peoples. They are mainly R1a-Z280, also R1a-Z2123, have little N1c, and lack Siberian ancestry, and represent thus the most likely original situation of Ugric peoples in 4th century AD (read more on Avars and Hungarians).
Among Samoyedic peoples, the Selkup, the southernmost ones and latest to expand – that is, those not heavily admixed with Siberian populations – , also have a majority of R1a-Z2123 lineages (see also here for the original Samoyedic haplogroups to the south).
To understand the relevance of Hungarians for Ugric peoples, as well as Estonians, Karelians, and Mordovians (and northern Russians, Finno-Ugric peoples recently Russified) for Finno-Permic peoples, as opposed to the Circum-Arctic and East Siberian populations, one has to put demographics in perspective. Even a modern map can show the relevance of certain territories in the past:
Summary of ancestry + haplogroups
Fennic and Samic populations seem to be clearly influenced by Palaeo-Laplandic peoples, whereas Volga-Finnic and especially Permic populations may have received gene flow from both, but essentially Palaeo-Siberian influence from the north and east.
The fact that modern Mansis and Khantys offer the highest variation in N1a subclades, and some of the highest “Siberian ancestry” among non-Nganasans, should have raised a red flag long ago. The fact that Hungarians – supposedly stemming from a source population similar to Mansis – do not offer the same amount of N subclades or Siberian ancestry (not even close), and offer instead more R1a, in common with Estonians (among Finno-Samic peoples) and Mordvins (among Volga-Finnic peoples) should have raised a still bigger red flag. The fact that Nganasans – the model for Siberian ancestry – show completely different N1a2b-P43 lineages should have been a huge genetic red line (on top of the anthropological one) to regard them as the Uralian-type population.
It is not hard to model the stepped arrival, infiltration, and/or resurge of N subclades and “Siberian ancestries”, as well as their gradual expansion in certain regions, associated with certain migrations first – such as the expansions to the Circum-Arctic region, and later the Scythian- and Turkic-related movements – , as well as limited regional developments, like the known bottleneck in Finns, or the clear late expansion of Ugric and Samoyedic languages to the north among nomadic Palaeo-Siberians due to traditions of exogamy and multilingualism. This fits quite well with the different arrival of N (N1c and xN1c) lineages to the different Uralic-speaking groups, and to the stepped appearance of “Siberian ancestry” in the different regions.
It is evident that a lot of people were too attached to the idea of Palaeolithic R1b lineages ‘native’ to western Europe speaking Basque languages; of R1a lineages speaking Indo-European and spreading with Yamna; and N lineages ‘native’ to north-eastern Europe and speaking Uralic, and this is causing widespread weeping and gnashing of teeth (instead of the joy of discovering where one’s true patrilineal ancestors come from, and what language they spoke in each given period, which is the supposed objective of genetic genealogy…)
As far as I know – and there might be many other similar pet theories out there – there have been proposals of “modern Balto-Slavic-like” populations (in an obvious circular reasoning based on modern populations) in some Scythian clusters of the Iron Age.
NOTE. I will not enter into “Balto-Slavic-like R1a” of the Late Bronze Age or earlier because no one can seriously believe at this point of development of Population Genetics that autosomal similarity predating 1,500+ years the appearance of Slavs equates to their (ethnolinguistic) ancestral population, without a clear intermediate cultural and genetic trail – something we lack today in the Slavic case even for the late Roman period…
We also know of R1a-Z280 lineages in Srubna, probably expanding to the west. With that in mind, and knowing that Palaeo-Germanic was in close contact with Finno-Samic while both were already separated but still in contact, and that Palaeo-Germanic was also in contact and closely related to a ‘Temematic’ distinct from Balto-Slavic (and also that early Proto-Baltic and Proto-Slavic from the Roman Iron Age and later were in contact with western Uralic) this will be the linguistic map of the Iron Age if R1a is considered to expand Indo-European from some kind of “patron-client” relationship with west Yamna:
My problem with this proposal is that it is obviously beholden to the notion of the uninterrupted cultural, historic and ethnic continuity in certain territories. This bias is common in historiography (von Falkenhausen 1993), but it extends even more easily into the lesser known prehistory of any territory, and now more than ever some people feel the need to corrupt (pre)history based on their own haplogroups (or the majority haplogroups of their modern countries). However, more than on philosophical grounds, my rejection is based on facts: this picture is not what the combination of linguistic, archaeological, and genetic data shows. Period.
Nevertheless, if Yamna + Corded Ware represented the “big and early expansion” of Germanic and Italo-Celtic peoples proper of the dream Nazi’s Lebensraum and Fascist’s spazio vitale proposals; Uralians were Siberian hunter-gatherers that controlled the whole eastern and northern Russia, and miraculously managed to push (ethnolinguistically) Neolithic agropastoralists to the west during and after the Iron Age, with gradual (and often minimal) genetic impact; and Balto-Slavic peoples were represented by horse riders from Pokrovka/Srubna, hiding then somewhere around the forest-steppe until after the Scythian expansion, and then spreading their language (without much genetic impact) during the early Middle Ages…so be it.
Even though proposals of an Eastern Uralic (or Ugro-Samoyedic) group are in the minority – and those who support it tend to search for an origin of Uralic in Central Asia – , there is nothing wrong in supporting this from the point of view of a western homeland, because the eastward migration of both Proto-Ugric and Pre-Samoyedic peoples may have been coupled with each other at an early stage. It’s like Indo-Slavonic: it just doesn’t fit the linguistic data as well as the alternative, i.e. the expansion of Samoyedic first, different from a Finno-Ugric trunk. But, in case you are wondering about this possibility, here is Häkkinen’s (2012) phonological argument:
The case of Samoyedic is quite similar to that of Hungarian, although the earliest Palaeo-Siberian contact languages have been lost. There were contacts at least with Tocharian (Kallio 2004), Yukaghir (Rédei 1999) and Turkic (Janhunen 1998). Samoyedic also:
a) has moved far from the related languages and has been exposed to strong foreign influence
b) shares a small number of common words with other branches (from Sammallahti 1988: only 123 ‘Uralic’ words, versus 390 ‘Uralic’ + ‘Finno-Ugric’ words found in other branches than Samoyedic = 31,5 %)
c) derives phonologically from the East Uralic dialect.
The phonological level is taxonomically more reliable, since it lacks the distortion caused by invisible convergence and false divergence at the lexical level. Thus we can conclude that the traditional taxonomic model, according to which Samoyedic was the first branch to split off from the Proto-Uralic unity, is just as incorrect as the view that Hungarian was the first branch to split off.
Late Uralic can be traced back to metallurgical cultures thanks to terms like PU *wäśka ‘copper/bronze’ (borrowed from Proto-Samoyedic *wesä into Tocharian); PU *äsa and *olna/*olni, ‘lead’ or ‘tin’, found in *äsa-wäśka ‘tin-bronze’; and e.g. *weŋći ‘knife’, borrowed into Indo-Iranian (through the stage of vocalization of nasals), appearing later as Proto-Indo-Aryan *wāćī ‘knife, awl, axe’.
It is known that the southern regions of the Abashevo culture developed Proto-Indo-Iranian-speaking Sintashta-Petrovka and Pokrovka (Early Srubna). To the north, however, Abashevo kept its Uralic nature, with continuous contacts allowing for the spread of lexicon – mainly into Finno-Ugric – , and phonetic influence – mainly Uralisms into Proto-Indo-Iranian phonology (read more here).
The northern part of Abashevo (just like the south) was mainly a metallurgical society, with Abashevo metal prospectors found also side by side with Sintashta pioneers in the Zeravshan Valley, near BMAC, in search of metal ores. About the Seima-Turbino phenomenon, from Parpola (2013):
From the Urals to the east, the chain of cultures associated with this network consisted principally of the following: the Abashevo culture (extending from the Upper Don to the Mid- and South Trans-Urals, including the important cemeteries of Sejma and Turbino), the Sintashta culture (in the southeast Urals), the Petrovka culture (in the Tobol-Ishim steppe), the Taskovo-Loginovo cultures (on the Mid- and Lower Tobol and the Mid-Irtysh), the Samus’ culture (on the Upper Ob, with the important cemetery of Rostovka), the Krotovo culture (from the forest steppe of the Mid-Irtysh to the Baraba steppe on the Upper Ob, with the important cemetery of Sopka 2), the Elunino culture (on the Upper Ob just west of the Altai mountains) and the Okunevo culture (on the Mid-Yenissei, in the Minusinsk plain, Khakassia and northern Tuva). The Okunevo culture belongs wholly to the Early Bronze Age (c. 2250–1900 BCE), but most of the other cultures apparently to its latter part, being currently dated to the pre-Andronovo horizon of c. 2100–1800 BCE (cf. Parzinger 2006: 244–312 and 336; Koryakova & Epimakhov 2007: 104–105).
The majority of the Sejma-Turbino objects are of the better quality tin-bronze, and while tin is absent in the Urals, the Altai and Sayan mountains are an important source of both copper and tin. Tin is also available in southern Central Asia. Chernykh & Kuz’minykh have accordingly suggested an eastern origin for the Sejma-Turbino network, backing this hypothesis also by the depiction on the Sejma-Turbino knives of mountain sheep and horses characteristic of that area. However, Christian Carpelan has emphasized that the local Afanas’evo and Okunevo metallurgy of the Sayan-Altai area was initially rather primitive, and could not possibly have achieved the advanced and difficult technology of casting socketed spearheads as one piece around a blank. Carpelan points out that the first spearheads of this type appear in the Middle Bronze Age Caucasia c. 2000 BCE, diffusing early on to the Mid-Volga-Kama-southern Urals area, where “it was the experienced Abashevo craftsmen who were able to take up the new techniques and develop and distribute new types of spearheads” (Carpelan & Parpola 2001: 106, cf. 99–106, 110). The animal argument is countered by reference to a dagger from Sejma on the Oka river depicting an elk’s head, with earlier north European prototypes (Carpelan & Parpola 2001: 106–109). Also the metal analysis speaks for the Abashevo origin of the Sejma-Turbino network. Out of 353 artefacts analyzed, 47% were of tin-bronze, 36% of arsenical bronze, and 8.5% of pure copper. Both the arsenical bronze and pure copper are very clearly associated with the Abashevo metallurgy.
The Abashevo metal production was based on the Volga-Kama-Belaya area sandstone ores of pure copper and on the more easterly Urals deposits of arsenical copper (Figure 9). The Abashevo people, expanding from the Don and Mid-Volga to the Urals, first reached the westerly sandstone deposits of pure copper in the Volga and Kama basins, and started developing their metallurgy in this area, before moving on to the eastern side of the Urals to produce harder weapons and tools of arsenical copper. Eventually they moved even further south, to the area richest in copper in the whole Urals region, founding there the very strong and innovative Sintashta culture.
Regarding the most likely expansion of Eastern Uralic peoples:
Nataliya L’vovna Chlenova (1929–2009; cf. Korenyako & Ku’zminykh 2011) published in 1981 a detailed study of the Cherkaskul’ pottery. In her carefully prepared maps of 1981 and 1984 (Figure 10), she plotted Cherkaskul’ monuments not only in Bashkiria and the Trans-Urals, but also in thick concentrations on the Upper Irtysh, Upper Ob and Upper Yenissei, close to the Altai and Sayan mountains, precisely where the best experts suppose the homeland of Proto-Samoyed to be.
The Cherkaskul’ culture was transformed into the genetically related Mezhovka culture (c. 1500–1000 BCE), which occupied approximately the same area from the Mid-Kama and Belaya rivers to the Tobol river in western Siberia (cf. Parzinger 2006: 444–448; Koryakova & Epimakhov 2007: 170–175). The Mezhovka culture was in close contact with the neighbouring and probably Proto-Iranian speaking Alekseevka alias Sargary culture (c. 1500–900 BCE) of northern Kazakhstan (Figure 4 no. 8) that had a Fëdorovo and Cherkaskul’ substratum and a roller pottery superstratum (cf. Parzinger 2006: 443–448; Koryakova & Epimakhov 2007: 161–170). Both the Cherkaskul’ and the Mezhovka cultures are thought to have been Proto-Ugric linguistically, on the basis of the agreement of their area with that of Mansi and Khanty speakers, who moreover in their Fëdorovo-like ornamentation have preserved evidence of continuity in material culture (cf. Chlenova 1984; Koryakova & Epimakhov 2007: 159, 175).
The Mezhovka culture was succeeded by the genetically related Gamayun culture (c. 1000–700 BCE) (cf. Parzinger 2006: 446; 542–545).
From the Gamayun culture descend Trans-Urals cultures in close contact with Finno-Permic populations of the Cis-Ural region:
[Proto-Mansi] Itkul’ culture (c. 700–200 BCE) distributed along the eastern slope of the Ural Mountains (cf. Parzinger 2006: 552–556). Known from its walled forts, it constituted the principal Trans-Uralian centre of metallurgy in the Iron Age, and was in contact with both the Anan’ino and Akhmylovo cultures (the metallurgical centres of the Mid-Volga and Kama-Belaya region) and the neighbouring Gorokhovo culture.
[Proto-Hungarian] via the Vorob’evo Group (c. 700–550 BCE) (cf. Parzinger 2006: 546–549), to the Gorokhovo culture (c. 550–400 BCE) of the Trans-Uralian forest steppe (cf. Parzinger 2006: 549–552). For various reasons the local Gorokhovo people started mobile pastoral herding and became part of the multicomponent pastoralist Sargat culture (c. 500 BCE to 300 CE), which in a broader sense comprized all cultural groups between the Tobol and Irtysh rivers, succeeding here the Sargary culture. The Sargat intercommunity was dominated by steppe nomads belonging to the Iranian-speaking Saka confederation, who in the summer migrated northwards to the forest steppe
[Proto-Khanty] Late Bronze Age and Early Iron Age cultures related to the Gamayunskoe and Itkul’ cultures that extended up to the Ob: the Nosilovo, Baitovo, Late Irmen’, and Krasnoozero cultures (c. 900–500 BCE). Some were in contact with the Akhmylovo on the Mid-Volga.
Parpola (2012) connects the expansion of Samoyedic with the Cherkaskul variant of Andronovo. As we know, Andronovo was genetically diverse, which speaks in favour of different groups developing similar material cultures in Central Asia.
Juha Janhunen, author of the etymological dictionary of the Samoyed languages (1977), places the homeland of Proto-Samoyedic in the Minusinsk basin on the Upper Yenissei (cf. Janhunen 2009: 72). Mainly on the basis of Bulghar Turkic loanwords, Janhunen (2007: 224; 2009: 63) dates Proto-Samoyedic to the last centuries BCE. Janhunen thinks that the language of the Tagar culture (c. 800–100 BCE) ought to have been Proto-Samoyedic (cf. Janhunen 1983: 117– 118; 2009: 72; Parzinger 2001: 80 and 2006: 619–631 dates the Tagar culture c. 1000–200 BCE; Svyatko et al. 2009: 256, based on human bone samples, c. 900 BCE to 50 CE). The Tagar culture largely continues the traditions of the Karasuk culture (c. 1400–900 BCE), (…)
The use of a map of “Siberian ancestry” peaking in the arctic to show a supposedly late Uralic population movement (starting in the Iron Age!) seems to be the latest trend in population genomics:
I guess that would make this map of Neolithic farmer ancestry represent an expansion of Indo-European from the south, because Anatolia, Greece, Italy, southern France, and Iberia – where this ancestry peaks in modern populations – are among the oldest territories where Indo-European languages were recorded:
Probably not the right interpretation of this kind of simplistic data about modern populations, though…
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).
However, this ‘something’ that some people occasionally find in some Uralic populations is also common to other modern and ancient groups, and not so common in some other Uralic peoples. Simply put:
I already said this in the recent publication of Siberian samples, where a renamed and radiocarbon dated Finnish_IA clearly shows that Late Iron Age Saami (ca. 400 AD) had little “Siberian ancestry”, if any at all, representing the most likely Fennic (and Samic) ancestral components before their expansion into central and northern Finland, where they admixed with circum-polar peoples of asbestos ware cultures.
I will say that again and again, any time they report the so-called “Siberian ancestry” in Uralic samples, no matter how it is defined each time: it does not seem to be that special something people are looking for, but rather (at least in a great part) a quite old ancestral component forming an evident cline with EHG, whose best proximate source are Baikal_EN (and/or Devil’s Gate) at this moment, and thus also East European hunter-gatherers for Western Uralic peoples:
So either Samara_HG, Karelia_HG, and many other groups from eastern Europe all spoke Uralic according to this ADMIXTURE graphic (and the formation of steppe ancestry in the Volga-Ural region brought the Proto-Indo-European language to the steppes through the CHG/ANE expansion), or a great part of this “Siberian ancestry” found in modern Uralic-speaking populations is not what some people would like to think it is…
PCA clines can be looked for to represent expansions of ancient populations. Most recently, Flegontov et al. (2018) are attempting to do this with Asian populations:
For some Turkic groups in the Urals and the Altai regions and in the Volga basin, a different admixture model fits the data: the same West Eurasian source + Uralic- or Yeniseian-speaking Siberians. Thus, we have revealed an admixture cline between Scythians and the Iranian farmer genetic cluster, and two further clines connecting the former cline to distinct ancestry sources in Siberia. Interestingly, few Wusun-period individuals harbor substantial Uralic/Yeniseian-related Siberian ancestry, in contrast to preceding Scythians and later Turkic groups characterized by the Tungusic/Mongolic-related ancestry. It remains to be elucidated whether this genetic influx reflects contacts with the Xiongnu confederacy. We are currently assembling a collection of samples across the Eurasian steppe for a detailed genetic investigation of the Hunnic confederacies.
There are potential errors with this approach:
The main one is practical – does a modern cline represent an ancestral language? The answer is: sometimes. It depends on the anthropological context that we have, and especially on the precision of the PCA:
The ‘Europe’, ‘Middle East’, etc. clines of the above PCA do not represent one language, but many. For starters, the PCA includes too many (and modern) populations, its precision is useless for ethnolinguistic groups. Which is the right level? Again, it depends.
The other error is one of detail of the clines drawn (which, in turn, depends on the precision of the PCA). For example, we can draw two paralell lines (or even one line, as in Flegontov et al. above) in one PCA graphic, but we still don’t have the direction of expansion. How do we know if this supposed “Uralic-speaking cline” goes from one region to the other? For that level of detail, we should examine closely modern Uralic-speaking peoples and Circum-Arctic populations:
The real ancient Uralic cluster (drawn above in blue) is thus probably from a North-East European source (probably formed by Battle Axe / Fatyanovo-Balanovo / Abashevo) to the east into Siberian populations, and to the north into Laplandic populations (see below also on Mezhovska ancestry for the drawn ‘European cline’, which some may a priori wrongly assume to be quite late).
The fact that the three formed clines point to an admixture of CWC-related populations from North-Eastern Europe, and that variation is greater at the Palaeo-Laplandic and Palaeo-Siberian extremities compared to the CWC-related one, also supports this as the correct interpretation.
However, judging by the two main clines formed, one could be alternatively inclined to interpret that Palaeo-Laplandic and Palaeo-Siberian populations formed a huge ancestral “Uralic” ghost cluster in Siberia (spanning from the Palaeo-Laplandic to the Palaeo-Siberian one), and from there expanded Finno-Samic on one hand, and “Volga-Ugro-Samoyed” on the other. That poses different problems: an obvious linguistic and archaeological one – which I assume a lot of people do not really care about – , and a not-so-obvious genetic one (see below for ancient samples and for the expansion of haplogroup N).
Unlike this PCA with ancient samples, where Bell Beaker clines could be a rough approximation to the real sources for each population, and where a cluster spanning all three depicted Early Bronze Age clusters could give a rough proximate source of European Bell Beakers in Hungary (and where one can even distinguish the Y-DNA bottlenecks in the L23 trunk created by each cline) the PCA of modern Uralic populations is probably not suitable for a good estimate of the ancient situation, which may be found shifted up or down of the drawn “Uralic” cluster along East European groups.
After all, we already know that the Siberian cline shows probably as much an ancient admixture event – from the original Uralic expansion to the east with Corded Ware ancestry – as another more recent one – a westward migration of Siberian ancestry (or even more than one). While we know with more or less exactitude what happened with the Palaeo-Laplandic admixture by expanding Proto-Finno-Samic populations (see here), the Proto-Ugric and Pre-Samoyedic populations formed probably more than one cline during the different ancient migrations through central Asia.
Apparently, the Corded Ware expansion to the east was not marked by a huge change in ancestry. While the final version of Narasimhan et al. (2018) may show a little more detail about other forest-steppe Seima-Turbino/Andronovo-related migrations (and thus also Eastern Uralic peoples), we have already had enough information for quite some time to get a good idea.
Mezhovska‘s position is similar to the later Pre-Scythian and Scythian populations. There are some interesting details: apart from haplogroup R1a-Z280 (CTS1211+), there is one R1b-M269 (PF6494+), probably Z2103, and an outlier (out of three) in a similar position to the recently described central/southern Scythian clusters.
NOTE. The finding of R1b-M269 in the forest-steppe is probably either 1) from an Afanasevo-Okunevo origin, or 2) from an admixture with neighbouring Andronovo-related populations, such as Sargary. A third, maybe less likely option is that this haplogroup admixed with Abashevo directly (as it happened in Sintashta, Potapovka, or Pokrovka) and formed part of early Uralic migrations. In any case, since Mezhovska is a Bronze Age society from the Urals region, its association with R1b-Z2103 – like the association of R1b-Z2103 in Scythian clusters – cannot be attributed to “Thracian peoples”, a link which is (as I already said) too simplistic.
The drawn “European cline” of Hungarians (see above), leading from ‘west-like’ Mansi to Hungarian populations – and hosting also Finnic and Estonian samples – , cannot therefore be attributed simply to late “Slavic/Balkan-like” admixture.
Karasuk – located further to the east – is basically also Corded Ware peoples showing clearly a recent admixture with local ANE / Baikal_EN-like populations. In terms of haplogroups it shows haplogroup Q, R1a-Z2124, and R1a-Z2123, later found among early Hungarians, and present also in ancient Samoyedic populations now acculturated.
The most interesting aspect of both Mezhovska and Karasuk is that they seem to diverge from a point close to Ukraine_Eneolithic, which is the supposed ancestral source of Corded Ware peoples (read more about the formation of “steppe ancestry”). This means that Eastern Uralians derive from a source closer to Middle Dnieper/Abashevo populations, rather than Battle Axe (shifted to Latvian Neolithic), which is more likely the source prevalent in Finno-Permic peoples.
Their initial admixture with (Palaeo-)Siberian populations is thus seen already starting by this time in Mezhovska and especially in Karasuk, but this process (compared to modern populations) is incomplete:
We know now that Samic peoples expanded during the Late Iron Age into Palaeo-Laplandic populations, admixing with them and creating this modern cline. Finns expanded later to the north (in one of their known genetic bottlenecks), admixing with (and displacing) the Saami in Finland, especially replacing their male lines.
So how did Ugric and Samoyedic peoples admix with Palaeo-Siberian populations further, to obtain their modern cline? The answer is, logically, with East Asian migrations related to forest-steppe populations of Central Asia after the Mezhovska and Karasuk periods, i.e. during the Iron Age and later. Other groups from the forest-steppe in Central Asia show similar East Asian (“Siberian”) admixture. We know this from Narasimhan et al. (2018):
(…) we observe samples from multiple sites dated to 1700-1500 BCE (Maitan, Kairan, Oy_Dzhaylau and Zevakinsikiy) that derive up to ~25% of their ancestry from a source related to present-day East Asians and the remainder from Steppe_MLBA. A similar ancestry profile became widespread in the region by the Late Bronze Age, as documented by our time transect from Zevakinsikiy and samples from many sites dating to 1500-1000 BCE, and was ubiquitous by the Scytho-Sarmatian period in the Iron Age.
Flegontov: Present day Turkic speakers fall into two clusters of admixture patterns (Uralic/Yenisean and Tungussic/Mngolic) based on genomic data with ancient Turks belonging almost exclusively to the first cluster. #ISBA8
The Ugric-speaking Sargat culture in Western Siberia shows the expected mixture of haplogroups (ca. 500 BC – 500 AD), with 5 samples of hg N and 2 of hg R1a1, in Pilipenko et al. (2017). Although radiocarbon dates and subclades are lacking, N lineages probably spread late, because of the late and gradual admixture of Siberian cultures into the Sargat melting pot.
The observed reduction in the genetic distance between the Middle Tagar population and other Scythian like populations of Southern Siberia(Fig 5; S4 Table), in our opinion, is primarily associated with an increase in the role of East Eurasian mtDNA lineages in the gene pool (up to nearly half of the gene pool) and a substantial increase in the joint frequency of haplogroups C and D (from 8.7% in the Early Tagar series to 37.5% in the Middle Tagar series). These features are characteristic of many ancient and modern populations of Southern Siberia and adjacent regions of Central Asia, including the Pazyryk population of the Altai Mountains.
Before the Iron Age, the Karasuk and Mezhovska population were probably already somehow ‘to the north’ within the ancient Steppe-Altai cline (see image below9 created by expanding Seima-Turbino- and Andronovo-related populations. During the Iron Age, further Siberian contributions with Iranian expansions must have placed Uralians of the Central Asian forest-steppe areas much closer to today’s Palaeo-Siberian cline.
However, the modern genetic picture was probably fully developed only in historic times, when Samoyedic and Ugric languages expanded to the north, only in part admixing further with Palaeo-Siberian-speaking nomads from the Circum-Arctic region (see here for a recent history of Samoyedic Enets), which justifies their more recent radical ‘northern shift’.
This late acquisition of the language by Palaeo-Siberian nomads (without much population replacement) also justifies the wide PCA clusters of very small Siberian populations. See for example in the PCA from Tambets et al. (2018):
For their relationship with modern Mansi, we have information on Hungarian conqueror populations from Neparáczki et al. (2018):
Moreover, Y, B and N1a1a1a1a Hg-s have not been detected in Finno-Ugric populations [80–84], implying that the east Eurasian component of the Conquerors and Finno-Ugric people are probably not directly related. The same inference can be drawn from phylogenetic data, as only two Mansi samples appeared in our phylogenetic trees on the side branches (S1 Fig, Networks; 1, 4) suggesting that ancestors of the Mansis separated from Asian ancestors of the Conquerors a long time ago. This inference is also supported by genomic Admixture analysis of Siberian and Northeastern European populations , which revealed that Mansis received their eastern Siberian genetic component approximately 5–7 thousand years ago from ancestors of modern Even and Evenki people. Most likely the same explanation applies to the Y-chromosome N-Tat marker which originated from China [86,87] and its subclades are now widespread between various language groups of North Asia and Eastern Europe .
The genetic picture of Hungarians (their formed cline with Mansi and their haplogroups) may be quite useful for the true admixture found originally in Mansi peoples at the beginning of the Iron Age. By now it is clear even from modern populations that Steppe_MLBA ancestry accompanied the Uralic expansion to the east (roughly approximated in the graphic with Afanasievo_EBA + Bichon_LP EasternHG_M):
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.
EDIT (28 OCT): Notice also the appearance of N1a1a1a1a1a1a1-L1025 in Lithuania (ca. 300 AD), from Damgaard (Nature 2018); the N1c sample of the Krivichi Pskov Long Barrows culture (ca. 8th-10th c. AD), and N1a1a1a1a1a1a7-Y4341 among late Vikings from Sigtuna (ca. 10th-12th c. AD) in Krzewinska (2018).
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]
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.
Interesting excerpts (emphasis mine, some references deleted for clarity):
The town of Sigtuna in eastern central Sweden was one of the pioneer urban hubs in the vast and complex communicative network of the Viking world. The town that is thought to have been royally founded was planned and organized as a formal administrative center and was an important focal point for the establishment of Christianity . The material culture in Sigtuna indicates that the town had intense international contacts and hosted several cemeteries with a Christian character. Some of them may have been used by kin-based groups or by people sharing the same sociocultural background. In order to explore the character and magnitude of mobility and migration in a late Viking Age town, we generated and analyzed genomic (n = 23) and strontium isotope (n = 31) data from individuals excavated in Sigtuna.
The mitochondrial genomes were sequenced at 1.5× to 367× coverage. Most of the individuals were assigned to haplogroups commonly found in current-day Europeans, such as H, J, and U [14, 26, 27]. All of these haplotypes are present in Scandinavia today.
The Y chromosome haplogroups were assigned in seven males. The Y haplogroups include I1a, I2a, N1a, G2a, and R1b. Two identified lineages (I2a and N1a) have not been found in modern-day Sweden or Norway [28, 29]. Haplogroups I and N are associated with eastern and central Europe, as well as Finno-Ugric groups . Interestingly, I2a was previously identified in a middle Neolithic Swedish hunter-gatherer dating to ca. 3,000 years BCE .
In Sigtuna, the genetic diversity in the late Viking Age was greater than the genetic diversity in late Neolithic and Bronze Age cultures (Unetice and Yamnaya as examples) and modern East Asians; it was on par with Roman soldiers in England but lower than in modern-day European groups (GBR and FIN; Figure 2B). Within the town, the group excavated at church 1 has somewhat greater diversity than that at cemetery 1. Interestingly, the diversity at church 1 is nearly as high as that observed in Roman soldiers in England, which is remarkable, since the latter was considered to be an exceptionally heterogeneous group in contemporary Europe .
Different sex-related mobility patterns for Sigtuna inhabitants have been suggested based on material culture, especially ceramics. Building on design and clay analyses, some female potters in Sigtuna are thought to have grown up in Novgorod in Rus’ . Moreover, historical sources mention female mobility in connection to marriage, especially among the elite from Rus’ and West Slavonic regions [41, 42]. Male mobility is also known from historical sources, often in connection to clergymen moving to the town .
Interestingly, we found a number of individuals from Sigtuna to be genetically similar to the modern-day human variation of eastern Europeans, and most harbor close genetic affinities to Lithuanians (Figure 2A). The strontium isotope ratios in 28 adult individuals with assigned biological sex and strontium values obtained from teeth (23 M1 and five M2) show that 70% of the females and 44% of the males from Sigtuna were non-locals (STAR Methods). The difference in migrant ratios between females and male mobility patterns was not statistically significant (Fisher’s exact test, p = 0.254 for 28 individuals and p = 0.376 for 16 individuals). Hence, no evidence of a sex-specific mobility pattern was found.
(…) As these social groups are not mirrored by our genetic or strontium data, this suggests that the inclusion in them was not based on kinship. Therefore, it appears as if socio-cultural factors, not biological bonds, governed where people were interred (i.e., the choice of cemetery).
Interesting from this paper is the higher genetic (especially Y-DNA) diversity found in more recent periods (see e.g. here) compared to Neolithic and Bronze Age cultures, which is probably the reason behind some obviously wrong interpretations, e.g. regarding links between Yamna and Corded Ware populations.
The sample 84001, a “first-generation short-distance migrant” of haplogroup N1c-L392 (N1a in the new nomenclature) brings yet more proof of how:
Admixture changes completely within a certain number of generations. In this case, the N1c-L392 sample clusters within the genetic variation of modern Norwegians, near to the Skane Iron Age sample, and not with its eastern origin (likely many generations before).
This haplogroup appeared quite late in Fennoscandia but still managed to integrate and expand into different ethnolinguistic groups; in this case, this individual was probably a Viking of Nordic language, given its genetic admixture and its non-local (but neighbouring Scandinavian) strontium values.
In recent decades, evidence has accumulated for comparable enclosures of later dates, including the Early Bronze Age Únětice Culture between 2200 and 1600 BC, and thus into the chronological and cultural context of the Nebra sky disc. Based on the analysis of one of these enclosure sites, recently excavated at Pömmelte on the flood plain of the Elbe River near Magdeburg, Saxony-Anhalt, and dating to the late third millennium BC
The main occupation began at 2321–2211 cal BC, with the stratigraphically earliest features containing exclusively Bell Beaker finds. Bell Beaker ceramics continue after 2204–2154 cal BC (boundary occupation I/II), although they were probably undecorated, but are now complemented by Únětice Culture (and other Early Bronze Age) types. At this time, with features common to both cultures predominate. Only contexts dating to the late main occupation phase (late phase II) and thereafter contained exclusively Únětice Culture finds. Evidently, the bearers of the Bell Beaker Culture were the original builders of the enclosure. During a second phase of use, Final Neolithic and Early Bronze Age cultures coexisted and intermingled. The material remains, however, should not be taken as evidence for successive groups of differing archaeological cultures, but as witnesses to a cultural transition from the Bell Beaker Culture to the Únětice Culture (Spatzier 2015). The main occupation ended 2086–2021 cal BC with the deconstruction of the enclosure; Bell Beaker finds are now absent. Finally, a few features (among them one shaft) and radiocarbon dates attest the sporadic re-use of the site in a phase of abandonment/re-use that ended 1636– 1488 cal BC.
How the above-ground structures possibly influenced perception may reveal another layer of meaning that highlights social functions related to ritual. While zone I was disconnected from the surroundings by a ‘semi-translucent’ post-built border, zones II/III were separated from the outside world by a wooden wall (i.e. the palisade), and zone III probably separated individuals from the crowd gathered in zone II. Accessing the interior or centre therefore meant passing through transitional zones, to first be secluded and then segregated. Exiting the structure meant re-integration and re-connection. The experience possibly induced when entering and leaving the monument reflects the three stages of ‘rites of passage’ described by van Gennep (1909): separation, liminality and incorporation. The enclosure’s outer zone(s) represents the pre- and post-liminal phase; the central area, the liminal phase. Seclusion and liminality in the interior promoted a sense of togetherness, which can be linked to Turner’s “communitas” (1969: 132–33). We might therefore see monuments such as the Pömmelte enclosure as important communal structures for social regulation and the formation of identity.
(…) The long-term stability of these connotations must be emphasised. As with the tradition of making depositions, these meanings were valid from the start of the occupation — c. 2300 BC — until at least the early period following the deconstruction event, c. 2050 BC. While the spatial organisation and the solar alignment of the main entrances were maintained throughout the main occupation, stone axes and ‘formal’ graves indicate the continuation of the spatial concepts described above until the twentieth to nineteenth centuries BC.
These layers of meaning mirror parallel concepts of space including, although not necessarily restricted to, the formation of group identities (see Hansen & Meyer 2013: 5). They can perhaps be better understood as a ‘cosmological geography’ manifested in the symbolism of superimposed levels of conceptual ideas related to space and to certain cardinal points (Figure 8). This idea is closely related to Eliade’s (1959: 29–36) understanding of “organized — hence comicized — territory”, that is territory consecrated to provide orientation within the homogeneity of the chaotic ‘outside world’, and the equivalence of spatial consecration and cosmogony. Put differently, the Pömmelte enclosure can be interpreted as a man-made metaphor and an icon of the cosmos, reflecting the Weltanschauung (a comprehensive conception of the world) of the people who built and used it. By bringing together Eliade and Rappaport’s ideas of meaningfulness in relation to religious experience (Rappaport 1999: 391–95), it may be argued that Pömmelte was a place intended to induce oneness with the cosmos. In combining multiple layers that symbolically represent different aspects of life (first-ordermeaning), the enclosure became an icon metaphorically representing the world (second-order-meaning). As this icon was the place to reaffirm life symbolism ritually, through their actions, people perhaps experienced a sense of rootedness in, or unity with, the cosmos (highest-order-meaning). Although we can only speculate about the perceptions of ancient people, such a theory aiming to describe general principles of religious experience can provide insight.
The circular enclosure of Pömmelte is the first Central European monumental complex of primarily sacred importance that has been excavated and studied in detail. It reveals aspects of society and belief during the transition from the Final Neolithic to the Early Bronze Age, in the second half of the third millennium BC. Furthermore, it offers details of ritual behaviour and the way that people organised their landscape. A sacred interior was separated from the profane environment, and served as a venue for rites that secured the continuity of the social, spiritual and cosmic order. Ancestor worship formed another integral part of this: a mound-covered burial hut and a square-shaped ditch sanctuary (located, respectively, within and near the enclosure’s south-eastern sector; cf. Figure 2)—dating to 2880–2580 cal BC and attributed to the Corded Ware Culture (Spatzier 2017a: 235–44)—suggest that this site was deliberately chosen. With construction of the ring sanctuary, this place gained an immense expansion in meaning—comparable to Stonehenge. Through architectural transformation, both of these sites developed into sanctuaries with increasingly complex religious functions, including in relation to the cult of the dead. The cosmological and social functions, and the powerful symbolism of the Nebra sky disc and hoard (Meller 2010: 59–70), are reflected in Pömmelte’s monumental architecture.
All of these features—along with Pömmelte’s dating, function and complex ring structure—are well documented for British henge monuments (Harding 2003; Gibson 2005). The continuous use of circular enclosures in Central Europe from around 3000– 1500 BC remains to be confirmed, but strong evidence indicates usage spanning from the fifth to the first millennia BC (Spatzier 2017a: 273–96). From 2500 BC onwards, examples in Central Europe, Iberia and Bulgaria (Bertemes 2002; Escudero Carrillo et al. 2017) suggest a Europe-wide concept of sanctuary. This indicates that in extensive communication networks at the beginning of bronze metallurgy (Bertemes 2016), intellectual and religious contents circulated alongside raw materials. The henge monuments of the British Isles are generally considered to represent a uniquely British phenomenon, unrelated to Continental Europe; this position should now be reconsidered. The uniqueness of Stonehenge lies, strictly speaking, with its monumental megalithic architecture.
The Classical Bell Beaker heritage
No serious scholar can argue at this point against the male-biased East Bell Beaker migrations that expanded the European languages related to Late Proto-Indo-European-speaking Yamna (see David Reich’s comments), and thus most likely North-West Indo-European – the ancestor of Italo-Celtic, Germanic, and Balto-Slavic, apart from Pre-Celtic IE in the British Isles, Lusitano-Galician in Iberia, or Messapic in Italy (see here a full account).
With language, these migrants (several ten thousands) brought their particular Weltanschauung to all of Western, Central, and Northern Europe. Their admixture precisely in Hungary shows that they had close interactions with non-Indo-European peoples (genetically related to the Globular Amphorae culture), something that we knew from the dozens of non-Indo-European words reconstructed exclusively for North-West Indo-European, apart from the few reconstructed non-Indo-European words that NWIE shares with Palaeo-Balkan languages, which point to earlier loans from their ancestors, Yamna settlers migrating along the lower Danube.
It is not difficult to imagine that the initial East Bell Beaker group shared a newly developed common cosmological point of view that clashed with other neighbouring Yamna-related worldviews (e.g. in Balkan EBA cultures) after the cultural ties with Yamna were broken. Interesting in this respect is for example their developed (in mythology as in the new North-West Indo-European concept) *Perkwūnos, the weather god – probably remade (in language as in concept) from a Yamna minor god also behind Old Indian parjányas, the rain god – as one of the main gods from the new Pantheon, distinct from *Dyēus patēr, the almighty father sky god. In support of this, the word *meldh-n- ‘lightning’, behind the name of the mythological hammer of the weather god (cf. Old Norse Mjǫllnir or Latvian Milna), was also a newly coined North-West Indo-European term, although the myth of the hero slaying the dragon with the magical object is older.
Circular enclosures are known in Europe since the Neolithic. Also, the site selected for the Pömmelte enclosure had been used to bury Corded Ware individuals some centuries before its construction, and Corded Ware symbolism (stone axe vs. quern) is seen in the use given by Bell Beakers and later Únětice at this place. All this and other regional similarities between Bell Beakers and different local cultures (see here an example of Iberian Bell Beakers) points to syncretism of the different Bell Beaker groups with preceding cultures in the occupied regions. After all, their genealogical ancestors included also those of their maternal side, and not all encountered males disappeared, as is clearly seen in the resurge of previous paternal lineages in Central-East Europe and in Scandinavia. The admixture of Bell Beakers with previous groups (especially those of similar steppe-related ancestry from Corded Ware) needs more complex analyses to clarify potential early dialectal expansions (read what Iosif Lazaridis has to say).
The popular “big and early” expansions
These syncretic trends gave rise to distinct regional cultures, and eventually different local groups rose to power in the new cultural regions and ousted the old structures. Social norms, hierarchy, and pantheons were remade. Events like this must have been repeated again and again in Bronze and Iron Age Europe, and in many cases it was marked by a difference in the prevailing archaeological culture attested, and probably accompanied by certain population replacements that will be seen with more samples and studies of fine-scale population structure.
Some of these cultural changes, marked by evident haplogroup or admixture replacement, are defined as a ‘resurge’ of ancestry linked to previous populations, although that is obviously not equivalent to a resurge of a previous cultural group, because they usually represent just a successful local group of the same supraregional culture with a distinct admixture and/or haplogroup (see e.g. resurge of R1a-Z645 in Central-East European Bronze Age). Social, religious, or ethnic concepts may have changed in each of these episodes, along with the new prestige dialect.
This must have happened then many times during the hundreds (or thousands in some cases) of years until the first attestation of a precise ancient language and culture (read e.g. about one of the latest branches to be attested, Balto-Slavic). Ancient language contacts, like substrates or toponymy, can only rarely be detected after so many changes, so their absence (or the lack of proper studies on them) is usually not relevant – and certainly not an argument – in scholarly discussions. Their presence, on the other hand, is a proof of such contacts.
We have dozens of papers supporting Uralic dialectal substrate influence on Pre-Germanic, Proto-Balto-Slavic, and Pre- and Proto-Indo-Iranian (and even Proto-Celtic), as well as superstrate influence of Palaeo-Germanic (i.e. from Pre- to Proto-Germanic) and Proto-Balto-Slavic into Proto-Finno-Saamic, much stronger than the Indo-Iranian adstrate influence on Finno-Ugric (see the relative importance of each influence) which locates all these languages and their evolution to the north and west of the steppe (with Proto-Permic already separated, in North-East Europe, as is Proto-Ugric further east near the Urals), probably around the Baltic and Scandinavia after the expansion of Bell Beakers. These connections have been known in linguistics for decades.
Apart from some early 20th century scholars, only a minority of Indo-Europeanists support nowadays an Indo-European (i.e. centum) substrate for Balto-Slavic, to keep alive an Indo-Slavonic group based on a hypothetical 19th century Satem group; so e.g. Holzer with his Temematic, and Kortlandt supporting him, also with some supposed Indo-European substrate with heavy non-Indo-European influence for Germanic and Balto-Slavic, that now (thanks mainly to the views of the Copenhagen group) have been linked to the Corded Ware culture, as it has become clear even to them that Bell Beakers expanded North-West Indo-European.
For their part, only a minority among Uralicists, such as Kuz’mina, Parpola or Häkkinen, believe in an ‘eastern’ origin of Uralic languages, around the Southern Urals. Genomic finds – like their peers – are clearly not supporting their views. But even if we accept this hypothesis, there is little space beyond Abashevo and related East Corded Ware cultures after the recent papers on Corded Ware and Fennoscandian samples. And yet here we are:
substitutes arrows for Kron-like colors (where danger red = Indo-European) with the same end result of many other late 20th century whole-Europe Kurgan maps, linking Sredni Stog and Corded Ware with Yamna, but obviating the precise origin of Corded Ware peoples (is it Sredni Stog, or is it that immutable Middle Dnieper group? is it West Yamna, or Yamna Hungary? is it wool, or is it wheels?);
relegates Uralic speakers to a tiny corner, a ‘Volosovo’ cultural region, thus near Khvalynsk/Yamna (but not too much), that miraculously survives surrounded by all-early-splitting, all-Northern Eneolithic Indo-Europeans, thus considering Uralic languages irrelevant not only to locate the PIE Urheimat, but also to locate their own homeland; also, cultures identified in color with Uralic speakers expand until the Iron Age with enough care not to even touch in the map one of the known R1a samples published to date (because, for some people, apparently R1a must be Indo-European); and of course N1c or Siberian ancestry are irrelevant, too;
and adds findings of wheels and wool probably in support of some new ideas based on yet another correlation = causation argument (that I cannot then properly criticize without access to its reasoning beyond cute SmartArt-like symbols) similar to their model – already becoming a classic example of wrong use of statistical methods – based on the infamously named Yamnaya ancestral component™, which is obviously still used here, too.
The end result is thus similar to any other simplistic 1990s Gimbutas (or rather the recently radicalized IE Sredni Stog -> Corded Ware -> BBC version by the Danish workgroup) + 2000s R1a-map + 2010s Yamnaya ancestry™; but, hard to believe, it is published in mid-2018. A lot of hours of senseless effort, because after its publication it becomes ipso facto outdated.
For comparison of Yamna and Bell Beaker expansions, here is a recent simplistic, static (and yet more accurate) pair of maps, from the Reich Lab:
If the Copenhagen group keeps on pushing Gimbutas’ long ago outdated IE Sredni Stog -> Corded Ware theory as modified by Kristiansen, with their recently invented Corded Ware -> Bell Beaker model in genetics, at some point they are bound to clash with the Reich-Jena team, which seems to have less attachment to the classic Kurgan model and the wrong interpretations of the 2015 papers, and that would be something to behold. Because, as Cersei would say: “When you play the game of thrones, you win or you die. There is no middle ground.” And when you play the game of credibility, after so many, so wrong publications, well…
NOTE. I have been working on a similar GIS tool for quite some time, using my own maps and compiled genetic data, which I currently only use for my 2018 revision of the Indo-European demic diffusion model. Maybe within some weeks or months I will be able to publish the maps properly, after the revised papers. It’s a pitty that so much work on GIS and analysis with genetic data and cultural regions has to be duplicated, but I intend to keep some decent neutrality in my revised cultural maps, and this seems impossible at this point with some workgroups who have put all their eggs in one broken basket…
I wanted to share here some interesting excerpts (emphasis mine):
NOTE. I have avoided many detailed linguistic discussions. You should read the whole chapter to check them out.
The origins of the Germanic subfamily of Indo-European cannot be understood without acknowledging its interactions with a language group that has been its long-time neighbour: the Finnic subgroup of the Uralic language family. Indo-European and Uralic are linked to one another in two ways: they are probably related to one another in deep time — how deep is impossible to say3 — and Indo-European has been a constant source from which words were borrowed into Uralic languages, from the fourth millennium BC up to the present day.4 The section of the Uralic family that has always remained in close proximity to the Indo-European dialects which eventually turned into Germanic is Finnic. I use the term Finnic with a slightly idiosyncratic meaning : it covers the Finno-Saamic protolanguage and both of its children, Saami and Balto-Finnic.(…)
Linguistically, the relationship between Indo-European and Uralic has always been asymmetrical. While hundreds of loanwords flowed into Uralic languages from Indo-European languages such as Germanic, Balto-Slavic, Iranian, and Proto-Indo-European itself, hardly any Uralic loanwords have entered the Indo-European languages (apart from a few relatively late dialectal loans into e.g. Russian and the Scandinavian languages). This strongly suggests that Uralic speakers have always been more receptive to ideas coming from Indo-European–speaking areas than the other way around. This inequality probably began when farming and the entire way of life that accompanies it reached Uralic-speaking territory via Indo-European–speaking territory, so that Uralic speakers, who traditionally were hunter-gatherers of the mixed and evergreen forest zone of northeastern Europe and gradually switched to an existence as sedentary farmers, were more likely to pick up ideas and the words that go with them from Indo-European than from anywhere else.
Farming requires a different mind-set from a hunter-gatherer existence. Farmers are generally sedentary, model the landscape, and have an agricultural calendar to determine their actions. Hunter-gatherers of the northern forest zone are generally nomadic, and rather than themselves modelling the natural environment they are modelled by it: their calendar depends on when and where a particular natural resource is available.(…)
All of this is no doubt a simplification of the thousands of years of associations between speakers of Uralic and speakers of Indo-European, but the loanword evidence strongly suggests that by and large relations between the two groups were highly unequal. The single direction in which loanwords flowed, and the mass of loanwords involved, can be compared with the relation between Latin and the vernacular languages in the Roman Empire, almost all of which disappeared in favour of Latin. It is therefore certain that groups of Uralic speakers switched to Indo-European. The question is whether we can trace those groups and, more particularly, whether Finnic speakers switching to Indo-European were involved in creating the Indo-European dialect we now know as Germanic.
Convergence of Finnic and Germanic
What both have in common is that the sound structures of Finnic and Germanic, which started from very different beginnings, apparently came to resemble one another significantly. If that is what we observe, we must conclude that both languages converged as a result of contact.
During the approximately five to six millennia that separate Proto-Uralic from Modern Finnish, there was only one episode during which the consonantal system underwent a dramatic overhaul. This episode separates the Finno-Saamic protolanguage, which is phonologically extremely conservative, from the Balto-Finnic protolanguage, which is very innovative.
By the time Finno-Saamic developed into Balto-Finnic, the consonant system was very different:
In Balto-Finnic, the entire palatal series has been lost, apart from j, and the contrast between dentals and alveolars has disappeared: out of three different s-sounds only one remains. The fricatives ð and γ have been lost, and so has the velar nasal ŋ. The only increase has been in the number of long (geminate) consonants by the appearance of ss, mm, nn, and ll. The loss of separate alveolar and palatal series and the disappearance of ŋ could be conceived as convergences towards Proto-Germanic, which lacked such consonants. This is not obvious for the loss of the voiced fricatives γ, ð, which Proto-Germanic did possess. However, this way of comparing Balto-Finnic and Germanic is flawed in an important respect: what we are doing is assessing convergence by comparing the dynamic development from Finno-Saamic to Balto-Finnic to the static system of Proto-Germanic, as if Proto-Germanic is not itself the result of a set of changes to the ancestral Pre-Germanic consonantal system. If we wish to find out whether there was convergence and which language converged on which, what we should do, therefore, is to compare the dynamic development of Finno-Saamic to Balto-Finnic to the dynamic development of Pre-Germanic to Proto-Germanic, because only that procedure will allow us to state whether Balto-Finnic moved towards Proto-Germanic, or Proto-Germanic moved towards Balto-Finnic, or both moved towards a third language. The Pre-Germanic consonantal system can be reconstructed as follows: 7
The slashes in the second and third rows indicate the uncertainty about the Proto-Indo-European nature of the sounds involved. (…)
What resulted was the following Proto-Germanic consonant system:
We are now in a better position to answer the question whether Proto-Germanic and Balto-Finnic have converged. Three striking developments affected both languages:
Both languages lost the palatalized series of consonants (apart from j), which in both languages became non-palatalized.
Both languages developed an extensive set of long (geminate) consonants; Pre-Germanic had none, while Finno-Saamic already had a few.
Both languages developed an h.
These similarities between the languages are considerable.
The idea that perhaps both languages moved towards a lost third language, whose speakers may have been assimilated to both Balto-Finnic and Germanic, provides a fuller explanation but suffers from the drawback that it shifts the full burden of the explanation to a mysterious ‘language X’ that is called upon only in order to explain the developments in Proto-Germanic and Balto-Finnic. That comes dangerously close to circular reasoning.
Verner’s Law in Pre-Germanic
As we have seen in the preceding section, Verner’s law is a sound change that affected originally voiceless consonants, so *p , t , k , kj , kw, s of the Pre-Germanic system. These normally became the Proto-Germanic voiceless fricatives *f, θ, h, h, hw, s, respectively. But if *p, t, k etc. were preceded by an originally unstressed syllable, Verner’s law intervened and they were turned into voiced consonants. Those voiced consonants merged with the series *bh, dh, gh of the Pre-Germanic system and therefore subsequently underwent all changes that the latter did, turning out as *b/v , *d/ð , g/γ in the Proto-Germanic system (that is, v, ð, γ after a vowel and b, d, g in all other environments in the word). When *s was affected by Verner’s Law, a new phoneme *z arose. In a diagram:
While it is very common in the history of European languages for stress to influence the development of vowels, it only very rarely affected consonants in this part of the world. Verner’s law is a striking exception. It resembles a development which, on a much larger scale, affected Finno-Saamic: consonant gradation.(…)
In all Finno-Saamic languages, rhythmic gradation has become phonemic and fossilized. The connection between rhythmic gradation and Verner’s law is relatively straightforward: both processes involve changing a voiceless consonant after an unstressed syllable. (…)
We can therefore repeat for Proto-Uralic the argument that persuaded us earlier that gradation in Saami and Balto-Finnic must go back to the common Finno-Saamic protolanguage: the similarity of the gradation rules in Nganasan to those in Finno-Saamic is so specific and so detailed, and the phenomenon of gradation so rare in the languages of the world, that gradation must be reconstructed for the Uralic protolanguage.
Verner’s law turns all voiceless obstruents (Pre-Germanic *p, t, k, kj, kw, s) into voiced obstruents (ultimately Proto-Germanic *b/v , d/ð, g/γ, g/γ, gw, z) after a Pre-Germanic unstressed syllable. Rhythmic gradation turns all voiceless obstruents after an unstressed syllable into weak-grade consonants, which means that *p, t, k, s become Finnic *b/v , d/ð , g/γ, z. This is striking. Given the geographical proximity of Balto-Finnic and Germanic and given the rare occurrence of stress-related consonant changes in European languages, it would be unreasonable to think that Verner’s law and rhythmic gradation have nothing to do with one another.
It is very hard to accept, however, that gradation is the result of copying Verner’s law into Finnic. First of all, Verner’s law, which might account for rhythmic gradation, in no way accounts for syllabic gradation in Finnic. And, second, gradation can be shown to be an inherited feature of Finnic which goes all the way back to Proto-Uralic. Once one acknowledges that Verner’s law and gradation are causally linked and that gradation cannot be explained as a result of copying Verner’s law into Finnic, there remains only one possibility: Verner’s law is a copy of Finnic rhythmic gradation into Germanic. That means that we have finally managed to find what we were looking for all along: a Finnic sound feature in Germanic that betrays that Finnic speakers shifted to Germanic and spoke Germanic with a Finnic accent. The consequence of this idea is dramatic: since Verner’s law affected all of Germanic, all of Germanic has a Finnic accent.
On the basis of this evidence for Finnic speakers shifting to Germanic, it is possible to ascribe other, less specifically Finnic traits in Germanic to the same source. The most obvious trait is the fixation of the main stress on the initial syllable of the word. Initial stress is inherited in Finno-Saamic but was adopted in Germanic only after the operation of Verner’s law, quite probably under Finnic influence. The consonantal changes described in section V.3.1 can be attributed to Finnic with less confidence. The best case can be made for the development of geminate (double) consonants in Germanic, which did not inherit any of them, while Finno-Saamic inherited *pp, tt, kk, cc and took their presence as a cue to develop other geminates such as *nn and *ll . Possibly geminates developed so easily in Proto-Germanic because Finnic speakers (who switched to Germanic) were familiar with them. Other consonantal changes, such as the loss of the palatalized series in both Germanic and Balto-Finnic and the elimination of the different s- and c-phonemes, might have occurred for the same reason: if Balto-Finnic had undergone them earlier than Germanic, which we do not know, they could have constituted part of the Balto-Finnic accent in Germanic. An alternative take on those changes starts from the observation that they all constitute simplifications of an older, richer system of consonants. While simplifications can be and often are caused by language shift if the new speakers lacked certain phonemes in their original language, simplifications do not require an explanation by shift: languages are capable of simplifying a complex system all by themselves. Yet the similarities between the simplifications in Germanic and in Balto-Finnic are so obvious that one would not want to ascribe their co-occurrence to accidental circumstances.
Grimm’s Law in Proto-Germanic (speculative)
Voiceless lenis pronunciation of b, d, g is typical of the majority of German and Scandinavian dialects, so may well have been inherited from Proto-Germanic. Voiceless lenis is also the pronunciation that has been assumed to underlie the weak grades of Finno-Saamic single *p, t, k. If Proto-Germanic *b, d, g were indeed voiceless lenis, the single most striking result of the Germanic consonant shift is that it eliminated the phonological difference between voiced and voiceless consonants that Germanic had inherited from Proto-Indo-European (…) Since neither Finno-Saamic nor Balto-Finnic possessed a phonological difference between voiced and voiceless obstruents, its loss in Proto-Germanic can be regarded as yet another example of a Finnic feature in Germanic.
It is clear that this account of the first Germanic consonant shift as yet another example of Finnic influence is to some degree speculative. The point I am making is not that the Germanic consonant shift must be explained on the basis of Finnic influence, like Verner’s law and word-initial stress, only that it can be explained in this way, just like other features of the Germanic sound system discussed earlier, such as the loss of palatalized consonants and the rise of geminates.
A consequence of this account of the origins of the Proto-Germanic consonantal system is that the transition from Pre-Germanic to Proto-Germanic was entirely directed by Finnic. Or, to put it in less subtle words: Indo-European consonants became Germanic consonants when they were pronounced by Finnic speakers.
The vocalic system, on the other hand, presented less difficulties for both, Indo-European and Uralic speakers, since it was quite similar.
Schrijver goes on to postulate certain asymmetric differences in loans, especially with regard to Proto-Germanic, Balto-Finnic, Proto-Saamic, Proto-Baltic, and later contacts, including a potential non-Uralic, non-IE substrate language to justify some of these, which may in turn be connected with Kroonen’s agricultural substrate hypothesis of Proto-Germanic, and thus also with the other surviving Scandinavian Neolithic cultures before the eventual simplification of the cultural landscape during the Bronze Age.
Conclusion on the origin of Germanic
The Finnic-Germanic contact situation has turned out to be of a canonical type. To Finnic speakers, people who spoke prehistoric Germanic and its ancestor, Pre-Germanic, must have been role models. Why they were remains unclear. In the best traditions of Uralic–Indo-European contacts, Finnic speakers adopted masses of loanwords from (Pre-)Germanic. Some Finnic speakers even went a crucial step further and became bilingual: they spoke Pre-Germanic according to the possibilities offered by the Finnic sound system, which meant they spoke with a strong accent. The accent expressed itself as radical changes in the Pre-Germanic consonantal system and no changes in the Pre-Germanic vowel system. This speech variety became very successful and turned an Indo-European dialect into what we now know as Germanic. Bilingual speakers became monolingual speakers of Germanic.
What we do not know is for how long Finnic-Germanic bilingualism persisted. It is possible that it lasted for some time because both partners grew more alike even with respect to features whose origin we cannot assign to either of them (loss of palatalized consonants): this suggests, perhaps, that both languages became more similar because generally they were housed in the same brain. What we can say with more confidence is that the bilingual situation ultimately favoured Germanic over Finnic: loanwords continued to flow in one direction only, from Germanic to Finnic, hence it is clear that Germanic speakers remained role models.
This is as far as the linguistic evidence can take us for the moment.
NOTE. The ‘common’ loss of certain palatals, which Schrijver interprets as a change of Pre-Germanic from the inherited Proto-Indo-European, may in fact not be such – in the opinion of bitectalists, including us, and especially taking the North-West Indo-European reconstruction and the Corded Ware substrate hypothesis into account – , so this effect would be a rather unidirectional shift from Finnic to Germanic. On the other hand, certain palatalization trends which some have described for Germanic could in fact be explained precisely by this bidirectional influence.
The indigenous populations of inner Eurasia, a huge geographic region covering the central Eurasian steppe and the northern Eurasian taiga and tundra, harbor tremendous diversity in their genes, cultures and languages. In this study, we report novel genome-wide data for 763 individuals from Armenia, Georgia, Kazakhstan, Moldova, Mongolia, Russia, Tajikistan, Ukraine, and Uzbekistan. We furthermore report genome-wide data of two Eneolithic individuals (~5,400 years before present) associated with the Botai culture in northern Kazakhstan. We find that inner Eurasian populations are structured into three distinct admixture clines stretching between various western and eastern Eurasian ancestries. This genetic separation is well mirrored by geography. The ancient Botai genomes suggest yet another layer of admixture in inner Eurasia that involves Mesolithic hunter-gatherers in Europe, the Upper Paleolithic southern Siberians and East Asians. Admixture modeling of ancient and modern populations suggests an overwriting of this ancient structure in the Altai-Sayan region by migrations of western steppe herders, but partial retaining of this ancient North Eurasian-related cline further to the North. Finally, the genetic structure of Caucasus populations highlights a role of the Caucasus Mountains as a barrier to gene flow and suggests a post-Neolithic gene flow into North Caucasus populations from the steppe.
On North Eurasians
In a PCA of Eurasian individuals, we find that PC1 separates eastern and western Eurasian populations, PC2 splits eastern Eurasians along a north-south cline, and PC3 captures variation in western Eurasians with Caucasus and northeastern European populations at opposite ends (Figure 2A and Figures S1-S2). Inner Eurasians are scattered across PC1 in between, largely reflecting their geographic locations. Strikingly, inner Eurasian populations seem to be structured into three distinct west-east genetic clines running between different western and eastern Eurasian groups, instead of being evenly spaced in PC space. Individuals from northern Eurasia, speaking Uralic or Yeniseian languages, form a cline connecting northeast Europeans and the Uralic (Samoyedic) speaking Nganasans from northern Siberia (“forest-tundra” cline). Individuals from the Eurasian steppe, mostly speaking Turkic and Mongolic languages, are scattered along two clines below the forest-tundra cline. Both clines run into Turkic- and Mongolic-speaking populations in southern Siberia and Mongolia, and further into Tungusic-speaking populations in Manchuria and the Russian Far East in the East; however, they diverge in the west, oneheading to the Caucasus and the other heading to populations of the Volga-308 Ural area (the “southern steppe” and “steppe-forest” clines, respectively; Figure 2 and Figure S2).
(…) The forest-tundra cline populations derive most of their eastern Eurasian ancestry from a component most enriched in Nganasans, while those on the steppe-forest and southern steppe clines have this component together with another component most enriched in populations from the Russian Far East, such as Ulchi and Nivkh. The southern steppe cline groups are distinct from the others in their western Eurasian ancestry profile, in the sense that they have a high proportion of a component most enriched in Mesolithic Caucasus hunter-gatherers (“CHG”) and Neolithic Iranians (“Iran_N”) and frequently harbor another component enriched in South Asians (Figure S4).
For the forest-tundra cline populations, for which currently no relevant Holocene ancient genomes are available, we took a more generalized approach of using proxies for contemporary Europeans: WHG, WSH (represented by “Yamnaya_Samara”), and early Neolithic European farmers (EEF; represented by “LBK_EN”; Table S2). Adding Nganasans as the fourth reference, we find that most Uralic-speaking populations in Europe (i.e. west of the Urals) and Russians are well modeled by this four-way admixture model (χ 2 p ≥ 0.05 for all but three groups; Figure 5 and Table S8). Nganasan-related ancestry substantially contributes to their gene pools and cannot be removed from the model without a significant decrease in model fit (4.7% to 29.1% contribution; χ 2 p ≤ 1.12×10-8; Table S8). The ratio of contributions from three European references varies from group to group, probably reflecting genetic exchange with neighboring non-Uralic groups. For example, Saami from northern Fennoscandia contain a higher WHG and lower WSH contribution (16.1% and 41.3%, respectively) than Udmurts or Besermyans from the Volga river region do (4.9-6.6% and 50.7-53.2%, respectively), while the three groups have similar amounts of Nganasan-related ancestry (25.5-29.1%).
The Caucasus Mountains form a barrier to gene flow
By applying EEMS to the Caucasus region, we identify a strong barrier to gene flow separating North and South Caucasus populations. This genetic barrier coincides with the Greater Caucasus mountain ridge even to small scale: a weaker barrier in the middle, overlapping with Ossetia, matches well with the region where the ridge also becomes narrow. We also observe weak barriers running in the north-south direction that separate northeastern populations from northwestern ones. Together with PCA, EEMS results suggest that the Caucasus Mountains have posed a strong barrier to human migration.
On the Botai individuals
The Y-chromosome of the male Botai individual (TU45) belongs to the haplogroup R1b (Table 411 S6). However, it falls into neither a predominant European branch R1b-L5165 nor into a R1b-GG400 branch found in Yamnaya individuals. Thus, phylogenetically this Botai individual should belong to the R1b-M73 branch which is frequent in the Eurasian steppe (Figure S9). This branch was also found in Mesolithic samples from Latvia as well as in numerous modern southern Siberian and Central Asian groups.
The Botai genomes provide a critical snapshot of the genetic profile of pre-Bronze Age steppe populations. Our admixture modeling positions Botai primarily on an ancient genetic cline of the pre-Neolithic western Eurasian hunter-gatherers: stretching from the post-Ice Age western European hunter-gatherers (e.g. WHG) to EHG in Karelia and Samara to the Upper Paleolithic southern Siberians (e.g. AG3). Botai’s position on this cline, between EHG and AG3, fits well with their geographic location and suggests that ANE-related ancestry in the East did have a lingering genetic impact on Holocene Siberian and Central Asian populations at least till the time of Botai.
The most recent clear connection with the Botai ancestry can be found in the Middle Bronze Age Okunevo individuals (Figure S6C). In contrast, additional EHG-related ancestry is required to explain the forest-tundra populations to the east of the Urals (Figure 5 and Table S8). Their multi-way mixture model may in fact portrait a prehistoric two-way mixture of a WSH population and a hypothetical eastern Eurasian one that has an ANE-related contribution higher than that in Nganasans. Botai and Okunevo individuals prove the existence of such ANE ancestry-rich populations. Pre-Bronze Age genomes from Siberia will be critical for testing this hypothesis.
So, to sum up:
Northern Eurasia forms a Uralic – Yeniseian cline from east to west, with contribution from Steppe, WHG, and Siberian ancestry. Siberian ancestry is represented by Palaeo-Siberian Nganasans, who adopted Samoyedic quite late. It was already known that the different waves of Siberian ancestry are too late and do not represent the spread of Uralic languages, so that leaves us with Steppe and WHG.
The Botai sample (ca. 3632-3100 BC) represents thus the furthest east that R1b-P297 subclades had expanded (we did know that, and that they didn’t have close genetic links with Khvalynsk, so the haplogroup spread there probably much earlier). It expanded R1b-M269’s sister clade R1b-M73 (also found in the Baltic region), and the Botai are on the ‘eastern’ end of an ancient genetic cline stretching from WHG to EHG to Afontova Gora.
EDIT (23 MAY 2018) Both samples share mtDNA, and the male one shares Y-DNA, with those reported in Damgaard et al. (Nature 2018); although dates are slightly different (3371-3354 calBC for BOT 14), it is within the range given for this one; for the female, the dates are similar (3521-3377 calBC for BOT2016, 3517-3367 cal. BCE for this one). The lack of data on their origin may point to the fact that we only have different bone samples from the same two Botai individuals. So probably still 50% R1b-M73 (with the other 50% being N2* from BOT15)…
It seems therefore not only that R1b-M269 is bound to split from the parent haplogroup in or around the steppe or forest-steppe: the Mesolithic spread of haplogroup R1b in North Eurasia is wider and its relevance thus greater than previously thought.
Featured image, from the supplementary materials: Frequency distribution map of the Y-chromosomal haplogroup R1b-P343(xM269) identified in the Eneolithic Botai individual. All modern Eurasian samples with this haplogroup tested to date for the downstream markers fall into R1b-M73 branch, suggesting Botai sample be one of its earliest representatives.