Baltic Finns in the Bronze Age, of hg. R1a-Z283 and Corded Ware ancestry

estonian-bronze-age-dna

Open access The Arrival of Siberian Ancestry Connecting the Eastern Baltic to Uralic Speakers further East, by Saag et al. Current Biology (2019).

Interesting excerpts:

In this study, we present new genomic data from Estonian Late Bronze Age stone-cist graves (1200–400 BC) (EstBA) and Pre-Roman Iron Age tarand cemeteries (800/500 BC–50 AD) (EstIA). The cultural background of stone-cist graves indicates strong connections both to the west and the east [20, 21]. The Iron Age (IA) tarands have been proposed to mirror “houses of the dead” found among Uralic peoples of the Volga-Kama region [22].

(…) The 33 individuals included 15 from EstBA, 6 from EstIA, 5 from Pre-Roman to Roman Iron Age Ingria (500 BC–450 AD) (IngIA), and 7 from Middle Age Estonia (1200–1600 AD) (EstMA) and yielded endogenous DNA ∼4%–88%, average genomic coverages ∼0.017–0.734×, and contamination estimates <4% (Table S1). We analyzed the data in the context of modern and other ancient individuals, including from Neolithic Estonia [13].

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Archaeological Information, Genetic Sex, mtDNA and Y Chromosome Haplogroups, and Average Coverage of the Individuals of This Study. Modified from the paper to mark distinct Y-DNA haplogroups in the LBA and IA.

We identified chrY hgs for 30 male individuals (Tables 1 and S2; STAR Methods). All 16 successfully haplogrouped EstBA males belonged to hg R1a, showing no change from the CWC period, when this was also the only chrY lineage detected in the Eastern Baltic [11, 13, 30, 31]. Three EstIA and two IngIA individuals also belonged to hg R1a, but three EstIA males belonged to hg N3a, the earliest so far observed in the Eastern Baltic. Three EstMA individuals belonged to hg N3a, two to hg R1a, and one to hg J2b. ChrY lineages found in the Baltic Sea region before the CWC belong to hgs I, R1b, R1a5, and Q [10, 11, 12, 13, 17, 32]. Thus, it appears that these lineages were substantially replaced in the Eastern Baltic by hg R1a [10, 11, 12, 13], most likely through steppe migrations from the east [30, 31]. (…) Our results enable us to conclude that, although the expansion time for R1a1 and N3a3′5 in Eastern Europe is similar [25], hg N3a likely reached Estonia or at least became comparably frequent to modern Estonia [1] only during the BA-IA transition.

A clear shift toward West Eurasian hunter-gatherers is visible between European LN and BA (including Baltic CWC) and EstBA individuals, the latter clustering together with Latvian and Lithuanian BA individuals [11]. EstIA, IngIA, and EstMA individuals project between BA individuals and modern Estonians, partially overlapping with both.

(…) EstBA individuals are clearly distinguishable from Estonian CWC individuals as the former have more of the blue component most frequent in WHGs and less of the brown and yellow components maximized in Caucasus hunter-gatherers and modern Khanty, respectively. The individuals of EstBA, EstIA, IngIA, EstMA, and modern Estonia are quite similar to each other on average, indicating that the relatively high proportion of WHG ancestry in modern Eastern Baltic populations compared to other present-day Europeans [15] traces back to the BA.

estonian-pca-published
Detail of the PCA, modified from the paper to label populations. Estonian Bronze Age and Iron Age samples cluster close to Early Corded Ware from the Baltic.. Principal-component analysis results of modern West Eurasians with ancient individuals projected onto the first two components (PC1 and PC2). BA, Bronze Age; EF, early farmers; HG, hunter-gatherers; IA, Iron Age; IMA, Iron/Middle Ages; LN, Late Neolithic; LNBA, Late Neolithic/Bronze Age; MA, Middle Ages

When comparing Estonian CWC and EstBA using autosomal outgroup f3 and Patterson’s D statistics (Table S3), the latter is more similar to other Baltic BA populations, to Baltic IA and Middle Age (MA) populations, and also to populations similar to WHGs and Scandinavian hunter-gatherers (SHGs), but not to Estonian CCC (Figures 2A and S2A; Data S1). The increase in WHG or SHG ancestry could be connected to western influences seen in material culture [20, 21] and facilitated by a decline in local population after the CCC-CWC period [20]. A slight trend of bigger similarity of Estonian CWC to forest or steppe zone populations and of EstBA to European early farmer populations can also be seen.

(…) When comparing to modern populations, Estonian CWC is slightly more similar to Caucasus individuals but EstBA to Baltic populations and Finnic speakers (Figure 2B; Data S1). Outgroup f3 and D statistics do not reveal apparent differences when comparing EstBA to EstIA, EstIA to IngIA, and EstIA to EstMA (Data S1).

estonian-ba-ia-ancestry
qpAdm results. Error bars indicate one SE. Central MN, Central European Middle Neolithic; EstBA, Estonian Bronze Age; EstIA, Estonian Iron Age; IngIA, Ingrian Iron Age; EstMA, Estonian Middle Ages; WHG, western hunter-gatherers.

These results highlight how uniparental and autosomal data can lead to different demographic inferences—the genetic change between CWC and BA not seen in uniparental lineages is clear in autosomal data and the appearance of chrY hg N in the IA is not matched by a clear shift in autosomal profiles.

EstBA individuals have no Nganasan-related ancestry and EstIA, IngIA, and EstMA individuals on average have 2% or 4% (Figure 3; Data S1). The differentiation remains when using BA or IA Fennoscandian populations [26] instead of Nganasans (Data S1). Notably, the proportion of Nganasan-related ancestry varies between 0% and 12% among sampled EstIA, IngIA, and EstMA individuals (Data S1), which may suggest its relatively recent admixture into the target population. Moreover, two individuals from Kunda (0LS10 and V10) have the highest proportions of Nganasan ancestry among EstIA (6% and 8%), one of them has chrY hg N3a, and isotopic analysis suggests neither individual being born in Kunda [34].

About these two males from Tarand-graves, ‘foreign’ to Kunda:

0LS10: Male from tarand III (burial 9; TÜ 1325: L777), age 17–25 years [34]. He had a fragment of a sheep/goat bone and ceramics as grave goods. This burial has two radiocarbon dates: 2430 ± 35 BP (Poz-10801; 760–400 cal BC) and 2530 ± 41 BP (UBA-26114; 800–530 cal BC) [34]. According to the isotopic analysis, the person was not born in the vicinity of Kunda; his place of birth is still unknown (but south-western Finland and Sweden are excluded) [34]. Sampled tooth r P1.

V10: Male from tarand XI (burial 24; TÜ 1325: L1925), age 25–35 years [34], date 2484 ± 40 BP (UBA-26115; 790–430 cal BC) [34]. He had a few potsherds near the skull. Likewise, this person was not locally born [34]. Sampled tooth l P1.

estonia-bronze-iron-age-steppe-siberian
Autosomal Analyses’ Results for Gyvakarai1 as the closest available Corded Ware source for Balto-Finnic populations.

The paper shows thus:

  • Major continuity of ancestry from Corded Ware to modern Estonians, with only slight changes in different periods. In fact, one of the best fits for the Late Bronze Age ancestry is Gyvakarai1, one of the Corded Ware “outliers” described as “closer to Yamna”, which I already said may be closer to Sredni Stog/EHG populations instead. Another interesting take is that the change from Bronze Age to Iron Age corresponds to an increase in Baltic Corded Ware-related ancestry, rather than being driven by Siberian ancestry.
  • pca-mittnik-gyvakarai
    File modified by me from Mittnik et al. (2018) to include the approximate position of the most common ancestral components, and an identification of potential outliers. Zoomed-in version of the European Late Neolithic and Bronze Age samples. “Principal components analysis of 1012 present-day West Eurasians (grey points, modern Baltic populations in dark grey) with 294 projected published ancient and 38 ancient North European samples introduced in this study (marked with a red outline). From Mittnik et al. (2018).
  • A Volosovo-related migration of hg. N1c with Netted Ware into the area seems to be discarded, based on the full replacement of paternal lines and continuity of R1a-Z283. It is only during the Tarand-grave period when a system of chiefdoms (spread from Ananyino/Akozino) brings haplogroup N1c to the Gulf of Finland. During the Iron Age, the proportion of paternal lineages is still clearly in favour of R1a (50% in the coast, 100% in Ostrobothnia), which indicates a gradual replacement led by elites, likely because of the incorporation of Akozino warrior-traders spreading all over the Baltic, bringing the described shared Mordvinic traits in Fennic.
  • finno-ugric-haplogroup-n
    Map of archaeological cultures in north-eastern Europe ca. 8th-3rd centuries BC. [The Mid-Volga Akozino group not depicted] Shaded area represents the Ananino cultural-historical society. Fading purple arrows represent likely stepped movements of subclades of haplogroup N for centuries (e.g. Siberian → Ananino → Akozino → Fennoscandia [N-VL29]; Circum-Arctic → forest-steppe [N1, N2]; etc.). Blue arrows represent eventual expansions of Uralic peoples to the north. Modified image from Vasilyev (2002).
  • The arrival of Akozino warrior-traders (bringing N1c and R1a lineages) was probably linked to this minimal “Nganasan-like” ancestry of some samples in the transition to the Iron Age. This arrival is supported by samples 0LS10 (the earliest hg. N1c) and V10 (of hg. R1a), both dated to ca. 800-400 BC, with V10 showing the highest “Nganasan-like” ancestry with 4.8%, both of them neighbouring samples showing 0%. This variable admixture among local and foreign paternal lineages might support the described social system of family alliances with intermarriages. In fact, a medieval sample, 0LS03_1 (hg. R1a) also shows a recent “Nganasan-like” ancestry, which probably points to the integration of different Arctic-related ancestry components among Modern Estonians, in this case related to Finnish expansions and thus integration of Levänluhta-related ancestry, as per the supplementary data.
  • NOTE. Such minimal proportions of “Nganasan-like” ancestry evidence the process of admixture of Volga Finns in Akozino territory through their close interactions with Permians of Ananyino, who in turn acquired this Palaeo-Arctic admixture most likely during the expansion of the linguistic community to hunter-gatherer territories, to the north of the Cis-Urals. This process of stepped infiltration and expansion without language change is not dissimilar to the one seen among Indo-Iranians and Balto-Slavs of hg. R1b, or Vasconic speakers of hg. I2a, although in the case of Baltic Finns of hg. R1a the process of infiltration and expansion of hg. N1c is much less dramatic, with no radical replacement anywhere before the huge bottlenecks observable in Finns.

  • The expansion of haplogroup N1c among Finnic populations, as we are going to see in samples from the Middle Ages such as Luistari, is the consequence of late founder effects after huge bottlenecks expected based on the analysis of modern populations. The expansion of N1c-VL29 is different in origin from that of N1c-Z1936 among Samic (later integrated into Finnish populations), most likely from the east and originally associated with Lovozero Ware.
haplogroup_n3a3
Frequency-Distribution Maps of Individual Subclade N3a3 / N1a1a1a1a1a-CTS2929/VL29, probably initially with Akozino warrior-traders. Map from Ilumäe et al. (2016).

In spite of all this, the conclusion of the paper is (surprise!) that Siberian ancestry and hg. N heralded the arrival of Finnic to the Gulf of Finland in the Iron Age… However, this conclusion is supposedly* supported, not by their previous papers, but by a recent phylogenetic study by Honkola et al. (2013), which doesn’t actually argue for such a late ‘arrival’: it argues for the split of Balto-Finnic around 1500 BC.

NOTE. I say ‘supposedly’ because Kristiina Tambets, for example, has been following the link of Uralic with haplogroup N since the 2000s, so this is not some conclusion they just happened to misread from some random paper they Googled. In those initial assessments, she argued that the “ancient homeland” of the Tat C mutation suggested that Finno-Ugrians were in Fennoscandia before Indo-Europeans. Apparently, since haplogroup N appears later and from the east, it is now more important to follow this haplogroup than what is established in archaeology and linguistics.

Even in the referred paper, this split is considered an in situ development, since the phylogenetic study takes the information – among others – 1) from Parpola and Carpelan, who consider Netted Ware, a culture derived from Fatyanovo/Abashevo and Volosovo, as the culprit of the Finno-Ugric expansion; and 2) from Kallio (2006), who clearly states that Proto-Balto-Finnic (like Proto-Finno-Samic) was spoken around the Gulf of Finland during the Bronze Age. Both of them set the terminus ante quem of the language presence in the Baltic ca. 1900 BC.

Anyways, as a consequence of geneticists keeping these untenable pre-ancient DNA haplogroup-based arguments today, I expect to see this “Finnic” language expansion also described for the Western Baltic, Scandinavia or northern Europe, when this same proportion of hg. N1c and “Nganasan” ancestry is observed in Iron Age samples around the Baltic Sea. The nativist trends that this domination of “Finns” all over Northern Europe 2,500 years ago will create will be even more fun to read than the current ones…

EDIT (10 May 2019) How I see the reaction of many to ancient DNA, in keeping their old theories:

Related

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

koszyce-gac-zlota-cwc

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Related

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

steppe-forest-tundra-biomes-uralic

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

Interesting excerpts, referring mainly to Uralic peoples (emphasis mine):

A model-based clustering analysis using ADMIXTURE shows a similar pattern (Fig. 2b and Supplementary Fig. 3). Overall, the proportions of ancestry components associated with Eastern or Western Eurasians are well correlated with longitude in inner Eurasians (Fig. 3). Notable outliers include known historical migrants such as Kalmyks, Nogais and Dungans. The Uralic- and Yeniseian-speaking populations, as well as Russians from multiple locations, derive most of their Eastern Eurasian ancestry from a component most enriched in Nganasans, while Turkic/Mongolic speakers have this component together with another component most enriched in populations from the Russian Far East, such as Ulchi and Nivkh (Supplementary Fig. 3). Turkic/Mongolic speakers comprising the bottom-most cline have a distinct Western Eurasian ancestry profile: they have a high proportion of a component most enriched in Mesolithic Caucasus hunter-gatherers and Neolithic Iranians and frequently harbour another component enriched in present-day South Asians (Supplementary Fig. 4). Based on the PCA and ADMIXTURE results, we heuristically assigned inner Eurasians to three clines: the ‘forest-tundra’ cline includes Russians and all Uralic and Yeniseian speakers; the ‘steppe-forest’ cline includes Turkic- and Mongolic-speaking populations from the Volga and Altai–Sayan regions and Southern Siberia; and the ‘southern steppe’ cline includes the rest of the populations.

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

For the forest-tundra populations, the Nganasan + Srubnaya model is adequate only for the two Volga region populations, Udmurts and Besermyans (Fig. 5 and Supplementary Table 8).

For the other populations west of the Urals, six from the northeastern corner of Europe are modelled with additional Mesolithic Western European hunter-gatherer (WHG) contribution (8.2–11.4%; Supplementary Table 8), while the rest need both WHG and early Neolithic European farmers (LBK_EN; Supplementary Table 2). Nganasan-related ancestry substantially contributes to their gene pools and cannot be removed from the model without a significant decrease in the model fit (4.1–29.0% contribution; χ2 P ≤ 1.68 × 10−5; Supplementary Table 8).

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

NOTE. It doesn’t seem like Hungarians can be easily modelled with Nganasan ancestry, though…

For the 4 populations east of the Urals (Enets, Selkups, Kets and Mansi), for which the above models are not adequate, Nganasan + Srubnaya + AG3 provides a good fit (χ2 P ≥ 0.018; Fig. 5 and Supplementary Table 8). Using early Bronze Age populations from the Baikal Lake region (‘Baikal_EBA’; Supplementary Table 2) as a reference instead of Nganasan, the two-way model of Baikal_EBA + Srubnaya provides a reasonable fit (χ2 P ≥ 0.016; Supplementary Table 8) and the three-way model of Baikal_EBA + Srubnaya + AG3 is adequate but with negative AG3 contribution for Enets and Mansi (χ2 P ≥ 0.460; Supplementary Table 8).

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

Bronze/Iron Age populations from Southern Siberia also show a similar ancestry composition with high ANE affinity (Supplementary Table 9). The additional ANE contribution beyond the Nganasan + Srubnaya model suggests a legacy from ANE-ancestry-rich clines before the Late Bronze Age.

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

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

Even among the earliest available inner Eurasian genomes, east–west connectivity is evident. These, too, form a longitudinal cline, characterized by the easterly increase of a distinct ancestry, labelled Ancient North Eurasian (ANE), lowest in western European hunter-gatherers (WHG) and highest in Palaeolithic Siberians from the Baikal region. Flow-through from this ANE cline is seen in steppe populations until at least the Bronze Age, including the world’s earliest known horse herders — the Botai. However, this is eroded over time by migration from west and east, following agricultural adoption on the continental peripheries (Fig. 1b,c).

Strikingly, Jeong et al. model the modern upper steppe cline as a simple two-way mixture between western Late Bronze Age herders and Northeast Asians (Fig. 1c), with no detectable residue from the older ANE cline. They propose modern steppe peoples were established mainly through migrations post-dating the Bronze Age, a sequence for which has been recently outlined using ancient genomes. In contrast, they confirm a substantial ANE legacy in modern Siberians of the northernmost cline, a pattern mirrored in excesses of WHG ancestry west of the Urals (Fig. 1b). This marks the inhospitable biome as a reservoir for older lineages, an indication that longstanding barriers to latitudinal movement may indeed be at work, reducing the penetrance of gene flows further south along the steppe.

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

Given the findings as reported in the paper, I think it should be much easier to describe different subclines in the “northernmost cline” than in the much more recent “Turkic/Mongolic cline”, which is nevertheless subdivided in this paper in two clines. As an example, there are at least two obvious clines with “Nganasan-related meta-populations” among Uralians, which converge in a common Steppe MLBA (i.e. Corded Ware) ancestry – one with Palaeo-Laplandic peoples, and another one with different Palaeo-Siberian populations:

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

The inclusion of certain Eurasian groups (or lack thereof) in the PCA doesn’t help to distinguish these subclines visually, and I guess the tiny “Naganasan-related” ancestral components found in some western populations (e.g. the famous ~5% among Estonians) probably don’t lend themselves easily to further subdivisions. Notice, nevertheless, the different components of the Eastern Eurasian source populations among Finno-Ugrians:

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

Also remarkable is the lack of comparison of Uralic populations with other neighbouring ones, since the described Uralic-like ancestry of Russians was already known, and is most likely due to the recent acculturation of Uralic-speaking peoples in the cradle of Russians, right before their eastward expansions.

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

A comparison of Estonians and Finns with Balts, Scandinavians, and Eastern Europeans would have been more informative for the division of the different so-called “Nganasan-like meta-populations”, and to ascertain which one of these ancestral peoples along the ancient WHG:ANE cline could actually be connected (if at all) to the Cis-Urals.

Because, after all, based on linguistics and archaeology, geneticists are not supposed to be looking for populations from the North Asian Arctic region, for “Siberian ancestry”, or for haplogroup N1c – despite previous works by their peers – , but for the Bronze Age Volga-Kama region…

Related

Fulani from Cameroon show ancestry similar to Afroasiatic speakers from East Africa

sahel-region-fulani

Open access African evolutionary history inferred from whole genome sequence data of 44 indigenous African populations, by Fan et al. Genome Biology (2019) 20:82.

Interesting excerpts (emphasis mine):

Introduction

To extend our knowledge of patterns of genomic diversity in Africa, we generated high coverage (> 30×) genome sequencing data from 43 geographically diverse Africans originating from 22 ethnic groups, representing a broad array of ethnic, linguistic, cultural, and geographic diversity (Additional file 1: Table S1). These include a number of populations of anthropological interest that have never previously been characterized for high-coverage genome sequence diversity such as Afroasiatic-speaking El Molo fishermen and Nilo-Saharan-speaking Ogiek hunter-gatherers (Kenya); Afroasiatic-speaking Aari, Agaw, and Amhara agro-pastoralists (Ethiopia); Niger-Congo-speaking Fulani pastoralists (Cameroon); Nilo-Saharan-speaking Kaba (Central African Republic, CAR); and Laka and Bulala (Chad) among others. We integrated this data with 49 whole genome sequences generated as part of the Simons Genome Diversity Project (SGDP) [14] (…)

afroasiatic-samples
Locations of samples included in this study. Each dot is an individual and the color indicates the language classification

Results and discussion

We found that the CRHG populations from central Africa, including the Mbuti from the Demographic Republic of Congo (DRC), Biaka from the CAR, and Baka, Bakola, and Bedzan from Cameroon, also form a basal lineage in the phylogeny. The other two hunter-gatherer populations, Hadza and Sandawe, living in Tanzania, group with populations from eastern Africa (Fig. 2). The two Nilo-Saharan-speaking populations, the Mursi from southern Ethiopia and the Dinka from southern Sudan, group into a single cluster, which is consistent with archeological data indicating that the migration of Nilo-Saharan populations to eastern Africa originated from a source population in southern Sudan in the last 3000 years [4, 23, 24, 25].

phylogenetic-relationship-africans
Phylogenetic relationship of 44 African and 32 west Eurasian populations determined by a neighbor joining analysis assuming no admixture. Here, the dots of each node represent bootstrap values and the color of each branch indicates language usage of each population. Human_AA human ancestral alleles

The Fulani people are traditionally nomadic pastoralists living across a broad geographic range spanning Sudan, the Sahel, Central, and Western Africa. The Fulani in our study, sampled from Cameroon, clustered with the Afroasiatic-speaking populations in East Africa in the phylogenetic analysis, indicating a potential language replacement from Afroasiatic to Niger-Congo in this population (Fig. 2). Prior studies suggest a complex history of the Fulani; analyses of Y chromosome variation suggest a shared ancestry with Nilo-Saharan and Afroasiatic populations [24], whereas mtDNA indicates a West African origin [26]. An analysis based on autosomal markers found traces of West Eurasian-related ancestry in this population [4], which suggests a North African or East African origin (as North and East Africans also have such ancestry likely related to expansions of farmers and herders from the Near East) and is consistent with the presence at moderate frequency of the −13,910T variant associated with lactose tolerance in European populations [15, 16].

Phylogenetic reconstruction of the relationship of African individuals under a model allowing for migration using TREEMIX [27] largely recapitulates the NJ phylogeny with the exception of the Fulani who cluster near neighboring Niger-Congo-speaking populations with whom they have admixed (Additional file 2: Figure S1). Interestingly, TREEMIX analysis indicates evidence for gene flow between the Hadza and the ancestors of the Ju|‘hoan and Khomani San, supporting genetic, linguistic, and archeological evidence that Khoesan-speaking populations may have originated in Eastern Africa [28, 29, 30].

afroasiatic-niger-congo-admixture
ADMIXTURE analysis of 92 African and 62 West Eurasian individuals. Each bar is an individual and colors represent the proportion of inferred ancestry from K ancestral populations. The bottom bar shows the language classification of each individual. With the increasing of K, the populations are largely grouped by their current language usage

About the Fulani, this is what the referenced study of Y‐chromosome variation among 15 Sudanese populations by Hassan et al. (2008), had to say:

  • Haplogroups A-M13 and B-M60 are present at high frequencies in Nilo-Saharan groups except Nubians, with low frequencies in Afro-Asiatic groups although notable frequencies of B-M60 were found in Hausa (15.6%) and Copts (15.2%).
  • Haplogroup E (four different haplotypes) accounts for the majority (34.4%) of the chromosome and is widespread in the Sudan. E-M78 represents 74.5% of haplogroup E, the highest frequencies observed in Masalit and Fur populations. E-M33 (5.2%) is largely confined to Fulani and Hausa, whereas E-M2 is restricted to Hausa. E-M215 was found to occur more in Nilo-Saharan rather than Afro-Asiatic speaking groups.
  • In contrast, haplogroups F-M89, I-M170, J-12f2, and JM172 were found to be more frequent in the Afro-Asiatic speaking groups. J-12f2 and J-M172 represents 94% and 6%, respectively, of haplogroup J with high frequencies among Nubians, Copts, and Arabs.
  • Haplogroup K-M9 is restricted to Hausa and Gaalien with low frequencies and is absent in Nilo-Saharan and Niger-Congo.
  • Haplogroup R-M173 appears to be the most frequent haplogroup in Fulani, and haplogroup R-P25 has the highest frequency in Hausa and Copts and is present at lower frequencies in north, east, and western Sudan.
  • Haplogroups A-M51, A-M23, D-M174, H-M52, L-M11, OM175, and P-M74 were completely absent from the populations analyzed.
fulfulde-fulani-language
Image modified from “Fulfulde Language Family Report” Author: Annette Harrison; Cartographer: Irene Tucker; SIL International 2003.

This is what David Reich will talk about in the seminar Insights into language expansions from ancient DNA:

In this talk, I will describe how the new science of genome-wide ancient DNA can provide insights into past spreads of language and culture. I will discuss five examples: (1) the spread of Indo-European languages to Europe and South Asia in association with Steppe pastoralist ancestry, (2) the spread of Austronesian languages to the open Pacific islands in association with Taiwanese aboriginal-associated ancestry, (3) the spread of Austroasiatic languages through southeast Asia in association with the characteristic ancestry type that is also represented in western Indonesia suggesting that these languages were once widespread there, (4) the spread of Afroasiastic languages through in East Africa as part of the Pastoral Neolithic farming expansion, and (5) the spread of Na-Dene languages in North America in association with Proto-Paleoeskimo ancestry. I will highlight the ways that ancient DNA can meaningfully contribute to our understanding of language expansions—increasing the plausibility of some scenarios while decreasing the plausibility of others—while emphasizing that with genetic data by itself we can never definitively determine what languages ancient people spoke.

EDIT (3 MAY 2019): Apparently, there was not much to take from the talk:

neolithic-pastoralist-africa
Pastoralist Neolithic in Africa, through a pale-green Sahelo-Sudanian steppe corridor. See full map.

This seminar (and maybe some new paper on the Neolithic expansion in Africa) could shed light on population movements that may be related to the spread of Afroasiatic dialects. Until now, it seems that Bantu peoples have been more interesting for linguistics and archaeology, and South and East Africans for anthropology.

Archaeology in Africa appears to be in its infancy, as is population genomics. From the latest publication by Carina Schlebusch, Population migration and adaptation during the African Holocene: A genetic perspective, a chapter from Modern Human Origins and Dispersal (2019):

The process behind the introduction and development of farming in Africa is still unclear. It is not known how many independent invention events there were in the continent and to which extent the various first instances of farming in northern Africa are linked. Based on the archeological record, it was proposed that at least three regions in Africa may have developed agriculture independently: the Sahara/Sahel (around 7 ka), the Ethiopian highlands (7-4 ka), and western Africa (5-3 ka). In addition to these developments, the Nile River Valley is thought to have adopted agriculture (around 7.2 ka), from the Neolithic Revolution in the Middle East (Chapter 12 – Jobling et al. 2014; Chapter 35, 37 – Mitchell and Lane 2013). From these diverse centers of origin, farmers or farming practices spread to the rest of Africa, with domesticate animals reaching the southern tip of Africa ~2 ka and crop farming ~1,8 ka (Mitchell 2002; Huffman 2007)

african-popularion-movements
Schematic representation of possible migration routes related to the expansion of herders and crop farmers during Holocene times. Arrow color indicate source populations; Brown-Eurasian, Green-western African, Blue-eastern African.

Similar to the case in Europe and the 1990s-2000s wrong haplogroup history based on the modern distribution of R1b, R1a, N, or I2, it is possible that neither of the most often mentioned haplogroups linked to the Afroasiatic expansion, E and J, were responsible for its early spread within Africa, despite their widespread distribution in certain modern Afroasiatic-speaking areas. The fact that such assessments include implausible glottochronological dates spanning up to 20,000 years for the parent language, combined with regional language continuities despite archaeological changes, makes them even more suspicious.

Similar to the case with Indo-Europeans and the “steppe ancestry” concept of the 2010s, it may be that the often-looked-for West Eurasian ancestry among Africans is the effect of recent migrations, unrelated to the Afroasiatic expansion. The results of this paper could be offering another sign of how this ancestry may have expanded only quite recently westwards from East Africa through the Sahel, after the Semitic expansion to the south:

1. From approximately 1000 BC, accompanying Nilo-Saharan peoples.

2. From approximately AD 1500, with the different population movements related to the nomadic Fulani:

sahel-nomadic-sedentary
Image from Sahel in West African History – Oxford Research Encyclopedia of African History.
  • Arguably, since the Fulani caste system wasn’t as elaborate in northern Nigeria, eastern Niger, and Cameroon, these specific groups would be a good example of the admixture with eastern populations, based on the (proportionally) huge amount of slaves they dealt with.
  • Similarly, it could be argued that the castes-based social stratification in most other territories (including Sudan) would have helped them keep a genetic make-up similar to their region of origin in terms of ancient lineages, hence similar to Chadic populations from west to east.

Reich’s assertion of the association of the language expansion with the spread of Pastoral Neolithic is still too vague, but – based on previous publications of ancient DNA in Africa and the Levant – I don’t have high hopes for a revolutionary paper in the near future. Without many samples and proper temporal transects, we are stuck with speculations based on modern distributions and scarce historical data.

fula-people-distribution
A distribution map of Fula people. Dark green: a major ethnic group; Medium: significant; Light: minor. Modified from image by Sarah Welch at Wikipedia.

About the potential genetic make-up of Cameroon before the arrival of the Neolithic, from the recent SAA 84th Annual Meeting (Abstracts in PDF):

Lipson, Mark (Harvard Medical School), Mary Prendergast (Harvard University), Isabelle Ribot (Université de Montréal), Carles Lalueza-Fox (Institute of Evolutionary Biology CSIC-UPF) and David Reich (Harvard Medical School)

[253] Ancient Human DNA from Shum Laka (Cameroon) in the Context of African Population History We generated genome-wide DNA data from four people buried at the site of Shum Laka in Cameroon between 8000–3000 years ago. One individual carried the deeply divergent Y chromosome haplogroup A00 found at low frequencies among some present-day Niger-Congo speakers, but the genome-wide ancestry profiles for all four individuals are very different from the majority of West Africans today and instead are more similar to West-Central African hunter-gatherers. Thus, despite the geographic proximity of Shum Laka to the hypothesized birthplace of Bantu languages and the temporal range of our samples bookending the initial Bantu expansion, these individuals are not representative of a Bantu source population. We present a phylogenetic model including Shum Laka that features three major radiations within Africa: one phase early in the history of modern humans, one close to the time of the migration giving rise to non-Africans, and one in the past several thousand years. Present-day West Africans and some East Africans, in addition to Central and Southern African hunter-gatherers, retain ancestry from the first phase, which is therefore still represented throughout the majority of human diversity in Africa today.

Related

Pre-Germanic and Pre-Balto-Finnic shared vocabulary from Pitted Ware seal hunters

corded-ware-pitted-ware

I said I would write a post about topo-hydronymy in Europe and Iberia based on the most recent research, but it seems we can still enjoy some more discussions about the famous Vasconic Beakers, by people longing for days of yore. I don’t want to spoil that fun with actual linguistic data (which I already summarized) so let’s review in the meantime one of the main Uralic-Indo-European interaction zones: Scandinavia.

Seal hunting

One of the many eye-catching interpretations – and one of the few interesting ones – that could be found in the relatively recent article Talking Neolithic: Linguistic and Archaeological Perspectives on How Indo-European Was Implemented in Southern Scandinavia, by Iversen & Kroonen AJA (2017) was this:

The borrowing of lexical items from hunter-gatherers into Germanic refers to the potential adoption of Proto-Germanic *selhaz “seal” (Old Norse selr, Old English seolh, Old High German selah) as well as Early Proto-Balto-Finnic *šülkeš “seal” (Finnish hylje, Estonian hüljes) from the marine-oriented Sub-Neolithic Pitted Ware culture.

kroonen-iversen
Modified from Kristiansen et al. (2017), with red circle around the hypothesized interaction of Germanic with hunter-gatherers. “Schematic representation of how different Indo-European branches have absorbed words (circles) from a lost Neolithic language or language group (dark fill) in the reconstructed European linguistic setting of the third millennium BC, possibly involving one or more hunter gatherer languages (light fill) (after Kroonen & Iversen 2017)”.

This is what Kroonen thought about this word in his Etymological Dictionary of Proto-Germanic (2006):

Gmc. *selha– m. ‘seal’ – ON selr m. ‘id.’, Far. selur m. ‘id.’, OSw. siæl m. ‘id.’, Sw. själ c. ‘id.’, OE seolh m. ‘id.’, E seal, OS selah m. ‘id.’, EDu. seel, seel-hont m. ‘id.’, Du. zee-hond c. ‘id.’, OHG selah m. ‘id.’, MHG sele m. ‘id.’ (GM).

A Germanic word with no certain IE etymology. The link with Lith. selė́ti ‘to crawl’ (Torp 1909: 436) is erroneous, as this verb corresponds to PGm. *stelan- (q.v.). The *h may nevertheless correspond to the PIE animal suffix *-ko-, for which see *elha{n)- ‘elk’ and *baruga- ‘boar’.

Focusing on this substrate etymon, coupled with archaeology and ancient DNA, in the recent SAA 84th Annual Meeting (Abstracts in PDF):

Kroonen, Guus (Leiden University) and Rune Iversen

[196] The Linguistic Legacy of the Pitted Ware Culture

The Scandinavian hunter-, fisher- and gatherer-based Pitted Ware culture is chronologically situated in the Neolithic. However, it challenges our traditional view on cultural and social evolution by representing a return to an otherwise abandoned hunter-gatherer lifestyle. In general, the Pitted Ware culture must be seen as an offshoot of the “Sub-Neolithic” societies inhabiting wide parts of northern and northeastern Europe in the fourth and third millennium B.C.E.

Isotopic and aDNA studies have shown that people of the east Swedish Pitted Ware culture, both dietarily and genetically were distinct from the early farmers in this region, the Funnel Beaker culture. Isotopic data shows a marked predominance of seal in the diet, which has given the Pitted Ware people the nickname “Inuit of the Baltic”.

As regards language, it is to be expected that people practicing a Pitted Ware lifestyle spoke a non-Indo-European language. In fact, there is some linguistic evidence that can support this claim. It is conceivable that both the Germanic and Finnish word for “seal” were ultimately borrowed from a language spoken in a Pitted Ware context. Once more, the linguistic evidence turns out to offer important information complementary to that of archaeology and archaeo-genetics.

prehistoric-seal-hunters
Stone Age Seal Hunters, by Måns Sjöberg.

Apparently, the idea of non-IE substrate languages in contact with Germanic in Scandinavia is fashionable for the Copenhagen group, probably due to their particular interpretation of the recent genetic papers, hence the multiple Germanic-Fennic connections to be reviewed through this new prism. While the ulterior motive of this proposal may be to try and connect yet again Germanic with CWC Denmark, I would argue that the effect is actually the opposite.

An early borrowing via Uralic

The word has always been considered a more likely loan from one language to the other, and – because of the quite popular idea of Uralic native to Fennoscandia – it was often seen as a likely borrowing of Germanic from Balto-Finnic. In any possible case, the borrowing in either direction must be quite early, for obvious reasons:

  • If the borrowing had been via late Palaeo-Germanic, the ending in *-xa– would have been reflected in Balto-Finnic, hence an early Palaeo-Germanic to Pre-Balto-Finnic stage would be necessary.
  • If the borrowing had been via late Balto-Finnic, the initial sibilant would be already aspirated, being adopted as *-x– in Palaeo-Germanic, while the ending in *-k– would have remained as such if it was adopted after Grimm’s law ceased to be active.
  • Similarly, a borrowing from a common, non-Indo-European & non-Uralic source would require that it happened during the early stages of both proto-languages to have undergone their respective phonetic changes, and both borrowings chronologically close to each other, to assume a similar vocalism and consonantism of the ultimate source.
wiik-indo-european-uralic-substrate
The idea of seal-hunting Uralic substrate of Pitted Ware is not new. Image modified from The Uralic and Finno-Ugric Phonetic Substratum, by Kalevi Wiik, Linguistica Uralica (1997).

Furthermore, regarding the most likely way of expansion of this loanword, due to the different vowels and sibilants present in Uralic but not in Indo-European:

  • A direct loan from Pre-Germanic **selkos – which shows a regular thematic declension – to Pre-Balto-Finnic *šülkeš doesn’t seem to be a reasonable assumption.
  • NOTE. A Germanic borrowing from alternative Gmc. genitive *silxis could only work in a Pre-Germanic to Pre-Balto-Finnic model, hence only if the Gmc. form can be reconstructed for an earlier stage. Even then, for the same reason stated above, the opposite could be more reasonably argued, i.e. that this form is the original one adopted in Germanic: Pre-PBF *šülkeš > Pre-Gmc. *silkis, reinterpreted as an -o- stem in its declension.

  • If we reconstruct an older Pre-Finno-Samic (i.e. with Finno-Permic-like vocalism) **šëlkëš, a borrowing into Pre-Germanic **selkos would work. Even though no Saami derivative exists to confirm such a possibility, this would be supported by the known common evolution of Finno-Samic dialects in close contact with Pre-Germanic.
  • Admittedly, even accepting the existence of a Finno-Samic stem, a potential substrate word could not be discarded. In fact, while **šëlkë- could perfectly be a Uralic root, the ending in *-š can’t be easily interpreted. Therefore, a third, non-Indo-European & non-Uralic source is a plausible explanation.

NOTE. Arguably, Proto-Finno-Samic could have adopted Gmc. *kh or *x exceptionally as PFS *k. However, early Palaeo-Germanic borrowings in Finno-Samic show a consistent regular consonant change as described above. For more on this, see Finno-Samic borrowings.

This likely Uralic first nature of the loanword is important for the discussion below.

Pitted Ware culture

pitted-ware-pyheensilta-ware-culture
Middle Neolithic A period. Distribution of Pyheensilta Ware, Funnel Beaker Culture in Sweden, and Pitted Ware Culture in northern Europe during the Middle Neolithic A period, c. 3300–2800 cal BC. Find locations with numbers demarcate sites where cereal grains have been found and later AMS radiocarbon dated. Figure was created by SV using QGIS 3.4. (https://www.qgis.org/) and Natural Earth data (https://www.naturalearthdata.com/). Image from Vanhanen et al. (2019).

About the Pitted Ware culture, this is what the recent paper by Vanhanen et al. (2019), from the University of Finland (including Volker Heyd) had to say:

The origins of the PWC are controversial. In one likely scenario, Comb Ceramic and Mesolithic hunter-gatherers first interacted with FBC during the last centuries of the EN and became specialized maritime hunter-gatherers. The PWC pushed south and westwards during the Middle Neolithic (MN), c. 3300–2300 BC, along the northern Baltic shoreline and adjacent islands, eventually reaching as far west as Denmark and southern Norway. Around 2800 BC, after the FBC ceased to exist, the Corded Ware Culture (CWC) migrated into the PWC area. The end date for the PWC and CWC is approximately 2300 BC, when the material culture was replaced by the Late Neolithic (LN) culture<. Spanning nearly a millennium virtually unchanged, the PWC maintained a coherent society and a successful economic model. PWC people lived in marine-oriented settlements, commonly dwelled in huts and produced relatively large amounts of ceramic vessels. This speaks to the partly sedentary nature of their habitation, at least for their base camps. These specialist hunter-gatherers obtained the great majority of their subsistence from maritime sources, such as seal, fish, and sea birds. Considering the amount of bones, sealing was of paramount importance, causing these peoples to be labelled ‘hard-core sealers’ or even the ‘Inuit of the Baltic’.

The Middle Neolithic Pitted Ware culture is dated ca. 3500–2300 BC, so we would be seeing here Pre-Germanic and Pre-Balto-Finnic peoples arriving near the Pitted Ware culture. That would leave us with one of both languages expanding with Corded Ware peoples, and the other with Bell Beakers. Since Battle Axe-derived cultures around the Gulf of Finland are associated with Balto-Finnic groups, and Bell Beakers arriving ca. 2400 started the Dagger Period, commonly associated with the Pre-Germanic community, I think the connection of each group with their language is self-evident.

pitted-ware-cored-ware-culture
Middle Neolithic B period. Distribution of Corded Ware Culture and Pitted Ware Culture in northern Europe during the Middle Neolithic B period, c. 2800–2300 cal BC. Find locations with numbers demarcate sites where cereal grains have been found and later AMS radiocarbon dated. Figure was created by SV using QGIS 3.4. (https://www.qgis.org/) and Natural Earth data (https://www.naturalearthdata.com/). Modified from Vanhanen et al. (2019).

NOTE. You can read some interesting information about prehistoric and recent seal hunting in the Baltic in the blog post “Själen” – Seal Hunting in the Northern Baltic Sea.

Germanic-Fennic phonetic evolution

The common Germanic – Balto-Finnic phonetic evolution, especially Verner’s law in Palaeo-Germanic and qualitative gradation in Proto-Balto-Finnic, has been variably interpreted as:

  • Uralic in Scandinavia influenced by Germanic (Verner’s law source of the gradation), by Koivulehto and Vennemann (1996).
  • Germanic over a Uralic substratum in Scandinavia, by Wiik (1997).
  • Both Germanic and Balto-Finnic influenced by a third language, an “extinct non-Uralic source” spoken in Fennoscandia before the arrival of Uralic and Indo-European, by Kallio (2001); maybe the same substrate proposed to have influenced the accent shift in Germanic similar to Uralic.
  • Balto-Finnic speakers adopting Pre-Germanic in Scandinavia, in contact with Balto-Finnic speakers retaining their language, by Schrijver in Language Contact and the Origins of the Germanic Languages (2014)– although first suggested by him in the 1990s.

NOTE. There are other (some much older) proposals of a Uralic substrate in Scandinavia, but I think those above summarize the most common positions tenable today.

If you add all linguistic, archaeological, and now genetic connections, it is really strange to keep arguing for so many surprisingly fitting common substrates and/or contact languages for both. Especially because the Pre-Germanic community – if originally from southern Scandinavia and not further south (see e.g. Kortlandt’s theory) – was marked by the Dagger Period, as accepted by most archaeologists (including Kristiansen), and we know that Bell Beakers – who triggered the Dagger period – might have arrived a little late to the Pitted Ware disintegration in most seal-hunting areas of southern Scandinavia.

bell-beaker-density
Density analysis based (Bell Beaker per km2) on the distribution of Bell Beaker per region (ca. 2700-2200 BC). Combination of different levels of b-spline interpolation. Exaltation of the values through square root usage. Modified from Michael Bilger (2018).

In other words, how many common substrate languages can we propose for Germanic (and Balto-Finnic)? Just from Kroonen we have already the Semitic-like TRB, and the seal-hunting Pitted Ware culture. Apparently, the culprit of the common phonetic evolution must be some (other?) culture that both Pre-Germanic and Pre-Balto-Finnic assimilated (or with which both were in contact) in Fennoscandia.

NOTE. I believe no data supports the attribution of those Germanic borrowings to the TRB culture, especially if one assumes they belong to an Afroasiatic branch, as did Kroonen. His initial assumption about an expansion of R1b-M269 associated with the Neolithic from Anatolia, and thus with Afroasiatic, must today be rejected. Much more likely is the incorporation of most of these loanwords during the expansion of North-West Indo-Europeans from Yamna Hungary.

How many “common” substrates from different regions and cultures is too much? Arguably, it’s not a question of quantity (because the overall probability remains the same), but a question of quality of arguments.

In my opinion, both a) the marked seal-hunting subsistence economy of the Pitted Ware culture and b) the difficult reconstruction of a fitting ‘natural’ PIE or PU stem warrant this proposal of a third source, just like the European agricultural substrate of North-West Indo-European and Palaeo-Balkan languages, as well as the Asian agricultural substrate of Indo-Iranian are the most logical interpretation of words not found in other IE dialects. The only problem in this case is the lack of other Scandinavian substrate words to compare its typology against.

scandinavia-neolithic-flint-daggers
Close contacts in Fennoscandia. The distribution of Scandinavian flint daggers (A) in the east and south Baltic region and possible trends of “down the line” trade (B). Good size and quality flint zone in the south-west Baltic region is hatched (C). According to: Wojciechowski 1976; Olausson 1983, fig. 1; Madsen 1993, 126; Libera 2001; Kriiska & Tvauri 2002, 86. Image modified from Piličiauskas (2010).

Common Scandinavian substratum

The theory of a Pitted Ware borrowing is therefore quite convincing from a cultural point of view, at the same time as it fits the linguistic data. However, one reason why I dislike the interpretation of a dual origin is that our knowledge of Uralic languages is fairly limited, whereas that of Indo-European branches and hence Proto-Indo-European is huge. To put it otherwise: if a common word appears in both, and it is most likely (culturally and linguistically) not Indo-European, it certainly means that it was borrowed in Germanic. What are the a priori chances of it coming directly from a third substrate language for both dialects, instead of coming directly from Pre-Balto-Finnic?

From Schrijver (2014):

What did happen, apparently, is that Finnic speakers had enough access to the way in which Germanic speakers pronounced Balto-Finnic in order to model their own pronunciation of Balto-Finnic on it. In other words, Balto-Finns conversed with bilingual speakers of Germanic and Balto-Finnic whose pronunciation of both was essentially Germanic. But access to the Germanic language itself was not sufficient to allow Balto-Finns to become bilingual themselves, either because social segregation prevented this or because contact with Germanic was severed before widespread bilingualism set in. This limited access to Germanic would allow us to understand why Balto-Finnic did not go the way of the vernacular languages that came in contact with Latin in the Roman Empire, where access to Latin was open to almost everybody and massive language shift in favour of Latin ensued.

NOTE. For a more detailed discussion, you can read the whole chapter dedicated to this question. I summarized it in Pre-Germanic born out of a Proto-Finnic substrate in Scandinavia.

On the other hand, about the ad hoc interpretation by Kallio (2001) of hypothetic third languages strongly influencing in the same way both the Palaeo-Germanic- and Balto-Finnic-speaking communities, Schrijver (2014) comments:

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.

early-bronze-age-nordic-dagger-period
Early Bronze Age cultures of Northern Europe (roughly ca. 2200-1750). Dagger period representing the expansion of BBC-derived groups from southern Scandinavia.

NOTE. The proposal of some kind of “SHG/EHG-based Fennoscandian substrate” seems funny to me, for two reasons: firstly, there is usually no talk about which culture spread that common language, how it survived, how it was in contact with both groups and until when, etc. (see below for possibilities); secondly, apparently the evident survival of West European EEF communities driven by at least two cultural groups – El Argar and the poorly known groups from the Atlantic façade north of the Pyrenees – is, for the same people proposing this simplistic SHG/EHG idea, somehow not fitting for the prehistory of Proto-Iberian and Proto-Aquitanian, respectively…

The same argument that one could use against the direct borrowing of both dialects from Pitted Ware, but much more strongly, can be thus wielded against a common, centuries-long phonetic evolution of both Balto-Finnic and Germanic caused by close interactions with (and/or substrate influence of) some third language. Which unitary culture and when exactly could that have happened around the Baltic Sea?

  • Was it Pitted Ware the mysterious substrate language? Seems rather unlikely, due to the early demise of the Pitted Ware culture in contrast to the long-lasting common influence seen in both dialects.
  • Was it Pitted Ware in southern Scandinavia, but Comb Ware in the Gulf of Finland? Is there a direct genetic connection between both cultures? And how likely is a common phonology of an ancestral Comb Ware-like substrate language surviving separately in Finland and Sweden? Even accepting these assumptions, we would be stuck again in the Indo-European Beakers vs. Uralic Battle Axe model.
  • Was it a succession of cultures, from some Scandinavian culture that was replaced by some incoming ethnolinguistic group, then influencing the other? This non-IE, non-Uralic substrate would then need to be proposed, given the chronological and archaeological constraints, as an effect of Pitted Ware over Pre-Finno-Baltic spoken by Battle Axe peoples in Scandinavia, then replaced by Pre-Germanic peoples arriving later with Bell Beakers. A reverse direction and later chronology (say, Germanic replaced by Balto-Finnic from Netted Ware arriving from the Volga) wouldn’t work as well.
  • Was it Asbestos Ware as a late Comb Ware group influencing both? How likely is such a continued influence in Southern Scandinavia and the Gulf of Finland? Even if we accepted this influence that miraculously didn’t affect Samic (most likely located between the Balto-Finnic-speaking Gulf of Finland and northern Fennoscandian Asbestos Ware groups), it would necessarily mean that Germanic and Balto-Finnic were spoken neighbouring exactly the same Asbestos Ware groups in Scandinavia. That is, essentially, that the BBC-derived Dagger Period represented Pre-Germanic, while Battle Axe-derived groups around the Gulf of Finland were Balto-Finnic.

Mixing linguistics with archaeology (now complemented with genetics) also risks circular reasoning. But, how else can someone propose a third substrate language for a phonetic change, necessarily represented by Fennoscandian groups potentially separated by thousands of years? In this age of population genomics we can’t simply talk about theoretical models anymore: we must refer to Fennoscandian cultures and populations in a very specific time frame, as Kronen & Iversen do in their proposal. Not only is such a third unknown language usually a weak explanation for a common development of two unrelated languages; in this case it finds no support whatsoever.

Seals and the Arctic

Another interesting aspect about this Fennic-Germanic comparandum is its relevance to the Uralic homeland problem.

uralic-languages-modern
Current distribution of Uralic languages. Nenets and Saami are among the best positioned to retain the ‘original’ Uralic seal-hunting vocabulary.

Since the publication of Mittnik et al. (2018), Lamnidis et al. (2018), and Sikora et al. (2018), the new normal is apparently to consider Corded Ware Finland as Germanic-speaking, the Gulf of Finland as Balto-Slavic-speaking, while the Kola peninsula and whichever Palaeo-Arctic peoples preceded Nganasans and Nenets as ancient Uralians. Uh-huh, OK.

But, if prehistoric Arctic peoples practiced specialized seal-hunting economies, and Uralians were one among such populations – supposedly one widespread from the Barents Sea to the Lapteve Sea…how come no common Uralic word for ‘seal’ exists? In other words, why would these True™ Uralic peoples expanding from the Arctic need to borrow a word for ‘seal’ from neighbouring populations in every single seal-hunting region they are attested?

grey-seal-distribution
Historical distribution of grey seals, an important part of the diet around the Baltic Sea. Image modified from Wikimedia to include Skagerrak and Kattegat regions.

About Saami, which some have recklessly proposed to be derived from Bronze Age N1c-L392 samples from the Kola Peninsula (against the good judgment of the authors of the paper), this is what we know from their word for ‘seal’, from Grünthal (2004):

Ter Saami vīrre ‘seal; wolf’ displays two meanings that refer to clearly different animals. Neither of them is borrowed from the source language because the word descends from Russian zver’ ‘animal’ (T.I.Itkonen 1958: 756). Another word, Skolt Saami näúdd ‘seal, wolf’, has been similarly used in the two meanings. The evidence of North Saami návdi ‘wolf; creature, fur animal; beast’ (Sammallahti 1989: 305; Lagercrantz (1939: 518) presents the alternative meanings in the opposite order; E. Itkonen (1969: 148) lists the meanings ‘wildes Tier; Raubtier (bes. Wolf); Pelztier’) suggesting that ‘wolf’ is the primary sense and ‘seal’ is a metaphorical extension of it. More precisely, it is an example of a mythic metaphor (cf. Siikala 1992). According to the old folk belief, seal was a wolf and the Skolt Saamis preferred not to eat its meat (T.I.Itkonen 1958: 906). Before that the metonymic meaning ‘wolf’ rose from the less specified meanings, and originally návdi is a Scandinavian or Finnic loan word in Saamic, cf. Old Norse naut ‘vieh, rind’, Icelandic and Norwegian naut, Swedish nöt < Germanic *nauta ‘property’ (Hellquist 1980: 721, T.I.Itkonen 1958: 275, Lagercrantz 1939: 518, de Vries 1961: 406; E. Itkonen (1969: 148) considers Finnic, cf. Finnish nauta ‘bovine’ (< Germanic) as a possible alternative source for the Saamic word).

NOTE. Possibly comparable, for the mythic metaphor proper of Scandinavian folk belief, are Germanic derivatives built as ‘seal-hound’ and/or ‘sea-hound’.

sea-distribution-arctic
Seals formed a great part of the diet for Palaeo-Arctic populations. Boundaries of regions used to predict sea ice, superimposed over the distributions of the five ringed seal subspecies. Image modified from Kelly et al. (2010).

About Nenets (quite close to the Naganasans of pure “Siberian ancestry”), here is what Edward Vajda, an expert in Palaeo-Siberian languages, has to say:

Nenets techniques for hunting the animals of the Arctic Ocean seem to have been borrowed from the first Arctic aborigines. Thus, the Nenets word for seal is nyak, the Eskimo word is nesak. Also, the Nenets word for a one-piece Arctic clothing is lu; the Korak word on the Kamchatka peninsula for clothing is l’ku. All of these groups may have borrowed the words from some original circumpolar aborigines. More probably, the first settlers of Arctic Europe were cousins of the present-day Eskimo, Chukchi and other residents of the far northeast region of Asia. Nenets folklore also speaks of the aborigines living in ice dugouts (igloos).

On the other hand, Proto-Uralic shows a Chalcolithic steppe-like culture, with common words for metal and metalworking, for agriculture, and for domesticated animals, most likely including cattle. They were close to Indo-Europeans since at least before the Tocharian split, and probably earlier than that (even if one does not accept the Indo-Uralic phylum). And there were clearly strong contacts of Finno-Ugric with Indo-Iranian, and especially of Finno-Samic with Germanic.

uralic-cline
Uralic clines from Corded Ware groups to the east. A clear reason for the lack of common seal-hunting vocabulary. Modified from Tambets et al. (2018). Principal component analysis (PCA) and genetic distances of Uralic-speaking populations. a PCA (PC1 vs PC2) of the Uralic-speaking populations. You can see another PCA including ancient samples.

Some among my readers may now be thinking about these totally believable proposals of prehistoric cultures around Lake Baikal representing the True™ Uralic homeland; because haplogroup N1c, and because some 0.5% more “Devil’s Gate Cave ancestry” in Estonians than in Lithuanians; despite the fact that 1) the so-called “Siberian ancestry” formed an ancestral cline with EHG in North Eurasia, that 2) N1c-L392 lineages seem to appear among many Asian peoples of different languages, and that 3) recent prehistoric N1c-L392 lines expanded clearly with Micro-Altaic languages.

Like, who would have hunted seals in Lake Baikal, right? The problem is, seals represented one of their main game, essential for their subsistence economy. From Novokonova et al. (2015):

One of the key reasons for the density of human settlement in the Baikal region compared to adjacent areas of Siberia is that the lake and its nearby rivers offer an abundance of aquatic food resources, including several endemic species, with perhaps the most well known being the Baikal seal. This freshwater seal is only found in Lake Baikal and portions of its tributaries. It shares lifecycle and behavioral patterns with other small northern ice-adapted seals, and is genetically and morphologically most closely related to the ringed seal (Pusa hispida). The nerpa can grow up to 1.8 m long and weigh as much as 130 kg, with the males tending to be slightly larger than the females.

Zooarchaeological analyses of the 16,000 Baikal seal remains from this well-dated site clearly show that sealing began here at least 9000 calendar years ago. The use of these animals at Sagan-Zaba appears to have peaked in the Middle Holocene, when foragers used the site as a spring hunting and processing location for yearling and juvenile seals taken on the lake ice. After 4800 years ago, seal use declined at the site, while the relative importance of ungulate hunting and fishing increased. Pastoralists began occupying Sagan-Zaba at some point during the Late Holocene, and these groups too utilized the lake’s seals. Domesticated animals are increasingly common after about 2000 years ago, a pattern seen elsewhere in the region, but spring and some summer hunting of seals was still occurring. This use of seals by prehistoric herders mirrors patterns of seal use among the region’s historic and modern groups.

Bronze Age movements in Fennoscandia

Regarding the shrinkage and expansion of different farming economic strategies in Scandinavia since the Neolithic, with potential relevance for population movements and thus ethnolinguistic change – either from Balto-Finnic peoples migrating back from eastern Sweden, or Germanic peoples moving to eastern Finland – from Vanhanen et al. (2019):

Cultivated plants at CWC sites in Finland were not discovered in the current investigation (Supplementary Results) or earlier studies. In Finland, the keeping of domestic animals is indicated by the evidence of dairy lipids and mineralized goat hairs. Charred remains and impressions of cultivated plants have been discovered at CWC sites in Estonia and east-central Sweden (Fig. 3: 12). In the eastern Baltic region, the earliest bones of domestic animals and a shift in subsistence occurred with the CWC. Whether CWC produced the cereals and other agricultural products found at PWC sites is difficult to estimate because only small amounts of plant remains have ever been discovered at CWC sites. The CWC seemingly reached east-central Sweden from regions further to the east, where there is evidence of animal husbandry, but only very few signs of plant cultivation.

For the Late Neolithic (LN), cereal grains have been found north of Mälaren and along the Norrland coast. In mainland Finland, the first cereal grains occur during the LN or Bronze Age, c. 1900–1250 cal BC. The earliest bones of sheep/goat from mainland Finland are earlier, dating back to 2200–1950 cal BC. Finds of Scandinavian bronze artefacts indicate an influx from east-central Sweden, which might well be a source area for these agricultural innovations. A similar development is found in the eastern Baltic region, where the earliest directly radiocarbon-dated cereals originate from the Bronze Age, 1392–1123 cal BC (2 sigma). Thus, agriculture was evident during the Bronze Age in the eastern Baltic, but at least animal keeping and probably crop cultivation were present earlier during the CWC phase.

It has been known for a while already that the only options left for the expansion of Finno-Saami into Fennoscandia are either Battle Axe (continued in Textile Ceramics) or Netted Ware (as proposed e.g. by Parpola), based, among other data, on language contacts, language estimates, cultural evolution, and population genomics. Data like this one on seal-hunting vocabulary also support the most likely option, which entails the identification of Corded Ware as the vector of expansion of Uralic languages.

NOTE. Also interesting in this regard is the lack of Slavic words for ‘seal’ – borrowed, in Russian from Samic, and in other Slavic dialects from Russian, Latin, or other languages -, and the coinage of a new term in East Baltic. Rather odd for an “autochthonous” Proto-Baltic (supposedly in contact with Pitted Ware, Germanic, and Balto-Finnic, then), and for a Proto-Slavic stemming from the Baltic. Quite appropriate, though, for a Proto-East Baltic arriving in the Baltic with Trzciniec and for a Proto-Slavic community evolving further south.

So, what new episode in this renewed 2000s R1b/R1a/N1c soap opera is it going to be, when eastern Fennoscandia shows Corded Ware-derived peoples of “steppe ancestry” (and mainly R1a-Z645 lineages) continue during the Bronze Age? Will the resurge and/or infiltration of I2 – maybe even N1c – lineages among Corded Ware-derived cultures of north-eastern Europe support or challenge this model, and why? Make your bet below.

Related

N1c-L392 associated with expanding Turkic lineages in Siberia

haplogroup-n1c-tat

Second in popularity for the expansion of haplogroup N1a-L392 (ca. 4400 BC) is, apparently, the association with Turkic, and by extension with Micro-Altaic, after the Uralic link preferred in Europe; at least among certain eastern researchers.

New paper in a recently created journal, by the same main author of the group proposing that Scythians of hg. N1c were Turkic speakers: On the origins of the Sakhas’ paternal lineages: Reconciliation of population genetic / ancient DNA data, archaeological findings and historical narratives, by Tikhonov, Gurkan, Demirdov, and Beyoglu, Siberian Research (2019).

Interesting excerpts:

According to the views of a number of authoritative researchers, the Yakut ethnos was formed in the territory of Yakutia as a result of the mixing of people from the south and the autochthonous population [34].

These three major Sakha paternal lineages may have also arrived in Yakutia at different times and/ or from different places and/or with a difference in several generations instead, or perhaps Y-chromosomal STR mutations may have taken place in situ in Yakutia. Nevertheless, the immediate common ancestor(s) from the Asian Steppe of these three most prevalent Sakha Y-chromosomal STR haplotypes possibly lived during the prominence of the Turkic Khaganates, hence the near-perfect matches observed across a wide range of Eurasian geography, including as far as from Cyprus in the West to Liaoning, China in the East, then Middle Lena in the North and Afghanistan in the South (Table 3 and Figure 5). There may also be haplotypes closely-related to ‘the dominant Elley line’ among Karakalpaks, Uzbeks and Tajiks, however, limitations in the loci coverage for the available dataset (only eight Y-chromosomal STR loci) precludes further conclusions on this matter [25].

yakutia-haplogroup-n1c
17-loci median-joining network analysis of the original/dominant Elley, Unknown and Omogoy Y-chromosomal STR haplotypes with the YHRD matches from outside Yakutia populations.

According to the results presented here, very similar Y-STR haplotypes to that of the original Elley line were found in the west: Afghanistan and northern Cyprus, and in the east: Liaoning Province, China and Ulaanbaator, Northern Mongolia. In the case of the dominant Omogoy line, very closely matching haplotypes differing by a single mutational step were found in the city of Chifen of the Jirin Province, China. The widest range of similar haplotypes was found for the Yakut haplotype Unknown: In Mongolia, China and South Korea. For instance, haplotypes differing by a single step mutation were found in Northern Mongolia (Khalk, Darhad, Uryankhai populations), Ulaanbaator (Khalk) and in the province of Jirin, China (Han population).

n1c-uralic-altaic-siberia
14-loci median-joining network analysis for the original/dominant Elley (Ell), Unknown Clan
(Vil), Omogoy (Omo), Eurasian (Eur) and Xiongnu (Xuo) Y-chromosomal STR haplotypes and that for a representative ancient DNA sample (Ch0 or DSQ04) from the Upper Xiajiadian Culture
recovered from the Inner Mongolia Autonomous Region, China.

Notably, Tat-C-bearing Y-chromosomes were also observed in ancient DNA samples from the 2700-3000 years-old Upper Xiajiadian culture in Inner Mongolia, as well as those from the Serteya II site at the Upper Dvina region in Russia and the ‘Devichyi gory’ culture of long barrow burials at the Nevel’sky district of Pskovsky region in Russia. A 14-loci Y-chromosomal STR median-joining network of the most prevalent Sakha haplotypes and a Tat-C-bearing haplotype from one of the ancient DNA samples recovered from the Upper Xiajiadian culture in Inner Mongolia (DSQ04) revealed that the contemporary Sakha haplotype ‘Xuo’ (Table 2, Haplotype ID “Xuo”) classified as that of ‘the Xiongnu clan’ in our current study, was the closest to the ancient Xiongnu haplotype (Figure 6). TMRCA estimate for this 14-loci Y-chromosomal STR network was 4357 ± 1038 years or 2341 ± 1038 BCE, which correlated well with the Upper Xiajiadian culture that was dated to the Late Bronze Age (700-1000 BCE).

eurasian-n-subclades
Geographical location of ancient samples belonging to major clade N of the Y-chromosome.

NOTE. Also interesting from the paper seems to be the proportion of E1b1b among admixed Russian populations, in a proportion similar to R1a or I2a(xI2a1).

It is tempting to associate the prevalent presence of N1c-L392 in ancient Siberian populations with the expansion of Altaic, by simplistically linking the findings (in chronological order) near Lake Baikal (Damgaard et al. 2018), Upper Xiajiadian (Cui et al. 2013), among Khövsgöl (Jeong et al. 2018), in Huns (Damgaard et al. 2018), and in Mongolic-speaking Avars (Csáky et al. 2019).

However, its finding among Palaeo-Laplandic peoples in the Kola peninsula ca. 1500 BC (Lamnidis et al. 2018) and among Palaeo-Siberian populations near the Yana River (Sikora et al. 2018) ca. AD 1200 should be enough to accept the hypothesis of ancestral waves of expansion of the haplogroup over northern Eurasia, with acculturation and further expansions in the different regions since the Iron Age (see more on its potential expansion waves).

Also, a simple look at the TMRCA and modern distribution was enough to hypothesize long ago the lack of connection of N1c-L392 with Altaic or Uralic peoples. From Ilumäe et al. (2016):

Previous research has shown that Y chromosomes of the Turkic-speaking Yakuts (Sakha) belong overwhelmingly to hg N3 (formerly N1c1). We found that nearly all of the more than 150 genotyped Yakut N3 Y chromosomes belong to the N3a2-M2118 clade, just as in the Turkic-speaking Dolgans and the linguistically distant Tungusic-speaking Evenks and Evens living in Yakutia (Table S2). Hence, the N3a2 patrilineage is a prime example of a male population of broad central Siberian ancestry that is not intrinsic to any linguistically defined group of people. Moreover, the deepest branch of hg N3a2 is represented by a Lebanese and a Chinese sample. This finding agrees with the sequence data from Hallast et al., where one Turkish Y chromosome was also assigned to the same sub-clade. Interestingly, N3a2 was also found in one Bhutan individual who represents a separate sub-lineage in the clade. These findings show that although N3a2 reflects a recent strong founder effect primarily in central Siberia (Yakutia, Sakha), the sub-clade has a much wider distribution area with incidental occurrences in the Near East and South Asia.

haplogroup-n1a-M2118
Frequency-Distribution Maps of Individual Sub-clades of hg N3a2, by Ilumäe et al. (2016).

The most striking aspect of the phylogeography of hg N is the spread of the N3a3’6-CTS6967 lineages. Considering the three geographically most distant populations in our study—Chukchi, Buryats, and Lithuanians—it is remarkable to find that about half of the Y chromosome pool of each consists of hg N3 and that they share the same sub-clade N3a3’6. The fractionation of N3a3’6 into the four sub-clades that cover such an extraordinarily wide area occurred in the mid-Holocene, about 5.0 kya (95% CI = 4.4–5.7 kya). It is hard to pinpoint the precise region where the split of these lineages occurred. It could have happened somewhere in the middle of their geographic spread around the Urals or further east in West Siberia, where current regional diversity of hg N sub-lineages is the highest (Figure 1B). Yet, it is evident that the spread of the newly arisen sub-clades of N3a3’6 in opposing directions happened very quickly. Today, it unites the East Baltic, East Fennoscandia, Buryatia, Mongolia, and Chukotka-Kamchatka (Beringian) Eurasian regions, which are separated from each other by approximately 5,000–6,700 km by air. N3a3’6 has high frequencies in the patrilineal pools of populations belonging to the Altaic, Uralic, several Indo-European, and Chukotko-Kamchatkan language families. There is no generally agreed, time-resolved linguistic tree that unites these linguistic phyla. Yet, their split is almost certainly at least several millennia older than the rather recent expansion signal of the N3a3’6 sub-clade, suggesting that its spread had little to do with linguistic affinities of men carrying the N3a3’6 lineages.

haplogroup_n3a3
Frequency-Distribution Maps of Individual Subclade N3a3 / N1a1a1a1a1a-CTS2929/VL29.

It was thus clear long ago that N1c-L392 lineages must have expanded explosively in the 5th millennium through Northern Eurasia, probably from a region to the north of Lake Baikal, and that this expansion – and succeeding ones through Northern Eurasia – may not be associated to any known language group until well into the common era.

Related

The cradle of Russians, an obvious Finno-Volgaic genetic hotspot

pskov-novgorod-russia

First look of an accepted manuscript (behind paywall), Genome-wide sequence analyses of ethnic populations across Russia, by Zhernakova et al. Genomics (2019).

Interesting excerpts:

There remain ongoing discussions about the origins of the ethnic Russian population. The ancestors of ethnic Russians were among the Slavic tribes that separated from the early Indo-European Group, which included ancestors of modern Slavic, Germanic and Baltic speakers, who appeared in the northeastern part of Europe ca. 1,500 years ago. Slavs were found in the central part of Eastern Europe, where they came in direct contact with (and likely assimilation of) the populations speaking Uralic (Volga-Finnish and Baltic- Finnish), and also Baltic languages [11–13]. In the following centuries, Slavs interacted with the Iranian-Persian, Turkic and Scandinavian peoples, all of which in succession may have contributed to the current pattern of genome diversity across the different parts of Russia. At the end of the Middle Ages and in the early modern period, there occurred a division of the East Slavic unity into Russians, Ukrainians and Belarusians. It was the Russians who drove the colonization movement to the East, although other Slavic, Turkic and Finnish peoples took part in this movement, as the eastward migrations brought them to the Ural Mountains and further into Siberia, the Far East, and Alaska. During that interval, the Russians encountered the Finns, Ugrians, and Samoyeds speakers in the Urals, but also the Turkic, Mongolian and Tungus speakers of Siberia. Finally, in the great expanse between the Altai Mountains on the border with Mongolia, and the Bering Strait, they encountered paleo-Asiatic groups that may be genetically closest to the ancestors of the Native Americans. Today’s complex patchwork of human diversity in Russia has continued to be augmented by modern migrations from the Caucasus, and from Central Asia, as modern economic migrations take shape.

pskov-novgorod-pca-eurasia-yakut
Sample relatedness based on genotype data. Eurasia: Principal Component plot of 574 modern Russian genomes. Colors reflect geographical regions of collection; shapes reflect the sample source. Red circles show the location of Genome Russia samples.

In the current study, we annotated whole genome sequences of individuals currently living on the territory of Russia and identifying themselves as ethnic Russian or as members of a named ethnic minority (Fig. 1). We analyzed genetic variation in three modern populations of Russia (ethnic Russians from Pskov and Novgorod regions and ethnic Yakut from the Sakha Republic), and compared them to the recently released genome sequences collected from 52 indigenous Russian populations. The incidence of function-altering mutations was explored by identifying known variants and novel variants and their allele frequencies relative to variation in adjacent European, East Asian and South Asian populations. Genomic variation was further used to estimate genetic distance and relationships, historic gene flow and barriers to gene flow, the extent of population admixture, historic population contractions, and linkage disequilibrium patterns. Lastly, we present demographic models estimating historic founder events within Russia, and a preliminary HapMap of ethnic Russians from the European part of Russia and Yakuts from eastern Siberia.

pskov-novgorod-pca-finno-permic
Sample relatedness based on genotype data. Western Russia and neighboring countries: Principal Component plot of 574 modern Russian genomes. Colors reflect geographical regions of collection; shapes reflect the sample source. Red circles show the location of Genome Russia samples.

The collection of identified SNPs was used to inspect quantitative distinctions among 264 individuals from across Eurasia (Fig. 1) using Principal Component Analysis (PCA) (Fig. 2). The first and the second eigenvectors of the PCA plot are associated with longitude and latitude, respectively, of the sample locations and accurately separate Eurasian populations according to geographic origin. East European samples cluster near Pskov and Novgorod samples, which fall between northern Russians, Finno-Ugric peoples (Karelian, Finns, Veps etc.), and other Northeastern European peoples (Swedes, Central Russians, Estonian, Latvians, Lithuanians, and Ukrainians) (Fig. 2b). Yakut individuals map into the Siberian sample cluster as expected (Fig. 2a). To obtain an extended view of population relationships, we performed a maximum likelihood-based estimation of ancestry and population structure using ADMIXTURE [46](Fig. 2c). The Novgorod and Pskov populations show similar profiles with their Northeastern European ancestors while the Yakut ethnic group showed mixed ancestry similar to the Buryat and Mongolian groups.

pskov-novgorod-yakut-admixture
Population structure across samples in 178 populations from five major geographic regions (k=5). Samples are pooled across three different studies that covered the territory of Russian Federation (Mallick et al. 2016 [36], Pagani et al. 2016 [37], this study). The optimal k-value was selected by value of cross validation error. Russian samples from all studies (highlighted in bold dark blue) show a slight gradient from Eastern European (Ukrainian, Belorussian, Polish) to North European (Estonian Karelian, Finnish) structures, reflecting population history of northward expansion. Yakut samples from different studies (highlighted in bold red) also show a slight gradient from Mongolian to Siberian people (Evens), as expected from their original admixture and northward expansions. The samples originated from this study are highlighted, and plotted in separated boxes below.

Possible admixture sources of the Genome Russia populations were addressed more formally by calculating F3 statistics, which is an allele frequency-based measure, allowing to test if a target population can be modeled as a mixture of two source populations [48]. Results showed that Yakut individuals are best modeled as an admixture of Evens or Evenks with various European populations (Supplemental Table S4). Pskov and Novgorod showed admixture of European with Siberian or Finno-Ugric populations, with Lithuanian and Latvian populations being the dominant European sources for Pskov samples.

direction-expansion-russians
The heatmaps of gene flow barriers show for each point at the geographical map the interpolated differences in allele frequencies (AF) between the estimated AF at the point with AFs in the vicinity of this point. The direction of the maximal difference in allele frequencies is coded by colors and arrows.

So, Russians expanding in the Middle Ages as acculturaded Finno-Volgaic peoples.

Or maybe the true Germano-Slavonic™-speaking area was in north-eastern Europe, until the recent arrival of Finno-Permians with the totally believable Nganasan-Saami horde, whereas Yamna -> Bell Beaker represented Vasconic-Caucasian expanding all over Europe in the Bronze Age. Because steppe ancestry in Fennoscandia and Modern Basques in Iberia.

A really hard choice between equally plausible models.

Related

Common Slavs from the Lower Danube, expanding with haplogroup E1b-V13?

late-iron-age-eastern-europe

Florin Curta has published online his draft for Eastern Europe in the Middle Ages (500-1300), Brill’s Companions to European History, Vol. 10 (2019), apparently due to appear in June.

Some interesting excerpts, relevant for the latest papers (emphasis mine):

The Archaeology of the Early Slavs

(…) One of the most egregious problems with the current model of the Slavic migration is that it is not at all clear where it started. There is in fact no agreement as to the exact location of the primitive homeland of the Slavs, if there ever was one. The idea of tracing the origin of the Slavs to the Zarubyntsi culture dated between the 3rd century BC and the first century AD is that a gap of about 200 years separates it from the Kiev culture (dated between the 3rd and the 4th century AD), which is also attributed to the Slavs. Furthermore, another century separates the Kiev culture from the earliest assemblages attributed to the Prague culture. It remains unclear as to where the (prehistoric) Slavs went after the first century, and whence they could return, two centuries later, to the same region from which their ancestors had left. The obvious cultural discontinuity in the region of the presumed homeland raises serious doubts about any attempts to write the history of the Slavic migration on such a basis. There is simply no evidence of the material remains of the Zarubyntsi, Kiev, or even Prague culture in the southern and southwestern direction of the presumed migration of the Slavs towards the Danube frontier of the Roman Empire.

Moreover, the material culture revealed by excavations of 6th- to 7th-century settlements and, occasionally, cremation cemeteries in northwestern Russia, Belarus, Poland, Moravia, and Bohemia is radically different from that in the lands north of the Danube river, which according to the early Byzantine sources were inhabited at that time by Sclavenes: no settlement layout with a central, open area; no wheel-made pottery or pottery thrown on a tournette; no clay rolls inside clay ovens; few, if any clay pans; no early Byzantine coins, buckles, or remains of amphorae; no fibulae with bent stem, and few, if any bow fibulae. Conversely, those regions have produced elements of material culture that have no parallels in the lands north of the river Danube: oval, trough-like settlement features (which are believed to be remains of above-ground, log-houses); exclusively handmade pottery of specific forms; very large settlements, with over 300 houses; fortified sites that functioned as religious or communal centers; and burials under barrows. With no written sources to inform about the names and identities of the populations living in the 6th and 7th centuries in East Central and Eastern Europe, those contrasting material culture profiles could hardly be interpreted as ethnic commonality. In other words, there is no serious basis for attributing to the Sclavenes (or, at least, to those whom early Byzantine authors called so) any of the many sites excavated in Russia, Belarus, Poland, Moravia, and Bohemia.

slavic-expansion-prague-korchak
Common Slavic expanding with Prague-Korchak from the east…or was it from the west?

Migrations

There is of course evidence of migrations in the 6th and 7th centuries, but not in the directions assumed by historians. For example, there are clear signs of settlement discontinuity in northern Germany and in northwestern Poland. German archaeologists believe that the bearers of the Prague culture who reached northern Germany came from the south (from Bohemia and Moravia), and not from the east (from neighboring Poland or the lands farther to the east). At any rate, no archaeological assemblage attributed to the Slavs either in northern Germany or in northern Poland may be dated earlier than ca. 700. In Poland, settlement discontinuity was postulated, to make room for the new, Prague culture introduced gradually from the southeast (from neighboring Ukraine). However, there is increasing evidence of 6th-century settlements in Lower Silesia (western Poland and the lands along the Middle Oder) that have nothing to do with the Prague culture. Nor is it clear how and when did the Prague culture spread over the entire territory of Poland. No site of any of the three archaeological cultures in Eastern Europe that have been attributed to the Slavs (Kolochin, Pen’kivka, and Prague/Korchak) has so far been dated earlier than the sites in the Lower Danube region where the 6th century sources located the Sclavenes. Neither the Kolochin, nor the Pen’kivka cultures expanded westwards into East Central or Southeastern Europe; on the contrary, they were themselves superseded in the late 7th or 8th century by other archaeological cultures originating in eastern Ukraine. Meanwhile, there is an increasing body of archaeological evidence pointing to very strong cultural influences from the Lower and Middle Danube to the Middle Dnieper region during the 7th century—the opposite of the alleged direction of Slavic migration.

When did the Slavs appear in those regions of East Central and Eastern Europe where they are mentioned in later sources? A resistant stereotype of the current scholarship on the early Slavs is that “Slavs are Slavonic-speakers; Slavonic-speakers are Slavs.”* If so, when did people in East Central and Eastern Europe become “Slavonic speakers”? There is in fact no evidence that the Sclavenes mentioned by the 6th-century authors spoke Slavic (or what linguists now call Common Slavic). Nor can the moment be established (with any precision), at which Slavic was adopted or introduced in any given region of East Central and Eastern Europe.** To explain the spread of Slavic across those regions, some have recently proposed the model of a koiné, others that of a lingua franca. The latter was most likely used within the Avar polity during the last century of its existence (ca. 700 to ca. 800).

*Ziółkowski, “When did the Slavs originate?” p. 211. On the basis of the meaning of the Old Church Slavonic word ięzyk (“language,” but also “people” or “nation”), Darden, “Who were the Sclaveni?” p. 138 argues that the meaning of the name the Slavs gave to themselves was closely associated with the language they spoke.

**Uncertainty in this respect dominates even in recent studies of contacts between Slavic and Romance languages (particularly Romanian), even though such contacts are presumed to have been established quite early (Paliga, “When could be dated ‘the earliest Slavic borrowings’?”; Boček, Studie). Recent studies of the linguistic interactions between speakers of Germanic and speakers of Slavic languages suggest that the adoption of place names of Slavic origin was directly linked to the social context of language contact between the 9th and the 13th centuries (Klír, “Sociální kontext”).

Avars

During the 6th century, the area between the Danube and the Tisza in what is today Hungary, was only sparsely inhabited, and probably a “no man’s land” between the Lombard and Gepid territories. It is only after ca. 600 that this area was densely inhabited, as indicated by a number of new cemeteries that came into being along the Tisza and north of present-day Kecskemét. There can therefore be no doubt about the migration of the Avars into the Carpathian Basin, even though it was probably not a single event and did not involve only one group of population, or even a cohesive ethnic group.

The number of graves with weapons and of burials with horses is particularly large in cemeteries excavated in southwestern Slovakia and in neighboring, eastern Austria. This was a region of special status on the border of the qaganate, perhaps a “militarized frontier.” From that region, the Avar mores and fashions spread farther to the west and to the north, into those areas of East Central Europe in which, for reasons that are still not clear, Avar symbols of social rank were particularly popular, as demonstrated by numerous finds of belt fittings. Emulating the success of the Avar elites sometimes involved borrowing other elements of social representation, such as the preferential deposition of weapons and ornamented belts. For example, in the early 8th century, a few males were buried in Carinthia (southern Austria) with richly decorated belts imitating those in fashion in the land of the Avars, but also with Frankish weapons and spurs. Much like in the Avar-age cemeteries in Slovakia and Hungary, the graves of those socially prominent men are often surrounded by many burials without any grave goods whatsoever.

early-avar-khaganate
Territory of the early Avar Qaganate and the location of the investigated sites in the Carpathian Basin in Csáky et al. (2019).

Carantanians

Carantania was a northern neighbor of the Lombard duchy of Friuli, which was inhabited by Slavs. According to Paul the Deacon, who was writing in the late 780s, those Slavs called their country Carantanum, by means of a corruption of the name of ancient Carnuntum (a former Roman legionary camp on the Danube, between Vienna and Bratislava). Carantanians were regarded as Slavs by the author of a report known as the Conversion of the Bavarians and Carantanians, and written in ca. 870 in order to defend the position of the archbishop of Salzburg against the claims of Methodius, the bishop of Pannonia.94 According to this text, a duke named Boruth was ruling over Carantania when he was attacked by Avars in ca. 740. He called for the military assistance of his Bavarian neighbors. The Bavarian duke Odilo (737–748) obliged, defeated the Avars, but in the process also subdued the Carantanians to his authority. Once Bavarian overlordship was established in Carantania, Odilo took with him as hostages Boruth’s son Cacatius and his nephew Chietmar (Hotimir). Both were baptized in Bavaria. During the 743 war between Odilo and Charles Martel’s two sons, Carloman and Pepin (the Mayors of the Palace in Austrasia and Neustria, respectively), Carantanian troops fought on the Bavarian side. The Bavarian domination cleared the field for missions of conversion to Christianity sent by Virgil, the new bishop of Salzburg (746–784). Many missionaries were of Bavarian origin, but some were Irish monks.

Moravians

Several Late Avar cemeteries dated to the last quarter of the 8th century are known from the lands north of the middle course of the river Danube, in what is today southern Slovakia and the valley of the Lower Morava [see image below]. By contrast, only two cemeteries have so far been found in Moravia (the eastern part of the present-day Czech Republic), along the middle and upper course of the Morava and along its tributary, the Dyje. In both Dolní Dunajovice and Hevlín, the latest graves may be dated by means of strap ends and belt mounts with human figures to the very end of the Late Avar period. (…)

The archaeological evidence pertaining to burial assemblages dated to the early 9th century is completely different. Shortly before or after 800, all traces of cremation—with or without barrows—disappear from the valley of the Morava river and southwestern Slovakia, two regions in which cremation had been the preferred burial rite during the previous centuries. This dramatic cultural change has often been interpreted as a direct influence of both Avar and Frankish burial rites, but it coincides in time with the adoption of Christianity by local elites. In spite of conversion, however, the representation of status through furnished burial continued well into the 9th century. Unlike Avar-age sites in Hungary and the surrounding regions, many men were buried in 9th-century Moravia together with their spurs, in addition to such weapons as battle axes, “winged” lance heads, or swords with high-quality steel blades of Frankish production.

morvaian-sites
Relevant Moravian sites mentioned in Curta’s new book.

When the Magyars inflicted a crushing defeat on the Bavarians at Bratislava (July 4, 907), the fate of Moravia was sealed as well. Moravia and the Moravians disappear from the radar of the written sources, and historians and archaeologists alike believe that the polity collapsed as a result of the Magyar raids.

Magyars

(…) although there can be no doubt about the relations between Uelgi and the sites in Hungary attributed to the first generations of Magyars, those relations indicate a migration directly from the Trans-Ural lands, and not gradually, with several other stops in the forest-steppe and steppe zones of Eastern Europe. In the lands west of the Ural Mountains, the Magyars are now associated with the Kushnarenkovo (6th to 8th century) and Karaiakupovo (8th to 10th century) cultures, and with such burial sites as Sterlitamak (near Ufa, Bashkortostan) and Bol’shie Tigany (near Chistopol, Tatarstan).* However, the same problem with chronology makes it difficult to draw the model of a migration from the lands along the Middle Volga. Many parallels for the so typically Magyar sabretache plates found in Hungary are from that region. They have traditionally been dated to the 9th century, but more recent studies point to the coincidence in time between specimens found in Eastern Europe and those from Hungary.

* Ivanov, Drevnie ugry-mad’iary; Ivanov and Ivanova, “Uralo-sibirskie istoki”; Boldog et al., “From the ancient homelands,” p. 3; Ivanov, “Similarities.” Ivanov, “Similarities,” p. 562 points out that the migration out of the lands along of the Middle Volga is implied by the disappearance of both cultures (Kushnarenkovo and Karaiakupovo) in the mid-9th century. For the Kushnarenkovo culture, see Kazakov, “Kushnarenkovskie pamiatniki.” For the Karaiakupovo culture, see Mogil’nikov, “K probleme.”

Given that the Magyars are first mentioned in relation to events taking place in the Lower Danube area in the 830s, the Magyar sojourn in Etelköz must have been no longer than 60 years or so—a generation. (…)

arrival-of-hungarians-feszty-slavs
A detail of the Arrival of the Hungarians, Árpád Feszty’s and his assistants’ vast (1800 m2) cyclorama, painted to celebrate the 1000th anniversary of the Magyar conquest of Hungary, now displayed at the Ópusztaszer National Heritage Park in Hungary. This specific detail is probably based on the account on The Annals of Fulda, which narrates under the year 894 that the Hungarians crossed the Danube into Pannonia where they “killed men and old women outright and carried off the young women alone with them like cattle to satisfy their lusts and reduced the whole” province “to desert”.

It has become obvious by now that one’s impression of the Magyars as “Easterners” and “steppe-like” was (and still is) primarily based on grave finds, while the settlement material is considerably more aligned with what is otherwise known from other contemporary settlement sites in Central and Southeastern Europe. The dominant feature on the 10th- and 11th-century settlements in Hungary is the sunken-floored building of rectangular plan, with a stone oven in a corner. Similarly, the pottery resulting from the excavation of settlement sites is very similar to that known from many other such sites in Eastern Europe. Moreover, while clear changes taking place in burial customs between ca. 900 and ca. 1100 are visible in the archaeological record from cemeteries, there are no substantial differences between 10th- and the 11th-century settlements in Hungary. (…)

As a matter of fact, the increasing quantity of paleobotanical and zooarchaeological data from 10th-century settlements strongly suggests that the economy of the first generations of Magyars in Hungary was anything but nomadic. To call those Magyars “half-nomad” is not only wrong, but also misleading, as it implies that they were half-way toward civilization, with social changes taking place that must have had material culture correlates otherwise visible in the burial customs.

Comments

The origin of “Slavs” (i.e. that of “Slavonic” as a language, whatever the ancestral Proto-Slavic ethnic make-up was) is almost as complicated as the origin of Albanians, Basques, Balts, or Finns. Their entry into history is very recent, with few reliable sources available until well into the Middle Ages. If you add our ignorance of their origin with the desire of every single researcher or amateur out there to connect them to the own region (or, still worse, to all the regions where they were historically attested), we are bound to find contradictory data and a constantly biased selection of information.

Furthermore, it is extremely complicated to connect any recent population to its ancestral (linguistic) one through haplogroups prevalent today, and just absurd to connect them through ancestral components. This, which was already suspected for many populations, has been confirmed recently for Basques in Olalde et al. (2019) and will be confirmed soon for Finns with a study of the Proto-Fennic populations in the Gulf of Finland.

NOTE. Yes, the “my parents look like Corded Ware in this PCA” had no sense. Ever. Why adult people would constantly engage in that kind of false 5,000-year-old connections instead of learning history – or their own family history – escapes all comprehension. But if something is certain about human nature, is that we will still see nativism and ancestry/haplogroup fetishism for any modern region or modern haplogroups and their historically attested ethnolinguistic groups.

balto-slavic-pca
Genetic structure of modern Balto-Slavic populations within a European context according to the three genetic systems. Image from Kushniarevich et al. (2015)

As you can see from my maps and writings, I prefer neat and simple concepts: in linguistics, in archaeology, and in population movements. Hence my aversion to this kind of infinite proto-historical accounts (and interpretations of them) necessary to ascertain the origins of recent peoples (Slavs in this case), and my usual preference for:

  • Clear dialectal classifications, whether or not they can be as clear cut as I describe them. The only thing that sets Slavic apart from other recent languages is its connection with Baltic, luckily for both. Even though this connection is disputed by some linguists, and the question is always far from being resolved, a homeland of Proto-Balto-Slavic would almost necessarily need to be set to the north of the Carpathian Mountains in the Bronze Age (or at least close to them).
  • NOTE. A dismissal of a connection with Baltic would leave Slavic a still more complicated orphan, and its dialectal classification within Late PIE more dubious. Its union with Balto-Slavic locates it close to Germanic, and thus as a Bronze Age North-West Indo-European dialect close to northern Germany. So bear with me in accepting this connection, or enter the linguistic hell of arguing for Indo-Slavonic of R1a-Z93 mixed with Temematic….

  • A priori “pots = people” assumption, which may lead to important errors, but fewer than the usual “pots != people” of modern archaeologists. The traditional identification of the Common Slavic expansion with the Prague-Korchak culture – however undefined this culture may be – has clear advantages: it may be connected (although admittedly with many archaeological holes) with western cultures expanding east during the Bronze Age, and then west again after the Iron Age, and thus potentially also with Baltic.
  • A simplistic “haplogroup expansion = ethnolinguistic expansion”, which is quite useful for prehistoric migrations, but enters into evident contradictions as we approach the Iron Age. Common Slavs may be speculatively (for all we know) associated with an expansion of recent R1a-M458 lineages – among other haplogroups – from the east, and possibly Balto-Slavic as an earlier expansion of older subclades from the west, as I proposed in A Clash of Chiefs.
r1a-m458-underhill-2015
Modern distribution of R1a-M458, after Underhill et al. (2015).

NOTE. The connection of most R1a-Z280 lineages is more obviously done with ancient Finno-Ugric peoples, as it is clear now (see here and here).

Slavs appeared first in the Danube?

No matter what my personal preference is, one can’t ignore the growing evidence, and it seems that Florin Curta‘s long-lasting view of a Danubian origin of expansion for Common Slavic, including its condition as a lingua franca of late Avars, won’t be easy to reject any time soon:

1) Theories concerning Chernyakhov as a Slavic homeland will apparently need to be fully rejected, due to the Germanic-like ancestry that will be reported in the study by Järve et al. (2019).

EDIT (3 MAY 2019). From their poster Shift in the genetic landscape of the western Eurasian Steppe not due to Scythian dominance, but rather at the transition to the Chernyakhov culture (Ostrogoths) (download PDF):

(…) the transition from the Scythian to the Chernyakhov culture (~2,100–1,700 cal BP) does mark a shift in the Ponto-Caspian genetic landscape. Our results agree well with the Ostrogothic origins of the Chernyakhov culture and support the hypothesis that Scythian dominance was cultural rather than achieved through population replacement.

scythians-chernyakhov-ostrogoths-jarve
PCA of novel and published ancient samples from Scythian/Sarmatian and related groups on the background of modern samples presented as population medians. Δ – ref. 1, ○ – ref. 2, □ – ref. 3, ◊ – this study. Embedded are the locations of some of the samples. Notice the wide cluster formed by the three samples, from Hungarian Scythians in the west to steppe-like peoples in the east.

2) Therefore, unless Przeworsk shows the traditionally described mixture of populations in terms of ancestry and/or haplogroups, it will also be a sign of East Germanic peoples expanding south (and potentially displacing the ancestors of Slavs in either direction, east or south).

It would seem we are stuck in a Danubian vs. Kievan homeland for Common Slavs, then:

3) About the homeland in the Kiev culture, two early Avar females from Szólád have been commented to cluster “among Modern Slavic populations” based on some data in Amorim et al. (2018).

Rather than supporting an origin of Slavs in common with modern Russians, Poles, and Ukranians as observed in the PCA, though, the admixture of AV1 and AV2 (ca. AD 540-640) paradoxically supports an admixture of Modern Slavs of Eastern Europe in common with early Avar peoples (an Altaic-speaking population) and other steppe groups with an origin in East Asia… So this admixture would actually support a western origin of the Common Slavs with which East Asian Avars may have admixed, and whose descendants are necessarily sampled at later times.

pca-medieval-avar-longobards
Procrustes transformed PCA of medieval ancient samples against POPRES imputed SNP dataset. AV1 and Av2 samples have been circled in red. Color coding of medieval samples is same as in Figs 1 and 2. Two letter and three codes for POPRES samples: AL=Albania, AT=Austria, BA=Bosnia-Herzegovina, BE=Belgium, BG=Bulgaria, CH=Switzerland, CY=Cyprus, CZ=Czech Republic, DE=Germany, DK=Denmark, ES=Spain, FI=Finland, FR=France, GB=United Kingdom, GR, Greece, HR=Croatia, HU=Hungary, IE=Ireland, IT=Italy, KS=Kosovo, LV=Latvia, MK=Macedonia, NO=Norway, NL=Netherlands, PL=Poland, PT=Portugal, RO=Romania, SM=Serbia and Montenegro, RU=Russia, Sct=Scotland, SE=Sweden, SI=Slovenia, SK=Slovakia, TR=Turkey, UA=Ukraine.

4) Favouring Curta’s Danubian origin (or even an origin near Bohemia) at the moment are thus:

  • The “western” cluster of Early Slavs from Brandýsek, Bohemia (ca. AD 600-900).
  • Two likely Slavic individuals from Usedom, in Mecklenburg-Vorpommern (AD 1200) show hg. R1a-M458 and E1b-M215 (Freder 2010).
  • An early West Slav individual from Hrádek nad Nisou in Northern Bohemia (ca. AD 1330) also shows E1b-M215 (Vanek et al. 2015).
  • One sample from Székkutas-Kápolnadülő (SzK/239) among middle or late Avars (ca. AD 650-710), a supposed Slavonic-speaking polity, of hg. E1b-V13.
  • Two samples from Karosc (K1/13, and K2/6) among Hungarian conquerors (ca. AD 895-950), likely both of hg. E1b-V13, probably connected to the alliance with Moravian elites.
  • Possibly a West Slavic sample from Poland in the High Middle Ages (see below).

A later Hungarian sample (II/53) from the Royal Basilica, where King Béla was interred, of hg. E1b1, supports the importance of this haplogroup among elite conquerors, although its original relation to the other buried individuals is unknown.

NOTE. You can see all ancient samples of haplogroup E to date on this Map of ancient E samples, with care to identify the proper subclades related to south-eastern Europe. About the ancestral origin of the haplogroup in Europe, you may read Potential extra Iberomaurusian-related gene flow into European farmers, by Chad Rohlfsen.

Even assuming that the R1a sample reported from the late Avar period is of a subclade typically associated with Slavs (I know, circular reasoning here), which is not warranted, we would have already 6 E1b1b vs. 1-2 R1a-M458 in populations that can be actually assumed to represent early Slavonic speakers (unlike many earlier cultures potentially associated with them), clearly earlier than other Slavic-speaking populations that will be sampled in eastern Europe. It is more and more likely that Early Slavs are going to strengthen Curta’s view, and this may somehow complicate the link of Proto-Slavic with eastern European BA cultures like Trzciniec or Lusatian.

NOTE. I am still expecting a clear expansion associated with Prague-Korchak, though, including a connection with bottlenecks based on R1a-M458 in the Middle Ages, whether the expansion is eventually shown to be from the west (i.e. Bohemia -> Prague -> Korchak), or from the east (i.e. Kiev -> Korchack -> Prague), and whether or not this cultural community was later replaced by other ‘true’ Slavonic-speaking cultures through acculturation or population movements.

slavic-origins
Common theories on Slavic origins.. After “The Early Slavs. Culture and Society in Early Medieval Europe” by P. M. Barford, Cornell University Press (2001). Image by Hxseek at Wikipedia.

5) Back to Przeworsk and the “north of the Carpathians” homeland (i.e. between the Upper Oder and the Upper Dniester), but compatible with Curta’s view: Even if Common Slavic is eventually evidenced to be driven by small migrations north and south of the Danube during the Roman Iron Age, before turning into a mostly “R1a-rich” migration or acculturation to the north in Bohemia and then east (which is what this early E1b-V13 connection suggests), this does not dismiss the traditional idea that Late Bronze Age – Iron Age central-eastern Europe was the Proto-Slavic homeland, i.e. likely the Pomeranian culture disturbed by the East Germanic migrations first (in Przeworsk), and the migrations of steppe nomads later (around the Danube).

Even without taking into account the connection with Baltic, the relevance of haplogroup E1b-V13 among Early Slavs may well be a sign of an ancestral population from the northern or eastern Carpathian region, supported by the finding of this haplogroup among the westernmost Scythians. The expansion of some modern E1b-CTS1273 lineages may link Slavic ancestrally with the Lusatian culture, which is an eastern (very specific) Urnfield culture group, stemming from central-east Europe.

An important paper in this respect is the upcoming Zenczak et al., where another hg. E1b1 will be added to the list above: such a sample is expected from Poland (from Kowalewko, Maslomecz, Legowo or Niemcza), either from the Roman Iron Age or Early Middle Ages, close to an early population of likely Scandinavian origin (eight I1 samples), apart from other varied haplogroups, with little relevance of R1a. Whether this E-V13 sample is an Iron Age one (justifying the bottleneck under E-V13 to the south) or, maybe more likely, a late one from the Middle Ages (maybe supporting a connection of the Gothic/Slavic E1b bottleneck with southern Chernyakhov or further west along the Danube) is unclear.

The finding of south-eastern European ancestry and lineages in both, Early Slavs and East Germanic tribes* suggests therefore a Slavonic homeland near (or within) the Przeworsk culture, close to the Albanoid one, as proposed based on topohydronymy. This may point to a complex process of acculturation of different eastern European populations which formed alliances, as was common during the Iron Age and later periods, and which cannot be interpreted as a clear picture of their languages’ original homeland and ancestral peoples (in the case of East Germanic tribes, apparently originally expanding from Scandinavia under strong I1 bottlenecks).

* Iberian samples of the Visigothic period in Spain show up to 25% E1b-V13 samples, with a mixture of haplogroups including local and foreign lineages, as well as some more E1b-V13 samples later during the Muslim period. Out of the two E1b samples from Longobards in Amorim et al. (2018), only SZ18 from Szólád (ca. AD 412-604) is within E1b-V13, in a very specific early branch (SNP M35.2), further locating the expansion of hg. E1b-V13 near the Danube. Samples of haplogroup J (maybe J2a) or G2a among Germanic tribes (and possibly in Poland’s Roman Iron Age / Early Middle Ages) are impossible to compare with early Hungarian ones without precise subclades.

east-slavic-expansion
East Slavic expansion in topo-hydronymy. Image from (Udolph 1997, 2016).

I already interpreted the earlier Slavic samples we had as a sign of a Carpathian origin and very recent bottlenecks under R1a lineages among Modern Slavs:

The finding of haplogroup E1b1b-M215 in two independent early West Slavic individuals further supports that the current distribution of R1a1a1b1a-Z282 lineages in Slavic populations is the product of recent bottlenecks. The lack of a precise subclade within the E1b1b-M215 tree precludes a proper interpretation of a potential origin, but they are probably under European E1b1b1a1b1-L618 subclade E1b1b1a1b1a-V13 (formed ca. 6100 BC, TMRCA ca. 2800 BC), possibly under the mutation CTS1273 (formed ca. 2600 BC, TMRCA ca. 2000 BC), in common with other ancient populations around the Carpathians (see below §viii.11. Thracians and Albanians). This gross geographic origin would support the studies of the Common Slavic homeland based on toponymy (Figure 66), which place it roughly between the Upper Oder and the Upper Dniester, north of the Carpathians (Udolph 1997, 2016).

EDIT (8 APR 2019): Another interesting data is the haplogroup distribution among Modern Slavs and neighbouring peoples (see Wikipedia). For example, the bottleneck seen in Modern Albanians, under Z5017 subclade, also points to an origin of the expansion of E1b-V13 subclades among multiethnic groups around the Lower Danube coinciding with the Roman Iron Age, given the estimates for the arrival of Proto-Albanian close to the Latin and Greek linguistic frontier.

Remarkable is also its distribution among Rusyns, East Slavs from the Carpathians not associated with the Kievan Rus’, isolated thus quite soon from East Slavic expansions to the east. They were reported to show ca. 35% hg. E1b-V13 globally in FTDNA, with a frequency similar to or higher than R1a, in common with South Slavic peoples*, reflecting thus a situation similar to the source of East Slavs before further R1a-based bottlenecks (and/or acculturation events) to the east:

* Although probably due in part to founder effects and biased familial sampling, this should be assumed to be common to all FTDNA sampling, anyway.

rusyns-map
Map showing the full geographic extent of the Rusyn people in Central Europe, prior to World War I (Carpatho Rusyn Society).

Repeating what should be already evident: in complex organizations and/or demographically dense populations (more common since the Iron Age), we can’t expect language change to happen in the same way as during the known Neolithic or Chalcolithic population replacements, be it in Finland, Hungary, Iberia, or Poland. For example, no matter whether Romans (2nd c. BC) brought some R1b-U152 and other Mediterranean lineages to Iberia; Germanic peoples entering Hispania (AD 5th c.) were of typically Germanic lineages or not; Muslims who spoke mainly Berber (AD 8th c.) and were mainly of hg. E1b-M81 (and J?) brought North African ancestry; etc. the language or languages of Iberia changed (or not) with the political landscape: neither with radical population replacements (or full population continuity), nor with the dominant haplogroups’ ancestral language.

Y-chromosome haplogroups are, in those cases, useful for ascertaining a more recent origin of the population. Like the finding of certain R1a-Z645, I2a-L621 & N-L392 lineages among Hungarians shows a recent origin near the Trans-Urals forest-steppes, or the finding of I1, R1b-U106 & E1b-V13 among Visigoths shows a recent origin near the Danube, the finding of Early Slavs (ca. AD 6th-7th c.) originally with small elite groups of hg. R1a-M458 & E1b-V13 from the Lower/Middle Danube – if strengthened with more Early Slavic samples, with Slavonic partially expanding as a lingua franca in some regions – is not necessarily representative of the Proto-Slavic community, just as it is clearly not representative of the later expansion of Slavic dialects. It would be representative, though, of the same processes of acculturation repeated all over Eurasia at least since the Iron Age, where no genetic continuity can be found with ancestral languages.

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