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


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.

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.

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.
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

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. ( and Natural Earth data ( 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.

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. ( and Natural Earth data ( 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.

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.

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 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.

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?

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’.

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 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.


N1c-L392 associated with expanding Turkic lineages in Siberia


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].

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).

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).

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.

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.

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.


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


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.

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.

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.

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.

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.


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


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.

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


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”).


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.

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


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.


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.

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.


(…) 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. (…)

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.


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.

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.
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.

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.

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.

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 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.

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.


Arrival of steppe ancestry with R1b-P312 in the Mediterranean: Balearic Islands, Sicily, and Iron Age Sardinia


New preprint The Arrival of Steppe and Iranian Related Ancestry in the Islands of the Western Mediterranean by Fernandes, Mittnik, Olalde et al. bioRxiv (2019)

Interesting excerpts (emphasis in bold; modified for clarity):

Balearic Islands: The expansion of Iberian speakers

Mallorca_EBA dates to the earliest period of permanent occupation of the islands at around 2400 BCE. We parsimoniously modeled Mallorca_EBA as deriving 36.9 ± 4.2% of her ancestry from a source related to Yamnaya_Samara; (…). We next used qpAdm to identify “proximal” sources for Mallorca_EBA’s ancestry that are more closely related to this individual in space and time, and found that she can be modeled as a clade with the (small) subset of Iberian Bell Beaker culture associated individuals who carried Steppe-derived ancestry (p=0.442).

Suppl. Materials: The model used was with Bell_Beaker_Iberia_highsteppe, a group of outliers from Iberia buried in a Bell Beaker mortuary context who unlike most individuals from this context in that region had high proportions of Steppe ancestry (p=0.442).

Our estimates of Steppe ancestry in the two later Balearic Islands individuals are lower than the earlier one: 26.3 ± 5.1% for Formentera_MBA and 23.1 ± 3.6% for Menorca_LBA, but the Middle to Late Bronze Age Balearic individuals are not a clade relative to non-Balearic groups. Specifically, we find that f4(Mbuti.DG, X; Formentera_MBA, Menorca_LBA) is positive when X=Iberia_Chalcolithic (Z=2.6) or X=Sardinia_Nuragic_BA (Z=2.7). While it is tempting to interpret the latter statistic as suggesting a genetic link between peoples of the Talaiotic culture of the Balearic islands and the Nuragic culture of Sardinia, the attraction to Iberia_Chalcolithic is just as strong, and the mitochondrial haplogroup U5b1+16189+@16192 in Menorca_LBA is not observed in Sardinia_Nuragic_BA but is observed in multiple Iberia_Chalcolithic individuals. A possible explanation is that both the ancestors of Nuragic Sardinians and the ancestors of Talaiotic people from the Balearic Islands received gene flow from an unsampled Iberian Chalcolithic-related group (perhaps a mainland group affiliated to both) that did not contribute to Formentera_MBA.

This sample, like another one in El Argar, is of hg. R1b-P312. So there you are, the data that connects the Proto-Iberian expansion (replacing IE-speaking Bell Beakers) to the Iberian Chalcolithic population, signaled by the increase in Iberian Chalcolithic ancestry after the arrival of Bell Beakers, most likely connected originally to the Argaric and post-Argaric expansions during the MBA.

PCA with previously published ancient individuals (non-filled symbols), projected onto variation from present-day populations (gray squares).

Steppe in Sardinia IA: Phocaeans from Italy?

Most Sardinians buried in a Nuragic Bronze Age context possessed uniparental haplogroups found in European hunter-gatherers and early farmers, including Y-haplogroup R1b1a[xR1b1a1a] which is different from the characteristic R1b1a1a2a1a2 spread in association with the Bell Beaker complex. An exception is individual I10553 (1226-1056 calBCE) who carried Y-haplogroup J2b2a, previously observed in a Croatian Middle Bronze Age individual bearing Steppe ancestry, suggesting the possibility of genetic input from groups that arrived from the east after the spread of first farmers. This is consistent with the evidence of material culture exchange between Sardinians and mainland Mediterranean groups, although genome-wide analyses find no significant evidence of Steppe ancestry so the quantitative demographic impact was minimal.

Another interesting data, these (Mesolithic) remnant R1b-V88 lineages closely related to the Italian Peninsula, the most likely region of expansion of these lineages into Africa, in turn possibly connected to the expansion of Proto-Afroasiatic.

We detect definitive evidence of Iranian-related ancestry in an Iron Age Sardinian I10366 (391-209 calBCE) with an estimate of 11.9 ± 3.7.% Iran_Ganj_Dareh_Neolithic related ancestry, while rejecting the model with only Anatolian_Neolithic and WHG at p=0.0066 (Supplementary Table 9). The only model that we can fit for this individual using a pair of populations that are closer in time is as a mixture of Iberia_Chalcolithic (11.9 ± 3.2%) and Mycenaean (88.1 ± 3.2%) (p=0.067). This model fits even when including Nuragic Sardinians in the outgroups of the qpAdm analysis, which is consistent with the hypothesis that this individual had little if any ancestry from earlier Sardinians.

Proportions of ancestry using a distal qpAdm framework on an individual basis (a), and based on qpWave clusters

Sicily EBA: The Lusitanian/Ligurian connection?

(…) While a previously reported Bell Beaker culture-associated individual from Sicily had no evidence of Steppe ancestry, (…) we find evidence of Steppe ancestry in the Early Bronze Age by ~2200 BCE. In distal qpAdm, the outlier Sicily_EBA11443 is parsimoniously modeled as harboring 40.2 ± 3.5% Steppe ancestry, and the outlier Sicily_EBA8561 is parsimoniously modeled as harboring 23.3 ± 3.5% Steppe ancestry. (…) The presence of Steppe ancestry in Early Bronze Age Sicily is also evident in Y chromosome analysis, which reveals that 4 of the 5 Early Bronze Age males had Steppe-associated Y-haplogroup R1b1a1a2a1a2. (Online Table 1). Two of these were Y-haplogroup R1b1a1a2a1a2a1 (Z195) which today is largely restricted to Iberia and has been hypothesized to have originated there 2500-2000 BCE. This evidence of west-to-east gene flow from Iberia is also suggested by qpAdm modeling where the only parsimonious proximate source for the Steppe ancestry we found in the main Sicily_EBA cluster is Iberians.

What’s this? An ancestral connection between Sicel Elymian and Galaico-Lusitanian or Ligurian (based on an origin in NE Iberia)? Impossible to say, especially if the languages of these early settlers were replaced later by non-Indo-European speakers from the eastern Mediterranean, and by Indo-European speakers from the mainland closely related to Proto-Italic during the LBA, but see below.

Regarding the comment on R1b-Z195, it is associated with modern Iberians, as DF27 in general, due to founder effects beyond the Pyrenees. It is a very old subclade, split directly from DF27 roughly at the same time as it split from the parent P312, i.e. it can be found anywhere in Europe, and it almost certainly accompanied the expansion of Celts from Central Europe under the subclade R1b-M167/SRY2627.

The connection is thus strong only because of the qpAdm modeling, since R1b-DF27 and subclade R1b-Z195 are certainly lineages expanded quite early, most likely with Yamna settlers in Hungary and East Bell Beakers.

In this case, if stemming from Iberia, it is most likely of subclade R1b-Z220 – or another Z195 (xM167) lineage – originally associated with the Old European substrate found in topo-hydronymy in Iberia, whose most likely remnants attested during the Iron Age were Lusitanians.

Left: Modern distribution of R1b-Z195 (YFull estimate 2700 BC); Right: Modern distribution of DF27. Both include later founder effects within Iberia, so the increase in the Basque country and the Crown of Aragon and the decrease in Portugal can safely be ignored. Contour maps of the derived allele frequencies of the SNPs analyzed in Solé-Morata et al. (2017).

We detect Iranian-related ancestry in Sicily by the Middle Bronze Age 1800-1500 BCE, consistent with the directional shift of these individuals toward Mycenaeans in PCA. Specifically, two of the Middle Bronze Age individuals can only be fit with models that in addition to Anatolia_Neolithic and WHG, include Iran_Ganj_Dareh_Neolithic. The most parsimonious model for Sicily_MBA3125 has 18.0 ± 3.6% Iranian-related ancestry (p=0.032 for rejecting the alternative model of Steppe rather than Iranian-related ancestry), and the most parsimonious model for Sicily_MBA has 14.9 ± 3.9% Iranian-related ancestry (p=0.037 for rejecting the alternative model).

The modern southern Italian Caucasus-related signal identified in Raveane et al. (2018) is plausibly related to the same Iranian-related spread of ancestry into Sicily that we observe in the Middle Bronze Age (and possibly the Early Bronze Age).

The non-Indo-European Sicanians and Elymians were possibly then connected to eastern Mediterranean groups before the expansion of the Sea Peoples.

For the Late Bronze Age group of individuals, qpAdm documented Steppe-related ancestry, modeling this group as 80.2 ± 1.8% Anatolia_Neolithic, 5.3 ± 1.6% WHG, and 14.5 ± 2.2% Yamnaya_Samara. Our modeling using sources more closely related in space and time also supports Sicily_LBA having Minoan-related ancestry or being derived from local preceding populations or individuals with ancestries similar to those of Sicily_EBA3123 (p=0.527), Sicily_MBA3124 (p=0.352), and Sicily_MBA3125 (p=0.095).

This increase in Steppe-related ancestry in a western site during the LBA most likely represents either an expansion from the Aegean or – maybe more likely, given the archaeological finds – a regional population similar to Sicily EBA re-emerging or rather being displaced from the eastern part of the island because of a westward movement from nearby Calabria.

Whether this population sampled spoke Indo-European or not at this time is questionable, since the Iron Age accounts show non-IE Elymians in this region.

Actually, Elymians seem to have spoken Indo-European, which fits well with the increase in steppe ancestry.

EDIT (21 MAR): Interesting about a proposed incoming Minoan-like ancestry is the potential origin of the Iran Neolithic-related ancestry that is going to appear in Central Italy during the LBA. This could then be potentially associated with Tyrsenians passing through the area, although the traditional description may be more more compatible with an arrival of Sea Peoples from the Adriatic.

Sad to read this:

This manuscript is dedicated to the memory of Sebastiano Tusa of the Soprintendenza del Mare in Palermo, who would have been an author of this study had he not tragically died in the crash of Ethiopia Airlines flight 302 on March 10.


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


We know that the Caucasus Mountains formed a persistent prehistoric barrier to cultural and population movements. Nevertheless, an even more persistent frontier to population movements in Europe, especially since the Neolithic, is the Pontic-Caspian steppe – forest-steppe ecotone.

Like the Caucasus, this barrier could certainly be crossed, and peoples and cultures could permeate in both directions, but there have been no massive migrations through it. The main connection between both regions (steppe vs. forest-steppe/forest zone) was probably through its eastern part, through the Samara region in the Middle Volga.

The chances of population expansions crossing this natural barrier anywhere else seem quite limited, with a much less porous crossing region in the west, through the Dnieper-Dniester corridor.

A Persistent ecological and cultural frontier

It is very difficult to think about any culture that transgressed this persistent ecological and cultural frontier: many prehistoric and historical steppe pastoralists did appear eventually in the neighbouring forest-steppe areas during their expansions (e.g. Yamna, Scythians, or Turks), as did forest groups who permeated to the south (e.g. Comb Ware, GAC, or Abashevo), but their respective hold in foreign biomes was mostly temporary, because their cultures had to adapt to the new ecological environment. Most if not all groups originally from a different ecological niche eventually disappeared, subjected to renewed demographic pressure from neighbouring steppe or forest populations…

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

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

Before the emergence of pastoralism, the cultural contacts of the Pontic region (i.e. forest-steppes) with the Baltic were intense. In fact, the connection of the north Pontic area with the Baltic through the Dnieper-Dniester corridor and the Podolian-Volhynian region is essential to understand the spread of peoples of post-Maglemosian and post-Swiderian cultures (to the south), hunter-gatherer pottery (to the north), TRB (to the south), Late Trypillian groups (north), GAC (south), or Comb Ware (south) (see here for Eneolithic movements), and finally steppe ancestry and R1a-Z645 with Corded Ware (north). After the complex interaction of TRB, Trypillia, GAC, and CWC during the expansion of late Repin, this traditional long-range connection is lost and only emerges sporadically, such as with the expansion of East Germanic tribes.

A barrier to steppe migrations into northern Europe

One may think that this barrier was more permeable, then, in the past. However, the frontier is between steppe and forest-steppe ecological niches, and this barrier evolved during prehistory due to climate changes. The problem is, before the drought that began ca. 4000 BC and increased until the Yamna expansion, the steppe territory in the north Pontic region was much smaller, merely a strip of coastal land, compared to its greater size ca. 3300 BC and later.

This – apart from the cultural and technological changes associated with nomadic pastoralism – justifies the traditional connection of the north Pontic forest-steppes to the north, broken precisely after the expansion of Khvalynsk, as the north Pontic area became gradually a steppe region. The strips of north Pontic and Azov steppes and Crimea seem to have had stronger connections to the Northern Caucasus and Northern Caspian steppes than with the neighbouring forest-steppe areas during the Upper Palaeolithic, Mesolithic, and Neolithic.

NOTE. We still don’t know the genetic nature of Mikhailovka or Ezero, steppe-related groups possibly derived from Novodanilovka and Suvorovo close to the Black Sea (which possibly include groups from the Pannonian plains), and how they compare to neighbouring typically forest-steppe cultures of the so-called late Sredni Stog groups, like Dereivka or partly Kvityana.

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

Despite the Pontic-Caspian steppes and forest-steppes neighbouring each other for ca. 2,000 km, peoples from forested and steppe areas had an obvious advantage in their own regions, most likely due to the specialization of their subsistence economy. While this is visible already in Palaeolithic and Mesolithic hunter-gatherers, the arrival of the Neolithic package in the Pontic-Caspian region incremented the difference between groups, by spreading specialized animal domestication. The appearance of nomadic pastoralism adapted to the steppe, eventually including the use of horses and carts, made the cultural barrier based on the economic know-how even stronger.

Even though groups could still adapt and permeate a different territory (from steppe to forest-steppe/forest and vice-versa), this required an important cultural change, to the extent that it is eventually complicated to distinguish these groups from neighbouring ones (like north-west Pontic Mesolithic or Neolithic groups and their interaction with the steppes, Trypillia-Usatovo, Scythians-Thracians, etc.). In fact, this steppe – forest-steppe barrier is also seen to the east of the Urals, with the distinct expansion of Andronovo and Seima-Turbino/Andronovo-like horizons, which seem to represent completely different ethnolinguistic groups.

As a result of this cultural and genetic barrier, like that formed by the Northern Caucasus:

1) No steppe pastoralist culture (which after the emergence of Khvalynsk means almost invariably horse-riding, chariot-using nomadic herders who could easily pasture their cows in the huge grasslands without direct access to water) has ever been successful in spreading to the north or north-west into northern Europe, until the Mongols. No forest culture has ever been successful in expanding to the steppes, either (except for the infiltration of Abashevo into Sintashta-Potapovka).

2) Corded Ware was not an exception: like hunter-gatherer pottery before it (and like previous population movements of TRB, late Trypillia, GAC, Comb Ware or Lublin-Volhynia settlers) their movements between the north Pontic area and central Europe happened through forest-steppe ecological niches due to their adaptation to them. There is no reason to support a direct connection of CWC with true steppe cultures.

3) The so-called “Steppe ancestry” permeated the steppe – forest-steppe ecotone for hundreds of years during the 5th and early 4th millennium BC, due to the complex interaction of different groups, and probably to the aridization trend that expanded steppe (and probably forest-steppe) to the north. Language, culture, and paternal lineages did not cross that frontier, though.

EDIT (4 FEB 2019): Wang et al. is out in Nature Communications. They deleted the Yamna Hungary samples and related analyses, but it’s interesting to see where exactly they think the trajectory of admixture of Yamna with European MN cultures fits best. This path could also be inferred long ago from the steppe connections shown by the Yamna Hungary -> Bell Beaker evolution and by early Balkan samples:

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


Happy new year 2019…and enjoy our new books!


Sorry for the last weeks of silence, I have been rather busy lately. I am having more projects going on, and (because of that) I also wanted to finish a project I have been working on for many months already.

I have therefore decided to publish a provisional version of the text, in the hope that it will be useful in the following months, when I won’t be able to update it as often as I would like to:

EDIT (20 JAN 2019): For those of you who are more comfortable reading in your native language, I have placed some links to automatic translations by Google Translate. They might work especially well for the texts of A Game of Clans & A Clash of Chiefs.

Don’t forget to check out the maps included in the supplementary materials: I have added Y-DNA, mtDNA, and ADMIXTURE data using GIS software. The PCA graphics are also important to follow the main text.

NOTE. Right now the files are only in my server. I will try to upload them to and Research Gate when I have time, I have uploaded them to and ResearchGate, in case the websites are too slow.

I would have preferred to wait for a thorough revision of the section on archaeology and the linguistic sections on Uralic, but I doubt I will have time when the reviews come, so it was either now or maybe next December…

I say so in the introduction, but it is evident that certain aspects of the book are tentative to say the least: the farther back we go from Late Proto-Indo-European, the less clear are many aspects. Also, linguistically I am not convinced about Eurasiatic or Nostratic, although they do have a certain interest when we try to offer a comprehensive view of the past, including ethnolinguistic identities.

I cannot be an expert in everything, and these books cover a lot. I am bound to publish many corrections as new information appears and more reviews are sent. For example, just days ago (before SNP calls of Wang et al. 2018 were published) some paragraphs implied that AME might have expanded Nostratic from the Middle East. Now it does not seem so, and I changed them just before uploading the text. That’s how tentative certain routes are, and how much all of this may change. And that only if we accept a Nostratic phylum…

NOTE. Since the first book I wrote was the linguistic one, and I have spent the last months updating the archaeology + genetics part, now many of you will probably understand 1) why I am so convinced about certain language relationships and 2) how I used many posts to clarify certain ideas and receive comments. Many posts offer probably a good timeline of what I worked with, and when.


I did not add this section to the books, because they are still not ready for print, but I think this is due somewhere now. It is impossible to reference all who have directly or indirectly contributed to this, so this is a list of those I feel have played an important role.

I am indebted to the following people (which does not mean that they share my views, obviously):

First and foremost, to Fernando López-Menchero, for having the patience to review with detail many parts on Indo-European linguistics, knowing that I won’t accept many of his comments anyway. The additional information he offers is invaluable, but I didn’t want to turn this into a huge linguistic encyclopaedia with unending discussions of tiny details of each reconstructed word. I think it is already too big as it is.

I would not have thought about doing this if it were not for the interest of Wekwos (Xavier Delamarre) in publishing a full book about the Indo-European demic diffusion model (in the second half of 2017, I think). It was them who suggested that I extended the content, when all I had done until then was write an essay and draw some maps in my free time between depositing the PhD thesis and defending it.

Sadly, as much as I would like to publish a book with a professional publisher, I don’t think ancient DNA lends itself for the traditional format, so my requests (mainly to have free licenses and being able to review the text at will, as new genetic papers are published) were logically not acceptable. Also, the main aim of all volumes, especially the linguistic one, is the teaching of essentials of Late Proto-Indo-European and related languages, and this objective would be thwarted by selling each volume for $50-70 and only in printed format. I prefer a wider distribution.

At first I didn’t think much of this proposal, because I do not benefit from this kind of publications in my scientific field, but with time my interest in writing a whole, comprehensive book on the subject grew to the point where it was already an ongoing project, probably by the start of 2018.

I would not have been in contact with Wekwos if it were not for user Camulogène Rix at Anthrogenica, so thanks for that and for the interest in this work.

I would not have thought of writing this either if not for the spontaneous support (with an unexpected phone call!) of a professor of the Complutense University of Madrid, Ángel Gómez Moreno, who is interested in this subject – as is his wife, a professor of Classics more closely associated to Indo-European studies, and who helped me with a search for Indo-Europeanists.

EDIT (1 JAN 2019): I remembered that Karin Bojs sent me her book after reading the demic diffusion model. I may have also thought about writing a whole book back then, but mid-2017 is probably too early for the project.

Professor Kortlandt is still to review the text, but he contributed to both previous essays in some very interesting ways, so I hope he can help me improve the parts on Uralic, and maybe alternative accounts of expansion for Balto-Slavic, depending on the time depth that he would consider warranted according to the Temematic hypothesis.

The maps are evidently (for those who are interested in genetics) in part the result of the effort of the late Jean Manco: As you can see from the maps including Y-DNA and mtDNA samples, I have benefitted from her way of organising data and publishing it. Similarly, the work of Iain McDonald in assessing the potential migration routes of R1b and R1a in Europe with the help of detailed maps was behind my idea for the first maps, and consequently behind these, too.

I should thank all people responsible for the release of free datasets to work with, including the Reich and Jena labs, the Veeramah Lab, and also researchers from the Max Planck Institute or the Mainz Palaeogenetics group, who didn’t mind to share with me datasets to work with.

Readers of this blog with interesting comments have also been essential for the improvement of the texts. You can probably see some of your many contributions there. I may not answer many comments, because I am always busy (and sometimes I just don’t have anything interesting to say), but I try to read all of them.

EDIT (1 JAN 2019) I think I should mention at least Chetan, Egg, or Robert George; but then I would leave out old europe, Sgr Ganesh, or Tileman Ehlen; and if I include them I would leave out others…

Users of other sites, like Anthrogenica, whose particular points of view and deep knowledge of some very specific aspects are sometimes very useful. In particular, user Anglesqueville helped me to fix some issues with the merging of datasets to obtain the PCAs and ADMIXTURE, and prepared some individual samples to merge them.

Even without posting anything, Google Analytics keeps sending me messages about increasing user fidelity (returning users), and stats haven’t really changed (which probably means more people are reading old posts), so thank you for that.

I hope you enjoy the books.

Happy new year!

Biparental inheritance of mitochondrial DNA in humans


New paper Biparental Inheritance of Mitochondrial DNA in Humans, by Luo et al. PNAS (2018).

Interesting excerpts (emphasis mine):


Although there has been considerable debate about whether paternal mitochondrial DNA (mtDNA) transmission may coexist with maternal transmission of mtDNA, it is generally believed that mitochondria and mtDNA are exclusively maternally inherited in humans. Here, we identified three unrelated multigeneration families with a high level of mtDNA heteroplasmy (ranging from 24 to 76%) in a total of 17 individuals. Heteroplasmy of mtDNA was independently examined by high-depth whole mtDNA sequencing analysis in our research laboratory and in two Clinical Laboratory Improvement Amendments and College of American Pathologists-accredited laboratories using multiple approaches. A comprehensive exploration of mtDNA segregation in these families shows biparental mtDNA transmission with an autosomal dominantlike inheritance mode. Our results suggest that, although the central dogma of maternal inheritance of mtDNA remains valid, there are some exceptional cases where paternal mtDNA could be passed to the offspring. Elucidating the molecular mechanism for this unusual mode of inheritance will provide new insights into how mtDNA is passed on from parent tooffspring and may even lead to the development of new avenues for the therapeutic treatment for pathogenic mtDNA transmission.

An example

Compared with Family A, a strikingly similar mtDNA transmission pattern was demonstrated in Families B and C. Taking Family B for illustration, II-3 having 29 heteroplasmic and seven homoplasmic variants should have inherited mtDNA from both his father (I-1, haplogroup of K1b2a) and his mother (I-10, haplogroup of H), who were supposed to possess 34 and nine homoplasmic variants, respectively. II-3 further transmitted his mtDNA that he inherited from I-1 to his son (III-2), who also inherited all of his mother’s mtDNA (II-30, carrying 34 variants and a haplogroup of T2a1a). However, III-2’s sister (III-1) and half-brother (III-5) only inherited the maternal mtDNA. Fresh blood sampling and repeated mtDNA sequencing in a second independent laboratory were also performed to rule out the possibility of sample mix-up for III-2 (III-2, column F-G and H-I). Additionally, these samples were further verified using Pacific Bio single molecular sequencing (see Materials and Methods) and by restriction fragment length polymorphism (RFLP) analysis of Family A, and these results were fully consistent with the previous sequencing.

Biparental mtDNA inheritance pattern shown in Family B. (A) Pedigree of Family B. The black filled symbols indicate the two family members (II-3 and III-2) showing biparental mtDNA transmission. The IDs of five family members tested by whole mtDNA sequencing analysis have been underlined in the pedigree. (B) Schematic of the mtDNA genotype defined by the homoplasmic and/or heteroplasmic variants aligned from the reference mitochondrial genome. Blue bars represent the genotype of paternally derived mtDNA, whereas purple-red and orange-red bars represent maternally derived mtDNA. Entries labeled (D) represent deduced mtDNA genotypes. (C) Summary of the haplogroup and mtDNA variant numbers in Family B.

A Resurgence of the Paternal Transmission Hypothesis

The results presented in this paper make a robust case for paternal transmission of mtDNA. Here, we report biparental mtDNA inheritance (either directly or indirectly) in 17 members in three multigeneration families. Thirteen of these individuals were identified directly by sequencing of the mitochondrial genome, whereas four could be inferred based on preexisting maternal heteroplasmy caused by biparental inheritance in the previous generation.

To further confirm these remarkable results and to exclude the possibility of sample mix-up and/or contamination, the whole mtDNA sequencing procedure was repeated independently in at least two different laboratories by different laboratory technicians with newly obtained blood samples. All results were reproducible, indicating no artifacts or contamination exist. More importantly, the multiple mtDNA variants that were paternally transmitted differ in both number and position among each of these three families as well as the related haplogroup (R0a1 in Family A, K1b2a in Family B, and K2b1a1a in Family C, respectively), providing two distinct forms of evidence supporting transmission of the paternal mtDNA.

Therefore, we have unequivocally demonstrated the existence of biparental mtDNA inheritance as evidenced by the high number and level of mtDNA heteroplasmy in these three unrelated multigeneration families. Most interestingly, the mixed haplogroups in these samples are very reminiscent of the mixed haplogroups found in the 20 studies that were dismissed by Bandelt et al. as due to contamination or sample mix-up. One is forced to wonder how many other instances of individuals with biparental mtDNA inheritance have been dismissed as technical errors, and whether Schwartz and Vissing’s original discovery has really been given the proper follow-up that it deserves. We suspect that these results will initiate a broader reassessment of the topic.

We propose that the paternal mtDNA transmission in these families should be accompanied by segregation of a mutation in one nuclear gene involved in paternal mitochondrial elimination and that there is a high probability that the gene in question operates through one of the pathways identified above.

If I have to be honest, I was stuck with the paternal transmission hypothesis which we were taught in class long ago. I didn’t know it was controversial or dismissed, I just thought it was really exceptional, and I never thought about learning more on the subject.

This paper proves it may be more complicated than that, especially for population genomics purposes, because biparental mtDNA transmission with autosomal dominant-like inheritance puts a serious barrier to a general, simplistic interpretation of mtDNA.

I don’t think it is a blow to all interpretations based on mtDNA, though, because the traditional interpretation should often work statistically. However, one has to be always very careful when saying “if it’s mtDNA from region X, it’s about female exogamy”, especially when samples are from neighbouring regions and similar periods.

The term “uniparental marker” for mtDNA is obviously misleading and shouldn’t be used, and many research papers and interpretations taking mtDNA as strictly uniparental should be taken with a pinch of salt.